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fy:

PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZQVOLOGICAL SOCIETY

OF LONDON.

1911, pp. 557-1213,

witH 48 Puatres and 86 TEx-FIGURES.

LLL585 |

PRINTED FOR THE SOCIETY,

AND SOLD AT THEIR HOUSE IN REGENT’S PARK.

LONDCN:

MESSRS. LONGMANS, GREEN, AND CO,

PATERNOSTER ROW.

Ss a

OF THE

COUNCIL AND OFFICERS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

WON Te

COUNCIL.

His Grace Tut Duxke or Beprorp, K.G., President.

Tue Karu or Auramont, F.S.A.

Sir Joun Rose BRADFORD,

KC MEGS) MED EDS,

FE.R.S., Vice-President.

Lt.-Col. Srp R. HaAvetock-

Cuartes, K.C.V.O., M.D.

ALFRED H. Cocks, Esq., M.A.

The Rt. Hon. the Ear or

@rommr, | 2-0 1G-C bs

G.C.M.G.

F. G. Dawtrey Drewirt, Ksq.,

WieA\o5 WilslD).

CuarLtes DrumMonn,

Treasurer.

Str Epwarp Duranp, Bt., C.B.

FREDERICK GILLETT, Esq., Vice-

President.

Srpney F. Harmer, Ksq., M.A.,

Se.D., F.R.S., Vice-President.

Ksq.,

Sir WALTER Roper LAWRENCE,

Bt. G.C.AnK-

Sir Epmunp G. Lopmr, Bt.

EK. G. B. Meapre-WaAtpo, Esq.,

Vice-President.

P. Cuaumers Mircuet, Esq.,

MEARS UDESc:.) JElonas ae

E.B.S., Secretary.

W. R. Ocitvie-Grant, Esq.

ADRIAN D. W. Poutock, Esq.

A. Trevor- Battys, Ksgq.,

M.A.

AntHony. H.

Ksa.

A. SmirH Woopwarp, Ksq.,

LL.D.,F.R.S., Vice-President.

Henry Woopwarb, Hsq., LL.D.,

F.R.S., Vice-President.

WINGFIELD,

PRINCIPAL OFFICERS.

P. CHatmers Mircnety, M.A., D.Sc., Hon.LL.D., F.B.S.,

Secretary.

Frank HK. Bepparp, M.A., F.R.S., Prosector.

R. I. Pocock, F.R.S., F.L.S., Curator of Mammals, and

Resident Superintendent of the Gardens.

D. Sera-Suiru, Curator of Birds and Inspector of Works.

Epwarp G. Boutencer, Curator of Reptiles.

Henry G. Purmimer, F.R.S., M.R.C.S., Pathologist.

F. H. Waternoust, Librarian.

JOHN Barrow, Accountant.

W. H. Coun, Chief Clerk.

dy?

LIST OF CONTENTS.

EXHIBITIONS AND NOTICES.

Page

The Secretary. Report on the Additions to the Society's

Menagerie during the month of February 1911 ......... 557

Mr. D. Sern-Sairs, F.Z.S. Exhibition of a living Hybrid

between the White-eyed Pochard (Aythya nyroca) and

the Marbled Duck (Marmaronetta angustirostris) ...... 558

Mr. R. 1. Pocock, F.RS., F.L.S., F.Z:S. -Exhibition of

the skin of a new Chacma Baboon (Papio porearius,

subsp. griseipes), a specimen of the North American

Black-footed Polecat (Putorius nigripes), ete. (‘Texti-

Sir E. Ray Lanxestsr, K.C.B., F.R.S., F.Z.8. Exhibition

of a special supplement to the ‘ Field’ newspaper ...... 620

Dr. R. T. Lererr, F.Z.8. A demonstration of Nematode

parasites obtained from animals in the Zoological

Garden sy hk eh isdn Wea ee. cae SpE AA EE 620

Mr. R. I. Pocock, F.R.S., F.L.8., F.Z.S. Exhibition of

a newly born Masked Palm-Civet (Paradoxurus larva-

tus), with an abnormal leg. (Text-fig. 147.) ............ 621

The Secretary. A letter from the Governor of the Seychelles

on Wand=Mortoises 1m those Islands 2.0.2... .........4....- 622

Dr. H. B. Fanruam, F.Z.S., and Miss Annie Porter, D.Sc.

On a Bee-disease due to a Protozoal Parasite (Vosema

The Secretary. Report on the Additions to the Society’s

Menagerie during the month of March 1911 ............

Mr. C. Tare Reean, M.A., F.Z.S. Exhibition of a series of

lantern-slides of scales of the Salmon (Salmo salar) ...

Mr. D. Seru-Suiry, F.Z.S. Exhibition of a nest of the

Grey Struthidea or Apostle Bird (Struthidea cinerea),

and lantern-slides of Penguins in moult and of wild

Swainson’s Lorikeets (Zvrichoglossus nove-hollandic).

(EGS NGS oP ern Ahi oS Seno udavadonasudoasesG

Mr. C. Curisry, F.Z.S. Exhibition of a collection of skins

from Uganda and of a loin-cloth taken from a native

Tow Northern Nieeria: ..ceeceeneerere had cena nee

Dr. Wit11AM Nico, M.A., F.Z.8. On a unique Patholo-

gical Condition ina Hare. (Text-fig. 165.) ............

Mr. R. I. Pocock, F.R.S., F.LS., F.Z.8. Exhibition of

some hair from the “puppy coat” of a Grey Seal

(Halichcerws giiypus) a eseee ace ene eee eRe ee

Mr. E. G. Boutencer. Exhibition of living male specimens

of the Midwife Toad (Alytes obstetricans) carrying the

COGS coerce cr cece cece eee ee rece eee cer eee nesessreecr scenes ceeeeene

Mr. A. E. Anperson. Exhibition of photographs of fossil

mammals in the American Museum of Natural History .

The Secretary. Report on the Additions to the Society’s

Menagerie during the month of April 1911 ............

The Rey. A. Mines Moss, M.A., F.Z.S., F.E.S. Notice of a

Mmemomon the Sphingides oll eru case eee eeere ee eeeee

Page

671

674

696

869

Mr. H. G. Purmmer, F.R.S. Report on the Pathological .

Examination of Rats (Jus decumanus) caught in the

Regent's Park and in the Society’s Gardens

eroesccecens

Dr. R. W. Ssurexipr, C.M.Z.S. Exhibition of a photograph

of a male albino Woodchuck (Arctomys POMTAS)) saecce a0:

Mr. R. E. Hotpine. Exhibition of the Horns of a High-

land Ram, a Fallow Deer, and a Roebuck, which were

fused at the base, and the skull of a coursing Grey-

hound with abnormal dentition

Seer eee eecces sesso ceo essere

Page

Mr. R. I. Pococr, F.R.S., F.L.S., F.Z.S. Exhibition of, and

remarks upon, the skin and skull of a specimen of the

Crested Rat (Lophiomys ibeanus Thos.). (Text-fig. 190.) 946

The Secretary. Report on the Additions to the Society’s

Menagerie during the month of May 1911 ............... 985

Mr. D. Sera-Suirn, F.Z.8. Exhibition of, and remarks

upon, two immature Black-backed Porphyrios (Por-

phyrio melanonotus) with wing-claws. (Text-fig. 200.) 985

Mr. J. Lewis Bonuots, M.A., F.L.S., F.Z.S. Exhibition of

a pair of abnormally coloured Egyptian Desert-Mice

(A GPUOTMES GRUSSOIS)) REN Roe nORe eos Ane odie soe danneteeee st 986

Dr. W. T. Catman, F.Z.8. Exhibition of living specimens of

the Brine Shrimp (Artemia salina), bred from Tidman’s

SOa Calc eee tet mem ace ae teres eee reat etea te tc cecal te erases centre 986

The Secretary. Remarks upon the eggs of Struthio massai-

' cus, S. australis, and S. molybdophanes, seen at Mr. Carl

Hagenbeck’s Ostrich Farm at Stellingen.................. 987

The Secrerary. Remarks upon a pair of young African

Rhinoceros seen at Mr. Carl Hagenbeck’s Zoological

JEpue enn tee) MEU. 0 roan Jane AB ac GAR AE AAlNBn noth eb 987

Mr. R. I. Pococn, F.R.S., F.L.S., F.Z.S. Exhibition of

photographs of, and remarks upon, two hybrid foals,

Equus asinus somaliensis x Hquus zebra and Hquus

asinus somaliensis x Hquus quagga chapmanni. (Text-

{Sea AD pears epee oman dee c aes aucei md atea eae ems ocibh 988

PAPERS.

26. On the Amphipod Genus Leptocheirus. By EK. W.

Sexton, Marine Biological Laboratory, Plymouth.

(Elates Revell SXGRXe and Wextoties WAG ee sss. ce anna cee 561

27. Notes on Marine Ostracoda from Madeira. By G.

STEWARDSON Brapy, M.D., LL.D., D.Se., F.R.S.,

OpWilAds. (IPI Nias OCC OS IUD) ei hansoccs sag sbnen sean Contes 595

28.

30.

ol.

33.

34.

36.

On Colour and Colour-pattern Inheritance in Pigeons.

By J. Lewis Bonuors, M.A., F.L.S., F.Z.8., and F. W.

Sud pune H ZS. ua(Plaibes 2OXUuEL —XOXGV cl) I Eee ee

. Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea.—I. On some Mammalian

Cestoidea. By Frank E. Bepparp, M.A., F.R.S.,

F.Z.8., Prosector to the Society. (Text-figs. 148-159.).

On the Natural History of Whalebone Whales. By

J. A. Morcu, Christiania, (Text-figs. 160-163.) ......

On Three New Trematodes from Reptiles. By WiLLrAm

Nicott, M.A., D.Sc., M.B., F.Z.S. (Plates X XVII.

& XXVIII.) |

Coe eee pret ese eres ees e terse e trees sere ee essere oesees

. The Duke of Bedford’s Zoological Exploration of Hastern

Asia.— XIV. On Mammals from Southern Shen-si,

Central China. By Otprienp Tomas, F.R.S., F.Z.S.

(Plate X XIX.)

freee ees e weer sere esr ese ese esse eseeeresesseseves

An Investigation into the Validity of Miillerian and

other Forms of Mimicry, with Special Reference to

the Islands of Bourbon, Mauritius, and Ceylon. By

Nevint—E Manpers, Lieut.-Colonel, R.A.M.C., F.Z.S.,

CSCC ee eee mw ene re reer essere seseseeessesserersesessessesesseoes

The Distribution of the Avian Genus I/egapodius in the

Pacific Islands. By J.J. Lisrer, M.A., F.R.S., F.L.S.,

OAS. ai Mexctiatigs NGG:)iss nC ie BER ene Renny aon Renvee

. Contributions to the Morphology of the Group Neritacea

of the Aspidobranch Gastropods.—Part Il. The

Heticinip#, By Ginpert C, Bourne, M.A., D.Sc.,

F.RS., F.Z.8. (Plates DOK I), Reese

On the Palatability of some British Insects, with Notes

on the Significance of Mimetic Resemblances. By

R, I. Pocock, F.R.S., F.LS., F.Z.S., Superintendent

of the Society's Gardens and Curator of Mammals.

With Notes upon the Experiments by Prof. E. B.

Poutton, F.R.S8., F.Z.S.

Coe eae eee e eee cesesesoeseevsceecsevere

. The Aleyonaria of the Cape of Good Hope and Natal.—

Gorconacea. By J. Stuarr THomson, Ph.D., F.R.S.E.,

F.LS., Lecturer and Senior Demonstrator in Zoology,

University of Manchester. (Plates X LIII.—_XLV. and

Wext-He, «NG7,).ccccin kee stes sates te ees nas LT eGeaee

Page

601

626

677

696

749

759

809

870

38.

39.

40.

4].

43.

44,

47.

Vil

On the Structure of the Skull in Cynodont Reptiles.

By R. Broom, M.D., D.Sc.,-C.M.Z.S. (Plate XLVI.

and Text-figs. 168-180.)

ew ree meee eee eee eee e erst ee sesesserere

Tooth-germs in the Wallaby (Macropus billardieri). By

A. Hoprweti-SmityH, L.R.C.P., M.R.C.S., and H. W.

VAREn iss) Moa MODE hase shEZS. (Plate

DIG Ls ewavch Mescpsintets pales | alCl)) a4. bad saceash ab seoeuaaeeee

On a New Species of Dinotherium (Dinotherium hobleyi)

from British East Africa. By C. W. AnpreEws, D.Sc.,

F.R.S., F.Z.S. (British Museum, Natural History).

(RP Tasbe XoTEIVATSIMIE esse 2 casts Narcuiestctinsiu es sic ae eersamelaticticl ae a

On an Amphipod from the Transvaal. By the Hon.

Pau A. MerHuen, F.Z.8. (Plates XLIX.—LI.) ......

. An African Rhinoceros, Klipspringer, and Gazelle. By

R. Lypexker. (Text-figs. 191-193.)

weet eee eee see tesco wee

The Subspecies of the Spanish Ibex. By Prof. ANGEL

Caprera, C.M.Z.S. (Uelaues 1 ye —LIV. and Text-figs.

194— 199.) EN ata clits Ae RR ama Pe Sa lal a Re

On Antelopes of the Genera Madoqua and Rhynchotragus

found in Somaliland. By R. E. Draxs-Brockmay,

IME ChS Wak C le) EeZeS (Plates 1a\Viids IVils)en

. Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea.—II. On Two New Genera of

Cestodes from Mammals. By Frank E. Bepparp,

M.A., F.R.S., F.Z.8., Prosector to the Society.

(Text-figs. 204-215.)

. Some Madreporaria from the Persian Gulf. By Rurn

Harrison, Oxford. With a Note on the Memoir and

some Further Notes on Pyrophyllia inflata by SYDNEY

J. Hickson, M.A., D.Se, F.R.S., F.Z.8. (Plates

VIL. & LUVIN. and Text-figs. 216—221.)...............

On Variation in the Medusa of Merisia lyonsi. By

CHARLES L. Boutencrer, M.A., F.Z.S., Lecturer on

Zoology in the University of Birmingham. (Plate

EXG en eM erate SD PE 228. esc ciwia tcl so ciewesiainine oui

Page

893

958

963

977

994

1018

Vill

Page

48. The Marginal Processes of Lamellibranch Shells. By

Cyrit CrossLanb, F.Z.S. (Plate LX. and Text-figs.

IOUS gl hE Mecctle cs ieiteiais inte wiet:o Sell eieitine ae «eae tae iS oer 1057

49. Warning Coloration in a Nudibranch Molluse and in a

Chameleon. By Cyrit CrossuanD, F.Z.S. ............ 1062

50. Chromodorids from the Red Sea, collected and figured

by Mr. Cyrin CrossuanD. By Sir CHAries E ior,

TEAC INING Io, Opleen Ia Ass (Celene IX.) «occossas0a0en00c 1068

51. On some New South African Permian Reptiles. By

R. Broom, D.Sc.,C.M.Z.S. (Plates LXII. & LXIIT.). 1073

52. On a New Tree-Frog from Trinidad, living in the

Society's Gardens. By Epwarp G. BouLENGErR,

(Chennai Ci Iaveroalles” (leleiws, IDOI 3) soncaeonssocoscn6 1082

53. A Contribution to the Ornithology of Western Colombia.

By C. EK. Hetimayr, Division of Birds, Zoological

MuseunosMumich’ aa. cere ec eeerer cece reece rer er eee reenter 1084

ADDENDUM.

Additional reference to Dr. P. CHatmers MircHe.’s

memo, “On Longevity and Relative Viability in

Mammals tamdaltirds'? 2) Sete Saeeaaie oe ene 868

siprateiesd cbratslat avelue | Srtais AES ee ae Eee ee XVI1ll

indexok Scrembitie Names s.45.-)) nee eee ee X1X

siseisar ang aomaseee Resco eee ae eee LEE XXXVIil1

AU RSIVAG BS MN CYAN pasar

OF TIIK

COND Wye Oks;

With References to the several Articles contributed by each.

1911, pp. 557-1218.

Anpprson, A. EK.

Exhibition of photographs of fossil mammals in the

American Museum of Natural History

ANDREWS, CHARLES W., D.Sc., F.R.S., F.Z.8. (British

Museum, Natural History).

On a New Species of Dinotherium (Dinotherium

hobleyt) from British East Africa. (Plate XLVIII.)

Bepparb, Frank E., M.A., F.R.S., F.Z.S., Prosector to the

Society.

Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea.—I. On some Mammalian

Cestodeawn Hexboiess U4 Salo) imme ttm ite ane vacultns

696

943

626

Page

BEDDARD, Frank E., M.A., F.R.S., F.Z.8. (Continwed.)

Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea—II. On Two New Genera of

Cestodes from Mammals. (Text-figs. 204-215.) ......... 994

Bonnore, J. Lewis, M.A., F.LS., F.Z:8.

Exhibition of a pair of abnormally coloured Egyptian

ID eseitin Wess (CUI IOIES GRUSSUIS) \sosascongsaaddoscoba5snonanAsosor 986

Bonnore, J. Lewis, M.A., F.L8., F.Z.S., and Smatiey,

FREDERIC W., F.Z.S.

On Colour and Colour-pattern Inheritance in Pigeons.

(Bilates XXII XC NVI ie ete nega Ne er hese ee ee 601

BouLeNnGer, CHARLES L., M.A., F.Z.S., Lecturer on Zoology

in the University of Birmingham.

On Variation in the Medusa of Merisia lyonsi.

(elenng JOD ane INex giles, WY2= 2) sn acadonoascconnancoce 50 1045

BouLencer, Epwarp G., F.Z.S., Curator of Reptiles.

Exhibition of living male specimens of the Midwife

Toad (Alytes obstetricans) carrying the [email protected] 696

On a New Tree-Frog from Trinidad, living in the

SoCs Ceiling, . Celis GIDC) sdohannonsscsocncnoaoscnaes 1082

Bourne, GinBert C., M.A., D.Sc., F.R.S., F.Z.8.

Contributions to the Morphology of the Group Neritacea

of the Aspidobranch Gastropods.—Part II. The Huts-

cinipm. (Plates XXX—XUIL.) w..n.0.. Bt ogas AOR 759

Page

Brapy,- G. Strewarpson, M.D., LL.D., D.Se., F.BS.,

C.M.Z.S.

Notes on Marine Ostracoda from Madeira. (Plates

DOGO). Seladiged ARORA? BERN aH See asa RMA ae ayaa Rin 595

Brockman, Raupy E. Drake-. See Draky-Brockman, R. E.

Broom, Rosert, M.D., D.Sc., C.M.Z.8.

On the Structure of the Skull in Cynodont Reptiles.

(late Xavi and Wext- figs hGS G05 ioe rcueeeaensee ans. 893

On some New South African Permian Reptiles.

(Elenite Bp Ue ca bp. WEE) tence bie: Renate bear cbadtanadnceosnac 10735

Caprera, Prof. ANcrn, C.M.Z.S.

The Subspecies of the Spanish Ibex. (Plates LII.—

JULI, uae ARexeiaitresi Ie ES Ie) eecna ge aeetiadade ssoubeboseokace: 963

CALMAN, Wii ho [email protected] 1GZelSy

Exhibition of living specimens of the Brine Shrimp

(Artemia salina), bred from 'Tidman’s Sea Salt ............ 986

Curisty, Curapert, M.B., F.Z.8.

Exhibition of a collection of skins from Uganda and

of a loin-cloth taken from a native in Northern Nigeria. 672

CRossLAND, Cyrit, F.Z.S.

The Marginal Processes of Lamellibranch Shells.

(Gelenie) IDG, eumel WMasxncsieys, P79) 5 A310), \eacemcandacodandde sobeae 1057

Warning Coloration in a Nudibranch Mollusc and in

a Chameleoignemey tytn cue tac n Ment Atk oot ecane 1062

Draxe-Brockman, Raupu E., M.R.C.S., L.R.C.P., F.Z.8,

On Antelopes of the Genera Madoqua and Rhyncho-

tragus found in Somaliland. (Plates LV. & LVI.) ......

Exsot, Sir Cuartes N. E., K.C.M.G., C.B., F.Z.S.

Chromodorids from the Red Sea, collected and figured

loyy Mitre, Cyysenll Oreogsllenaelee (Vee weey ILI) ccoasgnenonconaeocone

FantuamM, Haroxtp B., D.Sc., B.A., F.Z.8., and Porrsr,

Miss Annie, D.Sc.

On a Bee-disease due to a Protozoal Parasite (Vosema

Harrison, Ruru, Oxford.

Some Madreporaria from the Persian Gulf. With

a Note on the Memoir and some Further Notes

on Pyrophyllia inflata by Sypnny J. Hickson, M.A.,

D.Se., F.R.S., F.Z.S8. (Plates LVII. & LVIII. and Text-

figs. 216 -221.)....:

Hetimayr, C. E., Division of Birds, Zoological Museum,

Munich.

A Contribution to the Ornithology of Western

Colombia

Sihohoneseueless}ens}aieei sie) elese/eleleieielolsieleleisidie tiles) s/s late elelelnlisielsfolulateletalsichalalefersle

Hickson, Sypvey J., M.A., D.Sc, F.R.S., FZS. See

Harrison, Rutu.

Some Madreporaria from the Persian Gulf,

Page

S07

1068

625

1018

1084

xili

Ho.pine, R. E.

Exhibition of the Horns of a Highland Ram, a Fallow

Deer, and a Roebuck, which were fused at the base,

and the skull of a coursing Greyhound with abnormal

COTA GEG T OTIS ee ee ec ee ae a eR Aer. Sy ROLE

HopeEweE.u-SmirH, A., L.R.C.P., M.R.C.S.,and Tims, H.. W.

Marett, M.A., M.D., F.L.S., F.Z.8.

Tooth-germs in the Wallaby (Macropus billardier?).

(Bla tee xeliValiian diiexct “mies SIE G99) ls eeeeee sees eon

LANKESTER, Sir E. Ray, K.C.B., F.R.S., F.Z.S.

Exhibition of a special supplement to the ‘ Field’ news-

PBAPOV ocr e reece cece cece eee eee ee eee eee t eee c ence entre eee seeetees

Lerrrer, Rosert T., D.Sc., M.B., F.Z.S.

A Demonstration of Nematode parasites obtained from

animals in the Zoological Gardems .....:.......-.0c0cc.ee-0

ListEer, JoserH J., M.A., F.R.S., F.L.S., F.Z.S.

The Distribution of the Avian Genus Megapodius in the

Jee rouire, Ikslenaglsyy f(UMeraratves N50.) Gs sccncacndandocaosecedeqoesse

LYDEKKER, RicHARD.

An African Rhinoceros, Klipspringer, and Gazelle.

(Mextatigss VOLSIOS jak. esac. doussais Maar ohana nt

Manpers, Lieut -Colonel Nevitie, R.A.M.C., E.Z.8., F.E.S.

An Investigation into the Validity of Miillerian and

other Forms of Mimicry, with Special Reference to the

Islands of Bourbon, Mauritius, and Ceylon. ...............

Page

946

926

620

749

958

X1V

Page

Meruvuen, The Hon. Paut A., F.Z.S.

On an Amphipod from the Transvaal. (Plates XLIX.—

Jill) ae at Aas A AAR 1A AM RE AE ESE SNR RB cs za 948

Mircuett, P. Cuatmers, M.A., D.Sc., Hon. LL.D., F.B.S.,

F.Z.5S., Secretary to the Society.

Report on the Additions to the Society’s Menagerie

ching auinemmontinvot sue bruce lO) Ibe ae ernest ene nee nene 557

A letter from the Governor of the Seychelles on Land-

Hortorsesminkthosewlislancds 1c eer ee eee eee et Renee 622

Report on the Additions to the Society’s Menagerie

diuiemsuthemmonthweteMarchenl 9/0 ii asses eeeet eee nein anne 671

Additional reference to the memoir “On Longevity

and Relative Viability in Mammals and Birds” ......... 868

Report on the Additions to the Society’s Menagerie

ChB HAS WS TOD OH Wyaall WOM jo yseassacacso9osa9cugcosoce 869

Report on the Additions to the Society’s Menagerie

GlpuetNS tae rmo ain Wey WAI 5.5, co2ccons00ascn000q0osoa000 985

Remarks upon the eggs of Struthio massaicus, S. aus-

tralis, and 8. molybdophanes, seen at Mr. Carl Hagenbeck’s

OStrichyHarmatastellancenyeeeeeek: cote ree eens ee enn eeeeener 987

Remarks upon a pair of young African Rhinoceros seen

at Mr. Carl Hagenbeck’s Zoological Park at Stellingen... 987

Morcnu, J. A., Christiania.

On the Natural History of Whalebone Whales. (Text-

Fes OOS O32) a) peissinddwacieg « semsiss cd sruscacte ce ache eet eee eee 661

Moss, The Rev. A. Miuus, M.A., F.Z.S., F.E.S.

Notice of a memoir on the “Sphingidee of Peru” ...... 869

Niconi, WinuiaM, M.A., D.Sec., M.B., F.Z.S8.

On a unique Pathological Condition in a Hare. (Text-

HO LOD. Ay etme te Nees tds ce eorrk esta ce wed aw ig tale octane

On Three New Trematodes from Reptiles. (Plates

XXVIII: & XXVIII.)

Primmer, Henry G., F.R.S., F.LS., F.Z.8., Pathologist to

the Society.

Report on the Pathological Examination of Rats (J/us

decumanus) caught in the Regent’s Park and in the

Society’s Gardens

CEDOOIOIOIOIO POI IOC On ICICI ICO CICICICICECICEC IAI ICICNCECECECICICECHOIC

Pocock, Reernaup I., F.RS., F.LS., F.Z.8., Curator of

Mammals and Resident Superintendent of the

Gardens.

Exhibition of the skin of a new Chacma Baboon (Papio

porcarius, Subsp. griseipes), a specimen of the North-

American Black-footed Polecat (Putorius nigripes), ete.

eT MA Pi oiec aie oes occa intone sc on em nate seemnbenanos tthe

Exhibition of a newly born Masked Palm-Civet

(Paradoxurus larvatus), with an abnormal leg. (Text-

fig. 147.)

i i i ii i iii i i ii i ki i i i iii ii i i i ii i rice i i er ire aay

Exhibition of some hair from the “ puppy-coat” of a

Grey Seal (Halicherus grypus)

On the Palatability of some British Insects, with Notes

on the Significance of Mimetic Resemblances. With

Notes upon the Experiments by Prof. E. B. Potxron,

TS es Re ASie ea th oS ne aics dioltaiate ok EMU AN ar ci MR NE TaN

Exhibition of, and remarks upon, the skin and skull of

a specimen of the Crested Rat (Lophiomys ibeanus Thos.).

(Text-fig. 190.)

Oe ee ee ee i i i ee a

Page

674

677

558

621

696

809

XV1

Pocock, Reema I., F.RS. &e. (Continued.)

Exhibition of photographs of, and remarks upon, two

hybrid foals, Hquus asinus somaliensis x Equus zebra

and Hguwus asinus somaliensis x Equus quagga chap-

manm. (Text-figs. 201-203.)

Porter, Miss Anniz, D.Sc., and Fanrnam, Haron B.,

D.Sc., B.A., F.Z.S.

‘On a Bee-disease due to a Protozoal Parasite (Nosema

Poutton, Epwarp B., M.A., F.B.S., F.Z.8. See Pocock,

R. B.

On the Palatability of some British Insects, &e.

Reoay, C. Tare, M.A., F.Z.8.

Exhibition of a series of lantern-slides of scales of the

SHUMMTOM ((SOUMMO STGP) ococenrncesoepakocouacoocacvs.5hove0s06000 000

Seru-Smitn, Davin, F.Z.8., Curator of Birds.

Exhibition of a living Hybrid between the White-

eyed Pochard (Aythya nyroca) and the Marbled Duck

(Marmaronetia angustirostris) ............00cceceeseesseess eee

Exhibition of a nest of the Grey Struthidea or Apostle-

Bird (Struthidea cinerea), and lantern-slides of Penguins

in moult and of wild Swainson’s Lorikeets (7richoglossus

novee-hollandic). (Text-fig. 164.) ............cccceceseeeenes

Exhibition of, and remarks upon, two immature

Black-backed Porphyrios (Porphyrio melanonotus) with

wine=claws: (ext tis. 2005) vs. cceanss sis easeeen ce doemnCn eer

Page

988

625

671

5d8

671

Sexron, Mrs. KE. W., Marine Biological Laboratory, Ply-

mouth.

On the Amphipod Genus Leptocheirus. (Plates XVII.—

SXGIDXE: erm cleaner ule Oa) yee time ae geen oh aaron Maries

SHUFELDT, Ropert W., M.D., C.M.Z.S.

Exhibition of a photograph of a male albino Woodchuck

(CAP GHONDUIS TOOINRD) “pa oencobe neater MneuGsoncdoe echoes eh coHCBr enn

SMALLEY, FrepERIc W., F.Z.S., and Bonuorr, J. Lewis,

Wi oBloe TIGIUES 5 1WZAdse

On Colour and Colour-pattern Inheritance in Pigeons.

(Tete n gets 6) GD EPONA Ei os Ahen Nn ORE coc nce ese meee

Suirn, A. Hoprweni-. See Hopmweiu-Surru, A.

Smit, D. Seru-. See Sera-Surru, D.

THomas, OLDFIELD, F.R.S., F.Z.8.

The Duke of Bedford’s Zoological Exploration of

Eastern <Asia.—XIV. On Mammals from Southern

Shen-si, Central ‘China. ‘(Plate X XTX.) 1....0.0.0.....4..

Tomson, J. Sruart, Ph.D., F.R.S.E., F.L.S., Lecturer

and Senior Demonstrator in Zoology, University of

Manchester.

The Alcyonaria of the Cape of Good Hope and Natal.—

GorconacEa, (Plates XLITI.—XLV. and Text-fig. 167.)

Tims, H. W. Marerr, M.A., M.D., F.L.S., F.Z.S:, and

Horrwett-Suiru, A., L.R.C.P., M.R.C.S.

Tooth-germs in the Wallaby (Macropus billardieri).

(Plens CIEL. eiavel AMexceileiss JUS IS) s) gcse onceckoqubapacod

Proc. Zoou. Soc.—1911., 5

946

60]

XVI

NEW GENERIC TERMS

PROPOSED IN THE PRESENT VOLUME (pp. 557-1218).

Page Page

Anoplotenia (Vermidea) ...... 1003 Eriphostoma (Reptilia)......... 1078

Arctognathns (Reptilia) ...... 1079 Ictidognathus (Reptilia) ...... 1078

Arctosuchus (Reptilia) ......... 1079 Moschops (Reptilia) ............ 1073

Dasymetra (Vermidea) ......... 683 Taognathus (Reptilia) ......... 1076

Dizelurodon (Reptilia) ......... 1075 Thysanoteenia (Vermidea)...... 1002

SCLENTIFIC

Abraxas, 867.

grossulariata, 833.

Acanthogorgia

armata, 870, 880.

Acanthopneuste

viridanus, 718.

Accipiter

superciliosus, 1203.

tinus, 1203.

Aceratherium, 945.

Acontias, 705.

burtoni, 705.

Acrea, 714, 742,

Acridotheres

cristatellus, 860.

tristis, 698, 701, 720,

740.

Acrocephalus

dumetorum, 718.

stentoreus, 718.

Acrorchilus

erythrops griseigularis,

1149.

Acryllium

vulturinwn, 862.

Actinote, 705.

Adelomyia

cervina, 1211.

melanogenys maculata,

1211.

AXgithina

tiphia, 716.

A®lurosaurus, 1079.

fetinus, 1077, 1081.

tenuirostris, 1077,

1081, 1082.

whaitst, 1077, 1078,

1081, 1082.

fflurosuchus, 923,

924.

/#lurus

fulgens, 557.

Aapyprymnus, 940,

INDEX

Acelecyathus

persicus, 1019, 1034,

1035, 1039.

Aglaia

Fanny, 1107.

gyroloides, 1104.

peruviana, 1104.

Agyrtria

francie, 1182.

Alcadia, 761, 763, 776,

777, 780, 782, 783,

785, 794, 796, 797,

799, 800.

hollandi, 762, 766, 770,

TOE, TUB, HE WAY

781, 785, 787, 788,

793, 798, 807, 808,

809.

palliata, 762, 766, 768,

769, 770, 772, 773,

778, 787, 788, 789,

793, 798, 807.

Alcedo

amazona, 1192.

americana, 1192.

cabanisti, 1198.

inda. 1192.

torquata, 1192.

Algiroides

nigro-punctatus, 864.

Allantus

arenatus, 828, 850.

Aloposaurus, 919,

1078.

gracilis, 1081.

Alseonax

latirostris, 720.

muttut, 720.

Alytes

obstetricans, 696.

Amazilia

Juscicaudata, 1183.

riefferi, 1188.

NAMES.

Amazilia

tzacatl fuscicaudata,

1183.

— yucunda,

1183

Amblycercus

holosericeus, 1122.

Ampelis

natterertt, 1146.

Anabates

subalaris, 1151.

Anabazenops

lineatus, 1151.

mentalis, 1151.

subularis, 1151.

Anas

cyanoptera, 1209.

superciliosa, 755.

Androdon

equatorialis, 1176,

1177.

Anonchoteenia, 637.

Anoplocephala, 651,

653, 660.

Anoplops

bicolor, 1171.

— equatorialis, 1171,

1172, 1211.

— bicolor, 1171, 1173,

1212.

— dague, 1170, 1171,

1173, W211, 1212.

1182,

— olivascens, 1171,

1173.

— ruficeps, 1171,

1173.

leucaspis, 1171,

1172.

Anoplotznia, gen. nov.,

1003, 1018.

dasyuri, 1003-1007,

1009, 1010, 1012,

1013, 1OL5.

82 *

Anthrocera,

(Zygena) filipendule,

832.

Anthus

rufulus, 722.

Antilope

corrina, 962.

Aphanoconia, 761, 780,

785, 796, 797, 798.

andamnanica, 762, 773,

787, 801, 809.

gouldiana, 762, .7685,

778, 779, 785, 793,

808, 809.

merguiensis, 762, 779,

501, 808, 809.

pachystoma ponsonbyi,

763

rogersii, 762, 793, 801,

809.

(Helicina) theobaldiana,

60.

Aphantopus

hyperanthus, 8217,

828.

Aphelocoma

woodhousei, 869.

Apis

mellifica, 812, 850.

Aploparaksis, 652.

Apocrangonyx, 949,

Apodemus

aqrarius pallidior, 690.

speciosus peninsula,

690.

Appias, 730, 734, 787,

742.

galene, 709, 735.

paulina, 730.

Aptenodytes

pennant?, 558, 671.

Aramides

mangle, 1209.

wolfi, 1208, 1209.

Araschnia

levana, 823, 865.

prorsa, 823, 865.

Arctognathus, gen. nov.,

1079.

curvimola, 1079.

Arctomys

monax, 946.

Arctonyx

leucolemus, 688, 689.

— orestes, 688.

Arctophila, 854.

mussitans, 848, 851,

853.

Arctosuchus, gen. nov.,

1079.

tigrinus, 1079.

Argiades

sylvanus, 831.

Argilleecia

affinis, 595, 596,

601.

Argynnis

aglaia, 824.

castetst, 725.

euphrosyne, 820, 826.

hyperbius, 702, 703,

704, 706, 709, 735,

736, 737.

(Brenthis) ewphrosyne,

824,

(—) selene, 825.

(Dryas) paphia, $24,

827, 828.

Arion, 804.

ater, 813

hortensis, 813.

Arremon

aurantiirostris, 1118.

— erythrorhynchus,

1119.

— occidentalis, 1118,

— spectahiiis,

1119.

erythrorhynchus, 1118,

1119.

1118,

occidentalis, 1119.

spectabilis, 1118, 1119.

Arrenga

blight, 716.

Artamus

Fuscus, 720, 729.

personatus, 859.

superciliosus, 859.

Artemia

salina, 986.

Ascaris

canis, 620.

Aspidosiphon, 1023,

1025, 1027.

Aspidura sp., 716.

Asturina

schistacea, 1204.

Aulacorhamphus

albivitta, 1213.

— pheolemus, 1218.

ceruleiqularis,

1218.

petax, 1213.

pheolemus, 1213.

sulcatus, 862.

Aurelia

aurita,

1056.

Automolus

assimilis, 1150.

holostictus, 1152.

1047, 1053,

INDEX OF SCIENTIFIC NAMES.

Automolus

nigricauda, 1150, 1210.

(Rhopoctites) 7fo-

brunneus, 1149, 1150.

Avicula

zebra, 1058, 1059, 1060.

Aythya

nyroca, 5d8.

Bacillus

pestiformis, 626.

Bairdia

acanthigera, 595.

amygdaloides, 595.

dubia, 595, 596, 601.

mediterranea, 595.

obtusata, 595.

Balena

australis, 663.

Baleenoptera

borealis, 662, 668.

musculus, 662,

sihbaldii, 662.

Balanus

improvisus, 591.

Balistes

flavimarginatus, 1058,

1065.

viridescens, 1058, 1065

Basileuterus

auricularis, 1092.

Julvicauda semicer-

vinus, 1092.

melanotis, 1092.

— dedalus, 1092.

meridanus, 1092.

semicervinus, 1092.

tristriatus, 1092.

— melanotis, 1092,

1212.

— meridanus, 1092.

— tristriatus, 1092,

1212.

Bauria, 898, 899, 900,

901, 902, 905, 909,

913, 914, 915, 916,

917, 920, 923.

cynops, 895, 896, 897,

924, 925.

Belone sp., 1064.

Bertia

satyri, 649.

studeri, 649.

Bertiella, 994, 999, 1004,

1017.

cercopithect, 638, 640,

641, 643, 644, 647,

649, 650, 651.

conferta, 645, 646, 650,

edulis, 1017.

Bertiella

melanocephala, 639,

640.

mucronata, 639, 645,

646, 650. -

obesa, 1017.

plastica, 650.

polyorchis, 646, 651.

sarasinorum, LOL.

semonti, 1017.

Betta

splendens, 989.

Bison

bonasus, 557.

Bistylia, 1080.

Boeckia

typica, 563.

Boisscunneaua

flavescens flavescens,

1185.

— tinochlora, 1185.

Bombus, 841, 853.

agrorum, 846, 850,

803

hortorum, 812, 852,

854, 855.

joncellus, 891.

lapidarius, 845, 880,

852.

® terrestris, 851, 854.

Boreophausia

ineriis, 666, 667.

Bothriocephalus, 639.

Brenthis, 865.

Buarremon

brunneinucha, 1117.

Bubo

poénsis, 869.

Bucco

macrodactylus, 1196.

noaname, 1195, 1196,

1211.

pectoralis, 1194, 1195,

Uzi

subtectus, 1195.

tectus, 1195.

— subtectus, 1195,

We,

— tectus, 1195.

Buchanga

atra, 7217.

Bucorax

abyssinicus, 862.

caffer, 862.

Budorcas

bedfordi, 693, 695.

mitchelli, 695,

INDEX OF SCIENTIFIC NAMES.

Buthraupis

arcet, 1110, 1212.

aureocincta, 1110,

T1111, 1211.

edwardsi, 1111, 1211.

melanochlamys, 110,

1110, 1211, 1212.

rothschildi, 1110, 1211.

Bythocypris

reniformis, 595.

Cabera

exanthemaria, 832.

pusaria, 833.

Cabreta

leschenaultii, 706.

Cacicus

microrhynchus, 1212.

uropygtatis, 1121, 1212.

Cacomantis

passerinus, 723.

Calanus, 668.

Calliphora

vomitoria, 850.

Calliste

aurulenta, 1102.

cayanda, 860. .

emilie, 1103.

Jfrancesce, 1107.

Julvicervia, 1109.

gyroloides, 1104, 1107.

icterocephala, 1198.

Johanne, 1101.

labradorides, 1109.

larvata, 1108.

lavinia, 1103.

leucotis, 1109.

ruficervix, 1109.

rufigularis, 1102.

taylori, 1109.

Callithrix

Jacchus, 856.

Calophasis

ellioti, 862.

Calospiza

aurulenta aurulenta,

1102.

— sclateri, 11C3.

florida arcet, 1212.

— florida, 1212.

gyroloides, 1105.

— bangsi, 1105, 1106,

1107, 1212.

— catharine, 1106.

— deleticia, 1104.

— gyroloides, 1104,

1105, 1106, 1212.

XX1

Calospiza

larvata fanny, 1107,

1108, 1212.

— francesce, 1108.

— larvata, 1108.

lavinia, 1103, 1104.

—dalmasi, 1104,

1212.

— lavinia. 1103, 1105,

1212.

palmeri, 1107, 1211,

phenicotis, 1211.,

ruficervix, 1108.

— fulvicervix, 1109.

— ruficerviz, 1109.

— taylort, 1109.

rufigula, 1102.

Calotes, 727.

nigrilabris, 708.

ophiomachus, 707, 708,

739

polytes, 707.

versicolor, TOT, 708,

Ws 7

Calyptrophora

josephine, 886.

Canielus

bactrianus, 869.

Campophaga

sykesi, 719.

Capito

equatorialis, 1200.

bourcierti, 1199.

— eguatorialis, 1200.

— bourcierti, 1199,

1200.

— salvini, 1199, 1200.

maculicoronatus, 1198,

1199, 1211.

quinticolor, 1198, 1211.

shelleyi, 1200.

sguamatus, 1199, 1211,

Capra

cylindricornis, 973.

hispanica, 963, 970,

ibex, 963.

pyrenaica, 963, 964,

967, 969, 970, 972,

974, 975, 976.

— hispanica, 963, 966,

967, 969, 970, 972,

975, 976, 977.

— pyrenaica, 969, 970,

975.

— victorie, 975, 977

stbirica, 969.

sinensis, 695.

taxicolor, 694.

tibetanus, 687, 693,

694, 699. .

icterocephala, 1103.

johanne, 1101, 1102,

-1210, 1212.

labradorides, 1109.

Capreolus

bedtordi, 695.

Caprimulgus

rosenberg?, 1210,

Xx11

Carabus, 842.

violaceus, 816, 836,

841, 845.

Caranx sp., 1069.

Carbo

brasilianus, 1210.

vigua, 1209.

Cariama

cristata, 864.

Carpodectes

hopkei, 1147, 1210,

1212, 1218.

nitidus, 1212.

Caryophyllia

communis, 1088.

Casella

atromarginata, 1062.

Cassiculus

microrhynchus, 1121,

22.

Cassicus

persicus, 860.

pyrohypogaster, 4123.

uropygialis, 1121,1122. |

Cassidix

oryzivora, 1122.

—- oryzivora, 1122.

— violea, 1122.

Catenotenia, 637, 1018.

Catheturus

lathami, 863.

Catopsilia, 730, 734, 742.

crocale, 725.

florella, 727.

pomona, 709.

pyranthe, 727, 738,

739, 741.

Cebus sp., 815, 825, 829,

831, 832, 851, 856.

capucinus, 651.

loricatus, 1188, 1189.

- mentalis, 1189.

squamatus, 1189.

Ceophlcus

lineatus lineatus, 1189.

Cephalophus

ignifer, 673.

johnstont, 672.

weynst, 672, 673.

Ceratodus, 988.

forsteri, 942.

Ceratogymna

atrata, 861.

elata, 861.

Ceratoisis

ramosa, 870, 877, 892.

Ceratophora

myslaceus, 706.

negrilabris, 706, 708.

ophiomachus, 706.

steddarti, 708, 710.

Ceratophora

versicolor, 706.

Ceratopoma, 761.

Ceratosoma

cornigerum, 1068, 1065.

Cerchneis

punctata, 700.

Cercomacra

berlepschi, 1167,

1210.

carbonaria, 1166.

crepera, 1165.

maculicaudis, 1166.

nigricans, 1166.

tyrannina, 1165, 1167.

— rifiventris, 1165,

1166.

— tyrannina, 1169.

Cercopithecus

callitrichus, 688, 649.

centralis, 671.

denti, 856.

mona, 856.

pygerythrus, 856.

roloway, 856.

Certhia

cerulea, 1097.

Certhiola

chloropygia, 1098.

mexicana, 1098.

Cervus

bedfordi, 560.

hortulorum, 558.

canthopygus, 560.

Ceryle

amazona, 1192.

americana americana,

1192, 1198.

— cabanisii, 1193.

cabanistii, 1192.

inda, 1192.

torguata, 1192.

— torquata, 1192.

Cethosia

nietnert, 703.

Cheetura

indica, 7238.

Chaleidoseps, 705.

thwaitesir, 706.

Chalybura

buffont, 1184, 1185.

tsaure, 1212.

urochrysa, 1184, 1185.

1212.

Chama, 1059.

sp., 1057, 1061.

Joliata, 1057, 1061.

Chameleon, 922.

sp., LOG6.

Chamepelia

minuta, 869.

INDEX OF SCIENTIFIC NAMES.

Chameepetes

goudoti, 1207.

— goudotii, 1207.

— rufiventris. 1207,

Chapmannia, 652.

Charaxes

imna, 726.

Chelone, 921.

imbricata, 624.

mydas, 624.

Chilosia

illustrata, 848, 853,

854, 855.

Chiromacheris

aurantiaca,

1212.

vitellina, 1141, 1211,

PAN

Chlamydodera

naculata, 861.

1141.

| Chlorochrvsa

nitidissima, 1101,

1211.

phenicotis, 1211.

Chloronerpes

cecilie, 1188.

gularis, 1187.

rubiginosus, 1187.

— gularis, 1187, 1211,

1212.

— rubiginosus, 1187.

— uropygialis, 1212.

Chlorephanes

atricapilla, 1097.

cerulescens, 1097.

spiza . cerulescens,

1097.

— exsul, 1096, 1097.

| Chloropipo

holochlora, 1138.

— lite, 1138, 1210.

| Chloropsis

jerdoni, 716.

malabarica, 716.

Chlorostilbon

angustipennis, 1183.

pumilus, 1188.

Chlorothraupis

olivacea, 1114.

Chromodoris sp., 1070.

annulata, 1063.

diardui, 1062,

1064.

elizabcthina, 1062.

hilaris, 1063.

inopinata, 1070, 1072.

nigrostriata, 1062

quadricolor,1062.

reticulata, 1063, 1064,

1065, 1068, 1069,

1070, 1072.

1063,

Chromodoris

sykest, 1063.

tinctoria, 1069, 1072.

Chrysomela

polita, 841.

Chrysophanus

phleas, 830.

Chrysophrys

bifasciata, 1065.

Cicinnurus

regius, 861.

Cistecephalus, 1080,

1081.

microrhinus, 1081.

Cisticola

cursitans, 718.

Cistudo

carolina, 624.

Cittocincla

macrura, 721, 857.

Cladius

viminalis, 849.

Clavatella

(Eleutheria) prolifera,

1053.

Clematissa, 880.

Climacocercus

zonothorax, 1203.

Clisiocampa

neustria, &32.

Clytia, 1046.

folleata, 1048.

Cnipodectes

minor, 1128, 1129,

1130.

subbrunneus, 1128,

1129, 1180, 1213. »

— minor, 1129, 1130.

— subbrunneus, 1128,

1129.

Cobus

cob, 674.

Coccidium

cuniculi, 676.

Coccinella, 816, 845.

7-punctata, 812, 819,

841, 842, 845, 847.

Coccystes

coromandus, 724.

jacobinus, 724.

Coccyzus

melacoryphus, 1202.

Ceenonympha

pamphilus, 825, 829.

Ceereba

cerulea, 1097.

— luteola, 1098.

— microrhyncha, 1097,

1098.

chloropyga mexicana,

1098. ;

INDEX OF SCIENTIFIC NAMES.

Colias

nilgiriensis, 72d.

— Collocalia

francica, 698, 699.

' Colluricinela

heinet, 754.

Collyriocinela

harmonica, 859.

Columba

albilinea, 1205.

— albilinea, 1205.

— crissalis, 1205.

albitineata, 1205.

goodsom, 1205, 1206,

1210, 1218.

nigrirostris brunnet-

cauda, 1205, 1213.

plumbea bogotensis,

1206.

speciosa, 1205.

subvinacea berlepschi,

1206.

Compsocoma

notabilis, 1111.

Conchoderma

auritum, 667, 668.

Conocyathus, 1030.

zelandie, 1019, 1020,

1029, 1032, 1044.

Conopophaga

castaneiceps, 1176.

brunneinucha,

1176.

—- castaneiceps, 1176.

gutturalis, 1176.

nevutoides, 1167.

Conosmilia, 1088,

1041.

anomala, 1039.

elegans, 1039.

granulata, 1040,

lituolus, 1039.

striata, 1040.

stylifera, 1040.

Cophotis

zeytanica, 706, 708,

710.

Copsychus

saularis,

857.

Copurus

leuconotus, 112d.

Coracias

indica, 722.

Cordulegaster

annulatus, 835.

Coronula

diadema, 667, 668.

1039,

Ts TBR

| Corvus

cornia, 861.

scapulatus, 861.

XXU1

Coryphotriccus

albovittatus, 1134.

— distinctus, 1135.

parvus, 1139.

Cosmotricha

potatoria, 834.

Cossus

ligniperda, 831.

Cossypha

caffra, 857.

Cotinga,

cayana, 1147.

natterertt, 1146, 1147,

1211.

ridgway?, 1146, 1147.

simont, 1146.

Cotyle

uropygialis, 1093.

Cracticus

destructor, 861.

Crangonyx, 949, 951.

flageliula, 950.

subterraneus, 957.

Craspedoprion

equinoctialis, 1127.

brevirostris, 1127.

olivaceus, 1127.

Crateropus

canorus, 716.

cinereifrons, 715.

rufescens, 715.

striatus, 715.

Cratopus, 701.

Crax

alector, 863.

caruncuiata, 863.

globicera, 863.

Creciscus

alhigularis, 1208.

Cricetomys

gambianus, 651, 9995,

1003.

Cricetulus

andersont, 691.

triton, 691.

Crithidia, 626.

Crocidura

attenuata, 688.

coree, 688.

Crossarchus

fasciatus, 856.

Crosslandia

viridis, 1064.

Cuculus

canorus, 723.

micropterus, 723.

poliocephalus, 723.

Culcitella, 652.

Culicicapa

ceylonensis, 721.

| Cuscus, 921.

XX1V

Cyanerpes

cerulea microrhyncha,

1097.

Cyanocitta

pulchra, 1124.

Cyanocorax

affinis, 1128, 1124.

— affinis, 1123, 1124,

1212.

— zeledoni, 1124, 1212.

pileatus, 1124.

sclateri, 1124.

Cranolesbia

kingit, 1187, 1211.

— celestis, 1187, 1211.

— emme, 1187.

— kingit, 1187.

Cyanolyca

armillata, 1124.

pulchra, 1124.

Cyanomyia

francie, 1182.

Cyclorchida, 652.

Cyclorhynchus

e@quinoctialis, 1127.

cinereiceps, 1126.

subbrunneus, 1128,

1129.

Cycloseris

hexagonalis, 1036.

Cymbilanius

lineatus, 1157.

— fasciatus, 1157,

Cynanthus

celestis, 1187.

Cynochampsa

laniaria, 893.

Cynodraco

serridens, 893.

Cynognathus, 895, 899,

900, 904, 906, 907,

908, 910, 912, 923,

925, 1080.

berryi, 902, 905, 925.

erateronotus, 894, 900,

905.

leptorhinus, 901.

platuceps, 894, 901,

903.

seeley?, 893, 902.

Cyornis

rubeculoides, 720.

tickelli, 720,

Cyphorhinus

brunnescens, 1088

Cyphorinus

cantans, 1089.

lawrencii, 1039.

pheocephalus, 1088.

Cypselus

affinis, 723.

Cypselus

melba, 723.

Cythere

albo-maculata, 595.

cingulata, 595, 596,

601.

convexra, a95.

crispata, 595, 596, 601.

diffusa, 596.

elegans, 596.

emaciata, 595.

tuberculata, 595.

Cythereis

antiquata, 595.

deformis, 5965,

601.

Jonesit, 595.

runcinata, 595.

597,

Cytherella

ovalis, 595, 600, 601.

Cytheridea

elongata, 595.

Cytherideis

subulata — crenulata,

595, 600, 601.

— fasciata, 600.

Cytherura

cellulosa, 595.

eribriformis, 595.

cribrosa, 595, 599,

601.

fossulata, 595, 599,

Ol.

maculosa, 595, 599,

601.

striata, 595.

Dacelo

cervind, 861.

gegantea, 861.

leachii, 861.

| Daenis

cayana, 1095, 1096.

— callaina, 1096.

— cayana, 1096.

— cerebicolor,

1096, 1212.

— glaucogularis, 1096.

— napeéa, 1095, 1096.

— ultramarina, 1096,

1212,

cerebicolor, 1095, 1096.

napea, 1096.

salmoni, 1116, 1117.

venusta, 1094.

fiiginata,

1212.

Danais, 742.

aglea, 702, 725.

ceylonica, T7038,

1095,

1094,

INDEX OF SCIENTIFIC NAMES.

Danais

chrysippus, 702, 703,

704, 708, 725, 727,

731, 738, 739, 740,

744, 745.

Sumata, 709, 735, 736.

limniace, 704, 725, 737.

plexippus, 702, 703,

704,

septentrionis, 704, 725,

726, 727

( Amaur is) eee 702.

Dania

rero, 985.

Dasymetra,

683, 685.

conferta, 683, 686.

Dasy pus, 919.

villosus, 857.

Dasyurus

ursinus, 1003, 1017.

Davainea, 639, 652.

Dawsoniella, 762

gen. noyv.,

Delias 4

eucharis, 703, 708, 737,

740.

Delphinognathus, 919,

1074, 1075

conocephalus, 1073.

Dendrochelidon

coronata, 727.

Dendrocinela

atrirostris, 1156.

erythropygia, 1155.

Jumigata, 1156.

lafresnayet, 1155, 1156.

— pheéochroa, 1155.

meruloides, 1155.

— aphanta, 1156.

— lafresnayei, 1155,

1156.

— meruloides, 1155.

— pheochroa, 1155,

1156.

olivacea, 1155, 1156. .

— anguina, 1157.

— lafresnayei, 1155,

1156.

— pheochroa, 1156,

pheochroa, 1156.

Dendrocolaptes

atrirostris, 1156.

cuneatus, 1152.

triangularis, 1153.

Dendrocops

meruloides, 1155.

Dendrocopus

major, S61.

Dendrocygna

autumnalis, 1209.

discolor, 1209.

INDEX OF SCIENTIFIC NAMES.

Dendroica

estiva, 1091.

— estiva, 1091.

castanea, 1091.

Dendrohyrax, 673.

Dendrophyllia sp., 1019,

1036, 1039, 1044.

Dendropupa, 803.

Dendrornis sp., 1153.

equatorialis, 1153.

erythropygia @equa-

torialis, 1158.

lachrymosa, 1153.

— lachrymosa, 1212.

— rostrata, 1153, 1210,

1212.

punctiqula, 1153.

triangularis — e@qua-

torialis, 1153, 1154,

1212.

— punctigula, 1154,

— triangularis, 1154,

1155.

Dendrortyx

leucophrys, 863.

Dermochelys, 921.

coriacea, 9205.

Deuterosaurus, 1075.

Diadectes, 920.

Diademodon, 894, 923,

924, 920.

mastacus, 908.

Dizlurodon, gen.

1075.

whaitsi,

1082.

Diallactes

granadensis, 1158..

Dicheilonema

horrida, 620.

Dicrurus, 725.

ater, 717.

leucopygialis, 718, 728.

longicaudatus, 717.

Dictyocaulis

filaria, 620.

Dicynodon, 893, 1075,

1080, 1081.

sp., LOSI.

Didelphys, 921, 941,1017.

Diglossa

cerulescens, 1093.

Diglossopis

cerulescens, 1093.

— cerulescens,

_ 1094.

— pallida, 1094.

Dilepis, 652.

Dinotherium

curviceri, 944.

nov.,

1075, 1081,

1093,

Dinotherium

giganteum, 944.

hobleyi, 943, 944, 945.

Diorchis, 652.

Dissemuroides

lophorhinus, 718.

paradiseus, 718.

Disythamnus

rufiventris, 1165.

Dorcelaphus

savannarum, 869.

Doris

tinctoria, 1070.

Dromatherium, 924.

Dryocopus

lineatus, 1189.

Dryotriorchis

spectabilis, 869.

Dumetia

albigularis, 715.

Duneania, 1041.

Eccasaurus, 1075.

Echidna, 918, 919, 921.

aculeata typica, 924.

Echinomyia

ferox, 848. 855.

Elenia

cayenennsis, 1134.

cinerea, 1132, 11388.

— cinerea, 1132.

— parambe,

1183, 1210.

Elaphrornis

pallisert, 716.

Elymnias

fraterna, 702.

Embernagra

brunneinucha, 1117.

Empidonax

acadicus, 1186.

virescens, 1186.

Em pis

tesscllata, 848.

Enys

sculpta, 624.

Endothiodon, 1080.

bathystoma, 1081.

wniseries, 1075, 1081.

Engraulis

boelama, 1064.

Entameeba

apis, 626.

coli, 626.

Epenthis, 1048.

Hipimys

confiucianus, 689, 690.

— canorus, 690.

— luticolor, 689, 690.

— sacer, 690..

luticolor, 690. . »

1132,

XXV

Epimys

sacer, 690.

Epinephele

Jartina, 816, 820, 828.

Janira, 828.

tithonus, 829.

Equus

asinus, 993.

— somaliensis, 98,

993.

grevyt, 998.

hemionus hemionus,993.

kiang, 993.

quagga, 993.

= Sai 993.

— chapmanni, 992.

zebra, 989, 993.

Eranna

gucunda, 1182.

EKresia, 705.

Eriocnemis

aurelie, 1185, 1186. |

Eriphostoma, gen. nov.,

1078, 1079.

microdon, 1078, 1082.

Kristalis, $12.

Erythrura

trichroa, 985.

EHucephala

humboldti, 1210.

Euchelia, 867.

jacobee, 815, 820, 825,

8382, 845, 845.

Euchloé °

cardamines, 815, 820,

865.

Eucometis

cassini, 1116.

Eucope, 1046,

1055, 1056.

Kucrangonyx, 948.

robertsi, 950, 951, 957.

vejdovskyt, 949, 950,

951.

Eucythere

prava, 595.

Eulipoa, 751.

Kunicella

papillosa,

881, 892.

Euphonia

orevirostris, 1101.

fulvicrissa, 1100.

— fulvicrissa, 1211.

— purpurascens, 1100,

1048,

871, 880,

PAIN

purpurascens, 1100.

avanthogaster, 1100,

1101

— brevirostris, 1101.

— brunneifrons, 1101.

XXV1

Euphonia

xanthogaster chocoensis,

1100, 1211.

— wanthogaster, 1211.

Euplexaura

albida, 884. *

brauer?, 884.

media, 871, 883, 8938.

parciclados, 884.

Euplea, 713, 729, 734,

739, 742.

core, 702, 708, 704,

708, 709, 720, 725,

726, 730, 737, 738

740, 741, 743, 744.

coreta, 702, 704, 725,

726.

euphon, 702.

goudoti, 698.

klugit, 702.

Hupodotis

australis, 868.

Euripus

consimilis, 703.

Kurypyga

helias, 864.

Euscarthmus

cinereus, 1180.

Eutoxeres

aquila, 1180.

— aquila, 1180.

— heterura, 1180,1181.

baront, 1180, 1181.

heterura, 1180.

salvint, 1180, 1181.

Eutrochatella, 761, 777,

778, 779, 780, 782,

783, 785, 788, 796,

798, 799.

pulchella, 762, 763,

768, 772, 773, 786,

787, 788, 793, 797,

807, 808, 809.

Falco

cachinnans, 1204.

cayennensis, 1205,

palliatus, 1204.

superciliosus, 12038.

Felis

concolor, 671.

Sontanieri, 688.

lyne isabellinus, 557.

microtis, 688,

serval, 869.

Filaria

australis, 620.

Flabellum

laciniatum, 1088.

magnificum, 1019,

1020, 1039, 1044.

INDEX OF SCIENTIFIC NAMES.

Flabellum

rubrum, 1021.

Florisuga

mellivora, 1181.

— mellivora, 1181.

Formica

rufa, 812, 849, 868.

Formicarius

anatis, 1173.

— destructus, 1173,

1174, 1212.

— nigricapillus, 1174,

1212.

destructus, 1174.

migricapillus, 1174.

rufipectus, 11'74.,

— rufipectus, 1174,

1175.

— thoracicus, 1175.

saturatus, 1174.

thoracicus, 1175.

Formicivora

consobrina, 1168.

melena, 1162.

quadrivittata, 1160.

quixensis —_ houcardi,

1164, 1212.

— consobrina,

1164, 1212.

schisticolor, 1163.

Foudia

erythrocephala, 701.

Franklinia

gracilis, 718.

Fregilupus

capensis, 699.

varius, 699.

Frinyvilla

rubra, 1118,

Fulica

martinica, 1208.

Fundulus

gularis, 985.

Fungia, 1042.

patella, 1019, 10386,

1089.

1163,

Galbula

melanogenia, 1194.

ruficauda, 1194.

Galeoscoptes

carolinensis, 857.

Galepus, 919.

Galesaurus,

(00

894, 899,

planiceps, 893, 898,

928, 925.

Galictis

vittata, 857.

Gallinula

mangle, 1209,

Gammarus, 949.

Garrulax

leucolophus, 859.

picticollis, 859.

Gazella

cuvieri, 961.

dorcas, 961, 962.

hayi, 961, 962.

kavella, 962.

leptoceros, 961.

Genneus

nycthemerus, 862.

Geocichla

citrina, 857.

(Monticola) eyanus,

857.

( — ) sawatilis, 857.

Georissa, 760, 800.

Geotrupes

vernalis, 839,

eotrygon

bourcieri, 1206.

veraguensis, 1206.

— eachabiensis, 1206,

1210, 12138.

— cachaviensis, 1206.

— veraguensis, 1206,

Glaucidium

perlatum, 862.

Glaucis

e@neus, 1178.

columbiana, 1178.

Fraseri, 1177.

hirsuta, 1178.

— enea, 1178.

Glyphorhynchus

castelnaudit, 1152.

cuneatus, 1152.

— castelnaudit, 1152.

Gomphodontia, 894.

Gomphognathus, 894,

920, 923, 925.

rannemeyert, 908, 909,

910, 911, 912.

minor, 908, 909, 910,

911, 912.

polyphagus, 909.

Gonatodes

kandianus, 705.

Gonionemus, 1046, 1055.

Gorgonia

albicans, 881.

capensis, 871, 884,

887.

Hlammeoa, 871, 877,

888.

Gorgonocephalus, 874.

Gracula

intermedia, 860.

religiosa, 860.

Graculipica

melanoptera, 860.

Grallaricula

cucullata, 1176.

Grallina

australis, 858.

Graucalus

macit, 719.

Grison

Sura, 857.

Gryporhynchus, 692.

Guttera

pucherani, 863.

Guynia, 1038,

1040, 1041.

Gymnopithys

bicolor dague, 1170.

leucaspis, 1172.

ruficeps, 1172, 1173.

Gymnorhina

leuconota, 861.

1039,

Halcyon

sacra, T54.

smyrnensis, 723.

Halicherus

grypus, 696.

Hapalemur

griseus, 856.

Haplochilus

chapert, 985.

elegans, 985.

panchaz, 985,

Haplophyliia,

1041.

Harpactus

fasciatus, 723.

Harpalus, 836, 839.

ruficornis, 838.

Heleodytes

albobrunneus,

1211.

1038,

1088,

— albobrunneus, 1212. |

— harterti,

1212.

harterti, 1088.

Heliangelus

exortis, 1186.

Helicina, 772, 780, 783,

797, 798, 799.

brasiliensis, 760, 786,

787, 792.

japonica, 760, 784.

1038,

kubaryi, 59, 760,

Cae

sagraina, "760, 786,

787, 792.

substriata convexa, 761.

titanica. 7d9, 768,

786, 806.

INDEX OF SCIENTIFIC NAMES.

| Helicina

(Sturanya) beryllina,

760

| Heliconius, 742.

Heliothrix

auritus, 1186.

harrott, 1186.

Heliotrypha

parzudakii, 1186,

Helix, 762.

Helogale

varia, 557.

Hemipus

picatus, 719.

Hemithraupis

chrysomelas,

1212.

ruficapilla, 1117.

salmom, 1116, 1117,

1212.

IULN7/,

Henicorhina *

inornata, 1090, 1210,

1212;

leucosticta, 1090.

— eucharis, 1090.

— prostheleuca, 1090,

1212.

prostheleuca, 1090.

Hercynella, 803.

Herpetotheres

cachinnans, 1204.

Hesperia

antonia, 866.

carthami, 866.

side, 866.

Heterocyathus, 1021,

1031.

equicostatus, 1019,

1022, 1024, 1025,

1026, 1032, 1039,

1044.

alternatus, 1024, 1026,

1044,

cochlea, 1025.

heterocostatus, 1019,

1026, 1039, 1044.

lamellosus, 1025.

oblongatus, 1026.

parasiticus, 1025,

1026.

philippinensis, 1025,

1026.

pulchellus, 1026.

sulcatus, 1025.

wood-masom, 1026.

Heterodon

platyrhinus, 677.

Heteropsammia, 1031,

1039.

aphrodes, 1019, 1636.

michelini, 1032,

XXVli

Heterospingus

rubrifrons, 1115,

1212.

avanthopygius, 1114,

1115, 1212.

Hicksonella, 891.

spiralis, 871, 839.

Hirundo

chalybea, 1093.

ruficollis, 1093.

rustica, 722, 730.

viridis, 1099.

Hyalimax, 804.

Hyalonema, 877.

Hydrocena, 783,

800.

cattaroensis, 739, 760,

777, 806.

Hydrocorax

vigua, 1209.

Hyla

goughi, 1082, 1083.

misera, 1088.

rubra, 1032.

strigilata, 1083.

venulosa, 1082.

Hylocichla

mustelina, 857.

ustulata swainsonit,

1087.

Hyloctistes

subulatus — assimilis,

1150, 1151, 1212.

— virgatus, 1151,

1212.

virgatus, 1151.

— virgatus, 1151.

Hymenolepis, 652.

Hyperoodon

rostratus, 663.

Hypocnemis

neévioides, 1167.

— capnitis, 1168.

Hypolimnas

bolina, 702, '725, 735,

737, 738, 740.

diocippus, 727, 739.

misippus, 702, 703,

726, 727, 737, 738,

739, 740, 744.

Hypopyrrhus

pyrohypogaster, 1123,

Hypothymis

azurea, 721.

Hypsipetes

ganeesa, 717.

olivaceus, 700.

Hypuroptila

buffoni, 1185.

urochrysa, 1184.

784,

XXVill

Icterus

chrysocephalus, 860.

Jamaicai, 860.

mesomelas, 1123.

— taczanowskit,

1128.

salvinit, 1123,

vulgares, 860.

Ictidognathus, gen. nov.,

1078.

parvidens, 1078, 1081,

1082.

Ictodosaurus, 1079.

angusticeps, 1078.

Idiogenes, 652.

Tole

icterica, 717.

Irena

puella, 716.

Tthomia,

zelica, 705.

Jainides

bochus, 729.

Jonornis

martinica, 1208.

Juncella

elongata, 888.

flagellum, 838.

spiralis, 871, 889.

Junonia

lemonias, 727.

(Precis) almana, 738.

Kallima, 727.

philarchus, 728.

Kelaartia

penicillata, 717.

Lacerta

agilis, 864.

dugesit, 864.

muralis, 864.

— fjilfolensis, 864.

ocellata, 837.

viridis, 864.

vivipara, 837.

Lalage

maculosa, "Td4.

rufiventer, 699, TOL.

(Oxynotus) newton,

698, 699.

Lampides

batica, 701.

Lanius

collurio, 859.

cristatus, 719, 737,

739.

Lanius

erythronotus, 719.

lineatus, 1157.

nevius, 1158.

Laphyetes

satrapa, 1138.

Larvivora

brunnea, 716.

Lateriporus, 652.

Lathria

unirufa, 1144.

— castaneotincta, 1144

1145, 1212.

— clara, 1145, 1212.

Lechriorchis

elongatus, 681.

primus, 681.

validus, 677, 681, 682,

685, 685, 686,

(Renifer) — edongatus,

677.

Legatus

albicollis _albicollis,

1134.

Lemur

coronatus, 856.

Julvus albifrons, 856.

macaco, 856, 994,

1003.

mongoz, 850.

varius, 671.

Leontocebus

edipus, 856.

rosalia, 856.

rufimanus, 806.

Leptocheirus

aberrans, 561, 562,

563.

bispinosus, 562, 563,

572, 576, 585, 586,

587, 589, 594.

cornuauret, 561, 562,

563, 564, 569.

dellavallet, 561, 568,

576, 577, 582.

fasciatus, 576, 585.

guttatus, 561, 562,

563, 572, 576, 585,

587, 588, 589, 593.

hirsutimanus, 561,

563, 571, 588, 594.

massiliensis, 561,

589.

pectinatus, 561, 563,

572, 573, 574, 575,

576, 577, 582, 584,

585, 594.

ptiosus, 361, 562, 563,

564, 569, 57 0, ome

573, 576, 584, 585,

594.

INDEX OF SCIENTIFIC NAMES.

Leptocheirus

pm. ae 561, 562, Bee,

ee. 561, 562,

563, 564, 569. )

tricristatus, 561, ob?

572, 573.

Leptodira

annulata, 681.

Leptodon

cayennensis, 1205,

palliatus, 1204.

Leptopogon

poliocephalus, 1182.

superciliaris, 1132.

— poliocephalus, 1152.

— supercilaris, 1132.

— transandinus, 1182.

Lepus

jilchnert, 692.

swinhoei, 692.

— brevinasus, 692.

Lethe

daretis, 709, 710, 735,

737.

Leucolepis

pheocephalus lawrencii,

1089, 1213.

— pheocephalus, 1088,

1089, 1212.

Leucopternis

plumbea, 1204, 1210.

schistacea, 1204.

semiplumbea, 1204.

Linax, 804.

agrestis, 814.

arborium, 814.

maximus, 813.

Limenitis

sthylla, 865.

Limnoenida, 1046, 1047,

1049, 10538.

tanganice, 1056.

Linstowia, 636, 637.

Liothrix

luteus, 8d8.

Lipaugus

holerythrus, 1145,

1146.

— holerythrus, 1146.

1212.

— rosenbergi, 1145,

1146, 1212.

Lizzia, 1046.

Locusta

viridissima, 835.

Lonchodes sp., 839.

Lophiomys

ibeanus, 946, 947.

imhaust, O47,

948.

Loxia

grossa, 1121.

Loxoconcha

decipiens,

601.

impressa, 595.

obesa, 595, 597, 601.

‘subalata, 593, 598,

601

Lucanus

cervus, 839.

Lucidella, 761, 773, 776,

His, TD, TO, Wea,

729, 800.

aureola, 762, '763, 772,

780, 786, 793, 807,

808.

Lutreola

sibirica, 688.

Lycena

astrarche, 830.

icarus, 830.

595; (97,

(Zesius) chrysomellus, |

739, 740.

Lycosaurus, L078.

curvimola, 1079.

pardalis, 1079.

tigrinus, 893, 1079.

Lygosoma, 705, 706,

Lyriocephalus

seutatus, 700.

Lystrosaurus, 1080.

Mabuia, 705.

carinata, 706.

Macacus

cynomolgus, 651.

pileatus, 856.

radiatus, 650.

Macrocypris

decora, 595.

Macroglossa sp., 732.

Macropteryx

coronata, 723.

Macropus, 918, 938, 940.

bennetti, 927.

billardieri, 926.

brachyurus, 927.

eugentt, 927.

giganteus, 927, 939.

Macrorhinus

crozetensis, 671.

Madoqua

damarensis, 977.

hararensis, 979.

phillipsi, 978-984.

— gubanensis, 979,

983, 984.

— hararensis, 988,

984

INDEX OF SCIENTIFIC NAMES.

Madoqua

piacentinit, 981, 984,

swaynet, 980, 981, 982,

984.

Malacogorgia

capensis, 871, 885.

Malacoptila

mysticalis, 1197.

panamensis, 1196, 1197.

—- panamensis, 1197,

1212.

— poliopis, 1196, 1197,

IPARS

poliopis, 1196.

Malleus, 1060.

Mallotus

villosus, 666.

Mainestra

oleracea, 834.

persicarié, 832, 834,

835.

Mantis

religiosa, TOL.

Margaritifera

margaritifera, O57,

1058, 1059, 1060,

1061.

mauritit, 1052, 1060.

vulgaris, 1059, 1060,

1065.

Marmaronetta

angustirostris, 558.

Masius

chrysopterus _ bellus,

1138, 1139, 1211.

— coronulatus, 1139,

1211.

coronulatus, 1138.

Megacephalon

maleo, 757.

Megalama

virens, 862.

Megalobatrachus

maximus, 869.

Megapodius, 749, 750.

andersont, 758.

brazieri, 751.

brenchleyt, 751, 752,

cuming?, 756, 757.

duperreyt, 751, 753.

eremita, 752, Td8.

freycineti, 752, 758.

geelvinkianus, 757.

huttont, 756.

laperousit, 757.

layardi, 757.

macgillivrayt, 751.

pritchardi, "754, 755,

756, 757.

senex, 753, 757.

wallacei, 751.

XX1X

Megaptera

boops, 662.

Meiglyptes

loricatus, 1188.

- Melanargia

galathea, 816, 824, 827,

828, 865, 867.

Melanerpes

pucheramt perileucus,

1188.

— pucherani, 1188.

Melanitis

leda, 747.

tambra, TAT.

Meleagris

americana, 863.

ecellata, 863.

Melibe

jfinbriata, 1064, 1065.

Melierax

monogrammicus, 869.

Melinodon, 916, 923.

simus, 913, 923, 925.

Melita

artemis, 815, 820. 824,

825, 826, 829, 831,

866.

athalia, 866.

aurinia, 825, 866.

cinxia, 866.

Melitodes, 890.

dichotoma, 870, 877.

espert. 870, 874.

nodosa, 870, 876.

Melolontha

vulgaris, 839.

Meriones

crassus, 986.

Merops

ornatus, 869.

philippinus, 722.

swinhoei, 722, 733.

viridis, 722, %27,

732,

Merula

kinnist, 722.

tristis, LO86.

Mesopicos

eecilti, 1188.

Metroliasthes, 637.

Micrastur

guerilla, 1208.

— interstes, 1203.

— zonothorax, 1203.

interstes, 1203.

ruficollis, 1203.

Microconodon, 924.

Micromys

minutus, 690.

Micropogon

bourcierti, 1199.

XXX

Microtus

calamorum, 691.

— superus, 691.

johannes, 687, 691.

mandarinus, 691.

nux, 687.

(Caryomys)

692.

(—) nue, 691.

Milax

sowerby?, 814.

Millepora

alcicornis, 1058, 1060.

Mimus

orpheus, 858.

polyglottus, 858.

saturninus, 858.

Miniopterus, 910, 920.

Mionectes

oleagineus hederaceus,

1131.

olivaceus, 1131.

— galhinus, 1131.

eva,

— hederaceus, 1181,

1210, 1212.

— olivaceus, 1131,

1212.

— venezuelensis, 1131.

striaticollis, 1131.

— poliocephalus,

1131.

Mitrospingus

cassinit, 1116.

Mniotilta

varia, 1091.

Merrisia, 1048, 1050.

lyonst, 1045, 1046,

1047, 1049, 1050,

1058, 1054, 1055,

1056.

Molpastes

hemorrhous, 717.

Momotus

equatorialis, 1194.

— equatorialis, 1194.

— chlorolemus, 1194.

lessont, 1194.

martit, 1193.

semirufa, 1193,

subrifescens, 862.

Monasa

mystacalis, 1197.

pallescens, 1197, 1198,

1211.

Monopylidium, 652.

Mosvhops, gen. nov.,

1073

capensis, 1078, 1082.

Motacilla, 698.

estiva, 1091.

Motacilla

cayana, 1095.

noveboracensis, 1091.

ruticilla, 1098.

spiza, 1096.

varia, 1091.

Mungos

albicauda, 856.

Julvescens, 856.

galera, 857.

mungo, 856.

Murex, 1060, 1061.

ramosus, 1057, 1059.

Muriceides, 880.

Jusca, 870,

893.

Mus

decumanus, 945,

wagnert, 690.

Muscicapa

cayanensis, 1134.

Fulvicauda, 1092.

purpurata, 1148.

Muscipeta

borbonica, 701.

ralloides, 1087, 1088.

Myadestes

ralloides, 1087.

Mycalesis, 710.

ceyloniea, 728, 740,

TAL.

narcissus, 747.

Mycetes

878,

niger, 656.

Myiarchus

crinitus, 1137.

nigriceps, 1137.

tuberculifer, 1137.

tyrannulus, 1137.

Myiobius

barbatus, 1135.

— atricaudus,

1136.

— barbatus, 1136,

— mastacalis, 1135,

1136.

erythrurus, 1136.

— fulvigularis, 1136.

fulvigularis, 1136.

mastacalis, 1125.

sulphureipygius, 1135.

— aureatus, 1135,

1136, 1212.

— sulphureipygius,

1136.

1135,

— villosus, 1135, 1136,

1212.

villosus, 1185.

xvanthopygus, 1135.

— aureatus, 1135.

INDEX OF SCIENTIFIC NAMES.

Myiochanes

richardsonit

soni, 1136.

Myiodioctes

tristriatus, 1091.

Myiozetetes

eayanensis cayanensis,

1134.

— erythroptera, 1134.

rufipennis, 1134.

texensis, 1134.

richard-

| Myothera

analis, 1173.

Myotis

myosotis ancilla, 687,

688.

Myrmeciza

berlepschi, 1168.

exsul, 1169.

leucaspis, 1172.

zeledoni, 1169.

Myrmelastes

cassini, 1169, 1170.

exsul exsul, 1170,

1212.

— maculifer, 1169,

1170, 1212.

tmmaculatus, 1169.

— berlepschi, 1169.

— immaculatus, 1168.

— zeledoni, 1168.

Myrmothera

axillaris, 1162.

Myrmotherula

axillaris melena, 1162.

cineretventris, 1163.

fulviventris, 1162.

— fulviventris, 1162,

1212.

— viduata, 1162,

1212.

melena, 1162.

menetriestz, 1163.

— schisticolor, 1163.

ornata, 1162.

sancte-marte, 1163.

schisticolor, 1163.

— sancte-marte, 1163.

— schisticolor, 1163.

surinamensis, 1160,

1161.

— multostriata, 1159,

1159,

— surinamenis, 1159,

1160.

vidua, 1162.

Myzantha

garrula, 860.

INDEX OF SCIENTIFIC NAMES.

Navicella, 784.

Nematoteenia, 637.

Nemeobius

lucina, 831.

Nemorhzdus sp., 695.

Nemosia

rosenhergi,

1116,

. |

Neoniphargus, 949, 950, |

951.

Neptis

leucothoé, 735, 736.

lucilia, 865.

sapphe, 865.

varmona, 721.

Nerita, 764, 766.

Neritina, 762, 764.

Niphargus, 949, 950,

951.

Noctua, 818, 819,

835.

Nosema

apis, 625, 626.

bombyeis, 625.

Nothocercus

bonapartii, 1208.

Jrantzti, 1208, 1213.

intercedens, 1208,

1211, 1213.

Nyctipithecus

trivirgatus, 856.

Nythosaurus, 900, 901,

902, 905, 907, 908,

915, 919, 928.

larvatus, 894, 898,

899.

Obelia,

1355.

Oceania, 1049.

Ochetosoma

Jormosum,

686.

menstruosum, 681.

Ochotona

cansa, 692, 6935.

sorella, 692.

syrinx, 692.

Ocypode, 1066.

Ocypus

olens, 812, 816, 8836,

837, 838, 846, 855.

Odontophorus

cinctus, 1206, 1207.

parambe baliolus,

1211.

— parambe, 1211,

spodiostethus, 1206,

1207.

1046, 1048,

681, 683,

Ckdicnemus

bistriatus, 864.

Oligorchis, 652.

Oochoristica, 627, 630,

631, 632, 635, 6387,

654, 655, 658, 1000,

1005, 1017, 1018.

sp., 627, 628, 633.

rostellata, 1018.

tetragonocephala, 629,

638.

wageneri, 638.

Ophion

luteus, 848.

Ophisaurus

apus, 864.

Opisthoctenodon, 1075.

Orchistoma

pileus, 1051.

Oreocincla

imbricata, 722.

spiloptiera, 722.

| Oreotragus

saltator, 269.

— porteust, 960.

Oriolus

maculatus, 859.

oryzivorus, 1122.

Ornismya

kingtt, 1187.

Ornithoptera

darsius. 712.

Ornithorhynchus, 908,

910, 918, 919, 920,

921, 922, 942.

Orobophana, 761, 762,

773, 788, 726, 797,

798, 799, 800.

ponsonbyt, 772, 778,

780, 782, 783, 793,

807, 808, 809.

Ortalida

goudotii, 1207.

Ortalis

ruficauda, 863.

Orthogonys

olivuceus, 1114.

Orycteropus, 921.

Oryzoborus

ethiops, 1098.

funereus, 1098.

— ethiops, 1098.

— funereus, 1098.

Ostinops

_ atrocastaneus, 1211.

salmoni, 1211.

viridis, 860.

Ostracolethe, 804.

Ostrea sp., 1059,

1060.

XXX1

Otaria

californiana, 985.

Otis

ludwigi, 863.

vigorsti, 863.

Otocompsa

flaviventris, 858.

Otocryptis

bivitiata, 706.

Oudenodon, 1080.

bolorhinus, 1076, 1081,

1082.

strigiceps, 1076.

Ourebia

nuigricaudata, 869.

Oxynotus

JSerrugineus, 701.

Pachyrhamphus

atricapillus, 1144.

castaneus, 1144.

cinereiventris, 1143,

cinnamomenus, 1144.

dorsalis, 1143, 114,

1211.

marginatus, 1144.

niger cinereiventris,

1144.

ru,fescens, 1144.

rufus, 1144.

Paguma

larvata, 688.

Paleohelicina, 761, 762,

772, 113, 782, 787,

796, 797.

ide, 763, 770, 778,

780, 793, 807, 808,

809.

Panoplites

flavescens, 1185.

Pantodon

buchholzi, 985.

Papilio

agamemnon, 730.

aristolochie, 702, 709,

741, 748, 744.

clytia dissimilis, 702,

703, 705.

— lankeswara, 702,

704.

— panope, 705.

demodocus, 748.

demoleus, 08, 27,

748.

epiphorbas, 702.

hector, 702, 713, 714,

72%, Tdi, 40, 741,

744.

jason, 729.

exon

Papilio

lankeswara, 702, 704,

7A0.

limniace, 70).

machaon, 730.

manlius, 702.

phorbanta, 698, 700.

polytes, 702, 707, 708,

713, 714, 719, 729,

732, 743.

— romulus, 702.

surpedon, 727, 734.

(Menelaides) hector,

708, 709, 740, 743.

Papio

porcarius, 558, 559.

— griseipes, &58,

559.

sphinx, 842, 856.

Paracrangonyx, 949,

950.

compactus, 952.

Paracyamus

boopis, 668.

Paracyathus

cavatus, 1019, 1027,

1028, 1039, 1044.

erassus, 1039.

Paradoxostoma

arcuatum, 595.

cylindricum, 595.

flexuosum, 595.

_ gracile, 595, 600, 601.

- hibernicum, 595.

Paradoxurus

larvatus, 621.

Paranerita, 766,

779.

gagates, 764, 765.

Pareiasaurus, 919, 1079,

1080, 1081.

Pareronia

ceylonica, 702.

Parlotichus, 919, 920.

Parus

major, 858.

Paruterina, 652.

Passer

domesticus, 698.

Pellorneum

Suscicapillum, 715.

Pennella, 667, 668.

Penthoceryx

sonneratis, 724.

Perameles, 939, 942,

Perarge

megera, 815, 826,

Pericrocotus

flammeus, 719.

peregrinus, 719.

178,

' Phalanger, 1017.

Perisoreus

canadensis, 861,

Petanrista

alborufius, 689.

Petrogale, 940.

penicillata, 939.

Phethornis

berlepschi, 1179, ©

1180.

columbianus, 1179. °

syrmatophorus, 1179,

1180. i

— columbianus, 1180.

— syrmatophorus, °

1179, 1180,

yarugui, 1178.

— sancti-johanms,

1178, 1179, 1211.

— yaruqui, 1179,

PAD

Phalangista, 1017.

Phalangium sp., 814.

Phalera

bucephala, 832.

Phascolarctos, 1017.

Phasianus

principalis, 862.

reevesit, 862.

Pheasia

dicted, 833, 835.

tremule, 833, 834.

Phedina

borhonica, 698, 699.

Philydor

philydor

1151.

virgatus, 1151.

Pheenicothraupis

cristata, 1118, 1114.

Phrygilus

aldunatit, 558.

Phyllades

consolisma, 730.

Physeter

macrocephalus, 663.

Picumnus

canus, 1189, 1190.

dimotus, 1192.

granadensis,

1190, 1191.

olivaceus, 1189, 1190,

virgatus,

1189,

1192.

— dimotus, 1192.

— flavotinctus, 1190,

1191.

— granadensis, 1189,

1190, 1191, 1211,

1212.

— harterti, 1190, 1191,

211.

INDEX OF SCIENTIFIC NAMES.

Picumnus

' olivaceus _ olivaceus,

1190, 1191, 1192.

— panamensis, 1190,

Zale

Picus

lineatus, 1189.

rubiginosus, 1187.

(Chloropicus) iri,

1188.

_ Pieris

brassice, 816, 818, 823,

829, 834, 835, 846,

864.

napi, 815, 826, 827,

828, 829, 846, 865.

rapé, 815, 820, 826,

835, 865.

_ Pimelepturus

cinerescens, 1065.

Pionopsitta

hematotis, 1202, 1213.

pulchra, 1202, 1210,

1213.

Pionus

menstrius, 1102.

Pipra

chrysoptera, 1138:

coronata, 1140.

— velutina, 1140.

cyaneocapilla, 1140.

mentalis, 1139.

— ignifera, 1139, 1212.

— nmunor, 1139, 1140,

1212.

velutina, 1140.

vitellina, 1141.

Piranga

rubra rubra, 1113.

Pitangus

Ubovittatus, 1134,

parvus, 1135.

. sulphuratus, 861.

Pithys

bicolor, 1170, 1173.

— equatorialis, 1172.

— olivascens, 1173.

leucaspis, 1172, 1173.

Pittasoma

michleri, 1212.

rosenbergt, 1175, 1176,

1211, 1212.

rufopileatum, 1176,

1211.

Pitylus

grossus, 1121.

Planorbis, 762.

Platypsaris

homochrous, 11438.

Platyrhynchos

olivaceus, 1127.

INDEX OF SCIENTIFIC NAMES,

Platyrhynchos

virescens, 1136.

Platytriccus

albogularis, 1125,

1126.

cancroma, 1125.

insularts, 1126.

mystaceus, 1125.

— albogularis, 1125,

1126.

— insularis, 1126

— mystaceus, 1126.

Pleurotomaria, 803, 804,

805.

Plocamopherus

ocellatus, 1064.

Pneumatophilus, 683.

Podocoryne, 1046, 1047,

1049.

carnea, 1058.

Poéphagus

grunniens, 869.

Polydelphis, 620.

Polyerata

reint, 1181, 1182.

vosenbergi, 1181, 1182,

1210.

— reini, 1181.

Polyommatus

betica. 709, 740.

Polytes, 744.

Pomatorhinus

melanurus, 715.

Pontocypris

dispar, 595.

succinea, 595.

Porphyrio

melanonotus, 980,

martinicus, 1208.

Porzana

albigularis, 1208.

Pratincola

atrata, 721.

(Motacilla)

698.

Precis

almana, 740.

lemonias, 737.

rhadama, 747.

Prinia

Jerdoni, 719.

socialis, 718.

sylvatica, 718.

Prioneris

sita, 702,

704.

Prionites

martit, 1198.

Priotrochatella, 761.

risterodon, 1075.

sybilla,

703,

Procavia

dorsalis, 673.

emini, 673.

Procnias

cerulea

1099.

occidentalis, 1099.

tersa, 1099.

Eres opnen: 919, 920,

922, 1080.

Prodicynodon, 1076.

Progne

chalybea, 1093.

— chalybea, 1093.

Progynia, 652.

Proorchida, 652.

Propappus, 1080.

occidentalis,

Proserpina, 760, 761,

762, 799.

Protomedeia

fasciata, 568, 576,

585.

guttata, 561, 572, 573,

576.

hirsutimana, 561, 56

568, 571, 576, 58

589.

— massiliensis, 562.

pectinata, 576, 584.

pilosa, 563, 576, 577

585.

Psammogorgia, 883.

pulchra, 871, 881, 892,

893.

Psarocolius

mesomelas, 1122.

Pseudoclytia

pentata, 1048.

Pseudotrochatella

undulata, 761.

Psittacus

menstruus, 1202.

Psittospiza

riefferii, 1120.

—— elegans, 1120.

— riefferii, 1120.

Psophia

crepitans, 864.

Pteroglossus

erythropygius,

1202

ato

12018

— erythropygius, 1201,

1202, 1211.

— sanguineus, 1201,

1202, 1211.

sanguineus, 1201.

Pterostichus, 836,

855.

niger, 816, 837, 842,

846.

(Abax) striola, 837.

Proc, Zoou, Sov.—191L, No, LXX XIII,

XXX111

Pterostichus

(Steropus) madidus,

816, 838, 842.

Ptilocheirus

pectinatus, 576, 585.

tricristatus, 563,

576.

Ptilotis

carunculata, 754.

Putorius

nigripes, 097, 559.

Pyenonotus

hemorrhous, 858.

Jocosus, 701.

leucogenys, 808.

leuconotus, 858.

luteolus, (17.

melanicterus, 717.

rzanthopygus, 858.

Pyctorhis

nasalis, 715.

Pyrameis

atlanta, 823.

cardut, 709, 710,

738, 823.

Pyramidula, 803.

Pyranga

estiva, 1115.

Pyrgoma

stokesii, 1029.

Pyriglena

berlepschi, 1167.

maculicaudis, 1166.

tyrannina, 1165.

Pyrophyllia, 1088, 1041.

inflata, 1018, 1019,

1020, 1037, 1039,

1040, 1044.

Pyxis

arachnoides, 624.

O72,

736,

Querquedula

cyanoptera, 1209.

Querula

eruenta, 1148.

purpurata, 1148.

Quiscalus

gundlachi, 671.

Rallus

abbotti, 864.

Ramphocelus

icteronotus,

1113.

Rathkea, 1046, 1047.

Renifer, 677, 685.

ellipticus, 683.

sauronates, 681, 685.

Retorquata, 761.

1112,

XXXIV

Rhabdias

bufonis, 620.

Rhabdometra, 637.

RKhagonyche, 828.

Julva, 812, 840, 842,

850.

Rhamphastos

ambiguus, 1201.

swainsonit, 1200, 1201.

tocard, 1200.

tocardus, 1201.

Rhamphoeenus

cinereiventris, 1164.

— cinereiventris, 1164,

1212.

— semitorquatus, 1164.

semitorquatus, 1164,

1212.

Rhamphocelus

brasilius, 860.

icteronotus, 1112.

Rhinoceros, 945.

bicornis, 959, 985:

somaliensis, 9059,

960.

Rhinochetus

Jubatus, 864.

Rhbinolophus

Jerrum-equinum, 687.

Rbipidura

albifrontata, 721.

tricolor, 859.

Rhopocichla

nigrifrons, 715.

Rhopoctites

alogus, 1149.

Rhynchoeyclus

equinoctialis, 1 127,

cinereiceps, 1126.

— cinereiceps, 1212.

— flavotectus, 1126,

1127, 1210, 1212.

meg actphala, 1126.

— flavotectus, 1126.

sulphurescens, 1127.

— asemus, 1127.

Rhynchortyx

cinctus, 1206, 1207.

Rhynchotragus, 977,

978.

cordeauat, 983.

erlangeri, 983, 984.

guentheri, 980-984.

— wroughtont, 983,

984.

kirkii, 978, 982.

Rictularia

plagiostoma, 620.

Rivulus

flahellicauda, 985.

ocellatus, 985.

Rivulus

poeyi, 98d.

Salmo

salar, 671.

Saltator

atripennis, 1120.

elegans, 1120.

magnus, 1121.

maximus, 1121.

Sapayoa

enigma, 1141,

1213.

Saysia, 1046, 1056.

prolifera, 1066.

Sarsiella

capsula, 595.

Satyrus

semele, 830.

Saurosternon, 1080.

Sayornis

ardesiacus, 1125.

cineracea, 1125.

— cineracea, 1125.

Schcenicola

platyura, 718.

Scirpearella sp., 888.

aurantiaca, 888.

indica, 888.

Scirpearia sp., 888.

flagellum, 871, 888,

3g.

furcata, 871, 888.

ochracea, 839.

Sciurotamias

davidianus, 689.

Sclerochilus

contortus, 595.

levis, 695, 601.

Scops

leucotis, 862.

Scotothorus, 1141.

Scylacosaurus, 919.

constrictus, 1078.

Scymnosaurus sp., 1081.

Seiurus

noveboracensis, 1091.

— noveboracensis, 1091.

Semnornis

ramphastinus, 1200.

Septaria, 764, 760, 784.

Serpophaga

albagrisea, 1183.

cinerea, 1130.

— cana, 11380.

— grisea, 1130.

grisea, 1130.

parambe, 1138.

Sesamodon, 909,

923.

1210,

916,

INDEX OF SCIENTIFIC NAMES.

}

Sesamodon

browni, 913, 914, 915,

923, 925.

Setophaga

ruticilla, 1093.

Sialia

sialis, 857.

Sibia

capistrata, 859.

Siderastreea

radians, 1084.

Siphia

hyperythra, 720, 730.

Siptornis

erythrops erythrops,

1149, 1211.

— griseigularis, 1149,

1211, 1212:

— rufigenis, 1149, 1212.

Sitta

surinamensis, 1160.

Siurus

noveboracensis, 1091.

Spermophila

ophihalmica, 1088.

Sphecodes, 867.

Spheniscus

demersus, 571.

Sphenodon, 922.

Sphenuta

subulata, 1150.

Spilesoma

melanopsis, 709,

Spondylus, 1059.

sp., LOL.

Spongioderina, 872.

verrucosum, 870, 87-4.

Sporophila

aurita, 1099, 1212.

hicksii, 1099.

ophthalmica,

1099, 1212.

Stachvodes, 891.

gichristi, 871, 888,

893.

trilepis, 886.

Stelgidopteryx

ruficollis, 1093.

— uropygialis, 1098.

wropygialis, 1093.

Stenobothrus sp.,

846.

Stephanoseris

rousseaut. 1022, 1026,

Stilesia, 1002.

Stomotoca

dinema, 1051.

Stoparola

sordida, 720:

Strangalia

armatt, 839.

1098,

835,

muelanocephala, C39, |

650. {

Tetragonops

ramphastinus, 1200.

INDEX OF SCIENTIFIC NAMES. XXXV

Struthidea | Tenia | Tetrao

cinerea, 671, 859. mucronata, 649, australis, TAS.

Struthio tetragonocephala, 637. Thalassochelys

australis, 987. tuberculata, 635. | caretta, 624.

massaicus, 987. (Leniarhynehus) sagi- | Thalurania

molybdophanes, 987. | nata, 1018. funnyt, 1183, 1184,

Sturanya, 761. | Tamandua | 1,

Sturmopastor | tetradactyla, 636, 637, —_verticeps, 1184, 1211.

contra, 860. 638, 1018. Thamnobia

julle, 860. | Tanagara fulicata, 721.

Sturnornis | notahitis, 1111, Thamnophilus

senex, 720. | Tanagra atrinucha, L158.

Sturnus | maxima, 112). cachabiensis, 1167.

holosericeus, 1122, | melanoptera, 1112. immaculatus, 1163.

Stylocyathus, 1050. | palmarum., 1112. major, 1157.

Suastus — melanoptera, 1112. — granadensis, 1158,

gremius, 732, — vjolilavata, 1112. — melanurus, 115%.

Suberia, 872. peruviana, 1104, — transandeanus,

capensis, &70, 871, 873, | riefferti, 1120. 1157, 1158.

fohS)ZF, teh )5}, ruficer via, 1108. nevis, 1158.

clavaria, 873. (Aglaia) aurulenta, — atrinucha, 1158.

gentht, 873. 1102: | — nevius, 1159.

hellikeri, 873. (—) labradorides, 1109. | qurwensis, 1168.

Sulphurina, 761. (Calliste) ryfiverter, | fransandeanus, 1157,

Suricata 1108. | 1158.

suricatta, 856. | Tanagrella | Thanaos

~ Surniculus rufigula, 1102. tuges, 825, 831, 848.

lugubris, 724. | Taognathus, gen. nov., Therapha

Sus sp., 693. 1076. | — hyocyami, 846, 847.

Sylvia megalodon, 1076, 1081, | Vherjodontia, 894,

castanea, 1091. 1082. | Lhouarella

Symphalangus | Tapinocephalus, 919, | hicksoni, 886, 893,

syndactylus, 671, | 1074. striata, 871,

Synallaxis | Telehinia | Threnetes

castanea, 1148. viole, 704, 708, 709, | fraseri, 1177.

erythrops, 1149, G4, 726, 730; 740) | puckeri, 1177.

pudica, 1148. 7 ae 741. — fraser’, V77, W212)

rugigents, 1149. | Temenuchus — Tinea, — MAl7T

uniriufa, 1148. | pagodarumn, 720. 1212.

Synictis Vephrodornis Thrinaxodon, 899,

penicillata, 856, pondicerianus, 719. Thripadectes

Syrnium Teracolus flammulatus, 1149,

nuchale, 869. danae, 713. selateri, 1149, 1211.

Syrphus, 850. Jausta, 713, 727. Thryophilus

Terias leucapogon, 1089, 1090

Tachornis jfloricola, TAT. DUD

batassiensis, 723. hecabe, 708, 709, 710, nigricupillus, 1089.

Tachyphonus 727, 730, 735=747. — nigricapillus, 1089,

cassinit, 1116. libythea, 709, 735, ae

chrysomelas, 1117, Terpsiphone | — schattii, 1089, 1211,

delutrii, 1115, 1116. paradisi, 721. PA

melaleucus, 860. Tersina schotizi, 1089.

propingwus, 1115. viridis oceidcntalis, semibadius, 1089,

rubrifrons, 1115. 1099, NB,

xanthopygius, 1114, Testudo, 945. thoracicus, 1090, 1212,

1115. daudinii, 623. Thryothorus

Tenia, 994, 1003, 1017, elephantina, 628, | nigricapillus, 1089,

1018. greca, 624. | Thysanocephaluu

conferta, 6A9. nigra, 624, | crispum, 629.

Thysanosoma, 994, 996,

1000,

XXXV1

Thysanosoma

gambianum, 651, 652,

653, 658, 659, 995,

1002, 1008.

Thysanotenia, gen. nov.,

994, 1002.

gambiana, 997-1000,

1002, 1003.

lemuris, 994-1008,

1008.

Timarcha, 842.

tenebricosa, 819, 837,

841, 843, 845.

Tinamus

bonapartei, 1208.

Jrantzti, 1208.

Tinnunculus

alaudarius, 724.

punctatus, 700.

Tipula

oleracea, 848.

Tityra

albitorques, 1142.

— albitorques, 1142.

buckleyi, 1148.

personata, 1142.

semifasciata, 1142.

— columbiana, 1142,

1212.

— costaricensis, 1142, |

1212.

— semifasciata, 1142.

Todirostrum :

cinereum, 1128.

— cinerewm, 1128.

— sclateri, 1128.

Todus

cinereus, 1128.

Trachyphonus

caffer, 862.

Trematotrochus, 1038.

Jenestratus, 1030.

verconis, 1080, 1039.

zelandie, 1019, 1620,

1029, 1081-1084.

Treveleyana

erocea, 1064, 1068.

Tribonyx

ventralis, 864.

Triccus

sclateri, 1128.

Trichocephalus, 674.

Trichogaster

lahius, 985.

Trichoglossus

nove-hollandie, 671.

Trichosoma, 674, 676,

Trichostrongylus

retorteformis. 676.

Trichosurus, 918.

Trichuris, 674.

INDEX OF SCIENTIFIC NAMES.

Tridaena, 1059. |

gigas, 1058, |

Trionyx, 945, |

Trirachodon, 894, 905, |

908, 909, 913, 923. |

hannemeyeri, 906,

907.

Tritylodon, 898, 918,

919.

longevus, 925.

Trochalopteron

canorum, 859.

erythrocephalum, 859.

nigrimentum, 859.

Trochatella

chrysochasma, 798. |

Trochilus

aquila, 1180. |

aurelie, 1185. |

barroti, 1186. |

benjaming, 1186.

exortis, 1186.

Fannyi, 11838.

Navescens, 1185.

Jrancie, 1182.

hirsutus, 1178.

mellivorus. 1181. |

vuckeri, 1177.

feacatl, 1182.

yaruqui, 1178.

Trochocerecus

borbonicus, 698, 699,

7Ol.

Trochosmilia, 1040.

Trochus, 794.

Trogon

macrurus, 1193,

massena, 1193.

melanurus macrourus,

1195.

Trogopterus

xzanthipes, 689.

Tropicoris

rufipes, 840, 847.

Tropidonotus

rhombifer, 688.

Trypanosoma

lewis, 945.

Turbinolia, 1020, 1030.

Turbo, 794.

Turdus

ertnitus, 1137.

dague, 1086.

musicus, 857.

swatnsenii, 1087.

tristis cnephosa, 1087,

1212.

— dague, 1086, 1087,

1212:

ustulatus, 1087.

Turritopsis, 1049,

Tyrannula

ardosiaca, 1125.

cineracea, 1125,

erythroptera, 1134.

richardsonti, 1186.

sulphureipygius, 1135.

| Tyrannus

albicollis, 1134.

melancholicus, 1138.

— satrapa, 11388.

Udenodon, 923.

Upupa

epops, 862,

Uranomitra

Francie, 1182.

Urospatha

martit, 1198.

—— martii, 1193.

— semirufa, 1198.

Urosticte

benjamini, 1186.

| Valvata, 762.

Vanessa

haronica, 735.

Zo, 812, 822, 865.

urtice, 820,

865.

Veniliornis

kirkii cecilii, 1188.

— dariensis, 1188.

Viminella

Hagellum, 888.

Vincta sp., 748.

Viverra

jilchneri, 688.

etbetha, 688.

Volucella

bombylans, 812, 847,

853. 854, 855, 867.

Vulpes sp., 688.

822,

Waldemaria

Japonica, 784.

Xanthornus

mesomelas salvinii,

1122, 1128.

— taczanowshii, 1128.

Xenicopsis

mentalis, 1151, 1152.

subalaris lineatus,

1151.

— subalaris, 1151.

Xenops

genibarbis, 1152.

INDEX OF SCIENTIC NAMES. XXXVI1

Xenops Xestoleberis ; Zesius

genibarbis littovalis, nigromaculata, 595, | chrysomallus, 727.

1152; W212. 598, 6OL. Zizera, 722.

mexicanus, 1152, | Xiphophorus Zosterops

1212. helleri, 985. ceylouensis, 716.

littoralis, 1152. Xiphorhynchus chloronota, 701.

mecvicanus, 1152 lacrymosus rostratus, hesitata, 698.

ruficauda, 1152. 1153. mauritiana, TOL.

Xenopus, 919. palpebrosa, 716.

Xestoleberis Yyhthima, 710. (Malacirops) borbonica,

depressa, 595. 698.

latissima, 595, 598, GOL. | Zebrapicus Zschokkeella, 651, 660.

maryar ited, I95. | pucherant, 1188. Zygeena, 867.

INDEX

ILLUSTRATIONS.

Aiurosaurus tenuirosiris, Pl, UXII1.

p. 1073.

whattsi, P\. XIII. p. 1075.

Agelecyathus persicus, Fig. 220. y. 1035.

-Alcadia hollandt, Pils} XXX =—XiIL.,

p- 799.

palliata, Pls. XXX —-XXXII.,

p. 759.

Anopiotenia dasyuri, Figs. 208 215,

pp. 1004, 1006, 1007, 100%,

1010, 1012, 1018, 1015.

Aphanoconia andananica, Pl. XUL.

p. 759.

gouldiana, Pls. XXXVII., XLI.,

p. 790.

p- 799.

—— rogersti, P\. XLII. p. 759.

Argillecia affinis, Pl. XX. p. 595.

Avicula zebra, Fig. 280, p. 1058.

Bairdia dubia, Pl. XX. p. 595.

Bauria cynops, Pl. XLVI. p.

Figs. 168, 169, pp. 896, 897.

Bertiella cereopitheci, Figs.

pp. 640, 641, 643, 644, 647.

Budorcas bedfordi, P\. XX1X. p. 687.

893 ;

151-155,

Capra pyrendica hispanica, Pl. LIL.

p. 963 ; Figs. 195, 199, pp. 968,

973.

— —— pyrenaica, Bigs. 195, 198,

pp. 968, 972.

victorie, Pls. Lill, LIV.,

p- 963; Figs. 195-197, pp. 968,

970, 971.

Ceratoisis ramosa, Pl, XLIII. p. 870.

merguiensis, Pls. XXXV., XLIL, |

| Chama foliata, Pl. LX. p. 1057 ;

Fig, 229, p. 1057.

| Chromodoris inopinata, Pl. LXL

p. LOGS.

ene reticulata, P\. UXT. p. 1068.

| — ftinctoria, Pl. LXI. p. 1068.

| Columba

| Dasymetra

sp, Pls. XXIIL-XXVL.,

p- 601.

Conchoderma auritum, Big. 163, p 667.

Coronula diadema, Fig. 163, p. 667,

Cynognathus berryi, Pl, XLV E. p. 893.

-— platyceps, Figs. 171, 172, pp. 901,

903.

Cythere cingulata, Pl. XX. p. 599.

-—— crispata, Pl. XX. np. 499.

Cythereis deformis, Pl. XX. p. 595.

Cytherella ovalis, Pl. XXII. p. 595.

Cytheridets subulata, var. crenulata,

Pl XOX. p. 595:

Cytherura cribrosa, Pl, XXII. p. 598.

fossulata, Pl. XXII. p. 595.

—— maculosa, Pl. XXII. p. 595.

confenta, Pl. XOX VnLT.

p. 677.

Dendrophyllia sp., Pl. LVIT. p. 1018.

Diademodon, Pl. XLVI. p. 893.

Dielurodon whaitsi, Pl. UXIIL. p. 1073.

Diagram showing relative positions of

tooth-germs in Macropus hillardieri.

Figs. 187, 189, pp. 933, 936.

| Diagrammatic representation of two

species of corals which vary about

two distinct growth-modes, the

extremes of which converge towards

one another. Fig. 216 4, p. 1024.

representation of a single species

of coral varying about an imaginary

growth-mode. Fig. 216 2, p, L024.

INDEX OF ILLUSTRATIONS.

Dinotheriwn 1 OWI

p. 948.

hobleyi,

Equus asinus somaliensis, Fig. 202,

p. 989.

<x Hquus quagqga

chapmanni, Fig. 203, p. 992.

x Hguus zebra, Fig.

201, p. 989.

—— quagga chapmanni, Fig. 203,

p. 992.

zebra, Fig. 201, p. 989.

Eriphostoma microdon, Pl.

| p. 1073.

Eucrangonyx robertsi, Pls. XLIX.-LI1.,

p: 984.

Hunicella papillosa, Pl. XLII. p. 870.

Huplecaura media, Pl. XLV. p. 870.

Hutrochatella pulchella, Pls. XXXI.,

2LOOCUE, ZOO, XO-OOVINL, 20,

p. 799.

LXIIl.

Flabellun

p. 1018.

Play Vie

magnificum,

Gazella dorcas, Fig. 198, p. 962.

hayi, Fig. 193, p. 962.

Gomphcgnathus minor, Kigs. 175-178,

pp. 909-912.

Gorgonocephalus, Fig. 167, p. 874.

Heterocyathus equicostatus, Pl. LVIII.

p. 1018; Fig. 218, p. 1032.

alternatus, Pl. LVILTI. p. 1018.

heterocostatus, Pls. LVII., LVILII.,

p. 1018.

michelini, Fig. 218, p. 10382.

Hyla goughi, Pl. LXIV. p. 1082.

Ictidognathus parvidens, Pl. LXIII.

p. L073.

Lechriorchis validus, Pl. XXVII.

p- 677.

Leptocheirus bispinosus, Pl. XVIII.

p. 561; Fig. 146, p. 586.

guttatus, Pl. XVILI. p. 561.

—— hirsutimanus, Pl. XVIII. p. 561.

pectinatus, Pl, XIX. p. 561.

pilosus, Pl. XVII. p. 561,

pinguis, P\. XVIII. p. 561.

Lophiomys ibeanus, Fig. 190, p. 947.

XXX1X

Loxoconcha decipiens, Pl. XXT. p. 595.

obesa, Pl. XXI. p. 595.

subalata, Pl. XXI. p. 595.

LIucidella aureola, Pls. XXXII.,

XXXIV., XL, p. 759.

Macropus billardiert, Pl. XLVII. p. 926 ;

Figs. 181-189, pp. 927-929, 931-933,

9°75, 936.

Madoqua phillipsi, P\. LY. p. 977.

gubanensis, Pl, LY. p. 977.

—— —— fararensis: Pl, LY. p. 977.

—— piacentinii, Pl] LVI. p. 977.

-——— swaynei, Pl. LVI. p. 977.

Map illustrating the Distribution of

the Genus Megapodius. Fig, 166,

p- 750.

showing probable former and

present distribution of the Spanish

Ibex. Fig. 194, p. 965.

Margaritifera margaritifera, Pl. LX.

p- 1057.

Megaptera boops, Figs. 160-162, pp. 662,

664, 666.

Megapodius, Fig. 166, p. 750.

Melinodon simus, Pl. XLVI. p. 893.

Melitodes esperi, Fig. 167, p. 874.

Millepora alcicornis, Fig. 230, p 1088.

Merisia tyonsi, Pl. LIX. p. 1045; Figs.

222-228, pp. 1047, 1050, 1052, 1054,

1055.

Moschops capensis, P\. LXV. p. 1073.

Murex ramosus, Fig. 229, p. 1057.

Muriceides fusca, Pl. XLIV. p. 870.

Nythosaurus larvatus, Fig. 170, p. 899.

Ochetosoma formosum, Pl, XXVIII.

p. 677.

Oochoristica sp., Figs. 148-150, pp. 627,

628, 653.

Oreotragus saltator porteusi, Fig. 192,

p. 959.

Orohophuna pachystoma

PLL p: 759!

ponsonbyi, Pls. XXXII., XXXY.,

ponsonbyi,

XXXVIL., p. 759.

Oudenodon bolorhinus, Pl. LXIII.

p. 1078

Paleohelicina ide, Pls. XXXII.,

XXXV., XXXIX., XLI., p. 759.

Paracyathus cavatus, Pl. LVII.

p. 1018; Fig. 217, p. 1028.

| Paradoxostoma gracile, Pl, XX1.p. 590.

xl INDEX OF ILLUSTRATIONS.

Paradoxurus

p- 621.

Pigeons, Colour and Colour-pattern

Inheritaucein. Pls. XXITI.—-XXV1,

p. 601.

Porphyrio

p. 986.

Psammogorgia pulchra, Pls. XUIIL.,

XLYV., p. 870.

Putorius nigripes, Fig. 145, p. 559.

Pyrophyllia inflata, Pls. LVIL., LVIIL,,

p- 1018; Fig. 221, p. 1040.

larvatus, Fig. 147,

melanonotus, Fie. 200,

Rhinoceros bicornis, Fig. 191, p. 958.

somatiensis, Fig. 191,

p. 958.

Sclerochilus levis ?, Pl. XXII. p. 595.

Sesamodon browni, Pl. XLVI. p. 893;

Figs. 179, 180, pp. 914, 915.

Spondylus sp., Pl. LX. p. 1057.

Stachyodes gilchrist?, Pls. XLIV., XLYV.,

p. 870.

Struthidea cinerea, Fig. 164, p. 672.

= —_——_—~ —

Suberta capensis, Pls, XLIII., XLY.,

p- 870.

Taognathus LXII.

p. 1073.

Thouarella hicksoni, Pls, XLIV., XLV.,

p: 870.

Thysanosoma gambianum, Figs. 156-

159, pp. 652, 658, 658, 659.

Thysanotenia lemuris, bigs. 204-207,

pp. 995, 996, 998, 1001.

Tooth-germs in MMacropus billardieri.

Figs. 181-189, pp. 927-929, 931-

933, 935, 936.

Trematotrochus zelandie, Pls. LVII.,

LVIII., p. 1018; Figs. 218, 219,

pp. 1032, 1033.

Trichosoma sp, Fig. 165, p. 675.

Lrirachodon kannemeyeri, Figs. 173,

174, pp. 906, 907.

megalodon, Pl.

Aestoleberis latissima, Pl. XXI. p. 595.

-— nigromaculata, Pl. XXII. p, 595.

PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLELT STREET,

PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCIETY

OF LONDON.

1911.

PART, II.

CONTAINING Paces 557 to 868, witH 26 PuaTEs

AND 22 TEx?-FIGURES.

rs : .

J -xesonian tisty,

1" iy

VUI SO A9t 4

SEPTEMBER 1911. W/o)

PRINTED FOR THE SOCIETY,

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a [Price Twelve Shillings.

ES Ore CONT KIN es:

1911, Parr IID. (pp. 557-868).

EXHIBITIONS AND NOTICES.

ue Secretary. Report on the Additions to the Society’s Menagerie during the month of

February 1911

COE OO NON 8 Ce OO 0° OOOO DEO OOn Oh OO CHO dO Gatco noo ona oO mod OO COA mo ooUt

Mr. D. Szru-Suire, F.Z.S. Exhibition of a living Hybrid between the White-eyed

Pochard (Aythya nyroca) and the Marbled Duck (Marmaronetia angustirostris)

Mr. R. I. Pococs, F.R.S., F.L.S., F.Z.S. Exhibition of the skin of a new Chacma Baboon

(Papio porcarius, subsp. griseipes), a specimen of the North American Black-footed

Polecat (Putorius nigripes), ete. (Text-fig. 145.)

sete ee ee ee ee Oo ee ee eee eee ee te oe ee

Sir BE. Ray Layxesrer, K.C.B., F.R.S., F.Z.8. Exhibition of a special supplement to the

‘Field’ newspaper

Oe eee ee Ct Cc er we er err

Dr. R, T. Lurpzr, F.Z.8. A demonstration of Nematode parasites obtained from animals

in the Zoological Gardens

eC ee re i. rrr i, Prd

Mr. R. I. Pococs, F.R.S., F.L.S., F.Z.S8. Exhibition of a newly born Masked Palm Civet

(Paradoxurus larvatus)

eC Ce Cn CO ee i i i recy

ee ee eee eee OP ecw ee ee we we ow

Tux Secrerary. Letter on Land Tortoises in the Seychelles

Dr. H. B. Fayruan, F.Z.8., and Miss Annie Porrmr, D.Sc. Ona Bee-disease due to a

Protozoal Parasite (Nosema apis)

ee eee eee eee er PO Co eee re se eeeeee et ereesese ee thew

Tue Secrerary. Report on the Additions to the Society’s Menagerie during the month

of March 1911

ee ee ee eeco se Pe eee ese sees ee OH ee ee Cee FHP BOE ee oe PF eeseete es CO oe

Mr. C. Tare Ruean, M.A., F.Z.S. Exhibition of a series of lantern-slides of scales of the

Mallon (Sad720 Salary: ior ia oie as cale tele a tolere a wie teiotealas Cheb ce, wich chalet oleic cial aie eee

My. D. Sern-Ssirn, F.Z.S8. Exhibition of a nest of the Grey Struthidea or Apostle Bird

(Struthides cinerea), and lantern-slides of Penguins in moult and of wild Swainson’s

Lorikeets (Trichoglossus nove-hollandie). (Text-fig. 164.) ..........eseeececeeee

Dr. C. Curisty, F.Z.8. Exhibition of a collection of skins from Uganda and of a loin-cloth

taken from a native in Northern Nigeria .......... 0... 0ceeeceeeee f eievelepidereioes

Dr. Witutam Nicotn, M.A., F.Z.S. On a unique Pathological Condition in a Hare.

CMG KE tree GOS)! A26 Ne esters sate) slensieinialstauale Wiatieea fetetal choraso)o%= sic) wiehels ise sibys oh tenet ee en ee

Page

558

620

621

622

625

671

672

674

Contents continued on page 3 of Wrapper.

PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

EXHIBITIONS AND NOTICES.

March 21, 1911.

Dr. 8. F. Harmer, M.A., F.R.S., Vice-President,

in the Chair.

THE Secrerary read the following report on the additions made

to the Society's Menagerie during the month of February,

1911 :—

The number of registered additions to the Society’s Menagerie

during the month of February last was 123. Of these 38 were

acquired by presentation, 47 by purchase, 8 were received on

deposit, 24 in exchange, and 6 were born in the Gardens.

The number of departures during the same period, by deaths

and removals, was 138.

Amongst the additions special attention may be directed to :—

2 Northern Lynxes (Felis lyna isabellinus), from Tibet, pre-

sented by Capt. D. G. Oliver, on Feb. 17th.

1 Dwarf Mongoose (Helogale varia), new to the Collection, from

Lamu, presented by the Rev. W. D. Braginton, on Feb. 27th.

1 Black-footed Polecat (Putorius nigripes), new to the Collection,

from N. America, received in exchange on Feb. 16th.

2 Cat-Bears or Pandas (4lurus fulgens), from Nepal, purchased

on Feb. 20th.

1 European Bison (ison bonasus), from Lithuania, presented

Proc. Zoou. Soc.—1911, No. XL. 40

558 MR. R. I. POCOCK ON

by H.G. The Duke of Bedford, K.G., President of the Society, on

Feb. 23rd.

2 Dybowski’s Deer (Cervus hortulorum), from Manchuria, new

to the Collection, presented by Sir Edmund Loder, Bart., F.Z.5.,

on Feb. 23rd.

1 Aldunati’s Finch (Phrygilus aldunatii), from Chili, new to the

Collection, presented by Miss Phillis True, on Feb. 13th.

1 King Penguin (Aptenodytes pennanti), from the Antarctic Seas,

presented by Sefior Clemente Onelli, on Feb. 11th.

Mr. D. Seru-Smrru, F.Z.S., the Society’s Curator of Birds, ex-

hibited a living Hybrid Duck, which was believed to be a cross

between the White-eyed Pochard (Aythya nyroca) and the

Marbled Duck (JZarmaronetta angustirostris). It was hatched at

Scampston Hall, Yorkshire, from a clutch of eggs laid by a Mar-

bled Duck at Lilford Hall, Northamptonshire. - The bird displayed

the characteristic markings of Marmaronetta, but the general

colour was dark reddish brown. It had been presented to the

Society by Mr. W. H. St. Quintin, F.Z.S.

Mr. R. I. Pocock, F.R.S., F.L.S., F.Z.8., Superintendent of

the Gardens, exhibited :—

(1) A pair of Otter cubs about seven weeks old, which were

found under a landing-stage at Tewkesbury, and were presented

to the Society by Mr. W. Baring Bingham, F.Z.S.

(2) The skin of an adult female Chacma Baboon (Papio porcarius)

representing an apparently undescribed local race of that species,

which he proposed to name and diagnose as follows :—

PAPro PORCARIUS, subsp. GRISEIPES *.

Abstract P. Z.S. 1911, p. 17 (March 28th).

Of the size, general appearance, and coloration of the typical

form from Cape Colony, but distinguished by the absence of black

hairs from the upper sides of the hands and feet, the extremities

of the limbs and tail being clothed with grizzled or annulated

hairs.

Loc. of type: Potchefstroom in the Transvaal.

This animal was presented to the Society in 1904 by Mr. James

Adams, whose son had brought it from the above-mentioned

locality. She was adult at the time and died in 1910. On

arrival she was seen to differ in the characters enumerated from

the ordinary South African Chacmas, commonly received by

the Society, many of which are shipped from Algoa Bay and all

of which are believed to come from some part of Cape Colony.

This belief is borne out by Mr. W. L. Sclater’s t description of

* The complete account of this new subspecies appears here, but the name and a

preliminary diagnosis were published in the ‘ Abstract,’ No. 93, 1911.—Ep1ror,

+ ‘The Fauna of South Africa: Mammals,’ i. p. 14, 1900,

A NEW CHACMA BABOON. 559

P. porcarius, in which it is stated that “the lower part of the

arms and legs |are| darker than the rest of the body, almost black,

the upper part of the hands and feet quite black.” Since

Mr. Selater further adds that “there are in the South African

Museum skins and skulls from the Stellenbosch, Tulbagh,

Worcester, Beaufort West, and Albany divisions of the Colony,”

it may be inferred that his description of the typical form of the

species was taken from the material in question.

During its lifetime in the Gardens, the baboon did not show

any change in colour, so that there are no grounds for believing

that the differences above pointed out are due in any way to age.

The Society also possessed at one time a second specimen, an adult

male, deposited by the Hon. Walter Rothschild, which exactly

resembled the type ; but for this, unfortunately, no precise locality

was known. The exact northern range in 8. Africa of the typical

or black-handed Chacma appears to be unknown; but on the

evidence supplied by the type specimen of P. porcarius griseipes,

it may be supposed that the latter supplants the former at least

in the Southern Transvaal. Chaemas have been recorded by

Mr. Vaughan Kirby fron the country between Beira and the

Zambesi; and according to Mr. Sclater the species is abundant

in Natal and the Transvaal; but it does not appear that any

specimens from the latter country have been described or critically

examined hitherto.

Text-fig. 145.

Black-footed Polecat (Putorius nigripes).

(3) A specimen of the North American Black-footed Polecat

(Putorius nigripes) (text-fig. 145), recently received in exchange

from the Zoological Society of Washington.

The peculiar coloration of this animal, taken in conjunction

40*

560 ON A BLACK FOOTED POLECAT.

with its behaviour in captivity, suggests that it belongs to one of

the warningly coloured species of Mustelide. Although the upper

surface of the head, neck, and body is tinted with buff, owing

to the terminal portion of the long hairs being that colour,

the whiteness of the underfur, and of the basal portion of the

long hairs of these areas, together with that of the sides and

under surface of the body and of the greater part of the tail and

face, gives the impression of a whitish animal which must be

conspicuous against any dark background. Sharply contrasted

with the creamy tint of the body is the jet-blackness of the legs

and of the tip of the tail; while the whiteness of the face is

emphasised by a broad black band stretching across the lower

part of the forehead and the base of the nose and involving the

bright bead-like eye on each side.

Apart from its coloration, this Polecat exhibits in un-

mistakable manner that fearlessness which is so marked a

feature of protected animals. It never makes any attempt to

keep in the background or lie hid, like a savage cat, when

disturbed, but comes boldly to the bars of the cage with threatening

aspect, as if eager to attack the intruder, uttering every now

and again a shrill chattering scream. Its obstinate pertinacity,

indeed, in keeping to the front when any human being is in

sight makes the opening and shutting of the door of the

cage for cleaning and feeding purposes a matter of serious

difficulty. Nothing but a bass-broom, which few small mammals

will face, has any effect in making it beat a retreat. Finally,

when actively interfered with by being netted, it emits the

foetid odour so characteristic of species of the genus Putorius.

The combination of characters here mentioned—namely, the

repulsive smell, the courage and fearlessness of exposure, the

conspicuous coloration—are all found in the Skunks, the stock

examples amongst the Mammalia of protected self-advertisers.

Added to these attention-arresting characters is the penetrating

scream comparable as a warning signal to the rattling of a

Porcupine’s tail. These facts justify the placing of the Black-

footed Polecat in the category of aposematic Mustelines, an

account of which has already been published by the Society *.

(4) The antlers of an old male of the Manchurian Wapiti (Cervus

aanthopygus), that recently died in the Gardens, to show the

natural variation in the position and growth of the second or

“bez” tine. In one antler this tine was much smaller than the

first or ‘‘ brow” tine, and was situated nearly midway between it

and the third or “trez” tine, apparently as in the specimen of

this Deer described by Mr. Lydekker as C. bedfordi; but in the

other antler the bez-tine was as long as the brow-tine and placed

close above it as in normal antlers of the stags of this group.

* R. 1. Pocock, P. Z. 8. 1909, pp. 944-959.

duth, sc. et imp.

IGS VOCs US WULOSWS Aerveleleveta,

deg 20S), IS}, IAL OVAGUL,

EW. Sextom del. Huth,sc.etmp.

1S So nh PE OCEANUS GUl TLGWS Grube On 22 UNG UlS es umpsions

LSS iG. ERS U LEMANS) Bate 7520. Sis Pin OS tis iNicwemlean,

He a ae ae

PGS Oa go> aati

24, lke

EW. Sexton, del,

LEPTOCHEIRUS PHOTINATUS Norman:

ON THE AMPHIPOD GENUS LEPTOCHEIRUS. 561

PAPERS.

26. On the Amphipod Genus Leptocheirus.

By E. W. Sexton, Marine Biological Laboratory, Plymouth *.

[Received December 22, 1910: Read March 21, 1911.]

(Plates XVIJ.-XIX.+, and Text-figure 146.)

The genus Leptocheirus was instituted by Zaddach in 1844, with

the type species Z. pilosus. So much discussion has arisen over

this species that it seemed desirable to settle the matter by refer-

ence to the actual specimens, should it prove possible to trace them.

Dr. Braun, in whose keeping at the Konigsberg Museum they were

discovered, most kindly sent them to me for examination and also

granted permission to dissect and figure one of the specimens, so

that the vexed question might be finally set at rest. JI am deeply

indebted to him, and to all those also who have so generously

assisted me in this investigation : to Canon Norman, for the loan

of specimens of ZL. subsalsus, pinguis, hirsutimanus, guttatus, and

pectinatus ; to Monsieur Chevreux for specimens of L. cornwauret,

tricristatus, and dellavallei ; to Dr. Kiikenthal and Dr. Zimmer for

permission to examine Grube’s specimens of Protomedeia hirsuti-

mana? and P. guttata; to Dr. Hjalmar Théel and Dr. Holmquist

for the trouble they have taken in searching for Ohlin’s specimen

ot LZ. aberrans ; to Professor Steuer for his assistance in tracing

Heller’s specimen of P. hirsutimana; to Dr. Otto Pesta for the

description and figures of this same specimen, by which its

identification became possible; and to Professor Vayssiere and

Monsieur Collin, of Cette, for the help given in searching for

Catta’s specimen of massiliensis.

It will be noted that the number of species has been reduced.

L. cornuauret Sowinski and subsalsus Norman become synonyms

of the type species, the one being the full-grown male, and the

other the full-grown female. Norman’s record is very interesting

with regard to the distribution, confirming Zaddach’s account,

both being for fresh or almost fresh water. All the other species

of the genus, so far as yet known, are marine.

I have also shown, I hope conelusively, the identity of

L. pectinatus Norman with L. dellavallei Stebbing, whick indeed

Mr. Walker has always maintained. JL. pectinatus is, in my

opinion, the young form sexually mature but not full-grown,

while Z. dellavallei is the fully developed animal. Figures are

given of the characters on which Chevreux (16) p. 91, and

Norman (36) pp. 87, 88, separated them, to prove that the

differences are merely those due to sex and development. The

question of distribution is also discussed by these authors in

separating the species, pectinatus having always been found near

the shore, and dellavallei at greater depths, but our present

* Communicated by Dr. W. T. Cauman, F.Z.S.

+ For explanation of the Plates see p. 593.

562 MRS. E. W. SEXTON ON THE

knowledge of the bathymetrical limits of any species is far too

inddequate to base any conclusions upon. Grube’s hirsutimana

from the Adriatic is the same species.

L. tricristatus Chevreux becomes a synonym of guttatus Grube.

An interesting point in this species is the varying development of

different characters, notably the antenne, the last perzeopod, and

the last uropods.

L. bispinosus Norman I consider identical with the species

described and figured by Della Valle as Z. guttatus. Heller’s

P. hirsutimana is to be referred to this species (see p. 585).

I have been unable to trace the type specimens of two species,

L. aberrans Ohlin, and Protomedeia hirsutimana var. massiliensis

Catta. The only specimen of aberrans, dissected by Dr. Ohlin for

the purpose of description, appears unfortunately to have been

lost.

There is nothing to add to Mr. Stebbing’s definition of the genus,

‘Das Tierreich,’ p. 625, except that the outer ramus of uropod 3 in

all the species is 2-jointed, the terminal joint rudimentary.

The development of the secondary sexual characters in the male

can only be definitely stated in two species, LZ. pilosus and pinguis ;

a great deal more material must be examined before it can be

decided whether the rule which applies to these species holds good

for the whole genus or not. In pilosws and pinguis the first gna-

thopod in the male is longer than the second, the hand is greatly

developed, the palmar margin concave; while in the female, on the

contrary, the first gnathopod is shorter than the second, and the

palmar margin is convex. The full-grown female, in all the other

species, agrees with this description. I find another distinction

between the sexes in the type species (Z. pilosws), in the shape

of the basal joint of the 5th pereopod (see Pl. XVII. figs. 22

& 23); but as this is the only species in which I have had the

5th pereopods of both sexes, I cannot say if this distinction is

generic or only specific.

The species now included in the genus are as follows, arranged

in chronological order, with their principal distinguishing cha-

acters :—

1. L. pilosus Zaddach 1844, = L. cornwauret Sowinski, 1898, the 3,

and L. subsalsus Norman, 1908, the 2. Pleon-segments not

dentate: accessory flagellum 1-jointed: gnathopod 1 greatly

developed in ¢, longer than gnath. 2; shorter than gnath. 2

in 2; dth joint in ¢ powerful and curved, 6th bent inwards

at right angles to the 5th, palmar margin concave ; palmar

margin convex in ?: finger of gnath. 2 almost straight, apex

acute.

2. L. pinguis Stimpson 1853. Pleon-segments 4, 5, & 6 each with

2 dorso-lateral angles : accessory flagellum long, 6—8-jointed :

hind margins of sideplates 1—4 serrate, spiniferous : hand of

gnathopod 1 greatly developed in ¢, and much _ longer

than gnath. 2; much shorter than gnath. 2 in 9 ; palmar

margin concave in d, convex in ?: finger of gnath. 2 as in

pilosus.

3 LD,

(>)

“ID

(

Shp

i De

AMPHIPOD GENUS LEPTOCHEIRUS. 563

hirsutimanus Bate 1862 = Boeckia typica Malm 1871. Pleon-

segments not dentate, the 4th with a dorsal depression :

accessory flagellum 6- -jointed : sideplate 1 small, hidden by

the large sideplate 2: finger of gnath. 2 as in pilosus : fingers

of pereeopods 3-5 bifid : uropod 2 unusually massive.

guttatus Grube 1864 = Ptilocheirus tricristatus Chevreux

1886. Pleon-segment 4 with 3 medio-dorsal angles or teeth :

superior antenne not much longer than inferior ; accessory

flagellum 2-3-jointed ; palmar margin convex in 2: finger of

gnath. 2 unguiform, acute: falciform processes of uropods 1

and 2 of great length : inner ramus of uropod 3 tipped with

1 spine.

. pectinatus Norman 1869 = Protomedeia fasciata Costa 1864

and L. dellavallec Stebbing 1899: pleon-segments 4 and 5 each

with 2 dorso-lateral angles: sideplate 1 small, hidden by the

large sideplate 2: accessory flagellum 2—3-jointed : palmar

margin, gnath. 1, concave in ¢, convex in 2; finger of gnath.

1 exceeding the palm in length: finger of gnath. 2 straight,

laminar, tipped with setz.

aberrans Ohlin 1895.

bispinosus Norman 1908 = Protomedeia hirsutimana Heller

1866 and L. guttatus Della Valle 1893: resembles guttatus.

Pleon-segments 4 and 5 each with 2 dorso-lateral angles:

accessory flagellum 5-jointed: palmar margin, gnath. 1,

convex in 9: 2nd joint in gnath. 2 unusually long; finger

as in guttatus: falciform processes of uropods 1 and 2 of

great length: uropod 2 with two clusters of feathered sete on

the lower margin of the inner ramus.

Fam. PHorips.

For synonymy see Stebbing, Das Tierr. Lief. 21, p. 603.)

Gen. LEeprocHErRuS Zaddach.

(Stebbing, p. 625.)

LepTocHEIRus Prtosus Zaddach. (Plate X VII.)

1844.

1848.

1862.

1873.

1878.

1888.

1898.

1906.

1908.

1910.

Leptocheirus pilosus Zaddach (53) pp. 8 & 9.

wu Muller (32) p62:

Protomedeia pilosa Spence Bate (1) p. 168.

Leptochirus pilosus Mobius (80) p. 117.

Protomedeia pilosa Zaddach ( 54) 1 pp. 18 & 19.

Leptocheirus pilosus Stebbing (42) see p. 1707 for references.

45 cornuaureitSowinski(41) p. 470, pl. ix. figs. 9-22

3 pilosus Stebbing (44) p. 630.

é subsalsus Norman (38) p. 307, pl. xil. figs. 1-6.

Hf cornuauret = subsalsus Chevreux (18) p. 2.

This species was fully described by Zaddach in 1844, and the

accuracy of his description will be seen on comparing his account

with

the figures here given. In his later work, ‘Die Meeres-

564 MRS. E. W. SEXTON ON THE

Fauna an der preussischen Kiiste,’ is an interesting note on the

distribution of the species. It was found,so Zaddach states, both

in fresh and in salt water; by Rathke in 1843 in a large fresh-

water lake, the Geserich See, and by Zaddach himself, about the

same time, in the sea at Dantzig. It is not known under what

circumstances Rathke obtained his specimens, whether he collected

them himself near the banks, or whether they were given to him by

the fishermen. Zaddach, 34 years later, dredged the southern part

of the lake in search of the species, but his efforts proved fruitless.

The type * specimens are preserved in the Kénigsberg Museum,

and are labelled ‘‘ Protomedeia pilosa Zadd. Rauschen, Ostsee IX.

1866.”

The next record, also from the Baltic, is that of Miiller, who

noted the species as not rare in Greifswalder Bodden. He pointed

out the presence of a minute 1-jointed accessory flagellum on the

superior antenna, which had been overlooked by Zaddach.

The next authentic record of the species is in 1898, when

Sowinski described and figured the adult male as LZ. cornuwauret.

In 1908 Canon Norman described and figured the female under

the name of LZ. swbsalsus from specimens found by Mr. Gurney in

Norfolk, in fresh or almost fresh water.

In a note published in July, 1910, Monsieur Chevreux points

out that cornuaurei Sow. is the male of subsalsws Norman.

The examination of the type species which I have been enabled to

make and the comparison of it with specimens of both cornwawret

and subsalsws, prove beyond question the identity of all three forms,

pilosus being the young female, not sexually mature, cornuauwret

the full-grown male, and subsalsus the full-grown female.

The specimens examined were :—

Zaddach’s two types from the Baltic, measuring 4 mm.

around the curve; young females, not sexually mature.

4, 6 and 9, from muddy piles, R. Bure, Norfolk, Canon

Norman’s collection, measuring 5 mm. in a straight line.

4, $ and Q, from the coast of Algeria, sent by Monsieur

Chevreux, measuring 4—4°5 mm. in a straight line.

Description.

Body (Pl. XVII. fig. 1) smooth, moderately compressed ; pleon

without teeth, but last four segments with a seta inset on either

side of the median line.

Head about as long as the first two perzeon-segments ; lateral

corners rounded, not prominent.

Eyes oval in the small specimens, almost round in the large

ones. The ommatidia, which are large and few in number, are

* Prof. Braun informs me that these specimens were collected by Zaddach himself

in the Baltic at, Rauschen, about 35 km. northwest of Kénigsberg, in September 1866,

and are to be regarded as the types of the species. As the original description was

published in 1844, they are not, strictly speaking, types, but rather, perhaps, meta-

types.

AMPHIPOD GENUS LEPTOCHEIRUS. . 565

darkly pigmented in the centre of the eye; the outer ring in

the young animal is quite colourless, though this, of course, may

only be due to the fading of the pigment, little of which is left in

Zaddach’s specimens, while in Chevreux’s it is still fresh and

black.

Sideplate 1 free of sideplate 2, but not as deep and not quite

half its width ; obtusely rounded. Sideplate 2 the deepest of all ;

expanded inferiorly ; in Zaddach’s specimens and Norman’s small

one it is deeper than broad, but in the large specimens it 1s more

expanded in proportion to the other sideplates, in the largest of all,

a female, it is almost twice the width of sideplate 3; hind margin

straight, front angle produced and rounded. Sideplates 3 and 4

subequal, deeper than broad, of equal width throughout; front

and hind margins straight ; the 5th has the anterior lobe as wide

and as deep as sideplate 4, posterior lobe small; 6th and 7th

small ; 1-5 with a few piumose hairs and setules on the inferior

margin.

Pleon.—Segment 3 the largest; the 3rd—6th each with 2 sete,

one on either side of the median line, those of the 4th and 5th

segments the longest. These sete are inset submarginally on the

posterior margin in the same position as the angles or teeth of

the other species. Hind margin of the 2nd segment straight ;

that of the 3rd rounded ; inferior margins of 1-3 with plumose

sensory hairs, those of the 2nd longer and more numerous ; hind

margins 1-3 crenulate, 2-3 crenulations each with a setule inset.

ANTENNE (figs. 2 & 3).—Superior Antenna (fig. 2) much longer

than the inferior, about half the length of the body: 1st joint of

the peduncle stout; 2nd much more slender and longer than the

Ist in the large animal, in Zaddach’s smaller specimen it was

subequal to the Ist in length; 31rd not quite half the length of

the 2nd. Primary flagellum: Zaddach gives the number of

joints as 12, Norman as 13; one of Zaddach’s specimens, the

smaller of the two, had 11 joints, the other 12; Norman’s

largest specimen had 14; Chevreux’s 14, one specimen with 14

in one antenna and 15 in the other; the difference in number is

evidently due to development. The proximal joints are short,

the distal ones increasing in length, each, from the 5th or 6th to

the second last joint, being furnished with a stalked sensory

filament in addition to the small sete. Accessory flagellum not

as long as the Ist joint of the primary. It consists of 1 small

joint, which is so constricted near the apex as to give the appear-

ance of a minute second joint; furnished with 3 apical set, one

of great length, and 2 long setz above the constriction. Zaddach

completely overlooked this small appendage, which Muller was

the first to note, and indeed it might, as Miiller says, easily

escape attention, were it not for the long outstanding apical

bristles.

Inferior Antenna (fig. 3)—The proportions of the last three

joints of the peduncle are as given by Zaddach ; 3rd short, nearly

twice as broad as the succeeding joints; 4th the longest; 5th a

566 : MRS. E, W. SEXTON ON THE

little shorter than the 4th. The flagellwm is about half as long

again as the 5th joint; it is composed of 8 joints in Zaddach’s

specimens, of 10 in the larger ones, the terminal joint in all

microscopic.

OraL Parts.—Upper Lip (fig. 4): apex asymmetrically bilobed.

Lower Lip (fig. 5) large; outer lobes rounded, inner lobes

appear to be coalesced at the base.

Mandibles (fig. 6).—Both cutting-plates and accessory-plates

divided into 4-5 strong rounded teeth, the lowest being the

largest ; 7 dentate spines in the spine-row in Zaddach’s specimen,

9-10 in the large animals, the first two or three laminar. The

molar is large and cylindrical, crown ridged with rows of teeth,

and edged with flat spines, furnished with a long feathered seta

above ; in the figure the molar is not well represented, being

bent back to show the other portion of the mandible more clearly.

The palp is very large; 3rd joint the longest, tipped with 4

strong curved bristles, the distal inner margin furnished with a

double row of finely serrate bristles.

Masilla | (fig. 7) as described by Zaddach : inner plate large,

with 1 long plumose seta; outer plate with 11 strong spines on

its apex, 3 dentate ones at the upper angle, and the others

arranged in pairs, 1 bifurcate and 1 dentate together; in the

figure only one of the rows can be shown. The 2nd joint of the

palp widens towards the truncate apex, which is furnished with 4

strong short spines inset apically, and a diagonal submarginal

row of 4 sete.

Maxilla 2 (fig. 8) as large as maxilla 1; cnner plate the smaller ;

the apices of both plates carry long curved stiff bristles; the

inner margin of the inner plate has 2 rows of plumose sete,

setting out at different angles, one row containing 3 times as

many setz as the other.

Maxillipeds (fig. 9).—Inner and outer plates well developed ;

inner plates narrowed distally, the truncate apices inset with 3 flat

spines, outer margins with 3 setiform spines distally ; a row of 7

long jointed plumose setz crosses each of the plates diagonally

from the outer angle of the apex to the inner margin, extending

more than halfway down the latter. The outer plates are

widened distally ; in Zaddach’s specimen the plate on the right

side is furnished with 1 long plumose seta apically and 8 graduated

strong spines along the inner distal margin ; the plate on the left

has 2 of the plumose set apically, and 7 of the graduated spines ;

the outer surface carries numerous stiff curved bristles along the

inner margin. Palp, 2nd joint much the largest; 3rd much

produced on the inner surface over the insertion of the finger ;

finger small, obtuse, tipped with strong serrate setiform spines.

The 2 basal joints and the 1st-3rd joints of the palp are provided

on the outer surface with numerous long delicate plumose sete.

The Lirst Gnathopod (figs. 10-15) is longer than the second

in the male, shorter than the second in the female. It is

characterised by the remarkable development of the 5th—7th joints

AMPHIPOD GENUS LEPTOCHEIRUS. 567

in the male. In the female (fig. 11) the hand is shorter than the

preceding joint and lies in the same plane with it, but in the male

it is a quarter as long again as the 5th joint, and is carried bent

at right angles to it (fig. 12). The finger differs also: in the

male it is much stouter, more curved, and with a depression

in the inner margin proximally. The long 2nd joint is fur-

nished with a cluster of long delicate sete on the posterior

margin, and a row of short plumose setze on the anterior margin,

with another row of longer similar ones on the inner surface.

3rd joint bulging behind; 8rd and 4th fringed posteriorly with

long plumose sete ; 5th joint with transverse rows of plumose

sete, and 4 fan-like clusters of rigid serrate bristles on the

posterior margin. These bristles (fig. 15) are found on the 5th

joint of the first gnathopod in all the species of the genus. The

6th joint or hand is subequal to the 5th in length in the imma-

ture specimen (fig. 10); distally widened; palm slightly oblique ;

palmar margin microscopically pectinate, palmar angle with a

row of spines on the outer side, and 1 large spine on the under

side, with the tip of the finger fitting between them. The outer

row consists of 6 graduated slender spines, the first much the

Jongest, the shafts of which are produced apically into two unequal

processes, with a flat, delicate, feathered end-piece between the

processes. In one specimen there were 6 on one gnathopod and

7 on the other. On the inner side of the palmar angle a very

large stout sensory spine is inset, accompanied by 4 short stumpy

bristles similar in construction to those of the outer row. A row

of these stumpy bristles is found submarginally on the under

surface of the palm, and 3 or 4 rather larger ones on the outer

side. The finger in the young (fig. 10) is stout and curved, as

long as the palm, also microscopically pectinate, with a short

decurrent tooth near the apex, 2 sete inset in the notch, 1

smaller tooth behind, and 1 setule proximally. The full develop-

ment of the finger of the female is shown in fig. 13 and of the

male in fig. 14.

Second Gnathopod (figs. 16-18).—2nd joint very long, laminar,

with two rows of exceedingly long plumose set anteriorly,

one marginal, and the other submarginal on the under surface ;

posterior margin with only 3 or 4 long simple sete. A chitinous

ridge extends diagonally across the distal half of this joint, and

terminates at the posterior angle of the 3rd; this appears to be

the “crista” referred to by Zaddach (“in superficie externa

crista quadam a basi secundi articuli ascendente”). The 4th, 5th,

and 6th joints are practically subequal to each other in length ;

distal margin of the 4th joint with very long plumose sete on the

under side; posterior margins of the 5th and 6th with fan-like

clusters of short, stiff, finely serrated sete, anterior margins with

long plumose sete, the number of which increases a little with

growth, the smallest specimen having 9 on the 6th joint and

the largest 11, 3 on the 5th joint in all specimens. The

6th joint is slightly narrowed distally. The finger is almost

568 MRS. E. W. SEXTON ON THE

straight, the tip curved and 2 setules inset subapically. I have

given figures (see figs. 17 & 18) from both Zaddach’s and

Norman’s specimens drawn to the same magnification.

Pereopods 1 and 2 (figs 19 & 20) glandular; 4th joint very

much expanded, as noted by both Zaddach and Norman; 6th

joint as long as the 4th, but much narrower; 5th only half the

length; finger strong, two-thirds the length of the preceding

joint, with the gland-aperture at the tip.

Hinder Pereopods (figs. 21-23) rapidly increase in length; the

6th joint in all is much narrower and longer than the pre-

ceding. In pereopod 3 the joints are stout and short, furnished

with a few strong sensory spines ; 2nd joint obliquely oval, about

as wide as long, the anterior margin inset with 3 clusters of

jointed sensory ciliated hairs, posterior margin slightly crenulate,

with a setule in each crenulation, and with 4 plumose hairs

submarginally; finger about half the length of the 6th joint,

falciform. In pereopod 4 the 2nd joint is oval, longer than wide,

anterior margin with clusters of the sensory plumose hairs, pos-

terior Inargin crenulate, with a submarginal row of close-set long

ciliated hairs. In pereopod 5 there is a remarkable difference in

the shape of the basal joint in the male and female. In the

female the joint is oval, the posterior expansion narrowing distally,

while in the male it widens distally and is produced downwards

in a rounded lobe, but the curious part is that the ciliated sensory

hairs, instead of being inset close to the margin as in the female,

are at some distance from the margin on the under surface, but

yet give the same outline as in the female (cf. figs. 22 & 23).

Pleopods (fig. 24).—Inner rami twice the length of the peduncles;

outer rami the shorter, about three-quarters the length of the

inner. The peduncles of the 1st pair of pleopods carry each

on the outer side 9 long plumose hairs, on the inner side 1

exceedingly long plumose hair and 2 small coupling-spines with

recurved apices, the upper one (fig. 24) with 3 recurved teeth

on either side, and the lower with only 2. The peduncles of

the 2nd and 3rd pairs have only 1 or 2 long sete on the

outer side. The number of joints in the outer ramus is 11

in the Ist pleopods, 10 in the 2nd and 3rd; in the inner ramus

10 in the Ist, and 9 in the other pairs.

Uropods (fig. 25) extend backward to the same level; lst and

2nd pairs much alike in construction. The peduncle of the 1st

is as long as the inner ramus, with 3 or 4 spines on the

upper curve (3 in the small specimen, 4 in the larger), and

1 curved spine underneath, and the falciform apical process

reaching to half the length of the inner ramus ; outer ramus the

shorter, 1 spine inset on the upper margin, apex blunt with

a cluster of 4 stout spines; inner ramus with a similar apical

cluster and 2-3 along the upper margin. In uropod 2 the

peduncle is not as long as the inner ramus; it equals the

outer in length and carries 1 spine; rami as in uropod 1.

In uropod 3 the peduncle is subequal to the rami in length,

outer ramus, if anything, slightly longer than the inner, with a

AMPHIPOD GENUS LEPTOCHEIRUS. 569

cluster of setiform spines at the apex ; inner ramus with | spine

midway, and | spine and 2 setee at the apex; the large animals

have more spines ; all the spines are short and stout, “each with

an apical filament. The outer ramus is 2- “jointed, the terminal

joint rudimentary, carrying | long serrate spine or bristle.

Telson (figs. 25 & 26) short, broader than long, depressed

in the middle between the prominent lateral angles; apex

rounded ; 1 setaand 1 ciliated sensory hair on each angle, and

a pair of small ciliated hairs on either side.

Colour described by Zaddach as “ flavescens, dorso punctis

nigricantibus sparso.” In both his specimens and Norman’s the

colour has faded, but m Chevreux’s it is still fresh and vivid.

The whole animal is a beautiful golden yellow tint with stellate

markings in dark brown, These markings extend over the whole

dorsal surface of the head ; the anterior margin of the Ist, and the

posterior portions of the 3rd-7th pereon-segments and the Ist

pleon-segment are banded with them. The 5th and 6th pleon-

segments and the telson are entirely covered. An irregular band

of brown runs along each side of the perzeon just above the side-

plates and is continued along the epimera of the pleon to the

telson. All the side-plates and the posterior expansions of the

basal joints of the hinder pereeopods have each a patch of brown.

Distribution :—

GrsericH See, Prussia: and the Bauric: Zaddach (58)

I. pilosus, and (54) as Protomedeia pilosa.

Bauric : Greifswalder Bodden, Muller (32) as Z. pilosus.

Bauric: Mébius (30) as Leptochirus pilosus, depth 1-10 fins. ;

bottom, zostera and ulva.

R. Burn, Norfolk, England: Norman (38) as Z. swbsalsus.

The BospHorts: Sowinski (41) as LZ. cornuawrer.

Coast of Atgmria: Chevreux (18) as LZ. cornwaurei.

LEPTOCHEIRUS PINGUIS Stimpson. (Plate X VIII. figs. 10-12.)

For synonymy see Stebbing, Das Tierr. pp. 627 & 738, and

Norman (38) p. 309.

There is little to add to the summary of characters given by

Stebbing. The few notes subjoied were taken from three slides

kindly lent: to me by Canon Norman, and prepared by him from

specimens from Vineyard Sound and Long Island, N. America.

Head, 9 , lateral corners not much produced, truncate.

Superior Antenna.—I|st and 2nd joints in ¢ and @ practically

subequal to each other in length; 3rd a little move than one-

third as long as 2nd. Primary flagellum, 2, broken, 21 joints

remaining, all, except the first 6, with a small sensory filament

and short sete; 6 very long joints in the accessory flagellum which

equals 3 joints of the primary in length. In the male, the primary

flagellum is composed of 31 joints, Gah except the first 5, with

a small sensory filament. The accessory is broken, 6 joints

remaining, equalling 5 joints of the primary in length.

570 MRS. E. W. SEXTON ON THE

Inferior Antenna, 9 .—8srd joint short and broad, with 2 sensory

spines on the upper distal angle; 4th joint the longest; flagellum

longer than 5th joint, equal to 4th ; 11-jointed in this specimen,

each joint with a cluster on either distal angle of setiform spines

and very long set.

Lower Lip, 2, outer lobes rounded, densely setose ; inner lobes

coalesced, large, distally narrowed.

Mandible, 2 : cutting-plate on the right side produced below,

margin divided into 4 rounded teeth, the lowest the largest ;

accessory-plate much as in pectinatus, tapering from the base to

the long acute tip, with 2 very minute teeth above; 17 spines in

the spine-row, the first 4 large, the last 2 very small. Molar

prominent ; crown reniform in shape, edged with strong teeth,

and carrying a small accessory process on the side nearest

the cutting-plate. Palp: 2nd joint longer than Ist; 3rd

the longest, curved, falciform, attenuated distally, and armed as

in the type species with 4 long stiff curved bristles apically,

2 dense rows of similar shorter bristles on the inner margin,

and about 4 clusters on the outer margin. The left mandible

was broken.

Mazxilla, 1 2, much as in the type species; outer plate in one

maxilla with 11 strong spines, 3 dentate ones at the upper angle,

and the others set in pairs, 1 bifureate and 1 dentate together ;

in the other maxilla there were 12 spines.

Mazwilia,2 2, as in pilosus; inner plate narrowed distally.

Mavillipeds, 2 , covered with numerous long set. The basal

joints, and the Ist joint of the palp, on the outer side carry

fan-shaped groups of exceedingly long plumose sete; the outer

side of the 2nd joint of the palp is also covered with them, and

carries in addition a fringe along the inner margin. Outer plate

expanded distally, densely fringed on the inner margin with

slender spines, the apical ones of great length. Jnner plate

elongate, tapering distally to the narrow truncate apex; 3 flat

spines are inset in the apex, almost hidden by the plumose setz

surrounding them; the outer margin is furnished with a row of

about 10 simple setiform spines, and the inner carries a row of

18-20 very long, flexible, plumose setze. The 2nd joint of the

palp wuch the longest, much wider and about twice as long as

the 3rd; the 3rd produced over the 4th as in the type species ;

4th small, tipped with strong serrate spines,

First Gnathopod, 9 .—5th joint very long, provided on the

posterior margin with transverse rows of feathered sete, and

the clusters of stiff serrate bristles found in all the species of

the genus; 6th joint subequal to the 5th in length, as described

by Stebbing ; palmar margin convex, serrate. On the outer side,

extending from the angle nearly to the middle of the palm, is a

graduated row of bristles, 7 in number, similar in structure to

those of the type species; a row of smaller bristles is inset

submarginally along the palm. On the inner side, at the palmar

angle, is a short, very broad, sensory spine, and a submarginal

AMPHIPOD GENUS LEPTOCHEIRUS, 571

row of about 17 very small bristles, like those of the outer side.

The finger is exactly the length of the palm ; being closed in the

specimen examined, the detail could not be seen, owing to the

palmar spines obscuring it.

This gnathopod in the male differs from that of the female, as

in the type species. The 5th joint is longer in proportion ; the

2nd and 6th joints shorter. The 5th joint equals the basal joint

in length. The 6th is not quite two-thirds its length, and in the

natural position is held bent almost to a right angle with the

preceding joint. It is expanded distally; front margin very

thick; hind margin laminar, convex on the upper surface, and

furnished with six transverse rows of sensory sete ; palmar

margin transverse, concave instead of convex, furnished with the

submarginal rows of bristles along the margin, and the graduated

group at the angle as described for the female; the very stout

strong spine defining the angle is much larger than in the female.

On the under side of the hind margin of the 5th joint (not the

“basal joint” as given by Holmes (26) p. 522), distally, a chitinous

spine-like process is developed; another similar but smaller one is

on the 6th joint proximally, with a groove beside it, into which

the larger one appears to fit when the hand is bent in. In order

to show these processes, the hand is represented (fig. 10) with the

hind margin uppermost, and, in consequence of being placed in

this position, the true measurements of this joint cannot be seen ;

viewed from the side, it is exactly the same width proximally as

the 5th, expanding gradually to the palmar margin. The finger

in the male i is much more arched than in the female: and w fea

closed, the tip, instead of meeting the angle as in the female,

impinges against the under surface of the hand; it is finely

serrate, with a row of setules inset, and apparently a small

auxiliary tooth near the apex, but this was too obscured hy the

overlying palmar bristles to be ascertained with any certainty.

Second Gnathopod (fig. 11) as figured by Norman. The finger

is about half as long as the preceding joint, lightly curved,

acute, of the same structure as in the type species; with 4 sete

on the inner margin, and a cluster of 3 subapically in the male ;

fewer setee in the female.

Pereopod 5.—The finger is of unusual length, considerably

longer than those of perzopods 1 and 2; very slender, alinost

straight, with a setule near the apex.

Or -opod 3 (fig. 12), 9 A figure is given of the rudimentary

2nd joint of the outer ramus, with the spines omitted.

LEPTOCHEIRUS HIRSUTIMANUS Spence Bate. (Plate XVIII.

figs. 13-16.)

The synonymy as given by Stebbing, Das Tierr, p. 627, omitting

the references to Heller 1866, and Grube 1866, and adding : =

L. pilosus Norman and Scott (36) p.

The animal described by Heller as Bie Soe | hirsutimana is

572, MRS. E. W. SEXTON ON THE -

a female of Z. bispinosus (p. 585): Grube’s specimens proved on

examination to be pectinatus (see p. 577).

Sars has given an excellent description and figures of this

species, (40) pl. 197, to which it is only necessary to add one or

two details.

Superior Antenna, flagellum furnished with very long sensory

filaments.

First Gnathopod (fig. 13).—The palm is oblique, serrate, with

its limit defined, as in pectinatus, by a large sensory spine inset

on the under side. The tip of the finger reaches to this spine.

The palm carries a small strong spine on the outer side, close to

the insertion of the finger.

Second Gnathopod.— The finger (fig. 14) is exactly as in the

type species, cf. Pl. XVII. figs. 17 & 18.

Hinder Perceopods.—Sars figures the finger of perzeopod 5 only

as bidentate, but the finger in all three hinder pereopods is of

the same structure (see fig. 15). Spence Bate in his original

description notes it for the 3rd perzeopod, (1) p. 169; and Malm,

(27) p. 546, describes and figures it for all three.

Uropod 3 (fig. 16).—The outer ramus is 2-jointed, as in all the

other species of the genus, but the 2nd joint is exceedingly small

and very difficult to observe except from the dorsal view, owing

to the apex of the Ist joint being produced beneath it.

LEprocHEIRus Gurratus Grube. (Plate XVIII. figs. 1-9.)

1864. Protomedeia guttata Grube (23) p. 63.

1866. ms - », (24) p. 408, pl. x. fig. 3

1885. si Camus: (6) go 7.

1886. Prlnchere us tricristatus Chevreux (9) p. xl.

1887. ™" i LO) ies

1887. a oa (11)p. 310, pl.v. figs. 3 & 4,

and fig. 4 in the text, p. 578.

1888. Leptocheirus guttatus Stebbing (42) p. 366.

1898. 7. 7 Ghevreux (15) p. a

1900. he (16) p.

1906. 53 guitatus Stebbing (44) p. oh

tricristatus Stebbing (44) p. 629.

1906. = guttatus Norman & Seote (36) p. 85, pl. ix.

figs. 4-7.

1907. 5 guttatus Norman (87) p. 369.

1910. e tricristatus Chevreux (18) p. 2

The specimens examined were :—

1 Q, Grube’s type specimen, 5°5 mm. long, from Breslau

Museum.

3 2 L. trieristatus, taken by Monsieur Chevreux in the Bay of

Quiberon, the largest measuring 4°5 mm.

9 taken by Canon Norman at Falmouth, the largest 9?

measuring 6°25 mm.

The type specimen from Breslau has Dr, Grube’s original label

AMPHIPOD GENUS LEPTOCHEIRUS. 573

still on the bottle—‘* Protomedeia guttata Gr. m. Hiern, Lorenz,

Val Cassione 36.” It is a female, 55mm. in length from the

tip of the rostrum to the tip of the telson, with the pigmentation

still vivid after 48 years in alcohol. I have described this

specimen in detail, the account of the specific characters given by

Grube not being sufliciently adequate for the present system of

classification. In the figure of the whole animal it must be noted

that as all the appendages were drawn in situ, the ewact

measurements and proportions of the joints of the pereopods

cannot be expected. Hach character is compared with Chevreux’s

specimens of tricristatus to prove the identity of the two forms.

Description.

Body (Pl. XVIII. fig. 1) more stoutly built than in pilosus and

pectinatus, and sideplates shorter in proportion than in those

species.

Head longer than the Ist and 2nd pereon-segments taken

together ; lateral corners not prominent, rounded.

Hyes round, black or brownish-black in colour ; ommatidia large.

Sideplates 1-5 subequal to each other in length. Sideplate

1 not covered by sideplate 2; produced forward over the side of

the head, of equal width throughout ; inferior margin obtusely

rounded, with 3 or 4 stiff, sparsely feathered sets inset; hind

margin, as in the type species, not continuous proximally. Side-

plate 2 the largest, a little expanded distally, front angle and

inferior margin rounded, hind margin straight; inferior margin

fringed with a row of flexible, finely plumose sete. Sideplates

3 and 4 alike in form, deeper than broad, slightly wider proximally ;

sideplate 5 with the anterior lobe about as deep as preceding

sideplate, but not as wide; inferior margins of all three carry a few

setze, similar to those on sideplate 1. Sideplates 4 and 5 small.

Pleon.—Segment 3 much the largest, as long as segments | and

2 taken together. The 4th segment has the armature characteristic

of this species; the posterior margin is produced in 3 processes,

the median one upstanding, acute, shorter than the lateral ones;

the latter broad at the base, much produced over the succeeding

segments, apices acute, curving upwards, with a spinule inset in

each. Segments 5 and 6 are small, the posterior margin of the

5th with a few setules dorsally. The hind margin of the 2nd

epimeron is straight, with 1 crenulation and setule at the postero-

lateral angle; inferior margin densely fringed with long plumose

sete. The hind margin of the 3rd is produced and rounded, with

2 indentations, a setule in each; inferior margin lightly concave,

with 3 long plumose sete inset midway, and 3 short spines near

the antero-lateral angle.

Antenna. Superior Antenna (figs. 2-4).—1st joint of the peduncle

stout, shorterthan the 2nd; 3rd not quite half the length of the 2nd.

Primary flagellum 7—-12-jointed, the largest female had 11 joints ;

accessory flagellum 2—3-jointed. I must note here an occasional

Proc. Zoot. Soc.—1911, No. X LI. Al

574 MRS. E. W. SEXTON ON THE

curious variation in the number and proportions of the joints of

the flagella in the same animal. This variation occurs, I believe,

not infrequently in the Amphipoda, some species of Jassa, for

example, having usually one joint more on one side than on the

other. In Grube’s type specimen the primary flagellum on the

right side has 8 joints, 7 on the left; the right flagellum of the

inferior antenne has 4, while the left has only 3; the accessory

flagella are both 2-jointed, but the one on the left antenna is

longer than that on the right. Grube evidently counted the

joints of the antenne on the left side. In two other specimens

examined, females of the same size, the superior antennz were the

same length, but in one case there were 12 joints, and in the other

only 9; the accessory flagellum in both was 3-jointed, but whereas

in the animal with 12 joints it exceeded the 2nd of the primary

in length (fig. 4), in the other it did not reach to the level of the

2nd. Figures 2 and 3 are from another specimen, the right

accessory of which had 2 joints, the left 3. All the joints of the

primary, except the first 4, are provided with a sensory filament

in addition to the sete.

Inferior Antenna.—A4th joint of the peduncle the longest ;

flagellum a little longer than the 5th, subequal to the 4th in length.

Flagellum 3-6-jointed, the first the longest.

First Gnathopod (fig. 5).—Hand about twice as long as broad ;

palm oblique, defined, as in pectenatus, by a large sensory spine

inset on the under surface ; palmar margin convex, serrate, the

serrations turning the corner and ending at the level of the spine,

it carries 5 or 6 small sharp spines subinarginally on the outer

surface, and 8 smaller ones on the inner, all microscopically

serrate. The finger is as long as the palm, curved, serrate, with

2 auxiliary teeth.

Second Gnathopod (fig. 6).—2nd joint very long, equalling in

length joints 4-6 taken together. The 4th joint, as is usual in the

genus, is much produced over the 5th on the inner side, and if the

measurements are taken along this side, the 4th, 5th, and 6th are

seen to be subequal to each other ; Chevreux’s figure (pl. v. fig. 4

shows the outer side. The finger is about three-fifths the length

of the preceding joint, lightly curved, with 2 setules near the acute

apex.

MESReonae 1 and 2 alike in structure, glandular, resembling the

type species in the shape and proportions of the joints except that

the 5th joint is slightly longer in proportion than in pilosus, and

the finger is much longer and more slender, being equal in length

to the posterior margin of the preceding joint.

Hinder Per manos very like those of the preceding species, but

stouter and much less elongate. Norman gives ‘‘the greatly pro-

duced last pereeopod” as a specific character, but in most of the

animals examined by me the proportions are much as figured in

Grube’s type specimen (fig. 1); in the others the 6th joint is more

developed. The basal joint in all is large, rounded oval, produced

downwards in a lobe, furnished anteriorly with short spines at

intervals and ciliated tele s,and adense cluster of the latter at the

AMPHIPOD GENUS LEPTOCHEIRUS. 575

distal angle; the posterior margins of the 3rd and 4th joints are

crenulate, 6-9 crenulations each with a setule. In the 5th this

margin has 5-7 serrations, and 1-3 ciliated hairs inset in the

lobe. The 3rd joint carries a dense cluster of ciliated hairs at

the anterior distal angle; and the 5th joint a similar cluster

of feathered sete. Fingers short and curved. In peropod 3

the 4th joint is slightly longer than the 6th and wider;

5th shorter than 4th; 6th with groups of short spines. In

pereopod 4, the 4th and 6th joints are subequal to each other

in length; in one or two of the larger specimens the 6th is

slightly the longer; it carries numerous spines and a cluster of

very long setz at the distal angle. In perzeopod 5, the 4th, 5th,

and 6th joints rapidly increase in length; the 6th is slender and

lightly curved, inset on its posterior margin with exceedingly long

setze in addition to the spines.

Pleopods much as in pectinatus, except that the peduncles are

shorter in proportion to the rami, about one-third as long, and the

outer ramus is only two-thirds the length of the inner ; cleft spines

asin pectinatus, 5 in the 2nd pleopod; coupling-spines with 2 rows

of recurved teeth in addition to the recurved apex, 3 teeth in a row

in the lower spine, 2 in the upper.

Uropods (figs. 7 & 8).—There is a marked variation in the

length of the uropods in different specimens, most noticeable in the

3rd pair, connected apparently with the variation in the develop-

ment of the last pereeopod. In Grube’s specimen (fig. 8) the

peduncle of uropod 3 is short and broad, shorter than the outer

ramus; inner ramus not quite half the length of the outer. Some

of Norman’s and Chevreux’s specimens are like Grube’s type; in

others, the peduncle equals the outer ramus in length and the

inner ramus is nearly as long as the outer; while in the largest

specimens with the last perzeopods much produced, the peduncle

of uropod 3 is longer than the peduncle of uropod 2, and longer

also than the rami, both rami long and slender, inner quite equal

to the outer in length. Uropod 1, peduncle shorter than outer

ramus, the falciform process of unusual length, reaching almost to

the tip of the outer ramus, furnished with 5 spines along the

upper curve; outer ramus a little shorter than the inner, with

3 spines and an apical cluster of 5, 3 of which are longer than

the others; inner ramus with 4 spines and a similar apical group ;

these spines are longer and more slender than those of uropod 2.

Uropod 2 stoutly built, peduncle short and stout, with the falci-

form process equalling the outer ramus in length; outer ramus

shorter than the inner, with 2 spines, inner with 3, each with an

apical cluster of 5 strong stout spines (fig. 7). Uropod 3,

peduncle produced underneath in a flat laminar expansion; outer

ramus with a rudimentary 2nd joint carrying | long stiff sparsely

feathered bristle; the lst joint has a group of graduated similar

bristles almost concealing the terminal joint; inner ramusslender,

tipped with 1 stout spine. This description of uropod 3 applies

to all the specimens examined by me,

Telson (fig. 9) as figured by Chevreux and Norman: the apical

4\*

576 MRS. E. W. SEXTON ON THE

margin is convex; from the dorsal view it appears truncate,

cf. also bispinosus p. 089.

Colowr.—Chevreux in his original description gives the colour

as yellowish with some brown spots; in his later account he

describes it as generally uniform yellow, some specimens with

brown bands dorsally. In Grube’s type the colour is still vivid,

tawny yellow, with round stellate markings in light and fins

reddish-brown. These markings occur on the head ; in transverse

bands on segments 2-10; on the sideplates ; and on the posterior

expansion of the basal joints of the hinder peropods.

Distribution :—

CuanneL Isuanps: Norman (87) p. 369, as LZ. guttatus.

Fatmoutu Harsour: Norman & Scott (36) p. 86, as L. guttatus.

Oceanic Coast of France: Chevreux (11) p. 311,as Ptilocheirus

tricristatus, depth 7 m.; bottom deposit, gravel with coral-

lines; (12) p. 578, depth 10 m., bottom deposit, nullipores ;

(15) p. 481, as Z. guttatus, depth 10-50 m.

MEDITERRANEAN :—~ Coast of France: Chevreux (16) p. 92, as

L. guttatus. Coasts of Augmrta & Tunis : Chevreux (16) p. 92,

as L. guttatus; (18) p. 3. Apvriatic: Grube (24) p. 408, as

Protomedeia guttata.

LeprocHEIRus PecTINATUS Norman. (Plate XTX.)

The principal references to this species are as follows :—

1864. Protomedeia fasciata (non Kroyer 1842), Costa (19) p. 155,

pl. 1. fig. 8.

1864. Protomedeia Kr.

1864. Leptocheirus pilosus Grube (22) p. 73.

1866. Protomedeia hirsutimana? Grube (24) p. 402.

_ pilosa Grube (24) p. 417, pl. x. fig. 2.

1869, 3 pectinata Norman (34) p. 283.

1887. Ptilocheirus pectinatus Chevreux (11) p. 309.

1888. Leptocheirus pectinatus Stebbing (42) p. 1707 for references.

1893. A pilosus Della Valle (21) pp. 427-430, pl. iv.

fig. 10; pl. xii. figs. 1-14

1895. ss ,, Walker (47) p. 470.

1895. » (48) p. 310.

1899. f dellavallei. Stebbing (48) p. 350.

1900. 24 pilosus Chevreux (16) p. 90.

3 fasciatus Chevreux (16) p. 91.

1906. Be dellavallei Stebbing (44) p. 628.

i. pectinatus Stebbing (44) p. 629.

1906. . . Norman & Scott (36) p. 87, pl. ix.

figs. 1-3.

4 fasciatus Norman & Scott (36) p. 88, pl. v.

figs. 11 & 12.

1908. is dellavallei Norman (38) p. 310.

Bt pectinatus Norman (38) p. 310.

1909. x pilosus Walker (51) p. 341,

AMPHIPOD GENUS LEPTOCHEIRUS. 577

The specimens examined were :—

3 L. pectinatus, two females and one male, measuring 2°5—3 mm. ;

from Guernsey, sent by Canon Norman.

1 specimen, 9 , 4:°5 mm., taken by the ‘ Huxley’ in 109 fathoms,

during her cruise on the north side of the Bay of Biscay, 1906.

3 L. dellavallei, §, 5-6 mm., from the coast of Senegal, from

Monsieur Chevreux.

3 specimens from Breslau University Museum, referred to

above. These latter specimens were sent in two bottles with

Dr. Grube’s original labelsstill on them. One bottle, marked

““ Protomedeia pilosa Zadd. 9 . Luss. pice. Mundtheile. Grube,”

contained two tubes, with one specimen in each. One of these,

a female almost ready to moult, with a brood of young just

hatched, was dissected, and is evidently the specimen from

which Grube made his drawings. That this is so can be

proved by a comparison of the figures with the dissections:

e.g., the shape of the 2nd maxilla in his figure is due to the

fact that a portion of the inner plate with most of the

feathered bristles had been torn away (cf. (24) pl. x. fig. 2 m’,

with fig. 10); and again, in the Ist maxilla he notes the.

absence of the apical seta, which seta is, however, there, but

too completely masked by dirt to be seen, except under a high

power. The other tube contained a female 5°5 mm. long.

The second bottle was originally marked ‘“ Protomedeia

hirsutimana Sp. B. 2. Vollst. Luss. pice. Grube,” but over

the “hirsutimana ” is written in a different ink “ pilosa Zadd.”

and the words “‘m. Hiern ” added, apparently by Grube

himself. The tube in this bottle contains a large, brightly-

coloured female, the largest specimen I have yet seen.

The specimens form a most interesting developmental series 1n

the order in which I have arranged them above. Figures 14, 19,

& 23 are taken from a female pectinatus 2°75 mm., figs. 1 &

13 from a male 2°0 mm. long, Norman’s specimens: figs. 5, 16.

17, 25, & 28 are from the female specimen described by Grube:

the other drawings are from two males, dellavallei, 5°25 and

6 mm. respectively, Chevreux’s specimens. The pectinatus figures

are more magnified than the others for the purpose of comparison.

Description.

Head not quite so long as pereon-segments 1 and 2 taken

together ; lateral corners not prominent, truncate.

Eyes almost round, a little drawn out towards the lateral angle;

ommatidia few in number, large, with blackish-brown pigment

in the centre, outer row not so darkly pigmented.

Sideplates (figs. 13-16).—The first sideplate, which has been

the principal character for separating the two forms pectinatus

and dellavallei, is of exactly the same structure in all the

specimens. It is small, and completely hidden by sideplate 2.

578 MRS. E. W. SEXTON ON THE

(Through some mischance, Norman’s figure of the first gnathopod

((36) pl. ix. fig. 2) shows the second sideplate attached instead of the

first.) The first sideplate is subquadrate in the young animal, with

1 long sensory spine inset at the anterior angle, but with growth

this angle becomes more produced downwards, until in the largest

specimen of all the anterior margin of the sideplate is half as long

again as the hind margin. Four stages of development are

represented in the figures. The ‘Huxley’ specimen forms the

link between figs. 14 & 15; the anterior angle is not so much pro-

duced as in the latter, 6 setules are inset anteriorly, 1 inferiorly,

and 3 short sensory hairs at the posterior angle. In Grube’s

specimens, the anterior margin is lightly concave and the posterior

angle has a strong chitinous margin. The hinder portion of the

sideplate is firmly affixed to the basal joint beneath, so firmly in

fact as to make it impossible to separate them without destroying

the shape of the sideplate. A delicate laminar plate extends

beyond the sideplate behind, and is all but continuous with it

proximally, the line of demarcation being barely distinguishable.

Sideplate 2 is the largest of all, as deep as broad in the small

specimens, a little deeper than broad in the fully developed

‘ animal; front margin rounded; hind margin almost straight with

4 small sete inset; inferior margin rounded, thickly fringed

with delicate, sensory, cleft-tipped sete of varying lengths

(26 in the young, to about 41 in the full-grown). The remaining

sideplates are very like those of the type species. Sideplates 3

and 4 are subequal to each other in length, shorter. than the 2nd,

about half as deep again as broad. The 3rd is of equal width

throughout, front and hind margins straight, the latter with three

or four sete inset; inferior margin convex, fringed with 10-19 of

the sensory cleft sete. The 4th is a little wider proximally than

the 3rd, and the front margin is lightly convex; inferior margin

with fewer sensory sete, 5-15. In sideplate 5 the anterior lobe is

about as broad and as deep as the preceding sideplate; inferior

margin rounded, with only 4-7 sete; posterior lobe very small,

only one-third the depth of the anterior, 2 small sete inset.

Sideplate 6 small; posterior lobe about half as deep as the

anterior, with 1 small seta behind and 1 of the sensory

serrate spines similar to those found on the hinder perseopods

and uropods (see fig. 21); on the anterior lobe are 2 ciliated

hairs. Sideplate 7 small and subquadrangular.

Pleon._Segment 3 much the longest, as long as the 1st and 2nd

taken together; 4th segment a little depressed dorsally. The

hind margin of the 2nd is straight, that of the 3rd rounded;

inferior margins of 1-3 thickly beset with long plumose sete,

most numerous on the 2nd. On either side of the median line of

the 4th and 5th segments the posterior margin is produced in an

erect membranaceous lappet-like process, too delicate in structure

to be termed a tooth, each process having a setule inset in the

notch (fig. 26). Both Costa and Grube refer to these setules,

AMPHIPOD GENUS LEPTOCHEIRUS. 579

without apparently noticing the processes*, but this oversight is

very easily understood. The integument is so thin that if by any

chance, such as pressure or a little mucous dirt collected, the

process be flattened against the body, it is impossible to see it.

The 4th segment has a group of spines on either side, just above

the insertion of the uropods. The processes on the 5th segment

extend beyond the very small 6th segment.

ANTENNA. Superior Antenna (figs. 1—3).—1st joint of the peduncle

stout, shghtly longer than the 2nd in actual measurement, the

greater apparent length of the latter being due to its slenderness ;

in only one of the specimens, Grube’s largest, the 2nd joint

was a very little longer than the Ist; on the outer side are

several small ciliated hairs, and a cluster of sete with 1 long

ciliated hair at the distal angle; the inner angle carries a very

long, stout, outstanding sensory spine with 2 smaller ones inset

beside it. The 2nd joint is only half as broad as the Ist, with a

cluster of 3 or 4 sete and 1 long ciliated hair on either

distal angle. The 3rd joint in all the large specimens I have

examined barely reaches half the length of the 2nd; in the

smallest one it slightly exceeds half the length. The primary

flagellum is composed of 9-14 joints ; one young ovigerous female

2°5 mm. long had 9 joints; a male measuring 6 mm. had 14;

Grube’s large females 5°5 mm. and 6°25 mm. had respectively 12

and 10, the joints in the last-mentioned case being longer than in

the other animals examined. Each joint, from the 4th in the full-

grown and from the 2nd in the young, to the penultimate, is

provided with a very long sensory filament in addition to the

small sete. The accessory flagellum is usually 2-jointed, equalling

the Ist jot of the primary in length, and is so described by

Norman, Della Valle,and Walker. All the specimens examined

by me, except one, agree in having 2 joints only, though the

length varies a little, in the small specimen (fig. 1) being slightly

less than the Ist joint of the primary in length, and in the

medium-sized specimen (fig. 2) slightly more. The exception is

Grube’s largest specimen, which has 10 joints in the primary, and

a 3-jointed accessory flagellum equalling the first 2 joints of the

primary in length. Costa’s observation agrees with this, (19)

p- 155, “Il filetto composto di dieci articoli finamente pelacciuti ;

il filetto accessorio lungo appena quanto due articoli del primario.”

Grube in his description, (24) p. 403, says: ‘‘ Bei dem von mir

zuerst untersuchten Exemplar fehlte den oberen Antennen die

Nebengeissel..... An einem zweiten Exemplar fand ich die

Nebengeissel und zwar eine 3-gliederige.” The first specimen

examined by him (from Lussin-piccolo) had, however, an accessory

flagellum, a 2-jointed one, but this, in both antenne, had lost the

* “Quelli del quarto [abdominal segment] in oltre guerniti di piccole spine lungo

il margine dorsale.” Costa (19) p. 15.

“* Auf dem Riicken des 11ten und 12ten Segments vor dem Hinterrande sieht man

ein paar Borsten.” Grube (24) p. 407.

580 MRS. E. W. SEXTON ON THE

long apical setz, and, lying flat against the primary, was hidden

by a coating of dirt. In his third specimen both the superior

antennz are missing.

Inferior Antenna.—3drd joint of the peduncle short, as broad as

long, with a cluster of very long sete on the inner angle and 1

long sensory spine above. The 4th joint is twice the length of

the 3rd, slightly longer than the 5th, both beset with clusters of

the long, cleft-tipped setze and smaller setiform spines. Flagellum

shorter than the 5th joint, composed of 4 joints, the 1st much the

longest, almost equalling in length the two following taken

together, each carrying, in addition to the sete, a pair of strong

spines setting out on either side. In the small specimens the

flagellum is 3-jointed.

OrAL Parts.—The description is taken from the fully adult

specimens 5-6 mm. long, but the structure is precisely the same

in the small animals, the only difference being the lesser number

of spines and sete.

Upper Lip (fig. 4) thick, subquadrate in form, much arched

above; apex emarginate, slightly asymmetrical. In the figure the

lip is turned a little upwards to show the emargination of the

apex.

Lower Lip (fig. 5) as in the type species.

Mandibles (fig. 6).— Cutting-plate on the right mandible curved,

margin divided into 3 teeth, the lowest much produced; in

two specimens the margin was entire, the middle tooth not being

developed; accessory-plate narrow, in some specimens scarcely

wider than, and not as long as, the Ist spine of the spine row,

produced below to an acute tooth covered with microscopic

tubercles, and provided with a small tooth above. In the left

mandible the cutting-plate has 4 teeth, the second small, the

lowest the largest; the accessory-plate is strong, much broader

than that of the right mandible, the mandible divided into

3 teeth, the upper and lower of which are large, the middle one

small. There are 7 spines in the spine-row in Chevreux’s

specimens, 8 in Grube’s largest, the first 3 being unusually

large, wide at the base, flat and furry in appearance owing to

being covered with microscopic spinules. The molar is very

prominent; the crown transversely ridged with rows of teeth,

edged with flat spinules, with 1 long, feathered seta above.

The palp is very large; 3rd joint the longest, tipped with long

serrate bristles, with a double row of smaller ones extending

down the distal half of the inner margin; the outer margin has

4 groups of long serrate bristles inset.

Mazxilla | (figs. 7-9).—Jnner plate large, with 1 long plumose

seta inset; Grube notes the absence of this seta (p. 404),

but his specimen is so covered with mucous dirt as to render it

difficult to see details clearly ; not only is the seta present, but the

new one can be seen under the old loose skin, the animal, as

before stated, being ready to moult. Outer plate curved, with

11 strong spines, 4 of which are bifurcate (fig. 8), the rest

AMPHIPOD GENUS LEPTOCHEIRUS. 581

finely dentate (fig. 9); the outer margin is covered with fine

hairs. The 2nd joint of the palp is expanded apically, and

rounded, with a marginal row of 4 strong flat spines, and 3

plumose sete; 2 or 3 diagonal rows of set are inset sub-

marginally.

Maxilla 2 (fig., 10).—Outer plate the larger, covered with

exceedingly fine long hairs; expanded distally, bulging behind,

furnished with an apical cluster of stiff bristles, and a row of

sparsely plumose sete extending some distance down the inner

margin. The inner plate, distally narrowed, is provided with 2

rows of setz; one row, submarginal, consists of 20 long, delicate,

finely plumose set ; the second row contains an apical cluster of

stiff setae, with 8 inset along the margin, these latter of the same

structure as the plumose ones of the outer plate. In Grube’s

specimen the upper portion of the inner plate has been torn

away; the 2 apical setz represented in the figure 2 m‘ are two

of the plumose row of the outer plate.

Mawillipeds (figs. 11 & 12).—IJnner and outer plates well de-

veloped, but narrow. The apices of the inner plates are inset

with 4 long, feathered, setiform spines along the margin, with

a group at the inner angle of 4 long delicate plumose set

set together in a little hollow, 3 similar ones extending down

the inner margin, and 2 submarginally on the outer side. The

outer surface is longitudinally ridged. On this surface close to,

but just below, the inner angle, is a small cowpling-spine (fig. 12),

stout, and bent upwards. It appears to be tuberculated on its

inner side, but the detail is obscure, even with a ;4,th oil-immersion

lens. The ower plate carries a marginal row of flat feathered

spines (15 in Grube’s large specimen, 13 in Chevreux’s, 7 in the

smallest of all), the 2 apical ones much the longest, and sub-

marginally on the outer side a few long feathered set. The

2nd joint of the palp is the longest, in Grube’s specimen twice

the length of the Ist, furnished with numerous plumose sete ;

the 3rd is produced over the insertion of the finger as in the type

species; finger small, obtuse, carrying apically 1 long stout

dentate spine, 1 smaller one, and 2 sete in Chevreux’s specimen,

3 stout spines and 4 ‘sete in Grube’s, 1 spine and 2 sete in

Norman’s.

First Gnathopods (figs. 18, 15, 17, & 18).—2nd joint narrow at

the base, but widening distally; posterior margin convex, with

2-4 very long delicate setz inset midway; anterior margin

straight, carrying a row of delicate plumose setze, another row of

longer similar setz is found on the under surface as in the type

species ; 3rd joint bulging behind, fringed with numerous closely

set long feathered sete; 4th joint the smallest, also fringed

posteriorly with feathered sete shorter than those of the 3rd.

The 5th is considerably longer than the 6th, with 5-7 transverse

rows of plumose sete, and 5-7 clusters of rigid serrate bristles

along the hind margin. 6th oblong, about twice as long as broad,

with 4—5 transverse rows of sete onthe hind margin ; palm short,

582 MRS. E. W. SEXTON ON THE

truncate. The long spine on the hind margin in Chevreux’s

figure, (16) pl. xi. ig. 2a, is the longest of a group of 3 inset

on the under side of the hand, and projecting beyond the hind

margin. The palmar margin is mentioned by Chevreux (p. 91)

as affording a distinguishing character for the two forms—concave

for pectinatus, and convex for dellavallei,—but an examination of

a series of specimens shows conclusively that this difference is due

to sex. In the male (figs. 15 & 18) the margin is concave, and

the palmar angle forms a right angle with the hind margin;

in the female (fig. 17) the palmar angle is rounded, the

curve commencing at the insertion of the finger, and merging

imperceptibly into the hind margin. The palm is strongly

serrated in all; in the female the serrations turn the corner and

reach as far as the large spine Just referred to. The palmar spines

ave of the same structure as those of the type species. The finger

is more than twice the length of the palm (a little longer in pro-

portion in the young form), curved, with a strong auxiliary claw ;

a long, stiff, finely serrated spine and 1 setule are inset in the

notch, 1-2 setules behind the claw; the rest of the inner margin

is firmly serrated.

Second Gnathopod (figs. 19 & 20) as described by Grube,

Norman, and Della Valle. Figures of the finger, so characteristic

of the species, are given from Norman’s and Chevreux’s specimens

in order to show the identity of the two forms pectinatus and

dellavallet. Grube’s specimens agree exactly with these. In the

young form the finger is slightly longer in proportion to the pre-

ceding joint than in the older animals, not equalling quite half

its length in Norman’s specimens, and only about one-third the

length in Grube’s largest.

Pereopods | and 2 glandular, alike in structure ; 2nd joint large,

fringed posteriorly with long delicate sete, more numerous on the

2nd pereopod; anterior margins with from 4-6 similar sete:

Ath joint about half the length of the 2nd, slightly expanded

distally ; 5th joint narrower and shorter than 4th; 6th joint

slightly longer than the 4th; the two terminal joints tapering

gradually to the acute tip of the 7th; the 7th subequal to the 6th

in length, with the glandular aperture opening at the tip.

Hinder Pereopods (fig. 21) more strongly built than the pre-

ceding; rapidly increasing in length. Pereopod 3: 2nd joint

large, a little longer than broad, equalling in length the 3 fol-

lowing joints taken together; both margins convex, the anterior

beset at intervals with 7 small sensory spines, the first 3 un-

accompanied by sete, the 4th with 1 seta, the 5th with 2, 6th

with 3, and the distal one with 4; posterior margin produced

below in a rounded lobe, crenulate, 6 of the crenulations with

setules inset, the 2 crenulations on the lobe each with a ciliated

hair; 3rd joint short and broad, with a cluster of sete, 1 ciliated

hair, and 1 spine; 4th joint half the length of the 2nd, stout,

slightly expanded anteriorly, with the spines of the structure

peculiar to the hinder perzeopods and uropods (fig. 21); 5th joint

AMPHIPOD GENUS LEPTOCHEIRUS. 583

shorter than 4th, with two clusters of spines on either side;

6th about as long as the 4th, but much more slender, spinose,

with groups of spines on either side and | very long stout spine

inset behind the finger. Finger short, not half the length of the

preceding joint, much curved, with 2 specialised bristles towards

the apex, one on the inner side lying flat against the finger and

reaching to the apex, and the other on the outer side, setting out

at right angles to the finger. This one appears to have a flattened

tip, and the inner one to be feathered, but the detail is almost

impossible to see owing to their extreme tenuity and the angle

at which they are placed. Perzeopod 4 is noticeable for the great

length of the spines of the 5th and 6th joints; 2nd joint rounded

oval, produced behind into a rounded lobe, 8 spines along the

anterior margin, the distal 5 accompanied by increasing clusters

of ciliated hairs and sete; posterior margin with 10 crenulations,

and a submarginal row around the lobe of 5-8 ciliated hairs; 4th

and 5th joints practically subequal to each other in length, 5th

narrower than 4th; 6th considerably longer and more slender ;

the spines on this joint are longer than those on the 5th, 4 clusters

increasing in length and number, on either side, 2 of the clusters

behind the finger equalling the joint itself in length; they are

very brittle, several of the specimens not having a single perfect

one remaining. Finger lightly curved, of the same structure as

the finger of the preceding pereopod. Pereopod 5: 2nd joint

narrow proximally, widened distally ; the proximal end of the

posterior expansion is produced subacutely, the distal end, as in

pereopods 3 and 4, forms a rounded lobe; on the posterior

margin are 8 crenulations with a setule on each, and 20 ciliated

hairs in a submarginal row; the anterior margin has 8 spines as

in pereopod 4, and 3 long fine hairs proximally. The 4th—6th

joints rapidly increase in length and decrease in width ; 4th and

5th each beset with 3 groups of stout spines on either side;

6th twice the length of the 4th, with 5 groups of spines on

either side, the terminal posterior group of spines and setz of

extraordinary length. Finger as in pereeopod 4.

The smaller specimens agree in all details with this description,

the only difference being the lesser number of spines.

Pleopod 1 (fig. 22): peduncle short, hardly half the length of the

outer ramus, with about 10 long plumose setz on the outer side ; 2

small coupling-spines, and | long plumose seta inset together on the

inner. The outer ramus 12-jointed, considerably shorter than the

inner ; inner ramus 14-jointed, with 4 cleft spines. Pleopods 2

and 3 alike; peduncle more than half the length of the outer

ramus, with only 2 or 3 fine hairs. The rami are shorter than

in the lst pleopod ; outer ramus with 11 joints; inner with 13 ;

4 cleft spines in the 2nd pleopod ; 2 in the 3rd.

Uropods (figs. 23-26) extend backwards to the same level.

The peduncle of uropod 1 is shorter than the outer ramus, with

5 slender spines on the upper curve, and 3 inset diagonally midway

on the outer surface; the falciform apical process reaches to half

584 MRS. E. W. SEX'TON ON THE

the length of the outer ramus. The outer ramus is shorter than

the inner, both furnished with slender spines. In uropod 2 the

peduncle is almost as long as the outer ramus; outer ramus

the shorter; the spines are shorter and stouter than those of

uropod 1. I have figured uropod 3 in detail (figs. 23, 24, &

25), because it has been used by Norman (36, p. 89) as dis-

tinguishing the two forms. A comparison of the figures will

show the identity of structure. The outer ramus has a minute

apical joint carrying a long rigid spine, this joint being larger in

proportion in the young form; the detail of the spines in the

small specimen (fig. 23) could only be seen with the jth oil-

immersion.

The Telson (figs. 27 & 28) is of exactly the same structure in

all the specimens; lateral angles greatly elevated, each produced

to an acute point, each with 1 long spine and 1 short sensory

ciliated hair on the margin, and 2 ciliated hairs underneath. The

subacute apex curves upwards also, but not to the same degree

as the lateral angles. igure 27 is taken from Chevreux’s speci-

men ; in two of Grube’s, the tip was as represented in fig. 28, in

the third, the largest, it was as in fig. 27.

Colour.—Grube’s largest specimen still retains its vivid colour,

a yellowish tint, with markings in dark brown. The dis-

tribution of the pigment agrees perfectly with Costa’s figure (19)

pl. ui. fig. 8. The markings are composed of cloudy brownish

patches stippled thickly but irregularly with blackish-brown spots.

The dorsal part of the head is‘covered with dark brown pigment,

shading into black near the eyes, these dark patches causing

the eyes to appear larger than they really are. The lst perzon-

segment is plain; the 2nd has only faint, patches, one on the back,

and one on each side; the 3rd and 4th with their corresponding

sideplates, and the 5th are thickly covered with the pigment,

arranged in a band along the middle of the segment; the 6th is

plain ; the 7th and the first 3 pleon-segments with their epimera

are darkly pigmented ; the 4th pleon-segment has a faint patch

on each side; the rest of the animal is plain. The general effect

is very striking, the animal showing a dark head, 3 dark rings

anteriorly, and 4 farther back. The amount of colour probably

varies with the locality. | Walker, (48) p. 310, describes it as

‘“deep yellow, with transverse brown lines on all the segments.”

Distribution.—This species has a very wide distribution :—

SHETLAND Istus : Norman (84) p. 283, as Protomedeia pectinata ;

dredged; St. Magnus Bay, deep water.

Trish Sea: Walker (48) p. 310, as Leptocheirus pilosus ;

dredged; 17 fathoms.

W.&S. Coasts or IneLanp: Walker (50) p. 169, as Leptocheirus

pilosus; 40 fathoms ; bottom deposit, gravel.

CHANNEL Istanps: Walker & Hornell (49) p. 54; as Lepto-

cheirus pilosus Zadd.=L. pectinatus Norman; 7 fathoms;

bottom deposit, gravel, clinkers, and shells. Chevreux (16)

AMPHIPOD GENUS LEPTOCHEIRUS. 585

p. 91, as Leptocheirus pilosus ; tidal zone; bottom deposit,

rather coarse sand. Norman (37) p. 369, as Leptocheirus

pectinatus.

France: Ocranic Coasr: Chevreux (11) pp. 290, 309, as

Ptilocheirus pectinatus ; 10-19 m. ; bottom deposit, nullipores

and mud, Chevreux (15) p. 482, as Leptocheirus pilosus ;

10-20 m.

Bay or Biscay: ss. ‘ Huxley,’ 47° 48’ N.; 7° 46° W.; 109

fathoms.

MEDITERRANEAN : Chevreux (16) p. 91,as Leptocheirus fasciatus.

PROVENCE : coast of ALGERIA: Corsica, dredged in 12 fathoms:

Chevreux (17) p. 4, as Leptocheirus fasciatus.

Bay or Napues: Costa (19) p. 155, as Protomedeia fasciata.

Della Valle (21) p. 450, as Leptocheirus pilosus ; 10-20 m. ;

bottom deposit, sand.

Aprtatic : Grube (22) p. 73, as Protomedeia pilosa. Grube (24)

p. 403, as Protomedeia hirsutimana® and Pr. pilosa.

SENEGAL: Chevreux (in litt.).

Wasin, Brir. KE. Arrica: Walker (51) p. 341, as Leptocheirus

pilosus; 10 fathoms ; bottom deposit, mud.

LEPTOCHEIRUS BISPINOsUS Norman. (Plate XVIII. figs. 17-20.

Text-fig. 146.)

1866. Protomedeia hirsutimana Heller (25) pp. 34 & 35.

1893. Leptocheirus guttatus Della Valle (21) p. 430, pl. xii.

figs. 15-24.

1908. Leptocherrus bispinosus Norman (38) p. 308, pl. xii.

figs. 7-9; pl. xii. figs. 1-3.

The specimen described by Heller as Protomedeia hirsutimana

Sp. Bate is preserved in the Hofmuseum, Vienna. The de-

scription and figures of it, most kindly sent to me by Dr. Pesta,

prove it to belong to the same species as described by Norman

in 1908 under the name of ZL. bispinosus. -

The guttatus of Della Valle, in my opinion, must also be in-

cluded in this species. A comparison of the two accounts, Della

Valle’s and Norman’s, leaves no room for doubt. The only point

of difference is in the number of joints in the accessory flagellum :

Della Valle gives the number as 2, subequal in length, and as long,

taken together, as 2 joints of the primary flagellum; Norman

as 5, as long as 3 joints of the primary. It seems probable, aither

that Della Valle had a young specimen before him with only two

joints developed, or, what I think more likely, that the accessory

flagellum was broken. In the other specimens of this species (and,

indeed, in all the specimens of this genus that I have examined)

the terminal joint of the accessory flagellum is very small and

tipped with long sete, but Della. Valle says definitely that the 2nd

joint in his specimen was equal in length to the lst, and in his

figure he shows only 2 small sete instead of the usual long apical

cluster, I have added below some details to the description given

586 MRS. E. W. SEXTON ON THE

Text-fig. 146.

Leptocheirus bispinosus Norman.

The figures were drawn by Dr. Otto Pesta from Heller’s specimen

in the Hofmuseum, Vienna.

Fig. a. Superior antenna, accessory flagellum, left side, X 100.

Fic. 6. ee é 53 * right side, X 100.

Fig. ec. First pereopod, X 60. Fig. d. Hand of first enathopod, X 100.

Fig. e. Finger of second gnathopod, X 200. Fig. f Sideplates 2 and 3, x 60.

Fig. g. Pleon-segments 3-6, X 60.

AMPHIPOD GENUS LEPTOCHEIRUS. 587

by Canon Norman for the sake of a more complete comparison

with Della Valle’s account.

The species resembles guttatus Grube in many respects, the

principal distinguishing points

antennz ; the hand of gnathopod 1 ; sideplate 2 ; the shape of the

basal joints of the hinder peropods ; the armature of the pleon ;

and uropod 3

Description (taken from the type specimen of bispinosus, a

female, which Canon Norman kindly allowed me to examine) :—

Head \onger than pereon-segments 1 and 2 taken together ;

lateral corners rounded.

Hyes very large; pigment brownish black; ommatidia large.

Sideplate 1 free of sideplate 2, about half its width, but not

equalling it in length, a little expanded distally and fringed with

long setze. Sideplates 2-5 much as in the type species for shape.

The 2nd is the largest and deepest, much deeper than the corre-

sponding body-segment; expanded inferiorly, hind margin

straight, inferior margin rounded and fringed with numerous

plumose sete (cf. text-fig. 146, f, with Norman’s pl. xiii. fig. 2

and Della Valle’s pl. xii. fig. 17). The 3rd is deeper than the

4th ; and the 4th a little deeper than the 5th; all fringed inferiorly

with sete.

Pleon.—Segment 3, hind margin produced, rounded, crenulate ;

crenulations 8 in number, each with a setule; inferior margin

concave, furnished with numerous strong sensory spines sub-

marginally. Pleon-segment 4 as described by Norman, with ‘a

strong and acute angular backward projection on each side” ;

Dr. Pesta’s account agrees with this, “das 4. Pleonsegment hat 2

seitliche nach aufwiirts geschwungene ‘ angles’” (text-fig. 146, g) ;

Heller erroneously notes them as occurring on the 5th. The 5th,

indeed, carries 2 dorso-lateral angles (as stated by Della Valle),

but these are exceedingly small and difficult to see in situ, not

much produced, acute, each with a seta inset behind.

ANTENN (text- figure 146 a, b).—Superior Antenna, as figured

by Della Valle: 1st joint of peduncle shorter and broader than

the 2nd, carrying a stout spine on the inner distal angle; 3rd a

little more than one-third the length of the 2nd; primary

flagellum with 16 joints, each, from the 3rd, with a small sensory

filament ; Dr. Pesta says of Heller’ s specimen that the number of

joints on the right side is 17, on the left 18. The accessory

flagellum in Norman’s specimen extends to the 3rd joint of the

primary, 5-jointed, the first 4 joints subequal to each other in

length, the terminal one minute, tipped with long setze ; in Heller’s

specimen it does not reach to the distal margin of the 3rd joint of

the primary, ““4-gliedrig, jedoch ist das Endglied rechts kiirzere

als links.”

Inferior Antenna: Ath joint longer than 5th; flagellum about

the same length as the 5th, com posed of 7 joints, the first as long

as all the others taken together.

First Gnathopod (PI. XVIII. fig. 17): 3rd and 4th joints more

densely setose than in the other species, the plumose sete being

588 MRS. E. W. SEXTON ON THE

arranged in transverse rows in addition to the marginal fringe.

5th joint slightly longer than 6th; 6th twice as long as broad,

widening a little towards the palm. The palmar margin is convex,

its limit defined, as in the preceding species, by a long sensory

spine inset on the under surface; on the outer surface it carries

a submarginal row of about 10 strong sensory spines, and on the

inner side a thick row of setiform spines. The finger when closed

reaches to the large spine; it is curved, with a strong auxiliary

tooth subapically, and 6 smaller teeth on the inner margin, each

tooth with a setule beside it. In the notch near the apex 2 or 3

longer setules are inset.

Dr. Pesta’s description is as follows (text-fig. 146, d): “das

Handglied des 1. Gnathopoden ist vorne etwas verbreitert, die

Klaue so lang wie die konvexe Seite desselben ; der konvexe Rand

die ‘ Hand’ besitzt eine Reihe von kriiftigen Dornen, die gegen

die Klaue gerichtet sind.”

Second Gnathopod (fig. 18): 2nd jomt unusually long, equalling

in length the joints 3-6 taken together; 3rd, 4th, and 5th joints

subequal to each other measured along the inner side. The finger

is nearly straight, about half the length of the preceding joint ;

“leicht gebogen und spitz” (text-fig. 146, e).

Perceopods 1 and 2.—The length of the 4th joint forms a dis-

tinguishing character for this species. The peropods are about

the same length, but in perzopod | the basal joint is longer than

- in perzopod 2; and the 4th joint is longer in the latter than in the

former, being equal to the basal joint in length. Dr. Pesta says

“das 4 Glied des 1. Peraeopoden ist linger als das 6 Glied und fast

so lang wie das 2 Glied.” (Cf. text-fig. 146, ¢, with Norman’s pl. xii.

fig. 8 and Della Valle’s pl. xii. fig. 24.) The 2nd joint carries

posteriorly numerous delicate sensory set, some of great length ;

4th joint long, of equal breadth throughout; 5th about half as

long as the 4th; 6th longer than 5th, both with clusters of long

sparsely feathered sete posteriorly ; finger shorter than the 6th,

subequal to the 5th in length, with the glandular aperture

opening at the tip.

Hinder Perceopods (fig. 19) as figured by Della Valle. In perzeo-

pod 3 the basal joint has the posterior margin straight and

crenulate ; in pereopod 5 the posterior expansion of this joint is

much wider distally than at the base, and the margin is rounded

and serrate, the lower portion carrying numerous plumose sete.

Fingers curved and falciform. In Heller’s specimen, perzeopod 3

is missing, but Dr. Pesta says of the last two pairs “die End-

klauen sind einfach (nicht ‘ bifid’),” as in Spence Bate’s hirsuti-

mana. In Norman's figure, pl. xii. fig. 9, the terminal joints

are lying in such a position as to quite conceal thespines. I have

therefore refigured them to show the armature.

Uropods 1 and 2 (text-fig. 146, g) much as in gutiatus Grube ;

very spinose, the spines of uropod 2 shorter and much stouter than

those of uropod 1. The falciform process of the peduncle reaches,

in uropod 1, to the tip of the outer ramus; in uropod 2 beyond

the tip. The inner rami differ from the other species in having

AMPHIPOD GENUS LEPTOCHEIRUS. 589

slender, feathered set, or setiform spines, in addition to the other

spines. These are found on the under margin, 3 in uropod 1,

short, inset at intervals, and 2 clusters of long ones in uropod 2,

near the apex. In uropod 3 (fig. 20) the outer ramus is fur-

nished with an apical cluster of 5 spinesand 3 or 4 long feathered

setee similar to those of the 2nd uropod, and 1 cluster of small

spines midway, in Norman’s specimen (2 clusters figured by Della

Valle, probably from an older specimen) ; the inner ramus is tipped

with 1 stout spine and | seta, with 1 spine midway, in Norman’s

specimen (2 in Della Valle’s).

Telson.—Thé apical margin of the telson is represented in Della

Valle’s figure as truncate, but, as in guitatus Grube, this appear-

ance is due to the angle at which it is carried. The mar gin 1s

really convex, but as in the natural position it is a little bent i in

under, the convexity is not apparent from the dorsal view.

Colour given by Della Valle as greyish yellow without markings ;

by Dr. Pesta as “ gelbbraun, Auge schwarz” ; but, he adds, “ diese

Farbung diirfte nicht natiirlich sein.”

Distribution :—

Bay or Biscay: Norman (38) p. 309, as ZL. bispinosus,

depth 35-60 fathoms.

Narues: Della Valle (21) p. 482, as L. guitatus ; bottom deposit,

corallines.

Apriatic at Lesina: Heller (25) p. 35, as Protomedeia

hirsutimana.

Gutr or Bone and Coast of Tunis: Chevreux (18) p. 3, as

L. bispinosus.

The only details given by Catta concerning his species massili-

ensis, viz. the nature of the ground on which it was captured

“fonds coralligénes,” and the colour, “ un beau jaune, tandis-

que l’ceil était completement noir,” would seem to point to

L. bispinosus.

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. 561.

(47) Ween A. O.—The Amphipoda of Bate and Westwood’s

‘ British Sessile-eyed Crustacea.” Ann. & Mag. Nat. Hist.

(6) vol. xv. 1895, pp. 469-470.

(48) WALKER, A. O.—Revision of the Amphipoda of the L.M.B.C.

District. Trans. Liverpool Biological Society, vol. ix. 1895,

pp. 310, 311.

(49) WaLker & Horneti.—Report on the Schizopoda, Cumacea,

Tsopoda, and Amphipoda of the Channel Islands. Journ.

Marine Zoology and Microscopy, vol. ii. no. 7, 1896,

. 54,

(50) ae A. O.—Malacostraca from the West Coast of

Treland. Trans. Liverp. Biol. Soc. vol. xu. 1898, p. 169.

(50a) WALKER, A. O.—Malacostracan Fauna of the Mediter-

ranean. Journ. Linn. Soc. vol. xxviii. no. 12, 1901,

p. 305.

(51) Waker, A. O.—Amphipoda Gammaridea from the Indian

Ocean, Brit. East Africa, and the Red Sea. Trans. Linn,

Sce. vol. xii. pt. 4, 2nd series, Zool. 1909, p. 341.

AMPHIPOD GENUS LEPTOCHEIRUS. 593

(52) Waker, A. O.—Notes on Amphipoda. Ann. & Mag. Nat.

Hist. (8) vol. vi. 1910, p. 33.

(53) Zappacu, E. G.—Synopseos Crustacecrum Prussicorum

prodromus. Kéonigsberg, 1844, pp. 8, 9.

(54) Zappacu, E. G.—Die Meeres-Fauna an der preussischen

Kuste. Erste Abth. Schrift. d. Phys.-dkon. Ges. z. Konigs-

berg, 19. Jahrg. 1878, pp. 18, 19.

EXPLANATION OF THE PLATES.

PuatE XVII.

Leptocheirus pilosus Zaddach.

Fig. 1. Whole animal, Zaddach’s specimen, young SLR eats SEO aE Ne EAD

2. Superior antenna, seen trom above, young 2. Zaddach’s specimen. X ASO.

8. Part of inferior antenna............... eves i 3 X 50.

AM Wippersliplscrnsaesctexcgeee meee. basal ss ¥3 53 x 95.

Belhowertipissn5-eu eeu emer rcre. mane ces ue * XK 98.

Gap Man dibleped saat res eae Wee ae ene ee le 5 s x 95.

Teale xall aoe carte cara aie cence Mean Fess PH “i x 95.

Sat Texall Ate cea eet seacecasta coe cate ete i “ x 95.

Oro Mila xallamed sisunchescasedceteomemadteeeveri las ta. 95 Pa x 95.

10. First gnathopod .. SASS os 35 Xx 50.

11. Hand of the tirst enathopod. 9 . Norman’s POOVNIN coconsassconcen >< (0)

12. Gin (Chevréuxisy 8 os.) 4 ean ae x 50.

13. Finger of first enathopod. Large 2 3 5 x 95.

14, X 95.

3 ; 5 apni secaeces

15. Serrate bristle from 5th joint, first gnathopod. Large 3.

Chevreux’s spec x 175.

16. Second gnathopod. Young ie . Zaddach’s specimen ..... eae Oxo!

17. Finger of second gnathopod. Young 2. Zaddach’s specimen anes DLO!

18. 5 = PA 9. Norman’s Pen aces x 140.

19. First pereopod. Young 2. Zaddach’s AYNSOUNEN “UGescusosootace: éoacco, 4 GAO):

20. Second - ieee a "3 Rr om ieereatinae or <5) 0)5

21. Third Bn saedess 33 Say i uy, ates atrckoperynatna carseat x 50.

22. Fitth ae 55 55 adsisasdaacka eee 2 DADO!

23. Fifth pereeopod, under surtace. Large g. Chevreux’s specimen... X 40.

24. Upper coupling-spine, first pleopod. Young 9. Zaddach’sspecimen. X 290.

25. Telson and third uropods, dorsal view. ,,_,, 53 x 60.

26. Telson, lateral view. Young 9. Zaddach’s specimen . ise kVOO!

Poate XVIII.

Fig.1. Whole animal. Grube’s type specimen ... Leptocheirus guttatus. X 20.

2. Accessor y flagellum, right antenna. Nor-

man’s specimen .... atieeh. on 2 x 76.

3. Accessory flagellum, left antenna. Same

specimen ..... > x 75.

4, Accessory flagellum, right antenna. Another

SPECIMEN On erence neon crtie miea Nene = ss x 78.

5. Hand and finger, “first sgnathopod. Chev-

reux’s specimen, x 7bs

6. Finger of second enathopod. “ Chevrenx’s

specimen,? ... 3 3 x<a7ios

7. One of the smaller apical spines, uropod 2.

Norman’s specimen .... 55 5 X 265.

8. Third uropod, drawn in “situ. Grube’s

speciinen ... j PS x 76.

9. Apical margin of telson, ; a little upturned.

Grubesispecimentimyper ese meena cern 5 ¥ x 75.

594 ON THE AMPHIPOD GENUS LEPLOCHEIRUS.

Fig.10. Hand of first gnathopod, ¢, looking down

on the hind margin, from a ae from

Vineyard Sound . : sab . Leptocheirus pinguis. X 22.

11. Finger of second onatbopod, s same - specimen. a5 4 * 27.

12. Second joint, without the bristles, uropod 3,

QD, Wimesyemrdl SOWIE soo scocaseseosnosbe sn caneae ; % x 75.

13}, Finger of first gnathopod, small specimen,

4 mm., from Banff ........ : ....... Leptocheirus hirsutimanus X 58.

14. Tip of finger, second enathopod, small speci-

men, 4 1mm., from Banft.. Wenn:

15. Finger of third per: eopod, arge specimen,

10 mm., Shetland Isles, 1867 .................. 3 ep x 146.

16. Outer ramus, third uropod, small specimen,

Avmntn.; IB anittik tee eee Leen eee

17. Hand and finger, first pais aaa oF

Norman's type : specimen : Me

18. Finger of second enathopod, @ ‘Norman’s

type specimen ...... x

19. Terminal joints, fifth “peraopod, o, ~Nor-

man’s type specimen ........ : n 55 x 28.

20. Uropod 3, 9. Norman’s ty pe ‘specimen — x

» » x Mo.

a op < 58.

. Leptocheirus bispinosus. X 50,

be 99

32 »

PratEt XIX.

Leptocheirus pectinatus Norman.

Fig 1. Accessory flagellum, superior antenna. Norman’s specimen, 6 ...... x 1465.

2. i 3 33 Fs Chevreux’s specimen, large @. X 75.

3. ut * Py is Grube’s largest specimen, 2. X 75.

fe Wipper lip) eeeeaee tenes oe eeeen ee eeee a Oh evreuxes) specimens lansiend) sn Guor

SH ILOW EL: Niple cereale: ees eee ee Renee 5 op Se, Oe:

6. Mandible 35 3 hehe ate 1S 78.

7. Manilla 1 : 3 ‘ re eS, ae

. Biturcate spine, outer plate, maxilla 1. es aS as son

9. Dentate spine, 58 ” » 2 ) oy) ” x 435.

10), Wlesalllle Fo. cs0cs0 0: a 3 RT SES

11. Mawillipeds ........ HA be SAT Paso OM 00s

12. Coupling-spine, mner : plate, maxillipeds. 5 » x 430.

13. First gnathopod (the feathering of the setze on joints Be 6 not ‘show n).

Norman’s specimen, young ¢,2°5mm. X 9d.

14. First sideplate............. “5 young 9,275 mm. X 95.

15. First enathopod. “Chevreux’s specimen, large 6, 5:25 mm............. x 40.

16 Hirst sideplate. Giube’s specimen, large 9) ...)... x 40.

17. Finger and palm, first gnathopod. Grube’s specimen, large 9 ...... x 95.

(The other sete omitted in order to show the palmar spmes clearly. )

18. Finger and palm, first gnathopod. Chevreux’'s specimen, large g... X 95.

19. Finger of second gnathopod. Norman’s specimen, 2 .................. X 290.

20. ‘ 4 Chevreux’s specimen, 6... Agencies. WOM:

21. Sensory spine, 6th jomt, perseopod 4. ,, es atencdactocassaidanae | A ek)

22. Tip of 4th cleft spine, pleopod 1. as a Rt ied” oh Sle aa al X 300.

23. Third uropod. Norman’s specimen, young 2, 2° 75 mm. MY, BRO EE OC ZO0F

24. iy ap Chevreux’s specimen, large Sq DED WAV, ooo soncoooscnes x 95.

25. _ Grube’s specimen, large Q acae botenehmans | Oe

26. U ropods and telson. Chevreux’s specimen, large Be “6mm... x 40.

27. Apex of telson. an 3 A 35 SR x 58.

28. 55 5 Grulbeisispecimen. Oe volo amms ss eeee eee Doe

PAS, Leah IPL, 2OX,

anes eT Ra aca czecs

nae eae, AOL ae y ee a a sca

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ye er mea nt

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J S

ae a oa

penn Sarre etccereati = w

SS a

“S, Sr Fa

West, Newman lith.

OSTRACODA FROM MADEIRA.

PAS. isi, Ill, SOC.

ee a nett

oe eee

rattan cemnner RR oO sh

ties

eR meron te NAN A EE I

een

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spe EERIE RANE A PIP OEIC

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+: & 2 s

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ng, a 2 &

te ae a °

a a Ber -, Sy S o

Sn = iene ‘

Mtr rrccenseg apis

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West, Newman lith.

OSTRACODA EPROM MADEIRA:

fetidion.

bean

“On oie D

cag

EZ 8. eal, Jed, SOOT.

oer

seen PROTON sa

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eonnagneci

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West, Newman lith.

OSTRACODA FROM MADEIRA.

ON MARINE OSTRACODA FROM MADEIRA. 595:

27. Notes on Marine Ostracoda from Madeira. By G.

Srmwarpson Brapy, M.D., LL.D: D.Se., F.B.S.,

C.M.Z.S.

fReceived January 13, 1911: Read March 21, 1911.!

(Plates XX.-XXIL*)

T am indebted for the specimens which form the subject of the

following notes to my friend the Rev. Canon Norman, F.R.S.,

by whom they were collected in the spring of the year 1897.

The mountings submitted to me consisted altogether of dried

shells, and in no case was the contained animal preserved,

excepting in a few of those collected between tide-marks,

and only in some of these littoral forms was the original colouring

discernible. Apart from such as appear to be new to science,

the collection is interesting as extending the known range of

several species from the European and North Atlantic areas much

further southward, though not quite into the tropical zone. — All

the species belong, so far as can be ascertained from the shell-

characters, to well-known genera. The following lists show

separately the species found in the littoral zone and in deep

water.

Between tide-marks.

}

Cythere convexa Baird. Cytherura maculosa, sp. n.

eS albo-maculata Baird. 55 cellulosa Norman.

Xestoleberis depressa G. O. Sars. | Sclerochilus levis? G. W. Miller.

5 nigromaculata, sp. n. (Pl. XXII. fig. 10.)

ss margaritea Brady. | Paradoxostoma hibernicum Brady.

Loxoconcha impressa Baird. 59 arcuatum Brady.

Dredged in 30-70 fathoms.

Bythocypris reniformis Brady. | Loxoconcha obesa, sp. n.

Macrocypris decora Brady. | D decipiens G. W. Miller.

Pontocypris succinea G. W. Miller. sf impressa Baird.

35 dispar? G. W. Muller. | be? subalata, sp. 0.

Bairdia amygdaloides Brady. Xestoleberis latissima, sp. n.

» obtusata G. O. Sars. | Cytherura striata G. O. Sars.

» dubia, sp. n. | Bs cribrosa, sp. n.

> mediterranea G. W. Miller. | 5 eribriformis G. W. Miller.

» acanthigera Brady. | a fossulata, sp. n. |

Argillcecia atfinis, sp. n. Cytherideis subulata, var. crenulata

Cythere crispata Brady. Brady.

» cingulata, sp. n. Sclerochilus contortus Norman.

» tuberculata G. O. Sars. As levis? G. W. Miiller.

» emaciata Brady. (Pl. XXII. fig. 10.)

Cythereis runcinata Baird. Paradoxostoma gracile, sp. n.

» deformis, sp. n. fi flexuosum Brady.

» antiquata Baird. 4 cylindricum G. W.

>» jones Baird. Miller.

Cytheridea elongata Brady. Sarsiella capsula Norman.

Eucythere prava Brady & Robertson. Cytherella ? ovalis, sp. n.

* For explanation of the Plates see p. 601.

396 DR. G. S. BRADY ON

BairDIA DUBIA, sp. n. (Plate XX. figs. 1, 2.)

The outline of the shell as seen laterally is rhomboidal (fig. 1),

the greatest height situated near the middle and equal to half

the length; anterior extremity obliquely subtruncate, posterior

produced below the middle into a wide, obtuse beak; dorsal

margin forming a flattened arch, inferior nearly straight, gently

sinuated in the middle: seen from above (fig. 2) oblong, ovate,

twice as long as broad, greatest width in the middle, sides gently

curved, subparailel, extremities produced, the anterior obtusely

rounded, posterior prominent and submucronate. Surface of the

shell smooth, covered throughout with very small, closely-set,

impressed, circular punctations. Colour yellowish grey. Length

0°65 mm.

One specimen only, dredged in 30 fathoms.

ARGILL@CIA AFFINIS, sp.n. (Plate XX. figs. 9, 10.)

Shell, seen laterally, elongated, siliquose (fig. 9), greatest height

in the middle, equal to more than one-third of the length ;

anterior extremity evenly rounded, narrow, posterior scarcely

at all angulated ventrally; dorsal margin forming a continuous

even arch “throughout, ventral nearly straight through its whole

length: seen from above (fig. 10) the outline is narrowly subovate,

thrice as long as broad, greatest width in the middle, tapering

towards the subacuminate extremities, but more abruptly behind

than in front. Surface of the shell perfectly smooth. Colour

light grey. Length 0°54 mm.

Several specimens dredged in 70 fathoms.

The differences in form of shell between the various species of

Argillecia hitherto described appear to be extremely slight, but

that here figured does not seem fairly referable to any one of

them.

CYTHERE CRISPATA Brady. (Plate XX. figs. 3, 4.)

Specimens of C. crispata, an extreme form of which is here

figured, were dredged in 70 fathoms. This form differs from the

type in having the extremities—and to some extent the ventral

margins—irregsularly dentated. This condition, however, is found,

though to a much less marked degree, in some of the type-

specimens.

These specimens seem indistinguishable from a species described

by G. W. Miiller—C. diffusa*; perhaps also from C. elegans of

the same author.

CYTHERE CINGULATA, sp.n. (Plate XX. figs. 5, 6.)

Shell, seen laterally, oblong, subquadrate (fig. 5), of nearly equal

height throughout, height equal to at least half the length ;

extremities well rounded, the posterior rather the narrower, dorsal

margin nearly straicht, inclined gently from before backward,

* Die Ostracoden des Golfes von Neapel, p. 354.

MARINE OSTRACODA FROM MADEIRA. 597

ventral slightly sinuated in the middle: seen from above (fig. 6)

the outline is oblong, with straight, parallel, lateral margins and

strongly produced extremities ; the margins converge steeply in

front terminating in a stout median process, behind they converge

almost rectangularly and form a very wide central hump. Sur-

face of the shell covered with rounded and closely-set impressed

pittings, and bordered in front and behind with a smooth

depressed fillet. Length 0:78 mm.

One specimen dredged in 30 fathoms, and one in. 70 fathoms.

CYTHEREIS DEFORMIS, sp.n. (Plate XX. figs. 7, 8.)

Shell, seen laterally, oblong, subquadrate, greatest height

situated near the front and equal to nearly halt the length (fig. 7) ;

anterior extremity well rounded and bordered with a series of

short, blunt teeth ; posterior extremity much narrower, truncated,

prominently angular in the middle and divided below the middle

into a few irregularly tooth-like processes; dorsal margin sloping

gradually from the front backwards, ventral margin almost

straight: seen from above (fig. 8) the outline is compressed,

oblong, about thrice as long as broad, the lateral margins very

irregular, with a sharply prominent angle at the posterior third ;

extremities much produced, broadly truncate, with irregularly

dentate margins. Shell- surface irregularly rugose, with a wide

depressed band stretching round and within the anterior margin ;

a sharply elevated crest running obliquely across the posterior

half and terminating in a sharply produced angle on its dorsal

aspect. Length 0°5 mm.

Dredged in 70 fathoms.

It is impossible, by the shell-characters alone, to separate clearly

the species belonging to the two genera Cythere and Cythereis as

they are now under Stood by most authors, nor, as it appears to

me, are the distimetive characters of the contained animal much

more satisfactory.

LoxoconeHa OBESA, sp.n. (Plate XXI. figs. 3, 4.)

Shell, seen laterally, elliptical, greatest height situated m the

middle and equal to half the length (fig. 3); extremities evenly

rounded, the Cala ior somewhat the narrower of the two; dorsal

margin very slightly arcuate, ventral almost straight, not at all

sinuated : seen from above (fig. 4) the outline is very br oadly oval

with strongly produced mucronate extremities, width considerably

exceeding half the length, lateral margins very strongly convex.

Shell-surface perfectly smooth. Colour grey. Length 0°46 mm.

Dredged in 70 fathoms ; one specimen only.

LoxoconcHA DECIPIENS G. W. Miiller. (Plate X XI. figs. 1, 2.)

Several specimens, agreeing very accurately with the figures

given by Dr. G. W. Miiller, were dredged in 70 fathoms. One

quite characteristic specimen is figured here.

598 DR. G. 8. BRADY ON

LoxoconcHA SUBALATA, sp.n. (Plate X XI. figs. 5, 6.)

Shell, seen from the side, oblong, subrhomboidal, twice as long

as broad (fig. 5); anterior extremity obliquely rounded, posterior

very obliquely rounded below the middle ; ventral margin rather

deeply sinuated in the middle, curving suddenly upwards behind,

dorsal margin perfectly straight : seen from above (fig. 6) hastate

in outline, prominently angulated behind the middle, from which

point the sides converge with a gentle curve to the anterior

extremity, which is sharply acuminate; behind the two lateral

angles the margins converge rather sharply in an irregularly

sinuous curve to the posterior extremity : the general contour is

thus made of two wedge-shaped portions—an anterior larger

wedge and a posterior small one. Shell-surface rough, marked

by closely-set small fossze, sharply elevated in the postero-ventral

regions, beneath which it is depressed, forming a somewhat

flattened curved lip. Length 0°38 mm.

Dredged in 70 fathoms.

XESTOLEBERIS LATISSIMA, sp.n. (Plate XX1. figs. 10-13.)

Shell, seen laterally, oblong, subovate, quite twice as long as

broad (fig. 10), highest behind the middle; anterior extremity

narrowly rounded, posterior sloping with a steep curve to the

ventral margin, where it forms a rounded angle; dorsal margin

forming a somewhat flattened arch, sloping steeply behind, more

gently in front, ventral margin nearly straight: seen from above

(fig. 11) the outline is excessively tumid, ovate, widest behind the

middle, width equal to two-thirds of the length, mucronate in front,

broadly rounded behind; the end view (fig. 12) is very broadly

wedge-shaped, widest ventrally where the angles are moderately

rounded off, height equal to about two-thirds of the width. Shell-

surface perfectly smooth, marked in some cases with a very few

small papilliform tubercles. Colour white. Length 0°75 mm.

Dredged in 70 fathoms.

The outline shown in fig. 13 may perhaps be referable to the

male, the more tumid outline to the female.

XESTOLEBERIS NIGROMACULATA, Sp. n. (Plate XXII. figs. 1-3.)

Shell of the female, seen laterally, oblong, subreniform, greatest

height situated in the middle and equal to half the length (fig. 1) ;

anterior extremity depressed, rounded, posterior much wider and

evenly rounded; dorsal margin boldly arched, ventral sinuated in

the middle: seen from above (fig. 2) ovate, gradually tapering to

the anterior extremity, which is rather sharply pointed, much

broader and well rounded behind, width and height equal. Sur-

face of the shell smooth, yellowish in colour, with irregular

clouded dark patches, and bearing a few very minute distinctly

scattered circular papiile. Eye-spots very conspicuous. Length

0:55 mm.

Shell of the male (fig. 3) rather smaller, more markedly

depressed in front and more elevated dorsally.

Several specimens taken between tide-marks.

Caen tne

MARINE OSTRAGODA FROM MADEIRA. 599

Cy?HERURA MACULOSA, sp.n. (Plate XXII. figs. 6, 7.)

Shell, seen laterally, oblong, subreniform, highest in the middle

(fig. 6), height equal to half the length ; anterior extremity evenly

rounded, posterior produced in the middle into a wide obtusely

pointed beak; dorsal margin evenly arched throughout, ventral

rather deeply sinuated in the middle, prominent behind, thence

sloping with a sinuous curve up to the beak: seen from above

(fig. 7) oblong, three times as long as broad, lateral margins

parallel throughout the greater part of their length, converging

towards the front; anterior extremity broad and truncated,

posterior produced into a wide median beak. Surface of the

valves marked with faint longitudinal striz and with minute,

closely-set, impressed pits ; in most cases the middle of the valve

bears a dark transverse blotch of irregular shape, but this may be

absent or much reduced in size. Length 0°39 mm.

Several specimens from low-water mark.

CYTHERURA CRIBROSA, sp. n. (Plate XXII. figs. 4, 5.)

g, rhomboidal, height searcely equal

to half the length ; ainkierslert extremity well rounded, posterior

produced above the middle into a very large and rather sharp

beak (fig. 4); dorsal margin straight throughout almost its whole

length, ventral rather deeply sinuated in the middle, behind

which it bulges ventrally: seen from above, oblong, subovate,

width scarcely equalling half the length (fig. 5), rather abruptly

tapered and subacuminate in front, posteriorly abruptly truncated

and produced into a large central beak. Surface of the shell

honeycombed with large subrotund fosse, which are arranged in

irregular longitudinal rows. Length 0- 44 mm.

Many specimens dredged in 50 fathoms.

Shell, seen from side, oblong

CYTHERURA FOSSULATA, sp.n. (Plate XXII. figs. 8, 9.)

Shell, seen laterally, oblong, rhomboidal, of nearly equal height

throughout, length equal to more than twice the height (fig. 8) ;

anterior extremity obliquely subtruncate, often indented below

the middle, the sinuations bounded by two small nodules, two

rounded tubercles near the anterior extremity, just within the

superior margin, over the region of the eyes ; posterior extremity

oblique, produced above the middle into a large upward-pointing

beak; dorsal margin perfectly straight, ventral nearly straight,

with a slight ventral convexity: seen from below (fig. 9) the

outline is that of an arrowhead, the lateral acute prominences

situated near the posterior third, the width at that point equal to

more than half the length of the shell, behind these angles the

shell is suddenly narrowed, forming a rounded hinder end which

terminates ina lar ee sme en beak : " anteriorly the lateral margins

converge with a deep curvature to the front, ending in a wide

bluntly rounded extremity. The surface of the shell is variously

and very irregularly marked with furrowed undulations more or

less transverse in their direction, and by a conspicuous curved

longitudinal evest which ends behind the middle near the ventral

600 ON MARINE OSTRACODA FROM MADEIRA.

margin in a sharply angular projection: in some cases there is

a distinct longitudinal striation, more especially on the ventral

surface, and posteriorly near the base of the beak may be seen a

series of four nodules, the terminations of small curved carine

(fig. 9). Length 0-46 mm.

Several specimens were dredged in depths of 50-70 fathoms.

The surface-ornament of this species varies very much: the

foregoing description should be taken as belonging to a rather

strongly marked specimen. The variations of sculpture seem to

depend chiefly on conditions of age and sex.

CYTHERIDEIS SUBULATA, Var. CRENULATA, nom. n. (Plate X.XI.

fig. 7.) ,

Specimens, of which a figure is here given, seem to be identical

with a form deseribed in 1874 by myself and the late Dr. Robert-

son under the name Cytherideis subulata var. fasciata, the

varietal designation having been used on account of the presence

of a dark band across the shell. But the type specimens, taken

among the Scilly Islands and now in my collection, do not now

exhibit any such marking. It is possible that the markings may

have disappeared with exposure to the air, and as the published

name is inappropriate, I propose to substitute the varietal term

crenulata. In the Madeira specimens, which were taken in a

depth of 70 fathoms, the anterior crenulations are much more

developed than in those from Scilly, but in all other respects those

from the two localities are alike.

PARADOXOSTOMA GRACILE, sp. n. (Plate X XI. figs. 8, 9.)

Shell, seen laterally, oblong, subclavate, height equal to more

than one-third of the length ; extremities rounded off, the anterior

narrower and somewhat depressed (fig. 8), dorsal margin evenly

but not very strongly arcuate, ventral almost straight : seen from

above (fig. 9) extremely compressed, widest in the middle and

tapering evenly to the extremities, which are very acutely pointed,

width equal to about one-fourth of the length. Shell-surface

perfectly smooth and colourless. Length 0°55 mm.

One specimen only, taken in 70 fathoms.

CYTRERELLA (2) OVALIS, sp. n. (Plate XXII. figs. 11, 12.)

Shell, seen laterally, subelliptical. about twice as long as broad

(fig. 11); anterior extremity rounded above and below, almost

subtruncate, posterior narrower and somewhat oblique, dorsal

margin almost straight, ventral straight, upcurved toward the

posterior extremity : seen dorsally the outline is ovate (fig. 12),

twice as long as broad, greatest width behind the middle;

extremities well rounded, the anterior much narrower than the

posterior. Shell-surface perfectly smooth. Colour white. Length

0-42 mm.

Dredged in 70 fathoms.

PS SiS PE esi’

West, Newman chr.

3 CEE @ Uae

el eye Wide Tea

(Aisa ON ay

toy apevallte sulle iy) RV

West, Newman chr.

1. DARK GRIZZLE. Aan NATALIE TSH (Ey ICI, Mala

34 (ETRE A Ae aD) \Oella@NOMa IR.

PZ.S.1944. Pl. XXV._

wiles

West, Newman chr.

dy SDWVNIBNCe VATS. oy RINSE) WN TENE

oye) WORDT IMIR

PAS. 191 Pl Oa

West, Newman chr.

FEATHERS SHOWING PATTERN MARKINGS.

ON COLOUR INHERITANCE IN PIGEONS. 601

EXPLANATION OF THE PLATES.

Prate XX.

1, 2. Bairdia dubia. X 84.

3, 4. Cythere crispata. 110.

SO. 5 cingulata. X 110.

7, 8. Cythereis deformis. X 84.

9,10. Argillecia affinis. X 84.

EFratr XXI.

1, 2. Loxoconcha decipiens. X 90.

3, 4. 5) ohesa. X 84.

5, aa subalata. X 115.

7. Cytherideis subulata, var. crenulata. X 84.

Figs. 8, 9. Paradoxostoma gracile. X 84.

3. Xestoleberis latissima. X 110.

Puatre XXIT.

Figs. 1-3. Xestoleberis nigromaculata,? g. X 84.

4, 5. Cytherura cribrosa. X 100.

@, We 3 maculosa. X 100.

Saas - fossulata. .X 100.

Fig. 10. Sclerochilus levis? X 84.

Figs. 11, 12. Cytherella ovalis. X 100.

28. On Colour and Colour-pattern Inheritance in Pigeons.

By J. Lewis Bonsore, M.A., F.L.S., F.Z.8., and F. W.

Smauuey, F.Z.S.

[Received February 4, 1911: Read March 21, 1911.]

(Plates XXII-XXVL-*)

The following is a preliminary account of some experiments

undertaken by the authors to throw some light on the inheritance

of colour and colour-pattern in Pigeons. These experiments are

being continued, as the matter is a long and complicated one and

will of necessity take several years to complete.

The experiments on certain colours and colour-patterns, how-

ever, have been practically completed, and the results are briefly

given below.

Apart from the practical knowledge given in books on fancy

pigeons, no serious work has been published on the inheritance

of colour in Pigeons except Darwin’s (Animals and Plants under

Domestication, vol. 1. p. 197 et seg., 1868 ed.), and a recent paper

by Mr. Staples- Browne (P. Z. 8. 1908, p. 67).

The information in the ‘fancy’ books, valuable as a guidance for

practical breeding, is of little utility from the scientific point

of view, as details of pedigrees are often lacking. The work of

Darwin we have also had to pass over, for the present, owing to

the difficulty of making out with any accuracy the exact colour of

his birds from the terms he used. It is by no means intended to

imply imaccuracy in that most accurate of observers, but the

‘faney’ terms for colours, though well understood by breeders, do

* For explanation of the Plates see p. 619.

602 MESSRS. J. LEWIS BONHLOTE AND F. W. SMALLEY ON

not readily admit of a scientific interpretation unless the birds

themselves can be seen; frequently the same colour (from a scier-

tific point of view) may be known by different names according to

the particular breed of pigeons that may be under discussion.

Darwin’s experiments related chiefly to “reversion,” 7. e. the re-

appearance of the blue colour when distantly related breeds of

pigeons were crossed, and although our work has not been on the

same lines, the study of the blue colour should, none the less, bear

out the results arrived at years ago. We are not unmindful of

this, and for that very purpose hepe to mate up several pairs to

test Darwin’s conclusions.

Lastly there is Mr. Staples-Browne’s recent paper, in which he

has attempted to repeat Darwin’s experiments. This paper we

have been through very carefully, and find that in the main it

agrees with our results, but one or two little difficulties have to be

met, such as the occurrence of a Dun in Exp. 27, the absence of

Blacks in Exps. 9 & 11, the large proportions of white in

Exps. 16-23 and 30, and the occurrence of white feathers on a

homozygous blue.

We have no doubt that further work will clear up these slight

difficulties, which do not, however, greatly affect the main result ;

in fact a possible explanation of some of them has already suggested

itself to us, but the discussion of these is best deferred until the

results of our matings on the same lines as Darwin and Staples-

Browne have been obtained.

The majority of the birds used have been highly bred Dragoons,

but in a very few cases a Homer cross has been introduced.

The characters dealt with in this paper are :—

(i) Colour-patterns, ¢, e. Chequering, Grizzle, and Mealy.

(ii) Colours, i.e. Blue and Silver, with White and Red in those

eases where it is connected with Grizzle and Mealy.

Before detailing the experiments, however, it is necessary to

have a clear understanding of the terms used.

(i) Chequering. This is a pattern chiefly confined to the wing-

coverts, in which each chequered feather has a hght coloured

V-shaped patch at its distal end, the apex of the V being

nearest the base of the feather (see Pl. XXVLI. fig. 1).

The general appearance of a good chequered bird is shown

in Pl. XXIII. fig. 3, but it must be understood that

scientifically and in the experiments we have carried out,

a bird has been considered as chequered when it showed

the characteristic markings on its wings.

(ii) Grizzling. A grizzled feather is one in which the barbs are

partially white and partially coloured. This pattern is not

restricted to any particular part of a bird, but grizzled

feathers may be found in any feather tract including the

remiges and rectrices.

On a grizzled bird (.e.a bird with grizzled feathers)

whole coloured feathers are generally found as well as

feathers splashed with white (Pl. XXIV. figs. 1 & 2;

=a,

COLOUR INHERITANCE IN PIGEONS. 603

Pl. XXVI. figs. 2 & 4). The term ‘Grizzle’ is restricted

in the ‘fancy’ to birds whose pigment is Blue grizzled with

White.

(iii) Mealy. From the pattern point of viewa Mealy is identical

with a Grizzle but the White coloration is to a greater or

lesser extent replaced by Red. A Mealy (for Mealy

feathers see Pl. XX VI. figs. 5 & 6; the birds are figured

on Pl. XXV.) may therefore show Blue and Red, or Biue,

Red and White.

(iv) Blue (Pl. XXIII. fig. 1) is the colour of the Wild Rock

Pigeon, although in domestic breeds the rump is not

necessarily white. Ina Blue Chequer (Pl. XXIIT. fig. 3)

the light apical portions of the feathers are of the typical

blue colour, the rest of the feather and the general

appearance of the bird being very much darker. A very

dark Blue Chequer isalmost black, but this black is usually

dulland must not be confused with the glossy black (beetle-

black) characteristic of a pure black pigeon.

(v) Silver is a very pale blue with black bars and *‘ dun” flights.

In a Blue pigeon the flights are black (PI. XXIII. fig. 2).

(vi) Red. A Red pigeon is deep red all over including the flights

and tail. In a Red Chequer the dark portion of the

feathers are of the normal red and the light apical spots

white. The flights and tail are white.

It must be remembered that all these varieties show considerable

differences in shades of colour, and in the intensity and abundance

of the pattern markings. In this paper, however, we do not

propose to enter into these details, important as they are. Our

object for the present is to separate those characters which follow

apparently a Mendelian inheritance, from those whose mode

of inheritance is different. This paper therefore will only deal

with Mendelian inheritance, except to note in a few cases where

that inheritance is apparently subservient to other causes. None

the less these differences of shade and amount of pattern do

obviously follow a definite law of inheritance, and they are by no

means being disregarded by us, nor are we without hope of being

able at some future time to attempt some explanation of the laws

which govern their inheritance.

We may point out further that in the case of Mealies and

Grizzles (i.e. where normally there should be a small amount of

white), there is a great tendency to breed out in the course of a

few generations to practically pure white with only a few coloured

feathers. Our results, as far as they have gone, seem to show,

however, that these nearly pure white birds are still transmitting

to their offspring the colour which characterised their parents

and grandparents.

Blues and Silvers.

Blue is dominant to Silver.

A careful analysis of our results shows that Silver is in reality

a dilute Blue and that the colour factor in both is identical. The

604 MESSRS. J. LEWIS BONHOTE AND F. W. SMALLEY ON

allelomorph is therefore concentration and dilution (¢ and d) of

which the former is dominant.

Blue may therefore be represented as BBee or BBed, where B

represents the blue colour factor. Silver must. theretore be repre-

sented by BBdd.

Blues are said to almost always breed true, that is to say, that

they never throw any Silvers—this idea owes its origin, however,

to the fact that those Blues which are impure dominants (2. e.

containing the factor d) are of a poorer colour than the pure Blues

and have in consequence long been rejected by fanciers, so that a

race of pure dominants has thus been evolved *. Several of our

matings show that Blues or Grizzles that: are heterozygous as

regards cand d will throw Silvers.

The following are the results of our matings for these particular

characters, with the exception of Experiment 83, which is intro-

duced to show the reappearance of Silver from two heterozygous

Blues.

{Berenson bos IMGae alc ae lie ce:

| | Sales | ies eo foe ee EA DA Oe aR a as!

gs. |00/61 | Grizzle| BBed@g | — || 09/126 | Blue |BBedgg | — || 3B 1S.

14g. | 04/70 | Silver | BBdd — |}oa21 | Silver |BBdd | — | ANS

149. | 05/96 | BBdd — || 05/51 ‘! BBaa == |) ox

150. | 07/4 » |BBd@ | — [0656 | ,, | BBad =" Hh hs

151. | 05/96 so (eBiBad le Osan) Mines ea BBG: Zale os

152. | 03/5 i ee a BBddgg | 83 || 04/140 | Grizzle | BBee or ed | 157 All B or Equality

153. |0410 |, | BBdd_— | 140 {oat| » |BBecored| 77 || 3

154. | 04/14 | Grizzle | BBee or ed) 96 | 07/52 | Silver | BBdd | 151 4s se

185. | 08/1336 | Silver | BBdd | 148 | 08/1368 | Grizzle | BBee or ed| 137 | eS *

156. |03/5 | | BBaa | 88 |] 01/50 | Blue | _BBce | — | All Blue

fey 03sje- | 5, | BBdd 9) 83 (02/81 } 5. | BBee | — i

158. | 05/95 | Blue BBecored| 105 |/05/51 | Silver | BBdd | — ||, ,,

169. | 06/13 | ,, BBed 65 ||06/66 | ,, | BBad [es i Equality

160. | 03/5 | Silver BBdd 83 ||03/126 | Blue | BBcd = wpillind ss

Tete] OF . |. gl) PBBda > [i NOGereanerr | BBed: \ EB adie ds

162. | 04/52 Blue — BBed 156 || 06/56 | Silver | BBdd — | F

6B28

Result. |

7B —

4B 28

When the Gametic Formula is in italies it implies that it is not definitely known. Only the

characters under immediate consideration are given in the Gametic Formula.

* See also remarks on chequering.

+ We must, however, point out here that in Mr. Staples-Browne’s experiments,

one or two blues, apparently heterozygous, behaved as homozygous and vice versa.

~ With regard to the numbering of Experiments—the numbers refer to our stud

book, and rather than number the experiments quoted successively in this paper, it

was thought best to have only one set of numbers so that references might at any

future time be easily made without risk of confusion.

COLOUR INHERITANCE IN PIGEONS, 605

From these matings we see that :—

(i) Silver to Silver (Exps. 148 to 151) gives nothing but Silver,

according to expectation.

(ii) Blue to Silver (Exps. 152 to 159) should give nothing but

3lues or Blues and Silvers according to whether the Blues

are heterozygous or not,and wesee that when our knowledge

of the gametic for mul of the parents was known for cer-

tain, the results were in exact accord with expectations, and

no results ant: vgonistic to possible expectations occurred.

(iii) Heterozygous Blues to Silvers (Exps. 160-163) should give

equality of Blues and Silvers. In the case of Exps. 160 and

161 no Silvers appeared, but the numbers bred were very

small. In the other experiments, exact equality was

reached, so that we may well assume that a continuation of

the other experiments would have led to the appearance

of some Silvers,

(iv) Heterozygous Blue x Heterozygous Blue should give Blues

and Silvers in proportion of 3:1, which was the exact

result of this mating (Exp. 83).

The results therefore show clearly that Silver is recessive to

Blue. The arguments showing that the difference between these

colours is one of concentration and dilution (rather than a differ-

ence of the colour factor itself), depend on the study of this

dilution factor, which, as it concerns the inheritance of other

colours and shades, not dealt with in this paper and at present

only imperfectly understood, is best deferred for the present. Its

discussion in no way affects the proof of the dominance of Blue

over Silver.

Chequering.

Chequering is dominant to pure colour. It is difficult to

realize therefore how it can have originated, since the typical

wild pigeon shows no such markings. At the present time,

however, in many parts of the country wild birds show

chequeri ing, but it seems more than likely that in these cases the

marking has been introduced by a cross with the domestic bird, as

most of the Wild Rocks in this country are now intermixed with

feral ones. Once this cross had been effected the chequering would

of course frequently show itself.

In direct contradiction to the foregoing remarks the following

fact is worth noting. Some years ago a pair of pure wild bir ds

was taken by one of the authors from a remote district in the

West of Ireland. All the wild pigeons seen (and there were no

tame ones within a radius of at least 30 miles) were purely typical

Blue Rocks. This pair bred in an aviary for five or six seasons,

producing only typical wild birds like themselves. Two seasons

ago (in 1909) an attempt was made to establish some of their

progeny as semi-wild birds and they were allowed to fly at liberty

from a dove-cot. One pair remained and reared several young, one

of which proved to be chequered !

Proc. Zoou. Soc.—1911, No. XLITIT. AB

606 MESSRS, J. LEWIS BONHOTE AND F. W. SMALLEY ON

Scientifically this result is of but little value, as the hen bird

may have made a chance mating with a chequered pigeon ; on the

other hand, the youngster was in every other respect (shape and

size) a ty pically wild bird. Unfortunately it met with an accident,

so that we have not been able to breed from it.

Unsatisfactory as this case is, it is obvious that the chequering

must in the first place have onginated from the Wild Rock

Pigeon, and the above is probably a good example of its arising

as a mutation.

A further explanation may possibly be found in the fact that the

chequering cannot show itself except in the presence of two shades

of colour, and that it may be present in many of the pure wild birds

but cannot show itself until the colour factor producing the two

shades is present, when the chequering will immediately appear.

This, however, in no way affects the main issue, namely that

when once a Chequer has been produced it is dominant to the pure

colour. The mating of two Chequers should therefore produce

Chequers or Chequers and Self-colour, according to whether the

birds are homozygous or heterozgyous for that character.

As in the case of the Blues, so in the case of the Chequers it

appears that fanciers, by continually selecting the best birds for

breeding, have unconsciously been selecting only homozygous birds,

with the result that only pure Chequer dominants are to be found

in certain strains. We have for instance accurate records of

57 matings (Exps. 1-49; 169-172; 175-178) of Chequer to

Chequer from which 229 young were produced, and all these

without exception were Chequers like their parents. It theretore

became essential to carry out further matings with birds that

were known to be heterozygous in order to test the dominance

of the Chequer character. This has been done in the matings

detailed below, the results of which, as will be seen, approximate

very closely to the Mendelian expectation.

App. Formula. Exp. App. |Formula.| Exp. tions. | aa

06/104. 4 ace } Xx | 173 ‘|| 06/82 | Blue. XX 122 | Equality 2Ch :2B

650. F Once i) Xx | 148-B |} 06/69 Grizzle.| xx | 158 x 3:2

lc G : ; | |

07/5 if Gate ; Xx 196 |kaS/e | Blue || xx | 196 be es

ye; | § Grizzle ; ail Weer | Bive a S| ‘

O7/a a Chequer Xx 196 || 08/d Chequer, Xx 196 | 3:1 | 2:2 |

07/3 E Gane ; Xx | 196 | 08/13! Grizzle.| xx | 196 |Equality) 2:0

Areal alate 27) 2 li pgjoo | Blue | 7 | ie ire

05/19 | 2 Chequer. Xx 197 1 08/22 | Chequer, Xx 197 3:1 10:1

ee ue ae || eps | ue = i .3%

09/533) } Chaguer s| Xx | 197 || 08/36 ‘Chequer, X* | 188) 3:1 | 3:34

08 24. | { Ghee : Xx 198 | 08/30 | Grizzle. NX 197 ||Equality}; 7:2

Chequering and its absence are represented by X and x.

* Matings for the purpose of testing the extracted recessives from Exps. 200 and

201 have been undertaken this year (1911), and prove the recessives to breed true.

COLOUR INHERITANCE IN PIGEONS. 607

It should perhaps be noted that although the individual matings

show, considering the small numbers, a very close approximation

to the expected results, yet at the same time the tendency to vary

from the anticipated results isall in one direction, viz. to a greater

number of Chequered birds. This is most marked in those

matings where equality was expected, for of the 28 birds bred,

19 were Chequers; in those cases where the-expectation was 3: 1,

21 birds were bred of which 15 were Chequers, which is approxi-

mately correct. It is possibly due as much to this tendency as

to the unconscious selection by breeders, that this character has

become perfectly true and stable in some strains. As pointed out in

the earlier portion of this paper, we are restricting our remarks for

the present to the consideration of the Mendelian inheritance of

certain characters, and that theory seems to fit in well with the

main lines of inheritance as borne out by the facts, None the less

it is equally evident that there are other factors at work, which

are able to modify to some extent the results anticipated by the

Mendelian hypothesis. In addition to these definite matings we

have also notes of 6 matings Chequer to pure colour (Exps. 164—

168, 173) which gave 19 birds all chequered, In this latter set

of matings most of the Chequered parents were birds used in or

bred from the Chequer to Chequer matings; and therefore this

adds further proof that all those birds were homozygous

dominants, as otherwise we should have expected some self-

coloured birds to appear as they did in Exps. 174-203 (p. 606).

It may be as well to mention here that although some of the

matings referred to in this paper were not undertaken with the

special purpose of bringing out the facts which they are used to

interpret, yet they have all been conducted by one of the authors

in person. Special matings have, however, been made in every

case to prove the inheritance of the characters discussed *.

Grizaling.

Grizzling is dominant to Chequering and hence also to pure

colour. It probably originated from the cross between Blue and

White, although such matings usually give splashed birds, owin

probably to the true Grizzle character, in which individual barbs

show both white and blue, being absent. Cases, however, are

known in which the cross between pure White and pure Blue

have produced Grizzles, and in these cases there is little doubt

that the Grizzle character must have been present in one or both

of the parents but was unable to show itself owing to the bird

containing only one colour. Once, however, the Grizzle has shown

itself, the White and Grizzle characters seem to combine together

and to have a common inheritance. Furthermore, as already

stated, Grizzles when bred together tend usually, but not

invariably, to show an increase of white in successive generations

* Mr. W. Bateson has stated (Mendel’s Principles of Heredity, p. 43 (1909)) that

chequering is dominant to its absence; on writing to him for a reference to the

source of his statement, he says that he had in mind some experiments of Mr. Staples-

Browne, which have, however, not been published.

608 MESSRS. J. LEWIS BONHOTE AND F. W. SMALLEY ON

till eventually some birds will be produced showing only one or

two coloured feathers. This matter, however, we shall not

discuss at present.

For our present purpose we may ignore the White character,

assuming that it is always present and inherited in common with

the Grizzle. The gametic formula, therefore, of a grizzled bird

will be BBee or ed GG or Gg, where B is the Blue colour,

e & d concentration or dilution (the combination dd producing

a bird known as a Silver Girizzle), and G & g the presence or

absence of Grizzle. Therefore from the crossing of two Grizzlies

we may either get all Grizzles or Blues and Grizzle es, according to

whether the birds are homozygous or heterozygous to the Grizzle

character. Under the term Blue we here for simplicity’s sake

include Silver.

Again, as in the case of the Chequers, the results from pedigrees

are apt to be misleading owing to the difficulty of distinguishing

the homozygous from the heterozygous birds ; nevertheless such

results entirely bear out our hypothesis, since, as regards the

colours produced, the results are quite in accordance with ex-

pectations.

Grizzle to Grizzle*,

74. 02/46] Griale |G@GorGg] — |/99/104 Grizzle| GG | — || All Gor3:1/3 Grizzles

75. |03/21 | % Gg 87 {98/88 ; iGGorGg] — || ,, » | 5 Grizzles

76. |00/21 a GGorGg) — ||63/7 ss KC ore == 5 | & Grizzles

77. |00/61 - Gg | 99/104) 4G — || ,, » | 7 Grizzles

78. |04/21 » |GGorGg) 76 04/58 é Ge oy ll. | 4G 2a

79. 104/23 . Ge 99 ||06/102) Gg | 153] 3:1 Me WS

80. |04/1 rev. “3 Ge 99 1105/6 | " Ge 109 |) 333 IL 1G —

| 81. jowie |, Gg 103 |o1/15 [SO 16Gror Gel == i AniGeor se fate anne

* 82. |06/88 55 GGorGg 78 ||04/58 x Ge 94 | AllGor3:1] — 1B

% As Theos aane the Gharacten dealt tans Is Shows nin the Cancun Woumula,

Looking through these matings, we see that when both Grizzles

and Blues should have appear ed the expected proportion ought to

have been 3:1, and the results give 10:5 or a rather large

excess of Blues.

Grizele to Blue.

These matings (Exps. 83-138, 152-155) may be divided into two

groups: (1) those in which the. gametic formule of both parents

were known ; (2) those in which, owing to the gamietic formula

of one par ent coe doubtful, two expectations were possible.

Certain of the Grizzles were known to be heterozygous: this was

the case in 19 matings, in which therefore Grizzles and Blues

should have appeared in approximately equal numbers. Alto-

gether 64 young were reared, 37 Grizzles and 27 Blues, thus

showing a slight excess of Grizzles.

; COLOUR INHERITANCE IN PIGIONS, 609

Grizzle to Blue.

| | i

0 IWxt. |Gametic From | Ext. |Gametie! From |, ee é

t App. |Formula., Exp. | 3 ADD. ‘Formula. SD: | Expectation. Result.

00/61 Grizzle |GGorGe| —_ || 02/126 nice ge | == “All Ge or Kqui Ting 4G 4713 |

01/27 Blue oo a= | 01/28 Grizzle |(GGorGe) — | 3 P 5G 1B)

O1/10 u | Bo — || 03/26 , |IG@GorGe — || ,, ss 1G —|

00/21 Grizzle \GGorGe| —_ || 02/35 Blue OM) ieee IML gi Po 1G 2B)

00/6 rev. | Blue go — || 02/24 Grizzle |GGorGg) — - F 2G 423

02/9 a | gor — || 01/24 » \IaGorGg) — || ,, 3A 2G 4B

02/37 | Grizale |GGorGg| — _ || 02/41 | Blue ge | — A, vs 1G 2B

02/46 (a) Pr GGorGg) — 21 i Pape Wo IN pe 3 4G —

02/46 (b)| 4, IGGorGg| — 43 - oor — | 5 - ==) 6B!)

03/31 Blue go 84 || 02/23 Grizzle |GGorGg, — | ,, i — 5B

01/27 ” £8 cae | O1/15 ” ” eta ” ” 4G 23 |

03/19 _ 0" 86 || O1/24 3 as — |, mm 2G 2B

03/11 iy ow 51 || 02/77 ss rile ileal ta | caer . 4G 1B

Ol /13 ” £28 baa (1/28 ” ” ee! ” ” 4G 1B

03/37 Grizzle GGorGg| —_ || 02/38 Blue ge — || 4 3 — 8B

03/20 fr Ge — || 02/22 55 oor — || Wauality 1G 2B

O1/11 Blue eo — || 03/9 Grizzle Gg | 84 | - 5G 1B

01/10 is ep — || 00/53 in GGorGe — |AllGgor Hquality) — 1B

NORTON Ns oo — || 98/96 iy oI oi i i 3G 3B

02/34: Guizzle |GGorGe| — _ || 02/26 Blue ge | — | Bt is — 1B

98/9 | - i — || 02/20 i | Popes Soll y a WG eus

00/6 Blue oor — || 03/63 Grizzle | Ge | 88 - Jquality 1G —

02/52 Bs | po — || 03/7 » |GGorGs, — | AllGgor Equality) 2G 4B

01/27 re oo — || 01/24 * IGG orGe) — - rf — 1B

02/46 (6) | Grizzle |GGorGe| —_ || 02/25 Blue on : r - 1G 5B

03/75 2 5. 74, || 03/51 “bi ge BBE Nt ie # 3G 2B

02/46 (a) - i. — || 03/23 os gor 53 Ci|| 2G 5B

04/31 | Blue go 59 || 03/9 Grizzle |} Ge | 84 " Wqu: lity 2G 1B

04/7 I ge 95 || 00/53 » MGGorGe| = All Gg or Equality) 1G 3B

04./4 is oo — || 01/28 55 IGG orGe) — if Ai 2G 2B

03/29 As oor 88 || 02/77 4 GG orGe) — os s 3G 3B

00/21 Grizazle |GGor Ge} —_ || 04/84 Blue | gg 157 os He 2G 3B

04/3 rev. | Blue oo 61 || 03/8 Grizale |GGorGg) — a ‘ Gielen

00/61 Grizzle Ge — || 04/12 Blue | gg 61 % 3 6G —

05/14 Blue gor 65 || 01/53 Grizzle GGorGg) — . 3G 5B

| 04/121 EY) ge 59 || 01/24 ” ” nei ” oy) Cr |

01/7 Grizazle |GGorGe| — || 02/22 Blue | gg — ii (3 2G 8B

02/46 (a) * — || 04/53 4 ge 61 58 is 12G —

03/75 BS x 74, || 04/48 fe geo a= a es 2G 4B

02/46 (b) e 3 — || 03/78 ‘) go 53 % i 1G 5B

O4/4rev. | Blue ge 61 || 05/6 Grizale | Gg 95 Kquality 3G 4B

04/93 re oo 61 || 01/53 » |GGorGe| — | AllGgor Equality!) 4G 1B

04/115 Griazle | Gg 104 || 04/120 Blue | ge 59 || Equality 3G 2B

04/133 i Gy 90 || 02/22 i og — || 216 —

04/52 Blue ge 156 | 04/10 rev.) Grizale GGorGg 77 | AllGe or "Equality! 4G —

05/40 Grizzle Ge 108 || 04/53 Bich mare 61 || quality 3G —

05/119 Blue ge 65 | 00/53 | Grizzle GGorGg — — AllGgor Equality) 1G 1B

02/46(a) | Grizale |GGorGg| —_ || )4/27 Blue ge | Ql \2q —

04/14 nt Gg 96 || 06/87 Fe fe fsafosy fh iL} ! ” quality 4G —

05/92 Blue oe 105 || 03/9 Grizale | Gg | 84 || i Gra lls

04/121 . Bo 59 || 04/58 aly Ger, |) Oa i [ete

05/96 AA oor 105 || 06/22 Ah a AG yea alteattalis) of Pe |) A Bay

04/13 Grizale | Ge 96 || 04/48 Blue | ge | — || cs |\4G 1B

06/57 | 6 Gg 118 || 06/63 55 | ge 123° || 5 — 1B

04/121 | Blue ge 59 || 04/58 | Grizzle | Ge 94 | ey 2G —

06/57 Grizzle Ge 118 || 05/43 Blue yp) Mila ‘ |} — 1B)

03/5 | Silver eo 83 || 04/140 Grizzle | Ge lal 3 Gaels

04/10 si ro 140 || 04/l0rey.| _,, GGorGeg) 77 | AllGgor Hquality) 7G — |

04/14 Grizale | Gg 96 || 01/52 Silver we Uy Kquality 2G 4B

08/1336 | Silver gg | 148 | 08/1368 | Griade | Gg | 137 3 | 2G 4B

610 MESSRS. J. LEWIS BONHOTE AND F. W. SMALLEY ON

In only 8 of the other matings did Blues fail to appear, and

we may therefore consider that when they did appear the

expectation should have been equality as before. Deducting

therefore these 8 matings which only produced Grizzles, we have

left 33 matings producing 156 young, 63 being Grizzles and

93 Blues. Since only one expectation is possible if both colours

appear, We are justified in uniting the figures from these two sets

of matings and treating them together. We find, therefore, that

in the total we have 52 matings of Grizzle to Blue, which produced

220 young of which 100 were Grizzles and 120 Blues; and this

proportion, although not exact, is not unreasonably far from the

expectation (Equality), and certainly seems to show a Mendelian

basis of inheritance.

If we digest the facts still further, we find that actual equality

was reached in only 19 eases, and this by including the odd

numbered broods where the deviation was not more than one.

We have, therefore,

Actual Equality reached in roughly 36°/, of the

matings.

Blues outnumbered the Grizzles by more than one in 20°/, ,,

Grizzles ss ., Blues = Es 12°(0 ss

Blues only, appeared in ilo) Meee

Grizzles only, : _ PAS Joment Ez

This, therefore, seems to show that while only a moderate

percentage of matings gave the exact Mendelian expectation, the

variation to one side or other of the mean is fairly evenly balanced,

with however a slight but unmistakable tendency towards an

overproduction of Blues: a tendency which was also shown in

the Grizzle to Grizzle matings.

Thus, as in the case of the Chequers where we found the

Mendelian proportions fairly well maintained, but with a distinct

tendency to an overproduction of Chequers, so also in the Grizzles

we see a similar tendency to an overpreduction of Blues.

There remains for consideration the 8 matings in which only

Grizzles appeared. Two of these may be at once dismissed as

only one bird was reared, so that we have no hint as to the

probable expectation. From the other 6 matings 25 birds were

reared, so that there is a reasonable probability of a Blue having

appeared were either of the parents heterozygous. Unfortunately

for the simplicity of this reasoning, we find on investigating the

matter more closely that although in 3 of these matings the

same hen was used and we might therefore presume her to be

homozygous, yet by her progeny in another mating (Exp. 109)

she proved herself to be heterozygous. This then leaves only 3

cases out of 60 in which the Grizzle parent might be homozygous.

So that again, just as we found in the Chequers and Blues an

overwhelming proportion of homozygous birds, which we attributed

to the unconscious selection of breeders, similarly in this case the

* This last calculation is of course exclusive of the 8 matings mentioned above,

in which the gametic formula of the Grizzle parent was doubtful.

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612 MESSRS. J. LEWIS BONILOTE AND F. W. SMALLEY ON

number of the heterozygous birds largely predominates, and this

is probably due to the fact that Grizzle x Blue is the commonest

form of mating used by breeders to produce Grizzles.

We have no records of matings with 2 extracted (Grizzle-bred)

Blues, but of Grizzle-bred Blues to Blue we have made 9 matings

(Exps. 139-147). These matings produced 28 birds all Blue in

accordance with the expectation.

Tn order to still further test our hypothesis we have carried out

the foregoing experiments (see Table, p. 611).

The result of a ser utiny of these matings is sufficient to prove

the Mendelian inheritance of the Grizzle “character. In some of

this last batch the expectation was well defined ; in others—owing

to the impossibility of distinguishing homozygous and hetero-

zygous birds—the expectation was open to one or two, or in some

cases three, interpretations. In those cases where the expectation

was all Grizzles or Grizzles and absence, in proportion of 3:1, and

only Grizzles were produced, we have concluded that one of the

parents, at least, was homozy gous—similarly, if any of the pro-

geny in those cases lacked the Grizzle, we have presumed that

the expected result should have been 38:1. On this basis we

have tested the results and we find :—

No. of

Expectation. Matings. Result.

AUG a 27 1ESy es ccteuys 7 39 Grizzles.

BE ea Aa aire Helin s ar eee 32 Grizzles, 11 absence of Grizzle.

ADH GHP aeaen ace ec nncne cds 3 9 Grizzles, 9 absence of Grizzle.

INOMGIZZlesm Byaceen tones 2 18 absence of Grizzle.

Higuelityror 3: bean. Sit. aie 5 Grizzles, 1 absence of Grizzle.

This last set of matings places therefore beyond doubt the

Mendelian inheritance of the Grizzle character.

In the summary given above we have, however, left ont

Exp. 180, in which with an expectation of 3:1 or all Grizzles,

14 Grizzles and one pure Blue were produced. At the present

moment, we can offer no reason for this considerable deviation

from the expected result.

Grizales and Chequers.

The Grizzle character is dominant to the Chequer, although,

in almost every case, the heterozygotes may be easily recognised.

To test this inheritance we have made the following matings

(see Table, p. 613).

These matings show fairly clearly the mode of inheritance,

but from the smallness of the numbers the proportions of the

different colours are not always in exact accordance with the

expectation. It may be further noted that although, from the

above reason, certain expected colours have as yet not occurred,

on the other hand no unexpected colours or combinations have

appeared, Thus we see that in Experiments 200 & 201 the

recessive Chequers: gave us, as expected, 14 Chequers to 4 Selfs,

and no Grizzles; in Experiments 202 & 205 the pure Grizzles

gave us, 11 Grizzles to 1 Self, and no Chequers. These two latter

613

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614 MESSRS. J. LEWIS BONHOTE AND F. W. SMALLEY ON

matings (Exps. 202 & 205) may be compared with Exps. 203 &

204 *, ‘matings of heterozygote Grizzles, where two Chequered birds

turned up, although the whole number bred (12 birds) was far

too small to give any hope of an approximation to the expectation

being attained, Many of the Grizzles in these matings are very

white, a factor which would tend to conceal the Chequer markings

if present. It is more than likely therefore, that owing to this,

some of these so-called Grizzles are in reality Grizzle-Chequers.

Taking the Grizzles and Grizzle-Chequers (of Exps. 203 & 204)

together, the expectation is 12 Grizzles and Grizzle-Chequers,

4 Chequers and 2 Self, and our result, 12 Grizzles and Grizzle-

Chequers and 2 Chequers, i is reasonably near the anticipation.

The expectation in Experiments 179, 193, 195, 196 & 197,

was an aqme of Grizzle-Chequers, Chequers, Grizzles and Selfs,

and the result 5 Grizzle-Chequers, 12 Chequers, 5 Grizzles, 8 Selfs,

the only real discrepancy here being the overproduction of

Chequers. This apparently merely confirms the conclusion come

to in an earlier part of this paper, p. 607, which showed that

there is apparently some factor which overrides the Mendelian

inheritance, and leads to an increase in the number of Chequers

produced.

Mealies.

A ‘Mealy’ may best be described as a Red Grizzled Pigeon

showing Blue. The general appearance may be seen by merercnee

to Plate XXV, and it should be noticed that it has the red bars

and white flights characteristic of some varieties of red pigeons,

When dealing with the Grizzle-character (G) we had to consider

its relation to White, and we came to the conclusion that the

White was not a colour factor complementary to Blue, but a

separate allelomorph; so that the real gametic formula of a

erizzled bird was made up of a compound allelomorph containing

three characters—blue, white, and grizzling (B, W,'G).) Bor

practical purposes, however, the W & G combine in their inherit-

ance, and thus in the cases we have been considering they have,

for convenience, been regarded as a single character (G).

In considering the inheritance of the Mealies, however, we

must again pause to consider whether the factor for Red is to be

treated as a colour factor, complementary therefore to Blue, or as

complementary to the factor for White, or yet again as a separate

allelomorph.

At first sight it would seem natural to consider it an alternative

colour to Blue, but if this be the case a certain number of Self

Reds should have e appeared in our matings. None, however, were

produced, though a certain number of Blues have been reared.

The same argument, though in a lesser degree, should hold

good if it had an inheritance of its own, and we are thus driven

to the conclusion that the Red is a complementary factor to the

White. We have also had certain aberrant results (not deait

with in this paper) which point to a curious connection between

these colours.

* This pair has produced both CG and pure Chequers this year (1911).

COLOUR INHERITANCE IN PIGEONS. 615

Finally, the study of the Mealy itself bears out this contention.

A dark ‘ Mealy’ differs from a dark ‘Grizzle’ in the fact that

the white portions of the latter are replaced by red and the

flights and tail instead of being black are white. In the Grizzles

we noted that there was a great tendency for the white to

increase till an almost pure white pigeon, showing only a few

coloured feathers, was produced. Matings of Mealy to Mealy

show an increase of the white, as is the case with Grizzles, till we

eventually get a white bird showing a few coloured (Blue or Red)

feathers. We must then come to the conclusion that in Mealies

it is the white that is replaced by red, and not the blue, and

therefore that a Mealy is a Grizzled bird in which the white is

wholly or partially replaced by red.

We have had.to go into this matter thoroughly as it offers

certain difficulties, which cannot be entirely cleared up till the

relationship of white and red have been further investigated ;

nevertheless the results of our experiments will offer no difficulty

if the red is considered as an alternative factor to the white.

In onr experiments with Mealies, one character has been

present in all the matings, namely Blue, or in its dilute form

Silver; we may therefore dismiss it from our calculations.

The only Chequer which appears (the one in fact by which the

- red colour was originally introduced) is shown in Exp. 179;

the only Grizzle bird from that mating (Mealy ¢ 54), whose

descendants form the large bulk of the Mealies, emphasizes the

truth of his inheritance, since no chequered bird has appeared in

spite of the large number bred.

The Grizzle character in the Mealy or Mealy bred birds we

have already dealt with (Exps. 180-189, p. 611). The question

therefore left us to consider in dealing with the Mealy inherit-

ance is the question of Red and White. Red is apparently

dominant to White, and in consequence a Mealy is dominant to a

Grizzle.

The following matings (see Table, p. 616) show the inheritance of

this character. It must be borne in mind that W & w are in this

case practically equivalent to G & g in the earlier part of this paper.

According to our present knowledge, when the Grizzle character

meets with either white or red they combine in their inheritance

to give either a Grizzle or a Mealy.

Taken as a whole, it will be seen that the results come remark-

ably near the expectation. In 5 of the 11 matings, in spite of

the small numbers, the results exactly bear out expectations ; and

in all the others, with the exception of Exp. 181, the results are

sufficiently near to leave little doubt that a continuance of the

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Expectation 2M.1G.1S. No.of Matings 4. Result 18. 10.9

a 2M 2G 0S © 5; Pe peed 4-8-0

in both of which sets there is a slight tendency to fewer

MESSRS. J. LEWIS BONHOTE AND F. W. SMALLEY ON

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COLOUR INHERITANCE IN PIGEONS, 617

Grizzles than one would expect. So that here again, although

the numbers are smatl, the Mendelian proportion seems to be

upset by a slight tendency in a definite direction.

The occurrence of a pure Blue (no. 2) in Exp. 181 is the only

instance throughout the whole of our matings in which a colour,

other than one which was expected, has appeared. Possibly the

male parent of this mating (1326), although unlike his brother

(1354) in external appearance, had in reality the same gametic

formula. Such an occurrence would not be without precedent, as

Mr. Staples-Browne instanced a homozygous Blue which showed

a certain amount of white, and this ought to have indicated a

heterozygous bird. The difference between these two birds (i. e.

presence of white) is of the same nature, and in support of this

suggestion the result of the mating—7, 2. 1—seems to be following

on the same lines as Exp. 182.

There is one other possible suggestion for this abnormal result,

and this is that the White or Red character may not have been

inherited with the Grizzle.

The full formula of the parents on this basis is :—

2 BBed Gg [Ww|— ¢ BBed Gg [RW].

Now if the Grizzle has been independently inherited, we might

get BBedgg [RW, WW, wR, Ww] as the formula of one of

the offspring, and the last character | RW ete.] might not in the

absence of G be able to show itself, and this would give us what

we got, namely a pure Blue. The result of the mating of this

self-coloured bird (Exp. 192) throws no light on this, for if, as is

oS possible, her mate was a homozygous Grizzle, no selfs would

uppear.

It must be understood that these are merely possible explan-

ations, of which we are inclined to favour the first, but at present

we have no definite proof in support of either.

For the rest, we claim that our hypothesis is so closely borne

out by the facts that 1t may be accepted till further work

confirms or disproves it: and until we are clearly able to

differentiate between those characters which follow the Law of

Mendel and those which are apparently governed by other laws.

In this paper we are only touching on the fringe of colour inheri-

tance in Pigeons, as there still remains the question of Black,

Dun, Red, Yellow, and White inheritance, on which we are at

present continuing our researches. Our work, however, empha-

sizes the fact that there are three important problems which the

Mendelian hypothesis fails to meet :—

The differences of shades in the same colour ;

(ii) The predominance of one sex in certain colours * ;

(i) The gradual increase of the white in Grizzles and Mealies

in successive generations ;

* This most interesting question has not been dealt with in the present paper,

as we have not yet fully nivestic ated the results ; but we may mention that a large

proportion of the White Grizzles are 2’s, and in the Light Mealies by far the

larger number are @’s; we have also bred a certain number of Cream Mealies, and

these have all been ?’s.

618 MESSRS, J. LEWIS BONHOTE AND F. W. SMALLEY ON

and in addition to these the apparently large predominance of

homozygous Chequers and heterozygous Grizzles.

These questions have been very much before us during the

whole of our experiments, as Indeed must be the case, for they

cannot fail to bring themselves to the notice of every practical

breeder.

At the present moment we do not consider it advisable to

bring forward any attempted answers to these questions. Much

more work yet remains to be done among the other colours, and

until we know more of their inheritance any attempt to solve

these problems would be premature. On the other hand, we are

not without hope that the further experiments in which we are

at present engaged may help to throw some light on these

perplexing problems.

Summary.

This paper may strike a reader as having for its main object

the confirmation of the Mendelian Laws; that, however, is by

no means the case. Realizing that many details of inheritance

did not entirely accord with the Mendelian theory, our object has

been to extract, so to speak, from the results obtained by mating

on Mendelian lines, that portion of them which clearly shows the

Mendelian inheritance. ‘The residue must, we submit, have been

brought about by some law or series of laws, which overrides and

modifies (externally at all events) the expected Mendelian results.

By means of this extraction we are able to see the effect of that

law or laws untrammeled by the effects of inheritance as ruled

by the Law of Mendel, and thus we have been brought slightly

nearer to an understanding of them.

Now this paper deals with the Mendelian inheritance of the

characters considered ; consequently all details of shades of colour,

predominance of one sex in certain colours, and several other

similar matters have been entirely omitted.

On the other hand, certain points such as the superabundance

of Chequers in our Chequer matings and of Blues in the Grizzle

and Blue matings, have had to be brought forward and the results

are, we hope, sufficiently conclusive to prove to our readers that

although the characters dealt with follow in the main on the lines

of the Mendelian inheritance, yet it is equally certain that there

is another factor which is able to dominate and influence that

inheritance.

So far asthe matter has been dealt with in this paper there is

no evidence to show that the gametic inheritance has been

affected. Except in one doubtful case (Exp. 181) we have not in

the course of all our matings bred a single bird that was not a

possible result of the mating under the strictest expectation of

the Mendelian theory. What, however, we have been able to

show is, that in certain cases a consistent deviation from the

expected proportions occurs. It would thus seem at first sight as

COLOUR INHERITANCE 1N PIGEONS. 619

if some factor exists which has the power to influence but not to

alter the gametic inheritance.

The Mendelian conclusions reached in this paper may be briefly

summed up as follows :—

(1) Silver is dilute Blue.

(2) Blue is dominant to Silver.

(3) Chequering is dominant to its absence (7. e. a Self-colour),

(4) Grizzling is dominant to its absence (i. ¢. a Self-colour).

(5) Grizzling is dominant to Chequering ; the impure dominants

may however sometimes be easily distinguished.

(6) A Mealy is a Grizzled bird with the White wholly or

partially replaced by Red.

(7) Red in a Mealy is apparently dominant to White, and

hence a Mealy is dominant to a Grizzle.

(8) White and Grizzling when they have met combine together

and have a common inheritance.

(9) Red combines with Grizzling in the same way as does White.

EXPLANATION OF THE PLATES.

Prare XXIII.

. Blue Pigeon.

. Silver Pigeon.

. Chequered Pigeon.

ea a

PLATE XXIV.

. Dark Grizzled Pigeon.

. White Grizzled Pigeon.

. Grizzle and Chequer (Almond Grizzle) Pizeon—in nest feathering,

showing adult feathers appearing on the wing-coverts. (Note chequering

on the adult wing-coverts.) ;

wrt

PLATE XXV.

. Dark Mealy. Pigeon.

. Light Mealy Pigeon.

- White Mealy Pigeon. (Tricolor.)

wre

Pratt XXVI.

Feathers showing details of pattern-markings.

. Chequered feather.

. Dark Grizzled feather.

. Grizzle-Chequer feather, adult plumage, as shown in dark wing-coverts,

Pl. XXIV. fig. 3.

. White Grizzled feather.

- Light Mealy feather. (Note white. red, and blue in some barbs.)

. Dark Mealy feather. (Note absence of white.)

Fig.

On Whore

DR. R. T. LEIPER ON NEMATODE PARASITES.

EXHIBITIONS AND NOTICES.

April 4, 1911.

Dr. Henry Woopwarp, F.R.S., Vice-President,

in the Chair.

Sir E. Ray Lanxester, K.C.B., F.R.S., F.Z.S., exhibited a

special Supplement of the ‘ Field’ newspaper dealing with the

British non-migratory Trout, and called attention to this new

medium for the publication of scientific observations requiring

illustrations.

Dr. R. T. Lererr, F.Z.8., gave a demonstration of Nematode

parasites obtained from animals in the Zoological Gardens during

the vear ending November 1910.

The collection contained a number of new forms, of which a

systematic account will be published later. Among the more

interesting of the known forms were fictularia plagiostoma

from a Palm-Civet, a number of species of Polydelphis from

various Pythons, Dicheilonema horrida from the South American

Ostrich, and Dictyocaulus filaria from the lungs of Sheep.

It was noticed that whereas intestinal parasites were almost

wholly collected from animals that had not lived in the Gardens

for more than six months, those of which the normal habitat and

food were the internal tissues of the host occurred in animals

that had been confined in the Gardens for several years. Thus,

an undescribed /ilaria was found in a Lemur after four years’,

and Filaria australis in a Wallaby after two and a half years’

captivity.

In all these cases the number of parasites obtained was small,

and could have had little or no effect upon the health of the

host. There was a remarkable preponderance of female forms.

From these observations it appeared that the change of food

and general conditions obtaining in the Gardens were unfavourable

to the continued existence of the intestinal parasites an animal

may harbour on its admission. The number of cases of auto-

and re-infection during captivity was strikingly small, and bore

testimony to the cleanly surroundings in which the animals were

kept. In four cases only was there evidence of the occurrence

of accumulative infection in the Gardens :—

1. A number of Giant Toads died from lung infection with

Rhabdias bufonis.

2. The Wolves appeared to be heavily infected with Ascaris

canis.

3. A Sheep died from pneumonic condition resulting from

an intense infection with Dictyocaulus filaria.

4. The Tortoises had Oxyuriasis.

Tn all these cases repeated infection undoubtedly had followed

ON A NEWLY BORN CUB OF THE MASKED PALM-CIVET. 621

from contamination of food and drink with feces containing

eggs of the parasite. ‘The infection could be eliminated by steam

sterilisation of the cages, or still more easily by changing the

species of animal living in the particular paddocks or cages, for

Helminthes were often peculiarly selective as regards their hosts,

and those flourishing in one animal sometimes found it impossible

to continue their life even in closely allied forms.

Mr. R. I. Pocock, F.R.S., F.LS., F.Z.S., Superintendent of

the Gardens, exhibited the newly born young of the Masked

Palm-Civet (Puradoxurus larvatus), which had been born in the

Gardens from a pair from Szechuen, presented to the Society by

Mr. Thurlow Lay, and remarked that, although the specimen had

died soon after birth, two other individuals composing the litter

were alive and likely to do well. This was the first occasion on

which the species had bred in the Gardens. The coloration of the

Text-fig. 147.

Inner aspect of abnormal left fore-leg of a newly born Masked Palm-Civet

Paradoxurus larvatus.

p, pad; s, strip of naked skin; ¢, claw.

young resembled in a general way that of the adult, but was of a

more generalised type, the black and white pattern of the head

being less emphasized and the general colour of the body greyer

with less yellow; the greater part of the tail and the lower portion

of the limbs were sooty grey, the throat, chest, axille, belly, and

the inside of the thighs being white. Of special interest was the

presence of a pair of ill-defined dark stripes on the back and of

very indistinct traces of pattern on the sides of the body. The

Proc. Zoou. Soc.—1911, No, XLIV. 44

622 LAND TORTOISES IN THE SEYCHELLES.

head, which had the eyes and ears closed, was shaped very much

like that of a wolf-pup. The tail was short-haired and tapering

and as long as the body from the fore part of the shoulders

backwards.

Special attention was drawn to a peculiar abnormality of the

left fore-leg (text-fig. 147). The humerus appeared to be of

normal length, but the lower arm was quite short, and there was

no distinct elbow-joint ; the paw, although freely articulated at

the wrist, was axially rotated outwards so that its plantar surface

looked inwards. It was furnished with a single large pad repre-

senting the large pad of the normal foot, and was armed with a

terminal claw, a thin strip of naked skin passing from the claw to

the pad.

[Note added July 17th, 1911.—The two specimens of the litter

that survived grew with great rapidity as compared with dogs and

cats, and almost equalled the size of their parents when three

months old. Their eyes, however, opened, as in the former

animals, about the ninth day from birth. |

Land Tortoises in the Seychelles.

Tue Secrerary read the following dispatch from the Governor

of the Seychelles, a copy of which had been kindly sent him by

the Secretary of State for the Colonies, for communication to the

Society.

Government House,

Seychelles,

1st June, 1910.

My Lorp,

In view of enquiries made from time to time regarding

the conservation of the breed of land tortoises of the islands in

the Indian Ocean, I have the honour to append some notes

condensed from the entries in the stud-book of the herd at

Government House, Mahé. This book was opened by me soon

after my arrival here in 1904, and contains records of the annual

measurements of the specimens under my observation and of

their habits.

2. In June 1904 I found a herd of 42 adult land tortoises, and

17 young ones hatched out in 1902 and 1903; these were duly

marked, numbered and measured, and the particulars entered up

in a new stud-book. The bulk of the herd had been purchased

for the Government in 1892 from the late Mr. Nageon de!’ Estang

of Val des Prés, a proprietor of ancient family in the district of

Anse Aux Pins, Mahé. The animals were then transferred to

Curieuse Island, the property of the Crown, and were brought

back to Government House in 1902.

3. In addition to this herd, there are two large males : No. 1.

‘“Gordon,” presented by the late General Gordon (of Khartoum)

when he was stationed in Seychelles in 1881 after his transfer

from the Cape; this is the largest land tortoise in Seychelles, and

measures over the surface of the carapace 4’ 94". and 4’ 8". The

LAND TORTOISES IN THE SEYCHELLES. 623

plastron measures 3’ 9x 2'9". This is undoubtedly Testudo

elephantina according to Dr. Giinther’s monograph. The next

largest, No. 2. “Spurs,” was presented by Mr. Spurs, now of

Kuropa Island, a French possession in the Mozambique Channel :

I am indebted for many of my most curious notes on the habits

of all varieties of sea turtles and land tortoises to this gentleman,

who is an educated man trained in habits of observation and has

spent all his life among the islands of the Indian Ocean. The

present dimensions of ‘‘Spurs” are 4’ 5” x 4’ 4” on the carapace

and 3' 7" x 2'5'' on the plastron: he has grown slowly in breadth

since 1904 but not in length. This remarkably fine specimen (the

finest which Mr. Spurs has seen) is not of the elephantina variety,

and resembles the Testudo daudinii of Dr. Giinther’s monograph.

These are probably the finest specimens living of their race. My

recollection is that the Testudo elephantina which died at Colombo

in 1900, of a recorded age of 155 years, was of larger dimensions.

The large specimen at St. Helena was measured by Admiral Sir

J. Durnford in 1907 as 4’ 6” “ fore and aft,” but it is not certain

whether this measurement included only the shell of the back.

4. The adult females, which are readily distinguishable in

shape, are smaller than the males. The largest specimen in the

collection (No. 5) reached its present dimensions—3’ 8” x 3’ 10"

along the carapace and 2’ 8” x 2' 1” along the plastron—in 1906,

and has not grown during the last three years. Several others

have rather smaller dimensions and have not grown since 1904.

The breeding season extends from January to April: the

females carry their eggs for about 10 weeks and Jay them in holes

dug out by their hind legs and then covered over. The eggs in

each nest vary in number from 9 to 25 and are white, round, and

of the size of a lawn tennis ball. There may be two nests made

annually by one female. Sea turtles lay a much greater number

of eggs, e. g., the green turtle 250 eges at a time, and the hawks-

bill turtle 100 to 150.

5. The young hatch out in about 120 to 130 days and work

their own way out of the ground. At the Government House

“pare aux tortues” about half of the eggs are unfertile ; but in

some years of drought, very few young ones appear, being unable

(probably) to work their way up. They grow fast if well fed, and

at four years old measure 1’ 6" to 1’ 10” in length and breadth of

carapace. It is said that they attain full growth in 25 years. It

is a local custom to mark off a young one at any birth in the

family and to eat it at the child’s wedding day. The meat is

palatable and the liver is held to be a delicacy.

6. The number of young ones secured from 1904 to 1909 was

168; they are liable to be destroyed by rats before their shells

harden. Ina wild state at Aldabra practically all the young are

destroyed by floventins (cranes), rats, and wild cats.

7. Through the courtesy of Admiral Sir John Durnford and

Captain Dumas, R.N., six specimens from this herd have been

presented to various institutions, e. g., Groot Schnur, Pretoria, and

the Zoological Society of London.

44*

624 LAND TORTOISES IN THE SEYCHELLES.

8. Owing to the difficulty in providing food for the increasing

herd in the enclosures at Government House, I have drafted off—

in March and May 1910—4 adult males and 18 adult females

and 27 young ones of the “ récoltes” in 1903, 1904 and 1905 to

Long Island, a Crown property used as a quarantine station,

where they are placed under the charge of the Guardians and

where there is an ample supply of food.

9. “ Gordon ” shows likes and dislikes and is rather combative,

having successfully bitten some visitors who presumed on his

apparent lethargy, but generally the land tortoise shows little

intelligence.

10. There have been no deaths among the adults during six years

in the enclosures at Government House, but one male has been

killed by a fall at Long Island, where they have shown them-

selves to be capable of swimming. ‘Theremains of tortoises found

in the pits in the coral formation of islands in the Aldabra Group

points to the falling into pits as one of the principal causes of

death. ‘They live apparently to an extreme old age—probably for

200 years. No plan will effectively prevent the final extinction of

these curious survivals in a wild state in their natural habitats.

The archives of Seychelles, Vol. i., published in 1909, are full of

references to their size and number in Mahé and Praslin, where

they were speedily destroyed by the early settlers. But their

future existence is guaranteed by the fact that they breed in

captivity and that several large herds besides that at Government

House are kept and well cared for. It isa guarantee for their

being taken care of that there isa sale for living specimens for

zoological collections.

11. The best book in English on the subject is a monograph

entitled “‘ Gigantic Land Tortoises” by Dr. Giinther, published for

the British Museum about 1878. The names of the best known

varieties of the larger tortoises and turtles are as follows :—

Of Aldabra: Testudo elephantina.

Of Galapagos: Testudo nigra.

Of Greece: Testudo greeca.

Box Tortoise of Madagascar: Pyxis arachnoides.

Box Tortoise of North America: Ci%studo carolina (Brer

tarapin).

Lettered Tortoise of North America: Hmys sculpta.

Green Turtle: Chelone mydas. (The edible variety much

consumed in Seychelles.)

Loggerhead Turtle: Thalassochelys caretta. (1 do not know

this species by sight.)

Hawksbill Turtle: Chelone imbricata. (The tortoise-shell

variety.)

I have, ete.,

Tre Rigur HonovuRABLE Sioned W.E r

Tur SECRETARY OF STATE Pouginen) = 1 DASTDSOW,

FOR THE COLONIES. Governor.

DR. H. B. FANTHAM AND MISS A. PORTER ON BEE-DISEASE. 625

A Bee-disease due to a Protozoal Parasite (Nosema apis).

Dr. H. B. Fanruam, F.Z.S., and Miss Anniz Porter, D.Sc.,

exhibited some diseased bees and combs infected with a minute

pathogenic protozoal parasite, apparently the same as Vosema

apis found by Zander in diseased bees in Bavaria. Microscopic

preparations and drawings of the parasite, Vosema apis, were also

shown, as well as healthy bees and combs in contrast. The

material exhibited was obtained from Cambridgeshire and

Hertfordshire in. March, 1911. Some of the infected combs

were brown in colour instead of the normal yellow (combs of the

same age being compared), while the infected bees suffered from

a sort of dry dysentery which rapidly proved fatal.

The pathogenic agent of this dry dysentery, Vosema apis,

formed thousands of minute spores which fouled the hive, while

infection was probably spread to new hives by hungry, weakly

bees attempting to enter healthy hives. The spores, about 2 to

3p by 4 to 6u, were the resistant and cross-infective stages of the

Protozoon. The parasite Vosema apis was closely allied to that

of pébrine, the silkworm disease due to Vosema bombycis.

The trophozoite and pansporoblast stages of the Vosema apis

had been observed in the gut-epithelium of the bee. Some spores

with polar filaments extruded had also been found. It was very

probable that the young, growing and multiplicative stages of the

parasite were capable of killing the bees before the formation of

spores had been attained, for dead bees were often found in which

only young stages of the parasite could be detected, occurring

especially in the chyle-stomach and intestine. Like V. bombycis,

the bee-parasite was possibly capable of hereditary infection, as

infected bee-larve and a dead infected queen had been found

and examined. Maassen had recently found infected drones in

Germany, but the infection in drones was stated to be limited to

the intestine.

That Nosema apis was fatal to bees and allied Hymenoptera

had been shown by the exhibitors by feeding healthy hive-bees,

mason-bees, and wasps with honey infected with Vosema spores ;

also by placing hive-bees dead of the disease among healthy hive-

and mason-bees and wasps, and by direct contamination of healthy

bees with infected fecal matter. In each case the insects ex-

perimented upon succumbed to the effects of Nosema apis.

In Nature the method of infection is probably contaminative,

healthy bees becoming infected by swallowing the spores of the

parasite.

It should be noted that the virulence of the parasite appeared

to vary in bees at different times of the year and in different

localities. Bad seasons are usually followed by increase of disease.

Some bees became chronics, forming reservoirs of spores and so

acting as parasite-carriers.

The only certain destructive agent of the Microsporidian

spores was fire, and all infected bees and hives, and any débris

therefrom should be most carefully burned.

626 MR. F. E. BEDDARD ON

In the opinion of the exhibitors, the Microsporidian parasite,

Nosema apis, had been responsible for much of the bee-disease

recorded in this country since 1906, especially in 1906, 1907, and

1911. The exhibitors first noticed the parasite in 1906 in

diseased bees obtained from the Isle of Wight ; its full significance

was grasped in 1907, but owing to the difficulty of obtaining

material the exhibitors’ results were not published. As much

attention was now being directed to ‘“ bee-disease,” the exhibitors

briefly recorded their observations. It was not asserted that

microsporidiosis was the only disease of bees current in Great

Britain at present, as Dr. Malden had investigated a bacillary

infection in bees, the parasite being called Bacillus pestiformis

apis. ‘¢ Foul brood” also was a well-known and separate disease.

Microsporidiosis (due to Mosema apis) had probably been

introduced from the Continent into British apiaries.

Other parasites found in bees—chiefly in the gut—by the

exhibitors were various species of Gregarines, a Flagellate

apparently belonging to the genus Crithidia, a new Ameeba

(Entameba apis) very like Hntameba coli of the human intestine,

a Spirochete, and various Fungi.

PAPERS.

29. Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea. By Frank H. Bepparp, M.A.,

E.R.S., F.Z.8., Prosector to the Society.

[Received and Read April 4th, 1911.)

(Text-figures 148-159.)

I. On some MAMMALIAN CESYOIDEA.

I propose to communicate to the Society from time to time

reports upon the species of Cestoidea which have been collected,

and are being at present collected, from animals which have

died in the Society’s Gardens. The collection in my hands is the

result of nearly two years’ examination of (necessarily) a great

number of animals, but does not contain as yet a very large

number of species, either of known forms or of those which

I believe to be undescribed. Tapeworms are by no means so

common as other parasitic worms, particularly Nematodes, which

are most abundant among the animals in the Gardens. Of

the forms which I have identified as belonging to well-known

species, I propose at some date to give a complete list, which will

be useful, not only as indicating the species which are most

abundant in the captive animals, but as extending the range of

hosts. At present I could hardly give a long enough list to

SOME MAMMALIAN TAPEWORMS. 627

warrant publication in the ‘ Proceedings. The new forms,

however, [ intend to describe, a few at a time, as soon as I

can give a sufficiently comprehensive account of their structure

to make such publication useful. The present communication

contains a fairly full anatomical account of three new species, of

Text-fig. 148.

Oochoristica sp., X 2.

two of which I have abundant examples for study. Of the third,

IT have been able to study two specimens only. But I have

been fortunate enough to fill up most of the descriptions which

are chiefly required to determine its systematic position with

accuracy.

On a Species of Oochoristica.

I vefer a number of individuals from the Lesser Anteater,

Tamandua tetradactyla, to this genus, for reasons which will be

apparent in the course of the following account of this Tapeworm,

and which are summed up at the end of the description in con-

sidering its affinities. I believe, however, that the present species

is to be regarded as a new one, though it undoubtedly comes very

near to Oochoristica wageneri, described by von Janicki from the

same Edentate four or five years since*. The reasons for this

* y. Janicki, “Studien an Satigetiercestoden,” Zeitschr. f. wiss. Zool. lxxxi.

1906, p. 5.

628 MR. F. E. BEDDARD ON

opinion will also be given later, after I have detailed the

characteristics of these worms.

As is the case with other species of Oochoristica, the scolex of

the present species is entirely unarmed and without rostellum.

I have studied this and other regions of the body not only by

inspection with a lens and the microscope of specimens in spirit

mounted entire in Canada balsam, but also by means of transverse

and longitudinal sections. I find that the rostellum is not

represented by the smallest vestige that can be recognised. The

anterior end of the head or scolex is simply slightly and uni-

formly convex, the convexity representing a large circle and, there-

fore, not to be described asa papilla. The suckers appear to stand

out rather from the scolex, and are somewhat long in shape with

a narrow and slit-like orifice.

The general aspect of the scolex itself does not recall that of

such other members of the genus Oochoristica as have been

Text-fig. 149.

Scolex and a few anterior proglottides of Oochoristica sp.

figured, excepting in its tetragonal form in section. It is, more-

over, rather unlike that of other Tetracotylea. When viewed

superficially with a lens, or under a low power of the microscope,

SOME MAMMALIAN TAPEWORMS. 629

the scolex appears to be of an oval contour, passing thus gradually

into a neck, which is at first narrow and later widens out. There

is nothing abnormal about the shape of such a scolex. A close

examination, however, shows that such a description would be

quite incorrect. For the scolex itself is in reality almost semi-

circular in outline, and its outlines are carried on on either side

by the neck of the worm, which is, to begin with, rather wider

even than the thickest part of the scolex. This is clearly to be

seen in text-fig. 149. It seems to be imbedded in the first seg-

ments of the strobila, like an egg in an egg-cup. Thus the true

scolex is very short. The widening of the strobila, for a space of

some five or six proglottides, suggests the commencing formation

of a ‘‘ pseudoscolex,” such as is further developed in, for example,

Thysanocephalum crispwm *.

The first steps in the formation of a pseudoscolex might well be

imagined to occur in some such way as is shown in the anterior

proglottides of the present species. There is, however, no further

indication of the formation of a pseudoscolex. The anterior pro-

glottides are precisely like those which follow, save in their

greater diameter. The figure also suggests that the scolex is

retractile within the first part of the strobila. In a sense this

is the case. Longitudinal sections through the scolex and the

commencing strobila show a bulging of the latter. There is a

plain demarcation between scolex and strobila, and not much

curving back of the anterior proglottides when thus viewed.

That the scolex itself may be extended to a greater length is

possible enough. And yet in seven or eight examples which I

examined there was an identity of structure in this and other

particulars. It is to be noted, therefore, that this species,

unlike Oochoristica tetragonocephala, has no neck, as, indeed,

was asserted of the latter by Diesing, but denied later by Liihe

and Janicki.

In examining the scolex mounted entire, after clearing with oil

of cloves or in alcohol previous to mounting, the suckers are by

no means so conspicuous as in other Tapeworms. The slit-lke

orifice can, indeed, be detected, but it is rather shrouded and dim.

The reason for this is apparent when the scolex is studied by

means of transverse sections, in which the relations to the head

ave shown. It will be there seen that, except for the small

orifice of the sucker, that organ is not by any means entirely

external in situation ; the sucker is, in fact, covered by a thickish

layer of the ordinary parenchyma of the body. It is thus imbedded

in the head, and represents a still further pronounced sessile

condition of the sucker, which (inter alia) distinguishes the

Tetracotylea from the Tetraphyllidia. At the same time, the

sucker retains its cup-like form, the two edges although in

* Cf. Benham in pt. iv. of ‘A Treatise on Zoology,’ ed. Sir E. Ray Lankester,

p. 121, fig. xvi. 2.

+ Unless, indeed, the anterior bulging part of the body of the worm is the neck.

But as it shows transverse furrows (text-fig. 149) I do not think that this is the case.

630 MR. F. E. BEDDARD ON

contact not uniting, thus leaving aslit-like orifice, which is dimly

seen when the scolex is viewed in its entirety.

The general aspect of the body is like that of other Tapeworms,

in that the anterior region is much narrower than the posterior

region, where the segments are in every way larger. In the

present worm this distinction is very marked, and the posterior

segments are rather more than oval in section, and approach

a circular contour. At the same time, this region of the body is

much stronger and thicker in every way than the anterior region,

so that there is a tendency for this part to become broken off.

There seems to be a rather sudden transition between the anterior

and the posterior regions. The posterior region of the body, in

fact, has quite the contour of an Earthworm, and entirely lacks the

flattened appearance of a typical Cestode. The cause of this

differentiation is doubtless to be sought in the swelling caused by

the dissemination of the ripe eggs through the medullary layer.

It is, in fact, in the posterior region that the uteri of the pro-

glottides have entirely disappeared, and are replaced by eggs

scattered singly through the parenchyma of the middle layer of

the body.

This state of affairs was, however, only to be noted as existing

in a marked fashion in the larger and, therefore, presumably more

fully mature examples of the worms. Fresh specimens reached

about 60 mm. in length, though in none did the breadth of the

body exceed 2 mm. Indeed, 1-1-5 was the prevalent diameter.

In smaller worms of 30-40 mm. length, the posterior pro-

glottides were by no means so swollen and remained comparatively

flat. So far as can be seen from an examination of the dozen or

so of specimens of this worm which I have in my possession, there

is no detaching of proglottides. In the first place, I always found

the terminal proglottid in its place, and to be distinguished from

the others by its oval posterior boundary interrupted by a large

orifice (apparently of the water vascular canals) highly suggestive

of a proctodceal invagination.

Secondly, the posterior set of segments were not materially

different in their form from those preceding them. In those cases

where proglottides are shed singly, it is common for them to

assume an oval form at the end of the body, and at least to be

a good deal constricted at their junctions with each other. There

is, commonly, also a very considerable increase of length of the

individual proglottides, which favours an easy rupture of their

connection. In none of these features do the terminal proglottides

of this Oochoristica agree with those of the genera which con-

stantly shed thei proglottides singly. Indeed, I have observed

that the five or six terminal proglottides are occasionally even

shorter than those which immediately lie in front of them. These

facts, coupled with the absence of a “neck” where new proglottides

ave formed, incline one to the belief that the growth of a scolex

is limited, and that the whole posterior region dies and liberates

the eggs. But of this I have no direct evidence. The relative

SOME MAMMALIAN TAPEWORMS. 631

length and-breadth of the proglottides differ in different regions

of the body and in individuals. A small example (30 mm. in

length) mounted in two pieces in Canada balsam showed the

following characters of strobilisation. The diameter of the wider

posterior region of the body was not more than three times that

of the narrowest part of the neck. 1 made out in all 167 strobila,

and there were not, I am convinced, many more. The first 42

were very short, being very much broader than long. After this

point the diameter of the worm became greater, and the segments,

therefore, larger; they appeared to be longer in proportion to

their breadth; but the first 31 of the series were, I believe,

not so. From this point, however, to the end of the body the

length of the segments was proportionately a little greater. But

throughout the series the actual breadth was really considerably

greater than the length. In a second specimen, of which the

head end only was mounted, the state of strobilisation presented

quite a different appearance. The first 45 segments or so were

quite as in the last specimen; but after this point the length

grew very rapidly in proportion to the breadth, and in a few seg-

ments no measurements were needed to assist the eye in seeing

that the proglottides were very much longer than broad. A care-

ful examination of these elongated segments showed that the

broader hind margin which overlapped the narrower anterior

section of the ensuing segment was regularly crumpled, showing

a series of regularly arranged projections, and in the middle of the

segment these were represented by alternate furrows and elevations.

This looks very much like the effect of a muscular pulling out

of a segment from end to end. But in any case this variability

in the proportions of individual proglottides shows that it is

a difficult matter to define Tapeworms very accurately by the

proportions of length and breadth of the proglottides in different

regions. The instances given would appear to be those of pro-

glottides which had been unduly lengthened through pulling

longwise. But I have observed instances of the contrary. In some

proglottides from the middle of the body there was a marked

transverse wrinkling, a kind of division of the segment into

annuli, which shows from a different point of view how difficult

it is to rely upon the relative dimensions of proglottides as aids to

the definition of a species or of a genus, as the case may be. It

is certainly quite unsafe in the case of this Oochoristica.

Before proceeding to the consideration of the internal organs,

there is one other external feature that requires description,

namely, the orifices of the generative ducts. ‘These are alternate,

but not regularly so. The openings are, however, as a rule, pre-

ponderatingly upon one side of the body. Thus in a portion of

one worm I found two pores consecutively upon one side of the

body. The next was on the opposite side; the five following

reverted to the first state, and the last examined showed again an

alternation. In another specimen, from another worm, there was

one pore on one side, followed by three consecutive pores on the

632 MR. F. E. BEDDARD ON

other side; then came two on the opposite side to these, and then

two on the opposite side to the two last described. These instances

will be sufficient to illustrate the general facts. It is quite

possible that the proglottid which first shows these openings may

vary from individual to individual. In any case a careful

examination of a small individual mounted in two pieces upon

a slide, and whose strobilisation has been already described above,

showed no generative ducts in front of proglottid 113 or so. It

will be remembered that this worm had a limited number of

proglottides altogether, there being not many, if any, more than

167. In this particular case it was not difficult to ascertain

accurately the point at which the ducts began. For, where

present, they were extremely conspicuous through staining, and

the cells composing the conjoined ducts formed a rather thick mass.

And I am, therefore, sure that they did not exist, except perhaps

in small rudiments, before the 113th proglottid. It was easy to

ascertain that the gonads themselves—at any rate, the ovaries

—existed anterior to this segment. Furthermore, these facts

ave still further supported by the condition of a second specimen

which had been cut into longitudinal sections up to about the same

point of the body. Here, too, the generative ducts did not exist

much anteriorly to the 113th segment—if, indeed, at all anteriorly.

The actual pores were only clearly visible upon more mature pro-

glottides, and were quite conspicuous round orifices near to the

anterior border of the proglottid.

The excretory vessels would seem from the figures of Janicki

to be very characteristic of the genus Oochoristica, for in the species

figured by him there are additional lateral vessels varying

according to the species. I have found the same thing in the

species of Oochoristica which I describe in the present paper.

The conditions, however, seem to vary somewhat in different

regions of the body. Anteriorly there are eight, longitudinally

running, which in the very anterior and therefore very thin

segments are almost in the same plane, and therefore can be seen

ina single longitudinal section. Further back the eight tubes are

differently arranged. There are two nearer to the middle line

and dorsal, according to Janicki*, and two ventral more widely

separated. The two lateral tubes on each side are less markedly

dorsal and ventral respectively. In posterior segments I could

see only six longitudinal vessels, as is represented in text-

figure 150, of which the four median were placed alternately

with regard to each other, and not as in earlier segments.

The gonads are visible in sections rather early in the body.

I found them without any trouble in the rather wider segments

which follow immediately upon the anterior sixty or so segments

which form the anterior section of the chain of proglottides.

A somewhat leaf-shaped mass of cells, the apex directed posteriorly,

and which represents presumably the yolk-gland as well as the

* Loe cit.

SOME MAMMALIAN TAPEWORMS. 633

ovaries, reaches from near to the anterior boundary of the pro-

glottid to beyond the middle of a segment. The éestes occupy

a restricted area in the posterior region of the segment, and

I observed in these young proglottides something like 20 or 30 of

them. They are not in contact with the ovaries, but, like them,

Text-fig. 150.

Transverse section of a posterior segment of Oochoristica sp., showing

scattered embryos embedded in general parenchyma.

e. Embryo. ¢. Longitudinal trunks of water-vessels.

are median in position. I occasionally observed also 2 or 3 testes

on one side of the ovary. Further back, when the genital ducts

first appear, the testes still form a mass which is chiefly posterior

634 MR..F, E. BEDDARD ON

to the ovary, but which has also to some extent grown round the

sides and lies laterally of the ovary.

Examining, as an entire object stained and preserved in balsam,

some segments at about the same stage of development as

those last referred to, it will be noticed that the small and

numerous testes occupy in their extension from side to side of

each proglottid about half of each proglottid. The lateral areas

not invaded by the testes are together about equal in diameter

to the middle region where the testes lie. The testes do not

reach as far as the lateral excretory vessels, and naturally, there-

fore, not to the nerve-cord.

In the mature proglottides the generative organs have been

studied by me chiefly by means of transverse sections. The

ovaries may be said to be double and le anteriorly ; they occupy

a good deal of the available space in the medullary region, and

are by no means confined to the middle of the proglottid. The

yolk-gland is single and lies behind the ovaries. The shell-gland

lies medianly in the segment on the opposite side from the ovaries.

I could find no reeeptaculum seminis, and the narrow vagina runs

a straight course for some way before its external opening.

The testes in the mature segments occupy the posterior region

of each proglottid ; but they also extend forward on either side up

to the level of the anterior border of the yolk-gland, and even

a little beyond, so that in transverse section some ovarian tissue

is occasionally seen in the same field with one or more testes. In

transverse sections, the male gonads are seen to occupy pretty

nearly the whole of the medullary layer of the worm, and are

frequently in contact with each other. Insegments of individuals

which have apparently been stretched during life the testes were

in a single row only dorsoventrally, or, at most, arranged here and

there in the form of a “ W,” one testis being slightly dorsal of

another. In such proglottides I never counted more than ten

testes in a single row. On the other hand, in proglottides that

were rather contracted than extended, the testes were frequently

for a considerable extent of the proglottid in two rows, one above

the other. In such cases as many as fourteen or even fifteen

testes could be seen in one transverse section. The testes are of

fair size, and in proglottides which are depressed occupy the

whole of the medulla from above downwards. In the other case

mentioned above, the medulla accommodated two testes one above

the other, and not much room was left unoccupied by these bodies.

In fully mature segments the testes are all of approximately the

same size, and there must be some fifty or so in a single proglottid.

Tt is easy to trace a thin membrane surrounding each testis, and

completely shutting it off from the parenchymatous tissue of the

medulla from neighbouring testes. This is the state of affairs

that we find in fully mature segments, in which, however, there is

as yet no scattering of the eggs in the parenchyma, such as occurs in

the terminal segments of the body. In a young specimen mounted

entire, the testes were quite plain in certain anterior segments of

SOME MAMMALIAN TAPEWORMS. 635

the body, wherein the genital ducts had not attained their full

development. In such a segment the testes are seen to be

distinctly more numerous than in such segments as have just

been described, a transverse section would show fully twenty

testes in a given plane (in the region of the proglottid where

they are most abundant). At the same time, these gonads

are much smaller. I have sections of proglottides which show an

intermediate state of affairs, and in which the posterior testes are

large and apparently full-er own, while anteriorly the testes are

very small and quite similar to those of immature or only in-

completely mature segments. These immature testes had no

membrane that I could detect. It appears to me that the advance

in growth of these gonads is achieved by the inclusion within a

common membrane of areas of testicular growth which have

separately originated and which in the very “youngest stages can

be seen to consist of a single cell only. It will be observed that

the disposition of the testes of this species is quite like that of

some other forms belonging to this genus as described by

Janicki *

Several fortunate sections have enabled me to see very clearly

the arrangement of the vas deferens and its mode of termination.

This tube is loosely coiled not far from its entry into the cirrus

sac. Before it enters the latter it passes in a straight course and

enters at the very extremity of the sac. It is noteworthy that

this duct is quite double the width of the vagina at its opening

into the genital cloaca and for some distance behind this point.

'The vas deferens when it enters the cirrus sac at once swells out

into a small vesicula seminalis, which thus lies within the cirrus

sac instead of outside as in most cases. The vesicula does not

by any means fill even the end of the cirrus sac in which it lies.

Immediately after this the duct narrows again, and again imme-

diately swells out into asmaller dilatation. After this comes the

narrow cirrus itself. The walls of the cirrus sae are loose and

muscular.

Liihe, the founder of this genus, in some notes upon the

anatomy of species of Oochoristica from Lizards (for example,

Tenia tuberculata of Rudolphi), remarks that the development of

the uterus must be extraordinarily rapid, as so often no inter-

mediate stages are obvious between a fully developed ovary and

the scattering of the mature eggs in the parenchyma. I quite

agree with Liihe, for the reason that the uterus is not always to

be found and is at least by no means so characteristic of this

particular Tapeworm as it is of many others. There are, how-

ever, stages to be observed. I have never seen more than a small

elongated sac lying near to either the ventral or the dorsal side of

the segment. In longitudinal sections this sac shows a tubular

form, amg 3 is, Indeed, so narrow a tube that on first observing it I

mistook it for one of the excretory vessels, and imagined that the

* Zeitschr. f. wiss. Zool. Ixxxi. 1906.

+ “ Oochoristica nov, gen. Teniadarum,” Zool, Anz. xxi. 1893, p. 650.

636 MR. F. E. BEDDARD ON

gonads were passed tothe exterior through those tubes. However,

the uterus is actually present in this form.

From the above account of the external characters and internal

anatomy of this worm we may deduce the following brief

resumé :—

Length 80-GO mm., diameter 1 mm.—2 mm. Posterior region of

the body markedly differentiated in fully adult examples by its

great thickness, but anterior end not so thin as in many forms.

Terminal segment with deep slit-like depression always present.

Necolex unarmed with rudimentary rostellum. Suckers unarmed

and sunk within the head, which is tetragonal in section. No neck,

segmentation beginning at once ; the anterior five or six proglottides

wider than scolex and seeming to form a hood into which it can be

partly retracted. Proglottides variable in proportions of length to

breadth. In many examples all of them, save the last few, broader

than long. In others some of anterior segments longer than broad.

Dorsal and ventral excretory canals not parallel, the two dorsal

being nearer together than the two ventral ; in addition to these, two

lateral narrower canals on each side. Cortical layer as thick as

medullary. Genital pores alternate regularly, the preponderance

being on one side, near to anterior border of proglottid. Testes

numerous, posterior to and at the sides of ovary. Ovaries in

anterior part of proglottid commencing shortly after anterior border.

Genital ducts pass between dorsal and ventral water tubes. Vas

deferens coiled; no obvious seminal receptacle or seminal vesicle.

Uterus sac-like, with branches ; ultimately disappearing, the eggs

being imbedded singly in the parenchyma. Occasionally traces of

uterus in form of sacs contaiung two or three ova persist.

Hab. Small intestine of Tamandua tetradactyla.

We may now consider the systematic position of this Cestoid.

The lateral position of the genital pores, the unarmed scolex, the

four suckers without appendages or hooks, the anterior position

of the cirrus pouch, the absence of more than a suggestion of a

pseudoscolex, show that this worm is to be referred to one of the

three families Anoplocephalide, Hymenolepidide, or Teeniadz,

of Ransom’s systematic table * (which is with slight differences

the same as that of Fuhrmann?’). From many of the numerous

genera contained in the first two of these families and the very

few genera of the last, the present worm is to be differentiated

by the following assemblage of characters :—scolex unarmed, no

neck; genital organs one set to each proglottid, with irregular

pores; testes numerous, posterior in position ; uterus disappears ;

eges imbedded singly in parenchyma.

The characters of the worm rather suggest the Anoplocephalide,

particularly, of course, the unarmed scolex and the absence of a

“neck.” But there is no genus in this family to which it can be

referred. The nearest is Zinstowia, in which the genital pores

are alternate, the eggs are imbedded singly, and the cortical layer

* Bull. U.S. Nat. Mus. no. 69, 1910.

+ Zool. Jahrb. extra vol. x. 1908.

SOME MAMMALIAN TAPEWORMS. 637

of the body is thick. But in Linstowia the testes extend through-

out the proglottid, the genital ducts pass ventrad of the excretory

vessels, and there are other differences. Of the Hymenolepide

the following genera only have an unarmed seolex and a single set

of generative organs with alternate pores, viz., Catenotenia, Oocho-

ristica, Rhabdometra, Anonchotenia, Metroliasthes, and Nemato-

tema. Nematotenia may be set aside as only showing strobi-

lisation posteriorly. In Anonchotenia the testes are anterior

and the eggs finally pass intoa paruterine organ. In Rhabdometra

and Metroliasthes the testes are posterior and at the sides of the

ovary, and the genital canals pass between the excretory vessels

as in the species which forms the subject of the present com-

munication ; but in those genera, as In Anonchotenia, there is a

paruterine organ into which the eggs finally pass. There remain,

therefore, by this process of exclusion only Catenotenia and

Oochoristica. 'The former genus must be eliminated, since the

genital ducts pass dorsad of both excretory tubes* and the uterus

is persistent. The present species is therefore to be referred to a

new genus or is to be included in Oochoristica. More recent

information about this latter genus than was available to Ransom

when he drew up his table shows—what is, indeed, also apparent

from that table—that the worm with which I have occupied my-

self is an Oochoristica and does not need a new genus for its

reception. Ransom’s definition is: ‘ Scolex unarmed, without

.rostellum. Single set of reproductive organs in each segment.

Genital pores irregularly alternate. Testicles numerous, sur-

rounding female glands posteriorly and on the sides. Vas

deferens coiled; seminal vesicle absent. Uterus breaks down

early and eggs become enclosed singly in egg capsules.” TI shall

now deal with the question of the species to which these worms

should be referred.

So far as I am aware, only two Tapeworms have been described

from the gut of Tamandua tetradactyla. The first of these is Tenia

tetragonocephala of Bremser, described by Diesing ft, and later, as

well as more fully, by Lihe {, whose description disagrees in

several particulars with that of Diesing. The most important

external disagreement concerns the scolex, which is represented

by Diesing as having no neck, while Liihe finds a neck 2 mm. long.

This is obviously a matter of some importance ; and I am inclined

therefore to regard the Tapeworm described here by myself as

not identical with 7’. tetragonocephala for that reason alone. With

the general anatomical description added by Lithe to Diesing’s

account the worms studied by myself fully agree, and are clearly

of the same genus which Janicki§ more recently has shown to

be Oochoristica. Janicki’s memoir contains also additional facts

* This point is not referred to by Ransom in his definition of Oochoristica ;

I have not accidentally omitted it.

+ Denkschr. k. Akad. Wien, xii. 1856.

t Arch. f. Naturg. 1895, p. 199.

§ Zeitschr. f. wiss. Zool. Ixxxi. 1906, p. 524. See also Zschokke, “ Das genus

Oochoristica,” ib. vol. \xxxiii.

Proc. Zoou, Soc.—1911, No. XLV. 45

638 MR. F. E. BEDDARD ON

upon the anatomy of O. tetragonocephala, which confirm my

opinion that the species examined by myself is not the same. In

particular, I refer tothe much more complicated excretory vessels,

which are illustrated by a text-figure *.

The second species inhabiting the gut of Tamandua tetradactyla

is Oochoristica wagenert of Janicki, who had, however, only a

single not very well-preserved example to work upon. Unfortun-

ately one very important point, as I think it, viz., the condition

of the neck, was not ascertained, and, as I understand, could not

be ascertained by Janicki from the imperfection of his specimen.

There are some other points in which my specimens were not

identical with that described by Janicki. He described ripe

proglottides as longer than unripe ones. But this is not always

the case in my specimens, since I have found quite early in the

strobila long segments. But the examination of other examples

might have led Janicki to alter this statement. For, as I have

myself shown, there is some variation in the form of the pro-

glottides in different regions of the body and in different examples.

T have noted in describing the excretory tubes certain differences

from the apparently regularly arranged six tubes found in O. wa-

genert; but here, again, it is possible that the examination of more

material would have shown that these differences do not exist.

On the other hand, I am disposed to see differences in the uterus

in the two forms. Janicki describes that organ in O. wageneri

as an irregular sac lying anteriorly in the segment and ex-

tending back as far as or beyond the ovaries, and gives a figure

showing this arrangement. The uterus in my examples was much

more irregular, and here and there, as I have described, were

quite tubular portions of it.

Even in the very last segment of the body, which might be

supposed to be fully mature, the uterus had by no means quite

disappeared in all specimens, although the majority of the eggs

were strewn through the parenchyma, as Janicki states to be the

case in his species, and as is characteristic of the genus. In other

respects I can find no differences between the facts as I read

them and Janicki’s descriptions. I do not, therefore, from a con-

sideration of all the facts, come to the conclusion that the present

species is certainly distinct from O. wageneri, but that equally

it is not certainly identical with it. Since we know that two

undoubtedly different species are found in the Edentate Tamandua,

there is no @ priori reason against the existence therein of a third

or even of more species of this genus. I prefer, therefore, in view

of these doubts, to give no name to the worm which is here

described.

Bertiella cercopitheci, sp. n.

An example of the Green Cercopitheque, Cercopithecus calli-

trichus, which died in the Society’s Gardens in February 1911,

contained two examples of a Tapeworm, which were found,

* Loc. cit. p. 535, fig. 5.

SOME MAMMALIAN TAPEWORMS. 639

contrary to what is more usual, in the colon and not attached

to the walls of that gut. It may be, therefore, that the parasites

had been loosened from their attachment in the small intestine

and had drifted into the colon, of which portion of the ali-

mentary tract they are therefore not really inhabitants. However,

both specimens were quite alive and exhibited writhing move-

ments, and it is possible therefore that they are really parasitic in

the colon. ‘They were of about the same size, and shrunk con-

siderably on preservation in alcohol. One of the two specimens,

which I have preserved entire and regard as the type of the

species, measures in the alcoholic condition 150 mm. It is re-

markable for the extreme shortness and great width of the

segments, and agrees in this particular with species of the genus

Bertiella, of which other anatomical characters prove it to be

a member.

The anterior extremity (as is shown in text-figure 151) is very

minute, and the body gradually widens up to a diameter of some

10 mm. Its general appearance is thus not precisely, although it

is generally, like that of other species of thisgenus. Accompany-

ing the increase in width of the proglottides there is also an

increase in thickness, and the posterior end of the body is about

2-3 mm. thick. In addition to the two specimens of the worm

there was a detached piece, possibly of one of these, of about an

inch in length. It appears to me that, as in Bothriocephalus, for

example, the proglottides are not shed singly but in groups, The

head of the worm is black in parts, the arrangement of the pigment

being peculiar, as I shall describe shortly. This black-headed

condition suggested to me that we might be dealing here with

examples of P. van Beneden’s Tenia melanocephala*, a parasite

from another species of African Monkey. The other characters

given by van Beneden are in perfect harmony with this view of

the identity of the species, but, as Blanchard 7 has pointed out,

the characters given are really not enough to determine the genus

to which Tenia melanocephala belongs, let alone the species.

Nor does my discovery here recorded of a black-headed Tape-

worm found in an African Monkey, and clearly referable to the

genus Fertiella, in any way settle the point at issue. For, in the

first place. I have found in a species of Dawainea (or, at any rate,

an allied genus) the same distribution of the pigment in the head

that will shortly be described in the species now under consider-

ation ; so that the mere presence of pigment in the head is clearly

no criterion of the identity of the worm. In the second place,

another species of Lertiella, viz. b. nvucronata, also from a monkey,

has been described $ in which the head is likewise pigmented. It

will be shown later that my species is not Bertiella mucronata.

Moreover, there is no reason, owing to the defective description

of van Beneden, for the identification of B. melanocephala with

B. mucronata. Thusit is necessary, as [ think, to give new name

* Mém. sur les Vers intestinaux, Paris, 1859, p. 162.

+ Mém. Soc. Zool. France, 1891, p. 186.

{ Meyner, “ Zwei neue Tzenien aus Affen,” Zeitschr. f. Naturw. 1895, p. 1.

45*

640 MR. F. E. BEDDARD ON

to the present species, in spite of the fact that it may prove

ultimately to be identical with Bertiella melanocephala.

Tn any ease, I shall be adding some further facts to the general

anatomy of the genus by the following account of this form.

Text-fig. 151.

Bertiella cercopitheci, nat. size.

A few of the posterior segments are shown more highly magnified.

The head is rather narrower than the ensuing body, in which

the proglottides are evident from the very first, there being thus no

neck. In the two specimens which I have examined, the suckers

presented different conditions. In one they were apparently

SOME MAMMALIAN TAPEWORMS. 641

absent altogether, being really retracted almost completely within

the head ; there were four little tags only protruding from the

anterior extremity. This scolex I have examined further by

transverse sections. The second worm showed only two cup-like

suckers, with their cavity directed upward as in other species of

Bertiella, One of these is distinctly larger than the other, and

Text-fig. 152.

Bertiella cercopithect.

Transverse section through scolex.

S. Sucker. P. Pigment-sheath of the same.

I presume that the remaining two were completely retracted

within the head. In a series of very nearly accurately transverse

sections through the head of the first individual, the earliest

sections showed four equidistant grooves lined by a continuation

of the thick cuticle of the scolex, which expanded towards the

642 MR. F. E, BEDDARD ON

middle of the interior of the scolex into oval chambers lined by

an equally thick cuticle. Further down the canals lost their con-

nection with the exterior and were T-shaped, with still a very

thick cuticular lining and no recognisable sucker structure. This

canal in each of the four quarters of the scolex continued for

some sections without any change, and then the suckers themselves

became plainly visible, being thus entirely retracted within the

scolex and removed from the exterior. The cavity of the suckers

is here triangular, with the sides closely pressed together. The

outlines of each sucker ave sharply marked off from the sur-

rounding tissue of the scolex by a layer of black pigment granules,

which pigment is also found in the central portion of the scolex.

The lining cuticle of the suckers is much thinner than that which

covers the scolex, and lines the canal of invagination leading to

the suckers. The outline of the scolex is here approximately

circular.

The generative organs are visible in an immature condition

very early in the body, though I have not made an accurate deter-

mination of the exact segment in which they first occur. At first

I could find no trace of any testes, simply a mass of generative

blastema which occupies the position of the future ovaries,

vitelline gland, and shell-gland, from which leads a solid rod hardly

narrower towards but not to the edge of the proglottid ; this latter

is, of course, the vagina, &e. This mass of tissue lies just anterior

to the transverse vessel, uniting the two ventral excretory vessels,

and is therefore some way from the posterior boundary of the

proglottid. In transverse sections through these very anterior

and immature proglottides it is seen to lie upon the ventral side of

the transverse vessels and to cross it obliquely to the dorsal side,

whence it passes towards the edge of the proglottid to the dorsal

side of both dorsal excretory tube and nerve. This is the same

on both sides of the body, the generative organs being single and

alternate in this worm.

Only a segment or two further back than those just described

the testes become visible, though, of course, at first quite immature.

They form a row generally only one deep (when viewed in longi-

tudinal horizontal section) extending from the excretory tubes of

one side of the body to those of the other. I counted about fifty

small testes in such a row. Here and there the row is two deep.

I could detect no trace of the vas deferens. The row of testes was

anterior to the rudimentary female organs. These latter are by

this time somewhat more developed. They still present, however,

a perfectly straight line, but reach very nearly to the edge of the

proglottid. There is, however, no external aperture. The fact

that the vagina is a tubular formation is beginning to be evident,

and the shell-gland, with radiating cells, in which it ends is plain ;

the ovary and vitelline gland lie below it and thus not in the

same plane with the vagina.

The female organs extend over more than a quarter, but not

quite a third, of the diameter of the proglottid. In still later

proglottides the uterus is for the first time quite visible and can

SOME MAMMALIAN TAPEWORMS. 643

Text-fig. 153.

Bertiella cercopitheci.

Sections through sperm-duct and vagina at three points near to their external

orifices, to illustrate relative position and structure.

C. Cirrus. Sp. Sperm-duct. Va. Vagina.

644 MR. F. E. BEDDARD ON

be traced for a considerable distance either way as a solid cord of

cells of a fibrous appearance, being more darkly stained than the

surrounding parenchyma. I could detect nolumen in this young

uterus. It runs straight along the ventral side of the row of

testes; arrived near to the lateral vessels, it bends at right angles

Text-fig. 154.

Transverse section through immature segment of Bertiella cercopitheci.

D. Dorsal water-vessel. O. Ovary. TT. Testes. U. Uterus.

V. Ventral water-vessel.

and runs straight dorsally to the inside of, and close to, the water-

vessels, crossing the vagina at right angles on its way. I could

not ascertain definitely the opening of the sperm-duct into the

cirrus sac which lay parallel with and dorsally to the vagina. Nor

could I see any branches running from the sperm-duct to the

SOME MAMMALIAN TAPEWORMS. 645

individual testes at any point. It is for this reason that I regard

the cord of cells as the beginning of the uterus and not asa sperm-

duct, which it undoubtedly suggests in its appearance and

position.

In mature segments, when both the vas deferens and the re-

ceptaculum seminis contain sperm, but when the uterus is still of

moderate dimensions only, the ¢estes are seen to extend right

across the body, when there is room for them, up to the lateral

excretory vessels on either side. In transverse sections they are

seen to be not more than two or possibly three deep in the middle

region of the proglottides. They are rather more dorsal than

ventral in position and are above the uterus. In the lateral

regions of the segments they are more crowded and often rather

closely pressed together. I could not find that the testes existed

outside of the medullary region of the segments, a point which

J mention particularly, since Meyner* met with testes (in

B. conferta) which had traversed gaps in the transverse mus-

culature and had taken up their position among the longitudinal

muscles.

The ovaries in the young proglottides are quite distinct and even

some way apart from each other, being joined by a thin bridge

which is the oviduct and which dilates in the middle into a round

sac marking the point of junction of the two oviducts. Even in

young stages the ovaries are divided into numerous digitiform

processes, which are not in the young stages thicker at their free

extremities. These processes radiate out from a common centre,

fanning out away from each ovary respectively. The ovaries are

ventral in position and radiate out towards the dorsal side of

the segment. In mature proglottides the ovaries are apparently

nearer together, that is, the connecting bridge is thicker and forms

a continuous junction of germinal tissue between them. The

processes of the ovary are now club-shaped, 7. e. thicker at their

free ends. Their arrangement is otherwise the same, but they

are much more numerous. In such proglottides the two ovaries

together occupy about one-quarter of the breadth of the body.

They are massed towards the pore side of the proglottid. The

double character thus remains distinct and is more marked than

is figured by Meyner for his species Bertiella mucronata and

B. conferta.

The vitelline glands also exhibit a double character and are to

some extent a copy of the ovaries in their general form and rela-

tions to each other. Each gland in the mature proglottides lies

a little above and between the ovaries. The dorsal position of the

vitelline glands with regard to the ovaries is shown by the fact

that the latter, when fully developed, extend dorsally at their

lateral margins and thus come to encircle the vitelline glands

lying between them. Hach vitelline gland is lobate, with numerous

oval lobes tending towards a club shape radiating out from a

* ‘Toc. cit. p. 93.

646 MR. F. E. BEDDARD ON

common centre, there being thus a likeness to the ovaries. The two

glands are likewise united by a bridge of tissue. Both the ovaries

and the vitelline glands are in close contact with the walls of the

receptaculum seminis. The shell-gland lies again rather dorsally

to the vitelline glands; but here the growth of those glands

laterally causes the shell-gland to lie rather between than above

the vitelline glands. It is in close relation with the median end

of the receptaculum seminis, which does not extend beyond it.

The sperm-duct in the mature proglottides has a form apparently

like that of other species of Bertiella parasitic in Monkeys, and is

not to be distinguished, so far as I can see, from that of Bertiella

mucronata. The sperm-duct is of considerable width from the very

first, 7. e. where it emerges from the cirrus sac. It is probable, how-

ever, that this region is really to be looked upon as representing

the vesicula seminalis of other Cestodes. A part of this dilated

sperm-duct lies actually within the cirrus sac as is depicted for

Bertiella polyorchis by v. Linstow. This portion, which is quite

short, is wider than the section which follows. The latter, however,

is also wide and lies in pretty regular coils of three alongside the

vagina as far as the commencement of the wide receptaculum

seminis. It is gorged with sperm and its walls are thin, but very

plainly recognisable by their dark staining. The coiling com-

mences directly after the emergence of the sperm-duct from the

cirrus sac, and we have therefore here a coiled region of the

sperm-duct which corresponds to that characteristic of many other

Tapeworms, but with the addition that it is the vesicula seminalis

part of the sperm-duct which is coiled.

The sperm-duct appears to come to abrupt conclusion at about

the commencement of the wide receptaculum seminis. But in

favourable sections it may be traced further as a very slender

tube closely adherent to the ventral wall of the receptaculum.

In the posterior segments of the body, which are distended

with ova, the sperm-ducts do not degenerate; on the contrary,

they are somewhat larger than in the first mature segment. They

contain, moreover, more sperm, which has somewhat distended

them: the walls thus appear thinner. The ducts in this region

of the body lie quite as coiled as in the more anterior proglottides ;

it might be supposed that they would be straightened out by the

tension caused by the enclosed spermatozoa. Nor has the sperm-

duct in any way shifted from the normal position, lying, as it does,

alongside of, and in close contact with, the vagina, which has under-

gone in this region of the body considerably greater alterations.

The same triple arrangement of the coils is visible—that is to say,

in a given transverse section there are usually three tubes to be

seen, this being, of course, the expression of the coiling.

The vagina of this species is specialised into several regions, as

it is shown to be in B. mucronata and B. conferta in the figures

of Meyner. The proglottides, from an examination of which I

have compiled the following description, appear to be in much

the same stage of sexual development as those figured by

SOME MAMMALIAN TAPEWORMS. 647

Dr. Meyner *, for which reason, of course, I have selected them.

The terminal region nearest the external pore has a thick muscular

sheath and the lumen is narrow. This section widens abruptly

Text-fig. 155.

Transverse section through mature segment of Bertiella cercopitheci.

n. Nerve-cord (only one of the three strands visible). O. Ovary. RS. Receptaculum

seminis. WV. Ventral water-vessel. Va. Vagina, to the left of which is seen

the dilated and coiled sperm-duct.

* Loe. cit. pl. i. fig. 3, pl. il. fig. 9.

648 MR. F. E. BEDDARD ON

into a thin-walled and much wider section, the muscular walls of

which are not more than, if so much as, half of the diameter of

those of the previous section. The transition is not regular, as

is figured by Meyner in the two species examined by him. The

diminution in thickness of the muscular layer, however, is

rather gradual at the orifice of the terminal part of the vagina

into the middle region.

This middle region of the vagina contained sperm in the

mature segments. It apparently ends abruptly on the median

side, but is really connected by a very narrow tube, not wider than

the sperm-duct, with the distal section of the vagina, which may

be termed the receptaculum seminis. This section is twice the

width of the last, and into it opens the duct from the ovaries &c.,

which is about as wide as the intermediate passage connecting

the receptaculum and the vagina. The receptaculum seminis

thus begins and ends abruptly. It contains sperm, as does the

distal section of the vagina. At the orifice into the genital cloaca

the sperm-duct and the vagina lie obliquely with reference to each

other. The sperm-duct is anterior, but also dorsal to the vagina,

and a little further back is completely dorsal to it. In the very

posterior segments, which are otherwise filled with the distended

uterus, the vagina with its receptaculum seminis shows an alter-

ation. It has increased in size, owing to its being gorged with

sperm.

I presume, at any rate, that the granular, in places fibrous-

looking, contents of the vagina are sperm, though they have a

different appearance from, and stain differently to, the obvious

spermatozoa which fill the neighbouring vesicula seminalis. In

the posterior proglottides the vagina shows no differentiation into

regions such as have just been described, excepting the proximal

muscular region near to the external pore. The rest formsa wide

uniform tube, and in longitudinal horizontal sections is seen to be

as wide as the proglottid is long or nearly so. In exceptional

cases even this amount of differentiation in the vagina is lost and

the wide tube pushes itself as far as the external pore, crushing

the cirrus sac into a mere rudiment.

From the above detailed description of this Tapeworm there

may be abstracted the following réswmé of its characters :—

Length of about 150 mm., greatest breadth 10 mm. Shape an

elongated cone, gradually diminishing to scolex. No neck, the

strobilisation commencing immediately after scolex. Proglottides very

short and wide, not increasing posteriorly in length. Scolex with

four suckers looking wpwards, completely retractile into head, with

black pigment. No armed rostellum. Dorsal and ventral excretory

tubes at first subequal, later the ventral very much the larger ; the

two tubes are superposed dorso-ventrally, with a transverse trunk

connecting the two ventrals. Genital pores alternate, frequently

with regularity. Generative ducts dorsal to water-vessels and nerve.

Testes numerous, forming a layer two or three thick, reaching com-

pletely between water-vessels, anterior and dorsal in position.

Cirrus feeble, no sperm-sac (vesicula seminalis). Sperm-ducts

SOME MAMMALIAN TAPEWORMS. 649

open to side and in front of vagina and pass back along ventral

margin of rows of testes. In fully mature segments sperm-ducts

increase in volume. Ovaries double, ventral and posterior in posi-

tion, in front of transverse excretory vessel, formed of nwmerous

club-shaped masses radiating from common centre. Vagina

unusually wide, showing a marked receptaculum seminis in less

fully mature proglottides ; the width increases enormously in the

fully mature proglottides until it is nearly as wide as the proglottid

is long. Genital cloaca short. Uterus single, sac-like, without

definite outgrowths, but somewhat irregular in form, filling most of

the proglottid. The eggs without V-shaped apparatus, with a

thinner inner shell and a very wide and lax outer membrane.

Proglottides apparently not detached singly, but in groups.

Hab. Cercopithecus callitrichus.

It is clear from this definition that the Tapeworm now under

consideration has been rightly referred by me to the genus

Bertiella. There is, I think, no other genus which shows the

same assemblage of characters as those which I have just set forth

in brief. It remains to be seen what position the species occupies

within the genus, of which we now know a good many different

species.

It seems that we may select the following characters as

distinctive of this species, which will be thus definable :—

BERTIELLA CERCOPITHECI, Sp. n.

Length 150 mm. or more, breadth 10 mm. posteriorly, gradually

tapering to head. Scolew with black pigment scattered throughout

middle of scolexw and forming a special layer round each sucker.

Suckers directed anteriorly, and completely retractile. No neck,

strobilisation commencing at once. Lateral nerve-cord divided into

three, the middle one the largest. Lateral water-vessels superposed,

the dorsal much the smaller, Testes extending between water-vessels,

forming a layer two or three deep and three or four wide in middle

region of proylottid, more numerous laterally. WVesicula seminalis

very long and coiled, extending into cirrus sac. Vagina divisible into

two well-marked regions and communicating by very narrow interval

with wide receptaculum semis. Hggs without pyriform apparatus.

Hab. Cercopithecus callitrichus, in colon (2).

Following the subdivision of the species of Bertiella by

Zschokke * into three groups, the present species evidently

belongs to the first group, to which Zschokke assigned Tania

mucronata and 7. conferta of Meynery. Bertia studeri and

B. satyri of Blanchard} are doubtfully admitted by Braun § into

the genus Bertiella as recognised by the better-known species fully

described by Meyner, for Blanchard does not describe the genital

* “Neue Studien an Cestoden aplacentaler Satigethiere,” Zeitschr. wiss. Zool.

Ixv. 1897, p. 404.

+ “Zwei neue Tenien aus Affen,” Zeitschr. f. Naturw. 1895, p. 1.

¢ Mém. Soc. Zool. France, 1891, p. 186.

§ Bronn’s Klassen u. Ordn. des Thierreich, iv. Abth. 18, p. 1712.

650 MR. F, E. BEDDARD ON

organs, except in so far as to remark upon their alternating pores

and upon the structure of the ova. Nor does he say anything of

the pigmentation of the head, which is so marked a feature of my

species and of the otherwise unrecognisable Tania melanocephala

of van Beneden *.

B. mucronata was described from an American Ape, JJycetes

niger, and I gather from Meyner’s description that the Tapeworms

themselves were brought from Paraguay, and thus real inhabitants

of the Monkey in question, which, therefore, had not been

infected in a menagerie. This species shows the same pigmenta-

tion of the head as mine, and its general shape? is not very

different. Nevertheless, it appears to me that in the species

described in the present paper the anterior end is more tapering,

and has a very small diameter for a longer stretch. Indeed, the

general outline of the worm described in the present paper is

more like Meyner’s B. confertat, which is, moreover, a parasite

of an Old World Monkey, Macacus radiatus, and therefore,

perhaps, more likely on @ priori grounds to be identical with mine.

I think, however, that B. conferta may be put out of court in this

comparison, for the reason that no mention is made of a black

pigmentation in the scolex, which is so clear a feature of my

species and of B. mucronata. Nevertheless, 5. conferta agrees

with my species and differs from 4. mucronata in that the vagina

enters the receptaculum suddenly, there being no gradual widening

as in B. mucronata, where the tubes are continuous. The suckers

are said, however, to be lateral in position, as in B. mucronata §.

This is one of the principal reasons which lead me to regard my

species as distinct. Of B. mucronata Meyner writes :—“ Sind

nicht vollstiindig nach vorn, sondern mehr zur Seite gerechtet.”

In a series of transverse sections through B. cercopitheci, which I

have described above, it is very clear that the apertures of the

suckers are not lateral nor slit along the retracted sucker, such as

is so obvious in the other species described in the present paper.

Nor does Meyner describe the remarkable pigment-sheath to

each sucker which characterises my species. Furthermore, it

appears that B. mucronata has a “neck”; for Meyner writes ||:—

“ Schon etwa 0:9 mm. vom Skolexscheitel entfernt, also in einem

Stadium, wo der Kérper noch keine Spur einer Gliederung

erkennen lisst, gruppiren sich,” etc. Stiles, in reviewing 4 the

characteristics of this species, points out that B. mueronata is

also to be defined by the fact that the generative canal passes to

the exterior between the nerve-cord on the one hand and the

two excretory vessels on the other. My sections of B. cercopitheci

show very plainly that the generative canal passes dorsally to

both nerve and lateral excretory vessels. This is particularly

* Mém. sur les Vers intestinaux, Paris, 1859.

+ Loe. cit. pl. 1. fig. 1. t Loe. cit. pl. ii. fig. 8.

§ In other Bertiella (e.g. in Sluiter’s B. plastica, see Centralbl. Bakt. xix. 1896,

p- 941) the suckers look upwards.

|| Loe, cit. p. 81. € Proc. U.S. Nat. Mus. 1896, p. 145.

SOME MAMMALIAN TAPEWORMS. 651

obvious in young proglottides, which are so slender that the

disposition of these structures can be seen in a single section.

A final point of systematic importance concerns the horn-like

processes of the innermost egg-shell, which are figured by Meyner

and stated by him to be always very obvious. I have been abso-

lutely unable to detect these structures, even when using an oil

immersion-lens (4 in. Leitz). We must, therefore, agree that

this group of species of Bertiella is to be characterised, as the

two remaining groups into which Zschokke divides the genus, by

‘“birnformiger Apparat nicht constant.” Iam unable to compare

the species which I describe here as Bertiella cercopitheci with two

species deseribed by Gottheil * from Macacus cynomolgus and from

that species and Cebus capucinus, since they are not sufficiently

diagnosed, and Stiles is of opinion that they are only doubtfully to

be referred to the genus Dertiella. Indeed, the position of the

genital pores is not referred to.

Another Monkey parasite with which my Tapeworm might be

compared is v. Linstow’s Bertiella polyorchis + from Macacus cyno-

molgus. This is interesting, from the point of view of the ege-

shells, and confirms what I have said above concerning the absence

of the horn-like processes in 4. cercopitheci ; for it is hardly likely

that so experienced a helminthologist as Dr. von Linstow would

have overlooked these structures were they present, and his figure

of the egg of 4. polyorchis does not show them. Furthermore,

the suckers look forward and a dilation of the sperm-duct within

the cirrus sac is figured. The species, however, differs, as I

believe, from Lertiella cercopitheci by its less complicated vagina,

by the much greater number of testes which fill the middle of the

proglottid, and by the absence of any black pigmentation in the

head. At any rate, the latter point is not referred to.

Thysanosoma gambianum, sp. n.

I obtained from an example of the Gambian Pouched Rat,

Cricetomys gambianus, which died in June 1909, a considerable

number of Tapeworms which I regard as being of a new species

and belonging most probably to the family Anoplocephalide, The

Rodent had been one year and ten months in the Society’s

Gardens before its death, and it is, therefore, quite credible that

it was infected with these Cestodes when it arrived in London.

The material, as well as being abundant, was well preserved, and I

am therefore able to give a fairly comprehensive account of the

anatomy of the species, which presents certain new combinations

of characters.

The external characters alone appear to place this worm in

either the genus Anoplocephala or Zschokkeella. There are no

other genera in which the scolex is unarmed, the genital pores

are unilateral and the neck is absent, and the segments until the

* Journ. Comp. Med. & Surgery, 1887, vol. vii. The species are not named and

are referred to Tenia.

+ Arch. f. Naturg. xxi. 1905, p. 270,

652 MR. F. E. BEDDARD ON

very end of the body are broader than long. Unilateral genital

pores occur only in these two genera among the subfamily or

family Anoplocephalide or Anoplocephaline. There are, however,

several genera among the remaining Tetracotylea which possess

genital pores in sequence upon one side only. But the number

of genera is small and other external characters prevent a con-

fusion. Thus the Tapeworm which forms the subject of the

present section cannot be confused with Aploparaksis, Diorchis,

Gryporhynchus, Paruterina, Culcitella, Oligorchis, Lateriporus,

Dilepis, Davainea, Progynia, Idiogenes, Chapmannia, Pro-

orchida, Monopylidiwm, or Cyclorchida, since all of these have

a circle or more than one circle of hooks upon the rostellum.

There remains Hymenolepis, which is to be distinguished by

Text-fig. 156.

Thysanosoma gambianum, nat. size.

To the right are a few segments more highly magnified to show genital papilla.

possessing a “neck,” which the present species does not, and by

the limited series of testes in each proglottid. I am thus correct,

as I believe, in regarding this worm as a member of the Anoplo-

cephalide. ‘

There are reasons both for and against referring the Tape-

worm from the Gambian Pouched Rat to either of these genera

or to a new genus, into which I shall enter at length after

detailing its anatomy. This species isa large worm, measuring

SOME MAMMALIAN TAPEWORMS. 653

some 6 inches in length. A very marked characteristic is the

fact that the segments are wider than long throughout almost the

whole of the body. It is only at the extreme end that they

become longer. The segments overlap successively, so that it is

possible to detect the anterior margin of each segment. The

worm is not very stout, but flattened after the typical Tapeworm

fashion ; the most posterior segments alone being rather thicker

in a dorso-ventral direction. The diameter of the body in the

middle is some 6 mm. There is no marked colour, though, as will

be seen presently, there is some. internal pigmentation. The

general character of the segmentation suggests that the pro-

glottides are not shed.

This species like Anoplocephala has no “ neck,” as it is termed,

that is to say, there is no undifferentiated zone following the scolex,

Text-fig. 157.

Scolex of Thysanosoma gambianum.

where the limits of the proglottides are unrecognisable. The

definite proglottides are obvious from the very first. They are

at first rather narrower than they become posteriorly; but the

diameter of the body very soon attains to its full dimensions.

Proc, Zoou, Soc.—1911, No. XLVI. 46

654 MR. F, E, BEDDARD ON

The cirrus of each segment is not visible, or at least not con-

spicuous, when the worm is viewed with a lens, in a protruded

condition. This is due to the small size of these organs, which

will be more fully described later. I have not found it possible

to differentiate the ventral from the dorsal surface. The scolex, as

already stated, is unarmed. I investigated this part of the worm

by the section method, and am therefore able to state positively

that there is no rostellum nor any trace of hooks. Nor do the

suckers show any hooks that I could detect. I am convinced

that they are absent altogether from the scolex of this worm. A

regular series of transverse sections across the “ head ” shows a

rostellar region above the region of the suckers, which is oblong

in section ; it contains no pit or depression of any kind that was

obvious to me. The suckers are the usual four and as usual

symmetrical. When the scolex is viewed in its entirety under a

low power of the microscope the suckers seem to face rather

upwards ; but they do not lie on the upper side of the head as in

some Tetracotylea. There is only a slight obliquity. The orifice

of the sucker in such a preparation is very plain and rounded.

There is no appearance of a slit-like orifice such as is described

in the present paper in Oochoristica, and certain details in the

minute structure of the suckers in these two genera show corre-

sponding differences. I find in two series of transverse

sections of the suckers of the two Tapeworms, that in the present

species eleven or twelve sections displayed the entire sucker,

of which only five showed its cup-like orifice; in Oochoristica,

on the other hand, the entire sucker required seventeen sections

of equal thickness for its display, of which twelve or thirteen

showed the obviously more slit-like orifice.

As in Oochoristica the orifice of the external integument is

much more limited than that of the sucker itself. The two coin-

cided for only the space of two sections, rather less than in

Oochoristica. ‘This, however, I take to be simply a measure of the

state of contraction of the suckers. It is also important to notice

that the outer integument is grooved for a little space anteriorly

to the appearance of the sucker itself, which suggests a con-

firmation of a view already put forward, and that is the inde-

pendence of the suckers of the more specialised Tapeworms from

the bothria of the Dibothriata. It appears to be possible that

this grooving is the persistent trace of the bothrium, upon which

the sucker has been added as a subsequent development.

In series of transverse sections through the head there is

another feature of the suckers which deserves mention. It is

very clear from such sections that the growth of the sucker

has taken place in a definite direction.

Although I am unable to give histological details, it is certainly

the fact that anteriorly the sucker is in organic connection with

the general tissues of the head. ‘There is no break; it begins

gradually with a recognisable condensation of tissue marked by

its deeper staining with carmine. On the other hand, at the

SOME MAMMALIAN TAPEWORMS, 655

posterior end of the sucker its tissues come to an abrupt end, and

I invariably found a shallow cavity between the sucker and the

medullary tissue of the head. This suggests that the original

position of the sucker is more apical and that it is directed

upwards—a, state of affairs which is actually seen to persist in

many species of Tetracotylea.

The orifices of the generative organs are, as already stated,

completely unilateral. I have examined pieces of the worm

mounted whole, and many series of sections, and find that in

every case there is a regular sequence of the apertures which lie

upon one side of the body*. These orifices are not conspicuous

on examination with a lens, as they are in many Tapeworms, and

for two reasons. In the first place, the orifices are actually small,

and in the second place, I have never observed thecirrus to protrude.

Thus it is necessary to make a microscopical examination in order

to detect the genital pores. Apart from microscopic sections,

where, of course, it is easy to find the pores, I have only observed

them (that is, in pieces of the worm mounted entire) in the wider

posterior segments. Here they are seen to lie at just about the

middle of the proglottides. The orifices project slightly and are

quite circular.

The excretory system of this Tapeworm presents certain unusual

features. In transverse sections through some of the middle

segments of the body there are four longitudinal canals perfectly

obvious in the body and no more than four, unlike, for instance,

the genus Oochoristica, with which the present genus has some

characters in common. These four tubes are lateral in position,

and are all of them at about the same level. It is thus impossible

by position to distinguish the dorsal from the ventral excretory

canals. The two on each side are at a considerable distance apart.

The distance which separates the outer from the inner of the two

is about as great as that which separates the outer tube from the

nearest margin of the body. The tubes can, however, be dif-

ferentiated by their size. The outermost of the two excretory

canals is not more than one-third of the diameter of the inner-

most tube. The form is not, however, so small that it can be

missed in transverse sections. In the very anterior segments I

have noticed a third lateral vessel on each side; and in this region

of the body, moreover, the two main tubes have a greater incli-

nation respectively to the dorsal or ventral side. There is an

abundant plexus of vessels connected with these and traversing

the proglottides, but 1 am unable to give details. I think,

however, that I have seen numerous external pores.

The testes have a somewhat peculiar arrangement, which is

distinctive of this worm. The bulk of these very numerous

gonads lie upon that side of the body upon which the generative

* T cannot determine whether this is right or left with certainty, since in trans-

verse sections the gonads are not definitely dorsal or ventral in position, 7. e. nearer

to one surface of the proglottid or the other, while the dorsal and ventral excretory

tubes are parallel to each other and lie in the same plane with the nerve-cord.

46*

656 MR. F. E. BEDDARD ON

pores are not situated. They are naturally in the medullary part

of the body, and in transverse sections are seen to occupy the

greater part of this, being neither dorsal nor ventral in position,

but simply central. They reach towards the margin of the

segment which is nearest to them, far beyond the outermost of

the two excretory tubes of their side of the body—ain fact, up to

the nerve-cord ; medianly they do not reach the median line of

the body, but extend beyond the innermost of the two excretory

tubes. They occupy perhaps, when seen in this view, one-fourth

of the entire breadth of a proglottid. They are not very close

together and are, at most, in two rows dorso-ventrally, there being

thirteen or fourteen to each row, and thus something like thirty

may be visible in a single section. In longitudinal sections seven

or eight of these double or partly single rows are seen, and they

may be observed to occupy the greater part of the segment.

There is also a second set of testes at the opposite margin of the

proglottid, very much fewer in number—perhaps four or five in a

transverse row, which lie on either side of the outer excretory

tube.

The ovary may be regarded as a paired structure, and the two

are partly separated by the yolk-gland, which lies between and

behind them. In transverse sections through the ovaries, before

the other parts of the female reproductive system have been

reached, each ovary is seen to be distinct from its fellow and to

lie on either side and below the larger of the two excretory tubes,

which, in such a section, is seen to be not median in position.

The excretory vessel is here only just below the cortical layer of

the proglottid, though actually in the medullary layer. The two

ovaries are not quite in contact below the vessel, or, at any rate,

they can be recognised as two distinct bodies of a bushy form not

unlike that which has been figured in other Tapeworms. Within

a few sections from that which has served as the basis of the

above description the shell-gland is seen taking the place of part

of that ovary which lies median of, 2.e. not to the pore side of,

the excretory vessel, and the oviduct leading from it to the ovary

of that side is conspicuous. In a section or so further on in the

series the yolk-gland appears; although, as already said, this

gland divides the two ovaries, it does not lie symmetrically with

reference to the excretory tube. It lies almost entirely on the

median side of this tube but also below it, though it does not

extend at all on to the pore side of the excretory vessel. It

occupies nearly the whole of the space on the inner median side

of the excretory tube that, in previous sections, is occupied by the

ovary of that side. In the next proglottid to that whose ovaries

and associated glands have just been described, the ovary to the

outside reached nearer to the outermost and smaller of the two

laterally placed excretory tubes, but without arriving at it. In

this case it is important to notice that there was no accessory

group of testes lying between the ovary and the outermost of

the two excretory vessels such as occur in the proglottid first

SOME MAMMALIAN TAPEWORMS. 657

described, and which are referred to elsewhere in the description

of this species. Series of longitudinal (sagittal and horizontal)

sections brought out the position of the ovaries and their relation-

ship to adjacent glands still more clearly. Such sections also

showed that the gonads are quite in the middle of the proglottid,

2.e. dorso-ventrally. It should be remarked that the double

character of the ovary was not always so strongly marked, and

that it sometimes lay entirely upon the pore side of the more

median (and larger) excretory vessel. Furthermore, I have also

observed the yolk-gland to lie upon the median side of the larger

excretory tube as well as upon the pore side. There is thus some

variability.

It is, perhaps, noteworthy that the gonads of the pore side,

which consist of the ovary, yolk-gland, and the testes, together

occupy about the same space as the gonad and testes only, in this

case, of the opposite side of the proglottid. The female gonads, at

any rate, are visible very early in the body, within twenty seg-

ments, I dare say, of the head. Shortly after their commencement

the mass of tissue which is to form ovary, yolk-gland, and, as I

presume, oviduct, vagina, &e., is seen very plainly to extend to-

wards the periphery of the proglottid between the two longitudinal

excretory vessels, a feature of systematic importance in this group

which it is necessary to note. In the sexually mature proglottides

the oviduct runs straight from the ovary as a thin-walled delicate

tube not easy to see, which is sometimes wider, at least near to

its termination, in the ejaculatory apparatus, runs to near the

distal termination, and then suddenly narrows into a very fine

tube with thick, darkly-stained walls which, after a very short

course, again suddenly widens into a long sac which is as wide as

the cirrus sac and runs beside it and below in position. This

terminal reservoir is laxly surrounded by what appears to be an

adventitious sheath of muscular fibres, which have a circular

direction and thus appear cut across; they are very obvious

through their deep staining. This layer of muscles was some-

times, but not always, observed to commence with the very thin

region of the vagina. The wide terminal region of the vagina

bends towards the cirrus sac and again becomes narrow, opening

in common with the latter into the genital cloaca, which is very

short. This genital cloaca is quite distinct from an ingrowth of

the outer layer of the body which meets it and forms the actual

pore; this has been described in considering the external

characters. The cirrus sac, as already implied, is neither wide nor

long. The cirrus was never seen in a protruded condition, but

always lay a darkly staining rod within the pouch. The vas

deferens, directly it leaves the cirrus sac, is thrown into a large

and complicated coil, which reaches for a considerable distance

into the interior of the body. I think that it ends in a dilated

vesicula seminalis above the ovary; there is, in any case, a dilated

pouch in this region full of sperm, which does not appear to have

anything to do with the female ducts.

658 MR. F. E. BEDDARD ON

Tn sections through the posterior segment of the body, the ripe

eges, with their sacs, are seen to occupy the whole of the available

space, that being, of course, the medullary region of the proglottid.

The segments are, in fact, stuffed full of eggs, and, in accordance

with this, their dorso-ventral diameter has somewhat increased,

though not to so very great an extent as in some other Tapeworms,

for instance, in certain specimens of the Oochoristica described in

the present paper. A closer examination of the eggs shows that

they are imbedded a few together in a dense and darkly staining

Text-fig. 158.

Transverse section through proglottid of Thysanosoma gambianum to show

numerous paruterine organs (e). ¢. Water-vessels.

mass of tissue, which closely invests them. These sacs appear to

me to be, without doubt, the equivalents of the paruterine organs

of many Cestoidea. They are not precisely sacs, in that there is

no central lumen occupied by the eggs; they are rather concen-

trations of the medullary tissue round a series of eggs. These

bodies are of approximately equal size and contain much the same

number of eggs or, rather, embryos. There is no question here

of a circle of paruterine organs surrounding a centrally placed

SOME MAMMALIAN 'TAPEWORMS. 659

uterus. The uterus has entirely vanished, and the paruterine

organs are imbedded in the tissues of the body.

Text-fig. 159.

Paruterine organs of Thysanosoma gambianum more highly magnified.

S. Walls of paruterine sacs. O. Embryos.

So much for the condition in the fully ripe proglottides. Harlier

in the body this formation of paruterine sacs is preceded by a

uterus which is not very conspicuous, and consists of not much

more than a transversely running tube extending nearly right

across the proglottid in which the eggs occur, but with which the

uterus never appears to be stuffed. I could find no outgrowths

of this centrally placed uterus, and there was certainly nothing

in the nature of a reticular formation of its cavity. Gradually

the cavity of the uterus appeared, as it were, to dry up and the

eggs were found—to continue the simile—stranded in the tissue of

the body. At this time the formation of the paruterine sacs

became visible. Round each egg, or round two or three, as the case

might be, the tissue of the medullary region of the proglottid

became denser, this being shown by its darker staining. There

was thus a concentration of tissue round the ova. This concen-

tration of tissue had no relations that I could detect with the

660 ON SOME MAMMALIAN TAPEWORMS.

uterus. There was nothing like the formation of diverticula of

the latter walled by the condensed parenchyma.

The characters of this worm may be briefly summed up as

follows :—

Scolex unarmed, with no rostellum, only a raised area. Strobila

commencing without an intermediate neck. Proglottides much

wider than long, except at the very end of the body. Proglottides

very numerous, the length of worm being some 6 inches with a

greatest diameter of 6 mm. Genital pores unilateral. Eacretory

tubes four, parallel to each other. Cortical layer of body as thick

as medullary. Testes chiefly massed upon the side of the body

Surthest from the genital pore, very numerous, median in position.

In addition, a small number of similar testes on either side of the

outermost excretory vessel of the pore side of the proglottid. Vas

deferens with a large coil and a vesicula seminalis above ovary ;

cirrus sac not long. Genital cloaca small, with circular muscles.

Ovaries double, on either side of innermost of excretory tubes of pore

side of segment. Yolk-gland on one side of same excretory vessel

behind ovary. Shell-gland nearer to the middle of the body above

the yolk-gland. Seminal receptacle, long and not much swollen, |

begins soon after the terminal chamber of the vagina. Uterus broad

and sac-like, occupying a great deal of the middle of the proglottid.

Many paruterine organs present in later stages.

The characters given in the above paragraph are not distinctive

of any known genus of Tetracotylea. And I am, indeed, disposed

to think that ultimately it will be necessary to form a separate

genus for this worm from the Gambian Pouched Rat. In the

meantime, however, I do not burden zoological nomenclature with

an additional name until the possibility of its distinctness becomes

more settled. Besides it is also possible that the existence of this

species removes a barrier between the two genera Thysanosoma

and Anoplocephala. Until I had become aware of the numerous

paruterine organs, I was disposed to refer the worm to Anoplo-

cephala, with which genus it obviously has many points in

common. But the existence of these characteristic paruterine

organs—and in such great numbers—is a reason for removing it

from Anoplocephala and uniting it with Thysanosoma. On the

other hand, the latter genus has either double or single sets of

generative organs, and, correspondingly, either two pores upon each

proglottid or alternating pores, while the Tetracotylean described

in the foregoing pages has generative pores all upon one side.

Nevertheless, the double set of testes seems to be a last trace of an

originally completely double set of gonads and ducts, such as occurs

in some proglottides of other Thysanosomas. If the small set of

the testes existing in this species upon the pore side of the ovaries

were to disappear it would be, as I think, impossible to separate

this genus from Anoplocephala or Zschokkeella ; but the definition

of those genera would have to be enlarged in order to take in the

numerous paruterine organs, which is, after all, perhaps the chief

reason for referring this worm to the subfamily Thysanosominz

which is mainly thus characterised.

ON THE WHALEBONE WHALES. 661

30. On the Natural History of Whalebone Whales.

By J. A. Morcu, (Christiania) *.

[Received March 3, 1911: Read April 4, 1911.]

(Text-figures 160-163.)

No period in the annals of modern whaling exhibits such an

intense activity as that which has been developed during the

course of the last two years, after the pioneer expeditions of the

preceding years to antarctic, subantarctic, and other waters

in the Southern Hemisphere had demonstrated the wealth of

Cetacea which is to be found in some of these localities. The

rise in the price of oil to figures which, through shortage in the

crop of oil-seeds, have not been obtained for the last twenty-five

years, has had a further stimulating effect upon the development

of this pursuit in those southern waters.

The situation to-day, then, is that, after the short run of six

years, whaling in the Southern Hemisphere has attained a com-

mercial importance entirely overshadowing that of the industry

in our northern latitudes, which is now more than forty years old.

As an example, it may be mentioned that last season, from

the island of South Georgia alone, fourteen whaling steamers

brought 106,800 barrels of oil, which is more than the world’s

total production of whale-oil three years ago !

Kven during the latter part of the last century the attention

of Norwegian whalers was directed to these distant regions; and

in 1893 two expeditions started from Norway to try their luck

in the chase of Seals and Right Whales in southern seas. One,

in the ‘ Antarctic,’ proceeded to Australian waters, and the other,

in the ‘Jason, to the islands south-east of Cape Horn. The

pecuniary results were not encouraging, and further attempts

were given up for the time. Then came the Nordenskidld scientific

expedition in 1901, and Capt. C. A. Larsen, in the ‘ Antarctic’

belonging to that expedition, had an opportunity of continuing

his observations made during his previous voyage in the ‘ Jason.’

Having satisfied himself of the seemingly limitless numbers of

whales in those waters, he succeeded, in 1904, in interesting parties

in Argentina in his plan for establishing a whaling-station on the

island of South Georgia; and he began operations there about

the new year, 1905.

Meanwhile, in 1903, the Norwegian Storthing had passed a

law prohibiting whaling on the northern coasts of Norway, and

the now homeless whalers had to go in search of other fields for

their operations. In the spring of that year a comparatively small

vessel was fitted out as a floating factory and dispatched to

Spitzbergen as an experiment. This proved successful; and in

the next summer a larger vessel proceeded to these waters,

* Communicated by S. F. Harmer, Sc.D., F.R.S., V.P.Z.S.

662 MR. J. A. MORCH ON

with the same result. The floating factory, which was going to

play such an important part in the development of modern

whaling, became hereby an established fact; and in October 1905

this same vessel, together with two whaling steamers, was dis-

patched to the Falkland Islands and South Shetland. Upon the

vessel’s successful return with a nearly full cargo of oil in June

1906 the ice was broken ; and now followed with intense activity

the fitting out of a number of floating factories, as also of build-

ings and appliances for the erection of land stations in various

localities in the Southern Hemisphere.

Text-fig. 160.

Floating factory and whaling steamer in harbour, Deception Island,

South Shetland ; with floating carcases of Humpbacks.

From the southern coasts of South America, South Shetland,

the Falkland Islands, South Georgia, the coasts of South Africa,

and Kerguelen Island, whaling is now being prosecuted with an

ever-increasing number of whaling steamers and with returns of

oil undreamt of only a few years ago.

The species which are principally hunted are the Humpback

Whale (Megaptera boops), the Blue Whale (Balenoptera sibbaldi),

and the Finback Whale (B. musculus) ; and, in one locality (the

Falkland Islands), the “Seihval” or Rudolphi’s Whale (B. borealis),

which also occurs on the coast of Chili and the west coast of

South Africa without having been actually hunted there yet.

THE WHALEBONE WHALES. 663

The Sperm Whale (Physeter macrocephalus) and the Southern

Right Whale (Balena australis) are occasional visitors in some of

the localities; the latter species having been observed nearly

every year trekking in schools along South Georgia and towards

the Patagonian coast. At South Shetland, Bottlenose Whales

(Hyperoodon rostratus) have also been observed in small schools,

but have not been the objects of pursuit. The appearance of

this species near the coast in Bransfield Strait is interesting,

but as none were caught no observations could be made upon

what constitutes their food in this locality.

Humpbacks constitute the great bulk of the Whales caught in

most of the above-mentioned localities. The cow of this species

is supposed to go pregnant from ten to eleven months, and,

judging from their numbers there, must be the most prolific of

Whales. In February 1910 I observed at South Shetland two

Humpback feetuses about 23 feet long which, if we accept the

supposition of the late Prof. G. Guldberg as to the growth of

Humpback fcetuses, should be about 21 months old (impregnation,

therefore, about the first of November). At South Georgia,

L am informed, even at the commencement of the season, Hump-

backs with calves from about 12 feet long are met with, and

foetuses are also found here in their first months of development.

According to these observations, the indications are that the

Humpback Whales in these southern latitudes may be supposed

to give birth to their young at some time in September or

October, as against April and the neighbouring weeks in our

northern latitudes.

I may add that from the circumstance that in some of these

southern localities the Whales are only flensed and the carcases

left to be driven by wind and tide, it is unfortunately only by

the merest chance that observations upon the period of gestation

of the various species can be made. As, however, a more appro-

priate disposition of the huge masses of meat and bones may

be looked for in the future, opportunities should also present

themselves for acquiring enough material for examination in

this respect.

An interesting phenomenon is observed from South Georgia

relating to the Humpback Whales. At certain times all the

Humpbacks that are brought in have the belly nearly white ; this

variety may then disappear and those caught for some time may

have the belly marbled ; schools with their bellies entirely dark

may then put in an appearance, succeeded by the first variety, and

so on. At South Shetland I also observed these several varieties

of the Humpback, although they did not there appear in distinct

schools, but mingled. The bulls were here in a decided majority,

and the individuals observed in February and March were mostly

young.

A locality which is attracting serious attention at present 1S

the western coast of South Africa. During the months May to

October especially, the Humpback Whales have been observed in

664 MR. J. A. MORCH ON

great numbers and mostly off the coast of Portuguese West

Africa. I think the question may reasonably be raised whether

we have not here the great bulk of the Humpbacks from the

South Georgia region on their annual migratory route? In this

Text-fig. 161.

Humpback Whale (nostrils open).

locality, which is interesting in more than one respect, observa-

tions might probably be made upon fcetuses of Humpback Whales

in the later stages of their development. Large schools of

Rudolphi’s Whales and Blue Whales have also been observed

THE WHALEBONE WHALES. 665

along these coasts, feeding upon plankton, but the few Blue

Whales caught were all very lean.

The coast of Chili, from which whaling is at present being

prosecuted, also offers opportunities for interesting observations

upon the periods of gestation of various species of Whales, especi-

ally those of the Blue Whales and Rudolphi’s Whales, which are

very little known.

From a consular report which has come to my notice, it appears

that great numbers of the Finback Whale congregate in the

waters along the Brazil coast between South latitudes 12 and 18

every year during the period from May to November. Until

more definite information is obtained upon this subject, further

comment must, however, be reserved.

In the Northern Hemisphere, modern whaling is at present

being prosecuted from the following localities :—Spitzbergen,

Iceland, the Faeroe Islands, Shetland, the Hebrides, the western

coast of Ireland, Newfoundland (one station also in Labrador),

British Columbia, Japan, and Korea, while preparations are also

being made for an early start in Alaska and in the Sea of Okhotsk

from Saghalien Island.

When the veteran whaler Svend Foyn had perfected his

harpoon-gun in the latter part of the sixties, and had commenced

operations in Varangerfiord on the Finmark coast, the Blue

Whales were his only objects of pursuit; and, so long as the

hunting was carried on on a moderate scale, continued to be so for

a series of years. As, however, after the expiration of his patent

an increasing number of new companies entered the field, the

other species of Whales—Finbacks, Humpbacks, and occasional

schools of Rudolphi’s Whales—also became the objects of the chase.

Successively, however, the Blue Whales became scarcer and scarcer,

and the other species of Whales, especially the Finbacks, came

to play the principal part in the catches. The same order of

things has, as a rule, repeated itself also in most other whaling-

grounds in the North Atlantic.

As observations upon the breeding-season and the period of

gestation of the various species of Whales in our northern latitudes

can only be made during a limited part of the year, no exact

information may be said to exist bearing fully upon these ques-

tions. The results from the Finmark coast may very briefly be

summed up thus :—

The Blue Whales have often been observed in coition during

the summer ; the indications are that this act may take place also

at other times of the year. They have never been observed with

small calves in that locality, but occasionally with larger ones.

The period of gestation is supposed to last more than twelve

months, and it is believed that there is no fixed time of the year

when they give birth to their young; this probably takes place

in American temperate waters*. Contents of the stomach of

* The Sulphur-bottom variety has occasionally been obseryed in Finmark waters.

666 MR. J. A. MORCH ON

various Blue Whales have been observed by Prof. G. O. Sars to

consist chiefly of Boreophausia inermis. The Blue Whales are

remarkably free from parasites, this perhaps owing to the cir-

cumstance that their outer skin easily peels off.

Pinback Whales.—Impregnation is supposed to take place in

January—March, and the period of gestation to be about twelve

months. The length of the new-born calf is about 20 feet. Very

young calves are always seen together with full-grown individuals

in schools, older calves sometimes in schools by themselves. It is

supposed in the locality in question that among 50-60 Finback

cows 10-15 will have feetuses and a lesser number will be

accompanied by calves. Some cows give birth to their young

in those waters, some go westwards to other localities. It is

supposed that the cows do not become pregnant every year.

Text-fig. 162.

Typical site of Epizoic Crustacea on the ventral surface of a Humpback Whale.

As the Finback Whales in the North Atlantic feed on

plankton, caplin (J/allotus villosus),and herrings, their distribution

at most times of the year depends on the appearance of these

various sorts of food, The whalers distinguish three varieties:

THE WHALEBONE WHALES. 667

the blackish, the grey, and the yellowish. The blackish forms

follow the schools of caplin and herrings. The time of ap-

pearance of the two other varieties seems to indicate that they

feed chiefly on plankton. Further information upon the

migratory route of these varieties would be interesting, and

observations on the subject might be made during the progress

of the herring fisheries. I may add that at South Shetland, in

February and March, 1910, I observed several Finback and Blue

Whales which were covered by a muddy, yellowish deposit

which could easily be scraped off. As I had not’a microscope

with me, no information upon the nature of this substance

could be obtained. Penmnella is, so far as I know, the only

parasite that has been found, and this only occasionally.

Text-fig. 163.

Epizoic Crustacea (Coronula diadema and Conchoderma auritum)

from Humpback Whale.

Humpback Whales —During the early months of the year in-

dividuals of this species are met with on the Finmark coast going

west. They are at that time followed by large calves, are restless,

and approach the shores. About April the cows are supposed to

give birth to their young in subtropical waters. Impregnation

is supposed to take place shortly afterwards. The Humpbacks

arrive again in the waters north of Finmark in summer;

and around Bear Island they are found in July feeding on

plankton (Boreophausia inermis, &c.) and caplin, and are then

seen with calves about 20 feet long. The fcetuses found at

this time of the year are about 20 inches long. In September

they go east into the Barents Sea, but observations upon their

life during the last months of the year are wanting. They are

668 MR. J. A. MORCH ON

supposed then to feed on caplin and to follow the schools of these

fishes near the “ East Ice” and move westwards with them during

the winter. Those killed on the westward trek have been found

to have empty stomachs.

On account of the tough outer skins of the Humpback, parasites

can easily fasten themselves thereto, and this species is, as a matter

of fact, especially infested by various forms :—Coronula diadema,

Conchoderma auritum, and Paracyamus boopis. Pennella is rarely

found. ;

Rudolphi’s Whales (B. borealis) are very erratic in their

appearances. In 1884, for instance, only six were killed on the

Finmark coast; in 1885, 659. This is a typical plankton Whale,

and it appears on that coast only during the summer, feeding on

Calanus &c. The foetuses have a length of from 3 to 4 feet in

June, from which it may be inferred that the cows give birth to

their young during the latter months of the year in localities

at present unknown.

In Shetland in 1906 I observed on a Rudolphi’s Whale, which

had the front end of its lower jaw deformed, a colony of Concho-

derma auritum fastened thereto. ‘This is the only instance of

parasites on this species that I know of.

Plankton being the only or principal subsistence for the Whales

in question, an exceedingly interesting problem is suggested :—

What part do the great ocean currents play as highways and

feeding grounds for these Whales during their annual migratory

route ?

From investigations carried on by Prof. Nansen and _ his

assistants it has been proved that the cold polar water has a

beneficial effect upon the vegetable life in the open sea. The

cold polar currents, by mixing with waters of a higher temperature,

create favourable conditions for the growth of plankton and

higher marine life.

From these investigations, then, we may infer that it is along

the border layers of the great polar currents where these meet

and intermingle with warmer currents or waters that, given a

sufficient actinity of light for the production of vegetable plankton,

we may expect to find the most favourable conditions for the

subsistence of the plankton Whales.

We know from our northern latitudes that the waters

along the northern coast of Iceland, Finmark, and along the

western coast of Spitzbergen have attracted and been able to

maintain for a long series of years a considerable or even a very

great number of Whales.

On the contrary, we have seen that in localities which are under

the principal influence of a cold polar current—for instance, the

coast of Newfoundland—the stock of Whales has in the course

of only a very limited number of years been seriously reduced,

although the number of whaling steamers employed would not

have been excessive had the same favourable conditions in the sea

prevailed as, for instance, along the northern coast of Iceland.

THE WHALEBONE WHALES. 669

The considerable number of Whales which were met with along

the coast of Newfoundland during the earlier years of hunting

may probably have accumulated, slowly, during the course of time

by natural multiplication, z.e. the older ones have been followed

by their young and these also have come back to the same

locality. If the catches, then, had been proportional to the

approximate natural increase, the industry might most probably

have been carried on to the same extent for a long time to come.

It may be taken for granted that the great bulk of the Whales

which during the spring months migrate northwards through the

Atlantic take an easterly direction and Spread in a fan-like

manner towards Iceland, the Faeroe Islands, Shetland, Spitzbergen,

and the northern coasts of Norway. Now the question is: Does

the western part of the Atlantic bordering upon Nova Scotia,

Cape Breton, and Newfoundland offer them any inducement in the

form of sufficient food which they may be supposed to find at this

time of the year growing along the border of the Gulf-Stream in

various localities? This may, I think, be answered in the negative ;

and I believe that we have here also circumstances which may

have contributed to the condition of things which has manifested

itself off Newfoundland.

On the Murman Coast, also, there has been snails instance

of a similar case. In the eighties, two whaling stations were

started here by the Russians, and under apparently favourable

auspices as to the number of Whales which were seen in this

locality. After a few seasons of successful hunting, however, the

Whales became scarcer and scarcer, so that at last their pursuit

had to be given up. This agrees also very well with our own

experience on the Finmark coast, where the whaling stations in

the eastern localities had to be moved westwards at a com-

paratively early date, as the Whales became scarcer in the colder

areas towards the Murman Coast.

If the statistics of the Whale fisheries in some of our northern

localities are examined, it will strike an observer that the years of

good or poor returns generally run into periods of years of either

the one or the other kind. There may one year be foggy or

boisterous weather accounting for smaller catches, but these

obstructions do not generally last in periods of years. It will take

a closer study of the hydrobiological conditions of larger areas

during the particular years to give a satisfactory explanation of

this phenomenon, and I am only here indicating its existence.

With the latter part of August, or the first part of September,

most of the Whales in our northern latitudes have left their

summer haunts. The migratory routes of the Blue Whales and

the Rudolphi’s Whales between the seasons, and to some extent

also during these, may well be said to be shrouded in mystery.

The two American bomb-lances, which in the years 1888 and

1898 were found in Blue Whales on the coast of Finmark, do not

give us much clue as to the particular locality where they had

been fired into the animals.

Proc, Zoot, Soc.—1911, No. XLVII, AT

670 ON THE WHALEBONE WHALES.

Of the Finback Whales a considerable number must stay in our

home waters also between successive summer seasons, following

and feeding upon the shoals of caplin and herrings at various

localities and times of the year. But their numbers are not so

ereat as to account for the large schools which migrate to southern

latitudes. In the Mediterranean considerable numbers of Fin-

back Whales are seen, but further information from this locality

is lacking.

Tn addition to what has been mentioned above with regard to

the Humpback Whales, nothing is known about the migratory

route of those which visit the waters off Newfoundland and

northwards. It would be of interest to know if they follow

the caplin in these localities, like the FEST p/AGES in the Barents

Sea.

In the month of May, Humpbacks are found off the Azores,

Bermuda Islands, and occasionally the Antilles. In 1899 parts

of an American bomb-lance were found in a Humpback on the

Finmark coast.

The part played by the Gulf-Stream in the biology of Whales

in the North Atlantic is paralleled by the influence exerted by the

Antarctic current or great West Wind Drift in the South Sea.

The localities which have shown themselves to be rich whaling

grounds, such as the coast of Chili, South Georgia, and the couiheet

coast of West Africa, are under the influence of this current.

By the intermingling ‘of the cold waters from this current with

waters from the adjacent warmer currents, conditions must be

created favourable for the production of the masses of plankton

which are sometimes seen in these localities. What part the

Humboldt current along the western coast of South America and

the Benguela current along the western coast of South Africa

play as migratory routes and as feeding grounds during several

months of the year for those species of Whales which at the

present time are of the greatest commercial importance among

the Cetacea, must be left to future investigations to disclose.

The geographical positions of the various whaling grounds in

the Southern Hemisphere should offer special opportunities for

observations upon the migrations, breeding-season, food, and other

questions of biological and economical interest relating to these

Whales.

ON THE NEST OF A GREY STRUTHIDEA. 671

EXHIBITIONS AND NOTICES.

April 25, 1911.

Dr. S. F. Harmer, M.A., F.R.S., Vice-President,

in the Chair.

THE Secretary read the following report on the additions to

the Society’s Menagerie during the month of March 1911 :—

The number of registered additions to the Society’s Menagerie

during the month of March last was 270. Of these 73 were

acquired by presentation, 137 by purchase, 45 were received on

deposit, 8 in exchange, and 7 were born in the Gardens.

The number of departures during the same period, by deaths

and removals, was 158.

Amongst the additions special attention may be directed to :—

2 Siamang Gibbons (Symphalangus syndactylus) 9 2, from

Sumatra, purchased on March 10th.

2 Neumann’s Vervet Cercopitheques (Cercopithecus centralis),

new to the Collection, from Lake Kivu, Central Africa, deposited

on March 23rd.

2 Ruffed Lemurs (Lemur varius) $ 2, from Madagascar, pre-

sented by Frederick Burgoyne, Esq., F.Z.8., on March 15th.

2 Pumas (felis concolor), from Pernambuco, presented by John

Sparks, Esq., F.Z.S., on March 11th.

1 Elephant-Seal (J/acrorhinus crozetensis), new to the Col-

lection, from the Crozet Islands, presented to The King’s African

Collection by Dr. Louis Péringuey, F.Z.8., and deposited by

H.M. Tue Kine on March 23rd.

1 Gundlach’s Troupial (Quiscalus gundlachi), new to the

Collection, from the Greater Antilles, presented by the Countess

of Suffolk on March 2nd.

Mr. C. Tate Reean, M.A., F.Z.S., exhibited a series of lantern-

slides of scales of the Salmon (Salmo salar), and showed how the

life-history of the fish could be read from its scales.

Mr. D. Seru-Smitu, F.Z.8., the Society’s Curator of Birds,

exhibited :-—

(1) A nest (text-fig. 164, p. 672) of the Grey Struthidea or

Apostle Bird (Struthidea cinerea), composed entirely of mud, and

built on a branch in the Western Aviary.

(2) Lantern-slides from photographs of the King Penguin

(Aptenodytes pennanti) and Black-footed Penguins (Spheniscus

demersus) Showing the method of moulting.

(3) Lantern-slides of a number of wild Swainson’s Lorikeets

(Trichoglossus novee-hollandie), from photographs kindly sent

by Mrs, Innes, of Mackay, North Queensland. These birds came

47*

672 ON A COLLECTION OF SKINS FROM UGANDA.

in large numbers to feed daily at a table, on syrup provided,

settling without fear upon the head, shoulders, and arms of the

lady who fed them.

Text-fig. 164.

Nest of the Grey Struthidea.

Some Mammals from Uganda.

Dr. C. Curisty, F.Z.8., exhibited specimens from a collection

of skins of antelopes, hyrax, monkey, cheetah, serval and serva-

line cat, &e., obtained in the Chagwe forests in the south-east of

the Uganda Protectorate, and made the following remarks :—

“Perhaps the most interesting specimens in the collection

before you are those of Cephalophus weynsi. This red duiker

was first described by Mr. O. Thomas, F.R.S., from two skins

from eastern Congo. Subsequently a specimen was obtained by

Mr. L. M. Seth-Smith in the Budongo forest on the east of Lake

Albert. An imperfect skin in the National Collection, labelled

Cephalophus johnstoni, also obtained from the Lake Albert region,

and described some time ago by Mr. Thomas, may possibly also

belong to this species.

‘My own four specimens were shot in the Mabira forests more

ON A LOIN-CLOTH OF A NATIVE OF NORTHERN NIGERIA. 673

than 100 miles east of the Budonga, and with very little forest

intervening. The locality is a new one, and is probably the

easternmost limit of the species, which, after another gap of

150 miles of more or less open country still further east, gives

place to Cephalophus ignifer, the common red duiker of the East

African upland and rift-valley forests.

“The two species are closely allied. In both the body-hair is

short and close-laid, and the horns slope backward in a line with

the forehead. Amongst other characteristics C. weynsi may be

distinguished by the hairs on the nape being reversed forward.

“These sleek, heavy, short-legged duikers are very pig-like in

gait and appearance, carrying the head low. They are found only

in dense forest, and, so far as my experience goes, never even feed

in the open, unlike the grey duiker so commonly to be met with

at forest-edges and often seen in the open. My two best heads of

C. weynsi measure 44 and 43 inches in length respectively.

‘“‘T may also draw your attention to the specimens of hyrax—

Procavia emini and P. dorsalis (2). Naturalists do not seem to

quite realise that certain species of hyrax, the Dendrohyrax group,

are entirely arboreal, never living amongst rocks or holes in the

ground, but inhabiting the largest trees in dense tropical forests ;

whilst other species are rock dassies, and though able to run up

and down the face of a perpendicular rock or even to play about

the neighbouring bushes, yet are in no sense arboreal. The

members of the Dendrohyrax group do not even live in hollow

trees as a rule, but upon the branches.

‘On looking closely at the skins of these two animals, P. emane

and P. dorsalis (2), both killed in high trees, it is interesting to

observe that the long bristles amongst the fur, so numerous and

so conspicuous, especially on the hinder part of the body, in those

species which are not arboreal, are here obviously absent or

only to be found on the neck or shoulders. I find, as the result

of an examination of the skins in the British Museum, that this

distinctive peculiarity holds good for the two groups in almost

every instance, the arboreal Dendrohyrax group being almost

without them, while in the rock-inhabiting species they are very

conspicuously developed, mainly posteriorly.

“Tt, seems probable that these long stiff hairs are tactile organs

and of very considerable use in dark burrows and holes amongst

the rocks; whilst it is easy to see that they are of less utility

on the branches of trees, and in time, no doubt, would become

rudimentary or disappear altogether.

“he weird, nocturnal, ventriloquistic cries of both groups of

these animals are even more extraordinary than their powers of

climbing.”

Dr. Currsty also exhibited a loin-cloth taken in 1898 from the

dead body of a native in the Gando-Bornu district of Northern

Nigeria. In referring to it, he said :—‘‘ When first I caught

sight of this ornamental piece of wearing apparel it seemed so

peculiar that I stopped to secure it under considerable difficulties.

674 DR. WILLIAM NICOLL ON A UNIQUE

It is apparently made from the skin of a young Cobus cob; but

the white of the belly part and the inner part of the hind legs

you will see are transversely striped with inch-wide stripes, some

two or three inches apart, of a reddish-brown colouring exactly

resembling that of the animal. These stripes are doubtless made

with some native pigment or dye; but they are so placed and the

colour is so permanent, so fixed and difficult to remove by any

means, that serious doubts have been entertained, as Mr. Thomas

will tell you, as to whether they were artificial or natural

markings. one

“Tt will be also noted that the white hair on the striped belly-

parts is curled and not unlike sheep’s wool, but quite unlike any

known antelope. If this curling of the hair in this situation is

not natural, I can only suggest that it is produced by the heat

and chafing of the wearer’s thighs.”

Mr. Oldfield Thomas, in commenting upon Dr. Christy’s exhibit,

stated that there was little doubt that the stripes and curly

appearance of the hair were artificial, but the fact that the dye,

whatever it was, so closely resembled the natural colour of the

animal and appeared to be so “ fixed,” the possibility of the skin

belonging to some antelope hitherto quite unknown had been very

carefully considered at the Natural History Museum.

On a unique Pathological Condition in a Hare.

(Text-figure 165.)

Dr. Wiuit1am Nicout, M.A., F.Z.8., exhibited some pre-

parations from a Common Hare (Lepus europeus), which showed

an interesting and unique pathological condition. The hare was

obtained by W. Raphael Muckley, Esq., and sent by him to the

British Museum, whence it was forwarded to the Lister Institute

of Preventive Medicine. The manner in which the hare died

was somewhat remarkable. It was observed by Mr. Muckley to

pitch violently out of a hedge on to the roadside where it lay

struggling, and it died about three hours later. To him the

symptoms seemed to point to poisoning. At the post mortem

examination the liver was found to be extensively invaded with

small whitish chalky deposits of various sizes, especially con-

spicuous on the posterior surface. There was also a considerable

amount of chronic inflammation around the liver, with adhesions

to the diaphragm. On section. the liver was friable and gritty.

The concretions were amorphous and insoluble in acid. The bile-

ducts showed some fibrous thickening. On microscopic exam-

ination of a scraping from the liver, numerous ova of peculiar

structure were seen. They were fairly uniform in size, measuring

057 x ‘033 mm. At first sight they bore a considerable resem-

blance to the eggs of Trichuris (Trichocephalus) or Trichosoma,

but on more careful inspection they presented one or two unusual

features. The colour was much lighter, being greyish instead of

brown. The shell consisted of two layers, the outer of which

was marked with pronounced radial striations; the imner was

PATHOLOGICAL CONDITION IN A HARE. 675

constituted of fine concentrically-arranged lamelle. The two

layers were not in close apposition but were separated by a

narrow, somewhat irregular space. The egg is thus provided

with a double shell. At each pole of the egg there was a small

circular aperture, piercing both layers of the shell. The external

surface of the shell presented a well-marked papillated appear-

ance, the papille being irregularly rounded. Inside the shell

Text-fig. 165.

M.R., del.

Eggs of a Nematode worm from the liver of a Hare.

A. External surface of egg. bB. Optical section of egg: two-celled stage.

C. Egg in four-celled stage with thickened inner wall: from vagina.

there was a thin, continuous membrane investing the egg. The

polar apertures were each closed by a plug of homogeneous

material which was separate and distinct from the membrane

investing the egg-cells. They completely filled the apertures but

did not bulge beyond them. The eggs were mostly in the two-

celled stage, but a number of them showed four cells.

676 ON A UNIQUE PATHOLOGICAL CONDITION IN A HARE.

From the appearance of the eggs it is obvious that the lesions

were due to a Nematode worm of the 77ichosoma group. Search

was accordingly made and a number of worms were obtained. In

every case, however, they were fragments lacking the anterior

and posterior extremities. The longest specimen measured

27 mm. It was a female, full of eggs, and it was very slender.

It was narrowest at the anterior end (‘14 mm.), the breadth

increasing gradually towards the posterior end where it was

‘23mm. The cuticle was marked throughout with fine annular

striations. The vagina was of considerable length but no genital

aperture was seen, so that it must have been far forward. In

the vagina a number of eggs were present, which were much

larger than those first seen. They measured -070—078 x -:040—

045 mm. The shell, too, was much thicker, the increase being

almost entirely due to a great thickening of the inner layer. |

The polar apertures were somewhat reduced in size and the eggs

were for the most part in the four-celled stage.

The lesions in the liver showed much resemblance to those

seen in advanced stages of coccidiosis, a very common condition

in rabbits. For this reason they might readily be diagnosed as

such on cursory examination. This, so far as can be gathered, is

the first record of such an infection of the liver of hares or rabbits

by parasitic nematodes of this kind.

An analogous condition has not infrequently been met with in

rats, and has been reported from Europe, India, and Australia.

No record has been made of its occurrence in this country.

Whether this parasite in the hare is the same as that in the

rat must remain doubtful, for the descriptions of the latter have

hitherto not paid much attention to the character of the eggs

beyond mentioning that they resembled those of Z’richosoma.

An attempt is being made to hatch the eggs and to produce

infection in rabbits and rats, but the development is extremely

slow even at a constant temperature of 26° C.

In addition to the lesions in the liver the hare had a very

heavy infection of Z'richostrongylus retorteformis in the intestine

and a slight intestinal infection with Coccidiwm cuniculi. The

intestine and the appendix, moreover, showed a large number of

small caleareous patches, but on examination nothing of a parasitic

nature could be detected in the patches. This was of interest

from the fact that I had already seen similar patches in the

cecum of a Variable Hare, sent me from the Society’s Gardens.

The liver, unfortunately, was not submitted for examination.

That these patches might have something to do with the liver

condition was not impossible, for they might be considered as

degeneration following injury such as the passage of a worm

through the intestinal wall.

The hare, further, showed signs of recent parturition and there

was a septic condition of the uterus, and it must remain an open

question as to whether death was due to this or to the liver

disease.

PZ.S.19U.PLXXVI.

London Stereoscopic Co. imp.

M. Rhodes, del.

LECHRIORCHIS VALIDUS.

eee

Taian!

Rein

PZ.S.19M. PL XXVoL.

M. Rhodes, del. London Stereoscopic Co imp.

6. 7. OCHETOSOMA FORMOSUM.

8-10. DASYMETRA CONFERTA.

ON NEW TREMATODES FROM REPTILES. 677

PAPERS.

31. On Three New Trematodes from Reptiles.

By Witiiam Nicout, M.A., D.Sc., M.B., F.Z.S.*

[Received December 29, 1910: Read April 25, 1911.]

(Plates XX VII. & XXVIII.T)

The following notes were made on a collection handed over to

me by the Prosector of the Zoological Society. In no case was

the habitat noted, but from their affinities it may be presumed

that the specimens all came from the lungs, the mouth, or the

cesophagus. The specimens are interesting as forming an im-

portant addition to our knowledge of the large variety of forms

which inhabit the air-passages and anterior end of the alimentary

canal of reptiles and batrachians. To the taxonomy of this

particular group Odhner has recently (1910) made a valuable

contribution, which will be further referred to later.

The first species is from the Hog-nosed Snake (Heterodon

platyrhinus), and I include it provisionally in the genus Lechri-

orchis Stafford, 1904. Like most of Stafford’s genera, this genus

is insufficiently defined, although the fact that he includes in it

the well-described species LZ. (Renifer) elongatus Pratt, 1903, is

something to go upon. It is, therefore, necessary to amend

Stafford’s definition somewhat, as follows.

Genus LecHRiIorcHIs Stafford, 1904.

Resembling Renifer Pratt, 1903, except in the following

particulars. The intestinal diverticula extend a short distance

beyond the testes; the genital aperture is further from the edge

of the body, midway between it and the pharynx ; there is a well-

developed vagina.

These constitute, as far as appears, the only differences between

the two genera.

LECHRIORCHIS VALIDUS, sp. n. (Plate X XVII. figs. 1-5.)

This is a species of moderate size, measuring 3°6-7:-4 mm. in

length by ‘9-1-4 mm. in breadth. The smallest specimens had

just begun to produce ova, so that the minimum adult size is

probably about 3 mm. The body is elongated, the length being

4_5 times the breadth. The latter is comparatively uniform, but

the tail is distinctly pointed. The anterior end is more rounded,

while there is a tendency fora slight narrowing to occur about

or behind the ventral sucker. In young specimens the body is

flattened, but it becomes much thicker as the uterus increases in

size. In transverse section an adult specimen presents a strongly

convex dorsal surface and a fiat or slightly convex ventral surface.

* From the Lister Institute of Preventive Medicine, London.

+ For explanation of the Plates see p. 686.

678 DR. WILLIAM NICOLL ON

The cuticle is fairly thick, somewhat deciduous, and is studded

throughout its whole extent by stout salient spines. These are

regularly arranged, and become sparse towards the posterior end.

The underlying musculature has the usual formation.

In an average adult specimen of 6°5 mm. length the oral

sucker has a diameter of -45 mm. It is subterminal, globular,

and its aperture has a marked muscular rim. The thickness of

its wall is about ‘13 mm. In the same specimen the ventral

sucker measures 66 mm. It is also globular, slightly flattened,

and its wall is -18 mm. thick. The sucker ratio is, therefore,

very approximately 2:3. The diameter of the sucker relative to

the body-length diminishes as the animal increases in size.

Thus, in the smaller specimens the ventral sucker is + of the

body-length, while in the largest it is only j. The ventral

sucker is situated about 4 of the body-length from the anterior

end. In this respect, again, the young specimens differ from the

older ones in that they have the ventral sucker relatively further

back.

There is a very short prepharynx; usually the pharynx 1s

contiguous with the oral sucker, and it measures -23 x17 mm.

The cesophagus is about ? of the length of the pharynx, although

it is longer in young specimens. The intestinal bifurcation takes

place well in front of the ventral sucker. The intestinal diverticula

run almost parallel to the edges of the body. They bend in a

little just behind the ventral sucker, but they are pressed out

again by the testes. At their termination, just behind the

posterior testis, they usually turn in a little. Their ends are

about 2 of the body-length from the posterior end. They are

comparatively narrow tubes, and their wall is crmkled on the

inner side. The outer side is plain. They are lined by a layer

of low epithelium. The cesophagus has the usual cuticular lining.

Throughout their whole extent the diverticula are somewhat

ventral in position.

The excretory system is characteristic and agrees with Odhner’s

supposition as to its form in the group to which this species belongs.

The vesicle consists of a fairly broad main stem, opening at the

tip of the tail and passing forwards to the shell-gland, where it

divides into two limbs. These diverge to form a Y, and each passes

over the edge of the ventral sucker and terminates halfway

between the sucker and the intestinal bifurcation. The main stem

is pressed close to the dorsal surface, but the limbs are not quite

so dorsal in position. They eventually come to lie close up to the

intestinal diverticula and separate these from the uterus and

cirrus-pouch respectively (Pl. XX VII. fig. 4). The main stem is

much compressed dorso-ventrally, but the limbs are round or com~

pressed transversely. In addition to this central system, however,

numerous lateral twigs are given off from the stem and limbs.

These all radiate outwards towards the edges of the body, dividing

and subdividing and eventually forming an intricate anastomosis

(Pl. XX VII. fig. 3). The entire lateral fields of the body are thus

filled with a complex network of excretory tubes, and the amount

NEW TREMATODES FROM REPTILES. 679

of parenchymatous tissue is very much reduced. This is more

particularly the case in the post-acetabular region. It gives rise

to a reticulated appearance, which is strikingly seen in young

specimens (Pl. XX VII. fig. 1). In older specimens it is obscured

by the growth of the uterus.

The genital glands are difficult to distinguish in the adult, but

are easily seen in younger specimens. The testes lie not very far

behind the ventral sucker, near and internal to the ends of the ~

intestinal diverticula. They are obliquely situated, the left testis

being half its diameter in advance of the right, and they are

separated from each other by the uterus. In a young specimen,

the uterus being narrow, they lie quite close together, but they

are pressed further and further apart by the growth of the uterus.

They are flat, elongated oval bodies measuring at least *8 x 4 mm.

Their outer margin is plain, but their inner margin is indented

im one or more places, so that the outline is somewhat irregular.

At first they lie almost flat in the body, but the expansion of the

uterus pushes their inner border towards the dorsal surface so

that eventually they are considerably tilted.

The genital aperture is situated on the left side midway between

the pharynx and the edge of the body. It is always on the level

of the pharynx. The cirrus-pouch is of considerable length, and

is a conspicuous object. It is elongated, somewhat slender, and

extends to the middle of the ventral sucker. Its wall is remarkable

for the great development of the longitudinal muscular fibres,

which are stout and very prominent. The circular fibres are

much smaller. Within the pouch there is a more or less highly

convoluted vesicula seminalis. Usually it is simply bent double

(Pl. XXVIT. fig. 2), but frequently it is much more twisted. It

is small compared with the size of the cirrus-pouch. It is not

much dilated, and it is connected with the pars prostatica by

a narrow duct. The pars prostatica is of relatively great length.

It is an almost straight tube extending from a little in front of

the ventral sucker to the point where the cirrus-pouch crosses the

left intestinal diverticulum. It is uniform and fairly narrow.

Surrounding it are numerous prostatic cells which fill up the

greater part of the cirrus-pouch. The ductus ejaculatorius is

short and narrow, and the exsertile, unarmed cirrus is not very

long. The genital sinus is quite small.

The ovary is situated just behind the end of the cirrus-pouch,

over the right posterior quadrant of the ventral sucker. Half of

it lies beyond the sucker, and it frequently overlaps the adjacent

intestinal diverticulum. It is analmost globular body, somewhat

flattened dorso-ventrally, and is about half the size of the testes.

Behind and internal to it lies a compact shell-gland, with a short

ootype and a Laurer’s canal, but no receptaculum seminis.

Laurer’s canal.is short and opens dorsally in the middle line,

about the level of the ovary. A small yolk reservoir lies dorsal

to the shell-gland. The yolk-glands are of limited extent. They

are entirely lateral and lie close to the outer side of the intestinal

diverticula. On each side there are about half a dozen ill-defined

680 DR. WILLIAM NICOLL ON

groups of follicles, which extend from midway between the

intestinal bifurcation and the anterior edge of the ventral sucker

to near the ends of the intestines. The initial part of the uterus

is practically empty. In section it can be made out with difticulty

as an extremely narrow tube, passing backwards from the shell-

gland. Behind the right testis it widens out to form a recep-

taculum seminis uterinum, consisting of about four or five small

dilatations. Further back a few ova appear, but the uterus

still remains somewhat narrow. When it has nearly reached the

posterior end of the body it turns abruptly on itself to form an

ascending limb. Almost immediately this begins to dilate, and

it has only proceeded a short distance before it almost completely

fills the interior of the body. It passes forward, over and between

the testes, crosses the ventral sucker, and terminates ina straight,

thick-walled vagina, which is about half the length of the cirrus-

pouch, and lies on the left side. The uterus thus consists of

a descending and an ascending limb, the former being empty for

the greater part of its length, and the latter being enormously

dilated. The increase in size, therefore, takes place, not by an

increase in convolutions, but by a great dilatation of the ascending

limb. The ova are very numerous, rounded oblong in shape, and

dark brownin colour. They have a large well-marked operculum.

Many of them are more oval than oblong, and this gives rise to

some variation in dimensions. From a large number of measure-

ments the limits were found to be -038-:045 mm. for the length,

and -018-:023 mm. for the breadth, and the average ‘040 x :021

mm. The extreme sizes observed were 045 x-018 mm. for the

most oblong ova, and -038 x :023 mm. for the most oval.

No case of amphitypy was observed in any of the two dozen

specimens forming the collection. The nearest approach was in

one specimen where the testes were practically symmetrical, the

left being a trifle behind the right, but the ovary and genital

aperture were normal. In all the other specimens the position

of these structures was exactly as I have described. The species,

however, is extremely variable in one respect, namely, the

posterior limit of the yolk-glands. Hardly two specimens agree

in this respect. In some specimens they extend a short distance

beyond the testes, in others they reach the middle of the posterior

testis, and again in others they stop short of the testes. In

addition, they are very frequently asymmetrical, extending further

back on the right than on the left, or more rarely vice versa. The

fact, however, that their anterior limit is constantly symmetrical

induced me to consider a symmetrical posterior limit as the normal.

In two specimens also, the intestinal diverticula were of unequal

length, the left diverticulum being considerably shorter than the

right, which was normal. The position of the genital aperture

varied only very slightly, and most of the apparent variations

were due to contraction. The length of the cirrus-pouch was

practically constant, although in one specimen it extended nearly

to the posterior border of the ventral sucker. The size of the ova

was constant within the limits noted, and no increase in size takes

NEW TREMATODES FROM REPTILES. 681

place as the animal grows older. The average size of the ova in

young specimens was found to be the same as that in fully grown

specimens, and I am inclined to view with some doubt Odhner’s

statement (2. p. 59) that the ova in Renifer sauromates Poir.

increase in size as the animal grows older.

From Lechriorchis elongatus Pratt, this species is distinguished

by having more unequal suckers, the ventral being decidedly

larger, the yolk-glands being more extensive, being present some

distance in front of the ventral sucker, and in having slightly

larger eggs. From Z. primus Staff.,it appears to be distinguished

by its much smaller eggs, its smaller ventral sucker, and probably

in other respects.

« The second lot of specimens consists of five from an Annulated

Snake (Leptodira annulata). They bear a close superficial resem-

blance to Lechriorchis validus, but they do not belong to the same

genus. The only genus to which they can at present be referred

is Ochetosoma Braun, 1902, but they do not entirely agree with

the definition of that genus as given by Braun. In internal

anatomy they correspond quite closely, but they are not nearly so

flattened as O. monstruosum Brn., the only species of the genus.

The only other genus to which they could be referred is Renifer

Pratt, 1903, but from that they differ radically in the configuration

of the uterus.

OCHETOSOMA FORMOSUM, sp. n. (Plate XXVIII. figs. 6 & 7.)

The body is elongated and slightly flattened. The length is

3°7-5'7 mm., and the greatest breadth, about the ventral sucker,

is 11-16 mm. The breadth is therefore rather less than 4 of

the length. The body tapers gradually towards each end.

In an average specimen (length 4°5 mm.) the oral sucker has

a diameter of -45 mm., 7. e. ;/5 of the body-length. It is globular

and subterminal. The ventral sucker is situated 1:7 mm. ‘from

the anterior end. It is slightly oval, the transverse diameter

being ‘64 mm., and the longitudinal -57 mm. The sucker ratio

is therefore approximately 3: 4.

The cuticle is extremely deciduous; in most of the specimens

it was almost entirely stripped off. Only in one was it intact, and

even then not completely so; from this specimen the presence of

minute regular spines was determined.

There is a short prepharynx followed by an almost globular

pharynx, measuring “17x ‘16 mm. The cesophagus is about the

same length as the pharynx (:2 mm.), and the intestinal bifurcation

occurs well in front of the ventral sucker. The diverticula

diverge pretty widely, and they terminate almost immediately

behind the ventral sucker, the ends being somewhat turned in.

They are narrow and irregularly dilated, but there is no crenation

on their inner wall as in Lechriorchis validus. They are lined by

low epithelium. :

The excretory system is almost identical with that in Lechriorchis

validus, The vesicle consists of a similar dorsal main stem, which

682 DR. WILLIAM NICOLL ON

divides just behind the ovary into two limbs, which pass forwards.

a short distance in front of the ventral sucker. They are thus

not so long as in the above mentioned species. The main stem,

too, appears more expanded. There is the same system of

secondary lateral branches which anastomose freely in the sides

of the body.

The genital aperture is situated almost at the extreme left

margin of the body, on the level of the posterior end of the

pharynx. The genital sinus is very small. The cirrus-pouch is

very like that of Lechriorchis validus, but it is shorter. It is

usually disposed obliquely, and it terminates a short distance in

front of the ventral sucker, from which its end is usually separated

by a coil of the uterus. In one specimen it almost reached the

sucker. It contains a small convoluted vesicula seminalis, ending

in a narrow portion which runs into the pars prostatica. The

latter is not so long as in the foregoing species, and it 1s more

expanded, especially at its posterior end. The prostatic cells are

numerous. ‘There isa short ductus ejaculatorius, and an unarmed

cirrus of moderate length. The testes are situated just behind

the middle of the body (:3 mm. behind the ventral sucker). They

are symmetrical and lateral. They lie behind the ends of the

intestines, but are separated from them by folds of the uterus.

The length of each is about °5 mm. They are fairly thick and

elongated, and their outer margin is always distinctly divided

into three large lobes, which may in addition be slightly crenated.

The inner margins are completely obscured by the overlying folds

of the uterus.

The ovary is situated over the right posterior quadrant of the

ventral sucker and projects half beyond it. It is obliquely ovoid

and measures ‘°24x°17 mm. A large shell-gland lies close to its

inner side. The yolk-glands are entirely lateral and of limited

extent. They reach from the anterior border of the ventral

sucker to about the middle of the testes. Again in this species,

however, the posterior limit is extremely variable, and may be

anywhere between the anterior and posterior borders of the testes,

but never beyond them. The anterior limit is practically con-

stant. The uterus fills almost the whole of the post-acetabular

region, but its configuration is entirely different from that in

Lechriorchis. ' Here, again, the descending limb is small and

almost empty ; reaching the posterior end of the body it turns

into the ascending limb. In this case, however, accommodation

for the enormous number of ova is obtained not by excessive

dilatation, but by numerous convolutions, the diameter of the

uterus not being very greatly increased. The convolutions have a

markedly transverse disposition, extending from side to side of

the body. In the region of the testes the convolutions are shorter

and stouter. An additional small convolution is formed in front

of the ventral sucker. The uterus terminates in a well-marked

vagina, which is about a third of the length of the cirrus-pouch.

The ova are very like those of Lechriorchis validus, but are usually

more oval. They have a large distinctly-marked operculum, and

NEW TREMATODES FROM REPTILES. 683

they vary in length from -034 mm. to ‘042 mm. by -017 mm. to

°021 mm. in breadth. The average is about -4 x°2 mm.

In this species, again, no case of amphitypy was observed, and

the only pronounced variation was in respect of the posterior

limit of the yolk-glands as described above.

The species obviously presents a close resemblance to the genus

Renifer Pratt, sens. strict. In the shortness of the intestinal

diverticula, the symmetrical situation of the testes, and the

extreme lateral position of the genital aperture, the agreement is

complete. The essential difference lies in the configuration of the

uterus. Renifer ellipticus Pratt, the type species, is unfortunately

not fully grown and the ultimate disposition is not apparent. In

R. sauromates Poirier, the uterus is of the same type as in

Lechriorchis validus, and if this be taken as characteristic of the

genus, then O. formosum must be separated from that genus.

It is evident that the three genera Renifer, Lechriorchis, and

Ochetosoma ave somewhat closely related, and they differ from all

the other members of the family Lepodermatide in the extreme

lateral and forward position of the genital aperture. They

evidently form the nucleus of a group, but the extremely profuse

variety met with in the family renders it somewhat difficult to

divide it into definite subfamilies. Provisionally, however, these

three genera may be classified under Pratt’s subfamily Reniferinz.

That Pneumatophilus Odhn., and Leptophallus Liihe, are to be

included along with these, as Odhner has indicated (2. p. 56),

appears to me somewhat doubtful.

The third form which I have to describe here is one of very

great interest. It was obtained from a Diamond Water-snake

(Tropidonotus rhombifer) from North America. The habitat,

unfortunately, is not recorded. It bears a certain resemblance to

the foregoing species, and belongs to the family Lepodermatide,

but it possesses an individuality sufficiently marked to constitute

a distinct generic type.

DasYMETRA CONFERTA, gen. et sp.n. (Plate XXVIII. figs. 8-10.)

The collection consisted of about a dozen specimens, all of which

were mature, and measured 35-46 mm. in length. The body is

elongated, slightly flattened and of fairly uniform breadth. The

greatest breadth occurs about the middle and is 1-1-4 mm. The

length is therefore about 34 times the breadth. The cuticle is

beset throughout its whole extent by long straight spines. It

appears to be somewhat deciduous, and in many specimens is

absent from a considerable part of the body, especially towards

the posterior end. Several specimens, however, retained the

cuticle and spines quite intact.

In a specimen of average length (4:2 mm.) the oral sucker

measures °56 mm. in diameter. It is globular, almost terminal

and not very muscular. The ventral sucker is practically of equal

size, if anything a trifle less. It is somewhat transversely oval,

the dimensions being 52 x-57 mm, It is only slightly prominent,

684 DR. WILLIAM NICOLL ON

not very muscular, and is situated 1-7 mm. from the anterior end.

The neck, therefore, comprises 2 of the body-length.

The alimentary canal is highly developed. It consists of a very

short prepharynx, with an enormous pharynx measuring *28 mm.

in diameter. The esophagus is shorter than the pharynx, being

only about -2 mm. long. It is fairly wide, with well-developed

musculature and numerous peri-cesophageal cells. It divides into

two very wide diverticula, which extend along the sides of the

body to near the posterior end. From the latter they are

separated by a loop of the uterus. The ends are slightly inflated

and somewhat turned in.

The excretory system has the same general structure as in the

two previous species. The main stem of the excretory vesicle

divides close behind the sheli-gland into two limbs, which extend

a short distance in front of the ventral sucker. From the vesicle

numerous twigs are given off, which divide and subdivide in the

lateral fields to form an intricate anastomosis. ‘The most peculiar

feature of the excretory system, however, is the pigmented con-

dition of the excretory tubules, which renders them strikingly

conspicuous and marks out their course with great distinctness.

This feature renders thespecies uniqueamongst the Lepodermatide.

The pigmentation is due to the excretory granules, which are

almost black in colour, and which fill the tubules. Only a few of

these are to be met with in the vesicle. A main excretory tubule

runs along the greater part of the length on each side of the body,

ventral to the intestinal diverticula. In front of the ventral

sucker it divides into small branches, one of which runs in to join

the vesicle, and another runs forward to the oral sucker. At the

posterior end it also divides into several branches. It is impossible

to say whether this pigmented condition occurs in life or is

a post-mortem appearance, but it was certainly present in all the

specimens.

The genital aperture is median, just over the intestinal bi-

furcation (‘3 mm. in front of the ventral sucker). It shows

a tendency to be deflected very slightly to the left side. In every

specimen the long, thick cirrus was exserted. The cirrus-pouch

is short and stout, in some cases being almost globular. Its

posterior end lies dorsal to the middle of the ventral sucker, but

it may extend beyond this to almost the posterior border of the

sucker. The pouch has an external wall composed of very thick

longitudinal muscle-fibres, with an inner layer of much smaller

circular fibres. It contains a small, slightly-coiled vesicula semi-

nalis, a small bulbous pars prostatica, with numerous prostatic

cells, and a long ductus and unarmed cirrus. As already men-

tioned, the latter was exserted im every case, so that the

arrangement depicted in fig. 8 (Pl. XX VIII.) must be regarded as

hypothetical. The vesicula, prostate, and cirrus all have a very

well-marked layer of longitudinal muscle-fibres,

The testes are situated obliquely, the left being well in front of

the right, but not entirely so. The former les about °3 mm., and

the latter ‘8mm, behind the ventral sucker. They are large ovoid

NEW TREMATODES FROM REPTILES. 685

bodies, with entire margins, and their long axes lie nearly in the

longitudinal axis of the body. They measure *5—7 mm. in length

and ‘4—"5 mm. in breadth. They are most remarkable, however,

for their great thickness, which is equal to or greater than the

breadth. They thus occupy nearly the whole body-thickness, a

fact which prevents them being obscured by the uterus. ‘They

are separated from each other by the wide ascending limb of the

uterus, against which they press, and their outer margins are

closely apposed to the intestinal diverticula.

The ovary is situated over the right posterior quadrant of the

ventral sucker, a short distance behind the end of the cirrus-

pouch. It projects well beyond the sucker, and is transversely

oval, measuring °35 mm. xX ‘28 mm. It lies close up to the dorsal

surface of the body. Close to the inner side of the ovary and

a little behind it, les a large shell-gland, with a short ootype.

A receptaculum seminis is absent, but Laurer’s canal is present.

The yolk-glands are rather voluminous. ‘They are entirely

lateral and peripheral, and they extend from the level of the

genital aperture, or rather behind it, to near the posterior border of

the right testis. Here, again, the anterior limit is fairly constant,

but the posterior limit is somewhat variable, less so, however, than

in the two species already described. The follicles are large and

arranged in regular dendritic groups. All the follicles are

connected up by short ducts. They le close under the surface

of the body, extending widely under the dorsal surface, but being

much more restricted ventrally, where they do not overlap the

inner wall of the intestinal diverticula.

The uterus is large and voluminous, but isnot much convoluted.

It is intermediate in type between that of Renifer and Ochetosoma,

but resembling the former more than the latter. It consists of

a small descending limb, which has a dorsal position and contains

a considerable number of ova. This runs to the posterior end of

the body where it forms a small convoluted knot, a fairly sym-

metrical pair of loops, one on each side, being thrown up towards the

ends of the intestinal diverticula. From this knot emerges the

ascending limb, which rapidly widens but does not attain its

maximum width till it has passed in front of the testes. 1t is

never so enormously dilated as in Lechriorchis validus. Near the

middle of the ventral sucker it runs into a remarkably powerful

vagina, which is as long as or somewhat longer than the cirrus-

pouch. The vagina possesses unusually muscular walls, there

being a very thick layer of longitudinal fibres and an equally

thick layer of circular fibres. It is surrounded by a great mass of

peri-vaginal cells, and it is lined by a thick layer of cuticle from

which small regular cuticular processes extend into the lumen.

The ova are numerous, dark brown and oval, with large well-

marked operculum. ‘They measure ‘033—'037 mm. in length and

‘016-019 mm. in breadth, the usual size being -036 x ‘018 mm.

The genus Dasymetra may be defined as follows :—

Lepodermatide; with moderately flattened body, entirely covered

Proc, Zoou. Sov.—1911, No. XLVIII. 48

686 ON NEW TREMATODES FROM REPTILES.

with spines. Intestine with large pharynx and wide diverticula

which extend near but not quite to the posterior end. Ex-

eretory vesicle Y-shaped with numerous side twigs. Genital

aperture median, a short distance in front of the ventral sucker.

Cirrus-pouch short and plump ; vesicula and pars prostatica short ;

cirrus long. Receptaculum seminis absent ; Laurer’s canal

present. Yolk-glands extensive, dendritic, peripheral. Uterus

forming a small convoluted knot at the posterior end of the

body, with a wide unconvoluted ascending hmb. Vagina long

and very muscular. Ova about ‘035 mm. long.

Type, D. conferta, sp. n.

The type-specimens of these species are deposited in the

Museum of the Royal College of Surgeons, London. _Co-types at

the Zoological Society’s Gardens.

References.

(1) M. Brawn, 1902.—Fascioliden der Vogel. Zoolog. Jahrbiicher ;

Abt. f. Syst. xvi. pp. 64-67.

(2) T. OpHner, 1910.—Nordostafrikanische Trematoden, grossten-

teils vom Weissen Nil. i. Fascioliden. Results of the

Swedish Zoological Expedition to Egypt and the White

Nile, 1901. No. 23 A, pp. 22-76.

(3) J. Poirier, 1886.—Trematodes nouveaux ou peu connus.

Bull. Soe. Philomat. Paris, sér. 7, vol. x. pp. 24-6.

(4) H.S. Prarr, 1903.—Descriptions of Four Distomes. Mark

Anniversary Volume, pp. 23-38.

(5) J. Starrorp, 1905.—Trematodes from Canadian Vertebrates.

Zool. Anzeiger, xxviii. p. 691.

EXPLANATION OF THE PLATES.

The following letters apply to all the figures :-—

D.St. Yolk-glands. | R.S.Ut. Receptaculum seminis

Hx. Excretory vesicle. | uterinum.

Ex.T. Excretory tubules. | T.,T;, T2. Testes.

J. Intestinal diverticula. | Ut. Uterus.

K.St. Ovary. | Vg. Vagina.

P.Pr. Pars prostatica. | V.S. Vesicula seminalis.

Prate XXVII.

Lechriorchis validus.

Fig. 1. Young specimen. Ventral view. X 25.

. Adult specimen. Ventral view. X 20.

. Transverse section near ends of intestinal diverticula. X 53.

. Transverse section, immediately in front of ventral sucker. X 50.

. Ovum. X 500.

OP cw be

Prats XXVIII.

Ochetosoma formosum.

. Ventral view. X 30.

. Ovum. X 550.

Dasymetra conferta.

. Ventral view. x 30.

. Transverse section, a little in front of ventral sucker. X 60.

, Ovum. X 5600,

ics!

Hes

oy .

SOM WOH

Vee See nine WEALD ODS

H.Goodehild del et hth. Huth imp.

THE CHINESE TAKIN,

IBVUNSONRUCYATS) IEE IDE ORY.

ON MAMMALS FROM CENTRAL CHINA, 687

32. The Duke of Bedford’s Zoological Exploration of astern

Asia.—XIV. On Mammals from Southern Shen-si,

Central China. By Otprimip THomas, F.R.S., F.Z.S.*

[Received and Read April 25, 1911.]

(Plate X XTX. f)

As already indicated in paper No. XIII. of the present series +,

Mr. Malcolm Anderson and his party, working on behalf of the

Duke of Bedford, obtained a considerable number of mammals in

Southern Shen-si before going on to Kan-su and Sze-chwan,

whence the series described in that paper was collected. Owing

to delay in transport, however, a large part of the Shen-si

collection has only recently arrived, and the diagnoses of J/yotis

myosotis ancilla, Microtus nux and M. johannes § are all that have

been published upon it.

The present paper gives a list of all the specimens obtained in

8. Shen-si by Mr. Anderson between his second landing in China

in the autumn of 1909 and his move on into Kansu in 1910.

The regions explored were, firstly, the district round Shang-

chou, 8.E. Shen-si (about 33° 40’ N., 110° 20' E.), and, secondly,

the important mountain Tai-pei-san (about 34° N., 107° 30’ E.),

one of the sacred Chinese mountains, another being Omi-san,

Sze-chwan, where at a later period Mr. Anderson obtained the

many new species described in my Sze-chwan paper.

Of the majority of the Shen-si specimens there is little new to

record, as Mr. Anderson had obtained the same species on

his previous visit to the more northern part of the province. But

in any case their interest is dwarfed by the discovery on Tai-pei-

san of a magnificent species of Takin, quite different from the

known W. Chinese species Ludorcas tibetanus, and both in interest

and beauty one of the most striking {mammals that it has ever

been my good fortune to describe.

Besides this fine animal, of which a coloured figure is given

(Pl. XXTX.), Mr. Anderson obtained a new Badger, a new Pika,

and a new Vole ||.

1. RHINOLOPHUS FERRUM-EQUINUM Schreb.

@. 2079, 2080, 2081. Shang-chou District, S.E. Shen-si.

* Published by permission of the Trustees of the British Museum.

+ For explanation of the Plate see p. 695.

P. Z. S. 1911, p. 158.

» P.Z.S. 1910, p. 635.

|| The complete account of these new forms appears here, but the names and

preliminary diagnoses of the species underlined were published in the ‘ Abstract,

No. 95, 1911.—Epitor,

48*

Crt

688 MR. OLDFIELD THOMAS ON

9. Myoris MYOSOTIS ANCILLA Thos.

3. 2082, 2083, 2084. 9. 2085. Shang-chou Dist., S.H.

Shen-si.

The typical series (cf. P.Z.S. 1910, p. 636), no. 2082 (B.M.

No. 10.5,.2.4.) the type.

3. CrocipuRA ATTENUATA M.-Edw.

g. 2009. King-tze-Kwan, 8.W. Honan. 850’.

4, CrocipurA coREx Thos.

3g. 2185. 30 miles S. of Feng-hsiang-fu, 8. Shen-si. 7000’.

5. FELIS FONTANIERI M.-Edw.

2. 2038. Shang-chou District, 8.E. Shen-si. 3000’.

This fine Leopard is a valuable accession to the Museum

collections.

6. Fexis microtis M.-Edw.

$. 2176. 30 miles 8. of Feng-hsiang-fu, 8. Shen-si. 3600’.

7. VIVERRA ZIBETHA, subsp. ?

9192. Native skin. 40 miles N. of Han-chung-fu, Shen-si.

This specimen does not agree with the description of the Civet

called V. filehneri by Matschie, but how far the colour-characters

used by him are likely to be diagnostic in so variable a group I

am not at present prepared to say.

8. PAGUMA LARVATA Gray.

6. 2053. Shang-chou Dist., S.E. Shen-si. 2300’.

9. VULPES sp.

3. 2178. 30 miles S. of Feng-hsiang-fu, 8. Shen-si. 3600.

10. Lurreoa srpirica Pall.

3s. 2070. Shang-chou Dist., 5.E. Shen-si.

@. 2118. Si-ngan-fu, Shen-si. 1200’.

11. ARCTONYX LEUCOLEMUS ORESTES.

Thos. Abstr. P. Z.8. 1911, p. 27 (May 2).

Q@. 2191 (young adult). Tsin-ling Mts., 34° N., 107° 45' E.,

S.W.Shen-si. Alt.12,000'. 25 January,1910. B.M. No.11.6.1.6.

Type.

Distinguished from the true lewcolemus of Peking by the

following characters :—

Dark mark enclosing eye not broadly projected forwards and

downwards to the upper lip, but practically confined to a strong

spectacle-mark, barely half an inch broad, surrounding the eye ;

a narrow and indistinct line only running forwards to the base

of the whiskers, the upper lips quite white. Dark patch behind

mouth much reduced, a trace only of it running forward to the

MAMMALS FROM CENTRAL CHINA. 689

angle of the mouth, separated from its fellow of the opposite side

by a white interramial space over an inch broad. Light throat-

patch not really white, but brownish white, figured in lewcolemus

as snowy white. Light patch in front of ear less prominent,

continuous with but darker than that under the eye. White

ear-rim much broader and more prominent. Back more broadly

washed with whitish than appears to be the case in lewcolemus,

the dorsal hairs white for their terminal 15-20 mm., while only

their points are said to be white in lewcolemus. Tail wholly

white.

Dimensions of the type, measured in the flesh :—

Head and body 570 mm.; tail 195; hind foot 94; ear 45.

Skull: condylo-basal length 132 mm. ; basal length 123; greatest

breadth 72; interorbital breadth 29; palatal length 86 ; greatest

diameter of m’* 16,

Hab. and type as above.

The British Museum had previously possessed no examples of

the N. Chinese Arcéonyx, so that this fine specimen is a valuable

accession. On account of the differences above detailed I cannot

refer it to the true lewcolemus, but think it represents a special

subspecies, as is the case with so many other Shen-si mammals.

12, PETAURISTA ALBORUFUS * M.-Edw.

2194. Native skin. Near Pao-Ning-Fu, N. Sze-chwan.

13. Trocoprerus xANTHIPES M.-Edw.

2055. Shang-chou Dist., S.E. Shen-si.

14. SciuROTAMIAS DAVIDIANUS M.-Edw.

3g. 2022, 2037, 2052, 2065, 2066, 2067, 2069, 2087, 2095,

2104. 9. 2054, 2059, 2096, 2105. Shang-chou Dist., 5.H.

Shen-si.

3. 2106, 2115. Ching-ling Mts., Lo-nan-hsien, S. Shen-si.

@. 2107. Ching-ling Mts., Lo-nan-hsien, 8. Shen-si.

15. EPIMys CONFUCIANUS, subsp.

3. 2034, 2063, 2064. 9. 2024, 2033, 2035, 2036, 2039.

Shang-chou Dist., S.E. Shen-si.

6. 2114. Ching-ling Mts., Lo-nan-hsien, 8. Shen-si.

@. 2010. King-tze-Kwan, 8.W. Honan.

S. 2012,2014. 2. 2015,2016. Shan-nan-hsien, 8.E. Shen-si.

So. 2122, 2123, 2137, 2140, 2143, 2144, 2152, 2161. 2. 2162.

30 miles 8. of Feng-hsiang-fu. 3600’.

In my previous paper the Kan-su specimens of this group were

assigned to #. confucianus luticolor, though tending to intergrade

with the typical H. confucianus of Sze-chwan. But now, on

laying out the whole of the N. China representatives of the group,

* Petaurista is masculine, this being one of the numerous names which by their

deceptive form render desirable the suggested convention that all generic names in

zoology should be treated as masculine. See Stebbing, ‘ Knowledge,’ xxxiii. p. 259,

1910.

690 MR. OLDFIELD THOMAS ON

I find that the true Z. ¢. luticolor is a pale desert form peculiar

to the region towards the Ordos desert, that the present 8.

Shen-si specimens are, as is geographically correct, intermediates

between the /. c. sacer of Shantung on the east and the Kan-su

form on the west, and that the latter is sufficiently differentiated to

have a special subspecific name of its own. It may be called —

HPIMYs CONFUCIANUS CANORUS, subsp. n.

General colour approximating to “ clay-colour,” slightly darker

and more tawny than in sacer, much darker than in Jluticolor,

lighter than in confuctanus. Median darker dorsal line averaging

more distinct than in confucianus, less than in sacer. White of

under surface more markedly tinged with buffy or cream-colour

than in the other subspecies (but there is a doubt as to how much

this fades after death). Metatarsals without, or with but slightly

marked, darker patches. Tail with the dark colour of its upper

surface passing nearly or quite continuously to the end; its tip

hardly so heavily tufted as in sacer.

Skull about as in confucianus and luticolor, smaller than in

sacer.

Dimensions of the type, measured in the flesh :—

Head and body 120 mm.; tail 180; hind foot 27; ear 21°5.

Skull: greatest length 34°5 mm.: condylo-incisive length 30:5 ;

upper molar series 6.

Hab. Southern Kan-su, grading eastwards into sacer and south-

wards into true confucianus. Type from Wen-hsien Country,

S. Kan-su.

Type. Adult female. B.M. No, 11.2.1.110. Original number

2282. Collected 6 May, 1910.

16. Mus waGnert Eversm.

3. 2094. Shang-chou Dist., S.E. Shen-si.

17. APODEMUS SPECIOSUS PENINSUL& Thos.

3. 2046, 2076, 2077. ©. 2045, 2047, 2092, 2093, 2103.

Shang-chou Dist., S.H. Shen-si.

3. 2113. Ching-ling Mts., Lo-nan-hsien, 8. Shen-si.

Q. 2126, 2132, 2136. 30 milesS. of Feng-hsiang-fu, 8. Shen-si.

3. 2167, 2169, 2172. 9. 2166, 2168, 2174. Tai-pei-san.

10,600’.

18. APODEMUS AGRARIUS PALLIDIOR Thos.

3. 2030. 2. 2031, 2040, 2057, 2058, 2061. Shang-chou

Dist., S.E. Shen-si.

3. 2019. 9. 2020. Shan-nan-hsien, S.E. Shen-si.

@. 2108, 2109, 2112. Ching-ling Mts., Lo-nan-hsien, 8.

Shen-si.

96,99. 30 miles S. of Feng-siang-fu, S. Shen-si. 3600’.

19. Micromys minutus M.-Edw.

3. 2056. Shang-chou Dist., S.E. Shen-si.

MAMMALS FROM CENTRAL CHINA. 691

20. CriceruLus Trrron Thos.

@. 2011. King-tze-Kwan, S.W. Honan.

3. 2013, 2018. ‘Shan-nan-hsien, 8.E. Shen-si.

2. 2028, 2029, 2062. Shang-chou Dist., S.E. Shen-si.

21. CRICETULUS ANDERSONI Thos.

3S. 2116, 2117. Si-ngan-fu, 8. Shen-si. 1200’.

22. MicroTus CALAMORUM SUPERUS.

Thos. Abstr. P. Z.S. 1911, p. 27 (May 2).

6. 2121, 2127, 2128, 2129, 2138, 2139, 2147, 2154, 2163,

2181, 2182, 2183. .

On 2119, 2120, 2130, 2146, 2155, 2164, 2179, 2180. 30 miles

8. of Feng-hsiang-fu, S.Shen-si. 3600’.

A larger longer-tailed race of the Nanking Vole.

Fur longer and finer than in true calamorwm, the hairs of the

back about 16 instead of 10 mm. in length. Colour quite as in

calamorum, except that all the specimens have a dull buffy suffusion

in the surface-colour of the abdomen, this being only the case in

certain of the younger examples of calamorum, the older ones

having the belly clear greyish white. Tail longer than in cala-

morum, the usual length in adult specimens 60-63 mm., as against

52-53 in that animal, its coloration more prominently bicolor.

Skull like that of calamorwm, but slightly longer than in speci-

mens of similar age. Nasals longer. Bullw generally larger.

Dimensions of the type, measured in the flesh :—

Head and body 130 mm.; tail 63; hind foot 24; ear 13.

Skull : condylo- -basal lensth 33 mm. ; condylo-incisive length

33°2; zygomatic breadth 17:8 ; nasals 8°5 x 3°6.

Type. Adult male. B.M. No. 11.6.1.45. Original number

2163. Collected 7 January, 1910.

This Shen-si representative of the Lower Yang-tze Reed-Vole

is distinguishable by its longer tail and longer fur, the latter in

obvious correlation with the greater altitude at which it is found.

23. MIcROTUS MANDARINUS M.-Edw.

2. 2032. Shang-chou Dist., 8.E. Shen-si.

Skull. S.E. Shen-si.

The specimens referred to in the description of J/icrotus

johannes (P. Z.8. 1910, p. 637).

24. Microtus (CARyomys) Nux Thos.

3. 2041, 2042, 2043, 2072, 2073, 2088, 2089, 2090, 2097,

2098. 2. 2044, 2050, 2091, 2100. Shang-chou Dist., S.E.

Shen-si.

3. 2110, 2111. Ching-ling Mts., Lo-nan-hsien, S. Shen-si.

Described from these specimens, P. Z.5. 1910, p. 636. No.

2089 (B.M. No. 10.5.2.79) the type.

692 MR. OLDFIELD THOMAS ON

25. Microrus (Caryomys) EvA Thos.

3S. 2173. Tai-pei-san, Tsin-ling Mts., S.W. Shen-si. 10,600’.

The typical series was obtained near Tati-chow, Kan-su, at a

similar elevation.

26. Lepus swinHoE! Thos.

3. 2145. 30 miles 8. of Feng-hsiang-fu, 8. Shen-si. 3600’.

Represents L. filchneri Matsch., and also the Tai-pei-san

subspecies “ suppressed ” by Dr. Allen*, to which he, nevertheless,

attaches the name L. swinhoer brevinasus.

Of all the many examples of ZL. swinhoei obtained by

Mr. Anderson, whether topotypes from Shantung or from other

localities, this specimen is one of those that agree most closely

with the type collected by Swinhoe, owing to its being a dark-

coloured individual, with its general buffy coloration more or less

suffused with pinkish.

27. OCHOTONA SYRINX.

Thos. Abstr. P.Z.S8. 1911, p. 27 (May 2).

g. 2170. @. 2171. Tai-pei-san. 10,600’.

(2) S. 2071 (young). Shang-chou Dist., 8.E. Shen-si. 3300’.

Related to O. cansa, but larger. Bulle smaller.

Size distinctly larger than in O. cansa and O. sorella. Fur (in

winter pelage) long and soft, not very thick ; hairs of back about

16-17 mm. in length. General colour more blue-grey throughout

than in cansa; head grey tinged with clay-colour ; nape ‘‘smoke-

grey”; dorsal area darker grey tinged with brown ; rump nearly

“mouse-erey.” Flanks near ‘“broccoli-brown.” Under surface

greyish, the bases of the hairs slaty, their tips greyish white.

Ears greyish brown, their proectote blackish, their rims white ;

the tuft of long hairs at their inner base nearly “ cinnamon.”

Hands and feet white above, the thickly haired palms and soles

‘“‘ smoke-grey.”

Skull of the same general type as in O. cansa, of similar flat-

tened form, the palatal foramina not subdivided. Size distinctly

greater throughout ; nasals of about the same shape, not narrowed

as in QO. sorella. Interorbital space and brain-case broader.

Bulle conspicuously smaller or at least lower, those of O. cansa

being very large for so small a species.

Dimensions of the type, measured in the flesh :—

Head and body 142 mm.; hind foot 28:5; ear 16.

Skull: greatest length 36°5 mm. ; condylo-incisive length 33°5 ;

zygomatic breadth 18°5; nasals 11°8x 5-4; interorbital breadth

4-3; breadth of brain-case 15:7; palatilar length 12; palatal

foramina 9°2 x 3:7 ; upper cheek-tooth series (alveoli) 7:1.

Hab, Tai-pei-san.

* Bull. Am. Mus. N. H. xxvi. p. 427, 1909.

MAMMALS FROM CENTRAL CHINA. 693

Type. Adult male. B.M. No. 11.6.1.59. Original number

2170. Collected 9 January, 1910.

This is probably the species recorded as O. cansa by Dr. Allen *,

who had, however, only “skins with fragmentary skulls” for

comparison with the single type of Dr. Lyon’s species. The

present perfect specimens, compared with the topotypical series

of O. cansa obtained in Kan-su by Mr. Anderson, show that the

' species is readily distinguishable, both by its greater general size

and its much less swollen bulle.

No. 2071 is too young for certain determination, but it is

interesting as still retaining in place the minute milk-predecessors

of the small posterior incisors. The milk-teeth are pressed close

against the front face of the permanent teeth, between these and

the large incisors in front of them.

28. Sus sp.

@. 2186. 30 miles S. of Feng-hsiang-fu, 8. Shen-si.

In the state of confusion to which the late Pere Heude

reduced the systematic arrangement of the Chinese members of

Sus, as with every other genus with which he dealt at any length,

it is impossible to give a satisfactory determination of this Wild

Boar at present. It probably represents the Sus oxyodontus of

Heude, from the Upper Han, but whether that is or is not a valid

species I am unable to express an opinion.

29. Buporcas BEDFoRDI. (Plate X XIX.)

Thos. Abstr. P. Z.8. 1911, p. 27 (May 2).

3. 2175. 9. 2189, 2190. Tai-pei-san. 10,000’.

A wholly pale buffy species, practically without darker

markings.

Fur longer and richer than in the available specimens of B. tebet-

anus, but the dates of these are not known, while the examples of

B. bedfordi were killed in mid-winter; hairs of sides of neck

attaining 7-8 inches, and those of back 3-4. General colour a

beautiful glossy golden-buffy, more cream-buff in the females, more

tending towards ochraceous in the male; at least at the ends of

the hairs, their bases being still creamy or whitish. Darker

markings on muzzle, ears, hinder back, and limbs, characteristic

of B. tibetanus, practically absent. No. 2190 (2) entirely without

darker hairs on the muzzle, 2175 (¢) with a few, and 2189 (9?)

with more, but the imeonspicuous darker patch so formed very

different from the large and prominent black mask characteristic of

B. tibetanus. Dorsal line not darkened at all, its hairs elongated

and more strongly buffy in the male, not differentiated at all in

the females. Limbs not obviously darkened terminally, though a

few isolated darker hairs are present on the carpus and tarsus.

‘ * Bull. Am. Mus. N. H. xxvi. p. 427, 1909.

694 MR. OLDFIELD THOMAS ON

Tail bushy, a few darker hairs on it in No. 2189, soiled ochraceous

buffy in the others.

Skull apparently rather smaller, and its nasal region less vaulted

than in either tawicolor or tibetanus, but, owing to differences in

the ages of the specimens available, a satisfactory comparison 1s

not at present possible.

Dimensions of the male, measured by Mr. Anderson in the

flesh :—

Head and body 1925 mm. ; tail 23; hind foot 340; ear 127.

Skulls :—

3 2

(young adult). (adult).

Condylo-basal length ....................0006 389mm. 381 mm.

Ly Soma tich Ocead ble sne eae. oseee tae eere 176 164

Height of nasal convexity above middle

OL OL NCTC reer a Nan, arn ennm AR Ion Ltos 118 113

Length of muzzle, to front of p*......... 113 115

Palatal Mlemothim steers eee cere eran cere ter 245 24)

Greatest spread of horns, on outer edge... 401 308

Type. Adult female. B.M. No. 11.6.1.64. Original number

2190. Killed 15 January, 1910.

The discovery of this splendid animal, whose golden-bufty colour

renders it by far the most beautiful of its genus, is of the highest

interest, and it is with great pleasure that I name the species in

honour of the Society’s President, during whose exploration of

Eastern Asia it has been obtained. Mr. Anderson himself seems

to have thought the occurrence of Takin on Tai-pei-san of special

interest, and believed that they would probably prove to be new.

He says: ‘‘ The herds on Tai-pei-san are isolated by some hundreds

of miles from the nearest others we could hear of, and as I could

not learn that any other foreigner has hunted them on Tai-pei, I

believe the chance for a new species is good.”

As a matter of fact, however, specimens had previously been

obtained and had passed into the possession of the American

Museum of Natural History at New York. But these were quite

young, and showed, as it was not unnatural that the young should

show, more or less of the normal coloration of the group, with

blackish muzzle and extremities, and therefore in recording them

Dr. Allen* saw no reason to suppose them different from

B. tibetanus. The practically unicolor condition of b. hedfordi

proves therefore to be a characteristic of the adult, a fact which,

in view of the peculiar specialization of such a colour, is not at

all surprising.

Even in B. tibetanus, as shown by Milne-Edwards’s figure ft, the

young is very materially darker than the adult.

That the Takin of Tai-pei-san and other parts of the Pe-ling

range should be different from that of the mountains of Sze-chwan

* Bull. Am. Mus. N. H. xxvi. p. 4256, 1909.

+ Rech. Mamm. Atl. pl. 74.

MAMMALS FROM CENTRAL CHINA. 695

is only in accordance with the indications furnished by the

remainder of Mr. Anderson’s collections, as brought out in the

paper No. XIII. of the present series *.

With regard to other names that have been given in the genus

Budorcas, Mr. Lydekkert has shown that both sinensis and mitchella

are synonymous with tbetanus, so that there appears to be no

existing name which comes into question in now describing this

beautiful species.

It may be noted that on each side of the withers of the two

females there isa large patch of grey hairs, these hairs being horny

whitish, curiously ringed or beaded with black. Similar hairs

have been found on a specimen of B. tibetanus, and so would

appear to be natural and not discoloured artificially as I at first

supposed. No such patch is present on the male.

Mr. Anderson has sent me the following further notes on this

animal :—

“'Takin.—Found in large herds on Tai-pei-san, where it lives

in precipitous places at from 9,000 to 11,000 ft. altitude. Its

food is bamboo grass; a small bamboo very common at these

altitudes. Although apparently clumsy, I found these animals

very alert, and capable of picking their way very deftly and

rapidly down steep mountain sides. A herd my companions

and I saw seemed to contain about 40 individuals ; Chinese

hunters reported another herd of 80. These herds seem quite

isolated on the peaks of Tai-pei-san. Travelling westward we

did not hear of any Takins again till we reached Pie-kou in

Southern Kansu, and I am in doubt whether this was the same

animal or not.

Chinese name :—Pan-yang; at Tai-pei. The species in Sze-

chwan is called Yei-nu=Wild Cow. The name Pan-yang is also

applied to a mountain sheep or goat.”

30. NEMORHADUS sp.

@. 2188. 30 miles 8. of Feng-hsiang-fu, S. Shen-si. 10,000’.

Near the Long-tailed Goral (1. caudatus M.-Kdw.).

31. CAPREOLUS BEDFORDI Thos.

3. 2153 (young). 2. 2177, 2187. 30 milesS. of Feng-hsiang-

ily BOOP.

EXPLANATION OF PLATE XXIX.

The Chinese Takin (Budorcas bedfordi). Male.

* P.Z.S. 1911, p. 158. + P. Z.S. 1908, p. 795.

696 LT.-COL. N. MANDERS ON THE

: EXHIBITIONS AND NOTICES.

May 9, 1911.

E. G. B. Meapse-Wa po, Esq., Vice-President,

in the Chair.

My. R. I. Pocock, F.R.S., F.L.8., Superintendent of the Gardens,

exhibited some of the hair of the “puppy coat” of a Grey Seal

(Halicherus grypus), which was caught at Barmouth, in Merioneth-

shire, at the end of April. When received at the Gardens at

the beginning of May this Seal was covered, with exception of

the head and flippers, with longish woolly white hair, the last

of which was moulted on May 7th. Most authorities state that

Grey Seals are born in the autumn, not later than about the

middle of October, and that the puppy coat is shed from a month

to six weeks later. Allowing six weeks for the retention of its

puppy coat, this Welsh Grey Seal must have been born near the

middle of March, a date in tolerably close agreement with the

date, namely the end of February, given by Cneiff for the birth

of these Seals in the Gulf of Bothnia. It is, therefore, quite clear

that these Seals breed both in the early spring and the autumn.

My. E. G. BouLencer exhibited some living male specimens of

the Midwife Toad (Alytes obstetricans) carrying the eggs. He

also exhibited a number of the detached eggs to show the manner

in which they were strung together.

Mr. A. E. Anprrson exhibited a large number of photographs

of the more important fossil mammals in the Department of

Vertebrate Paleontology of the American Museum of Natural

History, New York, showing the methods of mounting fossil

skeletons. For comparison, a set of photographs was exhibited

with the skeleton supports eliminated from view, thus adding to

the pictorial value of pose in the specimens.

PAPERS.

33. An Investigation into the Validity of Miillerian and

other forms of Mimicry, with special reference to the

Islands of Bourbon, Mauritius, and Ceylon. By NEVILLE

Manpers, Lieut.-Colonel, R.A.M.C., F.Z.S., F.E.S.

[ Received May 8, 1911: Read May 9, 1911.]

Naturalists generally and the majority of entomologists are

disposed to accept a broad view that mimicry is caused by

VALIDITY OF SOME FORMS OF MIMICRY. 697

natural selection, but beyond this some are not prepared to go.

Others believe more or less implicitly m cryptic mimicry, either

active or passive, and that peculiar form of it known as Batesian

mimicry, that is, the resemblance for protection of a palatable to

an unpalatable species. And there are others again who, believing

in these, consider that Miillerian mimicry, that is, the resemblance

of unpalatable species for mutual protection brought about by the

tasting experiments of young reptiles and birds, plays a very

large part, even an overwhelming one, in the production of

mimicry.

The theories of the two great naturalists Bates and Miller

have now been before us for a great number of years ; but both,

the latter more especially, base their claims to recognition on

indirect evidence and not on experimentsand investigation in the

field. The exponents of these two theories maintain that though

direct evidence is largely absent, yet on no other reasonable

hypothesis can these remarkable cases of mimicry be explained.

The opponents, on the other hand, hold the view that as direct

evidence is possible though admittedly difficult to obtain, it ought

to be produced before either theory can be admitted as proved,

and until it is forthcoming they remain either actively hostile or

passively sceptical. It was with the feeling that both the sup-

porters of these theories and those opposed to them were equally

desirous of reaching some finality in this vexed question, which has

now been before them in one form or other for half a century—a

question which, with its periodical exhibition of violent eruption

and deceptive quiescence much resembles an Iceland geyser—that

I have during the past five years devoted as much of my time as

was practicable to the study of insectivorous birds and reptiles ;

and I have taken as my areas of investigation the islands of

Bourbon, Mauritius, and Ceylon, because the question is less

complicated on an island of small or moderate dimensions than on

such an extensive area as Africa or South America.

Before I left England in 1908 I had the great advantage

of being taken over the National Collection of Butterflies by

Mr. Guy Marshall, who, with his unrivalled knowledge of the

subject and persuasive powers, almost then and there made me

throw in my lot with the supporters of the Miillerian theory, but

in the midst of his arguments recollections of scenes in tropical

jungles obtruded themselves, and I was left in an irritating

condition of mingled belief and incredulity. A fairly extensive

reading of the whole subject consequently did little to convince

me, and my personal bearing towards both Batesian and Miillerian

mimicry was that the verdict must be the unsatisfactory one of

‘“‘not proven.”

I propose taking each of the islands in turn, enumerating the

reptiles and birds with their habits and distribution, directing

attention to the more striking cases of mimicry, and endeavouring

to ascertain on such data how far these theories are negatived or

sustained.

698 LT.-COL. N. MANDERS ON THE

Bourson.

Bourbon is a small circular island less than forty miles in

diameter, lying some three hundred miles from the east coast of

Madagascar. It has only twenty-two species of butterflies and

certainly one case of mimicry, which is very striking and quite

peculiar. It is that of a female Papilio (phorbanta) resembling

a Huplea. The group to whieh this Papilio belongs is green in

both sexes, but in this insect the female is dark brown and

resembles more or less closely the brown Huplea (goudoti)

occurring in the island. ‘There is no occasion to go into details,

as I have already brought them to the notice of the Entomolo-

gical Society in its Proceedings and Transactions, 1908, and have

figured both butterflies ; but I may add that they are essentially

insects of the littoral, common on one small portion of the coast,

particularly in gardens on the outskirts of. St. Denis, but very

rarely found above 1,500 feet or 2,000 feet. They are generally

associated.

Bourbon has no lizards with the exception of one introduced

species which is very rare. I was fortunate enough to find a

specimen; and I should say, judging by somewhat similar Ceylon

lizards, that in all probability butterflies would form part of its

diet, but it is far too rare to have any marked effect on the

butterfly population.

The following is a list of the insectivorous birds given to me by

a resident naturalist, which, so far as my knowledge extends, is

complete :—

1. The Sparrow. Passer domesticus.

. The Mynah. Acridotheres tristis.

. Zosterops (Malacirops) borbonica.

. Zosterops hesitata.

. Bec-Bee. Pratincola (Motacilla) sybilla.

. Coq des bois. Trochocercus borbonicus.

The Wheat Swallow. Phedina borbonica.

. The Little Grey-rumped Swiftlet. Collocalia francica.

. Le Merle cuisinier. Lalage (Oxynotus) newton.

With regard to these, two, the Sparrow and Mynah, have been

introduced; on the former I need make no remark, it has the

same habits as its English relations. Two efforts have been made

in the last hundred years to instal the Mynah, but without success,

and this because it is considered a desirable morsel by the natives

and is mercilessly trapped and consequently very scarce; I saw

only one pair during my stay in the country.

Both species of Zosterops are very small birds, no bigger than

the English wren, and are found either singly or in small

family parties of five or six, flitting and creeping about the shrubs

after the manner of our long-tailed tits; they feed on nectar and

small insects. The Motacilla or Chat is of the same size and

much the same colouring as the Whinchat, and quite possibly feeds

VALIDITY OF SOME FORMS OF MIMICRY. 699

on the smaller butterflies, but would scarcely tackle an insect

considerably larger than our Swallowtail, but of this I have no

evidence. I saw it frequently in the gorge leading up to Salazie

and on the hills round St. Denis, but I saw nothing of it in the

neighbourhood of the town.

The Flycatcher (Z'rochocercus borbonicus) is the same species as

oceurs in Mauritius; it is said, on the authority of M.de Charmoy,

to feed on diptera and by preference on mosquitoes. It is

quite a small bird.

The Wheat Swallow (Phedina borbonica) has precisely the same

habits as regards its food as the English species ; it. appeared to

me to be fairly plentiful. It also occurs in Mauritius.

The Little Grey-1rumped Swiftlet (Collocalia francica) is the well-

known species that forms a nest of inspissated saliva. It occurs

in Mauritius. It is quite a small bird, decidedly smaller than our

Sand-Martin, and may occasionally snap up a small Lycenid.

Le Merle cuisinier, or Tui-tuit —Lalage (Oxynotus) newton.

I am unable to say whether this bird is abundant or not,

or whether it destroys butterflies. I did not come across it

in Bourbon, so I am inclined to doubt its being particularly

common. It has the same habits as the Mauritius ZLalage

rutiventer.

In the absence of any living bird it occurred to me that

possibly the extinct Bourbon Starling might have been the prime

factor in producing this case of mimicry, and I therefore wrote

to my venerable friend Dr. Jacob de Cortimoy for information

concerning it. | He is now verging on his ninetieth year, and is

probably the only one now living who has seen this bird alive ;

his letter is so interesting that I need make no apology for

transcribing a portion of it.

“T have known the bird you ask me about since childhood,

namely the Mregilupus varius (old writers called it F. capen-

sis), which has in fact entirely disappeared ...... When I was a

boy this bird lived in the forests of the interior of the island and

never set foot nor wing in towns or inhabited places. It rermained

faithful to the forests where it was bred, which it enlivened

with its clear notes. I used to hunt it then at an age when one

is pitiless. I can seeit now, a little larger than the white black-

bird, with a white crest on the head in the case of the male,

the wings a blackish grey on the upper surface, the beak and

feet yellowish. By no means shy, it was not frightened even by

the sound of firearms, and after a regular slaughter one went

off with dozens of these poor victims in one’s game-bag.

“After ten years spent in Paris I did not find a single one in

the forests where formerly they flew about in flocks. All ruthlessly

destroyed. I shall never forgive myself for the part, slight

though it was, which I took in the matter. I lost my taste for

sport and the best bag would not tempt me....... We will

now consider the feeding habits of this bird. Having raised

several in the aviary, I can risk talking about it though I never saw

700 LT.-COL. N. MANDERS ON THE

one feeding in the wild state. In my aviary its food consisted of

bananas, potatoes, and choux-choux, Sechiwm edule (boiled). But

when left to its own instincts, it must, like the other winged

denizens of the forest, have eaten insects as is done by its

companion in the forests, the Bourbon Blackbird (Hypszpetes

olivaceus) *, and as is the habit of most fruit-eating birds.”

This is a sad commentary on our boasted civilization, and I have

only to add that not half a dozen skins are now in existence.

This bird cannot have been in any way the cause of this

mimicry, as it inhabited the forest-covered hills in the interior of

the island, where these butterflies do not occur.

Papilio phorbanta female was figured by Boisduval in 1833 and

differs in no way from recent specimens. We may therefore infer

that the factor or factors which primarily induced this change of

colouring are still active ; but as there is no reptile or any bird

now living which attacks these butterflies as adults, it is difficult

to accept this as an effect produced by them. And it would

seem that the young of existing birds, with possibly one or two

exceptions, would be too small and feeble to attack these large

butterflies during their tasting experiments.

T now turn to the island of Mauritius, which lies some eighty

miles to the north of Bourbon and which is visible from there on

a clear day at certain seasons of the year.

MAURITIUS.

Mauritius has no arboreal lizards, and but one species of ground-

lizard, in appearance very like the English Sand-Lizard. It is

confined to the coast, and is I believe somewhat uncommon. I

found it in some numbers on the uninhabited islet of the [le de

la Passe at the entrance of Mahébourg harbour. It was quite

tame, even confidential, and made no display of timidity in taking

and eating a small piece of boiled potato presented to it on the

end of a fork. We may, I presume, regard this lizard as an

indiscriminate feeder !

My friend M. d’Emmerez de Charmoy, Director of the Port

Louis Museum, a Mauritius gentleman who has an unrivalled

knowledge of the fauna, and who has tracked, shot, skinned,

dissected and mounted the whole of the splendid collection of

Mauritius birds in the Port Louis Museum, has very kindly

favoured me with the following list and notes on the insectivorous

birds; it can be taken as complete, and I doubt whether any

local fauna of a tropical island is so completely known as is this

to M. de Charmoy.

T have added a few notes of my own in square brackets.

1. Le Mangeur de Poule (Tinnunculus punctatus) [| Cerchneis

punctata|.

Feeds preferably on insects rather than on small birds.

* This bird occurs in Mauritius, but was not given to me by the Curator of the

Museum as an insectivorous bird,

VALIDITY OF SOME FORMS OF MIMICRY. 701

I have many times discovered in their stomachs the remains

of locusts, field-crickets, and also stick insects.

[This Kestrel is rather smaller than the English bird and, like

it, is persecuted persistently. It may also feed on butterflies, but

it is so rare that its influence can be little felt; it is entirely

confined to the small portion of indigenous forest now remaining. |

2. Le Merle cuisinier (Oxynotus ferrugineus) |Lalage rufi-

venter. |

Ts essentially insectivorous; I have seen these birds catching

Mantis religiosa and I have found in their stomachs Scarabzei

(Cratopus) and fragments of moths’ wings.

[This bird of late years has become exceedingly rare and is

verging on extinction. On my telling M. de Charmoy that I

had seen a pair in the forest, he congratulated me with as much

fervour as if I had seen a Dodo!|

3. L’Oiseau Banane (Youdia erythrocephala).

Frequents very persistently bananas when in flower, and

captures the minute insects which are attracted by the honey of

these flowers; lives also on the petals of flowers and on small

lepidopterous larvee. |M. de Charmoy considers from his dis-

sections that this bird is incorrectly placed in this genus, which is

essentially a grain-feeding one. |

4. All these species are indigenous and so also are the two

species of Zosterops, Z. mauritiana and Z, chloronota, which live

almost entirely on the larve of lepidoptera. I am unable to give

the names of the kinds they capture, but no doubt they take any

kind of caterpillar.

5. Le Coq des bois (Muscipeta borbonica) | Trochocercus borboni-

cus|. Also indigenous; is an inhabitant of the forests and is

found also along river-courses; it chases diptera by preference

and particularly mosquitoes.

6. Le Boulbul (Pycnonotus jocosus) was introduced in 1892 by

M. Gabriel Reynard and is now to be found everywhere. It is

certainly to be found in great numbers, being often a plague. It

consumes the best fruits and vegetables and the blossoms of

fruit trees. I have often seen it hunting for moths, especially

for Ophiuside, and in the fields of wild indigo it captures Lyceenide.

[The most common butterfly in these fields is Lampides betica. |

7. Le Martin (Acridotheres tristis) was introduced from the

Coromandel coast by M. Boueher des Friyes, and by Pierre a

Mainard (2) into Réunion, to destroy the crickets which ravaged

the islands in 1759. It is found in great numbers in newly

tilled fields hunting after all sorts of insects, and especially after

the eggs of crickets.

The Fringillide cannot be considered insect hunters though

they catch one on the wing when they come across it. They

Proc, Zoon, Soc.—1911, No, XLIX, 49

702 LT.-COL. N. MANDERS ON THE

have not the slightest share in the reduction of local species (of

insects).

There are thirty species of butterflies, but with the exception

of Hypolimnas misippus and Danais chrysippus there is no such

well marked case of mimicry as that of the Euploea and Papilio

in Bourbon.

Professor Poulton has however thrown out the suggestion

that the female Papilio manlius, though green, is approaching

Euplea euphon, as it is of a distinctly brownish green tint

compared for instance with the Madagascar P. epiphorbas. _ Mr.

Trimen also considers that Danais (Amauris) phaedon and the

Euploea mimic each other to a certain extent. Instances of

seasonal dimorphism (cryptic defence) are however numerous.

The evidence I have collected does not seem to justify the

contention that the above instances of mimicry are due to the

depredations of old birds or to the experimental tasting of young

ones. Though no doubt experimental tasting of insects generally

takes place, it would appear that the birds are of such a character

that butterflies would not be attacked except to the smallest

extent.

CEYLON.

T now turn to the island of Ceylon. If the problem we are

considering has so far been simple owing to the small size of the

islands dealt with and their very limited fauna, it is by no means

so in Ceylon, which is far larger, being about two-thirds the size

of Ireland, with a wonderful diversity of hill and plain and equal

diversity of climate. Its general characteristics are too well

known to require repetition, and there are two hundred and sixty

species of butterflies.

Mimicry among Ceylon Butterflies.

Mimicry is by no means uncommon among Ceylon butterflies

and the following will serve as examples of it.

Mimic. MODEL.

Hypolimnas bolina 2. Huplea (several species).

rs misippus 2. Danais chrysippus.

Elymnias fraterna 9. » plexippus.

Argynnis hyperbius 2. © 55

Pareronia ceylonica @. » aglea.

Prioneris sita. Delias eucharis,

Papilio clytia race

lankeswara 3 Q. Huplea (several species).

Papilio clytia form dissimilis. Danais aglea (and allies).

Papilio polytes Q. Papilio aristolochie.

5s ., form romulus Q. Papilio hector.

And the Miillerian combination of the three Eupleeas, core, coreta,

and klugit.

Euplea coreta and £, core, asa reference to Mr. Moulton’s plate

VALIDITY OF SOME FORMS OF MIMICRY. 703

in Trans. Ent. Soc. Lond. for 1908 will show, are extremely alike,

yet under certain circumstances I can recognize them when on

the wing without great difficulty. When herded together in

shady jungle, as is their frequent custom, it is impossible to differ-

entiate them; but when flying singly over an open space, the

former appears a blacker and broader insect with a rather more

flapping flight, by which I can usually distinguish it from £. core.

The resemblance between Danais chrysippus and Hypolimnas

misippus 9 is well known, and I have often found them mixed

together in local collections, but on the wing under ordinary

circumstances differential diagnosis is by no means impracticable :

the female of the latter is almost invariably seen flying close to

the ground selecting favourable situations for oviposition, and

her method of flight, difficult to describe, differs from that of

D. chrysippus, which oviposits on a shrub some four or five feet

in height; when not thus engaged the two are undoubtedly

difficult to distinguish at about ten yards distance, and it is

curious how often the male of H. misippus mistakes D. chrysippus

for a female of its own species. Prioneris sita can at once be

distinguished from Delias eucharis by its rapid darting flight.

A ease of resemblance, though not always stated to be one of

mimicry, is that-of Argynnis hyperbius female and Danais plex-

ippus or D. chrysippus. This was remarked on by Butler so long

ago as 1884 and has been repeatedly noticed since, particularly

by Longstaff and Bainbrigge Fletcher. The resemblance is,

however, entirely accidental, as their habitat in 8S. India and

Ceylon clearly shows. Broadly speaking, if observed above 4000

feet, it will assuredly be 4. hyperbius, if on the littoral and up to

about 4000 feet, almost certainly D. plexippus; it is only on the

confines of each other’s territory that they in any way come in

contact and where an error can be made. Cethosia mnietneri,

Danais ceylonica, and Papilio clytia (dissimilis) fly in the same

localities, that is, in jungle where the light is very flickering. It

is not only very difficult to distinguish ‘them apart, but they are

quite difficult to see, as their black and white marking tends to

make them invisible. They also frequent the outskirts of jungle

and even more open country, and here they are quite easy to

distinguish, particularly P. clytia (dissimilis), which is a larger

and much yellower butterfly.

Mr. T. Bell considers Cethosia to be an unpalatable genus owing

to the nauseous juices and leathery bodies of the species. Huripus

consimilis very closely resembles a Danaid in both sexes, and

would be considered an undoubted case of either Batesian or

Miillerian mimicry: it is not a Ceylon butterfly, and I first

made its acquaintance in the Nilgiris; by its lofty sailing flight,

particularly in the female, I recognized it at a glance from a

Danaid, which rarely or never ascends more than about ten feet

from the ground.

It is more in their peculiar manner of flight rather than in

any difference of colouring that model and mimic can as a rule

4.9*

704 L.-COL. N. MANDERS ON THE

be distinguished; when at rest the difficulty is considerably

greater, and I would draw particular attention how not infre-

quently butterflies of a similar pattern on the under surface of

their wings chose the same resting places, and often the same

bush or branch on which to settle for the night. The following

two notes indicate this.

“ Colombo, July 1909. TI observed at sunset sixteen Zelchinia

viole and some Danais chrysippus with their wings closed, at rest

on a leafless bush; they exactly resembled withered leaves, and it

was exceeding difficult at ten measured paces to distinguish the

species, at about twenty paces it was very difficult to recognize

them as butterflies at all, and at thirty paces they were practically

invisible.”

“Kullar, Nilgiris, 12.4.1910. In a grove of Areca palms

Euplea coreta, £. core, Danais limniace and D, septentrionis simply

swarmed, they were in hundreds and hung in festoons from the

palms. Though many of each kind were on each leaf, they usually

kept together. It was deep shade, and the Euploas seemed to

match the decayed vegetation and the Danaids the green leaves.

There were also many Danais plewippus but no other butterflies.”

With regard to the above quoted instances, Mr. R. C. Punnett,

after a study of them for about six weeks, writes: ‘‘ With the

exception of Argynnis hyperbius and Prioneris sita, 1 have had

frequent opportunities of observing all these cases, and in every

one it has appeared to me that the resemblance is far less striking

when the insects are seen alive than when they are exhibited

pinned out in the orthodox way on cork. I have found that with

very little experience the eye comes to distinguish the mimic from

the model without hesitation. As a rule it is in the mode of flight

that they differ from one another...... My impressions of all

these so-called cases of mimicry which I have been able to see, is

that the resemblances are certainly not sufficiently close to

deceive the eye of a civilized man with a little experience of them.

For that reason I am inclined to doubt whether they would

systematically deceive an enemy brought up among them, whose

means of earning a livelihood depended largely upon the readiness

with which he could distinguish between mimic and model. I

do not wish to deny that in some cases, and upon occasion, the

resemblance may be of service.”

All local entomologists would endorse the above remarks, but

there is this to be said, that Mr. Punnett was aware of these cases

of mimicry before he arrived in Ceylon, and was on the look out

for them; if he had been totally ignorant of them, it would

have taken him some time before he recognized the phenomenon,

but having once done so, he would no doubt have had no further

difficulty.

The detection of certain cases of mimicry would appear to be

greater in some countries than in Ceylon. Colonel Bingham,

writing of the Papilio clytia group, says: “They have nearly all

4 wonderful resemblance to forms of Huplea and Danais, and it

VALIDITY OF SOME FORMS OF MIMICRY. 705

requires a quick eye and some experience to discriminate between

Euplea and Papilio clytia vace panope and between Danas

limniace and the dissimilis form of P. elytia, especially when on

the wing.” In reply to a query of mine on this point Mr, W. F. H.

Rosenberg writes :—‘ London, August 29th, 1910. During my

travels in Colombia and Ecuador I found that mimicking species

did frequent the same places as their models. For instance, the

Dismorphias (Pierines) which mimic certain groups of Ithomiine

butterflies, such as /thomia zelica, were seen flying in clearings

in woods ete., in company with the Ithomias. It is curious to

note, however, that there is a slight difference, difficult to

describe, in the mode of flight of the two groups, so much so

that a trained collector would rarely mistake one for the other.

Again, the S. American Acrvines of the genus dActinote, which

settle in groups on damp patches of sand, have their mimics in

the shape of Nymphalines of the genus Hresia. But while the

Actinotes will allow themselves to be picked up with the fingers,

the Eresias always fly up on the approach of danger.”

The Rev. St. Aubyn Rogers also, in his well known paper on

Kast African butterflies, mentions how often at first he was

deceived by model and mimie, but how after a short acquaintance

he readily recognized them.

But confining myself to Ceylon, the ease with which every case

of mimicry occurring there can usually be detected, causes me to

be in sympathy with those who consider that this constitutes a

great difficulty in the acceptance both of Batesian and Miillerian

mimicry. Both reptiles and birds are well represented, and in

order to afford a complete study of the fauna I propose taking

them in order, first dealing with the reptiles. I can scarcely

hope not to have overlooked some species, but I trust there is no

serious error.

The Lizards of Ceylon.

Five families ave represented; namely Geckonidee, Varanide,

Scincide, Lacertidee, and Agamide. The first three of these are

almost unquestionably of no account in our enquiry. The

Geckonide are mostly nocturnal with but one diurnal species,

Gonatodes kandianus, common in houses at Kandy and confined

to that part of the island. The Geckos I have observed feeding

at night were quite indiscriminate in their captures. Of the

Varanide, two species are large carnivorous lizards, commonly

called iguanas, frequently attaining a length of three or even four

feet. One species when young ascends trees, and Mr. Rosenberg

has seen a Mexican species eating butterflies.

The Scincide has four genera, Acontias, Chalcidoseps,

Lygosoma, and Mabuia. The first has four species with limbs

rudimentary or absent ; in appearance they are very like our slow-

worms but smaller. One species, 4. burtoni, is usually found

under stones, and the others, so far as I have been able to

ascertain, inhabit similar situations. They appear to feed on

“06. OP LT.-COL. N. MANDERS ON THE

small worms. Chalcidoseps has but one species, C. thwartesti, a

small creature about two inches long with very short limbs. It

is not represented in the Colombo Museum collection, and I am

unacquainted with it. Zygosoma has three species, one found in

the hills, the other two common in the low country. Their

forelegs are very feebly developed, in fact almost rudimentary,

and judging by their general appearance they probably have the

same habits as the next genus, Mabwia, which contains two

species, one of which is rather rare; the other, J/. carinaia, is the

well known Brahminy Scink, which so far as my experience goes

feeds almost entirely on ants, I have invariably failed to get it

to feed on butterflies.

The family Lacertidee has but one genus, Cabreta, containing a

single species leschenwultii, very small and rave, found only at

Mullative in the arid north-western district.

The Agamide has five genera, but three are represented by a

single species in each. Otocryptis hivittata, a very small creature,

confined apparently to the wet districts up to 2,000 feet: I have

no personal acquaintance with it. Cophotis zeylanica, also small

and found only in the hills: in captivity it feeds readily on flies,

which it captures after the well known manner of the chameleon.

Lyriocephalus scutatus, a magnificent creature found only in the

outer hill-ranges, where the climate is hot and moist. My friend

Mr. Alervs Hankey, who has kept these species both in captivity

and at large in his garden, informs me that they feed on “ almost

anything— moths, flies, beetles, grasshoppers, worms, and even

boiled vice.” We may conclude that their taste in butterflies,

which they in all probability eat somewhat extensively, is

impartial.

The genus Ceratophora has three species only, one of which I

know, C. stoddartii, found only in the bigher ranges. It feeds

on worms and positively declined butterflies when in captivity.

The genus Calotes has seven species, three of which are some-

what rare; these I do not know nor one other, C. mystaceus. Two,

C. ophiomachus and C. versicolor, are extremely abundant from the

coast to about 3,000 feet, thence upwards nigrilabris takes their

place and is likewise very common. All members of the genus

have probably the same habits and are undoubtedly highly

destructive to insect life. In fact I have little hesitation in

saying that they are the greatest enemies that butterflies have

to contend against, and when in Nuwara Eliya in 1909 I should

have had no great difficulty in making a fair collection of butter-

flies mutilated by them. The injuries were of all kinds, but

mostly a semi-circular piece, the size and shape of the lizard’s jaw,

had been taken out of the hind wing. These mutilated butterflies

indicated no discrimination on the part of their enemy ; perhaps

Argynnis hyperbius was the most frequent victim. My experi-

ments, though not so complete as might be wished, show that

these lizards exercise no partiality; but the butterflies experi-

mented with are those commonly found in the gardens at Colombo,

VALIDITY OF SOME FORMS OF MIMICRY. 707

where Calotes is abundant, and almost all those which occur at

Nuwara Eliya, the fauna of which is very poor.

Experiments with Lizards im Colombo.

The two species of reptiles experimented on belonged to the

genus Calotes (C. ophiomachus and C. versicolor), or blood-sucking

lizards as they are commonly called. In appearance and habits

they are far more like Chameleons than ordinary Sand-Lizards,

and like them have the faculty of changing colour and assuming

on occasions brilliant scarlet, yellow and green, particularly about

the head; but on the other hand they are very quick in their

movements and can run with considerable speed. They are

usually to be found sitting on walls and palings, clinging to the

stems of the longer grasses, and frequently lying in wait for their

prey behind the umbels of flowering shrubs, where their gaudy

colours help to deceive visiting insects. In such positions they

will wait motionless for hours on the chance of prey coming

within reach. But as I was unable to devote a whole day for

such prolonged investigations, I adopted the following method as

being the nearest approach to natural circumstances. I attached

a defunct or moribund butterfly by a long line of fine silk toa

fishing-rod, and thus succeeded with the help of the wind in

bringing the fly within reach of the reptile. The following were

the results from the end of November to the end of December

1908, with the notes I made at the time.

November 24. A male Papilio polytes was waved over a green

Calotes ophiomachus which was clinging to some grass. The lizard

became slightly excited and made a grab at the insect, biting out

a piece of the fore wing and immediately eating it; a second time

it bit a piece out of the hind wing including the tail; the insect

if alive would certainly have escaped on both occasions. Finally

it seized it by the head and ate the remainder.

November 30. A female Papilio polytes of the black and white

variety offered in the same way to another Calotes. This lizard,

which was evidently hungry, became excited when he saw the

butterfly, and made a grab at it and bit out a piece of the fore wing

which it promptly began to eat; the butterfly would have

escaped. While this “fishing” was going on, a male butterfly

attracted by the female flew quite close to the lizard, which made

a dart at it and tore away a piece of its wing, and the butterfly

flew off. A lizard grabs at any part of the butterfly that comes

within reach, and as the grass sways when it moves it is quite

a chance what part is seized; the habits of Calotes therefore

negative any theory of so-called directing marks so far as they are

concerned.

November 25. The wings of the black and white variety of

P. polytes ave quite common in the garden, and I am quite sure

the Calotes devour great numbers of them. I saw C. versicolor

with a live one in its mouth ; on this occasion it was holding the

708 Lt.-CoL. N. MANDERS ON THE

insect by the fore wing—when it saw me it ran to a tree, and [

fancy dropped the insect which, as I could not find it, probably

flew away.

December 4. Saw C. versicolor seize Delias eucharis on the

wing. On this occasion the lizard was in the foliage at the top

of a bush, and sprang out and captured the butterfly as it flew

past.

December 6. Huplaa core was eagerly seized, held in the

mouth for a long time and then eaten.

December 7. Telchinia viole was eagerly seized by the body and

devoured.

December 9. Two Danais chrysippus taken one after the other

by a green Calotes ophiomachus. It watched the butterfly, seized it

by the body, and ate the whole of it after holding it in its mouth

for some minutes.

December 18. Papilio (Menelaides) hector seized as soon as seen,

held in the mouth for some time and eaten slowly. The lizard

seemed very puzzled at the dryness of its meal as the butterfly

had been dead five days, but finally ate it all.

December 20. C. ophiomachus ate a male Papilio polytes.

December 30. Saw Terias hecabe captured by Calotes versicolor.

During these two months butterflies and other insects were very

numerous owing to the rain at the break of the N.E. monsoon.

The tall Mauritius grass in the garden was a place. of refuge for

the butterflies during the heavy rain, and numbers of them could

be seen any morning sunning themselves and sitting with

expanded wings halfway up the grass stems. /P. polytes and

P. demoleus were particularly numerous and several had pieces

taken out of then wings, no doubt by lizards. It was an

interesting fact that so long as the butterflies remained perfectly

still, they were entirely unnoticed by the lizards, though they

might be in close proximity to them. Both these butterflies

rest throughout the night with wings widely expanded.

Experiments with Lizards in Nuwara Eliya, 6,200 ft.

Three species were experimented on, Calotes nigrilabris, Cophotis

zeylanica, and Ceratophora stoddartii. All three are peculiar to

Ceylon and confined to the hill-districts. C. n2grilabris is about

a foot in length including the tail, and is brilliant emerald green

with a black bar across the lips. It has the same habits as

C. versicolor and C’. ophiomachus but is much tamer ; in fact it is

not at all difficult to capture with the hand as it rests on a bush.

Calotes negrilabris.

3.3.09. Ate a ZVerias hecabe greedily, and another shortly after.

3.3.09. The same lizard ate another 7’. hecabe, and another

made frantic grabs at H. core, dangled as usual at the end of a

VALIDITY OF SOMB FORMS OF MIMIORY. 709

string ; when eventually I allowed it to take it, it ate it readily.

The same lizard ate 7’. hecabe.

7.3.09. Another lizard ran out from its bush and caught

Terias libythea. A Papilio aristolochic, a very scarce species in

N’Eliya, caused great excitement in a male and female on the

same bush; they rushed out to capture it, then drew back

apparently frightened at its size and at length would have nothing

to do with it. Offered to another it was seized by the fore wing

and a part only eaten, the reason being that it was frightened of

me. Offered again to the first pair they took no notice of it, but

one made a jump of quite six inches and snapped up a fly which

had settled on a leaf. This shows that they were hungry at the

time.

15.3.09. A fresh Appias galene 9 readily taken, but not so

eagerly as by another which had recently changed its skin and to

which some of the slough was clinging. This was very hungry

and made quite a respectable jump at A. galene ¢ , which it caught

round the body. Immediately afterwards it devoured Catopsilia

pomona.

15.3.09. Offered Papilio (Menelaides) hector to a remarkably

fine lizard, which caught it by the base of the hind wings ; these it

ate very slowly and dropped the rest, no doubt because it was too

dry. The same thing happened with Velchinia viole and another

lizard. These two butterflies do not occur in N’Eliya.

16.3.09. A female moth, Spilosoma melanopsis, Fam. Arctiide,

with remarkably large brilliant crimson body and pink hind

wings, the dull fore wings being removed, was at once taken by

a lizard, which ran some distance after it. It proved a very

considerable mouthful which took quite half an hour to get rid of.

During March, April, and May I noticed a very considerable

number of butterflies, more particularly Argynnis hyperbius, with

pieces taken out of their wings, usually the posterior portion of

the secondaries. I have no doubt that almost all these injuries

were caused by this species of Calotes which is very numerous at

N’Eliya. I might almost say there is a specimen on every bush.

19.3.09. A Huplea core 3 proved very attractive; a half-

grown lizard ran more than a yard out of the hedge and seized it

by the head as it lay on the ground. With the same species a

large male made a dart at Danais fumata attached as usual to a

line of silk, caught it by the hind wing and devoured the whole of

it. A smaller individual seated on a bush of salvia became highly

excited by Polyommatus betica with its wings closed, and ran all

over the plant after 1t, eventually catching it by the body; imme-

diately afterwards it ate three Zerias hecabe, one after the other.

24.4.09. A female lizard,and one I have frequently experimented

on, ran from its hiding place and caught Pyrameis cardui by

the head after considering for a moment its cryptic underside.

Another scrambled over its bush in the usual way and seized

Lethe daretis 2 by the hind wing.

These lizards were very tenacious in their grasp: asa rule, after

710 LT.-COL. N. MANDERS ON THE

capturing their prey, they hold it quietly in the mouth for some

time, but if it struggles they immediately begin eating it. A

renewal of a struggle leads to the same thing.

Cophotis zeylanica.

This lizard is found only in the hill tracts of Ceylon, and is

most frequently found resting on tree-trunks in shady places.

It has considerable power of adapting its colouring to its

surroundings, but not to the same extent as the Chameleon. It is

almost five inches long including the tail, which is usually two or

three inches. It is extremely sluggish in its movements, depend-

ing entirely on its cryptic colouring to escape observation. It is

quite harmless and can easily be taken in the fingers. It eats flies

readily in captivity, but all my efforts, both with specimens at

large and in captivity, to induce them to eat butterflies resulted in

failure, even those with their wings entirely removed seemed only

to frighten them.

The Horned Lizard. Ceratophora stoddarti.

This interesting reptile is also peculiar to the island, entirely

confined to the hills, and I do not think it occurs below 5000 feet.

It varies in colour from a vivid green with black bands (three

only seen) to a uniform brown. It has.the faculty of changing

its colour according to the nature of its environment. Only

once have I found it otherwise than on a tree-trunk in deep shade,

where butterflies very rarely penetrate. All attempts to make it

eat butterflies in the wild state failed, and the following is a note

on a captured specimen :—‘“‘13.4.09. Put numerous flies in its

cage, there appeared. to be a diminution the following morning.

Put Verias hecabe and Pyrameis cardui alive into its cage. It did

not notice the latter as it rested with closed wings on the gravel,

- though within three inches of it; but when it moved its fore

wings up and down—not in and out—it was on the alert and

crept up to it, but before it had made up its cautious mind to

attack, the butterfly flew off, hitting it on the snout in so doing,

which utterly disconcerted it.” I may say that about the only

butterfly in the upper hill district which frequents shady places

is Lethe daretis. Mycalesis and Yphthima do not extend so high.

The chance of a meal off a butterfly is therefore remote.

Dr. Willey informs me that its natural food is small worms.

It would seem then that those who assume that reptiles take no

part in the production of Batesian or Miillerian mimicry are

correct, though further experiments are required.

The Insectivorous Birds of Ceylon.

A few introductory remarks are needed before dealing with the

insectivorous birds. Speaking generally the Mimicrists, if I may

use the term, maintain that birds do eat butterflies largely; the

Anti-Mimicrists that they do not. The evidence of the latter is

VALIDITY OF SOME FORMS OF MIMICRY. (Mal

necessarily negative, and we must therefore try to find some

reasonable standard by which we can judge whether a particular

bird is a butterfly eater or not. We shall not be far wrong if we

employ the criteria accepted by Mr. Guy Marshall in his paper

‘“‘ Birds as a factor in the production of Mimetic Resemblances

among Butterfiies” (Trans. Ent. Soc. Lond. 1909), only in this

case in a contrary sense. Speaking of the want of real evidence

on the part of the Anti-Mimicrists, he says:——“* When a naturalist

who has spent some time in the tropics expresses a decided

opinion to the effect that birds do not normally eat butterflies,

because he has never observed them doing so, it is incumbent on

us, before accepting his evidence as having any real scientific

value, to satisfy ourselves that he has made a systematic and

thorough investigation of the subject, and that his views are not

based merely on casual and inadequate observations. For in a

matter of this kind there is grave danger that absence of evidence

may be due simply to lack of observation. If a collector main-

tains that birds do not eat butterflies, we are justified in asking

him for @ full list (italics mine) of the other insects which he has

seen captured by birds. And I venture to think that a closer

inquiry of this kind would reveal the fact that most of the

negative evidence which has been brought up against the Select-

ionist interpretation of mimicry is really of little worth.”

Passing over the obvious reply that as it was the Selectionists

who first asserted that birds ate butterflies, it is their duty to

prove it if they wish their theory accepted, it would appear

that My. Marshall does not consider it necessary for an observer

to be very accurate as to the species captured before attributing

butterfly-catching propensities to certain birds.

In the ease of the Redstart we read that ‘ They take flies, gnats,

small butterflies, and all sorts of small two- and four-winged insects,

partly on the wing and partly at rest”; and again, “ It feeds on

flies, gnats, small butterflies, and various other kinds of small

coleopterous and other insects, caterpillars, etc.” Now if such

evidence is accepted, namely, that the Redstart eats butterflies,

without the necessity of naming the individual species captured

(though it might well be suggested that these small butterflies

were really small moths), it would appear only just that when

such an authority as Legge states that the food of the large Indian

Cuckoo-Shrike consists of “ cater pillars, grasshoppers and various

other kinds of coleopterous insects” without mention of butterflies,

that such should be regarded as sufticient evidence that butterflies

are not destroyed by it in sufficient numbers to cause any form of

mimicry. But, on the other hand, when we read of a bird feeding

‘‘on beetles and the many larger members of the insect kingdom

which affect Ceylon forests,” it 1s quite possible that such a one

would produce a struggle for existence among butterfiies. It is

only by adopting some such standard as this, faulty though it may

be, that we are likely to come to any conclusion. Asto the actual

observation of insects captured by birds, no one who has not

m2, LY.-COL. N. MANDERS ON 'THS

experienced it can form an idea of its difticulty. Let any one who

would appreciate it, watch a blackbird or thrush in his own garden,

and even with the best field-glasses he will be unable in the great

majority of cases to name the species of insect caught, or more fre-

quently than not the order to which it belongs. In the tropics the

difficulties are increased a hundredfold : fortunately in the case of

butterflies the task is lighter, and it is often easy to name the species

owing to the habit some birds have of holding the insect in the

bill for a few moments before swallowing it, and the lengthy

time it takes others either to tear off the wings after the manner

of Shrikes, or beat them off against the ground as is the custom of

the Robin family. Bee-eaters and Paradise Flycatchers nip off the

wings close to the body as neatly as if done by a pair of scissors.

A certain number of birds are migrants, such as the Cuckoo

and Swallow, which pass a portion of the year in Ceylon and the

remainder in northern latitudes; such birds, as pointed out by

Mr. Marshall, would have to learn the distastefulness or otherwise

of the butterflies that inhabit such dissimilar countries, and

their influence on the butterfly population of both areas would

have to be taken into consideration. These are, however, few in

number compared with the partial migrants, which move from

one part of the Oriental region to another; and the butterflies

there being of the same character, the lessons they have learnt in

one part of their distribution are valuable to them in another.

A great many of the resident insectivorous birds move in an

irregular manner from one part of the island to another according

as their food supply varies, and as this consists of insects alone it

is largely dependent on the rainfall. Such a movement cannot

be strictly termed a migration and may be a few miles only ; for

instance, the rainfall in Colombo is about eighty inches, twenty

miles off it is nearly two hundred; when there is a comparative

paucity of insect life at Colombo the birds, or rather some of them,

find plentiful sustenance by moving a few miles inland. Some

birds again, such as the Robins, remain in and about the houses

and gardens year after year, and others, such as the Green Bee-

eater, are never found in the wet portion of the island.

It is this slight to and fro movement which makes me sceptical

of any insectivorous bird in Ceylon, and probably in any tropical

island, being ever really hard up for food.

Mr. Marshall quotes Dr. Franz Doflein as writing: ‘‘ From the

observations which I made in the jungles of Ceylon, it is quite

incomprehensible to me how naturalists who have spent years and

tens of year’s in the tropics can deny the fact” [that butterflies

are frequently attacked by birds]. I had very little doubt when

I read this passage that Dr. Doflein was speaking of the north of

the island, and his recorded observations confirm this; I should,

however, be very surprised if he could say the same of the whole

island, more particularly the hill districts. One point to which I

wish more particularly to draw attention, is his suggested

immunity from attack of the magnificent Ornithoptera darsius,

VALIDITY OF SOME FORMS OF MIMICRY. 713

attention being drawn by him to “its slow, lazy and almost

unwieldy flight,” the characteristics of an unpalatable butterfly.

It does not occur or very rarely in the north of the island, where

a bird, the Paradise Flycatcher, is very abundant, but at Kandy,’

to which place every visitor goes, and where possibly Dr. Doflein

made his observation, the butterfly is very common and the bird

rare, though [ have seen it. But it so happens that at the foot of

the Nilgiris, both bird and butterfly inhabit the same district and

the former is an inveterate enemy of the latter, it eats numbers

of them by nipping off the wings and swallowing the body. The

slow, lazy flight may possibly advertise its unpalatability to some

birds, but it makes it the easier victim to the Paradise Flycatcher,

which in my opinion is the greatest enemy butterflies have in

this part of the world.

I have always experienced considerable difficulty in under-

standing how a distasteful butterfly has acquired a slow sailing

flight *. It is easy to see how swift flight could be brought about

by natural selection, but the converse is not so clear. Presumably

Eupleeas, Danaines, etc., have always been distasteful on account

of the poisonous or nauseous nature of the food-plants, and those of

slower flight, as in the case of the palatable kinds, would naturally

be first captured, and we can understand how a race of quickly-

flying evil-tasting butterflies would be evolved by natural selection.

But if quick flight is of assistance in enabling a tasty butterfly

to escape capture, I find it difficult to believe that a nasty one,

with equal powers of flight, would not be equally benefited. And

if this be so, slow flight for the purpose of advertising unsaleable

goods seems unnecessary and the method by which it has been

evolved very obscure.

When we come to study fast-flying butterflies in their native

haunts, we find in every case a different rate of flight according

to the hour and weather. If it is a cloudy morning they fly much

slower than they do in hot bright sunshine; the majority fly

their fastest after 10 a.m. till 3 p.m., often resting between 12

and 2. But in the early morning and late afternoon, these same

butterflies can often be captured with the greatest ease, as at this

time they are usually feeding. This is the case with the rapidly

flying Teracolus (fausta, danae, &c.) and many Papilios, and this

may account for the different opinion Dr. Doflein and I hold

regarding P. hector and P. polytes. He considers them to be of

swift flight, and so no doubt they are in the middle of the day,

and particularly when flying over bare ground from one patch of

cultivation to another; but on a dull day or early in the morning

or evening they are particularly easy to catch. If Dr. Doflein is

correct we have here an instance of a Miillerian combination of

* Mr. Marshall speaks of “the acquisition of unpalatability,” by which I presume

he means that a slightly nauseous butterfly has become more so by a process of

evolution, and this has been accompanied by an increasingly slower flight. There

is no proof, so far as I know, that a Huplea for instance has undergone any such

process, and the assuinption appears to be entirely theoretical, :

714 LT.-COL. N. MANDERS ON THE

unpalatable butterflies adopting or having a naturally quick

manner of flight which enables them more or less successfully to

avoid the tasting experiments of young birds. I mention this in

order to show the difficulties that beset us even in the field, and

how two observers may form a diametrically opposite opinion on

apparently such a simple matter as the flight of a buttertly.

Dr. Longstaff is no doubt correct in saying that P. polytes has a

quicker flight than P. hector.

This varying velocity of flight makes me somewhat sceptical

that swiftness has been evolved in order to escape capture, as an

enemy by selecting the opportune moment can effect the seizure

of a fast-flying butterfly as easily as one of slow and laborious

flight. ‘Those foes of butterflies, the Bee-eaters, feed during the

hot hours of the day, and Drongos in the morning, late afternoon,

and often after sunset.

Many butterflies are conspicuous under one set of conditions

and the reverse in another, even at the same time of the day.

Telchinia viole is said to belong to the most distasteful group

of butterflies. Its flight is slow and deliberate, and it is very

conspicuous when flying over a grass field; its bright brick-red

colour forms a strong contrast against the green, and it thus has

the characteristics of an inedible insect. In the blazing dazzling

sunshine on the dried-up plains of India its colour so matches the

soil that it is decidedly difficult to see, particularly the female

which is almost invisible. Its under surface matches admirably

the dried-up leaves of the bush on which it frequently takes up

its position for the night, and under these circumstances it has all

the characteristics of an edible insect. On a day in March this

butterfly was flying over the green gardens of Colombo, and three

days afterwards I met it at Trichinopoly, with a shade temperature

of 104°, dazzling sunshine, and scarcely a blade of grass to be seen.

It will probably be held by Selectionists that in certain cases

such as in extremely dry weather, even an inedible butterfly

requires concealment and that conspicuousness is beneficial to it

as advertising the worthlessness of its goods in the wet season,

when insects are abundant. I have given my reasons for

believing that in tropical islands there is no real scarcity of insect

life that cannot be made good by birds. It may also be argued

that its invisibility one moment and conspicuousness the next may

be of advantage to it, but if this be so, it is difficult to understand

why such an extremely distasteful insect as an Acrea is held to

be, should be obliged to pass through such a complicated process

of evolution.

In compiling the following list of birds, I have followed Oates

and Blanford, as their nomenclature is more modern than that of

Legge; where not otherwise indicated, the notes in inverted

commas are quotations from Legge.

Captain Legge spent eight years in Ceylon, and those who were

there with him, now few in number, have a clear recollection of

his knowledge, zeal, and painstaking industry.

VALIDITY OF SOMf FORMS OF MIMICRY. all

Fam. CRATEROPIDA.

Subfam. Crareropin®. (The Babblers.)

“ All feed on the ground like thrushes. They ... probably

derive no portion of their food directly from trees, the fruit they

occasionally eat being picked off the ground as they forage for

insects.” (Oates.)

1. The Southern Indian Babbler. Crateropus striatus.

“Its food is entirely insectivorous, and is mostly taken by

scratching among leaves and débris on the ground.”

2. The Ceylonese Babbler. C. rufescens. Indigenous.

“T found the stomachs of several examples killed in the month

of August to contain portions of a large black beetle which was

affecting the jungle in large numbers at the time.”

3. The Ashy-headed Babbler. C. cineretfrons. Indigenous.

“Delights in exploring the mossy recesses of fallen trunks, in

which humid spots it finds an abundance of caterpillars, bugs,

hemiptera, and coleopterous insects.”

4, The Ceylonese Scimitar Babbler. Pomatorhinus melanurus.

Indigenous.

‘¢Goes about in small companies searching for its insect food

on low branches or clinging w oodpecker fashion to the trunks or

large branches, about ne it jumps and twists itself with con-

siderable agility.” ;

Subfam. TIMELIIN a.

5. The Small White-throated Babbler. Dwmetia albigularis.

‘‘Tts food consists of the larvee of various insects and minute

coleoptera.”

6. The Ceylon Yellow-eyed Babbler. Pyctorhis nasalis.

Indigenous.

‘“‘T have cone found its food to consist of small coleoptera

and various minute insects.”

7. The Brown-capped Babbler. Pellerneum fuscicapillum.

Indigenous.

‘Tt feeds on the ground in dense thickets, picking up beetles

and insects from amongst decaying herbage : it rarely shows

itself in the open.”

8. The Black-fronted Babbler. Rhopocichla nigrifrons.

Indigenous.

“This modest but active bird frequents underwood, anata

and tangled jungle. .... subsisting entirely on various insects

and their larve.”

There is no indication among the Ceylon Babblers of any

716 LT.-COL. N. MANDERS ON THE

butterfly-eating propensity, but Mr. Frank Finn experimented with

an Indian species (Crateropus canorus), and came to the conclusion

that they distinguished in time between a tasty and distasteful

butterfly. The note I made at the time I studied his experiments

is as follows :—It is evident that they had no notion at first as to

what was palatable and what was unpalatable, but as the experi-

ments proceeded they learnt gradually to discriminate ..... I

conclude that as these birds with one exception were adult when

captured, they could not have undertaken tasting experiments

when young, otherwise they would have recognized a distasteful

species.

Subfam. BRACHYPTERYGIN ®.

9, The Indian Blue-Chat. Larvivora brunnea. A migrant.

«“ Appears to feed entirely on the ground.” (Oates.)

10. The Ceylon Arrenga. Arrenga blight. Indigenous.

“The food consists of various insects and in the stomach of my

specimen I detected the bones of a frog.” Mr. Oswin Wickwar

tells me that he found a species of snake, Aspidwra sp., quite four

inches long in the stomach of the bird shot by him.

11. The Ceylon Short-wing. EHlaphrornis palliseri. Indigenous.

“Found in thick brushwood feeding on the ground.” (Oates.)

“ Tt feeds on ants and other minute insects and to some extent

on small seeds.”

12. The Indian White-eye. Zosterops palpebrosa.

13. The Ceylon White-eye. Zosterops ceylonensis.

These species have the same habits as those in Bourbon and

Mauritius.

Subfam. LiorRicHIN&.

14. The Fairy Blue-bird. Jrena puella.

“Tt feeds principally on fruit.” (Oates.)

15. Jerdon’s Chloropsis. Chloropsis jerdont.

“ Seeds may often be found in its stomach, though they are not

so generally partaken of as insects.”

16. The Malabar Chloropsis. C. malabarica.

“‘ Lives on fruit and insects, chiefly the latter.”

17. The Common Tora. “githina tiphia.

‘“‘T have occasionally seen it dart out and seize a passing moth

or butterfly on the wing and alighting again swallow it whole, a

habit which is testified to by the large Mantide and other winged

insects which are often found in its smallstomach.” ‘ Frequents

orchards ..... feeding on insects which it finds among the

leaves.” (Octes.) ; i

VALIDITY OF SOME FORMS OF MIMICRY. Ae

Subfam. BRACHYPODIN2.

18. The Southern Indian Bulbul. Hypsipetes ganeesa.

“Tts diet consists of fruits, seeds and berries....... 1b,

however, also feeds on insects, and I have observed it occasionally

dart at them from its perch, although its usual manner of cap-

turing them is to seize them from the branches of trees, to which

it will sometimes cling after the manner of a Tree-creeper.”

19. The Madras Red-vented Buibul. d/olpastes heemorrhous.

‘Weeds mostly on fruit.” (Oates.)

20. The Yellow-browed Bulbul. Jole ictertca. Indigenous.

“T have found it to be more insectivorous than frugivorous.”

‘Tt wanders about in small flocks, feeding almost entirely on

fruits and seeds.” (Bourdillon.) ‘In all the specimens I have

examined I have found fruit only in its stomach, but from the

strong bristles at the base of the bill I suppose it may, at certain

seasons, partake of insects.” (Jerdon.)

21. The Black-capped Bulbul. Pycnonotus melanicterus.

Indigenous.

‘Tt is chiefly insectivorous, small seeds are sometimes devoured

by it, and I have found snails of some little size in its stomach.”

22. The Yellow-eared Bulbul. Kelaartia penicillata. In-

digenous.

23. The White-browed Bulbul. Pycnonotus luteolus.

‘Tt is both insectivorous and frugivorous, chiefly the latter, and

there is nothing to which it is more partial than the seeds or

berries of the latana plant.”

Bulbuls are very frequently kept as cage birds, and have perhaps

been more often experimented with than any other tropical bird.

Those I kept years ago ate any butterfly given them, and I think

it is now generally acknowledged that those species which are

known to be insectivorous also attack butterflies, but they show

no discrimination in eating them.

Fam. Dicrurip#. Drongos or King Crows.

Ali the Drongos are known to eat butterflies.

24. The Black Drongo. Dicrurus ater.

‘¢ The principal food consists of coleoptera, grasshoppers, winged

termites, of which it is very fond, and ticks, which latter it takes

from cattle. It has been known to devour small birds.”

25. The Indian Ashy Drongo. D. longicaudatus. Migratory,

Proc, Zoou, Soc.—1911]1 No, L. 50

718 LT.-COL. N. MANDERS ON THE

26. The White-vented Drongo. D. leucopygialis. Indigenous.

“Tt is entirely insectivorous, its diet consisting chiefly of

beetles, bugs, termites and such like.”

27. The Ceylon Black Drongo. Dissemuroides lophorhinus.

Indigenous.

“Damp forests and even their most gloomy recesses are fre-

quented by this fine bird...... It feeds on beetles and the many

larger members of the insect kingdom which affect Ceylon

forests.”

28. The Larger Racket-tailed Drongo. Dissemurus

paradiseus.

‘“‘ Feeds chiefly at dusk when the bats come out.”

The distribution of this family in the island is to be noted.

The first species is confined to the Jaffna peninsula and north-

west coast: I found it abundant on the opposite Indian coast in

March. The second is migratory and does not breed in the island.

The third is of general distribution. The fourth is confined to

the heavy forests of the Western Province. The fifth to the

North and Eastern Provinces.

Fam. Syuvirp#. Warblers.

29. The Indian Great Reed-Warbler. <Acrocephalus stentoreus.

“The diet consists of small flies and minute insects.”

30. Blyth’s Reed-Warbler. 4A. dwmetorwm. Migrant.

31. The Rufous Fantail. Cisticola cursitans.

“The diet of this species in Ceylon consists of many sorts of

small insects and caterpillars.” ‘‘The indigestible parts of the

food, which consists of small beetles, caterpillars and little snails,

are thrown up in pellets.” (Jerdon.)

32. Franklin’s Wren-Warbler. Franklinia gracilis.

“The food consists of small insects, which it picks up among the

dead wood to which it is so partial.”

33. The Broad-tailed Grass-Warbler. Schenicola platyura.

“‘ Feeds on the ground.” (Ouates.)

34. The Greenish Willow-Warbler. Acanthopneuste viridanus.

A migrant. ;

35. The Ashy Wren-Warbler. Prinia socialis.

“‘TIts food consists of insects; but occasionally I have found

small seeds in its stomach.”

36. The Jungle Wren-Warbler. Prinia sylvatica.

*Tts food consists of small coleoptera and other minute insects,”’

VALIDITY OF SOME FORMS OF MIMICRY. 719

37. The Southern Wren-Warbler. Prinia jerdon.

‘“‘Tt is purely insectivorous.”

Fam. Laniip&. Shrikes.

Subfam. LANIINA.

“These birds live entirely on insects, the Tree-Shrikes oc-

casionally seizing a small bird or mammal. Some descend to the

ground to seize their prey, a few catch insects entirely on the

wing, and others, again, search branches and leaves for their

food.” ( Oates.)

38. The Rufous-backed Shrike. Lantus erythronotus.

Sew ocally bicdeen ener feeds on grasshoppers, which it entraps

on the ground, and also preys on Mantide and dragonflies.”

It almost certainly eats butterflies. I haveseen it occasionally

in Colombo.

39. The Brown Shrike. Lanius cristatus. A migrant.

Certainly eats butterflies. See experiments on birds in

Colombo (p. 737).

40. The Black-backed Pied Shrike. Hemipus picatus.

“They are rather flycatchers than shrikes in their habits . .

. continually darting out and seizing insects on the wing.”

(Oates.)

41. The Common Wood-Shrike. TZephrodornis pondicerianus.

“Moths and small butterflies form a considerable portion of

its food.”

42. The Orange Minivet. Pericrocotus flammeus.

“ Its diet consists of small butterflies and various winged insects.

In the woods of the Horton Plains I saw it catching insects

in the moss with which the trees are entirely covered in that

damp region.”

Mr. Ormiston tells me that he has known a small flock of these

birds completely clear off some dozens of Papilio polytes larve,

which he was hoping to rear on the fruit trees in his garden.

43. The Black-headed Cuckoo-Shrike. Campophaga sykesi.

‘‘ Its favourite food is caterpillars and other soft insects.”

44. The Large Cuckoo-Shrike. Graucalus macii.

“‘Tts food consists of caterpillars, grasshoppers and various kinds

of coleopterous insects.” Hodgson states its food to be ‘‘ Mantidee

Scarabeei, berries, vetches and seeds.”

45. The Little Minivet. Pericrocotus peregrinus.

“Tt feeds upon various larve (its favourite food) and small

insects.”

50*

720 LT.-COL. N. MANDERS ON THE

Subfam. ARTAMIINA.

“They catch their food entirely on the wing, either by darting

on it from a fixed perch or by flying about after the fashion of

swallows.” (Oates.)

46. The Ashy Wood-Swallow. <Artamus fuscus.

This bird has been frequently quoted as having been seen by

Colonel Yerbury to catch several Huplea core.

Fam. SturNip#. The Mynahs.

“They feed chiefly on the ground on insects and worms, but

they are fond of fruit and berries, which they pick off trees.”

( Oates.)

47. The Black-headed Mynah. Temenuchus pagodarum.

Widely distributed and common.

48. The White-headed Mynah. Sturnornis senea.

A rare and local resident.

49, The Common Mynah. Acridotheres tristis.

See experiments on birds in Colombo (infra, p, 740).

Fam. Muscicapip#. The Flycatchers.

‘The Flycatchers feed on insects which they either catch on the

wing, starting from a perch to which they usually return several

times, or by running with the aid of their wings along the limbs

of trees. They seldom or never descend to the ground.” (Oates.)

50. The Indian Red-breasted Flycatcher. Stphia hyperythra.

Partial migrant.

See experiments on birds at Nuwara Eliya (infra, p. 735).

50 (a). Tickell’s Blue Flycatcher. Cyornis tickelli. Food ?

51. The Blue-throated Flycatcher. Cyornis rubeculoides.

Partial migrant.

52. The Ceylonese Dusky-blue Flycatcher. Stoparola sordida.

Indigenous.

See experiments on birds at Nuwara Eliya (p. 735).

53. The Brown Flycatcher. Alseonaa latirostris, Migrant.

The habits of this bird are well known.

54. Layard’s Flycatcher. <Alseonax muttwi. Partial migrant.

‘‘ In the stomach of one example I found much larger insects

(moderately sized coleoptera) than I expected to find captured by

so small a bird,”

VALIDITY OF SOME FORMS OF MIMICRY. 72)

55. The Grey-headed Flycatcher. Culicicapa ceylonensis.

Jerdon says it feeds on small insects.

56. The Indian Paradise Flycatcher. TZerpsiphone paradist.

Partial migrant.

See correspondence (infra, pp. 728 & 730).

57. The Indian Black-naped Flycatcher. Hypothymis azurea.

Mr. Oswin Wickwar has given me the following interesting

note :—

‘“‘T watched two flycatchers of this species diving into a small

pond evidently in search of some aquatic insect. They both dived

in the most determined manner about five or six times, and,

although I looked carefully, I could not find any insects on the

surface of the water. They did not swoop down and just touch

the surface of the water in the manner of swallows, but deliberately

dived in with a splash like a kingfisher ; there was a momentary

pause, and then they fluttered back to the same perch or one near

about where they started from.”

58. The White-browed Fantail Flycatcher. Rhipidura

albifrontata.

“The chief food consists of mosquitoes and other small dipterous

insects, as also the small Cicadella.” (Jerdon.)

Fam. TURDID&.

Subfam. Saxtcotinz. The Chats.

‘“‘The Chats feed entirely on insects they capture generally on

the ground from a fixed perch, such as the summit of a stone, a

stalk of grass, or a branch of a bush, and then return at once to

their post of observation.” (Oates.)

59. The Southern Pied Bush-Chat. Pratincola atrata.

“The food consists of insects and larve of various kinds, which

they take chiefly on the ground.” This bird is also known as the

Nuwara Eliya Robin; it has very much the habits of the

Stone-Chat.

60. The Black-backed Indian Robin. Thamnobia fulicata.

Has been known to capture Veptis varmona; it usually feeds

just at sunset and as long as there is light.

61. The Magpie Robin. Copsychus saularis.

See experiments on birds in Colombo (p. 737).

62. The Shama. Cittocincla macrura.

“Those shot in Ceylon seemed to be entirely insectivorous,

the food consisting of small beetles, ants, flies, etc.”

722 LY.-COL. N. MANDERS ON THE

Subfam. TurDIN«».

63. The Ceylon Blackbird. Merula kinnisi. Indigenous.

Has almost precisely the habits of the English Blackbird.

64. The Ceylon Thrush. Oreocincla imbricata. Indigenous.

“Decidedly an uncommon bird.... it appears to feed on

insects which it procures beneath fallen leaves.” “‘ Thwaites says

it scratches much in rubbish thrown out at the border of his

plantation.”

65. “The Spotted-wing Thrush.” 0. spiloptera.

““Generally to be found in thick damp jungle picking up pups,

coleoptera and other insects.”

Three species of Swallow occur; one, the Common Swallow

(H. rustica),is migratory, the other two have similar habits.

Fam. MovTAciILuip#. Wagtails.

There are four species of Wagtails, three of which are migrants ;

the fourth, the Large Pied Wagtail, has the same habits as the

rest of the family.

66. The Indian Pipit. Anthus rufulus.

“Weeds on worms and various terrestrial insects and likewise

partakes of small grass seeds.” I have noticed it feeding on green

Aphides, and once saw it capture a Lycznid, a species of Zizera.

Fam. CorAcitAap#. The Rollers.

67. The Indian Roller. Coracias indica.

‘“‘T have on several occasions seen one pursue an insect in the

air for some distance and when the winged termites issue from

their nests after rain, the Roller, like almost every other bird,

catches them on the wing.” (Jerdon.)

“Its food is chiefly large insects, grasshoppers, crickets,

Mantids, and even beetles, occasionally a small mouse or shrew.”

(Jerdon.)

Family MEROPIDS.

68. The Common Indian Bee-eater. Jferops viridis.

69. The Blue-tailed Bee-eater. IL. philippinus.

‘A winter visitant..... it feeds on wasps, bees, dragonflies,

beetles, and even butterflies.” (Oates.)

70. The Chestnut-headed Bee-eater. Jf. swinhoei.

“ Locally distributed throughout Ceylon.” Not found in the

hill country.

These three species are well-known as butterfly-eaters; see

correspondence (p. 727).

VALIDITY OF SOME FORMS OF MIMICRY. 723

Fam. ALCEDINIDS#. Kingfishers.

71. The White-breasted Kingfisher. Halcyon smyrnensis.

“Tt occasionally, but rarely, catches fish by plunging after them,

it lives chiefly on insects and small lizards and sometimes on mice

and land crabs.” (Qates.) ‘‘ It subsists on lizards, grasshoppers,

locusts, and even small snakes.”

Fam. CypseLip#. Swifts.

72. The Alpine Swift. Cypselus melba.

73. The Common Indian Swift. C. affinus.

These birds have all the habits of the common European Swift. _

74. The Palm Swift. Zachornis batassiensis.

‘“‘ Feeds chiefly at dusk.”

75. The Brown-necked Spine-tail. Chetura indica.

“Mr. Carter says that those he shot had fed on beetles, green

bugs, sand-wasps and grasshoppers.” Mr. Spurway informs me

that he has more than once seen it snap up a butterfly. Legge

says it is fond of termites.

76. The Indian Crested Swift. J/acropteryx coronata.

I know nothing about the feeding habits of this bird.

Fam. Trogonip#&. The Trogons.

77. The Malabar Trogon. Harpactus fasciatus.

Mr. Butler has seen a large moth brought to the nest he was

observing.

“ Tt feeds chiefly on beetles, moths or cicades ; but it occasionally

feeds on insects on the ground.” (Blanford.)

Fam. Cucuuip#. The Cuckoos.

‘They feed chiefly on caterpillars and soft insects.” (Blanford.)

78. The Cuckoo. CO. canorus Migrant.

79, The Small Cuckoo. (C. poliocephalus. Migrant.

80. The Indian Cuckoo. C. mucropterus.

‘¢ Feeds on caterpillars.”

81. The Indian Plaintive Cuckoo. Cacomantis passerinus. A

migrant.

«“ Tt feeds on caterpillars, coleoptera and other large insects, and

may often be seen taking them on the ground.”

M24: LT.-COL. N. MANDERS ON THE

82. The Banded Bay Cuckoo. Penthoceryx sonneratis.

‘Feeds on coleoptera, Mantide, and caterpillars.”

83. The Drongo Cuckoo. Surniculus lugubris.

‘Locally dispersed .... the diet is mixed, consisting chiefly

of caterpillars and beetles but often combined with various seeds.”

84. The Pied Crested Cuckoo. Coccystes jacobinus. Migrant.

85. The Red-winged Crested Cuckoo. C. coromandus. Migrant.

‘‘The stomachs of those I have procured contained beetles,

grasshoppers, Mantide, and other large insects.”

Fam. FALCoNnIDs#. The Falcons.

86. The Kestrel. Zinawneulus alaudarius.

Has the same habits as the English bird, which has re-

cently been shown in some instances to have butterfly-eating.

propensities.

The above birds may be tabulated as follows; the arrangement

is artificial, and no doubt there are errors, but it is convenient.

Some may be inclined to transfer the Flycatchers from group 3

to group | on the evidence brought forward by Mr. Marshall in

the case of the European Flycatcher. I have put them in the

third group, as close observation for two months showed me

that two species at any rate only eat butterflies under unusual

circumstances.

May take Butterflies, |

ST ihre ta ee gee e afer atare |

| The Common Tora 1 | Chats and Robins... 4] Flycatchers ......... 8

| Dron Ose. ye eee eee | Swallows ............ 3) Sbabblersipaes sees

SIRAUEE” eceaneehbcouns eGapncencccuts Gy WES ee sch encodes ZI) WAVMIDERS. “lecodheoc: 9 |

The Orange Minivet ............ 1 | SURORQOIN soocccceaccdues Jb || ISGUNIS conpsadéeseonovace 2

The Ashy Wood-Swallow ...... 1 | Cuckoos oueeeeesee., 8} The Little Minivet 1

The Paradise Flycatcher _...... il DAMM socscune 9052 6 | ChloropSis ............ 2)

Way) OW Pecans ndoastcasteopcanccsca:’ L The Kestrel ......... 1

Beezeaters.... canoes oe eee: | Miynahs” jo. ase a

The White-breasted Kingfisher 1 | The Indian Pipit ... 1

The Brown-necked Spine-tail ... 1 |

Toran... 21 | Toran... 31 ToTat... 30

4 of the above are either mi- 9 of the above are 6 of the above are |

grants (2) or partial migrants (2). | either migrants (4) or | either migrants (4) or

partial migrants (5). partial migrants (2).

I would suggest this as a convenient place for a perusal of

VALIDITY OF SOME FORMS OF MIMICRY. 725

Mr. Marshall’s paper above quoted, more particularly that part

devoted to the Indo-Malayan region.

For the sake of convenience I shall designate the birds in

Group 1, “The Butterfly-Eaters,” and the evidence in support

of this term is given below under each species.

1. The Common Tora and Butterflies.

This bird belongs to the Bulbul family, another name for it

being “ The Common Bush Bulbul.”

I have frequently watched this bird and can only confirm

Legge’s observations. Being so closely allied to the Bulbuls it

is probable that, like its near relations, it is an indiscriminate

feeder.

2. King Crows (Drongos) and Butterflies.

“On the 14th instant I was with Mr. C. B. Antram in a very

small patch of wood surrounded by grass downs within a few

miles of Ootacamund. Running through this wood was a foot-

path, and this path was in one place a few feet long thickly

strewn with the wings of butterflies; on either side of this, for

some yards along the path, were scattered wings. Just above

this place three Drongos (Dicrurws) were to be seen on the trees.

The weather during my visit was misty, cold and rainy, and hardly

any butterflies were on the wing ; consequently I had not the

opportunity of seeing the birds hawking them. Dragonflies were

abundant and the Drongos made frequent excursions after them,

all unsuccessful so far as I saw. On examination, the wings were

found to belong to the following species:—Danais limmiace (or

septentrionis, or both), Danais aglea, Danais chrysippus, Huplea

core, Huplea coreta, Hypolimnas bolina 2 , and Catopsilia crocale.

At least 90 per cent. of the wings belonged to D. limniace or

septentrionis. There were several wings of Huplea, amongst

which I found those of the males of both core and coreta. The

remaining species were each represented by only one or two

wings. These Danaine are common in those parts, and in

sunny weather would be passing in numbers through and over the

wood, and the most common would almost certainly be limniace,

septentrionis, core and coreta. D. chrysippus would be commoner

probably on the outskirts of the wood. The only other butterflies

about in any numbers when I was there, were Argynnis castetsi

and Colas nilgiriensis, both eminently insects frequenting the

open country, and they would seldom or never be found in any-

thing heavier than scrub jungle. Some of the wings were

obviously quite fresh, others had been beaten on the ground by rain.

The three Drongos were almost the only birds about, and no

other birds that we saw were likely to be capable of capturing

these butterflies. It appears to me that the Drongos were

certainly the cause of this extensive shower of wings; the only

other possibilities are lizards or mantids, but no lizard, I should

726 LY. COL. N. MANDERS ON THE

imagine, has either the activity or voracity to make away with

sucha great number of butterflies, even assuming that they settled

within its reach. A mantid is even less probable, and I much

doubt if there is one large enough to tackle Danainz or Hypo-

Limnas in the Ootacamund region.

“One would not be surprised to find an occasional Danas or

Euplwa sampled by a bird, but to see evidence of a systematic

onslaught on butterflies which are so universally looked upon as

leaders of the army of distasteful insects, and which are so widely

mimicked by numerous ‘unprotected’ butterflies and moths,

tends to make one sceptical of the accepted theories founded on

the alleged value of this distastefulness. It is true that they may

disagree with other birds, lizards, &c., but if one enemy alone can

effect such wholesale destruction upon them, their immunity from

death by violence is so seriously impaired that it seems to me that

their numerous imitators amongst the ‘ Swallowtails,’ &c., are

simply asking for trouble.

“There was no lack of other food for the Drongos, and it can

only be assumed that they found the Danaine very much to

their taste. One can hardly think of a morsel more apparently

unpalatable than Telchinia viole, yet I saw a Bulbul feed its

young with one within a few feet of me. It would be interesting

if entomologists would give any evidence in the matter which

they can. I have seen the wings of Hypolimnas misippus some-

times scattered on the road near trees in considerable numbers,

and on two occasions I have seen the wings of Charaxes imna ;

this I imagine was not caught on the wing; if so, I must con-

gratulate the bird on its power of flight.

H. Lestre ANDREWES.” —

Barwood Estate, Nilgiris, 20th October, 1910.

(Journal Bomb. Nat. Hist. Soc., vol. xx. p. 850.)

This interesting observation may be held to support Mr.

Moulton’s proposition (Trans. Ent. Soc. Lond. 1908) that the

Kupleas of S. India are a Miillerian combination formed for

mutual protection against the onslaught of insectivorous birds ;

but evidence is required that at one time they differed materially

from their present day appearance.

I took the few notes following at Coonoor, 6000 feet, Nilgiris,

S. India, in April 1910. I asked my collector, a half-caste who

had spent all his life in the jungles, whether he had ever seen a

bird catch a butterfly, and he immediately said he had, describing

the Paradise Flycatcher very accurately. He said they eat the

brown butterflies (1. core and coretw) and white ones. While he

was speaking I happened to open an envelope containing Danais

septentrionis, and he exclaimed, ‘‘ I have seen them eat those too.”

He added the bird nips off the wings and swallows the body ;

also that they catch and eat the Ornithoptera. A few days after-

wards I was at Kullar at the foot of the hills, about thirty miles

from Ootacamund, where these birds and Drongos are common.

I saw a Drongo in a lane, and close together on the ground

VALIDITY OF SOME FORMS OF MIMICRY. (OAT

I found the wings of Papilio hector, Huplea core, and Danais

septentrionis, and a little distance further on another D. septen-

trionis. ‘These were the commonest butterflies about at the time.

12.4.10. Kullar, 1200 feet. Saw a Bulbul dart out at a

Papilio hector and miss it. Saw the following with notched wings :

P. demoleus, one or two Catopsilias, one or two Teracolus fausta,

several P. hector, many Junonia lemonias, these last usually with

oval pieces out of the secondaries, probably by Calotes.

10.4.10. Coonoor, Walking along a road I saw what I thought,

at first, was a leaf falling from a tree about twenty feet up, but on

going to look at it I found it was the fore wing of H. masippus 9,

diocippus form. It is curious that I have only seen one other

female during the many times I have been along this road. I could

not see the bird, but Bulbuls are plentiful. Found also on the

road the fore wing of Papilio sarpedon.

21.4.10. Kullar. Watched a Racquet-tailed Drongo for some

time hawking after flies. I did not see it chase a butterfly

though there were numbers ftying about, chiefly Euploeas.

Mr. T. N. Hearsy, Indian Forest Service, writes :—‘‘ Coimbatore,

GiOalONs. ks ee I have frequently seen the common green Bee-eater

(Merops viridis)and the King Crow (Buchanga atra) take butterflies

on the wing, the butterflies being Catopsilia pyranthe, C. florella,

Terias hecabe, and Papilio demoleus. The Bee-eater I have also

seen taking Danais chrysippus and Danais septentrionis, and I

remember to have been struck with their taste for those latter. .

I have also seen the Tree-Swift (Dendrochelidon coronata) take

Catopsilia pyranthe.” -

In another letter he mentions having seen Drongos attacking

dragonflies.

Mr. Ormiston, of Kalupahani, Hadumulle, Ceylon, 4500 feet,

writes :—“Kalupahani, 4.2.09. Of course I will try and get you any

notes I can on birds eating butterflies ....I can give you very

little assistance at present, as the only bird I have watched is the

Fork-tailed Drongo, who eats the whites during a flight and attacks

Kallima.

‘“‘ The Magpie Robin and black-and-white Flycatcher catch a great

lot of moths, but I have no notes of their taking butterflies. At

Ohiya bungalow, after a moth night, we used to bottle all the moths

we wanted, and then loose a tame Mynah who made short work

of the rest, but I never tried him with butterflies. I will try the

Robins in my garden, but the fact that they eat dead butterflies

will not prove that they catch them. Personally I do not think

birds make any appreciable difference to the number of butterflies

except possibly by eating them in the larval stage. It is quite

unusual to see a butterfly caught.”

“ Kalupahani, 3.1.10. I have kept my eyes open for butterfly-

eating birds, but I am sure that the slaughter, if any, must take

place in the larval stage. I have seen the Fork-tail Drongo

feeding on the flight whites and Kallimas, but that is all.

““T have seen a dragonfly catch and kill Zesius chrysomallus

and another fly killing Papilios, Lycenide, Syntomiide, with

728 LT.-COL. N. MANDERS ON THE

apparently preference for the last named. Some black-and-white

Flycatchers come close when I am beating for ‘plumes’ on the

road here, and catch lots of common Pyralids I put up. The

common Magpie Robin comes most mornings for the moths which

I have slain at the lamp and discarded, and I have seen a Sparrow

catch a gold-tail moth. But since you asked me a year ago

to look out, I have not seen half a dozen cases of birds eating

butterflies.”

“ Kalupahani, 26.6.10. I have kept my eyes open for birds eating

butterflies but have no new notes. During the last flight the

Fork-tailed Drongos were as usual feeding on the white butterflies,

but that is the only instance I have seen. Certainly few, if any,

birds eat the Terias family. I have watched them flying slowly

with Swallows, Drongos, and Flycatchers close to them and leaving

them alone. I fancy Bee-eaters take butterflies, but they are

searce here, and I have no data therefore to goon......

“ Kallima philarchus appears at Hadumulle in large numbers

and the flight lasts for about a week. The biggest flights are

’ very nearly always from about Christmas to the New Year, but a

smaller flight appears in July. About once in four years they are

especially common . . . They seem especially to favour Loquat trees,

but come readily to a mixture of jaggery (native sugar), beer and

rum. Instead of settling on the branches or trunk with their

heads towards the top, they seem usually to do the opposite, and

are therefore apparently very conspicuous; but the birds do not

seem to notice them till they move ren they come to sugar

they settle anyhow).

“The spot where I mainly catch them is about two acres of

Grevilleas planted with a few Loquats. When the flight is on,

the Grevilleas seem full of Forked-tailed Drongos, and as soon as a

butterfly moves a Drongo darts for him, but usually only takes a

big piece out of one wing. I have never seen the wings lying on

the ground, so fancy if the Drongo gets him he eats wings and all.

I have never, however, seen him catch one.”

“10.10.09. Paducka. Watched Drongos (Dicrurws leucopygialis)

hawking for flies ; though Mycalesis ceylonica and smali Lycznids

were flying plentifully beneath the birds, they did not take them.

“19.12.09. Paducka. Watched several Drongos and a Paradise

Flycatcher ; the former frequently caught small flies in the air and

occasionally came to the ground after bigger insects, but only once

did one catch a lepidopterous insect and this seemed to mea moth.

The Flycatcher took short flights on the ground picking up flies,

but certainly not a butterfly. Cameacross five fully fledged Ashy-

headed Babblers sitting all together on a branch ; they flew off only

when I approached quite close to them, with great chattering, very

much like the ordinary Babblers ; the old birds were hunting for

food in the thick bamboo jungle. This is very late in the year

for young birds.”

I give these merely as samples of negative notes; there is no

object in giving more.

VALIDITY OF SOME FORMS OF MIMICRY. 729

Mr. KE. Ernest Green writes :—

“* Peradeniya, 16th July, 1910.

“With regard to the capture of butterflies by birds, I was told

(in May last) by a lady who was staying with us, that she had

been watching the Drongos in these Gardens busily catching

butterflies. From her description, the victims seemed to be

species of Huplwa. She said that the birds bit off the wings, and

that the road was covered with the dismembered wings. I asked

her to collect some of the wings for identification, But, in the

meantime, they had either been swept or blown away, and

she could only produce one or two wings of Papilio jason and

Jamides bochus.”

The common brown Eupleea can hardly be mistaken for any

other Ceylon butterfly.

3. Shrikes and Butterflies.

Dr. Willey, F.R.S., writes:—‘‘ The late Grant Allen stated

positively that among the animals which he had seen in Butcher-

bird’s larders were mice, shrews, lizards, robins, tomtits, and

sparrows ; but he added that in spite of its occasional carnivorous

tastes, the Shrike is at heart an insect-eater.”

The few experiments I have been able to make leave little

doubt in my mind that they make little or no selection in their

butterfly diet. See below (p. 737).

4. The Orange Minivet and Butterflies.

I know nothing about this bird’s provender other than already

given ; Mr. Ormiston’s observation is, however, suggestive. The

larva of Papilio polytes lives in its earlier stages exposed on the

upper surface of the leaf of the orange or citron, looking exactly

like a bird’s dropping; when irritated it shoots forward two fleshy

“horns” emitting a pungent smell of orange, which is highly

disagreeable. I presume this is derived from the food-plant, and

if this be so the larva in all probability has a taste of unripe orange,

and consequently it would seem that the bird’s palate is not highly

educated and its taste in butterflies probably not selective.

5. The Ashy Wood-Swallow and Butterflies.

Mr. Walter A. Cave writes :—

“Colombo, 21st October, 1910.

‘“‘T am sorry I cannot help you much in regard to the question

of butterflies being taken by birds. On one occasion I observed

an Ashy Wood-Swallow (Artamus fuscus) tearing the wings off a

butterfly, then swallowing the body. This was in Peradeniya

Gardens a year or so ago. ‘There were many of these birds, which

are allied to the Shrikes, hawking over the Maha Weliganga river.

f did not see this particular bird actually catch the butterfly, but

I have every reason to believe it did, because I had a good pair

730 LT.-COL. N. MANDERS ON THE

of prism binoculars focussed on the bird as it alighted in a tree.

As I have said, the wings were first stripped, and as they fell I

was able to make them out. I am not well acquainted with

butterflies, but to make my notes complete I referred to the

Museum and determined the butterfly as the commonest brown

one we see about the roads here, and I put it down as Huplea

core,

“On another occasion I saw a Common Swallow (Hirundo

rustica) take a small white butterfly, but I was too far away, and

without my glasses, to determine the species. The Swallow was

hawking near a piece of very dark jungle, near the Nuwara

Eliya lake, and the white butterfly was very conspicuous against

the background.”

Colonel Yerbury’s note of this bird being seemingly partial to

Euplea has already been alluded to.

6. The Paradise Flycatchers and Butterflies.

Mr. John Pole, Scarborough, Maskeliya, 6000 feet, writes :

“ Maskeliya, 13.3.09. . . | seem to recall the attacks of the Odonata

and even Asylus (Diptera) on butterflies, but I never remember an

instance of a bird attacking one—I have watched the little Tailor-

birds eating the larve of Veritas hecube and that with seeming

distaste, and the Flyeatchers at work on Diptera from the shelter

of some leafy tree; I have had so large an insect as Phyllades

consolisma taken from under my very nose by a Drongo, have

had moths beaten from a fence in the daytime stolen by Swallows

ere I could net them, but have never seen a bird in Ceylon carry

off a butterfly. Im England I have seen a Swallow carry off

Papilio machaon whilst 1 was following it...I came out to this

Island in 1871.”

‘“‘Maskeliya, 17.3.09. Since writing on 13th we have had for two

days (16th and 17th) flights of butterflies, the first I have seen

this year. There has beenin my garden for the last three months

a bird, which generally goes by the name of the ‘Cotton Thief’

(the Paradise Flycatcher). This bird occupies a jak tree within

twenty feet of my window, and for the last two days he has been

obtaining all his meals from the flights of butterflies, and although

I have never actually seen him catch one, I have seen him

circling from the tree in pursuit and the wings of the insects he

captures falling around the base of the tree within a radius of

twenty feet. Should they be of any service to you, I can send

you the wings of Appias paulina and varieties ¢ and @? as

follows :—

Upper left wing, 4 g 2 @.

Upper right wing, 5 d 59.

Lower left wing, 5 Q.

Lower right wing, 2 3.

‘“« Maskeliya, 14.4... . I have only one more species of butterfly

to give you as its food— Papilio agamemnon, and this is the only

VALIDITY OF SOME FORMS OF MIMICRY, 731

butterfly to my best belief the bird has taken since my previous

letter on the subject. The bird is rare here ....We have had

no flights of the yellow and white Catopsilias this year—so no

wings ene the Appias victims were mostly the white forms 3g

and 92.”

7. The Roller and Butterflies.

Tam unable to add much regarding this bird’s dietary. It is

found only in the dry northern districts and is uncommon. In the

plains of India it is a familiar occupant of the telegraph-wire, and

I have often seen it chasing and no doubt catching various insects

from such a perch. I ‘have no doubt it catches butterflies, and

more than once am sure I have seen it do so, though I am unable

to name the species captured. Judging by its very varied menu,

I have little doubt that it pays little heed to the species it

manages to catch, but of this I have no direct evidence.

8. Lee-eaters and Butterflies,

Mr. Fred. Lewis, a well-known ornithologist, writes:—-‘‘ Colombo,

Sula ODN Ausra. 1 have noticed Swinhoe’s Bee-eater in particular

hawking after the common so-called ‘ Adain’s Peak butterfly ’

[ Catopsilia and App-as|, and it appears to prefer the white one to

the larger yellow fly. It does not, so far as I am aware, take any

of the large brown butterflies often to be found with the above

named, I have watched the bird when quite a selection of flies

could be made, but beyond taking the white and an occasional

yellow, I have never seen it feed on others. I am not prepared to

say, however, that the Bee-eater does not eat any other butterflies

than the two mentioned.

‘“¢ Our common black King-Crow appears to select the same flies,

taking them on the wing in the same way as the Bee-eater.

Swallows do not, so far as my observations go, ever touch any

butterflies.

“Tt isremarkable, however, on such occasions as one finds in the

dry zone, when vast masses of these ‘Peak’ whites and yellows

congregate round some wet pool or damp ground, that King-Crows

are not found taking the opportunity of a‘ square feed.’ I infer

therefrom that the butterfly is only a ‘side dish’ and not a

regular item of food. . . 1 cannot recall an instance of young birds

being fed with butterflies. I suspect the difficulty of swallowing

the wings is the reason, for I cannot say I have ever found small

nestlings with anything so difficult of mastication in their

mest,”

In another letter Mr. Cave writes :—

“ Belvedere, Colombo,

6th March, 1911.

“In reply to your letter I am sorry to say I have nothing

further to report on the subject of the capturing of butterflies by

732 LT.-COL. N. MANDERS ON THE

birds. I have the subject always in mind when I happen to get

out, and my friend Mr. Symons, of the Government Training

College, is also on the look-out, andif we should notice anything

it will be reported to you.

‘“‘ Being keen on birds there is very little that escapes my

notice when I happen to be out, and I must say the subject on

which you write is very exceptional so far as my observations go.

At Christmas time I motored round the south coast to Galle,

Hambantota and from thence to Haputale. The butterflies were

there in myriads, nearly all the way—none of us had ever seen

anything so remarkable in our lives. There were literally clouds

of butterflies—in fact we remarked that we now knew where all

the butterflies came from which used to appear here on migration

in the N.E. monsoon. But neither Mr. Symons nor I saw a

bird attempt to catch a butterfly, and we saw a large variety of

birds including Bee-eaters, Swallows, and Swifts.”

An argument of some force against the frequency of butterfly

victims may be advanced by the different behaviour of birds in

the presence of a flight of locusts and a migratory flight of

butterflies. I have been fortunate enough to witness both, and

the difference is very striking. In the former, every kite, crow,

and insectivorous bird in the district follows the locusts, gorging

to repletion; and it is a very remarkable sight the numbers, I

might almost say flocks, of birds following the swarm. In a

migratory flight of butterflies, on the contrary, with the exception

of a few Bee-eaters and Drongos, birds are conspicuous by their

absence.

Mr. Oswin Wickwar, F.E.S., sends me the following note :—

‘“When shooting in the Northern Province in May last, I was

crossing the dry bed of a river when a Bee-eater (Merops viridis)

swooped down and caught a butterfly on the wing within a

couple of yards of my face. The instant it snapped it up, the fore

and hind wings of the butterfly floated down in front of me, and

enabled me to prove its identity, Papilio polytes 3. This was

about 11 a.m. The following morning about 7 a.m. I had occa-

sion to cross the same spot, and saw what was probably the same

bird perched on a twig; I had hardly seen it, when it flew down

and entered a hole in the bank, but came out again in a few

seconds. This was evidently its nest, and I was anxious to look

for remains of insects, nor was I disappointed. By introducing a

twig, the hole seemed to be about two feet deep and to travel

in a horizontal direction, so a start was made to cut away the

bank, and before going a foot three insects were unearthed,—a

* humming-bird moth (MJacroglossa sp.?) which was still alive, a

‘skipper’ (Swastus grenvius) also alive, and a small green beetle

with white spots on the elytra. All these were secured and taken

away. The hole now took a turn and went a downward course

for about two feet, which meant cutting away a huge piece of the

bank to get at the nest, so I had to leave it,”

VALIDITY OF SOME FORMS OF MIMICRY. Gas

The following extract is of much interest ; it is from the ‘ Spolia

Zeylanica,’ 1910 :—

<< Bee-eaters as Fish-eaters.

“There are a pair of Chestnut-headed Bee-eaters (JJerops

swinhoet), which nest pretty regularly in a steep bank on a road

below my bungalow, and about 150 yards distant from my pond.

Almost any bright afternoon, between 2 and 3 p.m., they may be

seen fishing in the pond. They come down from a dead tree,

which stands on a knoll some 50 yards away ; sometimes hovering

for a moment over the water to locate their prey, but more

commonly marking it in their swoop, and dashing headlong into

the water like a kingfisher and very rarely missing their fish. IL

have seen the pair account for a dozen fish in as many minutes;

all quite small fry.

“When there is a flight of white butterflies on, these birds

devote most of their attention to them throughout the day, but

on warm bright days nearly always have a go at fishing in the

afternoon.

“‘T have always hitherto associated Bee-eaters with the one diet

of insects ; and I could not quite trust the accuracy of my eyesight

until I brought a strong pair of field-glasses to bear on the actors

at the short range of 15 to 20 yards. I think it probable that

many so-called insectivorous birds change their diet when some

chance has put them up to the taste of a new article which

happens to suit them. . .

‘¢ Since the pair, which I convicted of fishing, hatched out their

young, they have abandoned their fishing expeditions and may be

seen sitting on the tree facing their burrows catching insects

(chiefly white butterflies) to feed their nestlings.

E. Gorpon ReEgvss.”

Wiltshire, Matale, May 7, 1910.

9. The White-breasted Kingfisher and Butterflies.

1 have often seen this bird eating grasshoppers, and on one

occasion a butterfly which I was too far off to identify. Small

frogs and lizards, some of the latter of quite a respectable size, I

have also seen taken. Dr. Willey writes, ‘I remember being

much surprised, many years ago, to find a Kingfisher’s stomach

full of insect remains.” With such a mixed dietary its taste

for butterflies is probably impartial.

Sparrows and Butterflies.

Mr. R. D. Hodgins writes :—‘“ April 1911. These birds have

built their nests about my bungalow here at Matale, so I have

plenty of opportunity of watching them. JI have on three

Proc. Zoou. Soc.—1911, No. LI. 51

734 LT.-COL. N. MANDERS ON THE

occasions noticed the birds hawk and catch butterflies in mid-air,

and in two cases the butterflies caught were taken off the tree

tops, but I could find no trace of them on the ground.

‘“‘The flies on one of these occasions were brown and on the

other white, very like the common white cabbage butterfly of

England, but I was some distance away and couldn’t see them

properly [probably Huplea, and Catopsilia or Appias].

““On the third time the butterfly was caught while passing the

bungalow one sunny afternoon in February, but appeared to be

released the moment later and dropped to the ground. Only its

abdomen was taken and this was nipped off neatly at the waist.

These wings I collected and herewith enclose | Papilio sarpedon :

the butterfly was otherwise in remarkably good condition ; it is

a very rapid flier. |

‘“‘T have often seen a sparrow dive and catch a feather floating

in the air in a similar way to that in which a swallow does.

Whether the feather is mistaken for a butterfly or vice versa I

don’t know.

“In the case of Ceylon native birds, I have been unable to

detect any catches of butterflies, but have noticed that two of the

species will take and devour moths...... This bird [the

White-bellied Drongo| I have often seen catching flies from the

branch of a tree or telegraph-wire with downward swoop on to

the victim. I have often seen it catch small moths up to about

one inch in length, which it seemed to devour, wings and all, after

returning to its perch.”

It would be as well to refer to the distribution of the “‘ Butter-

fly-eaters.” The Drongos have already been alluded to; only

one is of general distribution. The Common Iora is widely

distributed. The Shrikes with two exceptions are found in the

wooded country of the upper and lower hill districts, the other

two I have seen only in the low country.

The Ashy Swallow-Shrike is a partial migrant in the island ; it

appears in small flocks in Colombo during the north-east monsoon,

at other times it is more an inhabitant of the north. The Roller

is found only in the dry districts, so also are the Bee-eaters,

particularly the Green Bee-eater, which is never found in the wet

country or above 300 feet. Swinhoe’s Bee-eater occasionally —

ascends higher, and the migrant Philippine Bee-eater appears for

a short time on the wet western coast at the break of the north-

east monsoon, but soon retires to the dry northern districts. The

White-breasted Kingfisher and the Brown-necked Spine-tail are

of general occurrence.

Tf this distribution is carefully studied, it will be noticed that

there is a distinct paucity of butterfly-eating birds in the wet

hill districts and that part of the coast subject to heavy rains ;

though mimicry occurs quite as commonly, if not more so, as

in the drier districts, where butterflies are less commonly met

with.

VALIDITY OF SOME FORMS OF MIMICRY. 735

Experiments on Wild Birds.

The following experiments were made on one bird in column 1,

two birds in column 2, and two in column 3 (see p. 724), a very

small percentage of the total number no doubt, but the best I

could do under the circumstances, and they indicate the lines

for future investigations.

Haperiments on Flycatchers at Nuwara Eliya, 1909.

The Indian Red-breasted Robin Flycatcher, Siphia hyperythra.

This bird is migratory and is found only in the hill districts ;

it is about the size of the European Spotted Flycatcher.

30.3.09. Deprived live Zerias hecabe one, Terias libythea one,

Hypolimnas bolina 3 one, Danais fumata one, Neptis leucothoé

one, and Appias galene one, of about two-thirds or three-quarters

of their wings, and put them on the ground near a tree from

which one of these birds was accustomed to feed. It first made off

with either the Veptis or Danais, I could not see which, then the

Appias. It then flew away, and J picked up one of the Terias

and Hypolimnas. This last had a good deal, perhaps half, the

wings left and fluttered about vigorously.

1.4.09. Lethe daretis one, Vanessa haronica one, Terias hecabe

one, and the same H. bolina, treated in the same manner as on

the last occasion and put in the same place. The bird carried off

the first two, but the H. Lolina seemed too large for it, as it was

for another Flycatcher which came along shortly after the first

had flown off. I could not see what became of the 7’. hecabe.

The same afternoon a similar experiment, but with no result.

9.4.09. This bird has evidently nugrated as I have seen none

since the last note. It nests in the Himalayas.

The Ceylonese Dusky-blue Flycatcher, Stoparola sordida.

This bird is peculiar to Ceylon but confined to the hill-tracts.

2.4.09. Placed Terias hecabe two dead, Argynnis hyperbius one,

Danais fumata three, Appias galene one, these latter alive but

largely deprived of their wings, near the cherry-tree much fre-

quented by Flyeatchers. The cock bird carried off one D. fumata,

its mate another ; the latter I was enabled to watch closely, and

it had extreme dithculty in swallowing the fly on account of the

wings. The former shortly afterwards returned and carried off

the remaining D. fumata, but the latter seemingly had had

enough of it. Shortly after a Red-breasted Flycatcher flew off

with the Appias. What became of A. hyperbius I do not know,

but the two dead Zerias were untouched.

Numerous specimens of the above butterflies were flying

about at the time, but I have never seen these Flycatchers molest

them.

5.4.09. Placed 7’. hecabe one, D. fumata one, and A. hyperbius

51*

736 LT.-COL, N. MANDERS ON THE

one, under the cherry-tree, having amputated both wings on

one side close to the body. The cock bird flew down almost

immediately and seized D. fuwmata, and as usual it had great

difficulty with the wings; about ten minutes afterwards it took

the A.hyperbius, but 7’. hecabe was left. I fancy the wings rather

‘‘ put off” the bird.

29.4.09. The birds have a nest with young in a cherry-tree

in the grounds. I put down 4. hyperbius 9, 7. hecabe, Neptis

lewcothoé, and one or two others; they had all been dead some

days and were very dry. After some time the bird noticed them,

flew down and seized the Argymnis, which had its wings closed

and showing the underside. It flew with it into a tree but very

shortly dropped it, it was evidently too dry. I found it had been

caught by the fore wings, one of which was gone with also a

portion of one hind wing. ‘The same afternoon put down live

Pyrameis cardui, N. lewcothoé, 7. hecabe, Appias nadina, and

Euplea core, but with both wings on oneside removed. The cock

bird flew down among them and caused a great flutter; 1t first

caught the Zerias, then the Neptis, and lastly the Huplea, which

provided a great chase. It carried them altogether to the nest,

but in feeding the young, the Huplwa escaped ; the bird was after

it in a flash, caught it again and carried it back to the nest. It

was very interesting to watch its efforts to get sucha large insect,

the size of our Camberwell Beauty, into the young one’s mouth.

Three or four times it had to take it out and manipulate it in its

own beak before another trial; eventually, it succeeded in forcing

it down the youngster’s gullet.

2.5.09. A half-winged live 4. hyperbius placed near the nest.

I am sure the bird noticed it, but beyond regarding it carefully

it did not molest it.

I am convinced from long and repeated observation that the

old birds never fed on butterflies themselves or fed their young

with them. <A critic, whose opinion I value highly, has objected

that because I never saw one of these birds capture a butterfly,

it is no proof that they did not do so and that very possibly

the difficulty of catching them would only induce pursuit

when the butterfly was off its guard and a capture possible.

I do not know why the birds should be more coy of capturing

a butterfly than a house-fly in my presence, and I can scarcely

believe they took the opportunity of my absence to do so.

Granted that difficulty of capture was the reason for non-

pursuit, what chance, it may be asked, would a young bird with

considerably feebler power of flight have of conducting a series

of tasting experiments on these butterflies? It is not infrequent

in the writings of advocates of mimicry to explain the rareness

of attack by difficulty of capture ; but by doing so they seemingly

forget that if such is the case with old birds, it makes tasting

experiments (with butterflies) very difficult for young ones.

VALIDITY OF SOME FORMS OF MIMICRY. 737

Haperiments on Birds in Colombo.

The Brown Shrike. anius cristatus.

28.12.08. Pinned a large Hypolimnas bolina 2 (a mimic of

HKupleeas) on a paling: in a few minutes the bird came along, and

directly it saw the butterfly it pounced upon it and carried it off,

and I could not see where it went, but I have no doubt it ate it.

29.12.08. Pinned a Hypolimnas misippus 2 on the paling.

The same Shrike saw it, seized it and held it in its claw, eating it

piecemeal and tearing off two or three wings. The following day

the same experiment was repeated with Danais chrysippus and

Telchinia viole, with the same result.

6.4.09. Nuwara Eliya. Noticed one of these birds, perched on a

twig, fly down and capture some insect on the ground. I watched

it for a long time, but though many butterflies flew past, it took no

notice of them. They comprised principally Argynnis hyperbius,

many Appias sp. ¢ Huplea core, Terias hecabe, and Lethe daretis.

28.11.09. Colombo. Put down Delias eucharis 2 , several Huplea

core, Danais limniace 2, Papilio hector, and Telchinia viole alive but

mutilated. A Shrike came and looked at them keenly from a tree

close by, but did not attack them. A small Cuckoo flew over

them twice, but took no notice of them.

21.10.09. Put down H. misippus $ 5, H. core 4. The Shrike,

perched in an oleander bush, evidently saw them, but for quite an

hour took no further notice though it took several insects close to

them. It eventually took one, perhaps two, Eupleas. I picked

up the others.

Magpie Robin. Copsychus saularis.

25.1.09. Put a number of Zerias hecabe (unpalatable) in the

porch of my house, where the Robin comes to feed morning and

evening; some had their wings removed, but it took no notice of

any of them.

4.2.09. Placed 1 7. viole, 1 FE. core, 2 T. hecabe, and 1 Precis

lemonias wingless, and 1 normal 7’. hecabe on the veranda. The

bird ate the Huplea with difficulty owing to its being very dry,

and it took the body of the wingless 7. hecabe in its bill, but

dropped it almost immediately as it was too dry ; it took no notice

of the others.

6.2.09. Placed specimens of the above on the veranda dead

but uninjured, and a wingless 7’. hecabe ; the bird ate the latter

but took no notice of the others.

Mr. Ormiston informs me that a Magpie Robin in his garden

has become so confidential as to take food from his fingers and

that it will eat ‘‘ almost any kind of butterfly when thrown to it,”

but he has never seen it catch one. Neither have I during the

seven years I have closely observed this species.

_ 21.10.09. Put down #. core 5 and D. chrysippus 1, with the

wings on one side removed. A young Magpie Robin, as shown by

its speckled breast, captured one Huplwa, and though evidently

738 LT.-COL. N. MANDERS ON THE

somewhat alarmed at its size, killed it and, after the usual difficulty

with the wings, swallowed it. It immediately captured and ate a

second, third, and fourth ; this last was a very vigorous insect and

fluttered a good deal before it succumbed. The bird was then

frightened and flew away, but carried the Huplea with it; it took

quite another ten minutes to get rid of the wings, and during

the process it twice flew off to capture small flies; it eventually

swallowed it. The D. chrysippus was, I have no doubt, eaten

by a Calotes which ran out of the grass close by where I had

put it.

22.10.09. Put down 2 Kupleeas, one dead and one moribund,

1 Hypolimnas bolina g dead with wings closed, and 3 H. mistp-

pus 3 alive and all lively. The young Robin immediately flew

down and tackled the Huplea, mangled it for some time and then

dropped it and flew away ; it returned shortly afterwards, picked

it up and flew away with it. Directly after, another young bird

flew off with and devoured the other Huplea. This attracted the

notice of the old birds, one of which, I think the cock, flew down,

but before he could seize a butterfly was hustled off by his mate,

who picked up two H. misippus and flew off with them. The

one H. misippus and H. bolina were left.

26.10.09. Put down H. misippus ¢ 2, H. bolina 2 1, Junonia

(Precis) almana 2, Pyrameis cardui and Catopsilia pyranthi, all

with wings on oneside removed. The H. misippus fluttered most

and attracted the attention of Calotes versicolor, which pounced on

and ate both of them; something then frightened it off. Next

an old cock Magpie Robin caught sight of the 1. bolina, seized and

killed it after a lively chase, and finally disposed of it. It was a

very long time beating off the wings and made many attempts to

swallow the fly, before it was finally successful. Three or four

times I thought the bird was going to leave it altogether. After

swallowing the fly, it went off and drank at the runnel close by.

Shortly afterwards, a young one of the same species caught sight

of the Junonia and captured it; this disturbed P. cardui close by,

and the bird dashed from one to the other, not able to make up

its mind which to take, when the old hen bird came and tried to

get one, but the youngster was too sharp for her and managed to

swallow both. I should have said that this bird hopped over the

Catopsilia, which was moribund and motionless, to seizethe Junonia.

Nore. I have now little doubt that so long as the butterfly is

motionless, resting, as these mutilated butterflies generally do,

with their wings expanded, they do not attract attention ; but

directly one moves, whatever the species happens to be, it is the one

to be seized and eaten, even though so-called palatable species are

close by. All these butterflies are flying commonly in the garden,

but I have never seen them molested.

28.10.09. Put down H. misippus 3 2, H. core 2, Catopsilia

pyranthi 2, Terias hecabe 1. The Robins came for them the

moment I went off; the old cock bird seized one Huplaa and made

off with it, and the young bird the other; this latter, after

VALIDITY OF SOME FORMS OF MIMICRY. 739

ineffectual efforts to break off the wings, left it and caught a

Catopsilia and ate it at once, and then the 1. misippus, afterwards

perching just above the place where I had put the butterflies.

The old bird saw the Huplea which had been killed by the young

one and regarded it for some time, then it flew down, pecked at

it, looked at it again and then flew off with it. The 7’. hecabe

managed to struggle into the grass and was lost.

Curiously enough, while this was going on, a Catopsilia

pyranthi was actually laying eggs within two feet of where I had

put down these butterflies, “and within twenty feet I found this

afternoon eges and larvee of Huplea on the oleander. This seems

to me to show that the butterflies, when whole, are not molested

because, I suppose, they are difficult to catch.

12.11.09. Put down three H. misippus 3, one a partial cripple,

the others with two wings off. An old hen Robin came at once,

and flew off with one of the wingless ones to a bush about twenty

yards off and ate it; but it did not seem very hungry. A young

one a few minutes after came and took the crippled fly and ate

it after the usual difficulty ; it came back in a minute or two and

ate the third one. The butterflies had emerged in the morning

and were consequently full of juice.

16.11.09. A young Robin made off at once with a newly emerged

H, nusippus 3.

18.11.09. Put down four mutilated recently emerged /7. misip-

pus $. The young Robin flew off with one from which the wings

had been removed and ate it; a few minutes after an adult cock

Robin came and ate the three others one after the other.

19.11.09. Put down four mutilated recently emerged H. misip-

pus 3S in the front garden on the drive. A Calotes ophiomachus

ran off with one anda Brown Shrike (Lanius cristatus) with two

others; the fourth, which had its wings only partially developed,

got into the grass, grew its wings, and eventually flew off. I

released three or four butterflies at the same time, and they flew

off strongly enough and were not chased by the Shrike, which was

sitting on a tree close by me.

21.11.09. Put down three H. misippus 2 form diocippus,

which resembles Danais chrysippus; the hen Robin came at once

and ate one and flew off with another ; a young bird followed its

mother, and flew off and ate the other. These butterflies had

hatched out that morning and the wings on one side had been cut

off. I next put down five more females, all with two exceptions

with the wings entirely removed. The cock bird took one of the

half-winged ones and then ate a wingless one. The young bird

then returned and finished off the remainder. ‘These female

butterflies evidently derived no protection from their resemblance

to D. chrysippus, and so far as two species of birds are concerned,

H. misippus is a palatable butterfly.

8.1.10. Found Lycana (Zesius) chrysomellus Q fluttering on

the ground ; it was headless and with a piece out of one hind

wing, probably caused by a Sparrow,

740 LT.-COL. N. MANDERS ON THE

During this month (November 1910) I have been breeding

Hypolimnas misippus freely, and the Magpie Robins come every

morning on the chance of getting one. I have tried them with both

males and females, crippled and perfect, and always with the same

result. They are immediately seized, well beaten, and swallowed

after considerable difficulty. 1 noticed on one occasion a perfect

female resting on the ground with wings widely expanded but

insufficiently strong to fly; the resemblance to D. chrysippus was

perfect, but the Robin seized it without hesitation.

One day the large Hill Crow—an occasional visitant—carried

off a crippled male with wings quite undeveloped ; it pecked at it

twice and then dropped it, shaking its head with every appearance

of disgust. I remembered that the insect had fallen into and was

well covered with the red liquid these butterflies always evacuate

on emergence, and thinking that this was the cause of the Crow’s

discomfiture, I covered a crippled female with the stuff and threw

it on the ground: a Magpie Robin soon came and saw it, and

shook its head once or twice after pecking at it, but it swallowed

it in a short time.

Mynah. Acridotheres tristis.

This bird belonged to Mr. O. Wickwar, F.E.S., who kindly

assisted me. The bird was quite young and was allowed perfect

liberty in a large garden, where it fed freely on grasshoppers

and other insects ; it had abundant insect food, and was also

accustomed to fill up its dietary by visits to the kitchen for

odd scraps. I have placed in brackets the presumed palatability

or otherwise of the species experimented on.

3.1.09. Given Huplea core (unpalatable), took it readily, but the

wings seemed to bother it considerably, so gave it another with

its wings shortened, this it ate readily enough and then went

back and finished off the first one. We then gave it Papilio lanke-

swara (palatable ?) which was also readily eaten. Half an hour

after gave Papilio (Menelaides) hector (unpalatable) ; this puzzled

it for a bit and it seemed disinclined to eat it, evidently on

account of the large wings, for when these were removed it ate

the body with relish, even hopping off the veranda after it when it

fell over the edge. <A couple of Telchinia viole (unpalatable) (an

Acreine) g and 2 followed, and then Hypolimnas bolina 2 (un-

palatable ?) and Delias euwcharis (unpalatable); all these received a

pinch on the head, were well pounded, wings partially removed,

and the remainder eaten.

24.1.09. The bird had been kept without food for some time

and was decidedly hungry. 3 P. hector, 1 7. viole, 1 Mycalesis

ceylonica (palatable ?), and 1 Polyommatus betica (palatable 2)

were put in a row outside the cage ; when this was opened the bird

hopped over them and made straight for the kitchen, where it was

accustomed to pick up odds and ends. After some persuasion it

ate 1 P. betica and 1 Lycena (Zesius) chrysomellus 2. Some

three hours afterwards it ate 1 Precis almana (unpalatable),

VALIDITY OF SOME FORMS OF MIMICRY. 741

1 Papilio hector and 1 P. aristolochic ; it seemed a good deal

worried by the wings. Afterwards neglected J/. ceylonica and

T. viole, but ate one wingless Huplea core.

The conclusion we arrived at from the above experiments was

that butterflies were not its natural food, but that when hungry

it would eat them indiscriminately, and that the palatability or

otherwise of butterflies was of no account with this species of

bird.

The dietary of Ceylon insectivorous birds is fairly well known,

and we are now in a position to discuss the questions—Do the

birds of this island eat butterflies largely ¢ If so, do they eat them

in sufficient quantity to produce any form of mimicry? and do

they show any discrimination in their attacks? In other words,

ean the terms palatable and unpalatable as applied to butterflies

be maintained.

As regards the first question, it will be granted that there is

a greater destruction of butterfly lfe than has hitherto been

supposed, and the following observation on a Bee-eater, though

necessarily a rough one, shows clearly that the destruction is

sufficiently severe to produce mimicry, provided of course that

the agents showed sufficient discrimination in their attacks.

The road from Trincomali on the north-east coast to Anuradha-

pura, runs through fifty-eight miles of thick forest which is cut

back some thirty paces on either side, thus affording a convenient

place for butterflies which avoid the dense jungle. Between the

hours of 9 A.M. and 10 a.m., I counted the number of butterflies

between the third and fourth milestones from Anuradhapura, and

they came to one hundred and ninety-five : the same day, in the

outskirts of the town I watched a Bee-eater feeding from 12.45

to 1.45 p.m., and during this time it caught twenty insects ; on

only one occasion could I be certain that the capture was a butter-

fly, and this was undoubtedly Catopsilia pyranthi. The bird feeds

from about eleven o’clock till five.

Motoring between these two places | calculated roughly that

there was a pair of these birds to the mile *, and consequently

the whole of the butterflies along this road would be cleared off

in about a fortnight unless they received an accession of strength.

The calculation is necessarily a rough one, but it gives a good

idea of the struggle for existence that is constantly going on.

This observation was made January 7th, 1909, at a time of year

when butterflies are less numerous than usual.

The question whether discrimination is shown by birds in their

attacks on butterflies is of the greatest importance in mimicry,

and on the answer depends the fate of both Batesian and

Miillerian mimicry.

I do not attach much importance to the fact, curious though it is,

* Bee-eaters are particularly fond of perching on telegraph-wires.

742 LT.-COL. N. MANDERS ON THE

that in the observations I have been able to collect, the Hupleines

and Danaines, popularly supposed to be highly distasteful, figure

more largely as victims than any other group. I believe this to

be simply due to the fact that these butterflies occur in very

large numbers, and not that distinct preference is shown for

them. Admitting that more evidence is needed, I doubt whether

future investigations will reveal any marked preference in those

birds which are mainly instrumental in the destruction of

butterflies, for the reason that their dietary is of such a mixed

character ; and if this were so, or if what I have here set forth

be considered sufficient to settle the question, it is difficult to

avoid the conclusion that the unpalatability of these butterflies

has been assumed on insufficient data. It is interesting to recall

Professor Meldola’s remarks written so long ago as 1879, when

Miller first propounded his theory of mimicry (Proc. Zool. Soc.

Lond. 1879) :—

“ ...it may be fairly asked how far we know that such

imitated groups as Heliconius, Huplwa, Danais, Acrea, etc., are

distasteful. But very few observations have, as far as | am

aware, been made even upon these groups which are generally

admitted to be the objects of imitation, and I certainly know of

no systematic exper iments conducted with these models and their

insectivorous foes.”

The Bee-eaters seem to show some partiality for the yellow and

white butterflies of the Catopsilia and Appias group ; but whether

this is more apparent than real is not clear. It may be that these

butterflies are more readily seen and easier to capture than

others ; but if it could be proved that there isa distinct preference

for them, it is noteworthy, considering the destruction that

undoubtedly takes place, that though very variable they do not

act as models or mimics, or form Miillerian combinations, either

in India or Ceylon.

Failing the butterfly-eaters, what evidence is there that the

birds of group 2, and group 3, show preference in their more or

less desultory attacks ? There is no doubt that those experimented

on showed none, and that they took no notice of butterflies unless

they were mutilated and rendered easy of capture. I should

much wish to see further experiments on wild birds of these two

groups undertaken, but if the butterfly-eaters do not conduce to

mimicry, it is doubtful to my mind if the partial feeders would

do so.

In the present state of our knowledge it is difticult to say what,

is or what is not an unpalatable genus, and the position is further

complicated by the proposition that unpalatable species are killed

in numbers sufficient to produce a special form of mimicry. It

is unfortunate that theoretical considerations rather than observa-

tions and experiments in the field have hitherto preponderated

in this matter. It seems to me that the terms palatable and

unpalatable are not justified at present,

VALIDITY OF SOME FORMS OF MIMICRY. 743

The Miillerian Theory.

The supporters of the Millerian theory hold the view that it is

chiefly by the attacks of the young inexperienced birds that this

form of mimicry is produced. Professor Poulton puts the case

as follows :—‘ The Miillerian theory presupposes that only young

birds test the palatability of a few members of each convergent

group in their locality and henceforward, except when driven by

hunger, avoid all the members, so that the recent tendency to

explain so many of the resemblances on Miillerian rather than

on Batesian lines is in harmony with the conclusion that the

members of such groups are not greatly attacked by adult birds.”

(Hssays on Evolution, p. 270.)

I have already expressed the opinion that it is unlikely that

young birds, except those in group 1, indulge in tasting experi-

ments on butterflies, but as IT am quite willing to admit that such

an opinion may be founded on insufficient data, and as I was un-

able to find the necessary evidence required by the Miillerian

theory, I approached the subject by another line of investigation,

which depends on the time of the nesting of the birds and the

broods of the butterflies.

The birds breed once a year, not twice as is the casein Mauritius.

They begin in March or April, sometimes early in May, according

to the season. When the March or April rains known as the

little monsoon bring out a large increase of insect life, the birds

immediately begin nesting amt the young birds are off the nest

and begin to forage for themselves i in May, June or early July.

The average life of an insectivorous bird is probably not more

than four or five years, and we may assume that tasting experi-

ments gradually grow fewer in number and are completed when

the bird is about six months old, 2. e. about the month of October.

In estimating the number of broods of butterflies in the year,

which vary much according to the species, I will direct attention

to two of the more striking cases of mimicry, that of the Kuplcas,

forming a Miillerian combination, and Papilio polytes with its

trimorphic female mimicking P. aristolochic and P. (Menelaides)

hector. They may be taken together. In January, February and

March, that is to say in the dry weather (I am speaking more

particularly of the plains), there is a very small but continuous

series of broods which depend on the weather for their develop-

ment. If it is very dry, the eggs, larvee or pups, as the case may

be, lie dormant, but with favourable meteorological conditions

such as a shower of rain, the eggs hatch, the larve shake off their

lethargy and feed, or the butterfly emerges. Mr. Mackwood in-

formed me that on March 24, 1908, in “his garden at Colombo,

eggs, larve and pupe of Euplea core could be found together on

the same tree. The majority of the pupe do not, however, hatch

out but remain quiescent until the April rains, when there is an

astonishing outburst of butterfly and other insect hfe. With the

onset of the south-west monsoon ut the end of May or beginning

744 LY.-COL. N. MANDERS ON THE

of June, the broods become larger in numbers and more frequent,

and this goes on until the end of August or beginning of

September, when there is a further spell of dry weather similar to

but not so pronounced as that in the early part of the year, when

the broods again become smaller and less frequent, but at the same

time produce the individuals which take part in the migratory

flights of the monsoon in November and December.

Strictly speaking, P. hector and P. aristolochic, though following

the above sequence of events, do not usually form part of the

flights, but they are nevertheless at their maximum at this time ;

the Eupleas and Polytes undoubtedly do so. We have now to

judge what influence the inexperienced young birds off the nest

in May, and their experiments concluded in October, can have on

these species. A butterfly the size of Huplwa core pairs during

its first flight, if we may judge by the cabinet condition of those

ovipositing, sail begins to lay its eggs three daysafterwards. The

usual number is about two hundred and fifty, which are deposited

according to the weather in about ten days (I have known one

hundred egos laid in five days). What becomes of the parent

after this? Whether she dies a natural death or becomes the

victim of a tasting experiment is immaterial, her time of danger

is a brief fortnight. As the females are less in evidence than the

males, fewer of them would be captured, especially if we agree with

Professor Poulton’s opinion that the Miillerian theory presupposes

that only young birds test the palatability of a few meinbers of

each convergent group in thei locality. To bring to such per-

fection the cases of mimicry I have selected, we must assume that

such a victim would be one having less converging characteristics

than the others ; and it must also be borne in mind that unless

she is killed within three days of her emergence, she will have laid

a certain number of eggs which will produce butterflies similar to

herself. It is difficult to understand how the broods of butterflies,

numbering some thousands of individuals, born between October

and the following nesting season, would be in any way affected

except in the very smallest manner. Nodoubt Nature is infinitely

slow in her methods, and we have no reason to suppose that these

cases of mimicry have been produced, otherwise than by a very

lengthy process of weeding out ; but even if we grant this, there

is a still greater difficulty in the case of Hypolimnas misippus, the

well known mimic of Danais chrysippus. In Ceylon the former

appears on the wing in October, when as I have said tasting

experiments are over. It remains on the wing until the end of

the year, when it disappears until the following autumn. There

are so far as I can see only two ways of getting over this diffi-

culty—either by assuming that the inherited tendency to produce

this form of mimicry has become so fixed that the withdrawal of .

the factor that produced it is immaterial, though there is no reason

for this supposition, or that there is a more or less constant influx

of the species from India. There is very little doubt that a certain

number of Ceylon butterflies in their annual migratory flights

VALIDITY OF SOME FORMS OF MIMICRY. 145

find their way to India, but I can find no evidence of a reverse

condition of things, and one can scarcely credit that the few chance

stragglers which possibly find their way across the sea could keep

up this perfection of mimicry in the south of the island. The

supposition that the sudden appearance of a previously unknown

species would produce further tasting experiments will not hold

good in this case, as the mimic so “closely resembles its model

D. chrysippus, which is on the wing all the year round.

Huperiments on Young Lirds wr Confinement.

I am extremely doubtful as to any real value accruing from ex-

periments on caged birds, whether nestlings or adult. No one, I

imagine, believes that all butterflies taste alike: no doubt some

are more tasty than others, and caged birds fed upon butterflies

even with other insect food would no doubt learn in time to dis-

tinguish the different kinds ; but this procedure to my mind begs

the question, as 1t assumes that butterflies are an ordinary article

of food in the wild state, a proposition regarding which the

evidence here brought forward does notaltogether support. The

ease 1s different with Coleoptera, Hemiptera, Diptera and the

like, which are known to be the staple food of birds. Lloyd

Morgan’s carefully conducted experiments leave no doubt that

certain species of birds, probably all, have very little instinct as to

what is good, and what not, and that they learn by imitation

and tasting experiments. My observations lead me to believe

that the former is very important. I briefly epitomize my own

conclusions.

1, Young birds probably learn at first in a general way

what is their natural provender by what is brought to them

in the nest.

2. That this is further developed when they have left the nest

but are too weak to accompany the parents when they are

foraging for food.

3. That when they accompany the parents, as they do for a

longer or shorter time according to the species, they notice the

insects caught and attempt to capture them themselves.

4. When they are left to shift for themselves they carry on

what they have learnt, and during this time they undertake

tasting experiments, but with the exception of the birds in group 1,

those on butterflies are few in number ; first, because they have

rarely or never had butterflies brought to them in the nest ;

secondly, because they have very rarely seen their parents catching

them, and so neglect them; thirdly, because they have considerable

difficulty i in catching them, and the process of getting rid of the

wings is tedious and lengthy and the morsel fluffy, and possibly

not always agreeable. If these observations are confirmed by

further experience, they would account for the fact that attacks

on butterflies are less frequent than those on other insects.

746 LT.-COL. N. MANDERS ON THE

Instances of Imitation by Young Crows.

July, 1910. 1 saw an old crow and two young ones on the Rifle

Green this morning; one of the youngsters had hold of a bone

with a piece of gristle attached to it. It was so firmly adherent

that the bird could not detach it as the bone constantly moved

with the bird’s efforts, and eventually it gave up. Then the old

bird, which had been standing by all the time, went to the bone,

put its foot on it, thereby gaining a purchase, and tore off the

gristle without difficulty; the young bird after two or three

attempts did the same.

September, 1910. An old crow had a piece of hard boiled

potato off which it was picking pieces and giving them to a

full-fledged young one close by. A goodly number of detached

pieces lay on the ground and attracted the attention of some

other crows, which flew down and began picking them up ; seeing

this the youngster did likewise, though it made no attempt to do

so before their arrival.

Seasonal Dimorphism—Cryptic Defence.

I should not conclude this study of mimicry without discussing

that form of it which is known as ‘‘ cryptic defence,” and especi-

ally that which is so noticeable in the seasonal changes of so

many tropical butterflies.

It is commonly believed to have been produced by natural

selection acting through the medium of insectivorous foes, the

more exact and perfect imitations found in the dry season being

due to the paucity of insect life at that time of the year, which

produces a greater keenness in pursuit and a greater struggle for

existence. The argument has been put forward in full by

Professor Poulton in his ‘ Essays on Evolution,’ page 203.

I hope in the near future to deal more exhaustively with this

subject, but at present will only direct attention to two species

occurring in these islands, a study of which does not favour the

usually accepted views. The contention for the production of the

dry season form rests upon the premiss that ‘the dry season is a

time of far greater pressure than the wet”; for although the

enemies of insects are fewer, the insects themselves are pro-

portionately even more reduced, and ‘‘the light thrown by recent

investigation leads us confidently to believe that the differences

between the seasonal forms—hitherto devoid of interpretation—

have a meaning and a value in the struggle for existence and

came into being under the sway of natural selection ” (Powlton).

Though it is probably correct to say that in countries such as

S_ Africa and India, which havea continental climate, the seasons

are such as to produce a wealth or poverty of insect life, it is de-

cidedly incorrect to assume the same with regard to the islands

we have been investigating. There is no doubt that at no season

of the year is there in any of them a paucity of insect life, and at

VALIDITY OF SOME FORMS OF MIMICRY. 47

no time would an insectivorous bird, or reptile, find any difficulty

in procuring its daily sustenance.

Bourbon and Mauritius are very largely under sugar cultivation,

and this necessitates constant manuring of the “fields! with a

consequent abundance or super abundance of flies of all sorts

throughout the year. The rainfall, though greater in the wet

season, 1s not infrequent during the dry season, and this also

favours insect life; and if we add to this the consideration of

the practical absence of butterfly-eating birds and reptiles, we

can estimate the difticulty of believing that these changes are the

outcome of natural selection in these two islands.

In Ceylon Mr. E. Ernest Green, who has lived thirty years in

the island and who knows every part of it, writes as follows :—

‘Though insects are more abundant at certain seasons, J have

never experienced any part of Ceylon where there was anything

approaching a dearth of them. I know that Iam always busy

pinning and setting throughout the year.

‘““T sometimes wish that pee was a short dead season, when one

could devote oneself to other work without being distracted by the

constant accumulation of material.

“T doubt if Melanitis is ever subject to much worry from birds.

It lives in the shade and never moves during the daytime, unless

flushed by some big animal. Jam now receiving (16.8.10) both

dry and wet season forms of J. tambra from Kandy.”

An allied species Melanitis leda occurs also in Bourbon and

Mauritius, and it is to be remarked that the dry season forms

begin to appear before the advent of the dry season, that is to say

before any form of stress would tend to make itself felt.

In explanation, it may be suggested that the butterfly was in-

troduced from the locality where natural selection produced these

changes and that it is simply carrying on an inherited tendency.

That it is an introduced species is highly probable, but it has been

known to entomologists in Bourbon and Mauritius for at least

sixty years, and it differs in no way now than in the time of

Boisduval. It is difficult to believe that the factor which pro-

duced this cryptic defence being removed and no longer required

would not have led to some other form of colouring, or a return

to that ancestral type from which these forms were evolved. The

above remarks apply equally to Wycalesis narcissus, Precis rhadama

(introduced 1858), and Terias floricola, and I have made a further

study of Terias hecabe in Ceylon. It is very frequently the case

that the wet form continues to appear well into the dry weather

and vice versa, but to a less extent; this has been remarked on

frequently, but so far as I know no exact observations have been

made. In Colombo there was no rain from November 19th till

December 10th, 1908, thence to January 6th, 1909,-70 of an inch,

but of this no less than ‘57 fell on one day (Dec. 19th); such an

absence of rain in a tropical country at once causes a general drying

up of vegetation and the assumption of dry weather conditions.

At weekly intervals I captured all the Terias I could, which were

748 ON THE VALIDITY OF SOME FORMS OF MIMICRY.

accustomed to breed on a hedge of Madras thorn in an isolated

position in my garden; the results of such counts showed 73 wet

forms, 1 intermediate, and 19 dry, and it was not until January

28th that the wet forms were entirely replaced.

The butterfly takes exactly a fortnight to pass from larva to

imago, and thus we have approximately three or four broods of

wet season forms produced under diy season conditions. We

must assume that as the butterfly is seasonally dimorphic it stands

in need of protection, yet so far as I could ascertain the wet forms

suffered no diminution though exposed to what were, or should

have been, adverse circumstances.

The butterflies rested during the night and in cloudy weather

on the under surfaces of the leaves of Vincéasp.?, a small flowering

shrub with pink flowers and small oval green or frequently yellow

faded leaves. It often collected gregariously, two or even three

being on the same leaf and perhaps ten on the same plant. The

position was an admirable one for protection from the wet, and

also from small predatory foes which seldom look upwards when

hunting for prey. The appearance of the plant is the same

throughout the year, and the butterfly derived no advantage from

its change from one seasonal form to the other.

The following experiment makes me still further doubtful of

these effects being due to natural selection.

If we take the pupa of a somewhat similarly coloured butterfly

which is not seasonally dimorphic, such as Papilio demodocus or

Papilio demoleus, and expose it to a hot dry temperature, we can

produce an insect with much of the yellow on the under surface

replaced by red. I am almost persuaded that these rusty red

spots are a vestige of a character at one time common to certain

Pierines and Papilionines which is more or less reproduced by

heat and dryness, if of sufficient duration and intensity under

natural conditions in the Pierines, but in the Papilionines oniy

under artificial stimulation of a like but exaggerated character.

CoNCLUSIONS.

1. It has been shown that in Bourbon and Mauritius there are

no butterfly-eating birds or reptiles; so that the cases of mimicry

occurring there cannot be due to their influence.

2. In Ceylon it has been conclusively shown that the butterfly-

eating reptiles are impartial feeders.

3. That a trained observer can distinguish the majority of

these mimetic butterflies at a distance of about twenty or thirty

feet and frequently at the same number of yards; and this being

so, it is certain that a bird which has to depend for its existence

on its powers of observation, could after a few failures be able to

discriminate them at the same and probably at a considerable

greater distance.

4. That Drongos feed largely upon Eupleeas, and this being so,

DISTRIBUTION OF THE GENUS MEGAPODIUS IN PACIFIC. 749

a Papilio mimicking them obtains no protection in the vicinity of

these birds.

5. There is no bird in Ceylon known to eat butterflies that dis-

tinctly discriminates as an adult between one species of butterfly

and another.

6. It has been shown that there is a great destruction of

butterfly life in the dry zone, and that here, if anywhere, Miillerian

or Batesian mimicry might be induced, but the destroyers are

largely migratory and their attacks are not selective.

7. That the number of broods of butterflies which occur be-

tween the termination of tasting experiments in one year and the

commencement of them in the next is so great that any influence

which could be wrought by such is almost inappreciable.

8. The little evidence available shows that young Ceylon birds

imitate their parents in their choice of food; but as regards butter-

flies, the fact that there is no discrimination shown by adults leads

one to conclude either that few or no tasting experiments were

undertaken in youth, or, what is more probable, that their taste

with regard to them is indifferent.

9. Itis questionable, and so far as an accurate knowledge of one

species goes it is definitely shown, that that form of mimicry re-

presented by wet and dry season forms (cryptic defence) is not

produced for the protection of the species, inasmuch as many

(four) succeeding broods of the wet weather form may be found

under dry season conditions without detriment to the species.

34. The Distribution of the Avian Genus Megapodius in the

Pacific Islands. By J. J. Lister, M.A., F.R.S., F.L.S.,

¥.ZS.

[Received and Read May 9, 1911. |

(Text-figure 166.)

The Megapodiide or Mound Builders are, as is well known,

large birds, with comparatively feeble powers of flight, con-

stituting a family of the order Galline. They are distributed over

the islands of the Kast Indian Archipelago and Western Pacific,

from the Philippines and Borneo to the New Hebrides, and are

found in several parts of the continent of Australia. Four out-

lying species of the genus J/egapodius are found in the Nicobar,

the Pelew, and the Marianne Islands, and, far out in the Pacific,

on the little island of Niuafou, belonging to the Tongan group.

As we cannot suppose that the birds found in these outlying

islands, remote from the other species, can have flown across the

intervening tracts of ocean, we are presented with the problem:

How did they reach these islands ?

The solution to which M. Oustalet gives his adhesion, in his

Proc. Zoot. Soc.—1911, No. LIT, 52

750 MR. J. J. LISTER ON THE DISTRIBUTION OF THE

monograph on the Family *, is that all these localities have been

at one time connected by land, which has since been to a large

extent submerged.

*Kermadec I$

“o

| rat Howell

Map illustrating the Distribution of the Genus Megapodius.

NEW ZEAL

160

Text-fig. 166.

ica

ep}

ayes

a <

= =|

She

Wallace had, in 1876 7, expressed the opinion that the Megapode

* “Monographie des Oiseaux de la famille des Mégapodiidés.” Ann. d. Sciences

naturelles, 6 sér. t. 10 & 11, 1880-81.

+ Geographical Distribution of Animals. London, vol. 11. p. 342.

AVIAN GENUS MEGAPODIUS IN THE PACIFIC. 751

of the Nicobar Islands had probably been introduced by the Malays,

but Oustalet cannot regard this explanation as plausible, on the

ground that ‘“‘ we possess no positive proof of the domestication of

Megapodes by the Malays or the savage people inhabiting the

Oceanic Islands.”

My object in this paper is to point out the reasons which appear

to me to make it probable that the distribution of the genus

Megapodius has been, as Wallace suggested in the case of the

Nicobar bird, considerably modified by human agency ; and that

the species found in these outlying Pacific islands have been

carried there by man.

Since Oustalet published his monograph the Megapodiide have

been again reviewed by Ogilvie-Grant in his Catalogue of the Game

Birds in the British Museum*; and in 1901 Rothschild and

Hartert t gave their revision of a portion of the genus Megapodius.

The determination of the limits of the species of this

genus is difficult on account of the variation in size and colour

presented by the birds inhabiting the same locality, and the fact

that the characters of those from different localities often merge

into one another. On comparing the results arrived at by the

authors mentioned, we find that nineteen species were enumerated

by Oustalet. If we take from these W. wallacei, which has been

placed by Ogilvie-Grant in a separate genus (Hulipoa), and M.

brenchleyt and M. brazieri, which had then been described only

from eggs or young birds, there remain sixteen species. Five of

these are united in two species by Ogilvie-Grant, and seven which

inhabit the area dealt with by Rothschild and Hartert are allowed

by these authors only subspecific rank under two specific names.

On the other hand, the species J. macgillivrayi, which is united

by Oustalet with the widely extended WZ. duperreyi, and regarded

as only a subspecies by Rothschild and Hartert, is reckoned a

distinct species by Ogilvie-Grant.

The peculiar nesting habits of the Megapodiide are well known.

Most of the species scrape together large mounds of earth or sand,

with or without vegetable matter, and the female deposits her

egos, which are very large for the size of the bird, at intervals of

several days, in excavations in these mounds. Incubation is

effected by the heat of the slowly fermenting mass, aided by that

of the sun, or by the sun alone; and the young are hatched in an

advanced state of plumage. They receive no attention from their

parents, and in some cases at least they are able to fly on the day

on which they are hatched. In some species many pairs of birds

frequent the same mounds or laying-grounds.

In almost all the countries where Megapodes occur, their large

eggs are highly valued by the natives as food, and their laying-

places are frequently visited for the purpose of obtaining them.

* Catalogue, Vol. xxii. 1893. Also‘ Game Birds’: Allen’s Naturalists’ Library,

London, 1895-7. :

+ “Notes on Papuan Birds.” Novitates Zoologicx, vol. viii. p. 135, 1901.

| 5o*

52 MR. J. J. LISTER ON THE DISTRIBUTION OF THE

There is clear evidence that the birds are at least semi-

domesticated in some localities.

Mr. C. M. Woodford *, speaking of J. brenchleyi (AL. eremita

of B. M. Catalogue) on Guadaleanar, in the Solomon Jslands, says :

“The birds lay in open sandy clearings, generally near the sea,

which are kept clear of shrubs and undergrowth by the natives,

and by the sand being constantly turned over by the birds... .

Many ‘thousands of birds congreg: ate at the same place, the laying-

yards being often some acres in extent.” Of the little island of

Savo, to the north of Guadaleanar, the same author writes } :

“although only about the size of a large pigeon ’ * the megapode

‘lays an ege bigger than that of a duck,” and “eggs form an

important item in the daily food-supply of the natives.” ‘The

megapodes lay their eggs on two large cleared sandy spaces and

nowhere else on the island. Upon these no weeds or grass can

grow as the ground is constantly being turned over by the birds

when digging ¢ holes to lay their eggs, and by the natives when in

search of them. The sandy spaces are fenced off in plots which

belong to different owners.” He adds that the natives are quite

indifferent as to the condition of the eggs when they eat them, it

is all the same to them whether they are newly laid or well

advanced towards hatching.

Mr. John Brazier, writing of a collection of eggs of Megapodes

exhibited before the Society: says t, ‘‘ When at San Christoval ”

(in the Solomon Islands) “I was shown an egg that Perry, a

white man living there these last five years, said was laid by the

‘Wild Fowl, and upon my visiting him a few days later, he had

just obtained another from the nest: of his domestic fowls.”

M. Freycinet, in his narrative of the voyage of the Uranie §, says

of the species which was discovered by this expedition on the

Marianne Islands: ‘‘ Espéce de gallinacée de couloir noir que jadis

les anciens Mariannais élevoient auprés de leurs cabanes ; elle est

aujourd’hui fort rare. Nos naturalistes lui ont donné le nom de

Mégapode la Perouse.”

In the volume on Zoology (p. 125) of the same work the

naturalists Quoy and Gaimard say of JIL. freycineti, which they

discovered on the island of Waigiou, to the west of New Guinea,

and which is now known to occur from the Moluccas to Western

New Guinea: ‘“ Sur les iles Vaigiou et Boni, ces oiseaux parois-

sent vivre dans une demi-domesticité, d-peu-prés comme les

canards qui habitent les marais que traverse la petite riviére de

Sévre, (Charente Inferieure).” One, brought by the natives, lived

sever: al days on the Uranie.

Professor J. Stanley Gardiner, whose investigations of the

Fauna and Flora of the Maldive and Laceadive Islands are well

* The Naturalist among the Head-hunters, PP. 100-101.

P. Z.S. 1888, pp. 249 & 260.

P.Z.S. 1874, p. 607.

Bi age autour du Monde..... sur les corvettes de S.M. PUranie et la

AVIAN GENUS MEGAPODIUS IN THE PACIFIC. 753

known, informs me that a sultan of the Maldive Islands, who

died in 1878, introduced Megapodes into an islet covered with

cocoanuts and scrub, forming part of the great atoll of Male in

that Archipelago. Where these were obtained is not known, but

Professor Gardiner thinks it probable that they were imported

from the Nicobar Islands, between which group and the Maldives

there is regular trade communication.

We have thus from four widely separated localities definite

evidence of the more or less complete domestication of Megapodes

by the natives. Finally Guillemard *, referring to Wallace’s view

that the Nicobar Island bird was introduced, says ‘‘ that this is

not impossible must be evident to every traveller in the Malay

Archipelago, for birds of this genus are often seen in captivity.”

With regard to the powers of flight of Megapodes, they are

compared by some authors with those of barn-door poultry.

Oustalet, however, recalls the fact t that a young specimen of

M. freycineti flew on board ‘ La Coquille,’ with a favouring breeze,

when that vessel was “plus de deux milles” (over two miles)

from land.

Le Souef ¢ mentions that IJ. duperreyi, although the birds are

‘very poor fliers,” occurs on the scrub-covered islands ‘a good

many miles” from the N.E. coast of Queensland. He surmises

that they may have been blown out during cyclones. .

Finsch § speaks of IW. senex, the species inhabiting the Pelew

Islands, as occurring on nearly all the sandy and rocky islands of

the group. Some of these are separated by intervals of some

three or four miles. He considers that the bird, ‘‘ which is a good

flier” (the term is of course used in a relative sense), may oc-

casionally fly from one island to the other. He also mentions

that the eggs are systematically taken by the natives.

But when all allowance is made for their powers of flight, it

would seem an extravagant suggestion, and one which I think

has never been made, that Megapodes could by this means have

reached the outlying islands in which they are now found. The

Pelew Islands are separated by nearly five hundred miles from the

Philippines, the nearest land to the west, and by a rather greater

distance from New Guinea to the south. The Marianne Islands

are some 600 miles to the E.N.E. of the Pelew Islands. Niuafou

is nearly 1000 miles to the east of the New Hebrides, the nearest

islands to it on which a species of Megapode exists.

We may now examine the geological nature and some other

conditions of these outlying islands in the Pacific on which Mega-

podes are found, as well as the characters of the species living

on them.

The island of Niuafou, or New Hope Island, although politi-

cally part of the Tonga group, is situated almost halfway between

* Cruise of the Marchesa, vol. ii. p. 122 (footnote), 1886.

+: IG. c, vols Xi. p69 ay

+ Ibis, 1899, p. 16.

§ “Die Vogel der Palau-Gruppe.” Journal des Museum Godeffroy, Heft viii.

Bd. iii.) 1875, p. 30 (p. 162 of the volume).

754 MR. J. J. LISTER ON THE DISTRIBUTION OF THE

the Fiji aud Samoa Islands. It is described in Findlay’s South

Pacific Directory * as a voleanic island with black lava rocks all

round the shores. It is 3 to 34 miles across, well wooded, and

some 500 to 600 ft. high. In the centre isa brackish-water lake

at sea-level in which are hot springs. Friedlander +, who visited

the. island in 1897, describes his visit to the nesting places of the

birds, which are on the shores of this lake. He had to swim

round some of the rocky points to reach them, and found the

temperature of the water that of a warm bath, and the rocks

under water too hot, in places, to rest his hands on them. He

says the island is an intermittently active crater, largely composed

of basaltic rock. An eruption occurred in 1886, when the whole

island was covered with ashes, and the Megapodes were nearly

exterminated. Owing to the ‘tabu’ imposed by the chief their

numbers had increased again, so that there was a fair number at

the time of his visit. The birds do not build mounds, as do many

of their congeners, but lay in holes which they excavate in the

voleanic sand. He is inclined to attribute the heat of the sand,

which he found on digging for the eggs, to the volcanic action.

We may note in passing that P. & F. Sarasin } found that the

Maleo (Aegacephalon maleo) of Celebes lays in the neighbourhood

of hot springs, as well as on the sea-shore.

The birds of Niuafou were collected by Mr. F. Hiibner and

described by Dr. Finsch §. With the exception of the Megapode

all belong to common Tongan species, but Finsch remarks on the

absence from the collection of four species (Ptilotis carunculata,

Halcyon sacra, Lalage maculosa, and Colluricinela heinei) which

besides being common in Tonga are present, or represented by

allied species, in Samoa and Fiji. At first sight 1t would appear

that the absence of these species from Niuafou might be at-

tributed to the destructive eruptions of the volcano, but as these

species are unrecorded from the not very distant group of Uvea

(Wallis Id.), their absence from Niuafou cannot certainly be

attributed to that cause. It must be admitted, however, that a

small volcano, still in intermittent activity, is the last place on

which the remnants of an ancient fauna would be expected to

survive. Had the bird been found on the high and ancient land

masses of Fiji or Samoa, the case against this view would not

have been so strong, but the birds of these islands have now been

so fully collected as to make it in the highest degree unlikely

that so large and useful a bird as a Megapode should have been

overlooked in them.

The Niuafou species, Megapodius pritchardi, was described by

the late Sir Walter Buller in his Supplement || to the ‘ Birds of

New Zealand,’ and included, under the name of ‘The Southern

* 3rd edition, p. 558.

+ Ueber die Nestlicher d. Megapodius pritchardii auf der Insel Niuafou.”

Ornithologische Monatsberichte, vil. p. 87, Berlin, 1899.

t Zeits. d. Gesellschaft f. Erdkunde, Berlin, 1894, pp. 375, 388, 396 & 398.

Z.S . 1877, p. 782.

|| Vol. i. p. 31.

AVIAN GENUS MEGAPODIUS IN THE PACIFIC. 755

Megapode,” in that fauna—on what appear to be wholly in-

adequate grounds.

In 1887, Mr. T. F. Cheeseman, the well-known Curator of

the Auckland Museum, to whose knowledge and kindness many

visitors to New Zealand ave indebted, visited the Kermadec

Islands, which are a scattered group lying nearly halfway between

the North Island of New Zealand and the Tonga Islands to the

N.N.E., and some 400 or 500 miles from either. Mr. Cheeseman

reported * that a Mr. Johnson, who had resided on Sunday

Island (a volcanic island, the most northerly of the group) about

fifteen years before, told him that “ prior to the eruption of 1876

a bird inhabited the floor of the large crater, which made

mounds of sand and decayed leaves, two to three feet high,

laying its eggs in the mounds. He was in the habit of visiting

the mounds for the sake of the eggs and young birds, and has

frequently taken 5 or 6 of the latter from the same nest at one

time.” The eruption of 1876 covered the floor of the crater and

apparently killed out the species. Mr. Cheeseman cautiously

observes that the evidence, such as it is, seems to point to the

former existence of a species of Megapodius on this island.

We may remark that the statement that five or six young

birds were taken from the same nest at one time is hardly

in accordance with the habits of the genus, for the eggs being

laid at some intervals the young ones are not of the same age,

and leave the mounds to feed for themselves soon after they are

hatched. The statement would be more appropriate to the young

of the Grey Duck (Anas superciliosa) which frequents this island.

It is the mound-building habit which, as Mr. Cheeseman says,

“seems to point” to the existence of a Megapode on Sunday

Island.

Sir Walter Buller + on the strength of this evidence includes

Megapodius pritchardi among the birds of New Zealand, in which

region the Kermadec Islands are included. He says: “1 have

no doubt whatever in my mind—notwithstanding the apparent

difference in their nesting habits—that Mr. Cheeseman was right

in his conjecture” that the Sunday Id. and Niuafou birds were

identical. (It will be noted that Mr. Cheeseman conjectured that

the genus, not the species, was identical.) On the discrepancy

that whereas the Niuafou bird lays in burrows, the Sunday Id.

bird is stated to have built mounds of sand and leaves two to three

feet high, he remarks (p. 33): “If the latter observation was

accurate it may have been due to circumstances of locality ana

environment, and by no means negatives the assumption of these

birds being one and the same species.” As we have seen, the

statement that the Sunday Island bird built mounds is the only

evidence we have of the existence of a Megapode on that island.

Through the kindness of Mr. Ogilvie-Grant I have had the good

* “Qn the Birds of the Kermadec Islands.” Trans. and Proc. of the N. Zealand

Institute, vol. xxiii. (1890) p. 219.

+ Supplement to the Birds of New Zealand, vol. i. (1905) p. 31.

156 MR. J. J. LISTER ON THE DISTRIBUTION OF THE

fortune to meet at the Natural History Museum Mr. Iredale, who

has recently resided on the Kermadecs with the object of studying

their fauna. He assures me that he was not able to obtain

any confirmation of the report of the existence of a Megapode

on Sunday Island which was given to Mr. Cheeseman, and finds

that the successors of his informant are not inclined to regard

that report as worthy of very serious consideration.

It therefore seems to me that we have no good evidence that

the genus Megapodius formerly inhabited the Kermadec Islands

and absolutely none that JZ. pritchardi lived there.

One good result, however, we owe to Sir Walter Buller’s

enthusiasm in claiming this species as a member of the New

Zealand fauna, and that is a plate representing the bird in a

condition of plumage not hitherto figured.

M. pritchardi belongs to the section of the genus with the

back and upper surface of the wings rufous brown, the breast

and belly lead or slaty grey, and in its general coloration perhaps

is nearest J/. cumingi, Dillw., of the Philippines and Borneo.

In the type specimen described and figured by G. R. Gray *,

and now in a somewhat dilapidated condition in the British

Museum, the bases of the quill-feathers, except the first, are

white; there is also some white among the upper tail-coverts.

A specimen in the Leyden Museum was described by Schlegel +

(and I have had an opportunity of examining it) which has,

as he observes, the upper tail-coverts pure white. The first

specimen which came to the Auckland Museum was described by

Buller ¢. It had no white on either quill-feathers or tail-coverts,

though the rectrices were white at the base§. As it was not

known then that this skin came from the same locality as

M. pritchardi, and it differed so considerably from the type of that

species, it was described by Buller as a new species—W. huttonz.

Buller’s plate (pl. ii.) in the Supplement to the ‘ Birds of

New Zealand’ shows no white in the plumage. In the descrip-

tion he says (p. 32):—“ Although absent in this specimen ”

(that named M. hattoni and perhaps the specimen figured in the

plate) ‘‘ most examples have a patch of white covering the basal

parts of the primaries and secondaries, the extent varying in

almost every individual. Some also have white markings on the

upper tail-coverts and basal part of the tail-feathers.”

It appears then that the feature by which M. pritchardi stands

apart from all other species of the genus—the occurrence of white

at the bases of the primaries and elsewhere—is a varying and

inconstant character.

To return to the comparison with MZ. cumingi, I find that

M. pritchardi has the top of the head slaty-brown rather than

brown, the sides of the head rather paler, the mantle brown tinged

with slate rather than olive-brown, and the belly a paler brown.

* P.Z.S. 1864, p. 41, pl. vi. + Mus. Pays-Bas, vii. p. 64.

Transactions of the N. Zealand Institute, vol. iii. (1870) p. 14.

Hutton, Trans. N. Zealand Institute, iv. (1871) p. 165.

AVIAN GENUS MEGAPODIUS IN THE PACIFIC. rey I

We may note that the species inhabiting the New Hebrides,

4M. layardi, belongs to the group of species having the mantle

and upper parts blackish grey, uot rufous brown as in

M. pritchardi.

There is one other fact which seems to point to the view

that the Niuafou bird is not indigenous to that island, viz., that

the native name, Mallow, is the same as that applied to several

species of Megapodiide i in the Malay Archipelago. This appears

in the specific name of Megacephalon maleo of Celebes. Mega-

podius cumingi is called moleo kitjil (= little moleo) by the

natives of that island, in distinction, probably, from the larger

Megacephalon maleo*. M. layardi is the Malow of the natives s of

the New Hebrides. Onustalet says that this name appears to be

applied indifferently to Megapodes by Malay hunters. Tis occur-

rence on Niuafon, far out in the Pacific among a population of

Polynesian speech, seems to suggest strongly that at some time

the name arrived at the island with the bird.

With regard to the Marianne and Pelew Islands, I have less

evidence to bring forward, but it was in the former group that

Quoy and Gaimard were told that the Megapode (i. laperousti)

had been domesticated. It is so closely similar to Uf. senex from

the Pelew Islands that M. Oustalet has regarded them as of the

same species T.

The geological structure of the Pelew Islands is discussed by

Semper 7, who shows that the islands are composed in part of

raised coral, in part of voleanic rock, formed during submarine

eruptions. If this is the case, there can be no remains on the

Pelew Islands of the fauna of a subsided land-mass, supposing

such a mass to have existed.

From the description of the Marianne Islands in the account

of the Voyage of the Uranie, above quoted, it is stated (Historique,

T. 2, p. 253) that they seem to have been formed in the remote

past by submarine eruptions, which have raised the floor of the

ocean, and that the reefs which have formed about the islands as

they have risen above the waves have since been raised with them.

So that it would appear that the same remark is applicable to

these as to the Pelew Islands.

4M. laperoustiand M.senex have the upper parts blackish grey, as

have M. freycineti, from the Moluccas and New Guinea, Jf. geel-

vinkianus from New Guinea and some adjacent islands, and

M.layard: from the New Hebrides. But they differ from these

and other species in the french-grey colour of the feathers of the

head. We must conclude therefore that this character has been

developed since their isolation, or else that the parent stock has

either not yet been discovered, or has become extinct.

In the British Museum Catalogue three “doubtful species”

* Meyer & Wiglesworth, Birds of Celebes, vol. ii. 1898, p. 671.

+ Of clic Guan Alea aes Library : Game Birds, vol. u. p- 182.

= The Natural Conditions Enxistenee as they affect Animal Life, by Karl

Semper, Chapter 8: International Scientific Series, vol. xxxi. London 1883, pp. 234

264.

758 DISTRIBUTION OF THE GENUS MEGAPODIUS IN PACIFIC.

are mentioned (p. 446) by Ogilvie-Grant, viz.: 1. Young birds

said to have been obtained on Lord Howe’s Island; 2. The

Megapode of Sunday Island in the Kermadec group; and

3. M.¢ andersoni of Gray from New Caledonia. The first of these

Mr. Ogilvie-Grant assures me is now known to have come not

from Lord Howe’s Island, but from New Hope Island, another

name for Niuafou. With the second I have already dealt. The

third is based on a reference in the MS. of Anderson, who

accompanied Cook’s third voyage, to a bird he called Tetrao

australis and briefly described as follows :—“ fusca nigraque ;

pedibus nudis.” The subsequent exploration of New Caledonia

has not revealed the presence of a Megapode on that island.

To sum up: There is evidence of the domestication or semi-

domestication of Megapodes in several parts of the area they

inhabit,—viz., in the Solomon Islands, Western New Guinea and

the Marianne Islands, and of their introduction into the Maldive

Islands. There is no satisfactory evidence that a Megapode has

ever existed on any Pacific island east of a line bordering the

Philippines, Solomon Islands, and New Hebrides except the

Pelews, Marianne Islands, and Niuafou. The geological character

of these islands, so far as we know it, lends no support to the

view that they could preserve the fauna of a sunken land-mass.

The birds of the Pelew and Marianne Islands are almost identical,

and on the latter group they were domesticated. The bird of

Niuafou is called by a Malay name.

When we consider the complex movements of the races of the

Western Pacific, of which there is much anthropological evidence,

and how easily Megapodes might be introduced into a new locality

by a canoe provisioned with their eggs, which are a staple native

food, it would appear that we have in human agency a probable

key to some of the anomalies of their distribution.

The analogy of the distribution by native agency of domestic

fowls, cousins of the Megapodiide, and of the dogs and pigs

which were found by the early voyagers on the Pacific Islands, is

obvious. :

How far the same cause may have been operative within

the main area occupied by the genus, and have given rise to the

anomalies in the distribution of the species alluded to by Oustalet,

is too large and complex a subject for me to attempt to deal with.

The Savo natives, says Mr. Woodford,* speaking of Jf. eremita

of the Solomon Islands, have a curious legend connected with this

bird. They hold the Shark in great veneration and say that their

island was made by the Shark, who brought stones together and

placed upon them a man, a woman, the Yam plant, and the

Megapodes. Things went well for a time and the people increased

and so did the Megapodes. At last the people went to the Shark

and complained that the Megapodes made havoc among the yam

* C. M. Woodford, P. Z. S. 1888, p. 249.

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ON THE MORPHOLOGY OF THE HELICINID#. 759

patches by digging holes to lay their eggs; so they asked the

Shark to take the Megapodes away. This was done, but now the

natives missed the Megapodes’ eggs, so they asked the Shark to

bring the Megapodes back but to confine them to one spot.

This request was also complied with, and the result may now be

seen. The Megapodes lay their eggs in two large and broad

sandy spaces, and nowhere else on the ccbnd.

I suspect that there is more than a grain of true history in

this legend, and that it records the fact that when the ancestors

of the natives came to the island, they brought with them two

main staples of their food-supply—yams and Megapodes.

35. Contributions to the Morphology of the Group Neritacea

of the Aspidobranch Gastropods.—Part Il. The Hett-

cinip#. By Givpert C. Bourne, M.A., D.Sc., F.R.S

F.Z.S.

[Received April 29, 1911: Read May 9, 1911.]

(Plates XX X.-XLIL*)

When, two years ago, the Society published the first part of my

contributions to the mor phology of the Neritacea (2), I had already

accumulated a number of observations on the anatomy of the

Helicinide, but deferred the publication of them until I was able

to obtain specimens of different species from various parts of

the Pacific region. Having experienced considerable difficulty in

obtaining specimens sufficiently well preserved for microscopical

examination, the publication of my results has been long delayed,

with the result that I lose the claim to priority for several minor

discoveries concerning the anatomical features of this family, for,

in the meantime, Thiele (10) has given an account of the anatomy

of Hydrocena cattaroensis in which is included a description of

the female generative organs of Helicina hubaryi, and the following

deser iptions lose much of the novelty they would have possessed

had they been published as soon as the facts were ascertained.

Previous to the publication of Thiele’s paper, our knowledge of

the anatomy of the Helicinide rested, for the most part, on

Isenkrahe’s (4) account of the anatomy of Helicina titanica.

Isenkrahe gave a sufficiently accurate description of the external

anatomy, the muscular system, the greater part of the alimentary

tract, and the pulmonary cavity, but he failed altogether to

distinguish the kidney, and his descriptions of the heart, the

nervous system, and the reproductive organs are defective. These

imperfections notwithstanding, Isenkrahe was able to confirm

Troschel’s opinion that Helicina, on account of its rhipidoglossate

dentition and other anatomical characters, was closely related to

the Neritide. .

Von Jhering (5) in 1877 placed the Helicinacea and Proserpinacea

* For explanation of the Plates see pp. 806-809.

760 PROF. G. C. BOURNE ON THE

in his class Orthoneura, order Rostrifera, sub-order Rhipidoglossa,

and gave a fairly accurate description, unaccompanied by a figure,

of the nervous system of Helicina (Stwranya Wagner) beryllina

Gld. E. L. Bouvier (3), in his great work on the nervous system

of Prosobranch Gastropods, gave a very complete account of the

nervous system of H. sagraiana d’Orb. and H. brasiliensis Gray,

laying stress on its close resemblance to the nervous system of the

Neritidz, and in addition he made some further observations on

the general anatomy, partly confirming and partly correcting and

adding to Isenkrahe’s descriptions. In 1902 Thiele (9) described

the male generative organs of Helicina japonica, and last year

he gave a description with a diagram of the female organs of

H. kubaryi, in addition to a succinct but sutticiently exhaustive

account of the general anatomy of Hydrocena cattaroensis.

The geographical distribution of the Helicinide, as is well

known, presents several interesting and ditticult problems. By

far the greater number of species are insular and confined to the

tropics. Such species as are found on continents are for the

most part limited to regions near the coast, very few being known

to occur any considerable distance inland. No Helicinide are

recorded from Africa. In Europe the group is represented only

by the genus Hydrocena from the Dalmatian coast, and this genus,

as Thiele’s recent work has shown, differs in several important

anatomical characters from Helicina and its more closely allied

genera. Georissa,a subgenus of Proserpina, is the only represen-

tative of the group in India, and no Helicinide have as yet been

recorded from Ceylon. The number of genera and species reaches

its maximum in the Antilles. The genus Helicina, as restricted

by Wagner, is fairly abundant in Mexico and the Centrai

American republics, and extends northwards into Texas and

Florida, southwards into Ecuador and Peru on the west coast and

to the south of Brazil on the east coast of S. America. Few

species, however, are recorded from the Pacific coast of S. America,

but, notwithstanding their comparative rarity on these shores,

the group reappears in great abundance in the Pacific islands,

extending as far east as the Marquesas and Paumotu Islands, and

having many representatives in the Society, Samoan, Friendly

and Fiji Islands, and in the New Hebrides and New Caledonia.

Several species occur on the east coast of Australia, and some

few are recorded from New Guinea, Celebes, Borneo, and Sumatra ;

none, so far as I can ascertain, from Java. But in this part of

the world the Helicinide attain their maximum in the Philippine

Islands, which are only second to the Antilles in the number of

species. From thence the group extends north, beyond the

tropical zone, to the Bonin Islandsand Japan. A few species

are found beyond the south-east coast of China and Siam, others

again in the Malay Peninsulaand Burma. Several species are

found in the Andaman and Nicobar Islands, but the group is

very poorly represented in the Indian Ocean. Aphanoconia

(Helicina) theobaldiana G. & H. Nevill is recorded from the

MORPHOLOGY OF THE HELICINID&. 761

Seychelles, and Psewdotrochatella undulata Morelet is a subfossil

form from Mauritius. None is known from Madagascar.

In a recent work of great value to the student of geographical

distribution A. J. Wagner (12) has revised the family Helicinide,

and, founding his diagnoses chiefly on the characters of the

operculum, has broken up the old genus Helicina into no less

than thirteen genera, reserving Lamarck’s appellation for the

American and Antillean forms which conform to the original

definition of the genus. Of the remainder I mention the largest

genera. Sulfurina has its centre in the Philippines and extends

thence to the Andamans, Nicobars, Moluccas, New Guinea, and

Tahiti. Aphanoconia, which also seems to be centred in the

Philippines, extends widely, to Japan, S$. China, the Malay

Archipelago, the Andamans, Nicobars, Seychelles, Moluccas, and

through Micronesia and Melanesia to the Paumotu and Sandwich

Islands. Stwranya has its centre in Fiji and Tonga, and extends

thence to the Carolines, Sandwich, Society, Hervey, and Solomon

Islands. Orobophana is found in Queensland and N.S. Wales and

extends through nearly the whole of Polynesia. Palcohelicina.,

with its subgenus Ceratopoma, is again a Philippine genus, and

extends to New Guinea, the Bismarck Archipelago, the Solomon,

Louisiade and Pelew Islands. The last-named genus is, according

to Wagner, closely allied to Helicina sensu restricto. Again, the

subgenus Letorquata of Helicina, which occurs in Mexico and

Central America with outliers in Florida and Texas, affords,

according to the same author, a transition to such a characteristic

Antillean genus as Alcadia. A consideration of these statements

leads to the conclusion that the Helicinidee are capable, by what

means we know not, of wide dispersal across seas and oceans, and

find conditions most suitable to their existence in proximity to

the sea. They appear to have originated in Mexico and Central

America, and to have spread eastwards to the Antilles, where

they found the conditions specially suitable, and have been

differentiated into several genera (Aleadia, L ucidella, Eutroch atella,

Priotrochatella, Proserpina) and numerous species, and one species

(H, substriata convexa Pf.) has found its way to the Bermudas.

Others have extended down the eastern coast of S. America, but

the Atlantic Ocean has proved an impassable barrier to their

further extension eastward. On the Pacific side the group has

been transported by some means unknown to us to the Pacific

Islands, and it would appear from the evidence that it did not at

first effect a lodgment in the more eastern islands, but in the

Philippines, from which centre it has spread in all directions—

eastward throughout Polynesia and to the Sandwich Islands,

southward to New Guinea and Australia, northward to Japan

and China, westward through the Dutch ‘Indies and Malaya to

the Andaman and Nicobar’ Islands. Very few have traversed

the Indian Ocean to reach the Seychelles and Mauritius.

Very little is known of the geological history of the group.

Helicina occurs in the post-Pliocene of N. America, but the

762 PROF. G. C. BOURNE ON THE

ancestral forms must have lived at a much earlier period, for

Proserpina is recorded from the Kocene of the Isle of Wight, and,

according to Kobelt (6), shells referable to the same genus have

been found along with Helia, Planorbis, Vi alvata, and three

species of Veritina inthe Lias of Somerset. There is some reason

for suspecting the correctness of the identifications in the last

case, and [ am unable to find any corroborative evidence of the

occurrence of Proserpina in the Eocene, but the distribution of

the Hydrocenids points to a geological history reaching well

‘ back into Tertiary times. Dawsoniella from the Carboniferous of

Illinois has been attributed to the Helicinide, but I have already,

in the first section of this memoir, discussed the affinities of this

genus and pointed out that it must be a case of convergence.

However this may be, paleontology throws very little light on

the origin and distribution of existing Helicinide, and rhe I

began this work I hoped, not only to give a full description of the

anatomy of a typical member of the family, but also, by the

comparison of the anatomy of Pacific and West Indian forms, to

discover some clue to the distribution of the group with its two

main centres in the Antillesand the Philippines. In this, as will

appear, I have been disappointed. From whatever part of the

world they may come, the anatomy of the different species and

even genera of Helicinidz is so closely similar that it is hard to

find any difference between them. It is true that I have not been

able to procure many species of Pacific Helicinide, but I have

examined fairly well preserved specimens of Or -obophana, Aphano-

conia, and Paleohelicina, and these three genera may be taken as

typical of the more widely distributed Pacific forms.

The material at my disposal was as follows :—

I. Antillean forms.

Alcadia palliata Ads. Contrivence, Walderston, Jamaica.

Alcadia hollandi Ads. Swing Hill, Walderston, Jamaica.

Lucidella aureola Fér. Bog Walk, Spanish Town, Jamaica.

Eutrochatella pulchella Gray. Bog Walk, Spanish Town,

Jamaica.

The above were kindly collected for me and preserved in

Perenyi’s fuid by Mrs. G. B. Longstaff, F.L.S.

Il. Pacific, Australian, and Indian forms.

Aphanoconia gouldiana Forbes, from Torres Straits: for

specimens of this species I am again indebted to Mrs.

Longstaff, who procured them for me from Mr. C. Hedley,

of the N.S. Wales Museum.

Aphanoconia andamanica Benson.

Aphanoconia merguiensis Pfeiffer.

Aphanoconia rogersii, sp. n.

These three species are from the British Museum and formed

part of the collection made in the Andaman Islands by Mr. G. Rogers.

MORPHOLOGY OF THE HELICINID®. 763

They were numbered respectively 16, 30,and 31. The first two I

have identified without difficulty, the third appears to be new

to science, and I will give a diagnosis of it in the latter part of

this paper. Iam indebted to Mr. KE. A. Smith for these and for

the two following species :—

Orobophana pachystoma ponsonbyi Smith. Admiralty Islands.

Paleohelicina ide Wagn. Amboina.

In describing the anatomy it will be convenient to take Alcadia

as the type, and to note such differences as may exist between it

and the other genera at the end of the description of each system

of organs.

External Characters, Mantle, Mantle-cavity,

and Muscular System.

Isenkrahe (4) has given an account of these so sufficient and

accurate that it is not necessary for me to do more than call

attention to some special features exhibited in fig. 1 (Pl. XXX.),

which is a representation of a left side view of Alcadia palliata :

the mantle has been cut through on the left side close above the

columellar muscle, the cut has been extended back to nearly the

extreme hinder end of the mantle-cavity, and the mantle has been

turned over towards the right. As compared with the Neritide,

in this and in all the other species of Helicinide that I have

examined the foot is attached to the head and body by a longer

and narrower. pedicle, the opercular lobe is relatively smaller, the

snout is narrower and longer, the columellar muscles of greater

antero-posterior length, and the whole body is longer, giving the

appearance of an increase in the coiling of the visceral mass, but

this last feature is more apparent than real, as I shall show.

A glance at the figure shows that the increased length of the

body is chiefly due to the elongation of the post-tentacular region

and the part of the body immediately following. Using

Amaudrut’s (1) phrases, we have an almost extreme case of

“allongement posttentaculaire,” followed by an “allongement

dorsal,” and many of the peculiar features of helicinid anatomy

are'to be explained by the excessive growth in length of these two

regions. The post-tentacular region lies above the anterior two-

thirds of the columellar muscle, and its posterior limit is marked

by two or three deep wrinkles of the body-wall. The body-wall

of this post-tentacular region is fairly stout and muscular, and

the epidermis is, as a rule, deeply pigmented. The colour differs

in different species. It is nearly black in Alcadia, grey shading

posteriorly into white in Hutrochatella pulchella, yellowish grey

in Lucidella aureola, a dark chocolate-brown in Paleohelicina ide,

and a bright chestnut-brown in Aphanoconia gouldiana. In the

post-tentacular region are contained the buccal bulb and the

greater part of the cesophageal pouches. In the dorsal region

following on the post-tentacular the body-wall is thin and nearly

764 PROF. G. C. BOURNE ON THE

transparent, the musculature is feeble, and the epidermis is not

pigmented, This dorsal region is relatively of considerable length ;

its concave lower border corresponds very closely in length with

the surface of insertion of the left columellar muscle; its upper

surface extends back to the pericardium. It contains nearly all

the coils of the intestine, the cesophagus, the radular sac, and the

hinder lobes of the ceesophageal pouches. Its roof forms the floor

of the hinder part of the mantle-cavity. In consequence of the

elongation of these two regions, but particularly of the dorsal

region, the mantle-cavity is continued very far back ; so far that,

measured from its most anterior to its most posterior limit, it

makes nearly a complete turn of a spiral, whereas in Verita and

Neritina it makes little more than half a turn. Broad in front

where its roof passes from the right to the left columellar muscle,

the mantle-cavity becomes narrower and narrower posteriorly and

ends ina pointed cul-de-sac below and somewhat to the right of the

lower surface of the visceral mass. Its extreme posterior limit is

not quite visible in fig. 1. With the hinder end of the mantle-

cavity the pericardium has also been carried very far back. It is

laid open in fig. 1 to show the position of the heart. It will be

seen that the single auricle seems to lie behind the ventricle, and

not in front of it as does the larger left auricle in the Neritide. In

horizontal sections, such as those depicted on Pl. XX XIII. figs. 18

and 19, this posterior position of the auricle forces itself so much

upon one’s attention that I was led to form the theory that the

single auricle of the Helicinide corresponds not to the left and

larger, but to the right and rudimentary auricle of the Neritidee ;

and in my memoir on the morphology of the latter family (2,

p. 833) I prematurely gave expression to this view, which seemed

to me the more probable because I found that in correlation with

an increase of the pallial vessels in Septaria the right auricle

became relatively larger and took an obviously larger share in

carrying blood from the pallial vessels to the ventricle. But since

then, after a careful study of the relative positions of the kidney,

the uropore, the rectum, and the heart in the Helicinide, I have

satisfied myself that this view was erroneous.

As a consequence of the dorsal elongation of the body all

these organs have been rotated through an angle of rather more

than 90°, in such wise that the pyloric end of the stomach, which

in the Neritide is directed forward and to the left, comes to lie

at the right posterior end of the body in the Helicinide, and

the swollen esophageal end of the stomach, which is posterior

and somewhat to the left in the Neritide, is directed anteriorly

and to the right in the Helicinide, forming a conspicuous

rounded prominence at the extreme end of the visceral mass

(Pl. XXX. fig. 1, St.'). To understand the nature of this rotatory

movement the reader should refer to fig. 42 of my memoir on

the Neritide. This figure represents a horizontal section of

Paranerita gagates, and shows the position of the cesophageal (St.)

and pyloric (St.') divisions of the stomach. It should be noticed

MORPHOLOGY OF THE HELICINIDA. 765

that the extremity of the visceral mass, lying to the right, is

wholly occupied by the liver and gonads. If, now, the reader will

lay a tobacco-pipe in front of him on the table, the bowl (repre-

senting the cesophageal division of the stomach) to his right, the

stem (representing the pyloric division of the stomach) to his left

front, it will occupy much the same position as does the stomach

in the figure referred to. If, while the bowl is kept pressed

against the same spot on the table, the stem is lifted up and

rotated through an angle of rather more than 90° till it points

over the observer's right shoulder, the whole pipe will have been

rotated through an angle which brings it into the position of the

stomach of the Helicinidea—the animal being supposed to be

placed foot downwards upon the table with its head turned away

from the observer. As all the organs of the left side of the

body, including the posterior end of the mantle-cavity, the peri-

cardium, the heart, the kidney, and the coils of the intestine,

have shared in this movement of the pyloric end of the stomach,

their positions have been nearly completely reversed, and the

left auricle instead of lying in front of the ventricle has come

to lie behind. The right auricle has entirely disappeared in the

Helicinide, and the rectum, having undergone some degree of

displacement in connection with the above-described movement

of rotation, is no longer enveloped by the ventricle.

The kidney has also undergone a curious and at first sight a

puzzling change of position. In the Neritide, as described in

my previous memoir (2, see Pl. XLVI. fig. 1 for its position in

Septaria, Pl. LIV. fig. 29 for its position in Paranerita gagates),

its glandular part lies to the left hand of and partly below the

rectum : posteriorly the glandular part opens into the spacious

non-glandular bladder or ureter, and the latter runs forward,

below the glandular part, to open by the uropore into the mantle-

cavity on the right of and close to the base of the ctenidium.

In this family the greater part of the kidney lies in the roof

of the mantle; it is only its posterior extremity that passes below

the rectum and invades the visceral mass, where it lies just above

the pyloric end of the stomach. The effect of the rotation of the

last-named organ in the Helicinide is that the kidney has been

carried round till it comes to le wholly in the visceral mass, on the

lower side of the latter, between the loop of the rectum which

passes round this region and the pyloric division of the stomach,

as may be seen in the series of sections (Pl. XX XIII. figs. 17 to 20)

and in the diagram (Pl. XXXIV. fig. 24), which is a reconstruction

from this series of sections. The kidney, in short, has been

turned completely round, so that its originally posterior end looks

to the left front and the uropore opens into the right hinder

corner of the mantle-cayity, the reno-pericardial canal, main-

taining its relation to the uropore, into the right posterior corner

of the pericardium. It is further to be observed that the visceral

mass, though apparently more coiled, is really less coiled in the

Helicinidee than in the Neritide. In all systematic werks stress

Proc, Zoou. Soc.—1911, No. LITI. 53

766 PROF. G. C. BOURNE ON THE

is laid on the fact that the internal partitions of the shell are

absorbed in both these families. This absorption has not pro-

ceeded quite so far in the Neritide as in the Helicinide. In

Nerita and Paranerita there is a recess in the upper right-hand

part of the shell which contains that lobe of the visceral mass which

consists wholly of liver and gonads and represents the visceral

spire of other Gastropods. This recess and the lobe of the

visceral sac corresponding to it are not found in the Helicinida :

the wall of the cesophageal division of the stomach comes very

near to the surface (Pl. XX XIII. figs. 17 & 18), and the liver and

gonads are disposed at the sides of and above the pyloric division

of the stomach. The more coiled appearance of the whole is

due to the elongation of the post-tentacular and dorsal regions,

not to the retention of a larger section of the visceral spire of the

presumed gastropod ancestor than in the Neritide.

From what precedes, it follows that most of the peculiarities of

the Helicinid organization are the result of excessive growth and

elongation of a particular region of the body, and it is an

interesting confirmation of the correctness of the above account

of the manner in which the Helicinid organization has been

derived from the Neritid that, if one makes a plasticine model

of the stomach, kidney, rectum, and intestinal coils as they occur

in Paranerita, and then rotates the stomach in the manner

described, the intestinal coils assume very nearly the position

found, with more or less variation in detail, in all Helicinids.

After this general explanation of the mutual relations of the

principal visceral organs in the Helicinide, I need only refer to

particular features in the several systems of organs which I have

to describe in detail. Before proceeding, it should be put on

record that there is not a rudiment of the ctenidium in the

Helicinide, and I cannot even find a trace of an osphradium.

The cephalic penis, characteristic of the males of the Neritide, is

also absent, and there is no externa] difference between the males

and females in any of the species that I have examined.

The Alimentary Tract.

The complex of organs formed by the buccal cavity, the pharynx,

the cesophagus with its smaller and larger glandular annexes, the

radula, the radular sac, and the odontophoral cartilages and their

muscles, can only be studied by dissection, and this is by no means

an easy task in animals so small as most species of Helicinide are.

The relations of the various organs to one another are far too

complicated to trace out in sections. The following description

applies chiefly to Alcadia palliata and A. holland, but will serve

almost equally well for any of the other species that I examined,

for all are very much alike except for the details of the radular

teeth.

The mouth is a gaping circular orifice, situated at the extremity

of the downturned snout: it is surrounded by folded muscular lips.

MORPHOLOGY OF THE HELICINIDA. 167

The buccal cavity occupies the snout, in front of the tentacles.

It is a simple funnel-shaped cavity bounded by a rather thick

muscular wall, the internal surface of which is thrown into about

19 or 20 longitudinal folds. The cavity is lined by a layer of

rather long columnar epithelial cells which secrete a thick cuticle.

Dorsally the buccal cavity is prolonged backward into a little

glandular diverticulum which lies above the median part of the

cerebral commissure. ‘The buecal cavity is separated from the

pharynx by a constriction, deepest on the dorsal side, where the

cerebral commissure lies in it. In a surface view, before dis-

turbance of the various parts, this constriction is not visible

from above, as it is covered over by the anterior salivary glands

shortly to be described, and muscle-fibres pass from the walls of

these glands to the walls of the buccal cavity and of the snout.

Consequently the cerebral commissure seems to be embedded in

the buccal mass.

The passage from the buccal cavity to the pharynx is narrow.

The pharynx is a relatively spacious sac, of which the cavity

is continued posteriorly into the esophagus above and into the

radular sac below. Beneath and at the sides of the anterior

end of the radular sac lie the odontophoral cartilages, the

anterior ends of which project forward into and occupy the

greater part of the lower moiety of the pharyngeal cavity. It

will readily be understood that, in consequence of the projec-

tion of the anterior ends of the odontophoral cartilages into

the pharyngeal cavity, the latter extends round them both at

the sides and below. Below the cartilages the pharyngeal

extension forms a broad flattened diverticulum, reaching back

nearly to the posterior ends of the anterior cartilages, as far

as the point marked a in fig. 3 (Pl. XXX.). Laterally, the line

of attachment of the pharyngeal wall to the anterior odonto-

phoral cartilages is roughly indicated in the same figure by the

curved line running upward and forward from the point « towards

the opening of the esophagus. It results from this arrangement

that in an oblique section, such as is represented in fig. 4

(Pl. XXX.), the pharynx appears to give off two posterior diver-

ticula, lying outside the anterior ends of the odontophoral

cartilages. The inner walls of these diverticula are thin and

composed of a single layer of cubical epithelial cells: they are

continued round the anterior and upper edges of the cartilages

into the lining membrane of the radular sac. The outer wall of

each diverticulum is strengthened by a thin plate of cartilage,

too small and transparent to be recognized in dissection, but

readily recognizable in sections. These lateral pharyngeal carti-

lages serve for the attachment of muscles, one set of which run

forward to be inserted on the walls of the snout, the other set

run backward and are inserted on the odontophoral cartilages ;

the former are protractors, the latter retractors, of the walls of the

pharynx.

A portion of the epithelial lining of the outer wall of each

53*

768 PROF. G. C. BOURNE ON THE

diverticulum is composed of very long attenuated epithelial cells,

among which are long club-shaped glandular cells. This glandular

strip may be traced upward and forward to the thickened lip of

the esophageal opening, where it forms a prominent ridge

which passes above into the anterior pair of salivary glands, to

be described shortly.

As is shown in P]. XXX. fig. 2 and in the section drawn in

fig. 4, the radular sac opens into the pharynx by a widely gaping

aperture situated in the trough-shaped depression between the

anterior ends of the anterior odontophoral cartilages. Posteriorly

the radular sac passes between the posterior odontophoral cartilages

and runs at first to the right of and below the cesophagus, but

soon mounts upwards and to the left, the two organs, oesophagus

and radular sac, being twisted round one another as shown in

fig. 2. The radular sac is short in Alcadia palliata and is of no

great length in any of the species that Ihave studied. The greater

or less length of the radular sac appears to be an individual rather

than a specific character. The characters of the radular teeth will

be dealt with in a separate section.

The odontophoral cartilages were described in some detail by

Isenkrahe for Helicina titanica, and I have but little to add to his

account, A ventral view of these structures in Alcadia palliata

is given in Pl. XXXI. fig. 5, and a sketch of a dorsal view of the

same structures in Hutrochatella pulchella in fig. 6: both figures are

drawn to the same scale. As may be seen from the specimens

figured, the odontophoral cartilages exhibit specific differences in

relative size and proportion, but these differences are of too slight

and elusive a character to be expressed in a description, and scarcely

important enough to make it worth while to give aseparate figure

for each:species examined. The essential structure is the same

in all. There are two pairs of cartilages, an anterior and a

posterior. Each member of the anterior pair is a plate having

the form of a more or less elongated isosceles triangle; the

margins of the plate ave thickened and rounded, the central

portion remains thin. The plate is bent in such a way that its

lower margin is bent inward posteriorly and its upper margin

outward. The posterior margin or base of the triangle, forming

the articular surface for the posterior cartilage, runs obliquely

from above downwards and inwards. ‘The lower margins of the

two cartilages are connected by a tough fibrous band, and their

thickened edges serve for the attachment of the intrinsic and

extrinsic odontophoral muscles. The posterior cartilages are

short conical masses; their apices directed backwards; their

ventral surfaces convex and their dorsal surfaces more or less

concave. Their broad anterior endsare shaped to correspond with

the articular surfaces of the anterior cartilages, and the two are

firmly held together by muscular fibres, whose arrangement is

indicated in fig. 5. It follows from the above description that

the odontophoral cartilages form the sloping sides of a V-shaped

trough, the concavity of which looks upwards and supports the

MORPHOLOGY OF THE HELICINID®. 769

anterior part of the broad radular ribbon. The median and

admedian radular teeth lie in the floor of this trough, the great

lateral teeth he in the angles between its floor and sides, and

the uncini form curved rows running upwards and backwards

along its sloping sides. The radular ribbon is attached by strong

muscular bands to the cartilages. These muscles run obliquely

forwards from the radula to be attached to the anterior cartilage

of either side, and obliquely backwards to be attached to the

posterior cartilages, these two sets of muscles causing the ribbon

to slide forward and backward over the smooth surfaces of the

cartilages.

The relation of the cesophagus to the radular sac and odonto-

phore is shown in fig. 3, which is a drawing of a dissection of

these structures in Alcadia palliata. The left cesophageal pouch

and the left side of the cesophagus have been cut away, and the

roof of the cesophagus has been lifted back to the right side to

show the entrance to the right cesophageal pouch and other

structures. In the angle of the deep fold between the cesophagus

and radular sac are seen the buccal ganglia (g.bwe.) lying just above

the middle of the anterior odontophoral cartilages. ‘The opening

of the cesophagus into the pharynx, situated just in front of and

above the buccal ganglia, is irregularly funnel-shaped, with thick-

ened and folded lips projecting forward into the pharyngeal

cavity. laterally, these lips are deeply grooved, and on either

side the groove is continued backward and downward into the

lateral pharyngeal diverticulum described above, and upward and

somewhat forward into the anterior salivary gland of its own

side. These anterior salivary glands are formed by a pair of pocket-

like forward projections of the cesophagus, which in their natural

position lie side by side and form a pair of pouches lying above

the cerebral commissure. Between them is an anterior cecal

diverticulum continuous behind with the median dorsal groove of

the anterior part of the csophagus. When separated by an

incision in the mid-dorsal line and turned outwards, the anterior

salivary glands present the appearance shown in fig. 2. Inter-

nally their walls are raised into a number of thick glandular

ridges: the outermost of these ridges is specially thick and is

continued downward, in the groove passing to the side of the

cesophageal orifice, into the glandular ridge on the outer wall of

the pharyngeal diverticulum, as has been described above.

The anterior section of the cesophagus, lying above the

odontophore, is fairly wide. Internally, its walls present a

number of longitudinal glandular ridges, and in the mid-dorsal

line there is a deep groove bounded internally by prominent

ridges; posteriorly this groove shallows and eventually dies out.

On either side of this anterior section of the esophagus is a

gaping oval orifice (figs. 2 & 3, [email protected].) leading into the large

cesophageal pouches, or, as some would call them, the posterior

salivary glands. The last-named structures are capacious irregu-

larly lobulated sacs with large cavities. Their inner walls are

770 PROF. G. C. BOURNE ON THE

lined throughout by a glandular epithelium, consisting, as far as

T was able to Gbeeenne! almost wholly of very long goblet-shaped

secretory cells containing zymogen granules, with very few attenu-

ated supporting cells lying between. The histological characters

of the epithelium were not, however, very well preserved in any

of my specimens. The two pouches are closely pressed against

the sides of the cesophagus, and in the species in which they are

longest follow the turns of the latter. Hence, as the esophagus

makes a turn towards the left before it passes down through the

loop of the rectum, the right esophageal pouch generally passes

over to the left side and above the cesophagus, the left pouch

passing to the right and below the esophagus and radular sac.

The cesophageal pouches are relatively short in Alcadia palliata

and of approximately equal width thr oughout, but they are much

longer and diminish in diameter towards their posteri lor extre-

mities in A. hollandi. They are particularly long and of a deep

chocolate colour in Paleohelicina ide ; in most species they are

white in spirit-specimens.

Behind the openings of the cesophageal pouches the cesophagus

narrows somewhat abruptly in diameter; the glandular internal

longitudinal ridges disappear, and are replaced by nine or ten

longitudinal ridges formed by columnar ciliated epithelial cells,

and these may be traced throughout its course to the stomach.

This course is a long one, for the stomach lies aslant on the

lower side of the visceral mass, below the greater part of the

lobes of the liver and below the coils of’ the intestine. As seen

from above and behind, it is a large pyriform sac, the narrower

end lying just above the posterior corner of the pericardium ;

the broader end forming on the right side of the visceral mass a

large rounded prominence which fits into the concavity on the

ventral side of the right columellar muscle. The cesophagus

enters the stomach on the upper side of its broader end, and its

course in the several species examined will be best understood by

reference to Pl. XXXII. figs. 10 to 15. Passing to the left as it

enters the visceral mass, the cesophagus lies above the first coils

of the rectum, then passes below the recurrent coil of the rectum,

and so arriving at the dorsal surface of the stomach runs along

the latter as a flattened tube and opens, as stated above, into

its broader end, dilating considerably just at its point of entrance.

This dilatation, which might almost be described as a diver-

ticulum of the stomach itself, receives right and left the wide

ducts of the liver (Pl. XXXT. fig. 7,and Pl. XXXIT. fig. 16, li.d.).

The internal structure of the eae is very complicated, It

attracted the attention of Isenkrahe, who gave a fairly accurate

description of it. Fig. 7 is a representation of a dissection of the

stomach made from behind and below the visceral mass. The

cesophagus is shown at &., and its entrance into the stomach is

indicated by the arrow. The entrance of the left liver-duct is

shown at li.d.; that of the right liver-duct lies on the far side

ef the prominent curved ridge guarding the entrance to the

MORPHOLOGY OF THE HELICINIDZ. 771

cesophagus. Both the esophageal aperture and those of the liver-

ducts are surrounded by complicated epithelial ridges, which, as

shown in the drawing, converge towards and pass into a deep

groove running along the dorsal side of the narrower pyloric

moiety of the stomach. The edges of this groove are bounded by

two preminent folds; that on the right side (the left in the

drawing) being continued towards the esophageal opening as a

projecting ridge, ending at the side of the aperture in a very

prominent crescentic projection covered by a thick iridescent

cuticle.

The internal surface of the stomach is lined by a mixed glan-

dular and ciliated columnar epithelium, the characters of which

are shown in P]. XX XI. fig.8; the glandular elements predominate

in the esophageal, the ciliated elements in the pyloric, moiety of

the stomach. ‘The epithelial cells are lower in the furrows, taller

and more slender in the ridges, these latter structures being

formed entirely by thickenings of the epithelium, and not by

foldings of the wall of the stomach. The crescentic projection to

the right of the esophageal opening is formed by a local modifi-

eation of the epithelium, the cells of which are here extraordi-

narily long, with nuclei placed about the middle of their length,

and with apparently homogeneous transparent cytoplasmic contents

(Pl. XXXI. fig. 9); they are all of one kind, without any admix-

ture of glandular cells, and, so far as one can judge, they are not

themselves glandular. The free ends of these cells are covered bya

very thick and tough cuticular coat, which stains deeply in hema-

toxylin, brazilin, and other ordinary dyes. The whole structure

corresponds to the “ fiéche tricuspide,” of which the characters

have been thoroughly described for Lamellibranch stomachs,

and which has been noted as occurring in several Gastropod

‘stomachs. There is no definite crystalline style in Helicinide,

but in several specimens that I dissected I found the cavity of

the stomach filled by a semitransparent gelatinous mass, which

appeared to be similar in origin and composition to a crystalline

style. In several specimens I found that the intestinal end of

the groove of the pyloric moiety of the stomach was occupied by

closely compacted fecal pellets or rods, while the cavity below

was filled either by the gelatinous mass above mentioned or by 4

loose mass of semi-digested food. It may be inferred from this

that digestion is effected in the general cavity of the stomach,

and that the indigestible materials of the food are collected into

the dorsal groove and passed into the intestine.

Morphologically, the stomach of the Helicinide closely resembles

that of the Neritide, and further bears a resemblance to the

stomachs of the Fissurellide# and Scissurellide#, which, as in this

case, have a groove leading from the hepatic ducts towards the

intestinal end of the stomach, but no spiral cecum.

The small intestine is a comparatively narrow tube, which after

leaving the pyloric end of the stomach runs back for a short

distance over the dorsal surface of the latter, parallel with the

172 PROF, G. C. BOURNE ON THE

cesophagus, and then turning upwards and forwards describes one

or more convolutions before it passes into the large intestine.

The small intestine can always be distinguished by its white

colour and narrow diameter; it varies considerably in length in

different species, being longest in Alcadia palliata and Orobophana

ponsonbyi, shortest in Hutrochatella pulchella. Internally it is

lined by a columnar ciliated epithelium containing a few gland-

cells, and its internal surface is increased by a well-marked

internal ridge or typhlosole formed by long ciliated cells. The

small intestine passes abruptly into the large intestine, the latter

being of much larger diameter and having pigmented walls. In

all species of Alcadia and Helicina examined, the large intestine

runs forward and to the left below the cesophagus, then bends

abruptly back, forms a wide circular loop which passes above the

cesophagus, in front of the anterior end of the stomach and along

the inner wall of the anterior end of the pericardial cavity: it then

turns downwards and to the right, passes right round the lower side

of the stomach, and mounting upwards again behind it runs in the

right side of the roof of the mantle-cavity to open by the anus,

opposite the right tentacle. In the first part of its course, 7. e. in

the short length between the small intestine and the recurrent

circular loop, the large intestine is lined by very evenly disposed

columnar ciliated cells, among which I could not detect any

gland-cells, and this part of the intestine is not, asa rule, full

of fecal matter. This section of the intestine may be described

as the large intestine proper, to distinguish it from the rectum,

into which, however, it passes without any obvious line of

demarcation. ‘The rectum is always full of feecal débris, and its

epithelium consists of (1) columnar ciliated cells; (2) goblet-

shaped gland-cells filled with coarse granules which stain deeply

in hematoxylin and are therefore probably mucinogenous ;

(3) smaller gland-cells containing small yellow granules. There

is no typhlosole either in the large intestine or the rectum.

Noticing that the coils of the gut differed in the different

species, I have been at some pains to work out this character in

detail, and figs. 10 to 15 (Pl. XXXII.) show the coils charac-

teristic of six different species. An examination of the figures

will give a better idea of the differences than any description.

It will be noticed that there are three main types. In Alcadia

palliata, A. hollandi, Orobophana ponsonbyi, and Paleohelicina

the intestinal coils, though differing in detail, are alike in this

respect, that the small intestine runs back more or less parallel

to the cesophagus over the dorsal surface of the stomach, and

the large intestine makes a bend to the right below the cesophagus

and then, turning back on itself, makes a second bend to the

right above the esophagus. The second type of arrangement

is shown in Lucidella awreola (fig. 14). In this species the

esophagus is not pressed against the dorsal surface of the

stomach, but runs obliquely down to it; the small intestine

passes forward from the pyloric end of the stomach, crosses over

MORPHOLOGY OF THE HELICINIDA. 773

the cesophagus, and curves round it till it nearly touches the

pyloric end of the stomach again; here it passes into the large

intestine, which turns sharply back, passes round the cesophagus

again, and coming to the surface sweeps round to the left in front

of the anterior corner of the pericardium to form the descending

loop of the rectum. This type is easily derived from the first by

the shortening of both the small and the large intestine, in

consequence of which the former is hooked round the cesophagus

and one of the bends characteristic of the first type is suppressed.

The third type, seen in Hutrochatella pulchella (fig. 15), differs

considerably from the other two. The small intestine is even

shorter than in Zuecidella, and the loop formed by the large

intestine and the first section of the rectum lies wholly on the

dorsal side of the esophagus; this condition is clearly due to

the gut being much shorter than in the other types, and it

appears to be quite a constant feature in Hutrochatella. The

intestinal coils of Aphanoconia andamanica ave singularly like

those of Hutrochatella.

I may appear to be giving an undue amount of attention

to characters of no obvious morphological or physiological

importance, but it is just because they may be claimed to be

of importance in the economy of the species that I have spent a

considerable amount of time in working out these details. Hach

species seems to have a characteristic arrangement of the coils

of the intestine, and the arrangement is remarkably constant in

individuals of the same species, allowance being made for

displacements due to the greater or less state of contraction of

the specimens. Closely allied species, such as Alcadia palliata

and A. hollandi, have a very similar arrangement, yet sufficiently

different to allow one to recognize them at a glance after

obtaining some familiarity with their anatomy. Lucidella and

Eutrochatella, both separated from Aleadia and from one another

by distinctive characters of shell, operculum, and radula, differ

in a nearly corresponding degree in the coils of the intestine.

Orobophana and Palcohelicina are Pacific forms which must

have been derived from American Helicine, the latter being

closely related to Alcadia, and they resemble the last named in

the coils of the intestine. To this extent it may be claimed that

two, three, or more characters vary together in these genera ; but

Aphanoconia presents a difficulty, for this genus is far removed

from Hutrochatella in shell and radular characters, and is in these

respects closely related to Paleohelicina, yet its intestine is as

nearly as may be that of an Hutrochatella. As the two genera

cannot possibly stand in close genetic relationship to one another,

the similarity in the pattern of the intestinal coils must be due

to parallelism, similar causes producing similar deviations from

type in the two organisms. It has been shown that the

differences in pattern are attributable to differences in the

length of the large and small intestines, and this is probably

connected with different forms of food. As we are ignorant of

774 PROF. G. C. BOURNE ON THE

the habits of the various species of Helicinide, and do not even

know for certain whether each or any species is restricted to a

particular kind of food, it would be rash to speculate on this

question, but such evidence as I have collected does seem to show

that such apparently trivial characters as the coils of the intestine

are of some physiological importance, and are therefore subject to

the action of natural selection.

The Celom.

The ewlom is represented by the pericardial cavity, which,

although it is of much smaller extent and less complicated than in

the Neritide, is nevertheless a cavity of relatively considerable size,

as may be seen by an inspection of figs. 17 to 20 (PI. XX XIIT.).

As is shown in fig. 1 (Pl. XXX.) it comes close to the surface of

the left side of the visceral mass, and extends forwards nearly

as far as the posterior end of the left columellar muscle and

backwards round the lower side of the visceral mass as far as the

end of the mantle-cavity. It is bounded externally by the very

thin body-wall, posteriorly by the inner wall of the mantle-

cavity, internally by the kidney (figs. 17 to 20). At about the

middle of its length it is a cavity of considerable depth, extending

some way into the visceral mass below the pyloric division of

the stomach. The reno-pericardial canal, which will be described

in connection with the kidney, opens into its right posterior wall,

at some little distance from its hindermost end (fig. 19); other-

wise it is a closed sac containing the auricle and ventricle of the

heart, and does not require further description.

The Hemocele, Circulatory and Respiratory Systems.

The blood-vascular system, as in all Molluscs, consists partly

of large lacunar spaces, which collectively are known as hemocele,

and partly of vessels with definite walls. The hemoccelic spaces

surround the viscera, and there is a specially large lacunar space

below the buccal bulb, in which lie the pleuro-pedal ganglia.

In the visceral mass and in the dorsal region of the body the

hemoceele is largely filled up by the peculiar form of connective

tissue which I have previously described (2, p. 861) as metabolic

tissue. In the Helicinide the tissue is of precisely the same

nature as in Neritidz, and it is not necessary to describe it again.

It evidently consists in large part of reserve tissue, for it is most

abundant in immature specimens in which the gonads and

gonaducts are but slightly developed, and is much less abundant

in sexually mature specimens. This metabolic tissue is specially

concentrated round the larger blood-vessels.

It is not possible to trace the course of the blood-vessels by

dissection of spirit-preserved specimens and only the larger vessels

can be traced in sections. The following account of the

circulation embodies as much information as I have been able to

MORPHOLOGY OF THE HELICINIDA. 775

obtain by reconstruction of sections. The ventricle is continued

forward into a short and wide aorta, which immediately after

passing through the pericardial wall—which it does at about the

level of the hinder end of the left columellar muscle—divides

into three principal branches. The one passes to the right

towards the stomach, and, entering the visceral mass, divides

into a number of branches which are distributed to the stomach,

the intestine, the right lobe of the liver, the gonad, and the

hypobranchial gland and gonaducts. The left branch runs

forward for a short distance and then turns downward into the

visceral mass and chiefly supplies the left lobe of the liver.

A branch is directed towards the posterior part of the left

columellar muscle. The third vessel is an almost direct forward

continuation of the aorta and runs up in the dorsal region of the

body towards the esophagus; it passes above this organ and runs

over the surface of the radular sac, to which organ it becomes

firmly attached at about the level of the hind end of the

pharyngeal bulb. Here it passes into a number of lacunar

passages, supplying the pharyngeal bulb and the cesophageal

pouches, and eventually makes communication with the large

blood-space surrounding the nerve-centres of the head. This

space in turn communicates freely with the lacunz surrounding

the pedal nerve-chords.

In this labyrinth of blood-channels I have not been able to

recognize those by which the blood is collected and brought back

from the various organs to the organ of respiration, the mantle.

The principal hemoceelic spaces or blood-sinuses are the

following: (1) a pedal sinus, surrounding the pedal nerve-cords ;

(2) a subcesophageal sinus, underlying the buccal mass and

esophagus ; this is continued back into (3) a cireum-intestinal

sinus, in which lie the coils of the intestine. and the posterior

part of the radular sac; (4) a peri-gastric sinus surrounding the

stomach ; (5) a recto-genital sinus, running the whole length of

the rectum and gonaduct. The last named is evidently the

pulmonary vein of Isenkrahe (“lings des Darmes zieht sich die

Lungenvene hin”), but it does not carry back blood from the

mantle to the auricle. On the contrary, it is easy to see that

blood passes from it to the numerous fine blood-vessels or

rather blood-spaces, for they have no definite walls, in the roof of

the pulmonary chamber. The efferent pallial vein that collects

blood from the mantle and returns it to the heart is on the opposite

or left side of the mantle-cavity. It isa direct continuation of the

auricle and can easily be traced forward in the left corner of

the mantle-cavity, running along the upper border of the left

columellar muscle (Pl. XXXII. fig. 16, v.pal.) nearly as far as

the thick muscular anterior border of the mantle. It receives

numerous vessels from the mantle, especially in the anterior

part of its course. The blood from the intestinal and perigastric

sinuses does not pass straight to the mantle, but is collected into a

large sinus provided with definite walls (Pls. XX XIII. & XXXIV,

776 PROF, G, C. BOURNE ON THE

figs. 19 & 21, v.ven.), whence it passes by an afferent renal vessel

to the glandular portion of the kidney. So far as I can ascertain,

the blood is returned from the kidney to numerous small vessels

running in the floor of the posterior half of the mantle-cavity,

and is conveyed from these to the auricle by a distinct vein

(Pl. XXXIITI. fig. 20, v.post.) which opens into the hinder part of

the auricle, and receives in addition blood from the roof of the

extreme hind end of the roof of the mantle-cavity. There are

thus two distinct vessels opening into the auricle, the foremost of

which brings back blood from the roof of the greater part of the

pulmonary chamber ; the hindmost brings blood that has passed

through the kidneys, then through the vessels on the floor of the

pulmonary chamber, and in addition a small quantity of blood

from the roof of the extreme hind end of the pulmonary chamber.

The Hxcretory Organs.

The topographical relations of the kidney, as compared with

that of the Neritide, have already been explained (p. 765). Re-

garded in detail, the kidney consists of a thiek-walled glandular

portion and a thin-walled non-glandular portion which serves as a

bladder and urinary duct. The glandular portion is a large and,

roughly speaking, quadrangular sac lying in the lower part of the

visceral mass, below the stomach but above and somewhat to the

left of the lower loop of the rectum. Its posterior and left wall

fits closely against the pericardium and forms the inner boundary

of the latter. The two ends of the sac are produced into large

pockets or recesses, which partly extend round and embrace the

walls of the pericardium (Pl. XX XIII. figs. 18 & 19), and partly

extend upwards round the sides of the pyloric division of the

stomach (Pls. XX XJIL. & XXXIV. figs. 17 & 24). The cavity of

the sac is spacious, and only partially subdivided by folds project-

ing inwards from the wall on the pericardial side; the opposite

wall is not folded. The renal blood-vessels run in these folds.

The whole cavity, including the folds, is lined by a uniform

glandular epithelium consisting of large irregularly shaped cells,

of varying length, their free ends rounded or club-shaped and

often projecting far into the lumen of the sac.

The characters of these cells are shown for Alcadia in fig. 22

and for Lucidella in fig. 23 (Pl. XXXIV.). In all the other

species that I have examined the kidney-epithelinm resembles

that of Lucidella ; it is only in Alcadia that the cells are as long,

irregular, and ameeboid-looking as those drawn in fig. 22. In

both cases the cytoplasm is clear and distinctly and coarsely

vacuolated ; the nucleus spherical, vesicular, with a few granules

of chromatin. The ureter or non-glandular part of the kidney

arises from the upper corner of the left-hand recess of the

glandular sac. Its walls are composed throughout of a non-

ciliated, very low, cubical epithelium, the cells of which are so

much flattened that they might almost be called a pavement-

MORPHOLOGY OF THE HELICINIDA. Ce

epithelium. The ureter is a widish tube which after leaving the

glandular sac turns back to run round the hinder wall of the

pericardium, interposing itself between it and the lower surface

of the visceral mass. After passing from the left to the right

side if mounts upwards again, passes under the reno-pericardial

canal, and opens into the right-hand side of the mantle-cavity

by a thick-lipped slit-like uropore. Asis shown in fig. 23, the

mantle-epithelium is invaginated at the lips of the uropore, and

this invaginated portion is ciliated, but there is no uropore-sac

such as I have described in the Neritide.

The reno-pericardial canal (figs. 19 & 22) opens out of the

lower part of the right-hand recess of the glandular part of the

kidney, and runs straight into the right side of the pericardium

opposite the middle of the expanded base of the auricle. The

canal is short, straight, and narrow, lined by a cubical ciliated

epithelium, the component cells of which are small and bear no

resemblance to the very large ciliated cells lining the long twisted

reno-pericardial canal of the Neritide. The cilia are fine and

directed towards the kidney. A thickening of the epithelium at

the pericardial opening of the canal is suggestive of the presence

of a pericardial funnel. The structure and relations of the kidney

and the reno-pericardial duct are remarkably uniform in all the

species of Helicinide that I have examined.

The Generative Organs.

Thiele (10) has shown that the female ducts are monaulic in

Hydrocena cattaroensis, but diaulic in Helicina kubaryi. Before

my memoir on the Neritide was published I had discovered the

diaulic ducts in Aleadia and Hutrochatella, but, as I omitted to

make mention of them in that place, I must yield priority to

Thiele, whose diagrammatic figure (loc. cit. text-fig. 2) gives a

correct representation of the general relations of the various sub-

divisions of the ducts. But it is almost impossible to construct a

life-hke picture of such complicated organs from a study of

sections, and as I have dissected out the gonaducts, both male and

female, in a number of species and have checked my observations

by the study of sections, I may be pardoned for again taking up

the subject and entering into it at some length. From the

analogy of the Neritidz, in which family the gonaducts exhibit a

considerable range of variation, I expected to discover equally

great differences in these organs in the various genera of Heli-

cinide, but have been disappointed. There are differences, it is

true, but they are slight and do not throw much light upon the

systematic affinities of the various genera studied.

The gonads in all Helicinide lie above and to the right side of

the liver. The ovaries are follicular, and the follicles open into a

large thin-walled chamber which in Alcadia and Hutrochatella is

situated on the right side of the visceral mass, just behind the

posterior end of the right columellar muscle and in front of the

778 PROF. G. C. BOURNE ON THE

muscular partition separating the visceral cavity from the complex

glandular mass formed by the hypobranchial gland and gonaducts

(Pl. XXXVI. fig. 32). The ova appear to go through their matu-

ration-phases in this chamber, as no ripe ova are to be seen in the

follicles. In Alcadia and Futrochatella this ovarian chamber is

sac-shaped and on the right side of the body, but in Pacific and

Oriental species, such as Paleohelicina ide, Orobophana ponsonbyt,

and Aphanoconia gouldiana, the ovarian chamber is produced into

a wide tube which stretches transversely across the body and

receives the products of the follicles of a left ovarian lobe, the latter

being a distinct triangular lobe, projecting from the surface of the

visceral mass and packed close under the left columellar muscle.

This lobe is possibly characteristic of Pacific and Oriental species:

it is absent in Alcadia, and scarcely represented in Luérochatella.

My specimens of Zucidella were all male, and therefore I cannot

say whether it occurs or not in this West Indian genus. The

female gonaducts of Alcadia holiandi, which are to all intents

and purposes identical with those of 4. palliata, are depicted in

Pl. XX XV. fig. 25; and figs.30 to 35 (Pls. XXXV. & XXXVI)

represent selected sections from a series passing horizontally

through the genital complex of a female of the same species,

fig. 30 being the uppermost and fig. 35 the lowest of the series.

Dealing first with the macroscopical characters, the following

organs or parts can be distinguished, and as their shape and

relative positions are clearly indicated in fig. 25a detailed de-

scription will be superfluous. (1) The oviduct od. is a very narrow

duct leading from the ovarian chamber to (2) the dilated or V-

shaped portion of the oviduct, in which a descending limb (od.')

and an ascending limb (od.*) can be recognized. In <dAlcadia

the ascending limb is as long and of approximately the same

diameter as the descending limb: in its lowest third it receives

the short and narrow duct of a globular receptaculum seminis.

(3) The ootype, oo¢., is a long, more or less dilated glandular duct

running parallel with the rectum in the right hand of the roof of

the mantle-cavity, and opening into the latter on the right side by

a relatively narrow aperture ona papilla which lies in a sort of

shallow cloaca formed by the expanded lips of the rectal opening.

Into the posterior end of the ootype open: the ascending limb of

the V-shaped part of the oviduct ; the caecum of the ootype, a sac-

shaped structure of considerable relative size, which lies parallel

to and to the inner side of the V-shaped duct ; thirdly, the short

and rather narrow duct of (4) the vagina, vag. The last named

is a slender duct with thin walls, opening into the mantle-cavity

close to the aperture of the hypobranchial gland. It is continued

posteriorly into a sac, which runs back on the outside of the

V-shaped duct and ends blindly (this sac has been displaced in

fig. 25 to show it more clearly, in its natural position it would be

concealed by the V-shaped duct). It may be called the vaginal

sac. Comparing these ducts with those of the Neritide, parti-

cularly with Paranerita (2, fig. 60), it is clear that the ootype is

MORPHOLOGY OF THE HELICINID. 779

homologous in the two forms, as is also the vagina. The vaginal

sac of Alcadia corresponds to the spermatophore sac of Paranerita,

but in the former genus the duct connecting the vagina with the

ootype has been shortened to such an extent that it is merely a

passage between the two. The V-shaped duct of Alcadia is only

a modification of the oviduct and has no exact homologue in

Paranerita; the position of the receptaculum seminis is also

somewhat different. There is no trace of an oviduco-celomic

funnel in any Helicinid, nor is there any representative of the

crystal-sac. On the other hand, the cecum of the ootype is not

represented in the Neritide, for it would be straining homology

too far to suggest that it is the equivalent of the lower dilated

part of the ductus enigmaticus. But if the morphological com-

parison between the female gonaducts of the Neritide and

Helicinide is fairly obvious, a physiological comparison is by no

means so clear. In the specimen of 4. fellandi whose ducts are

drawn in fig. 25 the hinder moiety of the ootype was filled and

greatly distended bya large mass of spermatozoa held together by

a coagulable substance, and, as shown in the figure, a string of the

same mass of spermatozoa and coagulum extended into and filled

the cecum of the ootype. The vaginal sac was empty, but the

receptaculum was full of spermatozoa. This suggests that the

“‘ootype ” is the copulatory canal or functional vagina, and that

the “‘vagina” may serve for the passage of the ova into the

mantle-cavity. But the probability of such a conclusion is

lessened by the fact that in Aphanoconia merguiensis and 4d.

gouldiana I found the vagina and vaginal sac full of spermatozoa.

[have not found ova either in the oviduct or in the ootype of any

of my specimens; and apparently in the Helicinide fertilization is

not effected by means of spermatophores, for I have found no

trace of such structures. The mass of spermatozoa and coagulum

in Alcadia hollandi cannot be called a spermatophore. The

evidence as to the function of the several parts being slender, and

what there is conflicting, I offer no definite theory on the matter,

but may add that the “ vagina” is evidently a distensible duct, as

it has thin walls, with a very feeble coat of circular muscle-fibres,

but with a number of muscular slips passing from its walls to be

attached to adjacent organs. On the other hand, the ootype has

a thick muscular coat, especially in its hinder moiety, and the

contraction of this muscular coat would expel any contained

material through the external aperture.

A comparison of the gonaducts of the several genera shows

certain differences in detail. The female gonaducts of Hutro-

chatella, a West Indian genus, are shown in Pl. XXXYV. fig. 26.

Their arrangement 1s clearly very similar to that of Aleadia. The

ascending limb of the V-shaped duct is shorter in Lutrochatella

and has more the appearance of a direct posterior continuation of

the ootype. The receptaculum seminis is small and ovoid, but in

the same position as in Alcadia. The vagina is short ; the vaginal

sac of moderate length. The cecum of the ootype is a large

780 PROF. G. C. BOURNE ON THE

flattened sac and extends some way behind the posterior end of

the V-shaped duct.

Of the Pacific and Oriental genera, Aphanoconia (Pl. XXXYV.

fig. 27) most nearly resembles Alcadia and Hutrochatella. 'The

descending limb of the V-shaped duct is large ; the ascending limb

short and scarcely differentiated from the hinder end of the

ootype; it bears a relatively large globular receptaculum seminis.

The vagina is fairly long, and opens by a very short transverse

duct rather high up into the ootype. The vaginal sac is short.

There is no cecum to the ootype. The female ducts are very

similar in all the four species of Aphanoconia that I have examined,

differing chiefly in the relative length of the vaginal sac and

the position of the vagino-ootypal connection.

On the other hand, Palwohelicina (fig. 28) and Orobophana

(fig. 29), while resembling one another, differ in some important

respects from the West Indian genera and from dAphanoconia.

In Paleohelicina ide the ascending limb of the V-shaped duct is

wide and scarcely differentiated from the hinder end of the ootype.

There is no receptaculum seminis, but the ootypal cecum is large,

bilobed at its extremity, lined by an epithelium of peculiar cha-

racter, and filled with spermatozoa. It evidently functions as a

receptaculum seminis. The vagina and vaginal sac are normal.

In Orobophana ponsonbyi the descending limb of the V-shaped

tube is unusually long ; the ascending limb very narrow and short,

and does not bear a receptaculum seminis. The cecum of the

ootype is large, bilobed, full of spermatozoa, and in every respect

similar to that of Palwohelicina. The vagina is long; the vaginal

sac leaves it about halfway between its external aperture and its

connection with the ootype.

I am unable to give a description of the female organs of

Lucidella aureola, as all my specimens were males.

Summing up the above facts, we see that as regards the

structure of the gonaducts there are two types in the genera dealt

with. In the first type there is a receptaculum seminis on the

ascending limb of the V-shaped duct. This type is divisible into

two sub-types: the one, found in Alcadia and Hutrochatella, is

characterized by the large cacal appendage of the ootype; the

other, found in Aphanoconia, has no such cecum. In the second

type, represented in Palwohelicina and Orobophana, there is no

receptaculum seminis on the ascending limb of the V-shaped duct,

but this organ is replaced functionally by the modified bilobed

cecum of the ootype. According to Wagner (12) Palcoheli-

cina stands nearest to Helicina sensu restricto, and therefore

nearer to Alcadia than other Oriental and Pacific forms, but in

the structure of the female gonaducts, dphanoconia stands nearest

to Alcadia, but Paleohelicina with Orobophana stand somewhat

apart.

"The gonaducts of the Helicinide evidently undergo great histo-

logical changes at the onset of sexual maturity. In sections of

immature females of Alcadia and Hutrochatella the ootype is a

MORPHOLOGY OF THE HELICINID®, 781

relatively narrow tube lined by a columnar epithelium, in which

glandular elements are hardly recognizable, and there is a similar

lack of differentiation of glandular cells in the V-shaped duct. On

the approach of sexual maturity the epithelium of the ootype is

enormously thickened, is rich in glandular elements, and is thrown

into complicated folds; at the same time, histological changes

occur in the V-shaped duct. In the breeding-season, when the

ootype is distended as shown in fig. 25 (PI. XX XV.), the epithelial

folds of the ootype disappear, the gland-cells have discharged their

contents, and the epithelium appears shrunken and thinner than

before. In consequence of these changes, it is difficult to give

a consistent account of the histological characters of the ducts :

what follows is based on a series of sections of a mature

specimen of Alcadia hollandi, in which the tissues, thanks to

Mrs. Longstaff’s care, are admirably preserved.

In the distal half of the ootype, that is in the portion marked a

in fig. 25, the epithelium is moderately long and of the mixed

glandular and ciliated kind. The ciliated cells are elongate

columnar, not much attenuated at their bases, their cytoplasm

elear and very finely granular, their nuclei rather large, oval,

situated rather to the basal side of the middle of the length of

the cell. The gland-cells are of nearly the same shape, but of

rather greater diameter than the ciliated cells, their nuclei slightly

larger, situated nearer the bases of the cells, and the cell-body

filled with rather small highly refracting spherules of a greenish-

yellow colour in preparations stained with hematoxylin and eosin.

The hinder moiety of the ootype—namely, that portion filled with

the mass of spermatozoa in fig. 25—shows somewhat different

characters. The supporting epithelial cells are, as before, columnar

and finely granular, but of greater length: they appear to have

lost their cilia over the greater part of the inner wall of the

ootype and to end distally in rounded and somewhat vacuolated ex-

tremities, but as I can find cilia in patches this appearance may be

due to maceration. ‘The gland-cells in this region (P]. XX XVII.

fig. 36) are not very abundant, but characteristic, and presumably

mucinogenous as they stain deeply in hematoxylin. Their basal

ends, resting on the basal membrane, are broad and in each is a

rather small subspherical nucleus, above which the cell tapers to

a fine tube filled with a darkly staining granular material: these

attenuated cell-bodies run between the supporting cells and ter-

minate in swollen extremities filled with coarse deeply staining

spherules. Throughout this region of the ootype the epithelium

is thrown into ridges and furrows, which are partly due to the

folding of the walls, but chiefly to the unequal length of the

epithelial cells. At the hinder end the folds increase and there is

a prominent valve projecting into the lumen and making a com-

plete spiral turn just above the opening of the V-shaped duct.

This opening, guarded by the above-mentioned valve, is narrow.

The upper part of the V-shaped duct is lined by an epithelium of

the same character as that of the hinder moiety of the ootype, the

Proc. Zoot, Soc.—1911, No. LIV. 54

782 PROF. G. C. BOURNE ON THE

supporting cells being here distinctly: ciliated and the deeply

staining mucinogenous cells conspicuous. This epithelium is

thrown into a distinct spiral ridge, which winds round the upper

part of the V-shaped duct nearly as far as the entrance of the

duct of the receptaculum seminis. Here the epithelium changes

its character; the deeply staining mucinogenous cells disappear

and give place to very numerous long tubular gland-cells with large

oval basal nuclei, the tube-shaped cell-body coarsely alveolar and

the alveoli containing large refringent non-staining spherules.

Wedged between these are the ciliated cells, with elongated nuclei

at about the middle of their length, very attenuated basal ends,

and somewhat expanded wedge-shaped distal ends, each with a

distinct striated border anda tuft of fairly long cilia. The recep-

taculum seminis (Pl. XX XVIT. fig. 37) is lined by a low columnar

ciliated epithelium of uniform character. The cell-bodies are finely

granular and stain readily; the nuclei spherical and deeply

staining. Each cell has a distinct striated border and bears a tuft

of long coarse cilia. The spermatozoa in the receptaculum are all

arranged with their heads directed towards the centre, their tails

outwards and entangled among the cilia of the epithelium. The

receptaculum and its duct are invested by a very stout coat of

muscular fibres.

The transition from the upper end of the ascending limb of the

V-shaped duct to the narrow tube of the oviduct is abrupt. The

oviduct is lined throughout by an epithelium consisting of long

columnar cells bearing specially long and coarse cilia. The cha-

racters of these cells are shown in fig. 38 (Pl. XX XVILI_).

The cecum of the ootype is lined by an epithelium differing

from that of the rest of the ootypein the absence of glandular cells.

The walls of the cecum, like those of the hinder moiety of the

ootype itself, are provided with a tolerably thick coat of muscular

fibres, mostly disposed circularly. The muscular coat is not folded,

but the epithelium is disposed in longitudinal ridges due, as seen

in fig. 39 (Pl. XX XVIT.), to the greater length of the cells com-

posing them: this figure is from Orobophana ponsonbyi and not

from Alecadia; in the latter genus the cells are somewhat longer

and more slender, but otherwise similar in character. Asis shown

in the figure, the cells are club-shaped with rounded ends projecting

into the lumen of the cecum; they do not bear cilia. It is the

presence of this characteristic epithelium in the bilobed sac full

of spermatozoa leading into the hinder end of the ootype in Paleo-

helicina and Orobophana which leads me to identify the sac in

question with the cecum of the ootype of Aleadia and Hutrochatella

rather than with the receptaculum seminis, and it is further to be

remarked that in the non-ciliated bilobed sac the spermatozoa are

arranged pell-mell, with their heads and tails in all directions

instead of being definitely oriented as they are in the ciliated

receptaculum seminis.

The distal third of the vagina is lined by an epithelium consisting

for the most part of highly vacuolated clear cells with basal nuclei,

MORPHOLOGY OF THE HELICINIDA. 783

and between these are very attenuated supporting cells. I could

not detect any cilia on the latter.

The proximal two-thirds of the vagina and the vaginal sac

are lined by a non-ciliated epithelium of uniform character, the

details of which were not well preserved in my sections of Alcadia

and Hutrochatella: apparently they had been injuriously affected

by the action of Perenyi’s fluid. In Orobophana ponsonbyi the

epithelial cells of the lower part of the vagina and of the vaginal

sac are squarish in outline, non-ciliated and clearly glandular, for

each contains a number of coarse non-staining spherules. The

cavity of the sac contains a number of spherules of similar

character and among them ropy masses of some coagulable sub-

stance staining faintly in hematoxylin.

In Orobophana the epithelial lining of the ootype differs in some

respects from that described for Alcadia. The supporting cells are

attenuated towards their bases, have long compressed nuclei

about the middle of their length, and are distinctly ciliated. The

gland-cells of the distal part of the ootype resemble those of

Alcadia, but are apparently differentiated to some extent, for the

granular contents of those on the inner side of the ootype nearest

the mantle-cavity are eosinophilous, those on the outer side are

not. Just above the entrance of the vaginal duct the eosinophilous

cells are replaced by mucinogenous cells staining deeply in

hematoxylin. A large spiral flap or valve separates the opening of

the cecum from that of the ascending limb of the V-shaped duct.

The last named is very narrow and invaginated for some distance

into the terminal part of the ootype: it has no spiral epithelial fold,

such as is seen in Alcadia. The gland-cells of the distal limb of

the V-shaped tube are highly eosinophilous.

Thiele (10) has shown that the female ducts of Hydrocena are

monaulic. The external aperture leads into a thick-walled glan-

dular duct, which is clearly homologous with the ootype of the

Helicinide. The lumen of this duct is continued posteriorly into

a fairly wide canal lined by a columnar epithelium devoid of

glandular cells, and this ends in a saccular dilatation of consider-

able size, which is apparently glandular; “sein Epithel enthilt

Kliimpchen von Kornchen.” Thiele identifies this sac and its

duct with the vaginal sac and vagina of Helicina, and regards it

as the homologue of the right kidney of the Trochid, which in

this case has not acquired an independent opening into the mantle-

cavity. In addition to this sac, three other. structures open into

the hinder end of the “ootype” in Hydrocena: on the right a

thick-walled glandular cecum; on the left the oviduct; and

between the two and dorsad of the “right kidney sac” a tubular

receptaculum seminis. Thiele’s homologies seem to be perfectly

just, and after his discovery of the conditions obtaining in Hydro-

cena, | must agree with him in regarding the vaginal sac of the

Helicinidee and the spermatophore sac of the Neritide as the

representatives of the right kidney. But I still beg leave to

differ from his interpretation of the vaginal aperture as the

54*

784 PROF. G. C. BOURNE ON THE

primitive aperture of the right kidney, and of the external aperture

of the ootype as a secondarily acquired separate genital duct. The

conditions in Hydrocena appear to me to be an ample justification

of the argument put forward in p. 873 of my memoir on the

Neritide. Hydrocena is in many respects more primitive and

therefore more nearly related to the ancestral Neritoid stock than

the Helicinide, and Thiele himself points tothe generative organs

as one of the evidences of primitive organization.

In his memoir entitled “« Die systematische Stellung der Soleno-

gastren und die Phylogenie der Mollusken” (9) Thiele, after de-

scribing the male organs of Septaria, gives the following account

of the male organs of Helicina (= Waldemaria Wagner) japonica :

“ Bei Helicina ist der miinnliche Geschlechtsapparat merklich

einfacher, der Samengang ist nur sehr wenig aufgekniuelt und

weiter, er mundet in den Driisengang von unter und rechts nicht

weit vor seinen Hinterende. Der letztere ist bedeutend einfacher

als bei Vavicella, durch Falten streckenweise etwas zertheilt,

doch scheint das driisige Epithel trotz geringer Verschiedenheiten

an manchen Stellen in Wesentlichen gleichartig zu sein. Die

Driisenzellen liegen durchweg zwischen den Stutzzellen. - Dieser

Gang reicht weit nach vorn in der Mantelhohle doch habe ich ein

besonderes Kopulationsorgane nicht wahrgenommen. Die ekto-

dermale Driise, welche rechts von Hinterende des Driisenganges

(Prostata) in die Mantelhéhle ausmiindet, erstreckt sich, in dem

sich allmihlich grosser wird, weit nach hinten, wo sie neben dem

Hinterende der Niere aufhért. Nach ihrem Bau ist an ihrer

ektodermalen Herkunft nicht zu zweifeln, da sie zwischen den

grobkornigen Driisenzellen deutliche Stutzzellen enthalt. Sie mag

als eine Art von Manteldriise ihnlich der Hypobranchialdriise

anzusehen sein: ihre Funktion ist unbekannt.”

The ectodermal gland referred to in this passage I have already

described as the hypobranchial gland, fully agreeing with Thiele’s

interpretation of it. It has no connection, with the generative

organs. In the six genera that I have examined the male ducts

are, with the exceptions to be mentioned hereafter, very much

alike, but not quite so simple as Thiele’s description would lead

one to suppose, and presumably Waldemaria japonica has undergone

some simplification in these organs, for so accurate an observer

cannot have overlooked the accessory organs that I am going to

describe.

The testis, like the ovary, is follicular in structure ; fine thin-

walled ducts converge from the follicles and unite in the mght

side of the visceral mass to form the sperm-duct. The last-named

organ occupies a position similar to that of the oviduct in the

female: in immature specimens it is slender, slightly convoluted,

and lined by a columnar ciliated epithelium. In mature specimens

its middle portion is greatly distended by spermatozoa, is consider-

ably convoluted, and the ciliated epithelial lining is no longer

distinguishable. The sperm-duct tapers somewhat and, as Thiele

clescribes, opens into the lower and right side of a long glandular

MORPHOLOGY OF THE HELICINID. 785

thick-walled sac, some little distance in front of the hinder end of

the latter. This thick-walled sac, which is evidently the repre-

sentative of the ootype of the female and of what I have called the

terminal chamber in the Neritide, runs forward in the roof of the

mantle-cavity, below and to the right side of the rectum, and

opens into the mantle-cavity by a terminal pore situated close to

the anus. At about one-third of its whole length from the

external aperture, the thick-walled sac—which I shall call the

terminal sac of the sperm-duct or, more shortly, the terminal sac

—~is Joined by another sac of considerable diameter. ‘This second

sac, which I shall call the diverticulum, opens into the terminal

sac by a wide aperture, and runs back close to the right side of

the latter, to end blindly, sometimes just in front of the entry of

the sperm-duct into the terminal sac (Aphanoconia gouldiana,

Pl. XXXVIT. fig. 41); in other cases, however, it extends as far

back as the hinder end of the terminal sac, and may even project a

little beyond it (Alcadia hollandi, fig. 40). The anterior third of

the terminal sac, in front of the entry of the diverticulum, exhibits

three or four deep transverse constrictions: in section it is round

or oval, and the lumen is partly occluded by deep longitudinal folds

projecting into it. These folds are covered by a mixed glandular

and ciliated epithelium: the ciliated cells of the familiar kind with

attenuated basal ends, the gland-cells tubular with basal nuclei and

vacuolated cell-bodies, in which no secretory granules could be

distinguished in well-preserved specimens of Hatrochatella. In its

posterior two-thirds the terminal sac is laterally compressed so as

to be elongate oval in section, and it gives off from each end of the

oval and from the adrectal side numerous short hollow cecal out-

growths which are sometimes branched, especially in Aphanoconia

(fig. 41). Internally the longitudinal epithelial ridges die out in

the posterior two-thirds of the terminal sac, but the epithelial

lining both of the cavity of the sac and of the cecal outgrowths

is of very nearly the same character as that of the anterior third.

The supporting cells are distinctly ciliated. The epithelium of the

diverticulum difters from that of the terminal sac only in the fact

that the gland-cells are full of eosinophilous granules, and the

cilia of the supporting cells are longer and rather coarser. The

above characters hold good for all the species that I have examined,

the differences between them being too slight to deserve mention.

It may be noted that in Alcadia the hinder moiety of the diverti-

culum is constricted at very regular intervals (Pl. XXX VIT. fig. 40).

Both in Hutrochatella and Lucidella I have found in sections a

second diverticulum in the form of a slender thin-walled tube

opening into a recess of the terminal sac at the same level as, but

on the opposite side to, the diverticulum above described. The

walls of the recess are lined by a glandular epithelium staining

deeply in eosin. The narrow tube runs back in the mantle-wall

nearly parallel to the terminal sac and ends blindly just in front

of the aperture of the hypobranchial gland. Its hinder end

touches and appears to be adherent to the sub-epithelial muscular

786 PROF. G. C. BOURNE ON THE

wall of the mantle-cavity, but there is no aperture into the mantle.

I have not been able to find this tube in other species, but this

may be due to the imperfection of my sections and to the fact that

it is too small to be recognizable in dissections. It is a well-defined

structure in Hutrochatella and is lined throughout. by a non-glan-

dular cubical epithelium. I am inclined to the opinion that the

anterior third of the terminal sac of the male is the equivalent

of the body of the ootype of the female; the diverticulum of the

male represents the cecum of the ootype, and the narrow tube

(Pl. XXX VIT. fig. 42, 4.7.) represents the vagina of the female,

but has lost its opening into the mantle-cavity. If this iden-

tification is correct, a relic of the right kidney-sac is retained in

the male, at least in Hutrochatella pulchella and Lucidella aureola.

It may be noted that Isenkrahe’s drawing of the male organs

of Helicina titanica is very nearly correct.

The Nervous System.

Nobody has given a detailed account of the nervous system of

any Helicinid since Bouvier dealt with this subject in his

classical memoir on the nervous system of Prosobranch Gastropods

(3). In that work he gives an elaborate figure of the nerve-

centres and principal nerve-trunks of Helicina sagraiana dOrb.,

and also several figures of the buccal ganglia of the same species

and of the cerebral and pleuro-pedal centres of H. brasiliensis

Gray. As is always the case, Bouvier’s figures possess a high

degree of accuracy, and if I have some criticisms to offer, they

must not be taken as depreviatory of his excellent work, but

as an elaboration of it, rendered possible by careful study of

sections and by the opportunities for exceedingly fine dissection

afforded by the Braus-Driiner microscope.

In the first place, it was necessary to determine whether the

supra-intestinal nerve exists in the Helicinide. Bouvier had

failed to find it in the Neritide, and when in a subsequent memoir

he announced its discovery in the latter group, he hazarded the

opinion that it would probably be found in the Helicinide.

But it does not exist in these pulmonate rhipidoglossates ; it has

‘disappeared in them as completely as the organs with which,

when present, it is associated, the ctenidium and the osphradium.

I can speak with certainty on this point, for I have made so

many dissections and have studied such a sufficient number of

serial sections that I could not have overlooked it if it were

present.

In the second place, I am unable to verify some of the details of

Bouvier’s figure of the nervous system of 7. sagraiana. In none of

the species that I have studied are the pedal, pleural, and subintes-

tinal centres as distinct as shown by him. As may be seen in

fig. 44 (Pl. XX XIX.), the pleural ganglia are ill-defined swellings,

scarcely distinguishable from the swollen anterior ends of the

pedal cords, and the subintestinal ganglion is so intimately fused

MORPHOLOGY OF THE HELICINID®. 787

with the pleurals that it is unrecognizable as a separate ganglion,

even in sections (Pl. XX XVIII. figs. 45 to 52). The pedal cords,

though not so widely separated in any of the species of Heli-

cinide that I have dissected as in the Neritide, are not so closely

approximated as shown in Bouvier’s figure. They are fairly

close together and nearly parallel to one another in Hutrochatella

pulchella, somewhat more divergent in Alcadia hollandi, further

apart in A. palliata, and widely divergent in <Aphanoconia

andamanica. The actual amount of divergence or approximation

is, however, undoubtedly dependent on the degree of contraction

of the muscular mass of the foot, and is a character cf no great

importance. If the foot is much contracted the pedal cords are

approximated and the numerous and slender pedal commissures

are arched : if the foot is relaxed the cords are turther apart and

the commissures are pulled out straight. I cannot but think

that Bouvier has exaggerated the length of the cerebro-pleural

and cerebro-pedal connectives. In none of the species that I have

examined are they appreciably longer than the antero-posterior

diameter of the cerebral ganglia, and in some species, e. g.

Lucidella aureola, they are very short, but it is, of course, possible

that they are unusually long in H. sagraiana. In respect of the

nerves issuing from the ganglionic mass formed by the fusion of

the anterior ends of the pedal with the pleural centres, Bouvier,

while otherwise exact, makes one important omission. He does

not figure or describe a relatively large pair of nerves which

originate one on each side of the most anterior pedal commissure

from the dorsal surface of the swollen anterior ends of the pedal

cords. Each of these nerves (Pl. XX XIX. figs. 43 & 44, m.op.)

passes outwards and backwards, penetrates the muscular wall of

the body, and passes to the muscles of the operculum, hence the

nerves in question may be called the opercular nerves. That of

the right side breaks up into a number of fine twigs in the

opercular muscles, that of the left side gives off a stout branch

which passes to a peculiar hollow organ connected with a plate of

cartilage near the left corner of the opercular Jobe. This organ

will be described in detail further on.

The otocysts are situated just above the origins of the two

opercular nerves, and are therefore on the dorsal side of the pedal

ganglia, as is shown in fig. 44. Bouvier, describing the otocysts

of Helicina brasiliensis says ‘elles sont situées sous les cordons

pedieux ”: and it is true that in a retracted specimen, in which

the head has been drawn back behind the foot, so that the

pedal cords appear to lie in front of the cerebral ganglia, one

does find the otocysts below the pedal cords, when making a

dissection from the dorsal side. But in such a case the sole of

the foot is uppermost, and the morphologically dorsal side of the

pedal cords is turned downwards, this change of position being

very puzzling to the observer both in sections and dissections.

It is worth remarking in this connection that in my experience

the small Polynesian genera, Aphanoconia, Palcohelicina, and, in

788 PROF, G. C. BOURNE ON THE

a lesser degree, Orobophana, when they withdraw themselves into

their shells, do not contract themselves as much as the West-

Indian species. The sole of the foot in these Polynesian genera

is longer and narrower than in the West-Indian genera, the

columella: muscles longer and inserted further back from the

mouth of the shell. ‘There is therefore more ample room for the

head and foot in the last whorl of the shell, and when the animal

retracts itself the foot is scarcely at all contracted, but simply

slides back with its sole applied to the outer side of the shell till

the operculum borne on the broad opercular lobe closes the

aperture. Specimens of these genera, when extracted from their

shells, do not present the deformed appearance of an Alcadia or

an Hutrochatella, the pedal nerve-cords are not turned forward

with their morphological surfaces reversed, and if the animals

were only a little larger they would be much easier to dissect

than their American congeners. It is possible, and even probable,

that these different modes of retreating into the shell, which are

themselves dependent on the varying length and points of

insertion of the columellar muscles, are correlated with the

different forms of operculum upon which Wagner has founded

his system. At any rate, they are consistent with it, but I have

not been able to follow out this problem in detail.

To return to the nervous system. The opercular nerves must

not be mistaken for the parietal nerves correctly described and

figured by Bouvier and labelled d', e’. The parietal nerves

(Pl. XX XIX, fig. 44, n.par.) are much more slender than the

opercular nerves and originate, as shown in fig. 44, from the

pleural centres, between the great pallio-columellar nerves and

the cerebro-pleural connectives. They pass to the muscular walls

of the head behind the tentacles. From the ventral side of the

swollen anterior ends of the pedal cords, in the same cross-section

as the opercular nerves, a rather stout pair of nerves originates

near the middle line; these nerves, which are shown in section

in fig. 46 (Pl. XX XVIIT.), pass to the pedal gland and appear to

be specially connected with that organ.

As regards the subintestinal nerve and its distribution, I am

unable to bring my observations into agreement with those of

Bouvier. The short nerve connecting the subintestinal with the

left pallio-columellar nerve-trunk does not appear to be a constant

feature. J have found such a connective in a single specimen of

Alcadia palliata, and in that one instance it is much closer to the

pleuro-pedal centres than is shown by Bouvier. But I can find

no trace of it in any other specimen that I have examined. I am

unable to find any trace of the visceral nerve labelled j’ in

Bouvier’s figure, which he describes as given off from the left side

of the subintestinal at some distance from the origin of the latter,

and in general my observations on the subintestinal and visceral

nerves differ so much from his that a detailed account is necessary.

The figure illustrating this account (PI. XX XIX. fig. 43) is founded

upon dissections, and the ultimate ramifications of the principal

MORPHOLOGY OF THE HELICINIDA. 789

nerve-branches have been traced in sections. The subintestinal

trunk in Alcadia palliata, A. hollandi, and Lutrochatella pulchella,

after leaving the subintestinal ganglion, which latter is unrecogniz-

ably fused into the pleural centres, courses along the floor of the

anterior division of the general body- -cavity, below the pharyngeal

bulb, the radular sac, and cesophagus. It keeps closer to the right

than to the left columellar muscle, and on reaching the posterior

end of the muscle it doubles the angle between it and the visceral

mass, and entering the latter turns to the left and enlarges to

form a ganglion of some size from which several nerves are given

off. That this isa true ganglion-centre, and not a mere nodal thick-

ening at a point from which several nerve-branches originate, is

denionstrated by the considerable sheath of nerve- ganglion cells.

The principal nerves issuing from the ganglion, in ade inn to the

main trunk, are the following :—a small nerve, v', which passes to

the right near the surface of the visceral mass and is distributed

to the gonads and liver. Another small nerve, v°, which passes

to the left of the visceral mass and appears to innervate the right

lobe of the liver and-surface of the stomach. A stout nerve,

n.gen., which runs to the right, passes above the oviduct or

sperm-duct, gives off a large branch to the mucous gland, and

turns forward to break up into twigs on the posterior part of the

complex of genital ducts: this is the genital nerve, and it

identifies the ganglion from which it originates as the visceral

ganglion. There is no separate genital ganglion as in_ the

Neritide. The hinder end of the visceral ganglion is continued

into a rather stout nerve, which may be regarded as the

continuation of the main trunk of the subintestinal: this passes

through the liver and helow the small and large intestine, and

turning towards the left it passes towards the lowes border of the

right moiety of the kidney, near which it enlarges to form a small

but distinct ganglion, which I take to be the representative of the

elongated visceral ganglion of the Neritide. From this ganglion

small nerves are given off to the liver and kidney, and a larger

nerve passes below the kidney, skirts the uropore, and can be

traced beyond as far as the auricle of the heart, at which point it

ceases to be recognizable. It is a matter of extreme difficulty to

follow the above-mentioned nerves through the liver and intestinal

coils by simple dissection, and I have only been able to make sure

of their ultimate course by the study of serial sections.

After this criticism of the general characters of the nervous

system, | may return to the consideration of some special details.

The pleuro-pedal centres and with them the subintestinal

ganglion are, as I have already said, so intimately fused as to be

practically indistinguishable as separate ganglia. This fusion is

brought out in a striking manner in sections. Figs. A5 to 52

(Pls XXXVIIL) represent selected members of a series of nearly

transverse sections through the pleuro-pedal centres of Alcadia

hollandi. Fig. 45 represents a section through the pedal cords at

the point where they begin to diverge from one another. Dorsad

790 PROF. G. C. BOURNE ON THE

of them are the otocysts, o¢.; ventrad of them is the pedal gland.

Each cord consists of a core of nerve-fibres and dendrites surrounded

by a cortical layer of nerve-ganglion cells. The latter send in

lateral horns at about the middle of the outer side of each cord

in such a manner as to divide the central core into upper and

lower moieties, which have been identified by French authors

with the pleural and pedal sections of the cords respectively.

This interpretation, however, does not appear to me to be well

founded. Fig. 46 represents a section somewhat further

forward than that in fig. 45; it passes through the hinder part

of the anterior pedal commissure, and includes the roots of the

two nerves of the pedal gland and of the right opercular nerve.

The nerve-fibres of the former are seen to be supplied from two

areas of the cortical layer lying respectively on the outer and

inner sides of the ventral side of the cord. The opercular nerve

receives its fibres partly from a centre on its own side, partly

from a centre on the opposite side of the cord, the latter fibres

crossing over in the commissure. Below them is a stout band of

commissural fibres connecting the lateral horns of nerve-ganglion

cells with one another. Above the median ventral raphe is a

thick mass of nerve-ganglion cells. In a section somewhat

further forward (fig. 47) the ventral raphe has disappeared and

the mass of ganglion-cells above it is only represented by two

small islets of nerve-ganglion cells, which are separated from the

ventral surface by two well-defined bands of nerve-fibres start-

ing from the ventro-lateral groups of nerve-ganglion cells and

passing towards the centre partly decussate, partly sweep round

the islets to curve round to the lateral horns of the ganglion-

cells. Above these curved bands is the well-defined transverse

commissural band of nerve-fibres, and above this again there is

ow either side a centre, consisting of nerve-fibres overlaid by a

layer of ganglion-cells, which is seen to be connected with the

origins of the opercular nerves. Above the nervous mass a pair

of muscular cords passing from the otocysts towards the centre

should be noticed.

In the next section (fig. 48), taken some little way further

forward, the two little islets of ganglion-cells lying opposite the

lateral horns are still visible. Below them are transverse bands

of commissural fibres. Between them is seen the most anterior

part of the decussating tract observed in the previous section.

After decussation the fibres sweep out right and left to the dorso-

lateral regions. In the mid-dorsal line is a deep and wide groove

into which the muscular cords noted in the last section are

entering. I regard the whole of the sections hitherto described

as belonging to the pedal centres. The next section (fig. 49)

shows that the dorsal groove containing the two muscular cords

has been converted into a canal by the upgrowth and dorsal

union of the nervous tissue. All that lies below this canal

belongs to the pedal centres; all that lies to the sides of and

above the canal belongs to the pleural centres.

MORPHOLOGY OF THE HELICINIDAD. 791

In the pedal centres a prominent bundle of nerve-fibres 1s

being formed on either side of the middle line: these when traced

forward prove to be the origins of the cerebro-pedal connec-

tives. Above them are the remains of the transverse anterior

pedal commissure. Laterally, above the lateral horns of the

ganglion-cells two other tracts of nerve-fibres are making their

appearance: these have evidently received large contributions

from the decussating fibres noticed in the previous sections, and

when traced forward they prove to form part of the cerebro-

pleural connectives. The central canal containing the two muscular

cords is surrounded by a layer of nerve-ganglion cells, thickest

on the ventral side, and above the canal are seen tracts of nerve-

fibres originating from the lateral horns of either side and passing

towards the middle line. Above them, again, is a small transverse

band of commissural fibres, and above these a fairly thick layer

of ganglion-cells, which, as may readily be seen by comparing

this with the preceding sections, is something added to what was

there before, and is, in fact, the layer of pleural ganglion-cells.

The next three figures (50, 51, and 52) explain themselves. In

fig. 50 the pedal centres are diminishing rapidly in volume. On

the left side, which, owing to the sections being somewhat oblique,

is rather behind the right side, a large tract of vertical nerve-

fibres is seen passing from the pedal to the pleural centres. To

the outside of this above the lateral horn are tracts of fibres some

of which run upwards and will pass into the left pallio-columellar

nerve, others will be continued into the right cerebro-pleural con-

nective and the right parietal nerve. On the right side the origins

of the cerebro-pedal and cerebro-pleural connectives are well

defined. Between them lie the ganglion-cells of the lateral horn,

and above the nucleus of the cerebro-pleural connective is a second

lateral ingrowth of nerve-cells, belonging to the pleural ganglion.

The pleural centre of nerve-fibres is well masked, and from it a

stout band of commissural fibres passes above the central canal to

the pleural centre of the other side. This is the pleural com-

missure, which I have already described in Neritide. Fig. 51

shows the cerebro-pedal and cerebro-pleural connectives in cross-

section. The central canal with its contained muscular slips,

having passed through the ring formed by the pleural and pedal

centres, emerges as a groove on the ventral side. The origins of

the right parietal and right and left pallio-columellar nerves are

clearly visible, and it may be seen that both pleural centres are

contributing fibres which, passing to the mid-dorsal line, form

the origin of the subintestinal nerve. It should be observed

that some fibres from the last named pass directly into the roots

of the two pallio-columellar nerves. In fig. 52 the origin of the

subintestinal nerve is distinct, and both it and the origins of

the two pallio-columellar nerves appear to be imbedded in a mass

of ganglion-cells in which the limits of the right and left pleural

and the subintestinal ganglia can be traced with the aid of a

little exercise of the imagination.

(2 PROF. G. CG. BOURNE ON THE

From what precedes, it follows’ that the pleuro-pedal centres are

extremely complicated and are largely composed of definite tracts

of nerve-fibres, some commissural, some decussating, some passing

into the nerves issuing from this region. It is noticeable that there

are several connections of a complex kind between the pleural and

pedal centres as well as between the right and left pleural centres

and the right and left pedal cords, and that there are evidences of

numerous nervous relays throughout the region illustrated. No

doubt, on analysis, these apparently complicated nerve-tracts can

be reduced to five groups: (1) the pedal commissure, containing

both direct and decussating tracts of fibres; (2) the pleuro-pedal

connectives ; (3) the cerebro-pedal and cerebro-pleural connec-

tives ; (4) the pleural commissure, peculiar to the Neritide and

Helicinide ; (5) the subintestinal connectives, derived from

both right and left pleurals. Thus, the paths of the nerve-tracts

might be described as normal, and consistent with our knowledge

of the usual connections between the chief nerve-centres of

Molluscs ; but.a study of sections will show that the arrangements

are not so simple as might be inferred from a superficial exami-

nation of the ganglia and their commissures and connectives.

For example, it appears that the cerebro-pleural connectives

contain fibres derived from the pedal centres and that all the

principal nerves contain fibres derived from two or more areas in

the fused pleural and pedal ganglion-mass. I have been unable

to pursue the subject further at the present time, and it is diffi-

cult to make any further progress because of our ignorance of

the physiology of the molluscan nervous system. ‘The main

nerve-trunks must contain both afferent and efferent nerves, and

it seems evident that these pass to different areas of the cortical

layer of ganglion-cells, but as at present we have no means of

distinguishing between these two kinds of fibres further analysis

of the details of the nervous system is impossible. It may be of

use to future workers on this subject to remark that there are two

kinds of ganglion-cells in the cortical layer: larger cells with

clear nuclei staining faintly in hematoxylin, and much more

numerous smaller cells with deeply staining nuclei. It should be

possible to trace the connection of the nerve-fibres with these

different kinds of cells, but such an investigation demands fresh

material, and could form no part of the present work on the

Helicinidee.

As Bouvier found considerable differences in the size and shape

of the cerebral ganglia in Helicina sagraiana and H. brasiliensis,

I have studied these centres with care in the hope that I might

discover characters of classificatory value, but I have been no

more successful here than I was in the case of the genital ducts.

The characters to which Bouvier draws special attention are the

relative size of the cerebral ganglia (enormous in /. brasiliensis) ;

the size and shape of the labial lobe; the origin of the labio-probos-

cidean nerves, which all spring from the labial lobe in H. brasz-

liensis, but only one has this origin in H/. sagraianu. As regards

MORPHOLOGY OF THE HELICINIDE. 793

the relative size of the cerebral ganglia I find that the proportion

of each cerebral ganglion to the pleuro-pedal mass is expressed by

the following figures :—in Alcadia palliata, 4; Alcadia hollandi,

+33 Hutrochatella pulchella, %; Lucidella aureola, 3; Paleo-

helicina ide, nearly +; Orobophana ponsonbyi, 4; Aphanoconia

gouldiana, +3; Aphanoconia rogersit, 2. The relative length of

the cerebro-pedal and cerebro-pleural connectives varies greatly :

they are longest in the two species of Alcadia and in Paleo-

helicona ide ; of moderate length in Orobophana, Aphanoconia,

and Hutrochatella ; extremely short in Zucidella aureola. I have

counted four labio-proboscidean nerves on each side with more

or less certainty in all the species examined with the excep-

tion of Aphanoconia rogersii, in which there appear to be five.

Of these, counting from above downwards, the first and third

are invariably stout nerves which branch soon after their origin ;

the second and fourth are slender and only divide into branches

at their extremities. The labial lobe in all the species at

my disposal has the form of a rounded boss projecting inwards

from the antero-inferior edge of the ganglion: the shape and

relative size of this lobe differ somewhat in the various species,

but the differences are too slight to express in words. Fig. 53

(Pl. XX XIX.), representing the left cerebral ganglion of Palco-

helicina ide, and fig. 54, representing the same ganglion in Aleadia

palliata, show the extremes of difference in shape observed by me,

and, on the whole, the cerebral ganglia of the Polynesian species

resemble those of Palewohelicina, those of the West-Indian species

those of Aleadia. ‘The cerebral ganglia of the Helicinide, wedged

in as they are between the anterior end of the pharyngeal bulb

and the walls of the head, are nearly flat, the labial lobe projecting

inwards beneath the pharyngeal bulb. Because of their flatness

they are very readily stained and mounted as transparent objects,

and figs. 53 & 54 give some idea of the complexity of the nerve-

tracts and centres within the ganglion. One may distinguish an

ocular centre, which is large relatively to the size of the ocular

nerve, a tentacular centre, small relatively to the size of the

tentacular nerve, and a relatively smaller commissural centre.

Hach of the labio-proboscidean nerves has a more or less well-

defined centre of its own, but the buccal and labial commissures

have no distinct centres at their origin. There is, further, a

median lobe which possibly serves as a relay for various nerve-

tracts running into and around it. ‘It is evident that the nerve-

fibres of the cerebro-pleural connective make direct and intimate

connection with the ocular and tentacular lobes, and that a

stout band of fibres curves round from the root ef the cerebro-

pleural connective to the base of the labial lobe, receiving on its

way an accession of fibres from the cerebro-pedal connective.

This nerve-tract makes connections with the centres of origin of

the buccal and labial commissures and the labio-proboscidean

nerves, and a well-defined curved band of fibres sweeps round

from the origin of the first labio-proboscidean nerve to enter the

794 PROF. G. C. BOURNE ON THE

cerebral commissure, the last named also having connections with

the tentacular and median centres. Evidently, the cerebral

ganglia have undergone a high degree of concentration and

integration, but it is to be remarked that if the labial lobe as

figured for Alcadia were pulled out towards the bottom of the

picture, it would form an elongated labial process from which

the labio-proboscidean nerves would be given off at intervals, as

is the case in Zrochus and Turbo. In other words, the labial

lobe, as it seems to me, is represented, not solely by the little

projecting boss from which the labial commissure originates, but

by all that part of the ganglion that hes below a line drawn from

the lower side of the origin of the cerebral commissure to a point

just above the origin of the buccal commissure.

The buccal ganglia, with their commissures and the nerves

originating from them, have been very correctly figured by

Bouvier.

The organ which I have mentioned above as connected with a

branch of the left opercular nerve is shown in section in fig. 55

(Pl. XL.). It is deeply embedded in the muscular tissue of the

opercular lobe and lies to the left of the anterior end of the

lobe close to the origin of the left columellar muscle. It consists

of a cartilaginous plate of subtriangular form, the edges of the

anterior apex inrolled ventrally and eventually fused so as to form

a short conical tube. This cartilage forms, as it were, the cover

of a flattened sac (fig. 55, sac.) lined throughout by an epithe-

lium which is thin and composed of a single layer of somewhat

flattened cells on the side attached to the cartilaginous plate, but

thick and composed of columnar cells bearing short stiff cilia on

the opposite side. Anteriorly this is continued into the tube

formed by the inrolled edges of the cartilaginous plate, and here

the columnar ciliated cells form a nearly continuous lining to

its cavity. Posteriorly, as shown in fig. 55, the ciliated cells

forming the floor of the sac (the reader must understand that the

figure is reversed, so that the ventral side is uppermost) rest on a

thick basement-membrane, from which a broad band of muscular

fibres spreads to be attached partly to the muscular wall of the

left side of the neck, partly to the bands of muscular fibres passing

over the otocysts. Into this muscular band the large branch of

the left opercular nerve penetrates. Taking an anterior course

this nerve gives off a branch to the opercular muscles, but its

main trunk is directed towards the anterior tubular end of the

organ under consideration, and there passes through a small

perforation in the cartilaginous wall and is distributed to the

ciliated columnar epithelium lining the cavity of the sac. Dorsally

the cartilaginous plate is connected with the left columellar

muscle by a stout muscular band passing obliquely outwards.

Above this band are seen in fig. 55 the sections of two convolu-

tions of a coiled glandular tube, which on the one hand communi-

eates by a very narrow duct with the anterior tubular end of the

above-mentioned sac, and on the other hand opens to the exterior,

MORPHOLOGY OF THE HELICINIDA, 795

near the left anterior edge of the operculum, between it and the

membranous flap that surrounds the opercular lobe. This tube is

lined throughout by a glandular epithelium composed of rather

tall goblet-cells with deeply staining basal nuclei and clear cell-

contents.

If, now, we enquire into the morphological significance of this

peculiar organ, I think there can be little doubt as to its homology.

It occupies the same position and receives the same nerve-supply

as the crypt into which fits the curved process of the operculum in

the Neritide. This process is no longer to be seen on the

operculum of the Helicinide, but from a study of the muscular

attachments in the two groups I am inclined to think that the

cartilaginous plate described above represents its inner extremity,

all direct connection with the operculum being lost. In

connection with it new structures have been formed, viz. the sac

and the glandular tube. What its physiological significance is it

is hard to say. It is clear that, by contraction of the muscular

bands attached to it, the sac may be widely dilated, and when

dilated, air must flow into it through the glandular tube. The

abundant nerve-supply and the character of the columnar epithe-

lium bearing short stiff cilia suggest that the sac has a sensory

function, and it is possible that it may be a special sense-organ,

likely enough of an olfactory character, which enables the animal

to receive impressions from the external world when retracted

into its shell. For the opening of the glandular tube is in such

a position that it would open to the outer air whenever the

opercular plug was ever so little loosened.

The pedal gland, an organ which is absent in the aquatic

Neritidee, is largely developed in the Helicinidee. Its position and

general structure are roughly indicated in fig. 45 (Pl. XX XVIII).

It consists of a main duct below the pleuro-pedal nerve-mass and

extending some little way but not far back below and between the

pedal cords. The duct is lined by a columnar ciliated epithelium

raised on the ventral side into two prominent ridges, one on either

side of the middle line. Surrounding the duct are bunches of

unicellular glands, which penetrate among the interlacing muscle-

fibres of this region of the foot. Kach unicellular gland is

prolonged into a fine duct which passes between the epithelial

cells of the main duct and opens into its lumen. The histology

of this gland is reminiscent of that of the byssus gland of

lamellibranchiate molluscs, described by me in another place.

The main duct opens at the anterior end of the foot, in the mid

line, below the snout.

Mention may be made here of the large mucous gland which I

agree with Thiele in identifying as the hypobranchial gland. It is

of relatively enormous size in the Helicinide, but in structure and

position does not differ much from what I have described for the

Neritide. It opens into the mantle-cavity (Pl. XX XV. figs. 28

& 30) just in front of the aperture of the vaginal duct and, as shown

in figs. 30 to 35, it forms a considerable glandular mass bulging”

796 PROF. G, C. BOURNE ON THE

into the mantle-cavity to the left side of the genital ducts. As

may be seen in figs. 16-20 (Pls. XXXII. & XXXIIT.), it extends

far beyond the hind end of the genital ducts, accompanying the

rectum in its course, and lying to the right side of the kidney

at the hinder extremity of the visceral mass. In the section

depicted in fig. 20, which passes throngh the rectal coil at the

bottom of the hind end of the visceral mass, the hypobranchial

gland on the right side of the rectum appears to form a pair with

the kidney on the left, but there is, of course, no relationship

between the two organs. Throughout its course the hypobran-

chial gland consists of an irregularly folded flattened sac, from

which short glandular diverticula are given off in all directions.

In its terminal part, as is shown in figs, 18-20, the gland appears

to be differentiated into two portions: one lying nearer the

cesophageal end of the stomach is lned by an epithelium loaded

with fine dark granules; the other portion, lying nearer to the

mantle-cavity, is lined by an epithelium of the character shown

in Pl. XX XIX. fig. 56. It is made up of large glandular cells,

oblong in outline, and filled with a highly refracting granular

substance which, when the cells are ready to discharge their

contents, is accumulated into oval pellets, as shown in the figure.

Between the gland-cells are long and very attenuated interstitial

cells, of which the outer ends! are expanded and produced to

form a cover over the outer ends of the gland-cells. The nuclei

of the interstitial cells lie in their expanded outer ends. I could

find no trace of cilia. In nearly every specimen examined the

mantle-cavity was full of a sticky gelatinous mass secreted by the

hypobranchial gland.

From what precedes it is evident that the Helicinide are a very

homogeneous group, so far as their anatomical characters are

concerned, and that such differences as I have been able to detect

are of very little assistance in classification, the resemblances and

differences in one set of organs suggesting one class of affinities,

those of another set of organs suggesting another class. Thus,

taking the arrangement of the coils of the intestine as a cr iterion,

we should place Alcadia, Or obophana, and Paleohelicina alse

together, with Zucidella related but somewhat apart : Hutrochatella

would go with Aphanoconia to form a distinct group. But if we

took the characters of the female gonaducts as a criterion we

should get a different arrangement. WHutrochatella would stand

nearest to Alcadia, with Aphanoconia more distantly related :

Paleohelicina with Orobophana would form a distinct group.

The nervous system is so similar in all the species that it affords

a very slight guide, but, as far as the cerebral ganglia give any

clue, the genera would fall into an arrangement consistent with

their geographical distribution; Alcadia, Lucidella, and Hutro-

chatella forming one group; Paleohelicina, Orobophana, and

Aphanoconia another. Finally, the radular characters, which

have attracted a considerable share of the attention of systematists

and in my experience afford the most reliable and readily

MORPHOLOGY OF THE HELICINIDA. 797

recognizable marks of distinction between different species,

give a totally different result. Taking these as a criterion,

Eutrochatella stands well apart: the remaining genera show a

sufficient amount of similarity to justify our placing them in a

single group, in which Paleohelicina stands nearest to Alcadia ;

Aphanoconia is closely related to Palewohelicina; Orobophana, 3

while showing relationship to the two last named, has distinctive

characters which keep it somewhat apart; and Lucidella, while

showing relationship to <Alcadia, has undergone modifications

which, in one feature at any rate, resemble those which distin-

euish Orobophana from Palwohelicina.

On the whole, the radular characters afford the safest clue to

affinity, agreeing as they do with the conclusions founded on

conchological characters and on geographical distribution.

In all the Helicinide, so far as is known, the radular formula

may be represented as co .1(3+1+3).1.0. The Proserpinide

have a somewhat different, and the Hydrocenide a very different,

formula, but they need not be considered here.

As different authors use different names in deseribing the teeth

of Neritoid gastropods, I must define my terms before proceeding

further. Of the teeth included in brackets in the formula, I call

the single tooth in the centre the median; the three teeth on

either side of it the admedians. he large tooth on either side of

the admedians | call the lateral, and the numerous teeth to the

outside of these the marginals. In Hutrochatella, as Troschel has

shown, the lateral teeth are relatively very large and of charac-

teristic shape, being mushroom-shaped, with the top of the pileus

hollowed out to form an articular cavity, into which fits the stalk

of the lateral tooth of the row next in front of it. The edge of the

pileus is entire. I have attempted in fig. 57 (PI. IL): to give

some idea of the very complicated str neues of this tooth in Hutro-

chatella pulchella. It should be noted that it has an external

rocess, or ‘‘ Basalanhang,” which Troschel described as charac-

teristic of the genus Helicina. For the rest, this mushroom-shaped

tooth more nearly resembles in shape the corresponding radular

tooth in the Neritide than is the case in any other Helicinid.

The marginal teeth of Hutrochatella ave simply pointed curved

bars, without denticulations at their free extremities. In the

genus Helicina Lamarck (non sensu restricto Wagner) the laterals

are not pileiform, but consist of a stout median portion which I

shall call the “‘ stalk”; from the inner side of this a more or less

broad aliform plate projects obliquely forward; the anterior

border of this plate is thickened, recurved, and bears a number of

denticulations, varying from 7 to 12 in number in the different

species that I have studied. Attached to the outside of the stalk

by an imperfect joint is the pointed external process (the

‘“‘ Basalanhang ” of Troschel), and the top of the stalk is excavated

to form an articular cavity for the hinder end of the stalk of

the corresponding tooth in the row next preceding. Also, in all

species of Helicina the marginal teeth have broadened recurved

Proc, Zoot, Soc.—1911, No. LY. 5o

798 PROF. G. 0. BOURNE ON THE

anterior extremities bearing denticulations. Thus, there is a

considerable difference between Hutrochatella and Helicina, but the

gap is bridged over by 7’rochatella chrysochasma, 1a which, accord-

ing to Troschel, the lateral tooth is pileiform, but with an oblique

anterior border bearing from 7 to 9 denticulations, and whereas

the proximal marginal teeth are simply pointed as in Hutrochatella,

the more distal marginals bear denticulations, increasing from one

to four innumber. I haveshown that anatomically Hutrochatella

bears the closest resemblance to Alcadia, differing only in the

arrangement of the coils of the intestine.

The genus Alcadia is characterized by a notch separating the

peristome of the shell from the columella. It is stated (Fischer,

‘Manuel de Conchyliologie, p. 795) that the operculum has a

dentiform process: I can only say that I cannot find a trace of

any such structure in A. palliata and A. hollandi. In all other

respects Alcadia is similar to the point of identity to Helicina.

Troschel declares that the only recognizable difference between the

radule of Aleadia and Helicina is in the form of the median tooth,

a character of very little value, for, as I shall show, the shape of

this tooth varies from species to species. But Troschel’s figure of

the radula of Alcadia is not very exact. J have given in Pl. XL.

fig. 58, a to f, large scale drawings of the teeth of A. hollandi;

those of 4. palliata differ only in minute particulars. Comparing

these with the drawings of the radule of Lucidella, Palecohelicina,

Orobophana, and Aphanoconia (Pls. XL.-XLII. figs. 59-65), it

will be seen that the first admedian tooth of Alcadia has a

characteristic shape, being subquadrangular in outline, with its

anterior outer angle produced into a knob on which are borne

four blunt denticulations. The large lateral has a short stalk,

hardly projecting behind the origin of the aliform plate: the

latter is large, expanded, bearing seven stout but blunt teeth on

its recurved anterior margin. The articular excavation is very

shallow: the external process long and pointed.

Of the other radule figured, those of Palwohelicina (Pl. XLI.

fig. 60) and Aphanoconia (Pls. XLI. & XLII. figs. 62-65) bear the

closest resemblance to Alcadia. Both these genera were included

in Helicina Lamarck, and have only recently been separated by

Wagner. In them the lateral teeth have the same general shape

as in Alcadia, but the stalk is longer, the aliform plate less

expanded, the denticulations on its anterior border vary in size,

shape, and number. The first admedian tooth is very similar in

the two genera, and differs from that of Aleadia. The median

tooth is very variable in size and shape. The similarity of the

radulee of Palcohelicina and Aphanoconia is remarkable, and leads

one to doubt whether Wagner is justified in placing these forms

in different genera. On the other hand, Orobophana (Pl. XLI.

fig. 61) is distinct: its lateral. tooth is that of Paleohelicina, but

the first admedian is relatively large, acutely triangular, its

anterior border thickened but without dentieulations; it is

feebly corneous and nearly transparent, suggesting that it is in

MORPHOLOGY OF THE HELICINID®. 799

course of disappearance. The median tooth is heart-shaped,

minute, and similarly feebly corneous. Lweidella (Pl. XL. fig. 59)

is quite distinct in radular characters: in the lateral teeth the

stalk is practically obsolete, the bulk of the tooth consisting

mostly of the aliform process with its thickened denticulate

border, which is continued posteriorly into the articular knob.

There is no anterior articular excavation, but a thin triangular

external piece which serves to support the articular knob of the

tooth of the row next in front, and for the attachment of the

external process. The third admedians are of the usual petaloid

shape; the second admedians stout, triangular, with a thickened

curved anterior edge, bearing on its outer surface a small minutely

denticulate trenchant process. The first admedians are rather

large, but feebly corneous, with a thickened anterior non-denti-

culate border; they have been modified in a manner analogous

to what has been observed in Orobophana. The medians are

broadly heart-shaped, feebly corneous, nearly divided into two by

a deep median anterior notch.

Summing up these details and taking into comparison Troschel’s

figures, which are mostly of species of. Helicina sensu restricto of

WwW agner, and confining our attention to the lateral tooth, which is

the largest and obviously of most functional importance in the

Ner itacea as well as in the Helinacea, we see that there is an easy

transition from LHtrochatella to Alleadia that the lateral tooth

of Alcadia is of the form characteristic of the Helicinide in

general, but shows a tendency to a reduction of the stalk, which,

as Troschel has shown, is common to many American and West-

Indian species. This tendency is exhibited in an extreme form

by Lucidella. But in the Pacific and Oriental genera the stalk

and the articular excavation connected with it are well developed.

But the lateral tooth of Hutrochatella bears an extremely close

resemblance to that of the Neritide, and there is this further

resemblance, that the first admedian tooth, which is of very large

size in the Neritide, is relatively of much lar ger size as compar ed

with .the second and third admedians in Hutrochatella than in

any other Helicinid. If such characters can be relied upon as a

guide to affinity, Hutrochatella is the most closely related among

the Helicinidee to the Neritoid ancestor of the group. From

Eutrochatella forms have been derived: on the one hand, the

Proserpinidee, which also have a large pileiform lateral fond: :

on the other hand, Helicina. We may infer that the earliest

Helicine retained the stalk and articular excavation which are

such marked features in the pileiform lateral tooth of Hutro-

chatella and Proserpina.

The forms which, as suggested in the earlier part of this paper,

were transported bys some unknown means across the Pacific

Ocean to the Philippines must have possessed these features

and transmitted them unchanged to their descendants which

now inhabit the Oriental and Indo-Pacific regions. But in

America and the West Indies there has been a tendency, more

55*

800 PROF. G. G. BOURNE ON THE

fully realised in some species than in others, to’a reduction of the

stalk and articular cavity, this reduction being shown to a slight

degree in Alcadia, to a marked degree in Laucidella.

In all the Helicinids there is a tendency to the reduction of

the central and admedian teeth: this tendency is shown in a

marked deeree in Lucidella and Orobophana, but must have

reached its present degree independently in these two genera.

The reduction is carried to an extreme degree in Hydrocena, in

which the second and third admedians have disappeared; the

median and first admedian are present, but in a rudimentary

condition, and the laterals are reduced to mere rods of no great

size. But I am disposed to think that the “Hydrocenide must

have branched off from the Neritoid stock independently of the

Helicinidee. They retain many primitive features, as Thiele has

shown, among others the process of the operculum which is quite

Neritoid in character, and their geographical distribution favours

this view. Mydrocena is contined to the marine littoral of

Dalmatia; Georissa lives at considerable altitudes on the Khasi

Hills in India. It is by no means improbable that pulmonate

forms may have been developed more than once from such

animals as the’Neritide, which show a predilection for migrating

as far as possible out of the water, and for the rest of it, the most

that can be said in favour of uniting the Hydrocenide with the

Helicinidee is that both display strong Neritoid affinities.

The main result of my researches is to show that in such a

limited group as the Helicinide the systematists are justified in

their methods. The visceral anatomy of all the forms that I

have examined is strikingly similar, and where deviations occur

they are contradictory and of uncertain value. The Helicinide

appear to have inherited an organization with marked Neritoid

characteristics, and to have maintained it, with little or no

change. Presumably that organization is well adapted to the

somewhat narrow range of the conditions of their existence, and

any deviation from it has been checked by the action of natural

selection. But there are a thousand deviations, 1m all directions,

among characters which cannot by any stretch of the imagination

be claimed to be of any importance in the struggle for existence.

Such characters are the texture and coloration of the shell; the

shape of the aperture; the extent and distinctness of the basal

eallus ; the presence or absence of folds at the aperture of the

shell; the presence or absence of a minute notch, such as occurs

in Aleadia; the arrangement of the growth-lines on the oper-

culum. It might be said that the operculum is an important

protective organ and therefore eminently susceptible to the action

of natural selection. But its function is simply to close the

aperture of the shell, and this it does equally efficiently in all

the species that I have examined, the number of these being

much larger than the few available for anatomical study. As

long as the operculum performs this function efficiently mmute

characters, such as the greater or less distance of its nucleus from

MORPHOLOGY OF THE HELICINIDA. 801

the anterior border, cannot possibly determine the question of

the death or survival of the animal. The same reasoning applies

to the variations of the radular teeth : the function of the radula

is to rasp, and any of the modifications shown in figs. 60 to 65

is equally efficient as a rasp. Nobody, I think, would venture

to assert that the minute differences in the four species of

Aphanoconia (figs. 62 to 65) could have had any value in the

differentiation of these species by natural selection.

Asa result of my somewhat elaborately minute studies, I am

driven, and, I confess, somewhat unwillingly driven, to the con-

clusion arrived at by a number of naturalists, that natural

selection is efficient in preserving characters of physiological

importance, but ineffective in producing new species by adding

together numerous minute successive variations. The only

conclusion justified by the facts seems to me to be that the

characters on which systematists rightly rely are of the nature

of deviations or mutations, of no consequence to the well-being

of the animals in which they appear, but inheritable, and there-

fore perpetuated under favourable circumstances by segregate

breeding. The Helicinide, inhabiting narrow areas, and often

segregated in remote islands, afford particularly favourable oppor-

tunities for segregate br eedin, g,

As to how far hase small deviations of functionally unimportant

structures may be due to the influence of external conditions I

do not venture to offer an opinion, but the following fact is

suggestive. Among the shells in the tube containing several

specimens of Viphaiacanae merguiensis was a specimen which

in size, shape, coloration, and marking so exactly resembled the

others ‘that I took it fon a Helicinid (as the colleetor must also

have taken it) and decalcitied it with a view to anatomical

investigation. It proved to be a Helicid, of what genus and

species I cannot say, as I had destroyed the shell and could not

find another specimen.

Among the collection of Helicinide made in the Andaman

Islands and presented to the British Museum of Natural History

by Mr. G. Rogers was a tube containing half a dozen specimens

which differ recognizably in the characters of the shell and

operculum from Aphanoconia andamanica Benson, but are clearly

closely related to that species. I have not been able to refer

them to any named species, and as the radular characters show

it to be distinct from anrdamanica, | describe it as a new species,

as follows :—

APHANOCONIA ROGERSII, sp.n. (PI. XLII. figs. 65-69.)

Shell oblately spheroidal, the surface marked with closely set

radial growth-lines ; colour ight orange-yellow marked with more

or less distinet reddish-brown radial bands ; spire of 47 whorls,

increasing regularly and somewhat rapidly in size, the last whorl

obtusely keeled, the keel produced into a prominent angular

802 PROF. G. C. BOURNE ON THE

projection at the peristome. Aperture semilunar, very oblique,

the outer margin thickened and expanded. Basal callus not

very thick, hardly differing in colour from the rest of the shell,

of rather small extent, its limits clearly defined above, as well as

below. ‘The whole shell deeper in proportion to its breadth than

in A. andamanica and the spire more prominent.

Operculum yellowish white in colour, the calcareous plate

rather thin, the sigmoid curve pronounced, the upper angle

produced.

Radula with small diamond-shaped median tooth; the first

admedian tooth more than twice as long as broad, the anterior

edge incurved and bearing four denticulations ; second and third

admedian teeth each with four denticulations; the lateral teeth

normal, the stalk rather long, the articular excavation deep, the

aliform process bearing seven round denticulations.

Closely as the shell of this species resembles that of A. anda-

manica, a glance at the drawings of the radule of the two species

(figs. 63 & 65) shows that they are distinct.

I must express my obligations to Miss Margaret Poole, both

for helping me in the determination of the different species of

Aphanoconia and for making the drawings of shells and radule

for figs. 62 to 68.

As I have discussed and offered an explanation of the geo-

graphical distribution of the Helinicide without either adopting

or criticizing the theories advanced by Dr. Simroth (7 and 8) on

this subject, I must, in conclusion, make some reply to the friendly

eriticisms that he has published on my paper on the Neritide.

I do not propose, in this place, to discuss the physiological

interpretation that he has given of the different arrangements of

the female ducts in the Neritide. For one thing, I have

obtained some new material and hope soon to publish further

observations throwing fresh light upon the problems to which

he refers: for another, J am inclined to accept much of what he

writes on that part of the subject.

_ But with regard to the ancestry of the Neritide, and with them

the Helicinide, which Dr. Simroth would derive from a pulmonate

stock, and with regard to the homologies that he wishes to establish

between the generative ducts of Neritide and Pulmonata, I am

unable to accept any of his conclusions. To do so would be

to throw the whole fabric of morphological reasoning to the

ground. Dr. Simroth’s views on homologies are largely influenced

by a theory of secular changes in the sea-level produced by a

swinging or “ pendulating” movement of the earth about an axis

which corresponds with the longest diameter of the earth and has

its poles in Sumatra and Ecuador. It is not my present intention

to discuss the difficult astronomical and geological problems in-

volved in the “ Pendulation theory,” and, indeed, I am sure that I

am incompetent to discuss them. The theory may be well founded

or 1t may not: I do not offer an opinion; but be it right or

MORPHOLOGY OF THE HELICINID. 803

wrong, I fail to see that the conclusions deduced from it by Dr.

Simroth are necessary. I will explain as briefly as possible why.

Unless I misunderstand him grievously, and if I do I beg his

pardon, one of Dr. Simroth’s chief conclusions is that, contrary to

the generally accepted doctrine, marine and freshwater animals in

general, the marine prosobranch Gastropods in particular, are

evolved from terrestrial forms which have been forced by the

above-mentioned secular inundations to adapt themselves to new

conditions of life and make their habitat inanother medium. As

the pendulation theory applies to all geological time, if the pre-

cursors of marine Gastropods were terrestrial in habit, we should

find evidence of this in geological deposits. The earliest-known

Gastropods, from the Cambrian to the Devonian, would bear

evidence of their terrestrial life, those found in later deposits

would indicate, in some periods at least, the change from a ter-

restrial to a marine existence. But, in point of fact, the geological

evidence points decisively the other way. In Cambrian, Ordovician,

Silurian, and Devonian deposits we get Gastropods belonging

almost exclusively to the Streptoneurous Aspidobranchia and

Pectimibranchia. There are, it is true, the pteropod-like shells of

the Conularida which, if they are really remains of Pteropods,

would demonstrate the great antiquity of highly specialized forms

of Kuthyneura. But the true systematic position of the Cornu-

larida is at the best doubtful, and it has been urged with much

reason and on high authority that the resemblance between the

shells of these archaic forms and the more modern Pteropoda is

due to parallelism. As so much doubt prevails as to their affinities,

the Cornularida cannot be brought into the argument. The

Aspidobranchiate and Pectinibranchiate Gastropods from these

earlier Paleeozoic deposits are without doubt marine forms. They

subsisted, without any important changes, through the four above-

mentioned geological epochs, and one genus, Plewrotomaria, has

survived to the present day. We know the habits and the

anatomy of Plewrotomaria, and they support in a most remarkable

manner the conclusions derived from an extensive knowledge of

gastropod morphology. On the other hand, with the exception of

Hercynella from the Devonian, undoubted Euthyneura first make

their appearance in the Car ‘bonifer ous. They belong to the

Acteonide and Pulmonata Stylommatophora. The first-named

family is marine, and anatomically displays so many strepto-

neurous characters that it might almost be included in the

Aspidobranchia. Of the Sty lommatophora we get forms hke

Dendropupa and Pyranidula, unquestionably terrestrial species,

and, according to views generally accepted, highly modified and

therefore indicative of a line of lost ancestry probably allied to the

contemporary Actzonide. But these pulmonate forms are few

and of rare occurrence in the Carboniferous, a period in which

the conditions for the preservation of terrestrial and freshwater

forms were particularly favourable. Had numerous Pulmonates

existed at that time their remains must have been more abundantly

804. PROF. G. G. BOURNE ON THE

preserved, Terrestrial pulmonates are still scanty in the Permian

and Trias, and only begin to show a considerable increase in the

Jurassic and Cretaceous. I need not labour the point further.

Clearly, paleontological evidence does not favour Dr. Simroth’s

theory of the orgin of marine from terrestrial Gastropoda.

But let us suppose that paleontological evidence may be ignored

on account of the imperfection of the geological record, and that

the Pendulation theory is so well supported by other evidence as

to compel us to give credence to Dr. Simroth’s doctrines as to the

origin of marine from terrestrial Gastropods. The Helicinide

are terrestrial and pulmonate. I have shown, and in so doing

have only corroborated the opinion of all other observers, that

they are Neritoid in almost every feature of their anatomy. If

the marine and fluviatile Neritids were to be derived from a ter-

restrial and pulmonate form, one would suppose that that form

must have been Helicinid in character, for the affinities between

the two groups are so very obvious. But Dr. Simroth does not

discuss this possibility. Making reference to Ostracolethe,

Hyalimax, Limax, and Arion, all highly specialized recent

Pulmonates, he boldly derives the Neritide from the Stylomma-

tophora, relying largely upon the supposed homology of their

generative ducts. This homology I do not admit: a resemblance

there i is, but not a close one, and, even if it were closer than it

actually i is, L should place very little reliance on the anatomy of

the generative ducts as indicative of relationship between groups

differing widely in all other respects. In the different phyla of

invertebrated animals the generative ducts are notoriously variable

in character. In the Platyhelmia, for example, their variety is

bewildering. Within the phylum Mollusca there are many in-

stances of variability and also of deviations which must have been

independently acquired but are in the same direction, as, for

instance, in the Doridomorpha and Elysiomorpha. The re-

semblances, such as they are, between the generative ducts of the

monecious Pulmonata and the dicecious Neritidee are just what

one might expect to find in animals in which a common plan of

organization, to wit a gastropod organization, is modified im

accordance oath etnies physiological requirements. The differ-

ences are of amply sufficient magnitude to betray a difference of

origin. In other words, the complex g gonaducts of Neritide and

Pulmonata are independently acquired structures, and such

resemblances as they display are due to parallelism.

T have already referred to the anatomy of the Pleurotomariide,

a family which existed in the Cambrian and survives to the present

day. Thanks to Bouvier and M. F. Woodward, we are well

acquainted with the anatomy of Plewrotomaria, which affords a

striking confirmation of the reliability of sound morphological

reasoning. Before Pleuwrotomaria had been studied, comparative

anatomists, as the result of extensive investigations of gastropod

structure, had come to an agreement concerning numerous marks

of primitive organization in the group. When this survivor from

MORPHOLOGY OF THE HELICINIDA. 805

the Paleozoic age came to be examined, all these marks were

found, some of them in a more pronounced degree than in any

other known Gastropod, and in no system of organs were these

marks more conspicuous than in the nervous system, the im-

portance of which Dr. Simroth seeks to minimize. Among these

marks may be enumerated—a cerebral commissure situated far

forward on the pharyngeal bulb; a distinct labial commissure ;

elongated and scalariform pedal nerve-centres ; a long crossed

visceral commissure; two auricles to the heart; the ventricle

lapped round the rectum; a rhipidoglossate dentition. Other

characters might be enumerated, but these suffice for the present

purpose. All these characters are absent in the Pulmonata : all

of them are present in the Nevritide. Moreover, by discovering

the oviduco-celomic funnel, I was able to demonstrate, beyond all

reasonable doubt, the homology of a part of the gonaducts to the

right kidney of Pleawsotomaria and other rhipidoglossate Aspido-

branchs, a homology which Thiele had already asserted on other

grounds. Now it is quite clear that, if structural resemblance is

of any value as a guide to affinity, we have a choice between two

alternatives. Bither the Neritide, to which we must add the

Helicinide, are descended from Aspidobranch ancestors, which

they resemble in all the points enumerated above, and have in-

dependently acquired genital ducts superficially similar to those

of Pulmonata; or,as Dr. Simroth will have it, they have descended

from stylommatophorous Pulmonata, have preserved the characters

of the genital ducts of the latter group, but have independently

acquired all the other characters enumerated above, characters

possessed by no Pulmonate, but invariably present in those

Aspidobranchs from which, on Dr. Simroth’s showing, the

Neritide are not descended. I am not quite sure whether he

would go so far as to assert that the remaining Aspidobranchs

possess those characters because they are descended from the

Neritide. To make such an assertion would, indeed, be flying in

the face of all reasoned opinion on this subject, and would amount

to a declaration that the geologically more recent Pulmonates are

the parents of their predecessors of Cambrian age !

T submit the alternative to the judgment of my readers, and in

doing so beg leave to enter a protest against the srowing tendency

to throw over long-established and carefully reasoned conclusions

founded upon morphological evidence, because of thei uncon-

formity with some new and as yet insufficiently tested hypothesis,

or because they do not help in the solution of certain limited

problems. I was quite aware, when I discussed the subject, that

the geographical distribution of the Neritide was a puzzle, and

that L had failed to find a solution to it. The distribution of the

Helicinide is scarcely less puzzling and awaits a final solution.

But with all respect for Dr. Simroth’s authority and deserved

reputation as a zoologist, | submit that the solution that he offers

is improbable, raises a crop of other puzzles, and throws

morphology into confusion.

806

(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

PROF. G. C. BOURNE ON THE

List or LireERATURE REFERRED TO.

Amauprur, A.—La partie antérieure du tube digestif et la

torsion chez les Mollusques gastéropodes. Ann. des Sci.

Nat. (8) vii. 1898, p. 1.

Bourne, G. C.—‘‘ Contributions to the Morphology of the

Group Neritacea of Aspidobranch Gastropods.—Part I.

The Neritide.” Proc. Zool. Soe. Lond., 1908.

Bouvier, EK. L.—‘‘ Systeme nerveux, Morphologie générale

et Classification des Gastéropodes Prosobranches.” Ann.

des Sci. Nat. (7) i1., 1887.

IsENKRAHE, C.—‘‘ Anatomie von Helicina titanica.” Arch.

f. Naturgeschichte, xxxiil., 1867, p. 50.

JHERING, H. von.—Vergleichende Anatomie des Nerven-

systems und Phylogenie der Mollusken. Leipzig, 1877.

Kopett, W.—Studien zur Zoogeographie. Wiesbaden, 1897.

Srurora, H.—‘“ Neuere Arbeiten iiber die Morphologie und

Biologie der Gastropoden.” Zool. Zentralblatt, Bd. xvi.,

ISOS),

SimrotH, H.—“The Anatomy of the Neritide.” Proc.

Malacol. Soc. Lond., ix. 1910, p. 27.

THIELE, J.—‘‘ Die systematische Stellung der Solenogastren

und die Phylogenie der Mollusken.” Zeitschr. f. wiss.

Zool., xxii. 1902, p. 249.

Ture, J.—“ Uber die Anatomie von Hydrocena cattaroensis

Pf. Abh. d. Senckenbergischen Naturf. Gesell., xxxi1.,

1910.

TroscHEL.— Das Gebiss der Schnecken. Berlin, 1856-1863,

Bd. 1. pp. 75-85.

Waener, A. J.—‘“ Helicinenstudien.” Denkschr. d. kais.

Akad. d. Wissenschaften, Wien, Bd. Ixxvii., 1905, and

Ixxviu., 1906.

(A complete list of the literature of the Neritoidea is given in

my previous paper [2 ]).

EXPLANATION OF THE PLATES.

Pruates XXX.-XLII.

Lettering in all the figures.

an. Anus. con. ped. Cerebro-pedal connective.

ao. Aorta. | con. pl. Cerebro-pleural connective.

ap. mg. Aperture of hypobranchial ce. ph. Lateral pharyngeal cartilage.

gland. div. Diverticulum of male gona-

Au. Auricle. duct.

bue. Buccal cavity. E. Hye.

c@. Cxcum of ootype. F. Foot.

cl. Cloaca. g. buec. Buccal ganglion.

e.m.l. Left columellar muscle. g. cer. Cerebral ganglion.

ce.m.r. Right columellar muscle. Gd. Gonaduct.

com. bucec. Buccal commissure. gl. pd. Pedal gland.

com. cer. Cerebral commissure. gi.r. Glandular ridge on floor of

com. lab. Labial commissure. the csophagus.

com. pd. Pedal commissure. g- pl. Pleural ganglion.

com. pl. Pleural commissure. Int. Intestine.

MORPHOLOGY OF THE HELICINIDA, 807

Ti. Liver. oot. Ootype.

li. d. Liver-ducts. Op. Opercultum.

M. Mantle. ot. Otocyst.

M. c. Mantile-cavity. ov. Ovary.

m. g. Hypobranchial gland. Pe. Pericardium.

n.cm.l. Left columellar nerve. ped. Pedal nerve-cords,

n.cm.r. Right columellar nerve. ph. Pharynx.

n. gen. Genital nerve. R. Rectum.

n. gl. p. Nerve to pedal gland. rd. Radula.

n. lpb. Labio-proboscidean nerves. rd. s. Radular sac.

m. oc. Ocular nerve. 7. p.c. Reno-pericardial canal.

n. op. l. Left opercular nerve. r. s. Receptaculum seminis.

n. op. r. Right opercular nerve. s. gl. Salivary gland.

n. ot. Otocyst nerve. Sn. Snout.

n. pal.-c.l. Left pallio-columellar nerve. sp. d. Sperm-duct.

n. pal.-c.r. Right pallio-columellar nerve. spz. Spermatozoa.

n. pal.l. Lett pallial nerve. St. Gisophageal moiety of

n. pal.r. Right pallial nerve. stomach.

n. par.l. Left parietal nerve. S¢!. Pyloric moiety of stomach.

n. par.r. Right parietal nerve. é. Tentacle.

n. sb.i. Subintestinal nerve. | é. s. Terminal sac of male gona-

n. ten. Tentacular nerve. duct.

od. Oviduct. Ur. Ureter.

od', Descending limb of V-shaped Ur.p. Uropore.

portion of oviduct. V. Ventricle of heart.

od*. Ascending limb of V-shaped vag. Vagina.

portion of oviduct. vag- ap. Opening of vagina into

od. c. a. Anterior odontophoral carti- mantle-cavity.

lage. | vg. s. Vaginal sae.

od. c. p. Posterior odontophoral carti- | V.G1. First visceral ganglion.

lage. | V.G*. Second visceral ganglion.

oe. (sophagus. | v. pal. Pallial vein.

ce. p. (Esophageal pouch. v. pst. Posterior pallial vein.

[email protected]. Opening of esophageal pouch | v. ren. Afferent renal vein.

into cesophagus.

Fig. 1. Left side view of Alcadia palliata. The mantle has been cut through close

to the left columellar muscle and turned back to expose the interior of the

mantle-cavity, the pericardium has also been opened.

Fig. 2. Dorsal view of the buccal cavity, pharynx, and anterior part of the

esophagus of Alcadia palliata: the buccal cavity, pharynx, and part of

the cesophagus have been laid open; gl.r., glandular ridge on the floor

of the cesophagus.

Fig. 3. A dissection: of the cesophagus and pharynx of Aleadia palliata, seen from

the left side.

Fig. 4. A horizontal section through the pharynx and buccal cavity of Alcadia

hollandi. ,

Fig. 5. The odontophoral cartilages of Aleadia palliata, viewed from below.

Fig. 6. The odontophoral cartilages of Hutrochatella pulchella, viewed from above.

lee Yo AN seo of the stomach of Alcadia palliata, viewed from the ventral

side.

Fig. 8. A portion of the epithelium of the stomach of Alcadia hollandi, showing

glandular and ciliated cells. about 960.

Fig. 9. Part of a section through the prominent ridge in the stomach of Alcadia

hollandi, < about 600; cu., the thick cuticle covering the ridge.

Fig. 10. The alimentary tract of Alcadia palliata, showing the arrangement of the

intestinal coils.

Fig. 11. Alimentary tract of Alcadia hollandi.

Fig. 12. Alimentary tract of Pale@ohelicina ide.

Fig. 13. Alimentary tract of Orobophana ponsonbyi.

Fig. 14. Alimentary tract of Zwcidella awreola.

Fig. 15. Alimentary tract of Hutrochatella pulchella.

Fig. 16. A horizontal section through the upper part of the visceral mass of

Alcadia hollandi. Figs. 17 to 22 are drawn from the same series of

sections.

Fig. 17. A section somewhat lower down, showing the origin of the ureter, Ur.,

from the kidney.

808 ON THE MORPHOLOGY OF THE HELICINIDA,

Fig. 18. A section still more ventral than the above, showing the deepest part of the

pericardium and its relation to the kidney.

Fig. 19. A more ventral section passing through the reno-pericardial canal,

Vp.c.

Fig. 20. A ee section passing through the ventral part of the visceral mass.

Fig. 21. Part of a section slightly ventral to that drawn in fig. 19, showing the

uropore, Uy.p., opening into the mantle-cavity.

Fig. 22. A section through the reno-pericardial canal. Magnified about 600.

Fig. 23. Part of a section passing through the uropore of Lucidella awreola and

showing the character of the renal epithelium. X 960.

Fig. 24. A diagram reconstructed from the series of sections drawn in figs. 16 to 20,

showing the relations of the kidney, ureter, stomach, pericardium, and

mantle-cavity.

Fig, 25. The genital ducts of Aleadia hollandi 2. 'The lower half of the figure is

drawn as it appears when viewed by transmitted light, the upper part as

seen by reflected light. In this and in figs. 26-29, 40, and 41, the

gonaducts are represented as seen from the ventral side, after the wall of

the mantle-cavity has been cut through by a dorsal incision and the

rectum and gonaducts turned over to the right side of the animal.

Fig. 26. The genital ducts of Hutrochatella pulchella 9 .

Fig. 27. The genital ducts of Aphanoconia merguiensis 2.

Fig. 28. The genital ducts of Paleohelicina ide ¢.

Vig. 29. The genital ducts of Orobophana ponsonbyi & .

Fig. 30. A longitudinal section through the genital ducts of Alcadia hollandi ?,

passing through the aperture of the hypobranchial gland.

Fig. 31. A longitudinal section from the same series, showing the aperture of the

vaginal duct.

Fig. 32. Another section from the same series, showing the origin of the oviduct

from the ovarian chamber.

Fig. 33. Another section from the same series showing the connection of the vagina

with the vaginal sac and ootype.

Fig. 34. Another section from the same series showing the opening of the oviduct, od.,

into the descending limb of the V-shaped tube, od.’

Fig. 35. Another section from the same series showing the receptaculum seminis

opening into the ascending limb of the V-shaped tube.

Fig. 36. Glandular epithelium from the wall of the ootype of Alcadia hollandt.

Fig. 37. Ciliated epithelium and spermatozoa from the receptaculum seminis of

Aleadia hollandi.

Fig. 38. A section through the oviduct of Alcadia hollandi. Highly magnified.

Fig. 39. An epithelial ridge from the bilobed cecum of the ootype of Orobophana

onsonbyi.

Fig. 40. ee of the genital ducts of Aleadia hollandi 6.

Fig. 41. A similar view of the genital ducts of Aphanoconia gouldiana 6.

Fig. 42. A drawing made from a combination of several serial longitudinal sections

through the genital ducts of Hutrochatella pulehella 8, showing the

narrow diverticulum, &.7., which may possibly represent the vagina of the

female and therefore be the homologue of the right kidney-sac.

Fig. 43. A dissection showing the pedal, pleural, and visceral nerve-centres in

Aleadia palliata, with the principal nerves issuing from them. The

cerebral ganglia have been removed. The dissection is made from the

dorsal surface; the foot, as is usual in contracted specimens, is turned

forward and lies in front of the head with the sole uppermost; the walls

of the head and the mantle have been cut away, and the visceral mass has

been dissected as far as is necessary to show the course of the subintestinal

nerve and its branches.

Fig. 44. The nerve-centres and principal nerve-trunks of Alcadia hollandi, viewed

from the right and above. The drawing was made with the camera

lucida, after removal of the nerve-centres from the body.

Figs. 45-52. A series of transverse sections through the pleuro-pedal nerve-centres of

Alcadia hollandi, showing the principal tracts of nerve-fibres in the fused

pedal, pleural, and subintestinal ganglia. For a full description of these

figures, see the text, p. 789. The position of the pedal gland is indicated

in fig. 45.

Fig. 68. The left cerebral ganglion of Paleochelicina ide, viewed from the inner

surface. The ganglion is stained with Mayer’s hemalum and drawn by

transmitted light ; 1, 2, 3, 4, the four labio-proboscidean nerves.

Fig. 54. A similar preparation of the left cerebral ganglion of Alcadia palliata.

ON THE PALATABILITY OF SOME BRITISH INSECTS. 809

Fig. 55. A drawing of a section showing the position, structure, and nerve-supply of

the opercular organ of Hutrochatella pulchella.

Fig. 56. A portion of the epithelium of the hypobranchial gland of Alcadia

hollandi. Wighly magnified.

Fig. 57. A left lateral tooth from the radula of Hutrochatella pulchella. Highly

magnified.

Fig. 68. Radular teeth of Aleadia hollandi, highly magnified: a, median; 6, c, d,

first, second, and third admedians of the left side; f; one of the marginals

or uncini; é, a lateral tooth of the right side showing the stalk, stk., the

aliform internal plate, al.p., the articular excavation, art., and the

process, ext.p.

Fig. 59. Three rows of teeth from the radula of Lucidella aureola. In this and

the following figures only the proximal members of the marginals are

indicated.

Fig. 60. Two rows of teeth from the radula of Palg@ohelicina ide.

Fig. 61. Two rows of teeth from the radula of Orobophana pachystoma ponsonbyi.

Fig. 62. Two rows of teeth from the radala of Aphanoconia gouldiana.

Fig. 63. Two rows of teeth from the radula of Aphanoconia andamanica.

Fig. 64. Two rows of teeth from the radula of Aphanoconia merguiensis.

Fig. 65. Two rows of teeth from the radula of Aphanoconia rogersii.

Fig. 66. Shell of Aphanoconia rogersii.

Fig. 67. Shell of the same species, showing the aperture.

Fig. 68. Shell of the same species, viewed from above.

Fig. 69. Operculum of Aphanoconia rogersii, viewed from the inner or ventral side.

36. On the Palatability of some British Insects, with Notes

on the Significance of Mimetic Resemblances. By

R. I. Pocock, F.R.S., F.L.S., F.Z.S., Superintendent

of the Society’s Gardens and Curator of Mammals.

With Notes upon the Experiments. By Pict. E. B.

Poutton, F.R.S., F.Z.S.

[Received and Read May 9, 1911.]}

INTRODUCTION.

At the request of Prof. E. B. Poulton, F.R.S., I undertook, in

the summer of 1909 and again in that of 1910*, to make a series

of experiments in the Zoological Gardens to test the palatability

of various species of British Insects. Much of the material was

sent to me by Dr. G. B. Longstaff from Morthoe in Devonshire.

Some I received from Prof. Poulton himself or from friends of

his. A few species I added on my own account; notably the

stick insects and the ants, of which we had an abundant supply

in the Insect House in the Gardens. Those that I supplied

I identified myself. The rest were in all cases named by the

senders. To the insects Dr. Longstaff added a number of slugs,

which were identified, I understand, by Mrs. Longstaff.

Since the majority of the experiments were made with English

Insects, it is regrettable that English, or at all events Palearctic

birds, were, for the most part, unavailable for the tests. There

were two reasons for this. In the first place, Palearctic insecti-

vorous birds were not strongly represented in the Society’s

* Records of a few experiments made in 1911 haye been incorporated in the text,

810 MR. R. I. POCOCK ON THE

collection. Im the second place, those that were in the Zoological

Gardens at the time were, in most cases, kept in a very large

flight aviary with plenty of cover in the way of shrubs, repre-

senting their natural environment as nearly as possible. Never

having been tamed by confinement in small cages, they were too

shy to come to the bars to take insects from my hand and too

seared to notice them if I entered the aviary. Once or twice

I tried the experiment of liberating butterflies in this aviary ;

but the frequency with which they escaped through the wire

mesh and were wasted for the purpose in hand, induced me to

abandon further experiments of that kind.

This reference to the shyness of birds in captivity brings me

to another of the limitations under which I was working. I

was forced to restrict my attention to particular birds, tame

enough either to take insects directly from me or sufficiently

accustomed to the presence of human beings in the aviary to

capture liberated insects in spite of my close proximity. If I

put the insects through the bars, myself standing outside, they

were either seized one after another by the boldest bird in the

place, or were carried by a timid bird to the back of the com-

partment, where | could not watch what befell them. I was

compelled, therefore, to be inside the bars. Since, moreover,

it was practically impossible to watch more than one bird at a

time, I was precluded from the method of experimenting with

the shyer specimens by giving insects to the bolder ones to

distract and monopolize their attention. Thus it comes about that

the same species appear over and over again in the experiments

below recorded, while many insectivorous birds, that might have

been tried but for their shyness, are omitted.

Two facts struck me very forcibly at an early stage of the

experiments. The first was the exceeding keenness of the birds

for the insects brought to them. This was no doubt due in a

measure to our inability in the Gardens to feed the birds on

living insects other than mealworms. ‘The living prey was

evidently a great treat to them; and over and over again I was

impressed with the persistence shown by birds in persevering

with insects that were obviously not to their liking, returning to

the morsels repeatedly as if food of such a nature was too good to

be wasted. From this I think it may be inferred that in a state

of nature hungry birds will eat nauseous insects which in times

of plenty they will reject after tasting, or will not take the

trouble to catch them if they have previously learnt their distaste-

fulness by experience. Furthermore, it is quite clear that the

plain record of an insect being eaten is no proof of its palatability.

Better evidence on this head is supplied by the behaviour of

the bird towards it. After a little experience in this matter, I

was able to satisfy myself at all events as to the approximate

correctness of my interpretation of the bird’s actions, and to

judge thereby of the comparative palatability of the insects they

tasted.

PALATABILITY OF SOME BRITISH INSECTS 811

The second fact has an important bearing upon the criticism

sometimes advanced against the theory of warning coloration

and mimicry as applied to butterflies, namely, that birds under

natural conditions are seldom seen to eat these insects.* Hence it

has been inferred that birds cannot be reckoned as serious enemies

of buttertlies. Whatever may be the explanation of the cireum-

stance, I am tolerably sure, from the behaviour of the two classes

of animals when pitted against one another, that the inference

drawn therefrom is erroneous. The insectivorous birds in our

aviaries seemed to know at once what the butterflies were; they

were on the alert the moment one was liberated and pursued 1t

with determination and precision, following its every turn and

twist, and either catching it upon the wing or pouncing upon it

after settling. It is true that this predatory deftness.may have

been acquired in relation to the chase of insects other than

Lepidoptera; but unless the birds recognised butterflies in

general—a group which cannot be mistaken for other insects—

as part of their natural prey, it is difficult to understand their

eager excitement at the sight of those I offered them.

Again, unless the species of butterflies used for the experiments

are, or were in the past, habitually preyed upon by birds,

whence comes the extraordinary skill the liberated specimens,

when undamaged or inexhausted by confinement, displayed in

dodging the swoop of the birds in mid-air? Having repeatedly

seen the aim of the pursuing bird baftled by the evasive twist

of the butterfly, I cannot doubt that the insect’s behaviour was

prompted by the instinct to escape an habitual enemy of its

species, of the same class and with the same predatory methods.

It cannot, I imagine, be seriously claimed that escape from the

upleap of insectivorous mammals, lizards, or frogs has been a

factor of sufficient importance in survival to be reckoned with

in this connection; and, a fortiori, the modernness of the

invention of the entomologist’s net puts this instrument of

capture out of court for consideration. The evidence, therefore,

seems to me to afford the strongest support to the conclusion

that the power to dodge in mid-air and the instinct to put it

in force have been fostered to subserve no other purpose than

the evasion of swift-winged insectivorous foes. Perhaps predatory

Pompilide must be regarded as a possible auxiliary influence ;

but apart from these hymenoptera, I can think of no enemies but

birds likely to have persecuted butterflies on the wing to the

extent presumably necessary to have guided their evasive tactics

to the pitch of proficiency they now exhibit.

Whatever be the value of this suggested explanation of the

facts, the facts themselves remain as I have stated them :—

(1) Caged insectivorous birds which, so far as is known, have never

been fed in captivity upon butterflies, are at once excited by

* Twice I have seen sparrows, which are not typically insectivorous, chase white

butterflies in London. Two birds acting in concert were successful on the first

occasion ; one singie-handed failed on the second occasion,

812 MR. R. I. POCOCK ON THE

their appearance, chase them with eager speed, catch them in

mid-air with precision, and eat them or taste them with avidity.

(2) Pursued butterflies when overtaken often avoid the birds,

not once only but twice or three times, by sudden turns up or

down to right or left.

Those who hold, on the negative evidence above stated, that

birds are not to be reckoned as serious enemies of butterflies,

must be called upon to sapply some explanation other than that

above proposed of the marked reactions between these two

classes of animals when brought into contact with one another,

and to show reason why what takes place in the aviary may

not be regarded as indicative of similar occurrences in nature.

With regard to the experiments on mimicry, especially those

made with Volucella bombylans and Bombus hortorwm, 1t appears

to me that they satisfy all that the theory, as propounded by

Bates, demands. They fully confirm Prof. Lloyd Morgan’s experi-

ments on birds, with the drone-fly (Hristalis) and the honey-bee

(Apis mellifica), as well as those with the banded and uncoloured

slips of glass holding respectively meal adulterated with quinine

and meal untampered with.* They show that several species

of birds, after learning by experimental tasting that Lombus

hortorum is unpalatable, refused to touch Volucella bombylans.

Other items of interest that may be briefly alluded to are the

experiment demonstrating, at least in the instance tried, the

attractive nature of the ocelli on the wings of the peacock butterfly

(Vanessa io); the experiments showing that Yormica rufa is not

protected from mammals and birds by its acid taste; that the

black members of the Carabide and Ocypus olens are unpalatable

to the ground-feeding mammals they were offered to; that

Coccinella T-punctata and the Telephorid beetle (? Rhagonyche

fulva)—belonging to families of beetles which are common

objects of mimicry in the tropics—are distasteful to nearly all

mammals and birds.

At the end of the part of the paper describing the experiments

made, I have added, at Dr. Longstaft’s suggestion, for the informa-

tion of those unfamiliar with the habits and distribution of the

mammals, birds, and reptiles to which the insects and other

invertebrates were offered, a list of the species of the former

groups giving a few particulars on those points.

Finally I have to thank Prof. Poulton for kindly annotating

the paper before it went to press, and for explaining more fully

than I could do the bearing of some of the results on the

theories of mimicry and of the connection between palatability and

coloration. I am also indebted to Commander J. J. Walker, R.N.,

for kindly giving me the scientific names of the Lepidoptera.

* Animal Behaviour, pp. 164-165, 1900.

PALATABILITY OF SOME BRITISH INSECTS. 813

THe EXPERIMENTS

MOLLUSCA.

(SLues.)

Large Black Slug (Arion ater).

Sept. 24,1910. Two taken and eagerly eaten bytwo Meerkats,

who wiped them down with their paws and rubbed them in the

sand apparently to remove the slime. -

Oct. 26, 1909. One given to Black-winged Grackle was

eaten.

One (larger specimen) given to the same bird was abandoned ;

offered to Sulphury Tyrant, but the bird would not touch it;

offered to Sun-Bittern, was pecked, but not eaten ; carried to a

perch by Harmonious Shrike-Thrush but was soon dropped ;

pecked and shaken about, and much hammered by Abbot’s Rail,

which managed to break the skin of the slug and getting at the

inside ate a large portion, but would not eat the outside.

One taken by Dial Bird which persevered for a long time,

hammering and wiping it in the sand; he was then driven off by

Black-chinned Laughing Thrush, which held the slug in his foot

and ate little pieces of the inside after breaking the skin, but left

the bulk of it.

Dial Bird tried another, but gave it up.

Common Hangnest took one, but left it after a few pecks.

Two offered to Kagu, a kind of Crane or large Rail, were

swallowed entire with very little delay.

One tried by Black-tailed Water-hen which, however, gave it

up; the same specimen given to Leach’s Laughing Kingfisher was

ultimately swallowed entire after being dropped many times.

Arion hortensis.

(Olive-brown Slug with orange-coloured foot.)

Oct. 26, 1909. One eaten by Yellow crowned Hangnest.

T'wo eaten by Dial Bird.

Two refused by Harmonious Shrike-Thrush.

One twice taken from my hand by Harmonious Shrike-Thrush

and dropped both times; but after taking it the third time the

bird ate it.

LTimax maximus. -

Oct. 26, 1909. One given to Harmonious Shrike-Thrush was

tried, but abandoned after one or two pecks. It was then taken

and eaten by a Black-winged Grackle after a great deal of wiping

of the bill.

Sept. 24, 1910. Two specimens tasted by Green Lizard, and

Black-spotted Lizard, but not eaten. The Lizards apparently

disliked the slime, because they wiped their mouths on the stones

after tasting. Both eaten without delay by Glass Snake.

Proc. Zoou, Soc.—1911, No, LVI. 56

814 MR. R. I. POCOCK ON THE

Limax agrestis.

Sept. 24,1910. One eaten after a good deal of pecking about

in the sand by White-crested Jay-Thrush.

Two eaten by a Shama.

One eaten by Kagu.

Tasted but rejected by Fantailed Flycatcher.

Tasted on two occasions by Hoopoe but rejected.

Tasted by Red-vented Bulbul but rejected.

Tasted but rejected by Yellow Hangnest.

Two taken, but not eaten, by Harmonious Shrike-Thrush.

Two taken, but not eaten, by Cuban Mocking Bird,

Limax arborum.

Sept. 24,1910. Four eagerly eaten by Wall Lizards, which

wiped their mouths to remove the slime after swallowing them.

Milax sowerbyt.

Sept. 24,1910. One taken and pecked and wiped about in the

sand for a long time by Indian Dial Bird, which finally left it.

Another specimen was eagerly taken by Sulphury Tyrant, which

after pecking and crunching it in his beak, and banging it from

side to side against a ledge, exactly as Laughing and other

Kingfishers do, finally swallowed it whole.

ARACHNIDA.

OPILIONES (Long-legged Spiders or Harvestmen).

Phalangium sp. ?

Sept. 1910. One (immature) tasted but immediately rejected

by Pekin Robin; the same specimen then taken and eaten by hen

Scarlet Tanager.

One (immature) put into cage with several Curassows was tasted

in turn by specimens of Yarrell’s and the Globose, and ultimately

eaten by one of the Globose Curassows, when crushed beyond all

recognition.

I was led to suppose these Arachnida would prove on experiment

to be unpalatable owing to their possessing a pair of glands, one

on each side of the dorsal area of the carapace, which are known

to secrete an odorous fluid. As elsewhere recorded *, I have seen

a Mason Wasp, hunting Spiders, run down a specimen of Phal-

angiwm, but turn aside and let it go unhurt the moment he

touched it with his antenne. More experiments with birds and

lizards are required fully to substantiate my belief ; but the refusal

of the Pekin Robin to eat the Phalangiwm is very significant, and

it is quite evident that the Arachnid was not to the liking of the

Curassows.

* Journ, Linn. Soc., Zool, xxx. p. 268, 1909,

PALATABILITY OF SOME BRITISH INSECTS. 815

INSECTA.

Order LEPIDOPTERA.

Butterflies.

Group PIERIN#.

THe SmMaLt Wuire (Pieris rape).

July 31,1909. One male (dead) given to Capuchin (Cebus sp. a)

was taken at once, and eaten without being removed from the

mouth for inspection. This specimen, given with Huchloé carda-

mines (see p. 820), was used as a check upon the behaviour of the

monkey towards Huchelia jacobec and Melitea artemis (pp. 825

and 832).

Sept. 6, 1910. One offered to a Red-handed Marmoset was

inspected, but not touched; but was eagerly taken and eaten by

another animal of the same species. This Marmoset then ate a

specimen of Perarge megera, his behaviour suggesting that the

two butterflies were equally palatable to him.

May 26,1909. One chased at once by Shrike-Thrush and Dial

Bird, but evaded them and escaped through the partition into

next cage, where it was promptly caught on the wing by a

Fantailed Flycatcher and eaten.

One caught at once on wing by Great Tit and eaten.

Aug. 21 to 27, 1910. One greedily eaten by cock Silver

Pheasant.

One let loose in aviary skilfully dodged the swoop both of a

Shama and a Wood-Swallow, and escaped.

One given to Dial Bird, which took it from my hands and

damaged it by the peck so that it was unable to fly away. Again

and again he pecked the butterfly as it fluttered about on the

ground, but would not hold it. Ultimately it escaped under the

partition into the next aviary, where it was pounced upon by a

Weaver, which held it in his foot and ate it, leaving the wings.

Sept. 6,1910. One taken by Masked Wood-Swallow and eaten

after much delay and pecking. The bird evidently was not very

keen on the insect; but he would not allow any other bird to

take it from him. He did not once shake his head or wipe his

beak as if there was any distasteful flavour.

One female taken and eaten by Ludwig’s Bustard.

Sept. 7, 1910. One male and one female taken and eaten

eagerly and with equal avidity by Green Lizard.

Larva of the Small White (P. rape), fed on cabbage.

_ Sept. 21,1910. One tasted but rejected by Yarrell’s Curassow

and Globose Curassow.

THE GREEN-VEINED WHITE (Pieris napi).

July 31, 1909, One offered to White-tailed Mongoose, to three

06*

816 MR. R. I. POCOCK ON THE

Meerkats and to two Banded Mongooses. All rejected it after

smelling it except the second Banded Mongoose, which took

it with his paw, rubbed it in the sawdust, but would not

eat it.

N.B.—The forceps with which this butterfly was offered had

been previously used for Ocypus olens, Carabus violaceus, Ptero-

stichus niger and P. madidus, and some Timarche as well as

Coccinella, and probably the scent of these beetles was adhering to

the steel.

May 26,1909. One taken and eaten by Dial Bird, by Har-

monious Shrike-Thrush, and by Blue Rock-Thrush.

July 26, 1909. One male given to Silver Pheasant, was taken

from my fingers and swallowed instantly without being first

deposited on the ground.

One female given to same bird was treated in exactly the same

way.

These two I used as checks upon two specimens of Melanargia

galathea, both of which the Pheasant treated very differently,

spitting them out upon the ground after taking them from my

fingers, and pecking them about a great deal before swallowing

them (p. 827).

Aug. 21,1910. One male greedily eaten by Silver Pheasant.

This bird ate at the same time a specimen of Hpinephele jurtina,

showing an equal liking for both.

One male taken eagerly by Pekin Robin, which, after much

pecking and tasting, left the butterfly uneaten.

July 31, 1909. One male eaten at once by Brazilian Hangnest.

Sept. 20,1910. One left untouched by Fantailed Flycatcher.

Taken and tasted but left uneaten by Dial Bird. Taken by

Yellow-crowned Hangnest, which held the butterfly in his foot

against the perch, pecked off its wings and finally picked it to

pieces, and ate at all events most of it.

Note.—The Hangnests which ate these butterflies are much less

typically insectivorous in diet than the Flycatcher, the Pekin

Robin, and the Dial Bird, which refused them.

Tae Larce Waite (Pieris brassice).

Oct. 26, 1909. One taken from my hand and greedily eaten

by Lion Marmoset.

May 26, 1909. One taken at once by Syrian Bulbul and eaten ;

also by Harmonious Shrike-Thrush.

Oct. 26,1909. One taken from my hand and greedily eaten

by cock Silver Pheasant and by Honduras Turkey.

One taken by Shama and finally eaten, but not with any ap-

proach to the readiness with which he had just previously eaten

a Tortoise-shell and the #. jurtina. At one time I thought he

was going to give it up; but finally he swallowed it.

One liberated in aviary was chased up and down by three

Wood-Swallows which, however, owing to hesitancy at the moment

PALATABILITY OF SOME BRITISH iNSECTS. 817

of coming to close quarters, did not catch it. It escaped into

another compartment, and was promptly seized by the Harmonious

Shrike-Thrush, which ate it after a deal of pulling about and

tasting.

Aug. 21, 1910. Two males greedily eaten by cock Silver

Pheasant.

One male caught by Pekin Robin and eaten after some time,

the delay being caused not apparently by distastefulness, but by

the difficulty of getting rid of the wings which were left uneaten.

This bird held the insect to the perch with his foot when

pecking.

One male eagerly taken by Pearl-spotted Owl, which held it up

in one foot while pecking it. He pecked away for some time at

the thorax and wings without making much headway. He then

shifted it and pecked off the end of the abdomen. But as soon as

he got the flavour of the exposed tissues he shook his head and

repeated the shake with every taste, showing unmistakable signs

of disliking the flavour. Finally he hopped to another perch, put

the butterfly down, and after looking at it for a little time, flew

away. I thought he had given it up; but upon returning to the

cage ten minutes later the butterfly had disappeared.

One put into an aviary of Tanagers was chased by several birds

which, however, hesitated at the critical moment to catch it, as if

a little doubtful as to its nature. At last a male Scarlet Tanager

took it in his beak, but not having the instinct to use his foot to

hold it or to put it into a cranny, went on masticating it for at

least five minutes without showing any signs of dislike. He

apparently refrained from swallowing it on account of the wings.

Ultimately he was robbed by a female of the same species, which,

after getting rid of the wings, continued pecking and tasting and

shaking her head in the intervals, quite obviously not enjoying the

flavour. She managed the insect better than the male, jamming

it first into a split orange, and then between the leaves of a palm

to peck it the better. Ultimately she ate what was left of the

body.

One male offered to a hen King Bird of Paradise. She looked

at it and as soon as she saw the legs move took it, but dropped it

at once to the bottom of the cage. After careful and long in-

spection, she pecked it once or twice, but showed no eagerness to

eat it. Ithen gave the same insect to a Larger Hill Mynah,

which soon swallowed it, wings and all.

One male taken and eaten at once by Ludwig’s Bustard.

One male offered to Fantailed Flycatcher, but he would not

touch it. Taken and tasted by Dial Bird, but left uneaten. Also

taken and tasted by Black-winged Grackle, and left and sub-

sequently refused twice. Quickly eaten up by Harmonious Shrike-

Thrush.

Sept. 18 to 20, 1910. One caught on wing by Fantailed Fly-

catcher, which had just eaten a ‘Blue.’ He carried it to a

window-sill, but after one or two pecks left it. Once or twice the

818 MR. R. I. POCOCK ON THE

bird, after waiting a short while, tried it again, but finally left it

alone.

It was then taken by a Dial Bird, which, after pecking it about

for a short time, was robbed by the Sulphury Tyrant. The latter,

after tasting it, left it alone. I then gave the remainder of the

insect, consisting only of the thorax and wings, to a Yellow-

crowned Hangnest, which took it toa perch, and holding it in one

foot gradually pecked away the wings and dropped them, and

then pecked the thorax to pieces, eating little bits of it and

dropping others.

Pupa of the Lance Wurte (Pieris brassice).

Oct. 26, 1909. One offered to the Dial Bird which had fifteen

minutes previously eaten the larva, but he would not touch it.

Offered to Yellow-crowned Hangnest which had tasted and

dropped the larva. He looked at it but would not touch it.

Given to Harmonious Shrike-Thrush, which behaved just as

the Dial Bird had behaved with the larva, pecking it and dropping

it repeatedly to shake his head. He was then robbed of it by a

Common Mocking Bird, which, however, dropped it in the grass

from the perch, and made no attempt to recover it.

One offered to a Black-winged Grackle, a Javan Pied Mynah, a

Fantailed Flycatcher, and a Sulphury Tyrant, all of which tasted it

once, but not asecond time. A Common Mocking Bird persevered

a little longer, but finally dropped it and made no effort to pick

it up again. Given to Harmonious Shrike-Thrush, was eaten

without much hesitation.

Larva of the Lares Wurte (Pieris brassice).

Food not recorded.

Oct. 26, 1909. One taken by Yellow-crowned Hangnest, but

soon dropped. Pounced upon by Dial Bird, which after many

trials, pecking it and shaking his head after every taste, at last

swallowed it; but he was evidently very uneasy for some twenty

minutes afterwards, periodically shaking his head and opening his

mouth and straining as if trying to vomit something nauseous.

Larvee of the same fed on 7ropeolum (so-called Nasturtium).

Sept. 13, 1910. Three eaten readily by Silver Pheasant and

Reeves’s Pheasant.

A small one given to Pekin Robin, which obviously did not

like the flavour. He pecked it about in the sand for a long time,

vigorously shaking his head after each taste. Ultimately, how-

ever, he ate it. I then gave him asa test the larva of a Noctua

(see p. 835), which he also took and very soon swallowed entire

without once shaking his head or evincing any sign of dislike.

He then took a second and larger brassice-larva, treating it just

as he did the first, but tackled it with still greater reluctance,

PALATABILITY OF SOME BRITISH INSECTS. 819

allowing himself to be robbed of half of it by another bird of the

same species. The two finally finished it between them.

One given to a Shama, which after pecking and tasting it for

a long time, with much headshaking, left it. It was then tasted

by a Wood-Swallow, which left it after one peck. The Shama

then tried it again, but left it. ‘Then a Red-vented Bulbul took

it, but soon dropped it. The Shama then tried it again and

ended by eating it. This Shama was the same bird that ate the

Coccinella 7-punctata (p. 846).

One given to Kagu, which after several attempts left it; and

immediately afterwards greedily ate the larva of a Voctua (p. 835).

This same Kagu ate 7imarcha tenebricosa.

One taken by Green Hangnest, which at the time was greedily

eating mealworms. ‘The bird finally ate it, but evidently did not

much like it, putting it down several times, and wiping it in the

sand.

One given to Pearl-spotted Owl, which dropped it at once.

One given to Butcher Crow, which dropped it directly; but

afterwards picked it up and swallowed it whole. Immediately

afterwards, however, he vomited it up and left it on the bottom

of the cage.

One smelt, but not touched by Common Marmoset, and by

Capuchin.

One eagerly eaten by Meerkat.

Sept. 21, 1910. Larve of the same, fed on cabbage (Brassice).

Taken and eaten by :—

Elliot’s Pheasant, Reeves’s Pheasant, and Silver Pheasant.

Vulturine Guinea Fowl. Crested Guinea Fowl. Ludwig’s

Bustard. Vigors’s Bustard. S. American Thicknee.

Cariama. Crested Curassow. Nigerian Ground - Horn-

bill.

Also by Meerkats and Banded Mongoose.

Tasted but rejected by:—Shama, Red-vented Bulbul, Green

Hangnest, Black Hornbill, Elate Hornbill, Trumpeter, Yarrell’s

Curassow, Globose Curassow, Crested Curassow, and Red-

tailed Guan.

Notes.—The nature of the food of the larve did not appear to

affect their taste. The Green Hangnest, it is true, refused larve

fed on cabbage, having a week earlier eaten one fed on Z’ropeolum,

but the bird was not eager for the latter, and I do not think this

refusal of the former can be taken as strong evidence that he

found them more unpalatable than the others. It is interesting

that the Pheasants and Guinea Fowl, that is to say, Asiatic and

African Gallinaceous birds, ate the larvee eagerly, while the

S. American Curassows and Guans, with the exception of one

Crested Curassow, refused them after many trials, and much head-

shaking. One Curassow eagerly ate the larva of the Woctua

(p. 835) after refusing that of P. brassice.

820 MR. R. I. POCOCK ON THE

THE ORANGE-TIP (Huchloé cardamines).

July 31, 1909. One male given to Cebus (sp. «) was seized at

once and stuffed into his mouth. He took it out, looked at it,

smelt it, then ate it without hesitation.

This was a check experiment upon the behaviour of the monkey

towards Huchelia jacobee and Melitea artemis. He showed much

greater alacrity in eating the cardamines than either of the others.

A Pieris rape given at the same time he ate without removing

it from his mouth.

May 26, 1909. One male taken by the Harmonious Shrike-

Thrush after a few moments’ inspection and eaten entire, wings

and all, with much less delay in the way of pecking and scraping

on the soil than the same bird displayed when dealing with

M. artemis and A. euphrosyne. Tested by this bird, H. cardamines

appeared to be more palatable ; but it is possible, though I do not

think probable, that he ate it with less delay because he had just

previously been robbed of the specimen of Argynnis euphrosyne

by not swallowing it at once.

Group NyYMPHALINA,

THE SMALL ToRTOISE-SHELL (Vanessa urtice).

Oct. 26, 1909. One taken and eaten by Shama which had just

previously eaten Hpinephele jurtina.

Hoopoe, Black-winged Grackle, and Harmonious Shrike-Thrush

very eager to take one, but it was secured by the Grackle,

which, however, was robbed by the Shrike-Thrush, the latter

eating the butterfly in about half a minute without any signs of

dislike such as shaking his head or wiping his beak.

Sept. 7, 1910. One taken and greedily swallowed, wings and

all, by Dial Bird.

Sept. 18, 1910. One caught on wing by Fantailed Flycatcher,

who carried it to a perch, but after a few tastes and pecks

dropped it to the ground. Whether this was done intentionally

or accidentally I cannot say, but the bird made no attempt to

follow up the insect. I then gave it to a Dial Bird, which, after

pecking it for a short time, was driven off by a Sulphury Tyrant.

This bird, however, did not touch the butterfly. I then offered

it to a Bulbul and a Yellow-crowned Hangnest; but neither

touched it. I then offered it again to the Dial Bird, who finished

it, but with no show of appetite. I am unable to say whether

the indifference shown by the birds to this butterfly was due to

its being distasteful or to the experiment being made at 5 P.m.,

when the birds had been feeding off and on through the day.

Pupa of Vanessa urtice.

June 24, 1909. One placed on a branch near a Shama was

taken after a good deal of preliminary inspection but was soon

PALATABILITY OF SOME BRITISH INSECTS. 82)

flicked away and fell to the ground. The bird made no attempt

to recover it. I then again put it on the branch by his side, and

on this occasion he pecked at the little stem to which the pupa

was attached. A hen Black Tanager was the next to try it. She

broke the shell and getting the taste flew away with the pupa and,

T think, ate it. At all events she flew up to the top of some

brickwork where I could not see her clearly, and presently came

down again without the pupa; and on going upa ladder to look

for the pupa, I could find no trace of it.

One offered to Syrian Bulbul was taken after some scrutiny.

The bird flew away with it and pecked it, but seemed greatly

bothered and puzzled by the tightness with which it adhered to

the twig. He was unable to detach it from the twig, and finally

left it. I then offered it to a Fantailed Flycatcher ; but

could not induce this bird to touch it, although he scrutinised

it carefully and was hovering round me the while, apparently

remembering that on previous occasions I had given him butter-

flies. I then gave it to the Harmonious Shrike-Thrush, which

took it, pecked away at it until he broke off the tail-end and ate

it. He then pecked off another piece and ate it, showing no sign

of dislike. He then left the larger piece; but soon returned,

broke it up, and finally ate it piecemeal.

From watching the behaviour of these birds, I should say that

these pupe are unpalatable only to the extent afforded by the hard-

ness and toughness of the chitinous integument. The birds that

tasted them after breaking the exoskeleton, showed no signs of

disliking the flavours. Those that took them—and the Flycatcher

could not be induced even to attempt it—did so after scrutinising

them in a way that suggested doubt as to their belonging to the

category of eatable things. They did not appear to me to know

what they were; and none of the many insectivorous birds in the

aviary showed the least sign of eagerness when I first put the

pupa on a perch, waiting to see which would be the first to come

down. It was only when I placed it about a couple of inches

from the Shama, a tame and fearless bird, that he took it. The

Tanager came, and after her the Bulbul, when they had seen the

Shama’s attempt, or at all events after the Shama had first tackled

it. These birds are accustomed to visitors and keepers bringing

food into the aviary; and I think it probable that the Shama

was induced to peck at the pupa merely because it was definitely

offered to him.

I suspect that this pupa is protected in the first place by its

likeness to things inanimate, and in the second place by the

toughness of its integument which does not readily yield to a

peck, and is quite in keeping with the general impression of life-

lessness suggested by the colour, shape, and immobility of the

whole pupa. J may add that I did not see the pupze move when

pecked by birds, although they did so when handled by myself.

829 MR. R. I, POCOCK ON THE

Young larve of Vanessa wrtice.

June 24, 1909. One eaten without hesitation by Brazilian

Hangnest, and by Common Mocking Bird; two by Shama; two

by Orange-headed Ground-Thrush, and one by Harmonious

Shrike-Thrush.

Two taken and tasted but whisked away by Larger Hill Mynah.

One taken and tasted but dropped by North American Cat-bird,

which refused to touch a second.

~ One pecked and tasted many times, but finally rejected, by

Fantailed Flycatcher.

THE Peacock (Vanessa 20).

May 26, 1909. One fluttered to ground and rested with wings

closed. A Fantailed Flycatcher flew down to inspect and was

preparing to peck, when the butterfly opened its wings and moved

them slowly up and down. The transformation seemed to dis-

concert the bird, which made no attempt to peck, but danced

round the insect at a distance of about three inches. A Shama

and, another Flycatcher, which joined the first, behaved in the same

way. A Syrian Bulbul then flew down and drove the three away.

After inspecting the butterfly for about half a minute, he pecked

the ocellus of the anterior wing of the left side; the second peck

struck the ocellus of the anterior wing of the right side ; the third

the ocellus of the posterior wing of the left side, tearing a piece

out. He was then driven away by a Sun-Bittern, which looked

at the butterfly for some two minutes, but made no attempt to

peck it, although it excited his interest. I then removed the

Bittern ; and the Bulbul returned at once, seized the butterfly by

the head and thorax, flew away with it, and devoured it.

One fell to floor of aviary with wings closed, and was at once

seized by Syrian Bulbul, before its wings opened, and was carried

away and eaten. A second Bulbul of this species pursued the

tirst; but I do not know which of these two birds was the one

that ate the zo first introduced.

The two features of interest in the first experiment with this

species were, first, the manifest disconcertedness of the three birds

by the sudden display of colour and the slowly waving wings of

io (my wife, who was with me, said at once, ‘‘They are afraid

of its eyes”); and secondly, the consecutive pecking of three of

the ocelli by the Bulbul. It can hardly have been by accident

that the ocelli were accurately struck three times running.

Aug. 21, 1910. A specimen let loose in aviary was chased by

a number of Tanagers and other small birds and was caught by a

Scarlet Tanager. The latter, however, was robbed by a Pekin

Robin, which ate the insect without showing any signs of dislike,

the delay of five minutes in finishing it off being caused by the

difficulty of managing the wings which the bird ultimately broke

off and left uneaten.

PALATABILITY OF SOME BRITISH INSECTS. 823

THE Rep Apirat (Pyrameis atalanta).

Aug. 21, 1910. One taken and eaten greedily by Lion

Marmoset.

One pursued by Shama, which grabbed it by the hind wing and

thereby lost the butterfly, which flew away and escaped through

the wires of the aviary.

Tue Patnrep Lavy (Pyrameis cardui).

Aug. 27,1910. One given to Pearl-spotted Owl was taken at

once and swallowed entire after a little preliminary pecking.

This was a test experiment to ascertain the meaning of the

bird’s behaviour towards Pieris brassicw (see p. 817).

Araschnia levana. Late summer form prorsa.

July 8,1911. One given to Harmonious Shrike-Thrush, an

Australian bird, was taken at once, but after being pecked and

tasted for some little time, was rejected. The remains were then

greedily eaten by a Wood-Thrush, from North America. A fresh

specimen given to this same Wood-Thrush was just as readily

swallowed ; but the Shrike-Thrush upon taking another, treated

it as before, wiped it in the sand, shook his head, and allowed

himself to be robbed by a Black-chinned Laughing Thrush, which

ate it and another without hesitation.

One taken and eaten, but very slowly and with much pecking

about, by a Hoopoe, which, after swallowing the last particle,

appeared to try to vomit it back but without success.

A Blue Rock-Thrush and a Common Rock-Thrush, both

European birds, each ate one greedily.

One pecked and tasted for some little time by Orange-headed

Ground-Thrush, which obviously did not care for the favour, and

allowed himself to be robbed by the Blue Rock-Thrush mentioned

above.

One liberated in aviary dodged the pursuit of a Shama and a

Sibia with great skill, and escaped.

One given to Shama was pecked and tasted for some time, but

the bird allowed himself to be robbed by a Wood-Swallow, which,

after much pecking, swallowed the butterfly.

This performance was repeated exactly when one was given to

the Sibia, the same Wood-Swallow taking it from him; but I

think the Sibia would have eaten it ultimately.

One given to Grey-headed Friar Bird, from Australia, was

taken and tasted for a long time and then dropped, given again

to the same bird, was again tasted and dropped. The remains

were then eaten without much delay by a Larger Hill Mynah.

One given to a Dial Bird was taken and after much tasting

was resolutely rejected. The remains were then given to a

824 MR. R. I. POCOCK ON THE

Sun-Bittern, which persevered for some time but finally rejected

them.

The only birds which ate the butterflies quite readily were

the two species of Rock-Thrushes, the Wood-Thrush, and the

Black-chinned Laughing Thrush. To the others they were

obviously more or less distasteful, the most significant rejection

being by the Shrike-Thrush, which on previous occasions has

eaten almost every insect offered to him.

N.B.—These experiments were made between 4 and 5 P.M.,

when the birds had been feeding throughout the day.

July 9, 1911. One eaten readily by Black-headed Lemur,

one by Meerkat, two by Common Indian Mongoose.

Three eaten readily by two Wall Lizards.

Two eaten readily by Silver Pheasant, and one fairly readily

by Mantchurian Crossoptilon (Pheasant).

One given to White-eared Scops Owl was taken at once but

dropped as soon as tasted.

Experiment repeated with same result.

Experiment repeated with same result with another specimen

of the same species of Owl.

One given to Pekin Robin was taken at once, but put down

upon the ground. For fully five minutes the bird continued to

peck it and shake his head. He would neither eat it himself

nor allow the other birds to take it from him. Ultimately he

pecked it to pieces; but I cannot say whether he ate particles or

wasted them on the ground. One thing was quite clear. He did

not find the flavour to his liking.

Dark Green Frivitiary (Argynnis aglaia).

July 21, 1909. One let loose in aviary was chased by Black-

headed Sibia and Fantailed Flycatcher, but eluded both and

escaped into a crevice. This is the first butterfly I have seen

dodge the Flycatcher, which is extraordinarily adept at taking

insects on the wing. I then gave it to the Spectacled Thrush, and

he ate it after he had succeeded in shaking off its wings. The

bird was keen not to lose it, and drove away the Flycatcher

whenever he ventured near.

SILVER-WASHED F'RITILLARY (Argynnis (Dryas) paphia).

July 26,1909. One caught on wing and eaten with avidity

by Fantailed Flycatcher.

Also used as check upon Melanargia galathea which the

Flycatcher had rejected (see p. 827).

July 31, 1909. One eaten readily by Brazilian Hangnest.

PEARL-BORDERED FRITILLARY (Argynnis (Brenthis) ewphrosyne).

May 26 to 31,1909. One eaten by Silver Pheasant. For

details see under Melitea artemis (see p. 826).

PALATABILITY OF SOME BRITISH INSECTS. 825

Two specimens given respectively to Brazilian Hangnest, and to

Saturnine Mocking Bird, were eaten much more readily than

were specimens of J/. artenvis offered to the same birds (see under

Melitceea artemis).

One female taken by Harmonious Shrike-Thrush, but not eaten

readily. While this bird was pecking the butterfly and wiping it

on the gravel, he was robbed of it by a Red-vented Bulbul; the

latter was in turn robbed of half of it by a North American

Mocking Bird. The two finished it between them.

SMALL PEARL-BORDERED FRittLuary (Argynmnis (Brenthis) selene).

May 31,1909. One taken and eaten by Capuchin, but without

relish.

One taken and eaten by Capuchin (Cebus sp. c), with obvious

avidity.

GREASY FRITILLARY (Melitcea aurinia or artemis).

May 26 to 31, 1909. One male given to same specimen of

Cebus that took the Huchelia jacobee five minutes previously.

He behaved in exactly the same way towards it. Stuffed it into

his mouth, but the moment he got the flavour or the feel, took it

out in his hands, pulled it to pieces, cautiously tasted it, and then

ate it, but with no great show of satisfaction.

One taken and eaten by Capuchin (Cebus sp. 6), but with great

hesitation and no particular signs of relish. This monkey also

ate one Cenonympha pamphilus, one Argynnis selene, and one

Thanaos tages; but treated them all in the same way, evidently

not caring much for any of them. In this particular he showed a

marked contrast to the two other examples of Cebus, sp. a and c,

used for these experiments.

One male offered to Meerkat, taken and eaten at once. Kager

for more.

One male offered to Capuchin, taken and eaten at once. Eager

for more.

One male offered to White-handed Lemur, which after carefully

smelling it, refused it.

Same one offered to Crowned Lemurand White-fronted Lemur,

was smelt and refused in the same way.

Offered to Black Lemur, was smelt, then carefully taken into

the mouth, but was then pulled out with the hand; then again

tasted, but rejected as if distasteful, the tongue being rapidly

protruded and drawn back through the front teeth as if to scrape

off something unpleasant, perhaps scales.

One offered to Diana Monkey, was taken and eaten piecemeal,

apparently with relish.

The mammals above mentioned had not been fed, and were

without exception hungry.

826 MR. R. I. POCOCK ON THE

One taken by Brazilian Hangnest, which pecked at it, ate a few

pieces as if testing its flavour, then let it fall from the perch to the

ground, and left it there.

One taken by Saturnine Mocking Bird, which shook it about,

pecked it, ate a fragment or two, then left it.

One taken by Brazilian Hangnest, which pecked it several

times, and finally ate it. The Mocking Bird then returned, and

after many trials finished off the remains of the first specimen

that had been left by the Hangnest and of the second that had

been left by himself. It was quite evident that neither of these

birds found much satisfaction in eating these butterflies.

One female liberated in aviary, caught on wing by Garrulous

Honey-eater, and eaten without delay.

One female taken by Blue Rock-Thrush, but left on the ground

after being pecked. Suspecting that his leaving it was due to my

propinquity, | moved away and told the keeper to throw it to him,

He then caught it on the wing, and ate it. He then came close

to me on a perch and eagerly took another specimen (male) from

my hand, then a third (female), and ate both greedily.

Two given to Silver Pheasant were taken and eaten, but with a

great deal of pecking and tasting. Comparing this bird’s behaviour

towards them with his manner of eating Pieris napi and rape, |

am quite sure he found them to a certain extent unpalatable. I

thought at first that he merely disliked the wings. To test this

I gave him immediately afterwards a specimen of napi. He took

it from my hand and put it on the ground; then tasted it, and

without more ado swallowed it. I then gave him a specimen of

rape. He took it from me, and without putting it on the ground

ate it up. I then gave him a specimen of Perarge megera, which

flew into a bush. He went after it, found it, caught it with the

dexterity of a ‘practised hand,’ but treated it exactly as he

treated the artemis, pecking and whisking it about, ultimately

after much delay eating it piecemeal, but with what might be

described as a very dubious air. He behaved in a precisely

similar manner towards an example of Argyniis ewphrosyne.

Tam convinced that no one who had seen this Pheasant eat

these five butterflies, could have doubted for a single moment that

he found the ‘ Whites’ pleasant to taste, and the ‘ Fritillaries’ not

altogether to his liking.

One male offered to Larger Hill Mynah was taken and eaten,

but with no great relish, being frequently dropped and picked up

again, and scraped in the sand.

One male offered to Levaillant’s Barbet, which took it and be-

haved towards it in exactly the same way as the Mynah. The

birds appeared to dislike the wings, and to want to get rid of

them.

One male offered to Fantailed Flycatcher, which after a little

inspection pecked it and took it, but was robbed by a Syrian

Bulbul, which ate it.

Two males taken and eaten by Shama,

PALATABILITY OF SOME BRITISH INSECTS. 827

One male taken and eaten by Cape Robin-Chat.

One male taken and eaten by Indian Orange-headed Ground-

Thrush, after being pecked and rejected by Hoopoe.

One male taken and eaten by Harmonious Shrike-Thrush.

One female taken and eaten after a great deal of pecking and

delay by Indian Black-headed Sibia, which was chased for it by a

Syrian Bulbul.

One female taken and eaten, after a few moments’ inspection

and biting at the wings before the position of the body was found,

by a Sand Lizard. A Dugés’s Lizard came up while the butterfly

was being chewed, and after tasting it once or twice, attacked the

Sand Lizard to make him relinquish his hold.

One male taken by the same Sand Lizard after he had finished

the first specimen. I then made him drop it; and offered it toa

Wall Lizard, which took it without delay and swallowed it.

Group SATYRIN#.

THe MarsieD Waite (Jelanargia galathea).

July 24,1909. Asa check I first of all offered a specimen of

P. napi to the cock Silver Pheasant. He took it from my fingers,

and without hesitation swallowed it and turned eagerly for more.

I then gave him a galathea, which he just as eagerly took, but

promptly lowered his head to the ground and spat it out. He

persevered with it, however, and after a little pecking and shaking,

ate it. Ithen tried him with another napi. He took it and

swallowed it at once, not hesitating for a single moment, exactly

as he had done with the first one. Then I gave him another

galathea, which he took but immediately put out of his beak upon

the ground ; but after some pecking and tasting he swallowed it.

I consider this bird to have rather a refined taste for insects:

and I can now tell tolerably accurately by his behaviour whether

he likes one or not. And Iam quite sure that he found napi

very palatable and galathea not so.

I then let a galathea loose in the aviary, and it was promptly

caught on the wing by a Fantailed Flycatcher, which flew with

it to the ground, and after pecking, pulling and shaking it about

for a minute or so, gave it up and took no further notice of

it. As a check I then tried him with Aphantopus hyperanthus,

which he caught in the same way, and very quickly demolished.

I then gave him another galathea, which he caught and pecked

and shook for some little time; but he would not eat it. Asa

further check I gave him Argynnis paphia, which he caught and

disposed of as quickly as he had disposed of the hyperanthus.

I noticed that some of these galathea had darker spots below

than the others. Thinking that perhaps this might be a sexual

difference, I gave one of each kind to the Pheasant and to the

Flycatcher; but the birds behaved in exactly the same way

_towards them.

One caught and eaten by Dial Bird; also by Orange-headed

828 MR. R. I. POCOCK ON THE

Ground-Thrush, and by White-cheeked Bulbul. The latter was

robbed by the Harmonious Shrike-Thrush ; but recovered the

butterfly and ate it.

One given to Sulphury Tyrant, who pecked it and shook it for

a long time, then allowed the Shama to take it from him.

The Shama ate it. This was a galathea with lighter spots

below.

One given tothe same Shama was also eaten, This was a darker

spotted specimen.

The specimen above alluded to that was rejected by the

Flycatcher, was eaten by a North American Cat-bird (Thrush).

One offered to Australian Bustard, was taken from my hand

and swallowed at once.

One given to Meerkat, was taken and eaten without hesitation.

One given to Capuchin, which by his rejection of the Telephorid

peetle (p. 840) had shown himself to be more particular in taste

than some others of his species, was eaten, but by no means

greedily.

With the exception of the Australian Bustard none of the

birds that ate the galathea did so with great alacrity. Swallowing

them was in all cases preceded by a varying amount of flicking

and shaking and pecking. When I began my experiments I

thought this behaviour was due to a wish to get rid of the wings ;

but I am now doubtful about this, and believe that in many cases

at all events it indicates dislike of the taste. When a butterfly is

really to the liking of a bird, he disposes of the insect as fast as he

can, without paying much attention to the wings. This struck

me to-day particularly in the case of the Silver Pheasant when

eating the api, and of the Flycatcher when eating the hyper-

anthus and the paphia. The paphia especially was a large-winged

butterfly for so small a bird; and yet he swallowed it, wings and

all, in a few seconds.

Tue Meapow Brown (Lpinephele jurtina x janira).

July 21, 1909. One female eaten at once by Lion Marmoset,

which had previously refused to taste the malacoderm beetle

(Rhagonyche) and the Saw-fly (Allantus arenatus).

One female taken from my hand by Spectacled Thrush, but

made his escape. Caught on wing by Fantailed Flycatcher and

eaten at once.

One female eaten at once by Common Pheasant.

July 31, 1909. One male caught on wing by Black-headed

Sibia and eaten at once; another (female) caught on wing and

eaten without delay by Fantailed Flycatcher.

Aug. 21,1910. One male greedily eaten by Silver Pheasant.

This bird at the same time ate with equal avidity a male specimen

of Pieris napi. Her behaviour indicated no difference of taste

between the two butterflies.

Oct. 26, 1909. One female taken and eaten fairly readily by

Shama.

PALATABILITY OF SOME BRITISH INSECTS. 829

Tue Larce Hearn or GATEKEEPER (Hpinephele tithonus).

July 31, 1909. One eaten at once by Brazilian Hangnest.

Aug. 25, 1910. One caught and quickly swallowed entire by

Pekin Robin.

One male taken and swallowed entire with scarcely any delay

by Pearl. spotted Owl.

Sept. 20, 1910. One caught on wing and eaten without delay

by Fantailed Flycatcher. This bird would not touch two White

Butterflies (P. brassice and napi), offered one just before and one

just after it took tithonus.

THe Rinewer (Aphantopus hyperanthus).

July 26, 1909. One caught on wing and eaten with ail speed

by Fantailed Flycatcher.

I used this specimen as a check upon galathea, which the bird

had just refused to eat after catching it and pecking it about for

some time.

One gobbled up at once by Silver Pheasant.

Tue Smatt Hearu (Cenonympha pamphilus).

May 26,1909. ‘Two taken and eaten at once by Fantailed

Flycatcher.

One taken by Fantailed Flycatcher which was at once chased

by Syrian Bulbul.

One seized by Orange-headed Thrush, which after carrying it

about gave it through the bars of the partition to another bird of

the same species. This was deprived of it by a Bower Bird, which

earried it about, perhaps as a possible ornament, since he made

no attempt to eat it.

THe WALL Burrerrty (Perarge megera).

May 31,1909. One taken and eaten by Capuchin (Cebus sp. b),

but without apparent liking (see under Melitea artemis).

One taken and eaten with avidity by Capuchin (Cebus sp. c).

Aug, 25, 1910. Kagerly taken and eaten by Red-handed

Marmoset.

May 31, 1909. One eaten by Silver Pheasant (see also under

Melitea artemis).

Aug. 25,1910. Two (male and female) caught and greedily

eaten, wings and all, by Pekin Robin.

Sept. 5 to 7,1910. One female taken at once’ by Ludwig’s

Bustard, which, however, let it escape. It was caught on the

wing and quickly eaten by a Larger Hill Mynah. Two more

specimens (female) eaten by Black-winged Grackle and by

Dial Bird.

Proc. Zoou, Soc.—1911, No. LVTI. 57

830 MR. R. I. POCOCK ON THE

THE GRravuine (Satyrus semele).

July 31, 1909. One caught on wing by Fantailed Flycatcher

and eaten with all speed.

Group Lyc#NID&.

Common Buus (Lycena icarus).

Oct. 26, 1909. One given to Shama, but it avoided him and flew

through into the next compartment, where it was captured smartly

by a White-browed Wood-Swallow, and eaten as soon as the bird

could get peace from the pursuit of two other Wood-Swallows in

the same compartment.

N.B.—These Wood-Swallows were desperately keen to get

the butterflies with which they saw me feeding the Shama in the

next compartment.

One male let loose in aviary containing Fantailed Flycatcher,

which I should describe as an expert butterfly-catcher. But the

Blue dodged him again and again, and got through into the next

compartment. Here again it avoided the swoop of one or two

birds whose identity I did not detect in my intentness in keeping

my eye on the butterfly. The latter then passed through to a

third compartment and settled on some yellow painted boarding,

which it did not match, and on which it was caught by a Brazilian

Hangnest, and quickly eaten.

One male caught deftly by Masked Wood-Swallow, which after

prolonged pecking and tasting, swallowed the body, having got

rid of the wings. This bird used its foot to hold the insect down.

One male caught by Pekin Robin and ultimately swallowed

entire; but the bird put the insect down many times before

swallowing it.

The behaviour of these two birds suggested that this ‘ Blue’

was not very palatable. Its size offered no obstacle to its being

swallowed at once ; but both birds delayed over the meal.

Two (male and female) given in succession to Pearl-spotted Owl

were taken and swallowed entire without delay.

Two (male and female) let loose in aviary were captured and

quickly swallowed entire by Pekin Robin.

Troe Brown Arcus (Lycena astrarche).

Sept. 18, 1910. One let loose in aviary was taken by a Cayenne

Tanager, which was quickly robbed by a Pekin Robin. The latter

ultimately swallowed it entire, after putting it down several times

before finishing it off.

One caught on wing and eaten at once by Fantailed Flycatcher.

One caught and eaten at once by Dial Bird.

THE SMALL Copper (Chrysophanus phleas).

Aug. 25,1910. One caught and swallowed quickly, wings and

all, by Pekin Robin.

PALATABILITY OF SOME BRITISH INSECTS. 831

Group NEMEOBIIN&.

THE Duke or Burcunpy (Vemeobius lucina).

June 15,1909. One given to Brazilian Hangnest was taken

and swallowed without any hesitation.

One given to Saturnine Mocking Bird was taken at once, but

not eaten eagerly. While she was pecking it about a Lesser Hill

Mynah flew up and took it away, but was in turn deprived of it

by the Mocking Bird, which then swallowed it quickly.

Two given to Silver Pheasant, which ate them with the same

eagerness as it had previously shown when tried with the

‘Whites.’

Group HESPERIIDA,

Tue Dryey Sxiprer (7hanaos tages).

May 31, 1909. One taken and greedily eaten by Dent’s

Monkey.

One taken and eaten cautiously by Capuchin (Cebus sp. 6). See

under J/. artemis.

THe Larce SKIPPER (Argiades sylvanus).

July 21 to 31, 1909. One caught and eaten at once by

Fantailed Flycatcher, and one by Brazilian Hangnest.

Moths.

Larva of Goat-Moru (Cossus ligniperda).

Sept. 20,1909. One taken first of all by Dial Bird, which after

a short time was driven off by Sulphury Tyrant. Both, after

pecking it, left it practically uninjured on the ground. It was

then taken by the Harmonious Shrike-Thrush. He kept it for

some time, pecking it about and was eager to prevent other birds

getting it; but was finally beaten in a ‘tug-of-war’ for it by a

Common Hangnest, which carried the grub to a bush, held it

against a branch with his foot, and pecked away for five minutes,

then voluntarily dropped it. It was then taken by a Dial Bird,

which persevered for a long time, pausing frequently between the

pecks, opening and shutting and wiping his beak. He was then

deprived of it by a Black-chinned Laughing Thrush, which kept

it for ten minutes, pecking and whisking it about without making

any visible impression on the skin. The head, however, was by

this time gone, and the bird pulled some soft tissues out of the

end and ate them. I then gave it to a Green Hangnest, but

after tasting it he let it drop and took no further notice of it.

I then gave it toa Leach’s Laughing Kingfisher, which after a

little delay swallowed it whole.

ihe

832 MR. R. I. POCOCK ON THE

Larva of the Lackny (Clisiccampa neustria).

Sept. 20, 1909. Seized at once by a Shama which flew away

with it, holding it by the head; but while he was adjusting it

for eating, the female Black Tanager grabbed the other end and

being victorious in the tug that ensued, carried away the larva

and, after a good deal of pecking, ate it.

Srx-sporteD Burnet (Anthrocera (Zygena) filipendule).

July 31, 1909. One placed on a branch was immediately seized,

but flicked away by a Black-headed Sibia, which made no attempt

to follow it up but flew away, shook his head once or twice, and

wiped his beak.

Sulphury Tyrant then pecked it and flicked it away; and tried

it again with the same result, and left it.

Harmonious Shrike-Thrush took it eagerly, wiped it on the

ground several times, then jammed it into a forked branch and

started gingerly pulling it to pieces with much shaking of his

head and wiping of his beak. He then broke it in two pieces;

flew away with one and pushed it into a cranny and still per-

severed. He then broke another piece off, and stuck it in a cleft

branch ; but finally left it. I did not see him eat any of the

moth although he may have swallowed small particles. In any

case there were pieces of it left in the places where he had

fixed them.

Tue Crnnapar Morn (Luchelia jacobee).

July 31, 1909. One given to Meerkat, which caught it on the

wing with a snap, devoured it with every sign of relish, and

seemed eager for more.

One given to Capuchin (Cebus sp. a), which stuffed it into his

mouth at once, chewed it, then hastily took it out again, ap-

parently finding he had something either unusual or unpleasant

on his tongue; smelt it, pulled it to pieces with his hands, and

finally ate it, but with a good deal of doubt as if undecided as to

whether it was nice or nasty.

July 5, 1909. One specimen offered to a Fantailed Flycatcher

was immediately seized and pecked and tasted, and then rejected.

The Shama then tried it, and treated it in the same way, finally

rejecting it. A second Flycatcher then tasted it, and rejected it.

Another specimen of the moth let loose in this aviary flew

through the wires into another compartment, and was captured

on the wing by a Pied Grallina. He pecked it once or twice, and

tasted it, then flew away. A Cuban Mocking Thrush then came

up, and while he was looking at it and hesitating to peck, the

Grallina came back, drove away the Mocking Bird, seized

the moth and gradually ate it, holding it in one foot and

pecking it to pieces,

PALAVABILITY OF SOME BRITISH INSECTS, 833

Larva of Cinnabar Moth.

Aug. 15, 1909. Inspected but not touched by English

Thrush.

Offered to many fowls, only one of which pecked it, but dropped

it at once and took no further notice.

Maepie Mors (Abraxas grossulariata).

Aug. 1909. Offered to fowls, was inspected by several, but

only pecked by one, which at once dropped it, and made no

further attempt.

Small Green Geometra larva, probably of Cabera

pusaria or exanthemaria.

May 26, 1909. One taken without any hesitation by a Shama ;

but dropped. Then taken a second time, and dropped. When

preparing to take ita third time, he was deprived of it bya

Black-headed Sibia, which after spending a few seconds adjusting

it in his beak, swallowed it. It appeared to me that the Shama

dropped this larva accidentally, owing to lack of skill in adjusting

it in his beak, rather than intentionally. He was just as eager

to take it, although dead, the third time, as the first.

THE SWALLOW PRoMINENT (Pheasia dicta or tremule).

July 12,1909. Flattened itself to the ground but was at once

pounced upon by the same Flycatcher that had just eaten the

Hemerobiid (see p. 835). The bird, without any hesitation, ate

it with all speed, being merely delayed by the trouble of adjusting

the wings. Both this moth and Mamestra persicarie betrayed

their identity as Lepidoptera by flying out of the boxes to the

ground, so I had no chance of judging whether the Flycatcher

or other birds would have been deceived by their procryptic

coloration.

THe Bure-tie (Phalera bucephala) (amago).

July 5, 1909. Not being aware of this moth’s propensity, I

picked it up by the wings, whereupon it immediately twisted its

abdomen round and ejected a stream of white fluid over my

fingers. | regret that I missed seeing this defensive device

practised on a bird. However, I placed the moth on a wooden

branch, and a Fantailed Flycatcher flew down to inspect it ; after

loooking at 1t for a few seconds, he flewaway. I then put it near

a Shama, who hopped up to it and almost immediately picked

it up by the thorax. The other birds in the aviary now became

interested and pursued the Shama, giving him no chance of

eating it. When on the wing he dropped the moth, and the

Fantailed Flycatcher, which had previously taken no notice of it,

834 MR. R. I. POCOCK ON THE

immediately pounced on it, and after one or two efforts swallowed

it at a gulp. :

My impression is that the Flycatcher did not suspect the moth

of being eatable until he saw the Shama take it. It certainly

looked very like an inanimate excrescence as it rested on the perch.

Presumably the moth had exhausted its intestinal artillery upon

me, because it shot out no more when seized by the birds, but

kept perfectly quiet without even flapping its wings, although the

Shama did not crush it, and having it end on by the thorax with

the moth’s head in his mouth, left the wings perfectly free to flap,

and the abdomen to wriggle, had the moth been disposed to

struggle. This behaviour, I take it, was a manifestation of the

deeply implanted instinct to keep absolutely still (commonly called

‘death-feigning’), which is so highly developed in many animals

with procryptic shape and colour.

THe Dor (Mamestra persicaric).

July 12, 1909. One flattened itself to the ground, and was

seized by the Fantailed Flycatcher that had eaten Pheasia tremule,

and was eaten with avidity, delay, however, being caused by the

bird’s desire to get rid of the wings as well as by being disturbed

by another Flycatcher and a Syrian Bulbul, which tried to de-

prive him of the moth. The Bulbul subsequently picked up the

pieces of wing and ate them.

Larve of the Briaut-LtivE Brown-rve (Mamestra oleracea).

Oct. 26, 1909. One given to Harmonious Shrike-Thrush was

taken after a moment’s scrutiny. He pecked it, and tasted it

three or four times, then swallowed it readily enough. His be-

haviour suggested to me a certain amount of caution at first, as if

he remembered the distastefulness of the pupa of Pieris brassice

which he had just previously eaten. The green hue of both gave

them a superficial similarity to one another. Having eaten the

one specimen of oleracea he was very keen to get the second.

This I gave to the Black-winged Grackle which a few minutes

previously had unhesitatingly left the pupa of P. brassice after

one taste. He took it, and after a taste or two proceeded to eat

it with avidity, not giving the Shrike-Thrush, who was hovering

near and following him up for an opportunity to snatch it, a

chance to do so.

Larva of Drinker (Cosmotricha potatoria).

May 26,1909. One thrown to floor of aviary, was followed

by many birds and secured by a female Black Tanager, which

earried it to a perch and proceeded to peck it and shake it for

about one minute. She then dropped it, and it was seized by

the Black-headed Sibia, but was dropped at once. The Tanager

thereupon tried it again; and again let it fall, this time almost

PALATABILITY OF SOME BRITISH INSECTS. 835

immediately. Two Fantailed Flycatchers then came up and in-

spected it. One of them pecked it, but let it alone after one

experimental taste. The larva was by this time dead. Thena

Sulpbury Tyrant came up, picked it up and after a peck or two

swallowed it.

Larva of Voctua (unidentified).

May 26, 1909. Two (fed on Tropeolwm, so-called Nasturtium)

eagerly eaten by Pekin Robin and by Kagu.

One (fed on cabbage) was readily eaten by Yarrell’s Curassow,

which had just rejected the larva of the Large White (Preris

brassice) and of the Small White (P. rape).

Order NEUROPTERA.

Hemeprosip (unidentified).

July 12, 1909. One turned loose in aviary was at once caught

on the wing by Fantailed Flycatcher and eaten without hesitation.

The bird wiped its beak two or three times ona branch afterwards ;

but I do not think this action can be regarded as a certain sign

that it wished to remove something unpleasant. It suggests the

possibility, however, especially in view of the fact that the action

was not repeated by the same bird after greedily eating Pheasra

dictea and Mamestra persicarie.

Large Black and Yellow Dragon Fiy

(Cordulegaster annulatus) male.

July 26, 1909. One pounced upon and eaten after a time by

Harmonious Shrike-Thrush.

Order ORTHOPTERA.

Common GRASSHOPPER (Stenobothrus sp.).

Sept. 6, 1910. One given to Pekin Robin was eagerly taken

and eaten, but not with great rapidity, the bird putting it on the

ground between the pecks, but without once shaking his head or

showing any signs of disliking the taste. He appeared to me to

be troubled by the insect’s legs.

Great GREEN GRAssHopPER (Locusta viridissima).

All the birds in the aviary were keen to get it. It was tackled

at once by a Dial Bird; but he was driven off by a Black Tanager,

who flew away with the insect and pulled it to pieces on the top

of a wall.

SumMATRAN Stick Insect (Lonchodes sp.).

Taken and eaten at once by:—Pinché Marmoset, Lion Mar-

moset, Douracouli, Capuchin, and Banded Mongoose.

836 MR. R. I, POCOCK ON THE

Taken in the hand, but put down untasted and unhurt by

Grey Lemur.

Taken and eaten at once by:—Silver Pheasant, Cartagenian

Motmot, Fantailed Flycatcher, two Dial Birds, Shama, Black-

chinned Laughing Thrush, two White-crested Jay-Thrushes,

Black-winged Grackle, Chinese Mynah, Brazilian Hangnest, and

Shrike.

The Shrike was too shy to take the specimen from my fingers,

so I threw it towards him on the sand, not seeing exactly where

it fell. He, however, saw the direction of the falling insect, and

hopped towards it, but somewhat to my surprise—for birds

seldom lose sight of thrown food—did not pick it up but looked

as if inquiringly up at me. After a little search I found the

small Stick Insect on the sand lying still with legs extended, and

looking exactly like a blade of green grass. When I stirred it

up and made it crawl, the Shrike was on to it in a moment; and

I have no doubt that he missed it in the first instance owing to

its resemblance to the grass blade.

I observed that several of the birds looked inquiringly, as I

should describe it, at the Stick Insects before taking them. One

in particular, the Harmonious Shrike-Thrush, usually one of the

keenest insect-eaters in the Gardens, hesitated on two occasions

so long before making up his mind to touch them that he was

promptly robbed of his prey, once by a Dial Bird and once by

the Black-chinned Laughing Thrush.

Two birds took them directly, but instead of eating them,

hopped about with them in their beaks. One of these, the Green

Hangnest, was deprived of his by a Chinese Mynah, which took

it from him through the partition bars of the next aviary; the

other, a Grey Struthidea, was similarly robbed by a White-crested

Jay-Thrush after a Collared Jay-Thrush had made several

attempts to get it from him.

Order COLEOPTERA.

Group GEODEPHAGA.

The species of this group used for the test belonged to the

Carabide, a family of carnivorous ground-beetles with an

exceedingly hard exoskeleton. Carabus violaceus is black with

blue reflections; the species of Péerostichus are dead black and

shine like pitch. Harpalus has pubescent elytra and is a little

less conspicuous.

Carabus violaceus.

July 51, 1909. One rejected, after being smelt by three Meer-

kats, two Banded Mongooses, and one White-tailed Mongoose.

The latter behaved towards it exactly as he did towards the

Ocypus oles (see p. 838).

PALATABILITY OF SOME BRITISH INSECTS. 837

Offered same specimen to Harmonious Shrike-Thrush, which

seized it eagerly but was robbed by the Dial Bird. I am sure by

the way they tackled the beetle that either of these birds would

have eaten it ; but the Spotted Bower Bird robbed the Dial Bird,

as in the case of O. olens, and finally finished it.

July 31, 1909. One dropped on to floor of cage of the

Meerkat which had just eaten a Zimarcha tenebricosa. He

pounced on it, but would not seize it as he did the Zimarcha. 1

think he bit it, but am not sure. However, by the way he

pawed it about I am convinced he did not care for it. While

he was holding and smelling it, he quite suddenly let it go and

vomited up the Zimarcha (see p. 841). The Carabus escaped

unhurt. I then gave it to a Capuchin which seized it, and was

proceeding apparently to eat it when another snatched it from

him and ate it without showing any marked signs of dislike, but

with no great avidity.

This species, like others of the genus Carabus, discharges from

its mouth when handled a most repulsive smelling fluid.

Mr. Beddard found that Lacerta ocellata ate this beetle.

Pterostichus (Abrax) striola.

July 26,1909. One taken by Sulphury Tyrant which shook it

and pecked it for some time until robbed of it by Spectacled

Thrush. This bird also pecked it and banged it about until

robbed by female Black Tanager, which ultimately ate it after

much pecking and tasting.

The delay in eating this beetle on the part of the birds that

tried it may have been due to its hard exoskeleton or to partial

unpalatableness from other causes. The hardness alone would, I

think, account for it.

One (dead) given to Silver Pheasant was swallowed entire with

very little delay. The bird, however, after taking the insect

from my fingers, put it on the ground as is his custom with

anything hard or with soft butterflies not quite to his liking.

July 31, 1909. One seized and bolted at once by Silver

Pheasant in exactly the same way that he had bolted the other

Carabide.

Pterostichus niger.

July 31, 1909. One smelt but rejected by three Meerkats;

snatched from the forceps by a Common Indian Mongoose,

which followed it and watched it, and smelt it as a cat does a

cockroach, but did not eat it, so I took it from the cage un-

injured. White-tailed Mongoose turned from it in disgust.

One seized and bolted at once by Silver Pheasant.

According to Mr. Beddard this beetle was eaten without

hesitation by Lacerta vivipara and another lizard: and with some

hesitation by Finches.

838 MR. R. I. POCOCK ON THE

Pierostichus (Steropus) madidus.

July 31, 1909. One smelt and refused by three Meerkats.

Seized and eaten by White-tailed Mongoose. This Mongoose is

a large animal approaching a cat in size.

One seized and bolted by Silver Pheasant.

One pecked twice by Elliot’s Pheasant, but escaped into the

grass unhurt.

One seized and eaten by Black-headed Sibia.

Harpalus ruficornis.

July 21, 1909. One pecked at twice by Silver Pheasant but

not eaten, the bird taking no further interest in it after the

second peck. The beetle escaped unhurt.

Group BRACHYELYTRA.

Devit’s CoacH-HorRsE or Cock-TaIL BEETLE (Ocypus olens).

(Uniformly velvety black in colour.)

July 31, 1909. One smelt and rejected at once by three

Meerkats, one Mongoose, one Banded Mongoose, and by the

White-tailed Mongoose that had just before eaten the Timarcha

(see p. 842). This Mongoose started away from the scent in a

way that reminded me of the behaviour of a person who finds a

bottle of smelling salts unexpectedly pungent.

Offered the same specimen to Harmonious Shrike-Thrush,

which tackled it at once, but while pulling it to pieces was

robbed by the Dial Bird, and this bird in turn was robbed by a

Spotted Bower Bird, which ate it.

Note.—The difference between the Viverrine mammals and the

birds in their behaviour towards the Ground Beetles (Carabide

and Ocypus olens) was very marked, and is to be in a measure

explained, I think, by the wide difference in their powers of

smell. ‘The beetles appear to be relished by the birds; but to be

nauseous to the mammalia, This perhaps is natural; because

the Passerine birds would seldom come across the Ground Beetles,

which are cryptozoic and largely nocturnal. The mammals like

the Meerkats, and the Mongooses, on the other hand, must

commonly find them as they grub about and hunt for food on

the ground. Therefore one would expect protective attributes,

if existing at all in these beetles, to be of a kind to guard them

against being eaten by Meerkats or insectivorous mammals of

similar habits.

The Silver Pheasant which ate these beetles is essentially a

diurnal feeder and would seldom find nocturnal beetles. After

seeing him eat the Péerostichi as if they were large seeds, I do

not understand why he did not eat: the Harpalus ruficornis

offered to him some time previously *.

* Mr. G. A. K. Marshall suggested at the meeting when this paper was read that

the Harpalus had retained while the Péerostichi had discharged their acrid juices.

PALATABILITY OF SOME BRITISH INSECTS. 839

Group LAMELLICORNIA.

Dune BEETLE (Geotrupes vernalis).

July 23, 1909. Offered to Pearl-spotted Owl, which blinked at

it, but refused to touch it. Offered to a White-eared Scops Owl,

was at once taken and held up in one foot ; but after a few pecks,

which removed some legs, it was let fall, no effort being made to

recapture it. Given to a Ludwig’s Bustard, was eagerly taken,

and swallowed whole after a few pecks.

The large CockcHaFER (JJelolontha vulgaris).

July 23, 1909. One dropped on floor of aviary was pounced

upon by Indian Dial Bird which had just before been trying the

Timarcha. He pecked it, hammered it with his bill, and after a

great deal of difficulty broke it in half. He evidently liked it,

because he would not give any other bird a chance of getting it.

However, when he had broken it up, the Harmonious Shrike-

Thrush secured one half and carried it away, and after pecking it

for a few minutes swallowed it. The Dial Bird in the meantime

finished off his portion.

Srac BEEerLe (Lucanus cervus) male.

July 31,1909. This I showed to some Capuchins, which evinced

the greatest eagerness to secure it, but no sign of fear. I gave it

to one, and his first act was to bite off the mandibles. This may

have been an accident, but it reminded me of the alleged action

of baboons in removing the stings of scorpions before they can do

any damage with them. He then bit off the legs, finding they

worried him, and sitting down munched up the beetle as if it had

been a bit of apple. On a previous occasion I gave a dead Stag

Beetle (male) to some Brush Turkeys. One seized it and was

promptly chased round and round the enclosure by the others,

which evinced the greatest keenness forashare. I could not wait

to see what ultimately happened to the insect.

Group LONGICORNTA.

Strangalia armata, the only species of this group experimented

with, is a black and yellow, somewhat wasp-like flower-haunting

diurnal beetle, with a very hard exoskeleton.

July 21 to 31,1909. One taken at once from my hand by

Silver Pheasant and eaten after a good deal of pecking and

breaking up. The way the bird persevered with this hard-shelled

beetle shows that his rejection of the Harpalus was not due to

its hardness (p. 838).

One offered to Fantailed Flycatcher, which, however, would

not touch it. Black-headed Sibia took it without hesitation, and

flying away with it pecked it to pieces and finally ate it. Further

840 MR. R. I. POCOCK ON THE

evidence of the bird liking the insect was shown by the way he

flew away with it when chased.

One taken and eaten by Dial Bird, which was apparently only

delayed in disposing of it by the hardness of the exoskeleton.

One taken and similarly disposed of by Great Barbet.

One eaten after being broken up and crushed by Brazilian

Hangnest.

Group MALACODERMATA,

The beetle of this group used for the experiment is a flying

diurnal flower-haunting species, with a soft exoskeleton. It is

quite fearless of exposure. Beetles allied to it commonly form

centres of mimetic attraction in the tropics.

TELEPHORID ( ? Rhagonyche fulva).

July 21,1909. Four offered to four Capuchins were eaten,

two readily and without examination, two after a good deal of

tasting and examination between the tastes.

Two offered to two Capuchins were taken into the mouth,

tasted, then taken out, wiped on the bars and left.

One refused by Ceylonese Macaque after being smelt.

One eaten by Mona Monkey after a good deal of tasting,

smelling and pulling about. This Mona also ate the bug 7’ropi-

coris rujipes (p. 847).

One offered to Lion Marmoset was taken in the hand, smelt,

and promptly dropped. The Marmoset then descended from the

perch, picked it up again, smelt it and dropped it. The beetle

crawled away unhurt.

One smelt once or twice by Meerkat, was rejected without

being tasted.

One taken by Silver Pheasant, was pecked twice and left alone.

Another offered to same bird was pecked once and left. One

taken by Fantailed Flycatcher was pecked and tasted, then left.

The same specimen was then pecked once or twice by a Shama

and rejected. Black-headed Sibia then tried it, but gave it up

and vigorously wiped his beak after a taste or two. Afterwards

he made another attempt with the like result. The Black

Tanager then took it, tasted it, wiped his beak and rejected it.

One caught on wing by Harmonious Shrike-Thrush was eaten

after much pecking and pulling about. Another was treated in

the same way by this bird.

Two specimens, one of which was dead, offered to and eaten by

Dial Bird.

One tasted two or three times by Shama but rejected.

One pecked by Black-chinned Laughing Thrush, but flicked

away. Pounced on and eaten by Dial Bird.

Although eaten by the Dial Bird and the Shrike-Thrush,

which ate most of the insects offered to them, and by some of the

PALATABILITY OF SOME BRITISH INSECTS. 841

Monkeys, there can be no doubt that this soft-shelled beetle

possesses distasteful attributes. Its rejection by the Meerkat,

which ate nearly all the insects offered to it with the exception of

Coccinella 7-punctata, was very significant, and suggestive of nasty

smelling secretions.

Group PHYTOPHAGA.

The three species of this group that were tested are well-known

species. They are slow-moving diurnal forms found on plants of

different kinds. ‘They are squat in shape, dorsally convex, and

have a very hard exoskeleton, the Ladybird (Coccinella) being in

addition exceedingly slippery and difficult to hold. The coloration

of the latter is orange with black spots. The others are uniformly

black or blue. TVimarcha tenebricosa, the familiar ‘ bloody-nose

beetle,’ is further notorious for the discharge from its mouth of

a crimson liquid, whence the trivial name is derived.

Chrysomela polita.

July 31, 1909. One oftered to Meerkat was smelt and refused.

Another Meerkat in the same cage took it in his mouth, but spat

it out; both then sniffed it as it lay on the ground, but would

not touch it.

The same specimen, offered to a Grison, was sniffed but not

touched. Snapped up by McCarthy’s Mongoose; but was at once

spat out and left. It was then taken and eaten by a Banded

Mongoose.

Query: Had the previous tasters exhausted the Beetle’s supply

of nauseous juices ?

One given to Dent’s Monkey was taken, rubbed between

the hands and in the sawdust, smelt, tasted, pulled about and

rejected. Picked up by Mona in the same cage, but rejected

after one taste. ‘This Mona had just eaten a living Bombus.

One given to Harmonious Shrike-Thrush was taken, pecked

and tasted for a little, then left. Picked up by Black-chinned

Laughing Thrush, was pulled to pieces, and rejected. This bird

may have eaten pieces of the beetle, but the other débris was left

on the turf. He did not appear to find it very unpalatable.

Possibly in this case the nauseous juices had been exhausted by

the Shrike-Thrush.

One pecked off a perch by Fantailed Flycatcher, but not

followed up. Pecked and tasted by Sulphury Tyrant, but left.

Then tried by Sun-Bittern, but also left, crushed but with

nothing missing.

Timarcha tenebricosa (=levigata).

July 23-31, 1909. One offered to a Meerkat was eagerly seized,

chewed up and swallowed without much hesitation. But while

this Meerkat was just afterwards occupied with the Carabus viola-

ceus (cf. supra, p. 837), he yomited the Timarcha, I do not know

842, MR. R. I. POCOCK ON THE

whether the sickness was caused by the smell of the Carabus,

which to me is nauseating, or to its taste, or by the irritation of

the stomach caused by the Zimarcha. I suspect the latter,

because the Meerkat refused to touch a second Zimarcha that

was offered to him.

One smelt and rejected untasted by two more Meerkats ;

taken by a third in the same cage, rubbed in the sawdust, but

left apparently uninjured.

One grabbed at once and eaten by White-tailed Mongoose,

which immediately afterwards heaved and went through the

action of vomiting without, however, ejecting the beetle. <A

second specimen was smelt and rejected with every show of

disgust by the same animal, which persistently refused for

the next two hours every beetle that was offered him, although

before eating the Zimarcha he had devoured a Pterostichus

madidus. One rejected without being closely smelt by a Banded

Mongoose which had eaten a Coccinella 7-punctata. Seized by a

second Banded Mongoose, and eaten after a good deal of rubbing

in the sawdust.

One offered to a Capuchin, one of the specimens which had

refused the Telephorid (hagonyche fulva) (p. 840), was taken,

smelt, and rejected.

One offered to another Capuchin was ultimately eaten piece-

meal, but with so much delay caused by handling, licking, and

inspection, that I am sure it was no great treat to him, especially

as he had every reason to eat it speedily because a bigger

Capuchin in the same cage, which had snatched the Carabus

from his grasp, was almost continually after him to get the

Timarcha. When monkeys like their food they gobble it up if

there is the least likelihood of another taking it.

One offered to a Vervet Monkey was accepted, pulled to pieces

and eaten, the exoskeleton being dropped to the ground.

This specimen of Zimarcha had been previously offered to a

Baboon (Papio sphinx) ; but he would not even touch it.

One put on the floor of aviary was pounced upon by Dial

Bird, which after continued pecking and hammering could

make nothing of it beyond breaking it in half at the waist.

Ultimately he left it. An Orange-headed Thrush then tried

the abdomen, but was driven off by a Hoopoe, which after

pecking and hammering it, gave it up. The Thrush then tried

again, and also gave it up. A Black-chinned Laughing Thrush

then had a turn ; but with the same result.

One given to Harmonious Shrike-Thrush which had eaten the

Coccinella. He persevered for a long time, but could not manage

it and flew away, leaving the beetle apparently unhurt. After

about five minutes the bird came back and tried again, this time

pecking off the legs and antenne of the beetle; but he would

not eat the body, and at last flew away and returned no more.

Sept. 18, 1910. One female taken by Kagu, well crushed,

then swallowed at a gulp. °

PALATABILITY OF SOME BRITISH INSECTS, 843

One male taken by Vigors’s Bustard, crushed and put down

with a head-shake ; then tasted by two Ludwig’s Bustards, the

three birds having alternate pecks at it, the Vigors’s Bustard

finally swallowing it.

One female well tasted, but rejected by Wood-Swallow, Black-

winged Grackle, Javan Pied Mynah, and Black-chinned Laughing

Thrush: also by Sun-Bittern, which persevered for a long time,

repeatedly washing the beetle in the water-trough, and taking a

drink at the finish.

Taken and pecked to pieces, and eaten bit by bit by Silver

Pheasant. The bird wiped his beak several times on the earth,

and for some little time afterwards stood opening and shutting

his beak like a monkey or a human being getting the flavour of

something tasty.

Some of the birds which tried to eat the Timarcha showed no

special signs of finding them unpalatable. It appeared to me

that they finally refused them on account of the hardness of the

exoskeleton. Probably this prevented them getting at the

softer tissues containing the flavour, whether unpleasant or

otherwise.

Larva of Vimarcha tenebricosa.

(A fat bluish-black grub.)

June 15 to 24, 1909. One eaten with apparent relish by

Meerkat, which only delayed seizing it for about two seconds to

rub it in the sawdust and smell it. ‘This was the same Meerkat

that on a previous occasion had eaten Huchelia jacobee and

rejected the Coccinella.

One taken at once by the same Capuchin that had eaten

E. jacobee and rejected Coccinella; but after crushing it between

his teeth and getting the flavour, the monkey at once took it out

in his hands, contemplated it for a few seconds, and moving his

lips the while as if sampling the flavour, then letting it fall,

retired to the back of his cage, salivated and heaved twice as if

going to vomit.

Another Capuchin in the same cage now picked up the crushed

larva, tasted it, and put it down; and neither of the monkeys

touched it again. So I gave it to the Meerkat, which ate it as

greedily as it did the first.

One given to Armadillo was eaten after a good deal of smelling.

A second was eaten without hesitation.

One given to Dent’s Monkey was eagerly taken and tasted,

but almost at once dropped. The monkey did not taste it again,

although he was interested in it and played with it for some

little time.

One given to Mona Monkey, which behaved in much the same

way as Dent’s Monkey, but played with the larva for a longer

time.

One given to Capuchin (sp. a) was taken and chewed up, but

844 MR, R. I. POCOCK ON THE

just as I thought he was going to swallow it, he spat it out with

profuse salivation.

One given to another Capuchin (sp. @) was licked and dropped.

One given to a third Capuchin (sp. 6) was chewed up and

swallowed without any signs of dislike, the larva being not even

taken from his mouth for examination.

Another given to the same monkey was also eaten without any

signs of dislike, although he held it in his hands and licked it

several times before finally putting it into his mouth and chewing

it.

June 24, 1909. Repeated experiments with monkeys.

The two Capuchins (sp. a), the Dent’s and Mona Monkeys

behaved exactly as before. They took the larve, smelt them,

tasted them once or twice, and finally rejected them. The

Capuchin (sp. 6) which had previously eaten two, again ate one

without signs of relish or the opposite. [I then offered a larva

to another Capuchin of the same species (}) and he treated it as

the specimens of the species @ and as the Mona had done, that

is to say smelt it, tasted it, rubbed it in his hands, repeated the

tasting once or twice, and finally dropped it. His behaviour

showed that the difference between the behaviour of the first

example of sp. b, which ate the larvee, and that of the examples

of sp. a, which rejected them, is not attributable to the specific

distinction between the Monkeys as might have been supposed, if

only one specimen of sp. 6 had been available for experiment.

One given to Canadian Jay, taken, pecked, jammed into a

cranny, and repeatedly pecked ; then dropped. When the bird

made no attempt to fetch it, the keeper picked it up and placed

it on the perch, when the bird again seized it, jammed it into a

cranny in the perch, and left it.

One given to Red-backed Shrike was eagerly seized, and after

one or two pecks was left, the bird retiring and wiping his beak

on the bars, as the Canadian Jay had also done.

Two given to Silver Pheasant were taken and pecked, and after

a good deal of rubbing in the earth were eaten.

One given to Prince of Wales’ Pheasant was taken, pecked and

rejected.

One given to Piping Crow was pecked and tasted and rejected,

after a good deal of shaking of the head and wiping of the beak

on the part of the bird. It was then picked up by a Magpie,

which after a taste or two stowed it away under a large stone,

and built up the hole with pebbles.

One given to Buff Laughing Kingfisher was taken and tasted,

but rejected with much bill wiping. Tried and rejected in the

same way by a second specimen of this bird.

One given to Common Laughing Kingfisher was taken and

tasted, but finally rejected.

One given to Dial Bird was finally rejected after a great deal

of pecking and tasting, accompanied by much shaking of the head

and wiping of the bill,

PALATABILITY OF SOME BRITISH INSECTS, 845

One given to White-collared Crow was taken, tasted, carried

about, and finally dropped. This bird refused to take a second

specimen offered immediately afterwards.

One given to Hooded Crow was treated in exactly the same

way as the one above-mentioned was treated by the White-

collared Crow. This Hooded Crow also refused a second

specimen.

One given to Wild Turkey was taken and pecked, but soon

rejected.

SEVEN-SPOTTED LADYBIRD (Coccinella T-punctaia).

July 5, 1909. I offered one to the Capuchin which was the

only one of these Monkeys to eat the 7imarcha-larve, thinking

he might be deficient in tasting powers. He took it at once from

my fingers into his mouth, and crushed it between his teeth;

but, presumably as soon as he got the flavour, removed it frora

his tongue with his fingers, and took no further notice of it.

I offered the remains to a Mona Monkey, but she only Smelt

them and pulled them to pieces, and would not taste them.

July 23 to 31, 1909. One was offered the Capuchin (sp. @)

that had eaten the Huchelia jacobee and Bombus lapidarius on

the previous day, and had so far vefused nothing in the way of

Lepidoptera. He took it from my hands directly, transferred it

to his mouth and crushed it; but instantly took it from his

tongue, wiped it on the perch and left it without a second look.

I then gave the crushed insect to the Meerkat that had eaten

E. jacobee and the Bombus lapidarius. He seized it at once, but

just as promptly spat it out, gave his mouth a wipe with his paw,

and never attempted a second taste.

One given to Vervet Monkey which had just eaten a Timarcha

tenebricosa (see p. 842). She took it, smelt, licked and examined it

thoroughly, rubbed it between her hands, then dropped it to the

floor and took no further notice of it. I had previously offered

this Coccinella to a Chacma Baboon. She smelt it but would not

take it from my fingers.

One given to the Capuchin which on a previous occasion had

tasted and rejected one. He took it, and after a great deal of

smelling, tasting, rubbing between his hands and on the boards

of the cage, finally ate it bit by bit, pulling it into many little

pieces. This Capuchin had just before eaten a Carabus violaceus.

One smelt but refused by three Meerkats. Grabbed by Yellow

Meerkat, tasted, but let go unhurt. Taken by Banded Mongoose,

and eaten after much rubbing in the sawdust, and with many

shakes of the head.

One offered to Grey Lemur, was smelt, taken in the hand and

dropped.

Sept. 20, 1910. One taken and quickly eaten by Meerkat;

but the same animal refused a second specimen.

One taken and rubbed about in the sand and repeatedly bitten,

Proc, Zoou, Soc.—1911, No, LVITL. 58

846 MR. R. I. POCOCK ON THE 2

and ultimately eaten by another Meerkat, but the same animal

refused a second.

One taken in the paws by a Marsh Mongoose, but rejected

after being repeatedly rubbed in the sand and smelt.

One taken by Banded Mongoose and crushed, but rejected with

much head-shaking; swallowed by a second animal also with

much head-shaking.

One refused after being smelt by three Yellow Meerkats.

One taken and licked by Capuchin, but rejected.

One licked but rejected by Red-handed Marmoset.

Another monkey of same species, and a Common Marmoset

refused even to taste it.

July 23, 1909. One examined by Spectacled Thrush, but not

touched. Pecked by Fantailed Flycatcher, which shook his

head and left it. The bird returned three times, however, and

pecked the beetle, but finally gave it up. I then offered it to

a Shama three times in succession, and upon each occasion he

flicked it away and made no attempt to follow it up. Next I

tried the Harmonious Shrike-Thrush. He took it, and after a

good deal of pecking, ate it.

July 31, 1909. Three eaten in succession by cock Silver

Pheasant. The first one he took from my hand, but put it out

of his beak on to the ground. After one or two pecks, however,

he swallowed it. The others he took from my fingers and bolted

entire as if they were grain, exactly as he had previously bolted

the beetles, Pterostichus niger and Ocypus olens.

Sept. 20, 1910. One taken by Pearl-spotted Owl, but dropped

at once.

One taken by a Pekin Robin, which after a few pecks and head

shakes left it and took a drink of water; tasted by another bird

of the same kind, but also left uneaten.

One taken and swallowed, after a deal of pecking about in the

sand and head shaking, by another specimen of Pekin Robin,

which had just previously eaten the grasshopper (Stenobothrus)

and the bug (Therapha hyocyami).

One given to the Dial Bird that had just eaten a Humble Bee

(Bombus agrorum). He took it at once, and after a little delay

swallowed it whole.

N.B. This is the bird that rejected the two White Butterflies

(Pieris brassice and nap?) after tasting them.

One taken but rejected by Masked Wood-Swallow ; then taken

and eaten by Shama.

Two taken and hkolted quickly by the same Shama, which

showed no signs of objecting to the taste, except a single shake

of the head on each occasion after swallowing the beetle.

Although some of these beetles were eaten both by mammals

and birds, there can be no doubt that they were distasteful to the

majority of the animals to which they were offered, even to some

of those that ate them.

PALATABILITY OF SOME BRITISH INSECTS. 847

The interest of the demonstration of the distastefulness of

Coccinella 7-punctata lies in the fact that Coccinellide of various

kinds are numicked in the tropics by insects of other orders, as

well as by spiders.

Order HEMIPTERA.

OxtvE-BROWN Bue (Zropicoris rufipes).

July 21, 1909. One (dead) given to Mona Monkey was eaten

after a great deal of handling, smelling and tasting.

One put on the ground was tackled by Fantailed Flycatcher,

which pecked it some half dozen times. He was then driven off

by a hen Black Tanager, which pecked it and pecked it again, and

then left it. A Syrian Bulbul then flew up and tried it, but after

persevering for some little time gave it up. Then the Tanager

had another attempt, but left it. I then gave the mangled

remains to the Harmonious Shrike-Thrush, and after a little

pecking about he swallowed them.

One (living) eaten with very little delay by Silver Pheasant ;

but put on the ground after being taken from my hand. This

specimen was immature on arrival; 1t moulted in the box, and

was apparently adult when given to the bird.

One (dead) treated in the same way and eaten by the same

bird.

Rep anD Buack Bue (Therapha hyocyami).

Sept. 20,1910. One given to Pekin Robin was at once taken

and ultimately eaten; but the bird took a long time over it,

putting it on the ground after each peck and vigorously shaking

his head before tasting it again. The behaviour of this bird was

exactly the same towards Coccinella 7-punctata (p. 846).

Order DIPTERA.

Bombus-like Fly (Volucella bombylans).

July 26, 1909. One taken by Fantailed Flycatcher but after

being pecked and pulled about for some time, was left. The

Sulphury Tyrant then tried it, but also left it alone after much

pecking. Finally it was taken by Spectacled Thrush, which ate

it after much pecking and wiping in the sand.

One given to Black-headed Sibia was eaten after a great deal

of pecking and breaking up.

These experiments, as Dr. Longstaff reminded me, suggest that

this fly is, at all events to a certain extent, unpalatable. If

future tests should prove it to be so, its likeness to Bombus will

be an instance of Millerian rather than of Batesian Mimicry.

See also below, pp. 854-855,

58*

848 MR. R. I. POCOCK ON THE

Bombus-like Fly (Arctophila mussitans).

See below, pp. 851 and 853.

Fly like a small Bombus (Chilosia illustrata).

See below, pp. 854-855.

Spiny Fry (Lehinomyia ferox).

July 31,1909. One (dead) taken by female Tanager, but after

a good deal of pecking, was left. A Black-headed Sibia then

tried it and finally ate it.

One also eaten by Sulphury Tyrant (see below, p. 855).

Dappy Lone-LeEes (Tipula oleracea).

Oct. 26, 1909. One taken from my hand and eaten readily by

Dial Bird; one taken and eaten, but not so readily, by a second

Dial Bird ; one eaten greedily by Fantailed Flycatcher.

One of these specimens of Tipula was taken twice by the

Harmonious Shrike-Thrush, but was dropped on both occasions.

Another was taken three times by Black-winged Grackle, but was

not eaten.

The rejection of this insect by the Shrike-Thrush, which ate

almost every insect other birds refused, was very surprising.

Fly (Zmpis tessellata).

July 31, 1909. Two (dead). Katen greedily by the Dent’s

Monkey that took the Thanaos tages with avidity (p. 831).

Order HYMENOPTERA.

Tipula-like Ichneumonid (Ophion luteus).

(Nocturnal species, mahogany-red in colour, with

very tough integument.)

Oct. 26, 1909. One taken and tried perseveringly by Fantailed

Flycatcher, but ultimately abandoned. Also tried but soon given

up by Yellow-crowned Hangnest; taken and after a little pulling

about swallowed entire by Dial Bird.

Noy. 7, 1909.—Taken by Black-winged Grackle; but so hard

was the insect that it shot away out of his beak. The bird

pounced on it at once on the sandy floor of the aviary and

ate it; but if the insect had not been very lethargic, or if it

had fallen amongst the undergrowth, it might have escaped

him. Hence probably the significance of its hard slippery

exoskeleton,

PALATABILIVY OF SOME BRITISH INSECTS. 849

Larvee of Saw-fly (Cladius viminalis).

These larve were yellow with black spots. They were sent to

me by Mr. Taylor.

Aug. 19,1910. Refused without tasting by Yellow-crowned

Hangnest, Crested Bulbul, Blue-bird, and Fantailed Flycatcher.

Tasted but rejected by Black-winged Grackle, Harmonious

Shrike-Thrush, Black-chinned Laughing Thrush, and Green

Toucanet.

Taken by Greater Spotted Woodpecker, placed in a hole in a

stump and hammered, but ultimately flicked away and lost.

Two taken and eaten after much pecking and tasting by a

Shama. One eaten fairly readily by a Dial Bird; but another

bird of the same species rejected a specimen after tasting and

flicking it from his beak about twenty times.

Woop-Ant (Yormica rufa).

May, 1910. Taken and eaten with avidity by the following

birds :—Pearl-spotted Owl; Orange-headed Ground-'Thrush ;

Dial Bird; Shama ; Black-headed Sibia ; Blue-bird ; Pekin Robin;

Harmonious Shrike-Thrush ; Spotted Oriole ; Larger Hill Mynah ;

Black-winged Grackle; Yellow-crowned Hangnest; Greater

Spotted Woodpecker.

A Capuchin Monkey also ate one after another, picking them

up in his hands and gobbling them as fast as possible.

Several specimens thrown into a cage containing three Wall

Lizards were tasted by two of them, but rejected at once without

being damaged in any way by the tasting.

Most of the birds showed no signs of objecting to the taste of

the ants, or even of perceiving anything peculiar in their flavour.

The Pearl-spotted Owl, however, shook his head, and the Spotted

Oriole wiped his beak on the perch after eating them. The

Pekin Robin and the Black-winged Grackle wiped the ants upon

their wings, presumably to remove the formic acid. It is inter-

esting to find the same device practised by two species so unlike

one another.

I found that the birds, like the monkey, would eat as many

of these ants as were given to them.

The unavoidable conclusion that these insects are palatable is

rather surprising in view of the frequency with which ants of

different kinds are mimicked in the tropics by Orthoptera, Coleo-

ptera, and other insects, as well as by spiders. Nevertheless, it

corroborates the opinion put forward by McCook and amplified

and endorsed by myself in 1909 *, before these experiments were

made, that ant-mimicry is mainly serviceable as a protection

against the predatory Hymenoptera of the family Pompilide, which

provision their nests with Arthropoda of various kinds, excepting

ants, and are certainly the direst enemies that spiders possess.

* Journ. Linn. Soc., Zool. xxx. pp. 265-268.

850 MR. R. I. POCOCK ON THE

Saw-rry (Allantus arenatus).

July 21,1909. One eaten by Mona Monkey fairly readily ; by

nee readily ; by the Capuchin which on the previous day

had refused the Malacoderm Beetle (Rhagonyche fulva); smelt,

but not tasted by Lion Marmoset.

One eaten fairly readily by Harmonious Shrike-Thrush ; by

Shama readily; by Silver Pheasant; refused without tasting by

Wild Turkey.

Honey-BEE (Apis mellifica). (Workers.)

May 8, 1911. One offered to Silver Pheasant was taken from

the forceps but immediately flicked away; the bird persevered,

however, and after much pecking and flicking ¢ about of the insect,

and wiping his bill on the ground, finally ate it.

One offered to Bornean Fire- backed Pheasant was inspected

carefully but rejected untasted.

One given to Pekin Robin was taken at once, but was quickly

flicked away. When pursued, however, by other birds in the

cage, the Pekin Robin pounced on the bee again and flew away

with it. Whenever he got a moment’s peace, he put it on the

eround, pecked and flicked it about, wiping it now and again in

the sand and repeatedly shaking his head. At length he flew to

a branch, and holding the bee against it with his foot, pulled it

in two pieces, dropping one piece to the ground. He still

persevered with the other piece, however, but an finally lost sight

of him and do not know whether he ate it or not.

One given to a Cayenne Tanager was taken and chewed for a

long time ; ; the remains, however, were finally jammed into a

banana and left.

One taken by a Blue Tanager which, however, allowed himself

to be robbed without resistance or flight by a Maroon Tanager.

This bird, after a deal of mastication, ate the bee.

One given to Wall Lizard was eagerly seized, but was left after

one or two attempts.

Another was twice darted at by another lizard of this species,

but was left alone the moment the lizard touched it. It was then

boldly seized by a third lizard, which with one bite disabled the

bee by crushing the head and thorax. This lizard persevered for

about seven minutes, biting at the bee, but stopping after each

bite to lick his mouth with his tongue and Pub it against the moss.

Finally he gave it up and went away.

Two Bluebottles(Calliphor a vomitoria) “ a Hover-fly (Syrphus)

given as a check experiment were seized and eaten in a few

seconds by the same lizards.

Humpte Bes (Bombus agrorum).

(See also infra, p. 853.)

Oct. 26, 1909. One eaten with avidity by Capuchin and by

Meerkat.

PALATABILITY OF SOME BRITISH INSECTS. 851

One given to Collared Jay-Thrush, which pecked it about and

scraped it in the sand for along time, wiping his beak in the

intervals, and ultimately left it. It was then picked up by a

White-crested Jay-Thrush, which treated it for some time in the

same way, but at last ate the mangled remains. This same bird

then took a specimen of the mimetic fly Aretophila mussitans, but

made just the same fuss over the eating of it as he had in the

case of the bee.

Sept. 18,1910. One taken at once by Dial Bird, and after a

good deal of pulling about, pecking and wiping in the sand, was

eaten. This bird had just previously eaten a small Tortoiseshell

Butterfly, and he took about the same time to finish off the one

insect as the other.

Sept. 20,1910. One offered to Dial Bird was taken at once and

eaten with very little delay, after being wiped once or twice in

the sand. The bird flew away with a second specimen and I did

not see what became of it; but he returned to me, and I had

difficulty in keeping him away from the bees with which I was

experimenting with other birds.

This Dial Bird was the one that ate the same species of Humble

Bee two days previously.

Humsie Bee (Lombus ? joncellus).

July 31, 1909. One offered alive to Mona Monkey was

snatched at once and eaten bit by bit.

Houms1e BEE (Bombus ? terrestris).

July 31,1909. One (dead) taken by Brazilian Hangnest and

pecked to pieces, the bird holding it the while in his foot against

the perch. The pieces pecked off were dropped about the cage

and not eaten.

HuMBLe BEE (Lombus lapidarius).

May 31, 1909. One dead specimen given to the Meerkat was

eaten bit by bit, after being rubbed im the sawdust by the

animal’s paws.

One dead specimen given to Capuchin (Cebus sp. a) was taken

in the hands and eaten bit by bit, just as the Monkey would eat

a piece of hard biscuit or sugar. Neither of these mammals

showed any signs of disliking the taste of the bees; quite the

contrary. Their molar teeth are evidently much better adapted

for crushing the chitinous exoskeletons than are the beaks of the

birds that tasted them.

One dead specimen offered to Syrian Bulbul was taken after

about a minute’s inspection. The bird pecked it and pulled it

about for at least five minutes and dodged away with it from

other birds that chased him. He grew, however, less and less

852 MR. R. I. POCOCK ON THE

keen, and ultimately allowed a female Black Tanager to rob him

of it. The Tanager behaved in just the same way, pecking and

pulling it about and breaking it to pieces, but gradually losing

her interest. At last she picked up a piece of the thorax and

flew to a bush with it, leaving the remainder on the ground. I

could not see what became of the piece she flew away with, but

she emerged from the bush without it, and wiped her beak on

a perch. She made no attempt to go back to the bits on the

ground. A Sibia tried these, but after a peck or two left them,

and no other insectivorous bird in that compartment took the

least notice of them. So I picked up the abdomen and gave it to

the Harmonious Shrike-Thrush which had just finished off the

example of B. hortorwm, mentioned below, and he ultimately ate

it after a great deal of pecking and pulling about.

Houmste Bre (Bombus hortorum).

May 31,1909. One living example fell to the ground of the

aviary when first liberated. Two Fantailed Flycatchers flew

down to it at once, but although interested would not touch it ;

while they were hesitating the bee took wing and escaped, none

of the birds in the aviary making any attempt at pursuit.

One dead specimen offered to a Shama. She allowed me to

hold it close to her beak, but would not touch it. None of the

other birds in the aviary would notice it when thrown to the

ground, though on a previous occasion they had shown great

eagerness in seizing dead butterflies. I then gave it through the

bars to the Harmonious Shrike-Thrush in the next compartment.

After pecking and pulling it about for six or seven minutes, he

ultimately ate it.

July 31, 1909. One sniffed at but rejected by two Meerkats ;

taken by a third and eaten.

One pecked and flicked away by Black-headed Sibia, by Shama.

and also by Sun-Bittern, each making two or three attempts.

Then carried off by female Black Tanager, but dropped to the

floor, where a North American Cat-bird tried it once or twice, but

gave it up. (The remains were now too mangled to be useful for

further experiment.)

T could not induce the Fantailed Flycatcher to take any notice

of this bee.

Conclusion. These experiments indicate that the Humble Bees

used for the tests were much more palatable to the mammals than

to the birds. With the exception of the one example of b. lapi-

darius which was smelt and left untouched by two Meerkats, all

the bees offered to the Monkeys and Meerkats were eaten without

any kind of dislike of the flavour being evinced. The Meerkat

that rubbed the B. lapidarius in the sawdust did so, I suspect, to

remove some substance offensive to his sense of smell. On the

other hand, of the birds to which the bees were oftered only three

PALATABILITY OF SOME BRITISH INSECTS. 853

ate them, namely a Dial Bird, a Jay-Thrush, and a Shrike-Thrush.

The Dial Bird ate one quickly with only one or two wipes in the

sand. In the other cases there was a great deal of pecking and

wiping before the insects were finally disposed of. From the

behaviour of the birds there could be no doubt that there was

something in the bees not to their liking, even to those that

ultimately ate them. The Bulbul, Sibia, and Tanager were

obviously keen to eat them, and gave them the fullest possible

trial before finally rejecting them ; but whether it was the hairs,

or the hard chitin, or the favour, or a combination of them that

made the insects unpalatable, I do not know.

Further experiments demonstrating the distastefulness of

Humbie Bees to birds of different kinds are given in the following

section :—

Experiments to test the significance of the resemblance between

Humble Bees (Bombus) and the Flies Arctophila nussitans,

Volucella bombylans, and Chilosia illustrata.

Bombus agrorum and Arctophila mussitans.

Oct. 26, 1909. Offered Bee to a Lion Marmoset which was

busily catching house-flies and bluebottles in his cage. He

looked at it, but would not touch it. I then offered the fly, but

he also refused to touch it. He did not, however, hesitate to take

a Red Admiral offered a moment afterwards.

Offered Bee to Leach’s Laughing Kingfisher. He took it at

once, but soon flicked it away. Six times in succession he took it

from my fingers and dropped it on each occasion. | could not

induce him to take it again. Instead he started pecking my

fingers. Thereupon I offered him the fly, and he just as reso-

lutely refused to take it.

Offered Bee to Kagu, a New Caledonian Rail. He inspected

it, and after a little hesitation tasted it. But he would not touch

it again; and when offered the fly, refused that likewise.

Offered Bee to Central American White-browed Partridge.

He took it without hesitation, but after a peck or two left it and

went away. I then threw it to him, and he tasted it again ; but

would not eat it. I then threw him the fly, but he would not

touch it.

A Douracouli (a South-American monkey) ; a Honduras Turkey ;

a cock and a hen Reeves’s Pheasant, and three hen Silver

Pheasants refused to touch both bee and fly, though they

inspected them intently for a few seconds.

Sept. 20, 1910. Bee offered to Hoopoe was taken at once and

tasted without being crushed, but was then left on the ground

uneaten. The bird refused the next one I offered, and then

refused to take the fly, although he stretched his head towards

it and inspected it.

Bee offered to Yellow-crowned Hangnest, which took it at once,

but soon dropped it. A second time he took it, and dropped it.

854 MR, R. I. POCOCK ON THE

The third time it was offered he refused it, and immediately after-

wards refused the fly.

Bee offered to Sulphury Tyrant. I importuned the bird into

taking it from my fingers no fewer than eight times, and each

time he flicked it away. The ninth time he refused to take it,

and then refused the fly.

Bee offered to Black-winged Grackle, which took it at once, but

dropped it. Twice more he took it and the last time flew a short

distance away and persevered with it for about three-quarters of

a minute, then leaving it returned to me; he refused the next

bee I offered, and then refused the fly.

Bee offered to Silver Pheasant was at once taken, put on the

ground, pecked and crushed almost past recognition, but left

uneaten. The bird then took from my fingers three more speci-

mens in succession, but dropped them uncrushed from his beak

at once. The fifth he looked at, but would not touch, and then

also refused the fly after inspecting it.

The experiments described above with the Lion Marmoset, the

Douracouli, the Turkey, and the Reeves’s and Silver Pheasants,

which would not touch either the bee or the fly after some seconds

of intent inspection, do not prove that the bee was known to be

distasteful, and that the fly was rejected in consequence. That

may be the explanation, The Douracouli, however, is nocturnal

and probably does not naturally feed upon diurnal-flying insects.

In the case of the Marmoset, the experiment does, however,

suggest very forcibly that the Arctophila was not recognised as

closely allied to the bluebottles the animal was hunting. The

other experiments speak for themselves.

Bombus hortorum, Volucella bombylans, and

Chilosia illustrata.

July 31,1909. Offered living Bombus hortorum to the Bra-

ailian Hangnest that had just pulled the dead Lombus terrestris

to pieces. He took it directly, but instantly flicked it away and

wiped his beak. The bee then crawled up the bars of the cage,

and he again pecked and flicked it away. It was now too injured

to crawl although still alive, so I picked it up and offered it in

my fingers. He took it again and flicked it away. Twice more

the trial was made, with the same result, although he was

patently tiring of the trials. The next time he refused to

touch it after inspection. I then substituted a dead Volucella

bombylans. We inspected it, but did not touch it, and hopped up

to the top perch.

I then offered the nearly dead Bombus to another specimen of

the same bird. He took it from my fingers three times in

succession, and each time flicked it away. The fourth time he

refused to touch it. I then substituted the same specimen of

Volucella bombylans, but after looking at it he would not take it.

PALATABILITY OF SOME BRITISH INSECTS. 855

I then again offered the Bombus to the first Hangnest. He

took it and flicked it away, and immediately afterwards refused

to touch the Volucella.

Next day I offered the first Hangnest a Bombusagain ; he took

it from me three times, and flicked it away without attempting

to eat it, but immediately afterwards took Chilosia illustrata and

ate it.

The second Hangnest took a dead Lombus, and flicked ite away,

and then ate Chilosia ilustrata, but refused immediately after-

wards to touch a live Bombus hortorwm.

Offered Bombus hortorum to North American Cat-bird, which

came up to me on seeing other birds being fed. He pecked it

several times, but flicked it away and gave it up. During the

next quarter of an hour I could not induce him to touch either

Volucella bombylans or Chilosia illustrata.

T then offered the Bombus on the forceps to a Sulphury Tyrant.

He pecked and flicked it away several times, then left it, and

refused it when offered again. I then offered him the Volucella

bombylans both in the forceps and by throwing it to him on the

ground, but he would not touch it. After a little hesitation,

however, he took an Hchinomyia ferow from the forceps and ate

it, and then took and ate Chilosia illustrata. I then offered him

Bombus hortorum again, and he took it but soon rejected it, and

immediately afterwards refused to touch Volwcella bombylans.

One Bombus hortorwm ottered to a Shama, which pecked it once

or twice, and flicked it away each time. He then refused to touch

the specimen of Volucella bombylans.

Tried the experiment with another Shama, which behaved in

exactly the same way towards the bee, and would not afterwards

touch the Volucella bombylans.

One Bombus hortorwm offered to Silver Pheasant was taken at

once, but left after some pecking and tasting. Then without

hesitation he took Chilosia wlustrata from the forceps and ate it ;

and promptly tried the Bombus again as it lay on the ground,

but would not eat it. Immediately afterwards he eagerly ate an

Ocypus olens and three specimens of Péerostichus (see pp. 837-838).

I made one Volucella bombylans do duty for all the experiments

described above and had it intact at the end. It was not pecked

by any of the birds, presumably because I never offered it to one

until he had tried Bombus hortorwm a sufficient number of times

to reject it as unpalatable ; and there is no doubt in my opinion

that they did not distinguish between the bee and the fly.

Although Chilosia illustrata is also very like Bombus, the difference

in size is well marked. I suspect that in this circumstance lies

the explanation of the birds not confusing this species of fly with

the bee. They could judge the difference in size quite easily,

because the insects were held at the same distance from them.

856 MR. R. I. POCOCK ON THE

List of THE MAmmats, Brrps, AND REPTILES USED FOR THE

EXPERIMENTS.

MAMMALS.

Mona Monkey (Cercopithecus mona), Nigeria. Diana Monkey

(Cercopithecus roloway), Gold Coast and Guinea. Dent’s

Monkey (Cercopithecus denti), Ituri Forest. Vervet Monkey

(Cercopithecus pygerythrus), Cape Colony. Yellow Baboon

(Papio sphinx), Nigeria. Ceylonese Macaque (J/acacus

puleatus), Ceylon.

Although feeding mostly upon fruits, roots, and vegetables of

various kinds, all the Monkeys of the Old World eat insects as

well.

Capuchins (Cebus, spp. 2).

Several immature specimens, belonging to undetermined species

inhabiting the forests of the northern parts of South America.

Douracouli (Wyctipithecus trivirgatus).

A nocturnal Monkey from the Amazons.

Lion Marmoset (Zeontocebus rosalia). Pinché Marmoset

(Leontocebus adipus). Red-handed Marmoset (Leontocebus

rufimanus). Common Marmoset (Callithrix jacchus).

Although vegetable feeders in the main, the South American

Monkeys and Marmosets seem more addicted to an insect diet

than the Monkeys of the Old World.

Grey Lemur (Hapalemur griseus). Crowned Lemur (Lemur

coronatus). Black Lemur (Lemur macaco). White-fronted

Lemur (Lemur fulvus albifrons). Mongoose Lemur (Lemur

mongoz).

Lemurs inhabit Madagascar. They do not appear to be partial

to insects.

Suricate or Meerkat (Suricata swricatta).

Cape Colony. Feeds on small animals of various kinds and

particularly insects and their grubs (W. LZ. Sclater).

Yellow Meerkat (Cynictis penicillata).

Cape Colony. Feeds on small birds, mammals, eggs, and insects

(W. L. Sclater).

Banded Mongoose (Crossarchus fasciatus).

South and East Africa. Feeds on insects, fruits, seeds, eggs,

snails, etc., according to Bohm.

Common Indian Mongoose (J/wngos mungo), from India, and

McCarthy’s Mongoose (Ifungos fulvescens), from Ceylon, live on

small mammals, birds, reptiles, insects, and fruit. The White-

tailed Mongoose (Mungos albicauda), from Africa south of the

PALATABILITY OF SOME BRITISH INSECTS. 857

Sahara, does not, so far as is known, differ in diet from the other

species just mentioned.

Marsh Mongoose (Mungos galera).

West and South Africa. An amphibious species feeding

mainly it is alleged upon crabs, fishes, frogs, and insects.

Grison (Grison furax=Galictis vittata).

A musteline carnivore from the Argentine, feeding upon

small mammals and birds but also fond of fruit.

Common Armadillo (Dasypus villosus).

Argentine. Feeds on insects, grubs, worms, carrion, and

vegetable matter.

BIRDS.

Cape Robin-chat (Cossypha caffra).

Range. East Africa to Cape Colony.

Food. Chiefly insects, spiders, and worms; also berries and

small fruit (Sclater d: Stark).

Common Thrush (Vurdus musicus).

Range. Palearctic Region, locally migratory.

Food. Insects, worms, fruit, etc.

Orange-headed Ground-Thrush (Geocichla citrina).

Range. The Himalayas up to 5000-6000 ft., Assam and

Tenasserim.

Blue Rock-Thrush (Geocichla (Monticola) cyanus).

Range. From South Europe and North Africa to Turkestan,

Tibet, the Himalayas, and Burma.

Common Rock-Thrush (Geocschla (Monticola) saxatilis).

Range. C. & S. Europe to C. Asia, N.E. Siberia and N. China.

Wood-Thrush (Hylocichla mustelina).

Range. Eastern North America, Central America to Guatemala.

Dial Bird (Copsychus saularis).

Range. Ceylon, India, ascending the Himalayas up to 5000 ft. ;

Burma and Tenasserim.

Shama (Crttocincla macrura).

Range. Ceylon, India and Burma.

Blue-bird (Szalia sialis).

Range. Eastern North America to a little west of the Missouri

River.

Food. Insects of various kinds ; also ripe fruits.

American Cat-bird (Galeoscoptes carolinensis).

Range. South-eastern United States to the Missouri, migrating

southwards in the winter.

Food, Insects, fruit and seeds,

858 MR. R. I. POCOCK ON THE

Mocking Bird (Wimus polyglottus).

Range. Southern United States from the Atlantic to the high

central plains ; locally migratory.

Food. Insects and fruit.

Cuban Mocking Bird (Mimus orpheus).

Range. Jamaica, Porto Rico, Haiti, Cuba.

Saturnine Mocking Bird (imus saturninus).

Range. Brazil.

Great Tit (Parus major).

Range. Widely distributed in the Palearctic Region. Locally

migrating but mostly resident.

Food. Insects and seeds.

Pekin Robin (Liothrix luteus).

Range. Himalayas from Simla to Bhutan ; extending also into

China ; resident.

According to E. W. Oates the food of this bird consists of

berries, fruit, seeds, and insects.

Pied Grallina (Grallina australis).

Range. Australia, generally distributed.

, Food. Insects (Gould).

White-eared Bulbul (Pycnonotus lewconotus).

Range. Persia; Sind, the Punjab, the N.W. Provinces of

India, and Central India as far east as Hoshargabad.

White-cheeked Bulbul (Pycnonotus leucogenys).

Range. Afghanistan ; the Himalayas from Murree to Bhutan,

up to 7000 ft.

Red-vented Bulbul (Pycnonotus hemorrhous).

Range. Ceylon; India roughly to the foot of the Himalayas.

According to E. W. Oates the Indian species of Bulbuls feed

chiefly upon fruit.

Syrian Bulbul (Pycnonotus xanthopygus).

Range. N.E. Africa, Arabia, Palestine, Cyprus.

Black-crested Bulbul (Otocompsa flaviventris).

Range. Nepal to Cochin China.

In the course of my experiments I noticed that Bulbuls of

different species were very keen on butterflies ; of beetles and

crawling insects generally they took little if any notice; but the

moment a butterfly was let loose in the aviary they were all on

the move. From this I infer that they are great butterfly-

hunters in their own countries,

PALATABILITY OF SOME BRITISH INSECTS. 859

Orange-headed Laughing Thrush (7rochalopteron

erythrocephalumy).

Range. Himalayas, from Chamba to Nepal up to 7000 ft.

Black-chinned Laughing Thrush (Zvochalopteron nigrimentum).

Range. Himalayas from Nepal to Assam (7000 ft.).

According to E. W. Oates the food of the Laughing Thrushes

(Trochalopteron) is the same as that of the Jay-Thrushes

(Garrulaz).

Spectacled or Melodious Jay-Tbrush (Trochalopteron canorwm).

Range. China; Shanghai, Amoy, Fokien, Chekiang.

Black-headed Sibia (Sibia capistrata).

Range. Himalayas from Hazira to Bhutan, 5000-8000 ft. ;

resident.

Collared Jay-Thrush (Garrulaz picticollis).

Range. China: Chekiang, Fokien.

White-crested J ay-Thrush (Garrulax leucolophus).

Range. Himalayas to Assam and Burma in the hill-tracts.

According to E. W. Oates the Indian species of Garrulax feed

upon every sort of insect and smaller reptiles, and probably also

on fruit.

Grey Struthidea (Struthidea cinerea).

Range. South-eastern Australia; resident.

Food, Insects, particularly beetles.

Spotted Oriole (Oriolus maculatus).

Range. Sumatra, Java, Borneo.

Harmonious Shrike-Thrush (Collyriocinela harmonica).

Range. Australia; N.S. Wales and 8. Australia.

Food. Insects (Gould).

White-eyebrowed Wood-Swallow (Artamus superciliosus).

Range. Interior of South Australia.

Food. Insects (Gould).

Masked Wood-Swallow (Artamus personatus).

Range. South Australia, locally migratory.

Food. Insects (Gould).

Red-backed Shrike (Lanius collurio).

Range. Europe, migrating in the autumn and winter into

Western India and to South Africa.

Food. Insects; small birds etc.

Fantailed Flycatcher (Lhipidura tricolor).

Range. Australia, widely distributed.

Food. Insects of various kinds (Gould).

860 MR. R. I. POCOCK ON THE

Garrulous Honey-eater (yzantha garrula).

Range. South Australia, Tasmania,

Food. Honey and insects (Gould).

Black Tanager (Z’achyphonus melaleucus).

Range. Costa Rica through Panama, Venezuela, Ecuador to

Bahia.

Scarlet Tanager (Rhamphocelus brasilius).

Range. South-eastern Brazil.

Cayenne Tanager (Calliste cayana).

Range. Guiana, Venezuela, Ecuador, Peru.

Green Hangnest (Ostinops viridis).

Range. Guiana, Brazil, Ecuador.

Yellow Hangnest (Cassicus persicus).

Range. Trinidad, Guiana, Ecuador, Bolivia, Brazil.

Common Hangnest (/cterws vulgaris).

Range. Colombia, Venezuela.

Brazilian Hangnest (Jcterws jamaicat).

Range. North Brazil.

,Yellow-crowned Hangnest (Jeterus chrysocephalus).

Range. Guiana, Venezuela, Kcuador, Brazil.

Larger Hill Mynah (Gracula intermedia).

Range. India: the south-eastern Central Provinces, the lower

ranges of the Himalayas from Kumaon to Assam, thence into the

Malay Peninsula.

Small Hill Mynah (Gracula religiosa).

Range. Ceylon and Southern India.

According to EK. W. Oates these two species of Mynah are

resident or only locally migratory and live exclusively upon

fruit. ,

Chinese Mynah (Acridotheres cristatellus).

Range, China: Shanghai, Hainan, Formosa; Philippine

Islands.

Pied Mynah (Stwrnopastor contra).

Range. Central and South India to Assam and Burma.

Javan Pied Mynah (Sturnopastor jalle).

Range. Sumatra, Java, Borneo.

Black-winged Grackle (Giraculipica melanoptera).

Range. Java.

PALA'TABILITY OF SOME BRITISH INSECTS. 861

Spotted Bower Bird (CAlamydodera maculata).

Range. New South Wales.

Food. Principally fruit and grain (Gould).

King Bird of Paradise (Cictnnurus regius).

Range. New Guinea.

Food. Fruit and insects.

Canadian Jay (Perisorews canadensis).

Range. Canada and the Northern States of the Union.

Food. Insects ; eggs, flesh ; leaves of fir trees (Audubon).

Hooded Crow (Corvus cornix).

Range. Palearctic Region.

Food. Omnivorous (eggs, carrion, young birds, etc.).

White-collared or Pied Crow (Corvus scapulatus).

Range. Atvica south of the Sahara.

Food. Omnivorous, with partiality for flesh food.

White-backed Piping Crow (Gymmorhina leuconota).

Range. 8S. Australia, New South Wales.

Food. Mostly insects (Gould).

Long-billed Butcher Crow (Oracticws destructor).

Range. Australia.

Food. Chiefly insects.

Sulphury Tyrant (Pitangus sulphuratus).

Range. Guiana, Ecuador, Peru, Brazil.

Food. Mostly insects and animal food of various kinds as well

as fruit.

Greater Spotted Woodpecker (Dendrocopus major).

Range. Palearctic Region.

food. Insects.

Common Laughing Kingfisher (Dacelo gigantea).

Range. New South Wales and South Australia.

Leach’s Laughing Kingfisher (Dacelo leachit).

Range. North-east coast of Australia.

Buff Laughing Kingfisher (Dacelo cervina).

Range. Kast and North Austvralia.

Food. These great Kingfishers feed mainly upon reptiles and

insects, but also upon rats and inice.

Hlate Hornbill (Ceratogymna elata) and

Black Hornbill (C. atrata).

Range. W. Africa, Nigeria, etc.

Food. Insects ; snakes, small mammals, ete.

Proc. Zoou. Soc.—1911, No. LIX. 59

862 : MR. R. I. POCOCK ON THE

Ground Hornbill (Bucorax abyssivicus).

Range. North Africa south of the Sahara. ny

Food. Insects, snakes, frogs, lizards (Stark and Sclater writing

of the closely allied southern species B. caffer).

Hoopoe (Upupa epops).

Range. Southern Palexaretic Region from Scandinavia and the

British Islands to Japan, migrating in winter to North Africa,

Arabia and India.

Food. Ground insects, beetles, grasshoppers and ants.

Cartagenian Motmot (Momotus subrufescens).

Range. From Panama, Colombia, and Venezuela to Matto Grosso.

Great Barbet (egalema virens).

Range. China and Upper Burma.

Levaillant’s Barbet (Zrachyphonus caffer).

Range. 8. Africa, Natal, the Transvaal, Rhodesia, ete.

Food. Fruits, berries, leaves, and insects such as termites

(Stark & Sclater).

Green Toucanet (Aulacorhamphus sulcatus).

Range. Venezuela; Colombia.

Pearl-spotted Owl (Glaucidiwm perlatum).

Range. Africa south of the Sahara.

Food. Mostly insects (grasshoppers, termites); also mice and

lizards (W. L. Sclater).

White-eared Scops Owl (Scops leucotis).

Range. Africa south of the Sahara to the Orange River.

Food. Chiefly insects, like grasshoppers; also rats and mice

(W. L. Sclater).

Prince of Wales Pheasant (Phasianus principalis).

Range. North-western Afghanistan and North-east Persia.

Reeves’s Pheasant (Phasianus reevesit).

Range. Mountains of Northern and Western China, extending

as far east as Kiu-Kiang.

Elliot's Pheasant (Calophasis elliott).

Range. Mountains of South-eastern China.

Silver Pheasant (Genneus nycthemerus).

Range. South China, Fokien and Chekiang.

Vulturine Guinea Fowl (Aerylliwm vulturinum).

Range. Kast Africa from the Pangani River westwards to

Kilimanjaro and northwards to Somaliland.

PALATABILITY OF SOME BRITISH INSECTS. 863

Pucheran’s Guinea Fowl (Guttera pucherani).

Range. Kast Africa: Zanzibar to the Tana River and thence

westwards into the interior.

N. American Wild Turkey (Meleagris americana).

Range. Formerly widely distributed in the United States of

America. Not migratory.

food. Beechnuts, acorns, berries, green-shoots, etc. ; also grass-

hoppers, and other insects (Bendire).

Honduras Turkey (Jeleagris ocellata).

Range. Central America: Guatemala, Yucatan, Honduras.

Food. Probably of a similar nature to that of JZ. americana.

Long-tailed Partridge (Dendrortyx leucophrys).

Range. Highlands of Guatemala and Costa Rica (Ogilvie-

Grant).

Brush Turkey (Catheturus lathami).

Range. North-east and Kast Australia.

Crested Curassow (C'raa alector).

Range. Northern part of South America: British Guiana,

Colombia, Rio Negro, ete.

Globose Curassow (Crax globicera).

Range. Central America: Western Mexico to Honduras and

Cozumel Island.

Yarrell’s Curassow (Craw carunculata).

Range. South-eastern Brazil from Rio Janeiro to Bahia.

Red-tailed Guan (Ortalis ruficauda).

Range. Venezuela and the island of Tobago.

Most Game-birds, especially when young, eat insects as well as

grain, nuts, and green-food.

Australian Bustard (Hupodotis australis).

Range. South and Western Australia.

Food. Seeds, vegetables, grasses, and insects (G'owld).

Vigors’s Bustard (Otis vigorsit).

Range. 8. Africa: Cape Colony, Natal, ete.

Food. Seeds, insects, small reptiles (Stark dg Sclater).

Ludwig’s Bustard (Otis ludwig?).

Range. 8. Africa: Cape Colony, Natal, Orange River Colony,

S. Transvaal; partially migratory within this area.

Food. Mostly beetles, caterpillars, and other insects (Stark &

Sclater).

The food of Bustards is probably much the same everywhere.

The diet is essentially mixed, and consists of grain, green-shoots and

leaves insects, smal] mammals (mice) and reptiles,

864 PROF. E. B, POULTON ON THE

Two-striped Thickknee (Hdienemus bistriatus).

Range. Mexico through Central America to Venezuela

Food. Insects, worms, snails, ete.

Trumpeter (Psophia crepitans).

Range. Brazil.

Food. Fruits, seeds, insects.

Cariama or Seriema (Cariama cristata).

Range. South-east Brazil.

Food. Reptiles and smal! mammals for the most part.

Abbott’s Rail (Hallus abbotti).

Range. Assumption Island.

Black-tailed Water-hen (Zribonyx ventralis).

Range. Australia, south of the 25th parallel ; locally migratory.

Kagu (Rhinochetus jubatus).

Range. New Caledonia.

Sun-Bittern (Hurypyga helias).

Range. Northern countries of the Neotropical Region.

Food. Mostly insects.

REPTILIA

The Green Lizard (Lacerta viridis), from Central and Southern

Europe; the Wall Lizard (Lacerta muralis), from Central and

Southern Europe; the Filfola Wall Lizard (LZ. muralis filfolensis),

from Filfola, near Malta; Dugés’s Lizard (Lacerta dugesii), from

Madeira; the Sand Lizard (Lacerta agilis), from North and

Central Europe ; and the Black-spotted Lizard (Algiroides mgro-

punctatus), from Dalmatia, feed mainly upon insects, worms, and

small slugs.

Glass Snake (Ophisaurus apus).

South-eastern Europe. Feeds on small mammals, reptiles,

slugs, ete.

Notes upon some of the above described Huperiments by

Prof. E. B. Poutton, F.R.S., F.Z.8.

Pages 815-820.

The experiments on the Pierinw support the conclusion that

the perfection of the under surface procryptic resemblance affords

a true criterion of the degree of palatability.

P. brassice, with its conspicuous gregarious larva, and imago

larger and less cryptically coloured than the other three species

PALATABILITY OF SOME BRITISH INSECTS. 865

(although nearly the same as P. rape in this respect), was

distinctly the least palatable of the four. On the other hand,

P. napi and EF. cardamines, in which the cryptic resemblance is

carried to its highest pitch, appeared to be most palatable; but

a larger number of experiments is greatly to be desired.

The results obtained in the two former species are of much

interest in relation to the experiments upon J/elanargia galathea

(p. 827).

Pages 820-822.

The evidence that J’. wrticee is not very palatable agrees with

my own experiments * with a Marmoset; and I obtained the

same results with V. 7o when offered in considerable numbers to

lizards. It is probable that the procryptic under surface of the

Vanessas is chiefly related to the attacks of mammals and of very

hungry birds during the long hibernating period. The special

interest in the eye-spots of V. io manifested by the Syrian Bulbul,

accords with previous observations on other insects and other

insect-eaters, Reptilian as well as Avian ft.

Pages 823-825.

The evidence of a certain amount of unpalatability in

Brenthis (Argynnide) is consistent with the degree of proeryptic

defence attained in this genus. It is also of much interest in

connexion with the experiments on Avaschnia levana, the early

or levana form of which is probably a mimic of the species of

Brenthis. The examples of the Araschnia tested by the author

(pp. 823-824) were of the form prorsa, belonging to the later

brood, and generally looked upon as mimics of the White Admiral

(Limenitis sibylla), which appears upon the wing at about the

same period. The experiments here recorded prove that the

mimic is certainly unpalatable to several birds, and support the

conclusion that the resemblance is Miillerian or Synaposematic.

The evidence, so far as it goes, points indeed to the inference

that Araschnia is more unpalatable than its Brenthis model. A

few experiments on the imago of L. sibylla made by Mr. Pocock

in the summer of 1910, also indicated that the prorsa form is

more unpalatable than the Limenitis. There is, however, nothing

improbable in a Miillerian mimic being more highly protected than

its model. The role of model is related to many characteristics,

and relative abundance, conspicuousness, and extent of range

may play their part as well as relative unpalatability. Thus it is

probable, from its habits and flight, that the Eastern Kuropean

Neptis lucilla, W.V.(=sappho Pall.) is more distasteful than its

Limenitis models, but the latter are widespread and abundant

species, and it is reasonable to suppose that the memories of

European insect-eating animals are more deeply impressed by

their pattern than by that of the Veptis.

* Trans. Ent. Soc. Lond. 1902, p. 442.

+ ‘Essays on Evolution ’ (Poulton), 1908, p. 210: see also p. 326,

866 PROF. E. B. POULTON ON THE

Pages 825-827.

The desirability of experiments upon the palatability of the

genus Melitea was suggested by the study in 1908 of a collection of

butter flies from the Tian Shan or Celestial Mountains in Western

Mongolia, Small as it was, the collection was sufficient to show

that Melitea is a dominant element in the insect fauna of the

locality. The large “ Skipper,” Hesperia antonia Spey, was also

abundantly represented, and I was at once struck with the marked

resemblance which its under surface would bear in the position of

rest to that of the species of Melitea. The striking feature of the

latter genus is supplied by two black-bordered orange bands

which cross the hind wings and stand out conspicuously against

the cream ground-colour. These two bands, the outer with its

festooned, the inner with its irregular borders, present a highly

characteristic appearance. ‘The small portion of the fore wing

under surface exposed in the position of rest conforms, as is

usual in butterflies, to the pattern of the hind and appears as a

slight extension of its area. In spite of differences in detail, the

two orange bands of the Hesperid closely resemble those of

Melitea, and in all essential respects the exposed under surface of

the former reproduces that of the latter. In the Skipper the

outer margin of the outer band is cut mto internervular con-

cavities, while the inner bulges into corresponding convexities:

in the Nymphaline, concavities are seen along both borders. The

orange of the bands and the tint of the ground-colour—white

between the bands, greyish elsewhere—are also much paler in the

Skipper, but the orange pigment is probably quite different from

that of Melitea and may rapidly fade. It is also interesting to

note that the orange bands of the under surface are represented

by black bands on the upper surface of the Skipper but by orange

bands on that of the Nymphaline. The allied Hesperia side Esp. Ls

with golden bands, is doubtless a co-mimie with H. antonia, while

im 2 fbr closely related species, 7. carthami Hiibn., the dark

bands have gained a bronzy greenish or yellowish tinge, probably

indicating the kind of variation out of which the pattern of the

two first-named species was produced by selection.

Probable evidence that Melitwa is a specially protected genus is

supplied by the well-known habits of the three British species

aurinia (=artemis) Rott., cinwia L., and athalia Rott. All are

known to be gregarious in the larval state, and so abundant in

confined localities that they may be described as gregarious in

the perfect state also. All are slow-flying and conspicuous on the

wing and at rest between the flights, while individuals have been

observed to “sham death” when captured. Putting all the facts

together, it appeared probable that we have an interesting addition

to the list of mimics among the Palearctic butterflies, a list

which is remarkably short in the western section of the Region.

Mr. Pocock kindly consented to test the hypothesis that J/elitcea

possesses the distasteful qualities of a model for mimicry, and

Commander J. J, Walker kindly helped to obtain material for

PALATABILITY OF SOME BRITISH INSECTS 867

the experiments which are recorded on pp. 825-827. The results

as a whole leave little doubt that Welitea is distasteful to many

birds, and that it does actually possess the qualities which would

render it an advantageous model for the Hesperiide.

Pages 827-830.

The experiments on Melanargia galathea are of peculiar interest.

The northern belt Satyrinz of this genus, with their white ground-

colour, stand out from the rest of their group. The under surfaces

are conspicuous, the species slow-flying and so abundant locally

that they may be called gregarious. The observations here

recorded show that they are also distinguished by their greater

distastefulness from other common northern Satyrines. The

appearance of the species of Melanargia, especially on the wing,

is markedly Pierine-like, and it is here also probable that a highly

distasteful genus has mimicked an assemblage of species which,

although generally less unpalatable, are excessively widespread and

abundant in individuals (see also p. 865).

Pages 830-831.

The experiments upon Lyczenide, Nemeobiine, and Hespertide

were not sufficiently numerous to form the foundation for safe

conclusions.

Pages 831-855.

The remaining experiments, for the most part, afford valuable

confirmation of previous work, but they also raise new questions

of great interest. Confirmation is afforded by the evident un-

palatability of Zygena, Hucheha, and Abrawas among the moths,

of the Saw-fly larve, of the Telephoride, Phytophaga, and Cocci-

nellidee among the beetles, and of the Hemiptera, as also by the

special and peculiar defensive secretions of the Carabide and, in

sharp contrast to all these observations, by the palatability of the

procryptically coloured moths and larvee.

Pages 847-848.

The apparent distastefulness of the humble-bee-like Volucella

bombylans suggests conclusions of so much importance and interest

that abundant confirmation is essential, and should be readily

available with so common a species.

Pages 848-852.

Experimental evidence that the Aculeate Hymenoptera possess

some special defence independently of the stings of the females is

now obtained for the first time. It was suggested as probable by

the present writer in 1904, as a result of the observation that the

males of the bee Sphecodes emerge in immense numbers and

form complex mimetic associations, before the appearance of the

females, as also from the consideration that the Braconide are

extensively mimicked *.

* Trans. Ent. Soc. Lond. 1904, pp. 645-6.

868 ON THE PALATABILITY OF SOME BRITISH INSECTS.

Page 849.

Although so many insectivorous animals in confinement dis-

regarded the special defence of Yormica rufa, there can be little

doubt that such defence is very effective in the wild state. It is

impossible on any other hypothesis to account for the conditions

under which the species exists—swarming in vast numbers in

restricted areas and an easy prey to any enemy that would dare

to attack.

A very important conclusion is suggested by several of the

experiments recorded in this memoir, namely, that the tastes of

mammals and birds are widely different. The author points out

that the defence of the ground-beetles appealed more strongly

to the mammals than to the birds, but it was also apparent in

many of the experiments that the unpalatability of conspicuous

Lepidoptera was, conversely, far more obvious to the birds than

to the mammals. Jn view of the part which birds are believed

to play in the production of mimetic resemblance, it is obvious

that this inference may be highly significant.

ADDENDUM.

Dr. P. Caatmers Mrrcneni’s Memoir ‘On Longevity and

Relative Viability in Mammals and Birds,” P. Z.8.

1911, p. 425.

[The Rev. F. C. R. Jourpatn has kindly called my attention to

the fact that I have overlooked a valuable paper “‘On the

Duration of Life of the Animals in the Zoological Garden at

Frankfort-on-the-Main,” by Director Dr. Max Scumunpt, P. Z.8.

1880, p. 299, and containing many valuable figures as to

longevity|.—P. C. M., Aug. 2, 1911.

No. 98.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

June 13th, 1911.

Epwin T. Newton, Esq., F.R.S., in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

Mr. H. G. Purmmer, F.R.8., Pathologist to the Society, pre-

sented a Report on the Pathological Examination of Rats caught

in the Regent’s Park and in the Society’s Gardens. 500 rats

had been examined between the lst of January and the 17th of

May, 1911, all in a precisely similar manner. The spleen, lungs,

glands, and blood were examined microscopically; and from

any animal which looked in any way unhealthy cultures were

made.

The results were summarized as follows :—5 rats were caught

in the Park, and 495 in the Gardens: 283 of these were males

and 217 females.

3 rats had tubercle, 10 had tapeworm cysts in the liver, 49 had

Trypanosoma lewisi in their blood, 2 had empyema (not tubercular),

1 had a tumour of the lower jaw (the result of an old injury), and

1 had pleuritis and hydrothorax (not tubercular).

Bacteria were found in 71 rats: in 40 in the lungs, and in 31

in the spleen.

Saccharomycetes were found in the lungs of 16 rats.

Fleas were found on 4 rats, and lice on 8 rats.

The general condition of the rats was very good, and in none

was anything at all suspicious found.

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent's Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications; but it may be obtained on the

day of publication at the price of Sixpence, or, if desired, sent post-free for

the sum of Six Shillings per annum, payable in advance.

Mr. R. I. Pocock, F.R.S., ¥F.Z.S., the Society’s Curator of

Mammals, exhibited the skin and skull of a Crested Rat

(Lophiomys ibeanus), sent from British Kast Africa by Mr. R. B.

Woosnam, and after drawing attention to the characters of the

skull in this genus, said that in his opinion the coloration was

for the purpose of self-advertisement and rendered the animal

conspicuous at night.

Dr. R. W. SHuFELDT, C.M.Z.S., sent for exhibition a photograph

he had taken of a living specimen of a male albino Woodchuck,

Arctomys monax, that had been sent to him from Virginia,

WESssA

Mr. R. E. Houpine exhibited and made remarks upon the

Horns of a Highland Ram, a Fallow Deer, and a Roebuck, which

were fused at the base, and also the skull of a coursing Greyhound

with abnormal dentition.

Dr. R. E. Draks-Brocxkmay, F.Z.8., read a paper on Antelopes

of the genera Madoqua and Rhynchotragus found in Somaliland,

specimens of most of which were exhibited at the Meeting. He

made general remarks on all the Dik-diks and gave a short

account of the species and subspecies, including the description

of a new form.

The Hon. Paut A. Meruuen, F.Z.8., communicated a paper

“On an Amphipod from the Transvaal,” in which he gave a

detailed description of a new freshwater Gammarid of the genus

Eucrangonyx, that had been found in caves in the Transvaal.

A paper was contributed by Mr. R. LypEKxKeEr on three African

animals. The first specimen was the skull of a Somali Rhinoceros,

a race for which the author adopted the name Rhinoceros bicornis

somalicus, Potocki. A Klipspringer skull from Northern Nigeria,

characterized by its great width and the peculiar form of the

lachrymal bone, was described as the type of a new race, Oreotragus

saltator porteust. Finally, a Gazelle from Algeria was referred

to a new species, Gazella hayi, agreeing approximately in size

with G. dorcas, but distinguished by the much straighter and

non-lyrate horns, each of which carried only about a dozen rings.

The face-markings were approximate to those of G. cuvierd.

A paper entitled ‘A Contribution to the Ornithology of Western

Colombia,” by Mr. C. E. HeLrtmayr, was communicated by Dr. P.

L. Scuarer, F.R.S. This memoir was based on a collection made

by Mr. M. G. Palmer in 1908 and 1909, which, though numbering

hardly 700 specimens, was of considerable interest and contained

many rare species, and also furnished information of importance

to students of zoogeographical problems.

No. 98.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

June 13th, 1911.

Epwin T. Newton, Esq., F.R.S., in the Chair.

The Minutes of the last Scientifie Meeting were confirmed.

Mr. H. G, Pumuer, F.R.S., Pathologist to the Society, pre-

sented a Report on the Pathological Examination of Rats caught

in the Regent’s Park and in the Society’s Gardens. 500 rats

had been examined between the Ist of January and the 17th of

May, 1911, all in a precisely similar manner. The spleen, lungs,

glands, and blood were examined microscopically; and from

any animal which looked in any way unhealthy cultures were

made.

The results were summarized as follows :—5 rats were caught

in the Park, and 495 in the Gardens: 283 of these were males

and 217 females.

3 rats had tubercle, 10 had tapeworm cysts in the liver, 49 had

Trypanosoma lewisi in their blood, 2 had empyema(not tubercular),

1 had a tumour of the lower jaw (the result of an old injury), and

1 had pleuritis and hydrothorax (not tubercular).

Bacteria were found in 71 rats: in 40 in the lungs, and in 31

in the spleen.

Saccharomycetes were found in the lungs of 16 rats.

Fleas were found on 4 rats, and lice on 3 rats.

The general condition of the rats was very good, and in none

was anything at all suspicious found.

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent's Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications; but it may be obtained on the

day of publication at the price of Sixpence, or, if desired, sent post-free for

the sum of Six Shillings per annum, payable in advance,

Mr. R. I. Pococx, F.R.S., F.Z.8., the Society’s Curator of

Mammals, exhibited the skin and skull of a Crested Rat

(Lophiomys ibeanus), sent from British East Africa by Mr. R. B.

Woosnam, and after drawing attention to the characters of the

skull in this genus, said that in his opinion the coloration was

for the purpose of self-advertisement and rendered the animal

conspicuous at night.

Dr. R. W. Suure.pt, C.M.Z.8., sent for exhibition a photograph

he had taken of a living specimen of a male albino Woodchuck,

Arctomys monax, that had been sent to him from Virginia,

U.S.A.

Mr. R. H. Hoipine exhibited and made remarks upon the

Horns of a Highland Ram, a Fallow Deer, and a Roebuck, which

were fused at the base, and also the skull of a coursing Greyhound

with abnormal dentition.

Dr. R. EH. Draxs-Brocremay, F.Z.8., read a paper on Antelopes

of the genera Madoqua and Rhynchotragus found in Somaliland,

specimens of most of which were exhibited at the Meeting. He

made general remarks on all the Dik-diks and gave a short

account of the species and subspecies, including the description

of a new form.

The Hon. Paut A. Meruuen, F.Z.8., communicated a paper

“On an Amphipod from the Transvaal,” in which he gave a

detailed description of a new freshwater Gammarid of the genus

Hucrangonyx, that had been found in caves in the Transvaal.

A paper was contributed by Mr. R. LyDEKKER on three African

animals. The first specimen was the skull of a Somali Rhinoceros,

a race for which the author adopted the name Phinoceros bicornis

somalicus, Potocki. A Klipspringer skull from Northern Nigeria,

characterized by its great width and the peculiar form of the

lachrymal bone, was described as the type of a new race, Oreotragus

saltator porteusi. Finally, a Gazelle from Algeria was referred

to a new species, Gazella hayi, agreeing approximately in size

with G. dorcas, but distinguished by the much straighter and

non-lyrate horns, each of which carried only about a dozen rings.

The face-markings were approximate to those of G. cuvieri.

A paper entitled ‘‘ A Contribution to the Ornithology of Western

Colombia,” by Mr. C. EK. HELLMAYR, was communicated by Dr. P.

L. Scuater, F.R.S. This memoir was based on a collection made

by Mr. M. G. Palmer in 1908 and 1909, which, though numbering

hardly 700 specimens, was of considerable interest and contained

many rare species, and also furnished information of importance

to students of zoogeographical problems.

No. 99.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

June 27th, 1911.

FREDERIOK GiLLeTt, Esq., Vice-President,

in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

The Secretary read a Report on the Additions that had been

made to the Society's Menagerie during the month of May

1911.

Dr. W. T. Caumay, F.Z.S., exhibited a number of living

specimens of the Brine Shrimp (Artemia salina) which had been

bred from Tidman’s Sea Salt. He remarked that this sea salt, as

sold in the shops, was found frequently to contain living eggs

of Artemia and that it was easy to obtain a supply of living

specimens. An 8 per cent. solution, allowed to stand for a few

days, produced a swarm of larve, and these could be fed on the

strained juice of green leaves and raised to maturity,

Mr. J. Lewis Bonunorz, M.A., F.Z.S., exhibited a pair of

Egyptian Desert-Mice (/eriones crassus) which showed a darker

and more rufous colour than normal examples. This coloration

had been artificially produced by keeping the animals in a moist

atmosphere at 80° Fahr. They were first exposed to these con-

ditions on the 7th of April and a month later were conspicuously

darker ; after that the darkening process still continued, but more

slowly, and they appeared for some time previous to being killed

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent’s Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications ; but it may be obtained on the

day of publication at the price of Sixpence, or, if desired, sent post-free for

the sum of Six Shillings per annum, payable in advance,

to have reached a limit to their darkening. During this same

period, and owing to the fine weather, other examples had been in

a temperature that rose during the day to 90°, falling at night to

60° or even lower. This had apparently produced no change in

their coloration. The change in the examples exhibited was

therefore due probably rather to the humidity than to the tempe-

rature. Mr. Bonhote was therefore inclined to think that the

pale colour of desert animals was due to the extreme dryness of

the atmosphere rather than to any special assimilation of their

colour to the surroundings.

Mr. R. I. Pocock, F.R.S., F.Z.8., Superintendent of the

Gardens, exhibited photographs of a hybrid between the Somali-

land Wild Donkey (Zquus asinus somaliensis) § and the Mountain

Zebra (Equus zebra) 2, and remarked that although Mountain

Zebras and domestic Asses had been previously crossed, no hybrid

had hitherto been produced between this Zebra and the Somaliland

race of Hgwus asinus. The period of gestation was twelve months

and three weeks exactly. The foal showed much greater resem-

blance to Hqwus asinus than to LH. zebra, the body being practically

unbanded, except for the spinal and shoulder stripes. The stripes

on the legs extended as high as the level of the belly and were

broader and more numerous than in the sire and narrower and

fewer than in the dam. In the presence of the shoulder stripe,

the spinal stripe, and the stripe on the base of the ear, the foal

resembled typical examples of Hquwus asiwus and differed from its

actual sire, which was without those marks.

Mr. D. Seru-Surru, F.Z.8., the Society’s Curator of Birds,

exhibited two immature Black-backed Porphyrios (Porphyrio

melanonotus) which had been bred in the Gardens, and remarked

upon their possession of a well-developed claw on the pollex.

Although these wing-claws were said to be functional only in the

Hoatzin amongst living birds, the exhibitor believed that they

were so also in the present species as also probably in the Common

Moorhen, these birds using these appendages to assist them in

climbing amongst reeds and other herbage.

The SECRETARY remarked that on a recent visit to the Ostrich

Farm of Mr. Carl Hagenbeck at Stellingen, near Hamburg, he

had seen in the incubator fertile eggs of Struthio massaicus from

German East Africa, S. australis from South Africa, and S. mo-

lybdophanes from Somaliland, the eggs all having been laid at

Stellingen. A. Reichenow (‘Die Vogel Africas,’ vol. i. p. 7) had

already described and figured certain specific differences in the

number and arrangement of the pits on the eggs of these species.

He himself had been interested to notice that the eggs of the

Masai Ostrich were much larger than those of the others, more

spherical in shape, and very smooth and porcelanous in texture.

Those of the Cape Ostrich were somewhat similar in shape and

texture, but were much smaller; Mr. Hagenbeck had informed

him that a pair of the Masai Ostrich bred by himself and sent out

to the Cape were regarded by expert ostrich farmers there as

unusually large birds. The eggs of the Somali Ostrich were

larger than those of the Cape Ostrich, but smaller than those of

the Masai species, and were markedly oval in shape with a rougher,

less polished surface.

The SzcrETARyY remarked that on his recent visit to Mr. Hagen-

beck’s Zoological Park at Stellingen, near Hamburg, he had the

pleasure of seeing a fine young pair of the common African

Rhinoceros, obtained from British East Africa, the exact locality

being unknown. The male closely resembled the ordinary figures

and mounted examples of the species, in that the skin appeared

to be smoothly stretched over the sides of the body, but the ears

were fringed with long tufts of hair. The female, on the other

hand, had ne hair on the margin of the ears, and the general

external appearance was very different. At first sight it seemed

as if it were in very poor condition, the ribs standing out through

the skin, but closer inspection showed that in reality the skin

of the flanks was disposed in thick, permanent folds, arranged

roughly like ribs. Thinking it possible that these differences

might indicate the existence of distinct races of the Rhinoceros,

on returning to London he had at once examined the Society’s

own pair of examples of this species, both of which had come from

British East Africa, probably somewhere near Nairobi. The

female, purchased in 1906, had the ears unfringed with hair, like

those of Mr. Hagenbeck’s female, but the rib-folds on the skin

were no more than indicated, although there were very heavy

permanent folds round the neck. In the male, obtained in the

current year from Nairobi as part of the King’s African Collec-

tion, the ears were fringed with hair as in Mr. Hagenbeck’s male,

whilst the rib-like folds on the skin were extremelystrongly marked,

as in the case of Mr. Hagenbeck’s female. The presence or

absence of the marginal fringe on the ears was therefore probably

either an individually variable or a sexual character. In the

absence of knowledge of the exact provenance of all the four

examples, nothing could be said as to whether or no the presence

of the rib-like permanent folds on the body were racial. Their

existence, however, as well as the presence of the heavy fold round

the neck, showed that it was not correct to distinguish the Asiatic

Rhinoceroses from those of Africa by the presence in the former

of permanent skin-folds. The neck-fold was almost identical in

both, whilst, although they were differently arranged, deep body-

folds occurred in both.

Dr. A. Suita Woopwarp, F.R.S., V.P.Z.S., communicated a

paper by Dr. R. Broom, C.M.Z.S., entitled ‘“‘On some new South

African Permian Reptiles.’

Mr. F. EK. Bepparp, M.A., F.R.S., F.Z.S., Prosector to the

Society, read a paper ‘“‘On two new Genera of Cestodes from

Mammals,” based on specimens collected from animals in the

Society’s Gardens.

A paper was received from Miss Ruta Harrison, entitled

“Some Madreporaria from the Persian Gulf; with a Note on the

Memoir and some Further Notes on Pyrophyllia inflata by

Sypney J. Hickson, M.A., D.Sc., F.R.S., F.Z.8.” This memoir

dealt with a collection of corals made by Mr. F. W. Townsend,

the most interesting species obtained being P. inflata and Tre-

matotrochus zelandice, the latter of which was identical with the

specimens from Cook’s Straits, New Zealand, described as Cono-

cyathus zelandic by Prof. Martin Duncan. A new species of

Heterocyathus was described, and Professor Hickson appended a

note on the affinities of Pyrophyllia.

Mr. C. L. Bounencer, M.A., F.Z.S., contributed a paper “ On

Variation in the Medusa of Merisia lyonsi,” based on an exami-

nation of 400 specimens. Nearly 14 per cent. of these were

found to be abnormal, and to fall naturally into two well-marked

groups containing completely distinct phenomena. The author

discussed these separately and in detail, and stated that he knew

of no-form in which such a variety of abnormalities occurred as

in Merisia.

Mr. Cyrit CrossLanD, F.Z.8., presented two papers, entitled

“The Marginal Processes of Lamellibranch Shells” and ‘“‘ Warn-

ing Coloration in a Nudibranch Molluse and in a Chameleon.”

Asa pendent to the second of these Sir CoartEs Exzot, K.C.M.G.,

C.B., F.Z.S., contributed a paper on ‘‘Chromodorids from the

Red Sea collected and figured by Mx. Crossland,” containing an

account of three species of Chromodoris, which were noteworthy

as being varieties of known species or forms hitherto imperfectly

described.

This Meeting closes the Session 1910-1911. The next Meeting

of the Society for Scientific Business will be held on Tuesday,

October 24th, 1911, at half-past Hight o'clock p.m.

The following Papers have been received :—

1. A. D. Imus, D.Sc., B.A.

On some Collembola from India, Burma, and Ceylon, with a

Catalogue of the Oriental Species of the Order.

2. Prof. P. P. Susuxin, C.M.ZS.

Ontogenetical Transformations of the Bill in Ardea cinerea.

3. EK. P. Stepsine, F.L.S., F.Z.8.

Game Sanctuaries and Game Protection in India.

Communications intended for the Scientific Meetings suould

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL SoctEty OF LONDON,

Recent’s Park, Lonpon, N.W.

July 4th, 1911.

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Exhibitions and Notices (continued).

- Page

Mr. R. I. Pocock, F.B.S., F.L.S., F.Z.8. Exhibition of some hair from the “ puppy coat”

of a Grey Seal (Halicherus grypus) ......++++0s Rertiblonenan do ac Ghd 6G cmH OUIDO He 696

Mr. EB. G. Bounencer. Exhibition of living male specimens of the Midwife Toad (Alytes

obstetricams) carrying the eggs... 0. os ee nsw e se nsec ne wee cee nee e en aes 696

Mr. A. HE. Anpersoy. Exhibition of photographs of fossil mammals in the American

Museum of Natural History..............- Peongour Bhs taeee ez adaicfaievabareas b netieborage. eyerels 696

PAPERS.

26. On the Amphipod Genus Leptocheirus. By E. W. Sexton, Marine Biological Labora-

tory, Plymouth. (Plates XVII.XIX. and Text-fig. 146.) ....-.....-.-2..20--- «. 561

27. Notes on Marine Ostracoda from Madeira. By G. Srzwarpson Brapy, M.D., LL.D.,

D.Se., F.R.S., C.M.Z.8. (Plates XX.—XXIT.) 1.0... cw eee eee eee eee eee eee Toy 8B)

28. On Colour and Colour-pattern Inheritance in Pigeons. By J. Lewis Bonuors, M.A.,

F.LS., F.Z.8., and F. W. Swanzey, ¥.Z.8. (Plates XXIII.—-XXVI.) .............- 601

29. Contributions to the Anatomy and Systematic Arrangement of the Cestoidea. By

Frank E. Bepparp, M.A., F.R.S., F.Z.S., Prosector to the Society. (Text-figs. 148-159.) 626

30. On the Natural History of Whalebone Whales. By J. A. Morcu, Christiania.

(Bex totes NGO UG ao) tare yeteneteiavel afegovs) ia" 20 -ths (olal-e’e “a. ale eleveies @tietutey nieleiieieme/ ae fey tela eats 661

31. On Three New Trematodes from Reptiles. By Witttam Nicoun, M.A., D.Sc., M.B.,

TRASH (GPileiais O10 ey DO. \ AEE) eee Sion dis Gincin bn OS bdbboUona eb edbomooo ne uDeo eS 677

32. The Duke of Bedford’s Zoological Exploration of Eastern Asia —XIV. On Mammals

from Southern Shen-si, Central China. By Oxprietp Tuomas, F.R.S., F.Z.S.

(CRIB IG) 2 O81 DO) nin a Beer be Rae Eo odie goon ace NEUF aha tenets ofchiay ten tat aerate eee oka 687

33. An Investigation into the Validity of Miillerian and other forms of Mimicry, with

Special Reference tothe Islands of Bourbon, Mauritius, and Ceylon. By NrviniE

Manpurs, Lieut.-Colonel, R.A.M.C., B.Z.8., FLHLS...--. 000.0 cc cece ce cece eee cane 696

34. The Distribution of the Avian Genus Megapodius in the Pacific Islands. By J.J. Listur,

MinAcuRb Ns Huy Se ebeZ omen (BexhticUOOs)ie watelsttnts +) se inlelser state exe e/eraistol ain els, oraieiaiels 749

35. Contributions to the Morphology of the Group Neritacea of the Aspidobranch

Gastropods.—Part Ii. The Hznicinips. By Gitperr C. Bourng, M.A., D.Sc., E.RBS.,

IE Za e (a ates NCNONG == hc DB ete ter cinta’ st ney eaves feet cheval eiataial suelo natavatg eters (ch alens “harbors 759

36. On the Palatability of some British Insects, with Notes on the Significance of Mimetic

Resemblances. By R. I. Pocock, F.R.S., F.L.S., F.Z.S., Superintendent of the Society’s

Gardens and Curator of Mammals. With Notes upon the Experiments by Prof. EH. B.

POUTINONS HERES. HLA Sy tevepcrter a aicrase te srcral clei nist stsnclov ale eats’ eVatatstevelovarct Sercvere(el ccc aletel ateleine 809

ADDENDUM.

Additional reference to Dr. P. Cuatuers Mircnent’s memoir, “On Longevity and

Relative Viability in Mammals and Birds” ........-.....++.... SOM LEpeU aI erKCr 868

DESEO PLATES:

1911, Parr III. (pp. 557-868).

Plate Page

XVII. Leptocheirus pilosus Zaddach ......-.2+.....0----.-00-> )

XVIII. i-9. Leptocheirus guttatus Grube. 10-12. L. pinguis |

. Stimpson. 13-16. LZ. hirsutimanus Bate. 17-20. ae

EE bispinosuswNOnManeyas eet siete aera

XIX. Leptocheirus pectinatus Norman .......--+..++ee-eeeee }

XX.

XXI. } Ostracoda from Madeira ..........0e.ceee Obie a HeNAr 595

XXII. |

XXIIL. 1. Blue, 2. Silver, 3. Chequer Pigeons ................. -)

XXIV. 1. Dark Grizzle, 2 - White (crue 3. Grizzled Chequer | |

TEGO) | Jobadodssabpesccsos Wan soooscsducdgg ss G0 L601

XXV. 1. Dark Mealy, 2. Light Mealy, 3. White Mealy Pigeons “+i

XXVI. Feathers showing pattern-markings in Pigeons .........-

XXVII. Lechriorchis CHGDIT AN ING HA A AEO OG o UO aa Oaoe Oo teens \ 677

XXVIII. 6,7. Ochetosoma formosum. 8-10. Dasymetra conferta

XXIX. The Chinese Takin, Budorcas bedfordi ...- 1.0. ..00+ee0es 687

XXX. \

KAKI

XXXII. |

XXXIII. |

XXXIV. |

XXXYV. |

XXXVI. } Morphology of Ge WERPI CITES) sWo seit b taghte im date hts. iets 759

XXXVII. Fe

XXXVITII.

XXXIX. |

XL: '|

XI: |

XLII. )

NOTICE.

The ‘ Proceedings’ for the year are issued in fowr parts, paged consecutively,

so that the complete reference is now P. Z. 8. 1911, p.... The Distribution

is as follows :—

Part I. issued in March

PEA te East June.

srg ep eche E BlD a September.

EESRTUS EAS NO EE ak December.

‘ Proceedings,’ 1911, Part II. (pp. 181-555), were published on

July 6th, 1911.

‘The Abstracts of the Proceedings,’ Nos. 98 and 99,

are contained in this Part.

PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCIETY

OF LONDON.

1911.

PART TV.

CONTAINING Paces 869 to 1213, witH 22 Prates

AND 64 TEXT-FIGUKES.

| FEB 96 1919

DECEMBER 1911.

PRINTED FOR THE SOCIETY,

SOLD AT THEIR HOUSE IN REGENT’S PARK.

LONDON :

| MESSRS. LONGMANS, GREEN, AND CoO., ‘

| PATERNOSTER ROW. |

pT Ki sala alt

s Teri } Sots *

}

Orn [Price Twelve Shillings. ] | ac

LSet Ol 0 IN: IN ans:

1911, Part IV. (pp. 869-1213).

EXHIBITIONS AND NOTICES.

{Pn Secretary. Report on the Additions to the Society’s Menagerie during the month

of April 1911

or gt PO ee OH OO PEE Eo ee ee EE ee Sr. sce ee er eens mrs OF eersteosce ee Our eeosae

The Rey. A. Mites Moss, M.A., F.Z.8S., F.E.S. Notice of a Memoir on the “‘ Sphingidze

of Peru ”

| os %) ex0) c|'e| e/'e\'8)@)/e a) 8) 8. 02 [sss 0) (sie ©) 6)'s, > s\» © e\ia: elie c/s le) e \eif/ip 0 ajc elle ee isvelie!a) jis) © mi elejieleialsl

Mr. H. G. Primer, F.R.S. Report on the Pathological Examination of Rats (Mus

decumanus) caught in the Regent's Park and in the Society's Gardens

Dr. R. W. Saurecpt, C.M.Z.S. Exhibition of a photograph of amale albino Woodchuck

(Arctomys moni)

Mr. R. E. Houpixe. Exhibition of horns of Deer and the skull of a Greyhound

ee eere

Mr. R. I. Pococn, F.R.S., 7.2.8. Exhibition of the skin and skull of the Crested Rat

(Lophiomys ibeanus Thos.), (Lext-fig. 190.)

i ee ee rc ee err rn

Tne Secretary. Report on the Additions to the Society’s Menagerie durine the month -

P y 8 g

of May 1911

CC a ie ee ek a i ir

Mr. D. Sern-Ssirno, F.Z.8. Exhibition of two immature Black-backed Porphyrios (Por-

phyrio melanonotus) with wing-claws. (Text-fig. 200.) .....-.......--

Mr. J. Lewis’ Bonnorts, M.A., F.L.S., ¥.Z.8. Exhibition ofa pair of abnormally coloured

Kgyptian Desert-Mice (Merzones crassus)

Ce i Ce Ci Ce rr ry

Dr. W. T. Catan, F.Z.S. Exbibition of living specimens of the Brine Shrimp (Artemia

salina)

SCC ee ee ee ee i a, i arid

Tne Secrerary. Remarks upon Ostrich eggs seen at Mr. Carl Hagenbeck’s Ostrich

Farm at Stellingen ......... Bass qososusc S66s0 06x SEA US SS suiao doo webs bo de dc

Tis Secretary. Remarks upona pair of young African Rhinoceros seen at Stellingen ..

g §

Mr. R. I. Pococs, F.R.S., F.Z.8. Exhibition of photographs of hybrid Zebra Feals.

(Dext-figs. 201=203))\ oe .e. oe 3 ee Bivens eee ndeleneasbatae Sinton choise Se eS eee ote

PAPERS.

37. The Alcyonaria of the Cape of Good Hope and Natal.—Goreonacna. By J. Srvant

Tuomson, Ph.D., F.R.S.E., F.L.8., Lecturer and Senior Demonstrator in Zoology,

University of Manchester. (Pls. XLIII.—XLV. and Text-fig. 167.) ...........

38. On the Structure of the Skull in Cynodont Reptiles. By R. Broom, M.D., D.Sc.,

OLIWIYAIS, (GHG IONE eral akesacltech IGS ASO)) Sodoscbdéodsaccos coodotgubbadoons

39. Tooth-germs in the Wallaby (Macrapus billardieri). By A. Hornweut-Ssitu,

L.R.C.P., M.R.C.S., and H. W. Marerr Tims, M.A.. M.D., F.LS., F.ZS.

(QE DOU \MULG eval Nexiairash NSW) Ak Gros sondodd0odooddnooosdno cous stosoiercnere

986

987

988

926

Contents continued on page 3 of Wrapper

THE ZOOLOGICAL SOCIETY OF LONDON,

eee

Tus Society was founded in 1826 by Sir Sramrorp RaFrtes,

Mr. J. Sasrnz, Mr. N. A. Vigors, and other eminent Naturalists,

for the advancement of Zoology and Animal Physiology, and for the

introduction of new and curious subjects of the Animal Kingdom,

and was incorporated by Royal Charter in 1829.

Patron.

HIS MAJESTY THE KING.

COUNCIL.

HIS GRACE THE DUKE OF BEDFORD, K.G., President.

THe Eart or ALTAMonrt, F.S.A.

Sir J. Rosp Braprorp, K.C.M.G.,

M.D:, DSe., F.R.S., Vice-

President.

Lr.-Cot. Srr R. Havetocx

Cuarues, M.D., K.C.V.O.

Atrrep H. Cocks, Esa., M.A.

Tue Rr. Hon. tar Earn or

Cromer, P.C.,G.C.B.,G.C.M.G.

F. G. Dawrrey Drewirrt, Esa.,

M.A., M.D.

Caartes DrumMonp,

Treasurer.

Sir Epwarp Douranp, Br., C.B.

Freperick Giiterr, Ese., Vice-

_ President.

Sipney F, Harmer, Ese., M.A.,

Sc.D., F.R.S., Vice-President.

Ksa.,

Srr Warter’ Roeper Lawrence,

Br., G.C.1.E.

Str Epmunp G. Loner, Br.

_E. G. B. Meave-Watpo, Esa.,

Vice-President.

P. Caatmers Mircuent, Ese.,

Mev Discs Ju Dever Rist.

Secretary.

| W. R. Ocitvie-Grant, Ese.

| Aprran D. W. Pottocg, Ese.

AvuBryn Trevor-Batrysg,

M.A.

AntHony H. WrneFrtetp, Esa.

A.Smira Woopwarp, Hse.,LL.D,

F.R.S., Vice-President.

Henry Woopwarp, Esea., LL.D.,

F.RS., Vice-President.

Ksa.,

vay

The Society consists of Fellows, and Honorary, Foreign, and

Corresponding Members, elected according to the By-Laws. It

carries out the objects of its foundation by means of the collection

of living animals, by its Library, and by its Scientific Publications.

The Office of the Society, Regent’s Park, N.W., where all com-

munications should be sent, addressed to “The Secretary,” is open

from Ten till Five, except on Saturdays, when it closes at Two p.m.

The Library, under the superintendence of Mr. F, H. Waterhouse,

is open daily at the above hours, except in September.

The Meetings of the Society for General Business are held in the

Meeting Room at the Society’s Office on the third Wednesday in

every month of the year, except in September and October, at half-

past Four o’clock p.m.

The Meetings for Scientific Business are held in the Meeting

Room at the Society’s Office fortnightly on Tuesdays, except in

July, August, September, and December and January, at half-past

Hight o’clock p.m.

The Anniversary Meeting is held on the 29th. of April, or the

nearest convenient day, at Four p.m.

The Society’s Gardens are open daily from Nine o’clock until

Sunset. Mr. R. I. Pocock, F.R.S., F.LS., is the resident Super-

intendent and Curator of Mammals, Mr. D. Seth-Smith is Curator

of Birds and Inspector of Works, and Mr. E. G Boulenger is:

Curator of Reptiles. The Prosectorium for Anatomical and Patho-

logical work is under the charge of Mr. Frank E. Beddard, M.A.,

F.R.S., Prosector, assisted by Mr. H. G. Plimmer, I.R.S., M.R.C.S.,

Pathologist to the Society.

TERMS FOR THE ADMISSION OF FELLOWS.

Frriows pay an Admission Fee of £5, and an Annual Contri-

bution of £38, due on the Ist. of January, and payable in advance,

or a Composition of £45 in lieu thereof; the whole payment,

including the Admission Ice, being £50.

No person can become a Friiow until the Admission Fee and

first Annual Subscription have been paid, or the annual payments

have been compounded for.

Fettows elected in November and December are not liable for

the Subscription for the current vear.

PRIVILEGES OF FELLOWS.

Ferttows have Personal Admission to the Gardens upon signing

their names in the book at the entrance gate, and may introduce

Two Companions daily.

The Wire or Hussanp of a Fattow can exercise these privileges

in the absence of the Fellow.

livery Frtiow is entitled to receive annually 60 undated Green

Cards, and, when no specific instructions are received, the supply

will be sent in this form. If preferred, however, 20 Green Cards

may be exchanged for a book containing 2 Orders for each

Saturday * throughout the year. A similar book of Sunday Orders

may also be obtained in lieu of 20 Green Cards. A Green Card

may also be exchanged for 2 Butf Cards for the use of Children

under 12 years of age.

It is particularly requested that Fellows will sign every Ticket

before it goes out of their possession, Unsigned ‘Tickets are not

available.

Green and Buff Tickets may be used on any day and in any year,

but in no case can two Children be admitted with one Adult's

Ticket, or an Adult be admitted with two Children’s Tickets.

The annual supply of Tickets will be sent to each Frtrow on the

Ist. of January in every year, upon filling up and returning the form

of Standing Order supplied to Fellows.

Frttows are not allowed to pass in friends on their written

order or on presentation of their visiting cards.

Futtows are exempt from payment of the fee for Painting,

Sketching, and Photographing in the Society’s Gardens.

Frttows have the privilege of receiving the Society’s ordinary

Publications issued during the year upon payment of the additional

Subscription of One Guinea. This Subscription is due upon the

Ist. of January, and must be paid before the day of the Anniversary

Meeting, after which the privilege lapses. Frtiows are likewise

entitled to purchase these Publications at 25 per ceut. less than

the price charged to the public. A further reduction of 25 per

cent. is also made upon all purchases of Publications issued prior

to 1881, if above the value of Five Pounds.

Frrttows also have the privilege of subscribing to the Annual

Volume of ‘The Zoological Record,’ which gives a list of the Works

and Publications relating to Zoology in each year, for the sum of

* The Saturday Orders are not available if the Fellow introduces friends

personally on that day.

One Pound Ten Shillings. Separate divisions of volumes 39 to

42 can also be supplied. Full particulars of these publications can

be had on application to the Secretary.

Frntows may obtain a TransrerasLe Ivory Ticker admitting

two persons, available throughout the whole period of Fellowship,

on payment of Ten Pounds in one sum, A second similar ticket

may be obtained on payment of a further sum of Twenty Pounds.

Any Frttow who intends to be absent from the United Kingdom

Curing the space of at least one year, may, upon giving to the

Secretary notice in writing, have his or her name placed upon the

“dormant list,” and will then be called upon to pay an annual

subscription of £1 only during such absence, but after three years

must make a further application to be retained on that list.

Any Frtiow, having paid all fees due to the Society, is at liberty

to withdraw his or her name upon giving notice in writing to the

Secretary.

Ladies or Gentlemen wishing to become Fellows of the Society

are requested to communicate with the undersigned.

P. CHALMERS MITCHELL,

Secretary.

Regent’s Park, London, N.W.,

December, 1911.

MEETINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON

FOR

SGILIB IN POI 1G) 1s) WES ICIS! IBS) Si

1912.

Truspay, Fepruary.. 6 & 20 | Tunspay, JuNF...... 4

55 INRCEH 2a) pOROg ee 3 OcroBER .. 29

“4 NPR See yOu Ze Ee November... 12 & 26

IMA ee 2k 7 & 21

The Chair will be taken at half-past [ight o'clock in the Evening

precisely.

ZOOLOGICAL SOCIETY OF LONDON.

LIST OF PUBLICATIONS.

Tue scientific publications of the Zoological Society of London

are of two kinds—“ Proceedings,” published in an octavo

form, and ‘‘ Transactions,” in quarto.

According to the present arrangements, the “ Proceedings”’

contain not only notices of all business transacted at the scien-

tific meetings, but also all the papers read at such meetings

and recommended to be published in the ‘‘ Proceedings ”’ by

the Committee of Publication. A large number of coloured

plates and engravings are issued in the volumes of the

“ Proceedings,” to illustrate the new or otherwise remark-

able species of animals described therein. Amongst such

illustrations, figures of the new or rare species acquired in a

living state for the Society’s Gardens are often given.

The “Proceedings” for each year are issued in four

parts, paged consecutively, in the months of March, June,

September, and December. From January 1901 they have

been issued as two half-yearly volumes, indexed separately.

An “ Abstract of the Proceedings” is published by the

Society on the Tuesday following the date of Meeting to

which it refers. It is issued along with the “ Proceedings,”

free of extra charge, to all Fellows who subscribe to the

Publications, but it may be obtained on the day of pubii-

cation at the price of Sixpence, or, if desired, sent post free

for the sum of Six Shillings per annum, payable in advance.

The “Transactions ”’ contain such of the communications

made to the scientific meetings of the Society as, on account of

the nature of the plates required to illustrate them, are better

adapted for publication in the quarto form. They are issued

at irregular intervals.

Fellows and Corresponding Members, upon payment of

a Subscription of One Guinea before the day of the Anni-

versary Meeting in each year, are entitled to receive the

Society’s Publications for the year. They are likewise

entitled to purchase the Publications of the Society at 26 per

cent. less than the price charged for them to the Public. A

further reduction of 23 per cent. is made upon purchases of

Publications issued prior to 1881, if they exceed the value of

five pounds.

Fellows also have the privilege of subscribing to the

Annual Volume of the Zoological Record for a sum of 30s.

(which includes cost of delivery), payable on the Ist. of July

in each year; but this privilege is forfeited unless the

subscription be paid before the 1st. of December following.

The following is a complete list of the publications of the

Society already issued.

TRANSACTIONS* OF THE ZOOLOGICAL SOCIETY OF LONDON.

4to. 19 vols. and Index. Eicetto Price tote

ellows. ublie.

Vol. I., containing 59 Plates.... (1833-35) .... £3 138 6 .... £418. OF

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PROCEEDINGS OF THE COMMITTEE OF SCIENCE AND

CORRESPONDENCE OF THE ZOOLOGICAL SOCIETY OF

LONDON. 8vo. 2 vols. (Letterpress only). 22 ' Pace

Pave Me ISOS ILO G@s “Gesaooccagocas AS Gs) ee BOS

4 Ue TEED. Ean ene Ms Gi os

PROCEEDINGS OF THE ZOOLOGICAL SOCIETY OF LONDON.

8vo. 15 vols. (Letterpress only) and Index. (First Series.)

: Price to Price to the Price to Price to the

Fellows. Public. Fellows. Public.

Part J. 1833. 1 vol. 8vo. 4s. 6d. .. Gs.f | Part IX. 1841.1 vol. 8vo. 4s. 6d. .. 68.F

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ViIL. 1840. . ° 4s.6d... 6st | Index 1880-1847. , 4s.6d. .. 6s.

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8vo. 18 vols. and Index. (Second Series.)

Letterpress only. With Plates coloured.

Price to Price to the Price to Price to the

Fellows. Public, Fellows. Public.

Part SWAT, WSS. Ul swwoll, Bras 4's, Ge 45 CS. ooscaocen £10058) 6 SIG

25 XVI. 1849. 5 AS ROG ancy OSI ee ciate tence 1 (0. 68 ooo) DGG

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Index 1848-1860. . ds. 6d. 6s,

t+ Out of print.

* In consequence of a re-arrangement of the stock of the ‘Transactions,’ the Society is

now able to offer for sale, at the reduced price of £30, sets of Vols. v.-xvi. inclusive, and

separate papers, of which a list can be supplied, at about one-fourth their published price.

PROCEEDINGS OF THE SCIENTIFIC MEETINGS OF THE

ZOOLOGICAL SOCIETY OF LONDON. 8vo. 40 vols. and 4 Indices.

Letterpress only. With Plates uncoloured. With Plates coloured.

Price to Price to the Price to Price to the Price to Price to the

Fellows. Public. Fellows. Public. Fellows. Public.

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* No perfect copies in stock. ft Out of print.

PROCEEDINGS or roe GENERAL MEETINGS ror SCIENTIFIC

BUSINESS or tar ZOOLOGICAL SOCIETY OF LONDON.

8vo. 22 vols.

Price to Price to the

Fellows. Public.

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LISTS OF THE ANIMALS IN THE SOCIETY’S GARDENS.

List of the Vertebrated Animals now or lately Living in the Gardens

of the Zoological Society of London. (Mighth Edition.) 8vo.

1883. Cloth, 4s. 6c.

List of the Vertebrated Animals now or lately Living in the Gardens

of the Zoological Society of London. (Ninth Kdition.) 8vo.

1896. Cloth, 6s.

Catalogue of the Library of the Zoological Society of London

(Fifth Edition.) Svo. 1902. Cloth, 6s.

THE OFFICIAL ILLUSTRATED GARDEN GUIDE— 9th Edition

—with (1) a Railway and ‘Street Map, showing a direct

route to the “* Zoo” from all parts of London and the Suburbs ;

(2) a Plan of the Grounds, showing at a glance the location of

the animals; (3) a short description of some of the principal

animals in the Collection (now containing over 3000 spe-

cimens), together with 50 Photographic Illustrations and

Index. Price 6d. in Stiff Paper Cover, postage 13d., or in

Green Cloth Cover price 1s. 2d. post free.

P. CHALMERS. MITCHELL,

Secretary.

Regent's Park, London, N.W.,

December, 1911.

These publications may be obtained at the Socrrry’s Orrice,

at Messrs. Lonemans (Paternoster How, £.C.), or through any

bookseller.

AVOLOGICAL SOCIETY OF LONDON.

THE ZOOLOGICAL RECORD.

HE object of the Zoonoeican Kecorp is to give, by means of an

annual Volume, complete lists of the Works and Publications

relating to Zoology in all its branches that have appeared during

the year preceding the issue of the Volume, together with full

information as to the points they deal with, arranged in such a

manner as to serve as an Index to the literature of Zoology in all

parts of the globe, and thus to form a repertory that will retain its

value for the Student in future years.

The ‘ Zoological Record’ having been amalgamated with the

International Catalogue of Scientific Literature, Zoology, Volumes

from 43 onwards cam now be obtained only from Messrs. Harrison

& Sons, except when purchasing complete sets from the Zoological

Society.

Under the scheme of amalgamation, Fellows of the Society, and

Institutions already on the subscription-list, have the privilege of

subscribing at the old rate of 30s. per annum, which covers the

cost of carriage of the volume. The subscription becomes due on

July Ist. in each year, and lapses if not paid by the 1st. of December

following.

The Society is able to supply complete sets of the Record on the

following terms :—

Vols. 1 to 37, price £14 10s. net.

Vols. 38 to 42 at 10s. each net.

Vol. 43 and onwards at 40s. each.

The prices for separate volumes ere as follows :-—

Vols. 1 to 42 (except Vols. 4 and 6) 10s. each net.

The price of the ‘Zoological Record,’ Vol. 43 and subsequent volumes,

published now by Messrs. Harrison and Co., is 40s. each.

Inpex Zootocicus. An alphabetical list of names of genera

and subgenera proposed for use in Zoology, as recorded in the

‘Zoological Record,’ 1880-1900; together with other names not

included in the ‘ Nomenclator Zoologicus’ of S. H. Scudder. Com-

piled (for the Zoological Society of London) by CHarrus Ownn

Wartrrnouse and edited by Davin Sarr, Editor of the ‘ Zoological

Record.’ London, 1902. Price to Fellows, 18s.; price to the

public, 20s., or if sold with a set, 10s.

Divisions of the ‘ Zoological Record’ of Vols. 39 to 42 ean be

supplied by the Society, but after Vol. 42 they can be had only of

Messrs. Harrison & Sons, 46 St. Martin’s Lane, W.C.

[Rs OEHO,

SEPARATE DIVISIONS OF THE ZOOLOGICAL RECORD.

Divisions of the ‘Zoological Record, Vols. 39-42, containing

the literature of the years 1902-1905, may be obtained separately

as follows :—

8. ah

list of abbreviations of journals, cte. .. .. 2 © Met.

Special Records, viz. :—

lls CGH SMI cs so) oe ge 6 BG oy

eee Miaimaciliane weet crests eee aches oo es On sae

ELMAN CSI re Shee eee! Peers aapraces Matec GO Oy ee

JAWS Ikejomuien, aval iateeelue., o5 of 24 © 5.

V. Pisces 20) es

VI. Tunicata Degen Oe

VII. Mollusea AU aL

VIII. Brachiopoda .. le POM ee

IX. Bryozoa Wh Sot a

X. Crustacea 1M ee

XI. Arachnida 2 Orne

XII. Myriopoda DP O28 ss

XIII. Insecta 120 = 3

XLY. Echinoderma 3 VCs

XV. Vermes .. 3) 0) 5g

XVI. Coelenterata .. A AG ae

XVII. Spongiee 2 Os

XVIII. Protozoa nO aes

Index of new names of genera and subgenera. 2 O ,,

Divisions from Vol. 43 onwards are now supplied t

Harrison & Sons, 46 St. Martin’s Lane, London, W.C.

y Messrs.

P. CHALMERS MITCHELL,

Secretary.

Recents Park, Lonpon, N.W.

December, \911.

ON ADDITIONS TO THE MENAGERIE. 869

EXHIBITIONS AND NOTICES.

May 23, 1911.

Dr. A. Smrra Woopwarp, F.R.S., Vice-President,

in the Chair.

THE Secrerary read the following report on the additions made

to the Society’s Menagerie during the month of April, 1911 :—

The number of registered additions to the Society’s Menagerie

during the month of April last was 274. Of these 75 were

acquired by presentation, 147 by purchase, 34 were received on

deposit, 5 in exchange, and 13 were born in the Gardens.

The number of departures during the same period, by deaths

and removals, was 169.

Amongst the additions special attention may be directed to :—

1 Black Serval (Felis serval), from Nairobi, presented by the

Marchioness of Donegal to H.M. Tue Krne’s African Collection,

on April 25th.

2 Bactrian Camels (Camelus bactrianus), from Central Asia, and

1 Yak (Poéphagus grunniens), from Tibet, presented on April 4th

and 29th respectively, by H.G. The Duke of Bedford, K.G.,

President of the Society.

1 Klipspringer ( Orectragus saltator), from Somaliland, presented

by R. E. Drake-Brockman, Esq., M.R.C.S., F.Z.S., to H.M. Tax

Kine’s African Collection, on April 28th.

1 Savannah Deer (Dorcelaphus savannarum), from Iquique,

Chile, presented on April 24th by Miss Peggy Lomax,

1 Black-tailed Oribi (Ourebia nigricaudata), from the Ivory

Coast, presented by F. W. F. Jackson, Esq., to H.M. Tue Kina’s

African Collection, on April 4th.

2 Australian Bee-eaters (Jerops ornatus), from New South

Wales, new to the Collection, purchased on April 30th.

2 Woodhouse’s Jays (Aphelocoma woodhousei), from North

America, and 3 Pigmy Ground-Doves (Chamepelia minuta), from

South America, new to the Collection, received in exchange on

April Ist.

2 Sharpe’s Wood-Owls (Syrnium nuchale), 1 Fraser’s Eagle-Owl

(Bubo poénsis), | Beautiful Wood-Hawk (Dryotriorchis spectabilis),

and 1 One-streaked Hawk (Melierax monogrammicus), from

Dunkwa, Gold Coast, presented by Dr. H.G. F. Spurrell to

H.M. Tae Kine’s African Collection, on April 2nd.

4 Gigantic Salamanders (Megalobatrachus maximus), from Japan,

purchased on April 28th.

The Rev. A. Mines Moss, M.A., F.Z.S., F.E.S., gave a short

account of his memoir on the Sphingide of Peru, based on studies

of Lepidoptera, with special reference to the larvee, which he had

made during a three years’ residence at Lima.

This paper will be published in the ‘Transactions’ in due

course.

Proc. Zoou. Soc.—1911, No. LX. 60

870 DR. J. STUART THOMSON ON

PAPERS.

37. The Aleyonaria of the Cape of Good Hope and Natal.—

GorconaceaA. By J. Stuart THomson, Ph.D., F.R.S.E.,

F.L.S., Lecturer and Senior Demonstrator in Zoology,

University of Manchester *.

[Received May 4, 1911: Read May 23, 1911.]

(Plates XLIII.—XLV.? and Text-figure 167.)

The Aleyonaria here reported on were collected during the years

1898 to 1907, off the coasts of Cape Colony and Natal. In

a previous paper I dealt with the Alcyonacea, and in the

present report I confine myself to the Gorgonacea. This con-

tribution is not, however, to be regarded as an exhaustive one, as,

unfortunately, the specimens have come into my hands at different

times. Professor Hickson has contributed two papers on South

African Aleyonaria to the publication, ‘ Marine Investigations in

South Africa,’ and I have to thank him for allowing me to

examine his type-specimens and microscopic slides of the spicules

of various forms.

I have also to thank Professor J. D. F. Gilchrist, formerly

Government Biologist at the Cape of Good Hope, who kindly

entrusted these Aleyonarians to me for description.

Tn this paper, the following 19 species are described or recorded,

of which 6 are new :—

Family BRIAREIDA.

Suberia capensis, sp. n., p. 871.

Spongioderma verrucosum Mobius, p. 874.

Family MEurropip#.

Melitodes esperti W. & S., p. 874.

Melitodes nodosa W. & S., p. 876.

Melitodes dichotoma Pall., p. 877.

Family Istpm.

Ceratoisis ramosa Hickson, p. 877,

Family Muriceis.

Muriceides fusca, sp. n., p. 878.

Acanthogorgia armata Verrill, p. 880.

* Communicated by Prof. ARTHUR DENpDy, D.Sc., F.R.S., F.Z.S.

+ For explanation of the Plates see pp. 892-893.

+ “The Alcyonaria of the Cape of Good Hope and Natal.—Alcyonacea” (with

our plates). ‘Trans. Roy. Society Edinburgh, vol. xlvii. part ii. (No, 19), 1910,

P.Z.S. 1911. Pl. XLII.

= VS

fi “ * e

4 ,

Ree

I ¢ 3

Soy! :

Sy &

~ s .

e 3% vip ey

ALCYONARIA OF

THE CAPE OF GOOD HOPE.

P.Z.S8.1911.P1. XLIV.

ALCYONARIA OF THE CAPE OF GOOD HOPE.

P.Z.8.1911.P1. XLV.

CAPE OF GOOD HOPE.

ALCYONARIA OF THE

CORALS FROM SOUTH AFRICA. 871

Family PLEXAURIDS.

EKunicella papillosa Esper, p. 880.

Psammogorgia pulchra, sp. n., p. 881.

Huplexaura media, sp. n., p. 883.

Family MALAcocoreiiD&.

Malacogorgia capensis Hickson, p. 884.

Family Primnoms.

Stachyodes gilchristi, sp. n., p. 885.

Thouarella hicksoni, sp. n., p. 886.

Family GorGoNntD&.

Gorgonia capensis Hickson, p. 887,

Gorgonia flammea HK. & S., p. 888.

Family GoRGONELLID#.

Scirpearia furcata emend. Simpson, p. 888.

Scirpearia flagellum emend. Simpson, p. 888.

Hicksonella spiralis Simpson = Juncella spiralis

Hickson, p. 889.

Family BRIAREIDS&.

SUBERIA CAPENSIS, sp. n, (Pl. XIII. fig, 4; Pl. XLV.

figs. 4a—c.)

The colony is of a dark red colour, resembling that of Gorgonia

flammea K. & 8.,and consists of (1) an encrusting base, (2) a more

or less cylindrical trunk, and (3) an expanded apical part. The

basal part is 10 mm. long, 13 mm. broad, and 6 mm. high,

The trunk is about 27 mm. in height, 6 mm. in diameter basally

and medially, and about 7 mm. apically. The lowest polyp is

situated on the trunk about 13 mm. from the base. About

fourteen polyps are situated near the apex of the trunk, and these

ave almost entirely confined to three sides. The largest polyps of

this part are 2 mm. in height and 3 mm. in diameter. The

distance of the polyps from one another varies considerably,

nainely from about 1 to 8 mm. The apical part of the colony is

about 14 mm, in height and 15-17 mm. in diameter (including

the polyps). The polyps are in this part more prominent than

in the trunk region. They vary from 2-3 mm. in height and

breadth. The polyps are irregularly scattered over the surface of

the apical part, and the interval between them varies from about

1to5mm. There are thirty to forty polyps in the apical part.

The entire surface of the colony is rough to the touch and has

a tough consistency. In the centre of the colony there is a horny

and limy axis; in other words, the axis consists of horny tissue

and in association with this a large number of spicules. The

spicules of the axis usually differ in form from those of the cortex.

60*

872 DR. J, SEUART THOMSON ON

There is a ring of canals immediately surrounding the axis and

another ring occurs in the cortex. The polyps are retractile

within the verruce. In my specimen all the polyps are retracted.

Spicules are present in the anthocodie.

The spicules of the polyps are (1) small tri-radiate spicules with

sharp spines on the blunt ends of the rays, this being the

commonest type of spicules in the polyps; (2) rods with spines on

the expanded ends, some of which approach a dumbbell shape ;

(3) approximately straight rods; and (4) spindles. The spicules

are arranged in eight longitudinal bands on the anthocodiz and

do not appear to be in any definite arrangement in relation to one

another. They are all red in colour.

The spicules of the axial part of the trunk are (1) colourless rods

with few spines, (2) spindles with blunt processes, the latter with

spines or thorns, and (3) irregularly-shaped spicules.

The spicules of the cortex of the trunk are most frequently

spindles (with a similar shape to those of the axial part), tri-

radiate spicules, short rods, and small irregularly-shaped spicules.

The spindles are more characteristic of the cortex, and the rods of

the axial part. The wall of the stomodeum contains numerous

small red spicules, mostly tri-radiate forms.

The dimensions of the spicules are as follows :—From the polyps :

(1) large spindles, from 0-119 x 0-085 to 0-188 x O19) mamas

(2) tri-radiate forms, from 0:051 x 0:0357 to 0-068 x 0-068 mm. ;

(3) 4rayed forms, from 0:051 x 00425 to 0-068 x 0-051 mm. ;

(4) small dumbbell-like forms, but with double ends, about

0-051 x 0:020 mm. .

From the cortex of the trunk: (1) large spindles, from

0:0935 x 0:0595 to 0:221 x 0-119 mm ; (2) small tri-radiate forms,

from 0:0357 x 0-051 to 0:0595 x 0:022 mm.; (3) dumbbell-lke

forms approximately the same size as those of the polyp.

From the axial part of the trunk: (1) large spindles (only a few

occur), from 02040-1105 to 0:289 x 0:0935 mm.; (2) rods,

from 0°085 x 0°0255 to 0°272x0:0765 mm.; (3) irregularly-

shaped spicules, from 0-238 x 0-119 to 0:340 x 0-119 mm.

Locality, ete.—‘ Pieter Faure,” No. 13139. Cape Morgan,

N.N.E. 92 miles. Depth, 47 fathoms. Procured by dredge.

Nature of bottom, broken shells. Only one specimen of this

species was collected.

Studer gives the following diagnosis of the genus Suberta :—

“Stamm einfach oder verzweigt, aufrecht, mit einer Achse, die

aus unverschmolzenen, von Hornsubstanz umgebenen stabformigen

Spicula gebildet wird und der Ernihrungscanale entbehrt. Rinde

dick, enthalt spindelférmige stachlige Spicula, Die Polypenwarzen

sind gross, senkrecht vom Stamme abstehend, die Offnung an der

Spitze der Warzen achtstrahlig. Die Polypen von der Basis bis

in die Tentakel mit feinen spindelformigen Spicula erfwllt. Um

die Achse ein Kranz von Langscaniilen.”

Studer adds that the genus Suwberia stands near Spongioderma,

but differs from it in the form of the calyx and spicules.

CORALS FROM SOUTH AFRICA.

873

Three species of the genus Suberia have been described, and the

following table shows the differences between them so far as these

can at present be stated.

Suberia capensis, sp. n.

Colour, dark red.

Simple, unbranched.

Height 53 mm.

Trunk cylindrical,

expanded at the apex.

Suberia genthi Str.

Branched.

80 mm.

Cylindrical.

Polyps at the apex but Polyps not terminal.

not terminal, 2-3 mm.

in height.

The rind is firm and

rough.

The spicules are :—

(1) rods, (2) spindles,

(3) tri-radiate forms,

(4) 4-rayed forms,

(5) irregular spicules.

No canals in axis.

(1) straight warty

spindles,

(2) curved warty

spindles,

(3) 4-rayed forms,

(4) double crosses,

(5) irregularly branch-

ed spicules.

Canals in the axis.

Suberia koellikeri Str. Suberia clavaria Str.

Rose-red, disappearing Flesh-coloured during

in alcohol. life, disappearing in

alcohol.

Branched. Simple, unbranched.

About 110 mm. 160 mm.

Cylindrical, branches Cylindrical, expanded

expanded at the at the apex.

apices.

Polyps thickly crowded

together at the apex,

1-2 mm. in height.

The rind is thin and The rind is soft and

1-15 mm. in height.

rough.

(1) rod-like spicules,

(2) cross-shaped or

triplets.

smooth.

(1) slightly spined

spindle-shaped rods,

(2) crosses,

(3) spinous spindles.

Canals immediately out- Canals in the circum- Axis surrounded by a

side the axis.

Axis of horny tissue with

numerous associated

spicules.

Dimensions of Spicules :—

Large spindles, from

0:0935 X 0°0595_ to

0'0289 X 0'0935 mm. ;

tri-radiate forms, from

00357 X 0:01 to 0°068

<0°068 mm.

rods, from 0°085 X 0°0256

to 0°272X0:0765 mm. ;

irregularly - shaped

spicules from @'238X

0119 to 0:340X0119

mm.

dumbbell-shaped forms

about0°051 X 0°020 mm.

Habitat :—South Africa,

Cape Morgan, N.N.E.,

92 miles. Depth, 47

fms. By dredge.

Bottom, broken shells.

ference.

Straight, warty spi-

cules, from 0°09 X

0°08 to 0'34X0°10

mm.

curved, warty spindles

from 0°24X012 to

0°30X0'10 mm. ;

long, spiny, curved

spindles, from 0°'4X

0°04 to 0°24 XK O04

mm.;

irregularly branched

spicules, from 0:24

to 0°06 mm. in widest

diameter ;

4-rayed forms, 0°32 to

0:06 and 0°20 to 0°12

mm, ;

double crosses, from

0'1 to 0°025 and 0°08

to 0°01 mm.

Shallow water, off

Port Jackson

(Australia).

wreath of canals.

Axis consists of smooth Axis as in the last

rod-like spicules species “* well separ-

(with few spines) ated” and consists of

lying close to one long, slightly spined,

another. spindle-shaped rods ;

crosses also occur.

Spinous spindles from

the rind, 0°71 to 0°18

mm. in length ;

Warty, thorny spi-

cules, 0°24 and 0°021

mm. in length ;

rods (‘ Drillinge”)

from axis, 0°35 mm.

long, slightly spined,

spindle-shaped rods,

0°237 and 0°265 mm.

in length.

North of New Zealand. Atlantic, off Monte

Video

874 DR. J. STUART THOMSON ON

SPONGIODERMA VERRUCOSUM Mobius.

This species has already been described from South African

waters by Hickson in his paper on the Alcyonaria and Hydro-

corallinz: of the Cape of Good Hope, Part I.

Localities, etc.— Apparently a shallow-water form.

“Pieter Faure.” 12310. Port Shepstone, north, 8 miles.

By dredge. Depth, 36 fathoms. Nature of bottom, broken shells

and stones. Date, October 30, 1902.

‘“‘ Pieter Faure,” 11127. Umkomass River mouth, N.W. by W.

2 W.54 miles. By large dredge. Depth, 40 fathoms. Nature

of bottom, broken shells and stones. Date, December 30, 1900.

Family MeLITODIDA.

MELITODES ESPERI W.&S8. (Text-fig. 167.)

The specimen is about 104 mm. in length and 117 mm. in

breadth, but these dimensions are only approximate as several of

the terminal branches were broken away. The colony had three

beautiful young specimens of Gorgonocephalus intertwined among

its branches.

Text-fig. 167.

Melitodes esperi and Gorgonocephalus.

The colony is red in colour, and is attached to the substratuin

by a broad, trunk-like base. The branches are mostly in one

plane, although a few are slightly turned inwards in a direction

CORALS FROM SOUTH AFRICA. 875

vertical to the main axis. There is a slight anastomosis of the

branches; the meshes thus produced are very irregular in shape.

Sometimes branches, which may be slightly divided, are seen

growing into the meshes; these offshoots may arise from the

internodes, but this is rare.

The nodes and internodes are very distinctly marked, the former

being dark red and the latter hght red in colour. The internodes

vary in length from 2 mm. near the base to 9 mm. towards the

apex of the colony. The nodes are more or less ring-shaped, but

they have a slight projection in front. The nodes are smaller

towards the apices of the branches than near the bases and are

not so easily distinguished from the internodes. Near the base

the nodes may be 4 mm. in length and 3 mm. in diameter ;

towards the apex they are about 1 mm. in length and diameter.

The axis consists of long needle-like spicules densely compressed

together. The lower part of the axis is not perforated by canals.

On a branch of average size, the diameter of the axis is about

1 mm. The internodes of the axis are pale pink, the nodes

darker in colour.

The base of the colony is a thick trunk about 5 mm. in

diameter, but it soon divides and gives rise to the branches. The

branches are disposed in a more or less fan-shaped manner.

Various organisms, such as Polyzoa, Brachiopods, and Hydroids,

are thickly grouped at the base of the colony.

The coenenchyma covering the axis is about 4 of a millimetre

in thickness: its surface is densely covered by an extremely large

number of spicules of varied size and shape. A rough surface is

produced by the slightly protruding spicules.

The contracted polyps appear as yellow papille on the surface

of the ceenenchyma, and are about °6 mm. in height and 1 mm. in

breadth. On the large, basal branches of the colony, the polyps

are only distributed on three sides, but on the apical branches

they occur on all sides. Polyps are situated on the nodes, and

not simply confined to the internodes. The spicules of the polyps

are yellow, and are arranged in eight areas.

The spicules of the coenenchyma are of the following forms :—

Clubs, with broad leaf-like processes; curved spindles with

narrow processes, some of these spicules are so much curved as

almost to form a semicircle; straight spindles of two forms:

(a) those with small processes, and (6) those with broad, expanded

processes; other spicules are irregular in shape, owing to the

presence of broad processes, which produce a more or less branched

appearance; one of these spicules has, for example, three apical

prongs or processes.

The spicules of the ccenenchyma are all red in colour.

The spicules of the polyps are as follows :—

Straight spindles fairly similar in form to those of the ccenen-

chyma; curved spindles, also fairly similar in shape to those of

the coenenchyma, some of these spindles are bifurcated at the end

876 DR. J, STUART THOMSON ON

in a similar manner to that shown by Wright and Studer for this

species ; straight rod-like spicules with rounded processes ; clubs

and spicules of an irregular shape.

The spicules of the polyps are yellow in colour.

The spicules of theccenenchyma have the following dimensions :—

(a) Clubs, from 0:102 x 0:054 to 0:187 x 0-076 mm. ; (6) curved

spindles, from 0-085 x 0:0204 to 0:238 x 0-047 mm. ; (c) straight

spindles, from 0°0765 x 0:0204 to 0-272 x 0:0646 mm. ; (d) irregu~

larly-shaped spicules, from 0-085 x 0'068 to 0:119 x 0-068 mm.

The dimensions of the polyp spicules are as follows :—(a) straight

spindles, from 0°081 x 0-037 to 0°153x0:025 mm.; (6) curved

spindles, from 0:122 x 0-084 to 0°187 x 0-034 mm. ; (c) clubs, from

0-085 x 0:034 to 0-136 x 0-051 mm.; (d) irregular spicules, from

0:076 x 0:110 to 0°170 x 0:034 mm.

Locality, etc—P.F. 852. About 25 miles east of Hast London.

Tat, 32° 48’ 30" S.; Long. 28° 11! 15” KE. By shrimp trawl.

Depth, 22 fathoms. Nature of bottom,mud. Date, January 11,

1899.

Me.iroprs noposa W. & 8.

The specimen was not complete, the base being absent and many

of the terminal branches had become broken away. ‘The entire

colony probably measured 230 mm. in length and 120 mm. in

breadth. The branches all arise in one plane. The nodes are

very prominent, measuring as much as 6 mm. in diameter and

5 mm. in height near the base of the colony, and 2 mm. in

diameter and 1-5 mm. in height on the small apical branches.

The internodes vary considerably in length, namely, from 2 mm.

near the base of the colony to 12-13 mm. on the terminal

branches. The internodes are more or less cylindrical, and their

diameter varies from about 3 mm. near the base to | mm. at the

apex. Several of the branches originate from the internodes, as

in the specimens of this species collected by the ‘Challenger.’

There is a slight anastomosis near the base of the colony, produced

by branches springing from the nodes.

On the upper branches the polyps are distributed on all sides ;

on the lower part, however, they are mostly confined to three

sides. The polyps are retractile within well-defined verruce,

which measure about 1 mm. in diameter and 0°75 mm. in height.

The ground colour of the colony is yellow, the nodes are brown,

the verruce yellow. The axis, when deprived of the overlying

cenenchyma, is brown at the nodes and white at the internodes.

The conenchyma is rough owing to the presence of numerous

spicules and has a thickness of 1-2 mm. ‘The surface of the

cenenchyma has longitudinal grooves. The spicules of the

cceenenchyma are (1) irregularly-curved spindles, (2) irregularly-

branched spicules, (3) curved, spinous spindles, (4) 4-rayed forms.

‘heir dimensions are as follows :—Irregularly-curved spindles,

from 0:204x0:059 to 0:238x0:051 mm.; irregularly-branched

spicules, from 0'0765 x 0:068 to 0°136x0-085 mm.; curved,

CORALS FROM SOUTH AFRICA. 877

spinous spindles, from 0°153x0-025 to 0170x0°025 mm. ;

4-rayed forms, 0-051 x 0-051 mm.

The spicules of the polyps are (1) curved spindles, (2) straight

spindles, and (3) irregular spicules. The dimensions of these

spicules are as follows:—Curved spindles, from 0:136 x 0017 to

0-250 x 0°034 mm.; straight spindles, from. 0°023 x 0°051 to

0-255 0-056 mm.; irregular spicules, from 0-068 x 0-051 to

0-170 x 0-085 mm. The spicules of the polyps may be contrasted

with those of the ccenenchyma by the predominance of straight

and curved spindles.

The ‘Challenger’ specimens were collected (1) off the New

Hebrides at a depth of 60-120 fathoms, (2) on Hyalonema-ground,

off Japan, at a depth of 345 fathoms. These specimens differ

from mine in the colour of the cenenchyma and axis; the

cenenchyma being reddish brown and the axis yellowish red.

In both, the polyps were yellow and the nodes darker than the

internodes. ‘The shape of the spicules of my specimen rather

differs from that of the ‘Challenger’ forms, but, on the other

hand, the spicules seem to vary so much within this species that

I do not feel inclined to lay great stress on this point.

Locality, etc—P.F. 18381. Off Flesh Point, N. 6 miles to

Flesh Point, N. 2 W. 64 miles. By large trawl. Date,

January 15, 1904.

MELITODES DICHOTOMA Pall.

Fragments of a red variety of this species. This species has

already been recorded from the Cape. It is interesting to note

its occurrence in such shallow water, namely from 6—14 fathoms.

At Gordon’s Bay, in all probability it may be thrown up on the

shore during storms, as is so much the case with Gorgonia lammea

HE. & 8. iz

Locality, etc—P.F. 15725. Off Gordon’s Bay. By dredge.

Depth, 6-14 fathoms. Nature of bottom,rock. Date, October 20,

1902.

Family Isip &.

CrRAToISIS RAMOSA Hickson. (Plate XLIII. fig. 1.)

This beautiful form has already been described by Hickson

from South African waters. The species is very fragile, and the

delicate terminal branchlets are very liable to become broken

away. In Hickson’s specimen the base was wanting, but my

specimens are more complete in this respect. The calcareous base

measures about 8 mm. in length, 4 mm. in breadth, and 2 mm. in

height.

The height of the complete specimen would be about 100 mm.

The first part of the main stem immediately above the base is

brown or bronze in colour.

My specimens agree with Hickson’s description of forms col-

lected off Vasco da Gama Peak at a depth of 230 fathoms.

The distribution of colour on the colony appears to vary. The

878 DR. J. STUART THOMSON ON

ivory-white internodes, as described by Hickson, appear to become

reddish in the upper part of the colony, but this is apparently not

always the case.

Locality, ete.—P.F. 13197. Cove Rock, N.W. ? W. 134 miles.

By dredge. Depth, 80-130 fathoms. Nature of bottom, coral

and rvecks. Date, July 30, 1901.

Family MuRICEID&.

Muricerpes FuscA, sp.n. (Plate XLIV. figs. 4 a—c.)

The colonies are about 102 mm. in height and 95 mm. in

breadth. The branches are all in one plane. The main trunk

has a strong horny axis and an expanded base measuring 14 mm.

in length and 12 mm.in breadth. The main trunk has a diameter

of 5mm. and gives off its first branch 13 mm. from the base.

The branches of the first order are given off irregularly from the

trunk, and these, in their turn, give off branches of the second

order, which may also bear small terminal branches. ‘The

branches of any order do not, as a rule, originate opposite one

another. The axis of all parts is thick, the cenenchyma thin.

The axis is brown near the base, and light yellow at the apices.

The main trunk is fairly cylindrical, but the branches are

slightly flattened. This flattening of the branches appears to be

due to the manner in which the ccenenchyma grows over the

axis, as the latter is fairly cylindrical in shape. The upper branches

are thin and flexible, but their apices are expanded. The polyps

are mainly confined to three sides of the axis. The general

surface of the ccnenchyma is covered by small, protruding

spicules. These spicules do not overlap, but there are only

minute spaces between them.

The polyps are also well protected by spicules. There is a

crown of spindles, in which the spicules are disposed en chevron,

in eight triangular areas which are very pointed towards the

apices. In each of these triangular areas there appear to be

about 8-12 spindles, which are placed more or less vertically to

the surface. At the base of those areas the spindles are arranged

in a continuous ring surrounding the polyp. The spindles

forming the ring also overlap one another, but not to such an

extent as those in the triangular areas. There appear to be about

ten spicules at any one place in this ring, enumerating in a

proximo-distal direction. Distally to this ring, spicules of

another form are situated on the polyp wall. These are very

protuberant, and stand more or less in a vertical direction.

These spicules are ‘“* Blattkeulen” and pass over basally into small

spicules of the same general form as those of the general surface

of the ccenenchyma, namely “‘ Kalkkorper.” The entire polyp is

thus well protected by spicules.

The polyps are capable of being withdrawn within the verruce,

but the latter are small and do not rise any distance beyond the

surface of the cenenchyma. The polyps vary considerably as to

CORALS FROM SOUTH AFRICA. 879

the degree of extension, but the apices of the tentacles are never

seen to any extent.

In all the better expanded polyps, the crown of spindles is very

apparent. This crown has frequently a reddish or brownish

coloration. The general surface of the colony has a greyish

colour, which is produced by the spicules.

Owing to the varying degree of extension, the polyps naturally

vary considerably in size,

The verruce have minute lobes, measuring as much as 2 mm.

in height and 1:6 mm. in diameter.

The ceenenchyma is thin at all parts and sometimes is 1 mm. in

thickness.

The axis of the secondary branches varies in diameter, from

about 1:3 mm. in the lower branches to 1 mm. in the apical

branches.

The interval between the polyps varies from about -1 mm.

to 19 mm. There is a terminal polyp at the apices of the

branches.

The spicules of the crown of the polyp are straight or curved

spindles, armed with blunt tubercles. The majority of the

spindles are very much curved, but a few are straight. The

tubercles have a tendency to great irregularity of shape, and

vary very considerably. The spicules from this part also vary

notably in shape, some being rod-like, others club-like, but

there are all transitions between the different forms. The

number of tubercles or spines also varies considerably, in some

spicules there are scarcely any, in others they are numerous. In

some of the more curved spindles there is a secondary offshoot

from the main axis about the middle of its length.

The spicules of the lower part of the polyp differ, as a rule, in

shape from those of the upper crown. Many of the spicules from

this part are “DBlattkeulen,” others are more of the form termed by

Kolliker “ Kalkkorper.’ There is much variety in the shape of

those spicules, probably every stage grading the one into the

other. There are also a few spicules similar to those occurring in

the crown of the polyp, namely curved spindles. There are other

spicules which may be more appropriately termed spinous clubs.

The superficial spicules of the coenenchyma are much smaller

than those of the polyp. They are mostly small “ Aalkkorper”

with irregular blunt processes coming off in all directions, less

frequently there are short spindles with broad processes.

The dimensions of the spicules are as follows :—Spindles of the

polyp crown, from 0:102 x 0-017 to 0°340 x 0:085 mm.; ‘ Biatt-

keulen” of the lower polyp, from 0°17 x 0°119 to 0-289 x 0°085 mm. ;

“ Kalkkorper” of the lower polyp, from 0°153 x 0-119 to 0°204 x

0:153 mm.; ‘“ Kalkkorper” of the eenenchyma, from 0-047 x 0-030

to 0105 x 0:076 mm. ; spindles of the cenenchyma, from 0:030 x

0:023 to 0-088 x0-061 mm.

There was unfortunately no bel attached to this specimen,

and thus the exact locality cannot be stated.

880 DR. J. STUART THOMSON ON

Nutting has given the following diagnosis of the genus

Muriceides :—

“ Muricemes W. & 8. (emended).

Muriceides Studer + Clematissa Studer, Archiv f. Naturgesch.,

Jahrg. liii. Bd. i. pp. 54, 55. .

Muriceides Wright & Studer + Clematissa Wright & Studer,

‘Challenger’ Reports, the Alcyonaria, 1889, pp. lu, lin, 105,

106.

‘“‘Calyces cylindrical, or in the form of truncated cones, their

walls filled with vertically-placed spindles, often modified into

clubs, discs, or triradiate forms vertically placed, but not ‘en

chevron,’ and not forming a true crown of points around the

margin. The opercular spindles are placed ‘en chevron’ on the

tentacle bases. The ccenenchyma contains spicules of various

forms, and the branches may, or may not, end in calyces.”

ACANTHOGORGIA ARMATA Verrill.

The colony is irregularly branched. The ccenenchyma is thin,

and filled with conspicuous, white spicules. The verruce are

elongate, often curved, capitate or clavate, surmounted by eight

groups of long, divergent, sharp spicules, with an irregular

““chevroned ” arrangement.

The axis has a yellowish-brown colour. The spicules are white,

rough, curved, and fusiform. The colour of the colony is ash-

rey.

; This species has been previously recorded by Hickson from

off Vasco da Gama Peak, 230 fathoms. There are three good

examples in the present collection.

Locality, etc—P.F. 18857. Cape Morgan, N. 2 W. 13 miles.

Depth, 250-320 fathoms. By shrimp trawl. Nature of bottom,

broken shells. Date, July 8, 1906.

Family PLEXAURID2.

EUNICELLA PAPILLOSA Esper. (Plate XLIII. figs. 2 & 3.)

The colony measures 162 mm. in length and 112 mm. in

breadth. It has a basal encrusting part measuring 11 mm. in

length and 15 mm. in breadth. The branches are all disposed

in one plane. The verruce occur over the entire surface of the

colony, even on the flat encrusting base. The main axis and

most of the branches are cylindrical, but occasionally some of the

upper branches are slightly flattened. Proceeding from the main

stem, the branches gradually decrease in diameter but expand

again near the apices. Some of the branches have a few local en-

largements, but these are probably not of specific importance.

The verruce give to the colony its papillated appearance; the

largest are about 2 mm. in height and 1:2 mm. in diameter,

the smallest are only fractions of a millimetre. The surfaces of

the verruce are covered bya mass of spicules. A rough transverse

CORALS FROM SOUTH AFRICA. 881

section through a branch shows (1) the horny brown axis in the

centre, (2) a ring of canals grouped round the axis, and (3) ex-

ternally, the cavities of the polyps arranged in a circumferential

series.

The spicules are situated on the verrucz, on the septa between

the polyp cavities and surrounding the horny axis. These spicules

are very abundant and are as a rule placed in a radial direction ;

they are of two kinds, viz. (1) the characteristic torch-like spicules

and (2) spindles. The torch-like spicules measure about 0-068 x

0:025 and the spindles from 0°0935 x 0:034 to 0-136 x 0°0374 mm.

A portion of a colony of Hunicella papillosa was dredged in

Simon’s Bay during the voyage of the‘ Challenger,’ and Hickson

has also recorded it from Rij Bank (Algoa Bay) at a depth of

25 fathoms. Hickson also draws attention to the remarkable

resemblance between the spicules of Gorgonia albicans and those

of Hunicella papillosa, more especially in regard to the torch-like

spicules.

He says, “The examination of my preparations of spicules

alone would lead any one to the conclusion that they were taken

from the same species. Yet the specimen of Hunicella papillosa

obtained on Rij Bank, 25 fathoms, has a cylindrical axis and

prominent verruce, whilst the specimen of Gorgonia albicans

picked up on the beach at Port Alfred has a very much flattened

axis, thin cenenchyma, and inconspicuous verruce.”

Hickson thinks that there is justification in transferring

Eunicella papillosa to the family Gorgoniide,and that Lunicella

papillosa and Gorgonia albicans should probably be included in

the same genus.

Locality, ete.—P.F. 15801. Off Seal Island, 8.S.E. 24 miles,

and Seal Island, S. by E. 2} miles. Procured by dredge. Depth,

9-10 fathoms. Nature of bottom, broken shells. Date, October 30,

1902.

PSAMMOGORGIA PULCHRA, sp.n. (PI. XLIII. fig. 5; Pl. XLV.

figs. 3a & b.)

This very beautiful form has an orange ground-colour, with

the red tips of the tentacles projecting. The colonies are not

all complete. A small colony (P.F. 858) consists of a fairly

flat, expanded, basal part which gives rise to two upright shoots,

one of which branches into two, the other being simple. The

basal part of the colony is 11 mm. in length, 6 mm. in breadth,

and 3mm. in height. The simple, upright shoot is 9mm. in

height, 2 mm. basally and 41mm. apically in diameter. The other

(divided) shoot is 28mm. in height, 3mm. in diameter at the

base and 5mm. at the apex. The branches are thus expanded at

the apex. Another specimen (15345), which is probably not quite

complete, has a very small basal part. The base gives rise to a

main shoot (2mm.in diameter), which at a distance of 9 mm. from

its origin sends off a lateral branch about 12 mm. in height; a

second small branch is given off from the main shoot after an

882 DR. J. STUART THOMSON ON

interval of 12 mm. This second branch measures 11 mm. in

length and 3 mm. in diameter.

The main shoot is 21 mm. in height and has a diameter of

5—6 mm. at the apex.

The surface of the cenenchyma is covered with large, yellow

spicules, which are easily seen with the naked eye. They lie ad-

jacent to one another but do not overlap, the boundaries of each

being clearly defined. These superficial spicules of the ccenen-

chyma do not appear to be arranged in any very definite manner

in relation to one another; they are of such varied sizes that any

great regularity is prevented. It may, however, be observed that

all the superficial spicules of the coenenchyma have their long axes

in the direction of the longitudinal axis of the colony; in the

verruce, the spicules have a corresponding position in relation to

the polyps.

The spicules of the cortex are (1) broad spindles, yellow in

colour, (2) long narrow spindles, of a pale colour. The spicules

project very much on the surface at all parts. The spicules of

the anthocodie are red and are long spindles arranged ‘ en chev-

ron’ in eight longitudinal, triangularareas. At the base of these

strips, a layer of spicules forms a ring round the polyp. This ring

is composed of two or three spicules lying one behind the other in

a proximo-distal direction.

The verruce have a circle of projecting spicules, only one layer

deep. This circle consists of 14-16 yellow spicules which lie with

their long axes parallel to the length of the polyps. These pro-

jecting spicules appear to be more prominent towards the apex of

the colony. The general appearance of the polyp spicules reminds

one, at first sight, of the setting of a stone in a finger-ring; the

anthocodia with its red spicules resembling a jewel which is

surrounded by little rivets, namely, the large, yellow spicules of

the verruce. The degree to which the anthocodize are exposed

beyond the verruce varies to some extent; as a rule, eight trian-

gular areas are easily seen, in other cases there is only a minute

area, with spicules or only an opening at the apex of the verruce.

The axis is horny in the centre, but with long narrow needles or

spindles on its surface. ‘These axial spicules are about 0-340 mm.

in length and 0:051 in breadth ; their margins are almost smooth.

Tn a rough transverse section, the axis is seen to be composed of

a large central grey area surrounded by a brown ring. The axis

is not penetrated by canals.

The polyps occur on all parts of the colony including the basal

encrusting part; they are arranged in a spiral manner on the

branches. The polyps are about 15mm. in height and 2 mm. in

diameter.

The dimensions of the spicules are :—(1) large, yellow spindles,

from 0-081 x 0:039 to 0-123 x 0-021 mm.; (2) small, yellow

spindles, from 0-042 x 0-024 to 0-093 x 0°033 mm.; (3) red spindles,

from 0:021 x0:0015 to 0°114x 00135 mm.; (4) long needle-like

spicules of the axis, 0°340 mm. in length by 0-051 mm. in breadth.

CORALS FROM SOUTH AFRICA. 883

Localities, etc.—P.F. 13345. Off Cape Morgan, N.N.W. 7

miles. Depth, 52 fathoms. By dredge. Nature of bottom, rocks,

sand, and shells. Date, August 12, 1901.

P.F. 858. Off and east of Cape Morgan. Depth, 36 fathoms.

By dredge. Nature of bottom, stones. Date, January 12, 1899.

Verrill’s diagnosis of the genus Psammogorgia is as follows :—

*Corallum dichotomous or subpinnate, with round branches.

Axis horn-like. Coenenchyma moderately thick, the surface finely

granulated with small rough spicula. Cells scattered, sometimes

flat, more frequently raised in the form of rounded verruce.

Polyps with rather large, elongated, slender warty spindles at the

bases of the tentacles. Spicula of the ceenenchyma mostly short,

thick, and very rough, warty spindles and rough, warty clubs of

moderate size.”

Eleven other species of Psammogorgia have been described.

KUPLEXAURA MEDIA, sp. n. (Plate XLIV. figs. 2 a-c.)

The colour of the colony is bright red, with white polyps. A

large part of the colony is concealed beneath a thick growth of

Hydroids, and thus the exact dimensions of the specimen are

rather uncertain. It was probably about 25 cm. in length.

In the lower part of the colony the axis is much flattened, but

towards the apex it becomes rounded. The large axis near the

base of the colony is 3°5 mm. in one diameter and 1:6 mm. in the

narrow diameter, The axis is very small towards the apex of the

colony, and is surrounded by the canals in a circular manner.

The axis is hollow, and has the structure described by Wright and

Studer for the genus Huplexaura.

The base of the colony is missing. The branches are all in one

plane and vary in diameter at different parts of their length.

They are thick near the base, narrower medianly, and expand

again near the apex.

The polyps are arranged in an irregular spiral round the

branches. The polyps are in many cases fairly well extended, but

others are retracted and appear only as minute pores on the

surface of the cenenchyma. These pores vary in shape, some are

circular or oval, others are slit-like and elongated in the direction

of the long axis of the branch. The pores are from °3 to ‘5 mm.

in diameter. A well-expanded polyp is about 1-2 mm. in length

and *5 mm. in diameter.

Each tentacle has about ten pairs of pinnules. The spicules are

situated at the bases of the tentacles and extend in a distal

direction. There is, firstly, a series of spicules on the polyp sur-

face in a line with the tentacles, and, secondly, another series

placed intermediate to the first. The second series has not so

many spicules as the first. The spicules in each of these series are

not arranged in a very regular manner.

The cenenchyma is dense and granular. The spicules are

arranged in a very dense layer in the cortex, but there are fewer

towards the axis.

884 DR. J. STUART THOMSON ON

The spicules of the cortex are mostly spindles with irregular,

blunt processes. Some have a simpler form than others. The

spicules of the centre resemble those of the cortex, but simpler

forms occur such as rods (with small processes) and crosses. ‘The

spicules of the polyps are very minute, needle-like or rod-like

spicules, with blunt processes.

The dimensions of the spicules are as follows :—Spindles of the

cortex, from 0:059 x 0-034 to 0°085 x 034 mm.; spindles of the

centre, from 0:051 x 0:034 to 0:085 x 0°051 mm.; rod-like spicules

of the polyps, from 0-064 x 0-008 to 0:112 x 0-034 mm.

This species resembles Hupleaaura brauert Kiikenthal, /. albida

Kiikenthal, and Z#. parciclados Wright & Studer. It has the

closest resemblance to Huplexaura brauert Kiikenthal, the spicules

of which are almost identical with it except in size. My specimen

differs, however, from Huplexaura brauert in the arrangement of

the polyp spicules and in other points. From the shape of the

spicules alone, one would probably conclude that they belonged to

the same species.

Locality, etc.—P.¥. 742. Between Roman Rock and Cape

Recife. Depth, 17 fathoms. By dredge. Nature of bottom,

corals. Date, December 12, 1898.

Family MALACOGORGIIDS.

MALACOGORGIA CAPENSIS Hickson.

The colony consists of a main stem (with an expanded, basal,

attaching part) which divides at some distance from the base into

two primary branches; one of the latter gives rise to three, the

other to eleven secondary branches. ‘The polyps are situated on

the secondary branches. ‘The main stem is cylindrical and

measures 50 mm. in length by 4mm. in diameter. The basal,

attaching part has a diameter of 6 mm. The polyp-bearing

branches are long and slender, the longest in my specimens being

80 mm., the shortest 27 mm. in length.

The branches originate in the manner shown in Hickson’s

figure, coming off right and left in one plane.

There are no polyps at the bases of the secondary branches, thus

leaving bare areas, 3-7 mm.in length. The younger polyps are

situated near the bases of the branches. The polyps are fully ex-

panded ; a large one measures 1 mm. in length and about °75 mm.

in diameter. The general appearance of the polyp reminds one

superficially of a contracted Hydra. On the lower part of the

branches the polyps have a bilateral arrangement, but higher up,

and especially near the apices, they originate on all sides and form

a dense cluster.

The tentacles are about 0:4 mm. in length and have 12 pairs of

pinnules.

The axial part of the secondary branches has a diameter of 0:5

to |] mm.

CORALS FROM SOUTH AFRICA. 885

IT am able to confirin Hickson’s statement regarding the absence

of spicules in this genus. My specimens are larger than that

described by Hickson, and the polyps in his form are also smaller.

My specimens, however, agree well with his description.

Hickson gave the following diagnosis of the genus J/alaco-

gorgia :—

“Colony slightly branched. Axis horny with no trace of lime.

No spicules in any part of the colony. Pelyps arranged bilaterally

in the plane of branching at the basal two-thirds of the secondary

branches and on ali sides of the terminal one-third of the

secondary branches.

“ Malacogorgia capensis, with the characters ef the genus.

Colour in spirit, white.”

Hickson placed this form in a new family, the Malacogorgiide.

He gave the characters of the new family as follows :—‘“ Colony

branched and upright. Axis slender, horny. Spicules and all

other forms of calcareous skeleton absent.”

Localities, etc. P.F. 18729. Bird Island (near Cape Seal),

EK. by N. + N. 5 miles. By large trawl. Depth, 40 fathoms.

Nature of bottem, mud. Date, August 29, 1905.

P.F. 703. Lat. 33° 53! 15" S., Long. 25° 51! 45" EK. By large

trawl. Depth, 26 fathoms. Nature ef bottem, mud.

Family Primnorps#,

STACHYODES GILCHRISTI, sp.n. (PI. XLIV. fig. 1; Pl. XLV.

figs. 2a & 6.)

The specimens are not complete. The branching is in one plane,

but is neither dichotomous nor regular. The polyps are arranged

in verticils which-are separated from one another by an interval

of about a millimetre, but the degree ef separation varies con-

siderably at different parts ef the celony. Thereare onan average

five verticils te every ten millimetres. The length of a large

verticil is about 3 mm. on the ad-axial and 2°5 mm. on the ab-

axial side. The verticils are about 5°5 mm. in diameter,

The axis, which is brown on the lower part of the colony and

yellow nearer the apex, is covered by sclerites which vary con-

siderably in size and shape. These sclerites covering the horny

axis are thin and fairly trausparent. The diameter ef the axis

is about *35 mm.

In each verticil there are usually five polyps. The polyps are

protected by three pairs of sclerites, namely an ad-axial pair,

amedian pair, and an ab-axial pair. The ad-axial pair consists of

long sclerites which are produced outwards in a lateral direction,

the median pair are smaller. The ad-axial and median spicules

have sharp, rather angular margins, and when the latter are

viewed from the side, they appear like spines. The ab-axial pair

of sclerites have slightly ribbed or dentate margins. The ad-axial

sclerites of the two lateral polyps form a ring or tunnel embracing

Proc. Zoou, Soc.—1911, No. LXI. 61

886 DR. J. STUART THOMSON ON

the main axis. The sclerites vary considerably in size, but they

probably range from 0:085 x 0:153 to 4:0 x 2:0 mm.

The diameter of a branch in the intervals between the

verticils is about 3 mm.

The sclerites covering the axis have the following forms :—(1)

triangular, (2) quadrangular, (3) forms with curved sides, and

(4) irregular forms.

The systematic position of this species is near Stachyodes

trilepis Pourtalés and Calyptrophora josephine Lindstrom.

Locality, etc—P.F. 11966. Cape Vidal (Natal), N.N.E. 4 N.

94 miles. Depth, 80-100 fathoms. By dredge. Nature of bottom,

rocks. Date, February 27, 1901.

THOUARELLA HICKSONI, sp. n. (Pl. XLIV. figs. 3a & 5;

TP; ROU die, Js)

The colony is more or less bottle-brush-like, but it gradually

tapers off towards the apex. Thedimensions of the colonies vary,

the largest specimen was 66 mm. in length and 25-28 mm. in

diameter. The central stem gives rise to the polyp-bearing

branches in a spiral manner.

These branches vary in length, they are longer near the base

of the main stem, and gradually decrease in size towards the apex.

The branches are as a rule simple, but in some cases there are

secondary and tertiary branches. The: primary branches are not

arranged in a regular spiral as the distance between their points

of origin is not the same in all cases. The interval between the

origin of two primary branches on the central stem is frequently

about 1 mm.

The axis is horny, flexible, yellow and iridescent, and its surface

is covered with imbricating spicules. It is more or less oval in

transverse section. When the spicules are removed from the

surface of the axis, fine longitudinal lines or strieareseen. There

are generally three spicules in a transverse row on the axis of a

branch. The diameter of a branch is about 0°391 mm.; that of

the horny axis about 0°136 mm.

Most of the polyps are situated on the primary, secondary and

tertiary branches, but sometimes they occur on the central stem.

On the branches they originate singly, and are disposed in a spiral

manner. ‘The polyps are pear-shaped, and their surfaces are

covered by imbricating scales. The size of the polyps varies con-

siderably, the following dimensions may be noted :—(1) 1-088 x

0°629 mm.; (2) 0-476 X0°306 mm.; (3) 1°343 x 0-595 mm. ; (4)

0:595 x 0°374 mm. ; (5) 0°935 x 0°663 mm. ; (6) 0°612 x 0-391 mm.

The interval on the branches between the polyps also varies but

not to any extent. ‘The distance is generally from °5 to °6 of a

millimetre. In some polyps, the tentacles were extended beyond

the verruce, and the pinnules were seen, though not sufticiently

well to determine their number. There are on an average about

twenty-six spicules on each verruca and these are disposed in six

transyerse (excluding the opercular spicules) and four or five

CORALS FROM SOUTH AFRICA. 887

longitudinal rows. The spicules of the verruce show considerable

variation; they may be grouped in two sets, namely (1) those

covering the lower or general surface of the vérruéz, and (2) the

apical or opercular spicules. The lower spicules of ‘the verruce

are triangular, rectangular, scale-like or plate-like spicules. In

their imbricate ar rangement and also in their shape, these spicules

remind one of the scales of fishes. Their surface is marked, except

near the margin, by a number of small, rounded pores, which are

arranged more or less in rows running in the direction of the long

axis of the polyp. The margins oe these scale-like spicules is

frequently, wholly or partially dentate. The opercular spicules

situated at the apices of the verruce differ in shape from those

last described. They are more or less triangular, with a

pr ojecting spine in front, the entire spicule resembling a Skate

(Raia) inform. These oper cular spicules are éight i in number and

their spines project towards the central opening of the verruca.

The surface of these opercular spicules have pores similar to those

of the lower part of the verruca.

The dimensions of the spicules are as follows: a) lower

spicules of the verruce from 0°1105 «x 0°1615 to 0°187 x 0:221 mm.

(>) the apical or opercular spicules (including the spine) nee

0-289 x 0°136 to 0°561 = 0°255 mm.

The spiculés covering the main stem of the colony are similar to

those of the lower part of the verruce, but are frequently much

smaller. They have a more or less ir regular arrangement and do

not as a rule everlap oné another. Their size is very variable,

ranging from 0-068 x 0-025 to 0:221 x 0°153 mm.

Around the basal part of the colony a Polyzoan is encrusted,

and intertwined among the upper branches are small Ophiuroids.

Locality, éte.—P.¥. 14265. Off Cape St. Francis, N.E. by E.

32 miles. By dredge. Depth, 74 fathoms. Nature of bottom,

rocks, Date, February 19 19, 1902.

Family GoRGONTID A

GorGoniIA CAPENSIS Hickson.

This is a béautiful example of a species described by Hickson

from the Cape in 1900.

Hickson’s specimen is larger, namely 250 mm. in length, while

my example is only 140 mm. long.

The spicules are warty spindles (*“ Doppelspindeln” of Kolliker).

The ENTE length of these spindles in Hickson’s specimen is

0-1 mm. ; in mine they are smaller, namely, from 0-0544 x 0:034

to 0° 0935 x 0-034 mm. .

In 1900 Hickson recorded this species as a viviparous Alcyo-

narian, and in 1905 Thomson and Henderson corroborated this

discovery i in specimens from Ceylon.

The Ceylon specimens were collected in deep as well as in

shallow water. They were larger than either of the Cape

specimens, and were practically white. Hickson’s specimen was

61%

888 DR. J. STUART THOMSON ON

collected off Cape St. Blaize, S.W. 3 W. 10 miles, at a depth of

40 fathoms. My specimen was collected farther to the north :—

Between Knysna Head and Nutze River, 3 to 4 miles off shore.

Depth, 35-49 fathoms. By large trawl. Nature of bottom, sand

and mud, The specimen had entertwined among its branches,

the string-like projections of a ‘“‘Mermaid’s Purse” and two or

three Brittle-stars.

GORGONIA FLAMMEA E. & 8S.

This species is collected in great abundance on the shore,

especially after storms, at many parts of the coast. It is

extremely common for example at Gordon’s Bay, False Bay.

Family GORGONELLID&.

ScIRPEARIA FURCATA Hickson, emend. Simpson.

1903. Scirpearia furcata Hickson.

1903. Scirpearia furcata var. ? Hickson.

1903. Scirpearella indica Hickson.

1905. Scirpearia sp., Thomson & Henderson.

1905. Scirpearella sp. B, Thomson & Henderson.

1905. Juncella elongata (Val.) Hickson.

1909. Scirpearella aurantiaca Thomson & Russell.

This species has recently been thoroughly revised by Simpson

and I therefore include his list of synonyms. In preparing his

valuable monograph on the Juncellid Group of the Gorgonellide,

Simpson had an opportunity of examining my specimens from

South Africa.

The South African localities are as follows :—

P.F. 13081. Hood Point, N. 53 miles. Depth, 42 fathoms,

Nature of bottom, saad and shells.

P.F. 858. Offand east of Cape Morgan. Depth, 36 fathoms.

Nature of bottom, stones.

P.¥.12377. Umblangakulu River mouth, N.W. by N.

74 miles. Depth, 50 fathoms. Nature of bottom, sand,

shells and sponge fragments.

P.F. 10841. Umbloti River mouth, N. by W. 2 W. 83 mules:

Depth, 40 fathoms. Nature of bottom, sand, shells, and

hard ground.

PIE 13030: Beacon east of East London, N. 7 E. 10 miles.

Depth, 52 fathoms. Nature of bottom, sand and shells.

P.F. 12033. Cone Point, N.W. 3 W. 4 miles. Depth, 34

fathoms. Nature of bottom, br folkea shells.

P.F. 11543. Tugela River mouth, ING Wey eee

miles.

Depth, 47 fathoms. Nature of bottom, broken shel

]

am Ce)

SCIRPEARTIA FLAGELLUM Studer, emend. Simpson,

1863. Juncella flagellum Johnson.

1864. Juncella flagellum Johnson.

1870. Viminella flagellum Gray.

CORALS FROM SOUTH AFRICA, 889

1881. Scirpearia flagellum Studer

1891. Scirpearia ochracea Studer.

1901. Scirpearia flagellum Studer.

1901. Scirpearia ochracea Studer.

1909. Scirpearia flagellum Thomson & Russell.

T have quoted the preceding synonyms from Simpson’s paper.

The following South African localities at which this species

occurs, may be noted ; ——

P.F. 12855. Buffalo River, East London, N. 15 miles. Depth,

310 fathoms. Nature of bottom, coral and mud.

P.E. 12061, O'Neil Peak, NW 2 W. 92 miles. Depth, 90

fathoms. Nature of bottom, broken shells.

P.F. 12107. O’Neil Peak, N.N.W. + W. 8 miles. Depth, 55

fathoms. Nature of bottom, broken shells.

P.F. 11586. Amatikulu River mouth, N.W. 2? N. 20 miles.

Depth, 62 fathoms. Nature of bottom, rocks and sponges

(hard ground).

HICKSONELLA SPIRALIS Simpson.

Juncella spiralis Hickson,

This species was described from the Cape by Hickson, later

revised by Simpson and named /icksonella spiralis. Unfortunately

the generic name, Hicksonella, is already preoccupied for an

entirely different Aleyonarian genus.

Simpson has given the following diagnosis of this species :—

“Colony unbranched; in the larger forms spirally twisted. The

ecenenchyma is thin and densely packed with scale-like spicules ;

the axis is composed of concentric laminz of a horny substance

in which a calcareous deposit is embedded. The polyps are re-

stricted to a region occupying two-thirds to three-fourths of the

circumference of the ceenenchyma,; a longitudinal bare tract oc-

cupies the remaining part. The verruce are long and club-shaped,

and are evidently not retractile into the coenenchyma; they are

closely packed together, and are covered with minute overlapping,

seale-like spicules. The flat thin scales on the aboral surface of

the tentacles form a sort of pseudo-operculum to the partially

retracted polyp.

The chief types of spicules are :—(1) In the ceenenchyma, very

thick spindles with close-set irregular warts, passing by gradual

transitions to almost spherical warty forms: (2) in the polyps, (a)

long thick spindles with a few long warts, (}) irregular forms and

crosses, (c) small, flat, thin scales.”

Localities, ete —P.F. 13152 4. Cape Morgan, N. 3 W. 104

miles. Depth, 77 fathoms. Nature of bottom, rocks aid broken

shells.

P.F. 13121. Cape Morgan, N.N.E. 92 miles. Depth, 47

fathoms. Nature of bottom, broken shells.

Hickson’s specimens were also collected off Cape Morgan, at a

depth of 36 fathoms.

890 DR. J. STUART THOMSON ON

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CORALS FROM SOUTH AFRICA. 891

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892 ON CORALS FROM SOUTH AFRICA.

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EXPLANATION OF THE PLATES,

PruatE XLIII.

. Ceratoisis ramosa Hickson. X 1}.

. Hunicella papillosa Wsper. Nat. size.

. Eunicella papillosa, part of branch of (with cephalopod egg-capsule

encircling the branch). X 6.

. Suberia capensis, sp.n. Nat. size.

. Psammogorgia pulchra, sp.n. X 1.

n> Go to a

IAS), eH PIL OIL AVAL,

ae PTT

9 : Huth lith.eb tap .

SCUENENS) (Ol {ESANOIDIO INGE IE0e 2 NES).

ON THE SKULLS OF CYNODONT REPTILES. 893

Prare XLIV.

Fig. 1. Stachyodes gilchristi, sp. n., part of branch of. X 8.

2a. Hupleraura media, sp. u. Spicules of central trunk.

2b. 33 6 i Spicules of external trunk.

2c. y x Spicules of polyps.

3a. Thouar ella hicksoni, sp. n., verruca of. X 45.

3b. 5 apex of verruca of. x 100.

Aa. Muriceides fusca, sp. n. Spicnles of upper part of polyp.

i 5 +5 Spicules of lower part of polyp.

4e. 5 on - Spicules of coenenchyma.

Pratt XLV.

Fig. 1. Thouarella hicksoni, sp.n. Spicules of verruea.

2a & b. Stachyodes gilchristi, sp. u., sclerites of.

3a. EA hg pulchra, sp. n., red spicules of.

35. 53 » yellow spicules of.

4a. Suberia capensis, sp. n. Spicules of central trunk.

4b. 5 fi 5 Spicules of external trunk.

Ac. oe ne 3 Spicules of polyp.

38. On the Structure of the Skullin Cynodont Reptiles *.

By R. Broom, M.D., D.Sc., C.M.Z.S.

[Received April 10, 1911: Read May 28, 1911. ]

(Plate XLVI.7 and Text-figures 168-180.)

Historical and Introductory.

In 1853 the British Museum received from Andrew Bain the

first known skulls of fossil reptiles with a mammal-like arrange-

ment of the teeth. These ultimately became the types of

Lycosaurus tigrinus and Cynodraco serridens. In 1858 Sir

George Grey presented the skulls which were shortly afterwards

described by Owen as Galesaurus planiceps and Cynochampsa

laniaria. The Galesawrus skull though crushed was nearly

complete, and being so very vemarkably mammal-like Owen

almost immediately described it 1n a paper read before the

Geological Society on 20th April, 1859.

Although for seventeen years nothing further descriptive of

any of the reptiles with a mammal-like dentition was published,

it is necessary to briefly consider some of Owen’s other work in

the interval to clear up a certain confusion of nomenclature. In

1859 Owen gave to the world his famous classification of the

fossil reptiles, and though he formed the Order dAnomodontia

for the South African reptiles of the Dicynodont type, he care-

fully omitted all reference to those reptiles, like Galesawrus and

Cynochampsa, with a mammal-like dentition. When in 1861 he

published his ‘ Paleontology,’ feeling compelled to put the

remarkable Galesawrus somewhere, he made it the type of a

“family ” of the Anomodontia, calling it the Cynodontia, doubtless

* On p. 902 Dr. Broom names a new species, viz. Cynoguathus seeleyi.— Kh pitToR.

+ For explanation of the Plate see p. 925.

894 DR. R. BROOM ON THE

recognising that Galeswurus was in some way related to Dicyno-

don, but hesitating to make a new Order on the evidence of a

single skull. As he still defined Anomodontia as reptiles with

‘ teeth wanting or limited to a single maxillary pair,” it is mani-

fest he did not regard Galesaurus as really an Anomodont.

In 1876, when Owen issued his‘ Catalogue of the South African

Fossil Reptiles ’ he put all the forms with a mammal-hke dentition

into a new Order, the Theriodontia. In 1903 I showed that Owen's

Theriodontia is not a natural order, for it included two groups

which, though agreeing in having the dentition specialised into

incisors, canines, and molars, and possibly the one being ancestral

to the other, were yet so dissimilar that they could not be well

kept together. The more primitive group, which occurs only

in Permian beds, has simple molars, an open Rhynchocephaloid

palate, a transpalatine bone, large angular and surangular bones,

a single occipital condyle, no acromion process, and apparently a

digital formula of 2, 3, 4,5, 3. The higher group, which is known

only from Upper Triassic beds, has usually specialised molars, a

secondary palate asin Mammals, no transpalatine, small angular

and surangular bones, two occipital condyles, an acromion process,

and a digital formula 2, 3, 3, 3, 3. As Cynodontia was the name

first applied to animals of the Galesaurus type, this title should

be retained for the higher group. For the lower forms I proposed

the name 'Therocephalia. The name Theriodontia should be

dropped, as only likely to lead to confusion.

Among the new forms described by Owen in his Catalogue is a

badly weathered small Cynodont skull somewhat resembling that

of Galesaurus and named WVythosaurus larvatus. In 1887 he

described another small but well-preserved skull which he believed

to be an additional specimen of Galesaurus.

Most of our knowledge of the Cynodonts, however, is due to

Seeley, who, as the result of his expedition to South Africa, not

only came across the skulls of many new types, principally in the

collections of Dr. Kannemeyer, Mr. A. Brown, and the Albany

Museum, but for the first time obtained most of the skeleton of

some Cynodonts. In one paper issued in 1896 he described a

very fine skull with most of the vertebral column, the lmb-

girdles, and portions of the limbs of a large carnivorous type,

which he called Cynognathus crateronotus, also a fine skull of an

allied form called Cynognathus platyceps from the Albany Museum

collection. In other papers he described new types of Cynodont

reptiles with flat-topped molars. Of these the best known types

are Gomphognathus, Diademodon, and T'rirachodon. These were

regarded by Seeley as belonging to a distinct Order, which he

called Gomphodontia; but as, apart from the specialisation of

the molars, there are no characters of any importance to distin-

guish the Gomphodonts from the Cynodonts, it seems to me

impossible to regard them as forming more than a Family of

the Cynodontia.

Within the last eight years I have been so fortunateas to come

SKULLS OF CYNODONT REPTILES. 895

across a considerable number of new Cynodonts, mostly collected

by Mr. A. Brown, and also to add a good many facts to our

knowledge of the anatomical structure. Probably the most

important of the recent finds has been the discovery by myself

of the nearly perfect skull which I have called Bauria cynops.

Of all extinct reptilian groups there is probably no one of

greater interest than the Cynodontia. Many years ago Owen

recognised the remarkable mammalian characters in the Permian

and Triassic South Afvican reptiles, and though the Cynodonts

were so little known, he ventured to suggest that certain of the

Anomodonts were fairly closely allied, and perhaps ancestral, to

the Monotremes. Cope held much the same view. When the

very much more mammal-like Cynodonts were described by

Seeley, many recognised in this higher group the looked for Sauro-

Mammalia. Osborn has been the chief advocate of this opinion.

Seeley himself, though at, first inclining to it, afterwards came to

the conclusion that the Mammals were in no way nearly related

to the Cynodonts, but sprang from some unknown ancestor that

lived in Devonian or Silurian times.

If the Cynodonts are not nearly related to Mammals, the group

is still of great interest as showing a marvellous parallelism with

the Mammals in skull, teeth, girdles, limbs, and digits; but if,

as all recent work seems to indicate still more clearly, the mam-

malian ancestor was probably a Cynodont, the group becomes

vested with an interest altogether unique, and everything bearing

on it becomes worthy of the most careful study. I have fortu-

nately been able to examine every known skull, and in the present

paper I give the results of my researches. As the paper is

morphological rather than systematic, | propose to give a detailed

account of the skulls of only the principal Cynodont types, and

to consider more fully those points which seem to have a special

bearing on the question of mammalian descent.

Bauria.

(Pl. XLVI. figs. 6, 7, 8, and text-figs. 168, 169.)

Though Bauria cynops occurs in the same horizon as Cyno-

gnathus, it is the most primitive Cynodont at present known, and

may be regarded as the type of a distinct family which may be

called the Bauride.

As I have just recently, at considerable length, described the

only known skull of Bawria cynops, it will be unnecessary here

to do more than supplement that description in a few details

and to consider its relationships to the other known Cynodonts,

the Therocephalians, and the Mammals.

Further development and examination of the. skull has revealed

one or two points not previously noted. Under the nostril and

forming not only its floor but covering a considerable part of

the premaxillary is a large septomaxillary bone. The lachrymal

and prefrontal bones cannot in the specimen be clearly separated

896 DR. R. BROOM ON THE

from one another, but it is quite manifest that the lachrymal

is small and the prefrontal only moderate-sized. The nasal

extends well back and forms a broad suture with the frontal.

The frontals form the greater part of the interorbital region

and most of the supraorbital ridge. There are no postfrontals,

and the postorbitals are remarkable in forming only a postorbital

process and in not meeting the jugal to form a postorbital arch.

The jugal is slender and passes back nearly to the articular region.

The portion of the squamosal that supports the quadrate is well

developed, but the zygomatic portion is slender.

Text-fig. 168.*

Side view of the type and only known specimen of Bawria cynops. Since the speci-

men was first figured it has been considerably further developed at the British

Museum and by myself. The jaws are represented as closed. ‘The molar teeth

must meet one another as shown in the figure. When first described the teeth

were regarded as round, but further development shows that they are about

twice as broad in one diameter as in the other. Though the incisors are mostly

broken the impressions of the greater part of each is preserved, and the lower

must have met the upper as shown in the figure. The jugal arch is represented

in its central part only by the impression, but there is no doubt it must have

been practically as restored. It certainly did not meet the postorbital, which

is perfectly preserved on both sides. All the sutures shown in unbroken line

can be clearly made out.

The palate is as in typical Cynodonts, the secondary palate

being as well developed. The vomer, palatines, and pterygoid, so

far as can be seen, all are of the ordinary Cynodont type. The

lower part of the alisphenoid appears to be of the same type as in

higher Cynodonts, articulating with the basisphenoid and passing

out to the quadrate. It is just possible, however, that this

outward extension may be, as in the Therocephalians, entirely

formed by the pterygoid. The basisphenoid is unlike that of

either the Anomodonts, Therocephalians, or higher Cynodonts.

It is short, with a broad articulation for the basioccipital behind,

and a narrow articulation in front for the alisphenoid and

probably the vomer. From its under surface there passes down

* For explanation of the lettering in the: text-figures see p. 925.

SKULLS OF CYNODONT REPTILES. 897

a narrow deep median ridge, which is nearly as deep as the bone

is long. The basioccipital resembles considerably that of the

Anomodonts in having a pair of short postero-lateral processes

Text-fig. 169.

~Pmx.

757

ait,

FO.

Upper view of the skull of Bauria cynops.

which meet the basisphenoid. The condyle is unique (Pl. XLVI.

fig. 8). Itis asingle condyle, only partly divided into two by a

deep median groove. It is thus in type intermediate between the

condyle of the Therocephalian and that of the higher Cynodont.

898 DR. R. BROOM ON THE

The large foramen for nerves 1x., X., x1., and xi). lies by the side

of the basioccipital and in front of a bone which is probably part of

the opisthotic. Nerve xii. enters the foramen exactly as it does

in the higher Cynodonts. On the inside of the skull it has two

small distinct canals, which pass forwards and both unite with

the large foramen.

The bone which is supposed to be the stapes is shown in fig. 8

(Pl. XLVI). It is apparently a little displaced forwards.

The lower jaw has a fairly large surangular and angular, the

dentary being considerably in front of the articular region.

Taking all the characters into consideration, Bauria becomes

one of the most interesting intermediate types ever discovered.

Though an undoubted Cynodont, it retains many of the Thero-

cephalian characters. On the other hand, though on the whole

it is less mammal-like than the higher Cynodonts, it has some

mammalian characters which the others have lost.

The following are Therocephalian characters usually lost in

Cynodonts but retained in Lawria :—-

1. Large septomaxillaries forming part of the facial surface.

2. Moderate prefrontals.

3. Large frontals forming most of the interorbital region.

4. Feeble zygomatic arch.

5. The two occipital condyles so imperfectly separated as to

represent practically a single condyle.

6. Large size of angular and surangular.

7. Shape of the articular.

8. Simple condition of the molar teeth.

In the following characters Bawria is nearer to the mammalian

ancestor than are the higher Cyiodonts :—

1. Large size of septomaxillaries and development on face.

A somewhat similar condition is found in primitive -

Multituberculata (e. g. Tritylodon), also in Monotremata,

as shown by Gaupp in Lchidna embryo.

. Large frontals.

Complete loss of parietal foramen,

Absence of postorbital arch.

5. Simple condition of molar teeth.

bm Gols

Nythosaurus,

(Text=fig. 170.)

The type of Galesaurus planiceps is a somewhat céushed skull

with the bones in an unsatisfactory condition for showing sutures.

No second specimen of Galesaurus has ever been discovered.

In 1876 Owen des¢ribed an imperfect skull as Vythosaurus lar-

vatus. In 1887 he described another skull in fairly good preserva-

tion which he believed to be another specimen of Galesaurus.

Seeley in 1894 showed that this supposed second specimen of

SKULLS OF GYNODONT REPTILES. 899

Galesaurus differed greatly from the type, and gave it the name of

Thrinaxodon liorhinus.

There is, in my opinion, not the least doubt that Seeley was

right in regarding the 1887 skull as belonging to a very different,

animal from the 1859 one. In fact it seems strange that any one

should ever have thought them the same. Galesaurus has a

dental formula apparently of 1. 2 c. a m. ae the 1887 specimen

G. , m. q Tn the 1859 specimen 10 molars

occupy 20 mm.; in the 1887 specimen 7 molars occupy 20 mm.

But while the two supposed Galesaurus specimens represent

different genera, two other imperfect specimens in the British

Museum show that the 1887 specimen is the same animal as was

described in 1876 as Nythosauwrus larvatus. Hence the well-

known skull which is figured in various text-books as Galesaurus

must in future be called Vythosaurus.

Nythosaurus is a much higher type than Bawria: but though

it comes fairly close to the higher Cynodonts such as Cynognathus,

it should, I think, be taken as the representative of a distinct

family, the Galesauride. From the various specimens in the

British Museum it is possible to make an almost complete restora-

tion of the skull.

has a formula of 1.

Text-fig. 170.

Side view of the skull of Nythosawrus larvatus. The drawing is mainly that of the

best preserved specimen in the British Museum, compared with the other

specimens and slightly restored from them. ‘The teeth are represented in the

mature condition.

The septomaxillary though smaller than in Bauria still appears

on the face. The nasal is large and very broad at its upper end.

The lachrymal is large, and though the prefrontal is only of

moderate size, it joins with the postorbital and completely shuts

out the frontal from the orbit. The postorbital forms with the

900° DR. R. BROOM ON THE

jugal a rather feeble postorbital arch. The zygomatic arch is

formed by the jugal and the squamosal. ‘The jugal extends

nearly back to the articular region, and the squamosal nearly

forward to the base of the postorbital arch. The squamosal is

not unlike that of Bauwria, but the zygomatic portion is much

better developed ; so that as regards the squamosal Vythosaurus

is intermediate between Bauria and Cynognathus. The quadrate

is of the same type as in the better known Cynognathus.

The palate, so far as known, agrees fairly well with the Cyno-

gnathus type, and the occipital condyie is double.

The lower jaw has a large dentary, but there is no trace even

of a condylar process. The angular and surangular are fairly

large and still resemble considerably the Therocepbalian type.

The articular also resembles that of the earlier rather than that

of the later types.

Nythosaurus is perhaps the most mammal-like of the known

Cynodonts. The zygomatic arch is exceedingly like that of most

primitive mammals, and if the prefrontal and postorbital bones

were lost and the internasal process of the premaxilla aborted

there would be nothing left in the side view of the skull to

distinguish it from that of a mammal. The lower jaw with its

fairly large angular and surangular is still much less like the

mammalian condition than what we see im the higher Cynodonts,

and the articular is of the same primitive type seen in Lauria.

The dentition though very primitive is considerably more

1 7

3) D mM.

comes very near to that of the typical mammal, and that of

highly evolved than in Baeria. The TOLEDO, ty Sy Ce

o 4) 1 i2

Galesawrus, i. 5, C- 72 M. 7, 1S near that of the ancestral mammal.

The difference in the teeth in some of the specimens of Vytho-

saurus is apparently due to the fact that in some the teeth belong

to the first set and in ethers to the second.

Cynognathus,

(Pl. XLVI. figs. 1 & 2, and text-figs. 171, 172.)

The genus Cynognathus is known by the very fine skull of

(. crateronotus in the British Museum, the type skull of C. platy-

ceps in the Albany Museum, a fairly good skull of C. berryiin the

S. African Museum, and three or four less satisfactory specimens.

Seeley has given a fairly full account of the skull of Cyno-

gnathus crateronotus, but unfortunately a number of his figures are |

so indifferently reproduced that they convey no more to the

student than does the plaster cast. And further, while most of

his determinations are correct, he unfortunately suggests so many

alternative possibilities that the morphologist is left compara-

tively helpless.

The figure given by Seeley of the side view of the skull of

C. crateronotus gives an excellent idea of the general form of the

skull and of the structure of the temporal region, except that the

SKULLS OF CYNODONT REPTILES. 901

supposed perforation in the zygomatic arch is, in my opinion, not

a natural feature and has been produced post mortem. The side

view which I give of the skull of C. platyceps is fairly similar,

except that the skull is here much broader and flatter, and in

this species, at least, there is no trace of an opening in the

zygoma.

The snout of Cynognathus differs from that of Sauria and

Nythosaurus chiefly in the fact that the septomaxillary does not

appear on the face, though it can be readily seen within the

nostril.

Text-fig. 171.

hip,

Vi Less

Side view of the skull of Cynognathus platyceps. With the exception of the

tront of the snout the drawing is made trom the very fine type skull in the

Albany Museum. The front of the snout is from the specimen described by

Seeley as ? Cynognathus leptorhinus. As I showed some years ago, this is

unquestionably the snout of a nearly full-grown specimen of Cynognathus

platyceps.

The premaxilla is relatively rather larger than in Vythosaurus,

while the maxilla is about equally well developed in the two

genera. The canine is, however, much further forward in Cyno-

gnathus than in the smaller genus.

The nasal bone is fairly similar in the two genera, being broad

both in front and behind and narrow in the middle.

The lachrymal extends further forward than in WVythosaurus,

the portion showing on the face being nearly as large as the

orbit.

The prefrontal is a long narrow bone which forms the greater

part of the upper margin of the orbit, and by meeting with the

postorbital completely shuts out the frontal from the orbital

margin.

The frontals are each about four times as long as_ broad.

In front they meet the nasals. Laterally they are in contact

with the prefrontals and postorbitals. Posteriorly they taper

away to narrow points, which mect the anterior ends of the

parietals.

The postorbital is a large triangular bone. In front it meets

Proc. Zoou. Soc.—1911, No. LXII. 62

902 DR. R. BROOM ON THE

the prefr ontal and forms part of the orbital margin. Externally

it gives a large articulation to the jugal and a small articulation

to the squamosal, Internally it overlaps the parietal for only a

very short distance.

The jugal is relatively considerably larger than in Vythosaurus.

Anteriorly it meets the maxilla and jachrymal, but extends further

forward thanineither Vythosaurus or Bauria. Immediately below

the orbit there is a small but very distinct tubercle. The

ascending process of the jugal is unusually broad and forms a

large articulation with the postorbital, the two together forming

a very strong postorbital arch. The posterior portion of the jugal

extends to the articular region.

The parietal is a narrow bone which forms a low median crest.

There is a small pineal foramen.

The squamosal is the largest bone in the skull, with the excep-

tion of the dentary. The inner and posterior portion forms

nearly the whole of the back wall of the temporal fossa, and has

a large articulation with the parietal. This back portion of the

squamosal is very thin and is closely united with the flat upper

expansion of the opisthotic. The squamosal forms the outer wall of

the lateral occipital foramen. Inferiorly it meets the exoccipital

and on passing outwards supports the small quadrate. The

zygomatic portion is much larger than in Vythosaurus, articulating

with the whole of the upper side of the posterior limb of the

jugal and meeting the postorbital. There is a deep groove along

the middle of the back part of the zygomatic portion, whieh

curves downwards and inwards, and, most probably, the pos-

terior part was for the support of the external auditory canal.

The occiput is best known from the specimen of Cynognathas

berryt in the 8. African Museum, which though imperfect shows

the sutures very distinctly. The lar “ge occiput figured by Seeley*

(p. 130) and doubtfully referred by him to @. hoe yi, 1S, IN my

opinion, considerably too large. It also differs from the known

occiput of C. berryi in the ‘shape of the foramen magnum, the

slope of the exoccipitals, and the moulding of the interparietal

region, Itis pretty clearly not the occiput of C. crateronotus,

while it is much too lar ge to be that of C. platyceps. As it thus

seems to belong toa new species, it may appropriately be named

Cynognathus seeler Ye

The occiput proper (Pl. XLVI. fig. 2) is made up of seven

bones, or, if we include the squamosals, nine. About a quarter

of the occiput proper is formed by a median bone, which is

apparently the interparietal. I have not seen any specimen which

enables me clearly to differentiate this bone from the parietal in

front, but, judging by analogy and by the direction of the fibres

of the bone, the probability seems much in favour of its being a

distinct interparietal.

On either side of the interparietal is articulated a large bone,

which is apparently the opisthotic. Its occipital por tion is com-

paratively thin and to a considerable extent covered in front by

* Phil. Trans. Vol. elxxxvi. B. (1895).

SKULLS OF CYNODONT REPTILES. 903

the squamosal and parietal. Inferiorly it articulates with the

exoccipital and to a large extent with the supraoccipital, It

Text-fig. 172.

Upper view of the skull of Cynognathus platyceps-

From the same specimen as text-fig. 171.

forms the upper and inner wall of the lateral occipital foramen,

and passes well forward below the edge of the parietal and appears

to articulate with the large prootic.

62*

904 DR. R. BROOM ON THE

The supraoccipital is completely anchylosed to the exoccipitals,

but its probable limits are indicated in the figure given, [mme-

diately above the foramen magnum is a small ‘rounded bony knob.

‘The exoccipitals form the greater part of the oceipital condyles,

whieh are remarkably mammal-like. Above each condyle is a

narrow fissure or groove leading into the foramen magnum, but

it seems too small to have been for the transmission of either an

artery or a vein, and most probably it was not for a nerve.

Possibly ib is merely owing to the elevation of a portion of the

bone for the attachment of a ligament. The exoccipital passes out

aconsiderable distance under the lateral foramen and articulates

with the squamosal.

The quadrate is well shown in Seeley’s figure 8. It is a

relatively small bone, which is tirmly fixed on to the squamosal by

the main part being in front and two long processes being behind

its lower projection. Referring to the posterior delicate processes

Seeley says: “ I am unable to attirm that they represent auditory

ossicles.” They are unquestionably parts of the quadrate and

have nothing to do with the auditory function. The quadrate

forms the greater part of the articular surface for the lower jaw,

but not the whole of it, part of the squamosal also forming

a portion of the articulation. This is particularly interesting in

view of the fact that in the Monotremes the lower jaw hinges

directly on the squamosal bone.

The palate of Cynognathus is nearly wholly known, the only

points concerning which we are still ignorant being the relations of

the bones round the anterior palatine foramina, the nature of the

middle part of the basicranium, and the relations of the palatine

and pter ygoid to the jugal.

‘There is a large secondary palate formed by the maxille and

palatines exactly as in Mammals. In a recent paper Seeley

describes what he believes to be teeth on the palate of

Cynognathus. I have, however, had an opportunity of examining

Seeley’s specimen and believe the supposed teeth to be merely

irregularities of the bony surface, possibly pathological. Certainly

in the other specimens I have examined there is no trace of

anything like teeth. The hard palate ends in the middle line

opposite the front of the third last molar.

The pterygoids are large and have well- -developed pterygoid

processes, which lie close along the inside of the jaws as in reptiles

generally. There is no transpalatine or ectopterygoid bone.

Instead of, as in most reptiles, the pterygoid having a posterior

process which extends to the quadrate, it here ends near the

middle of the inner wall of the temporal fossa, the posterior

continuation which looks like pterygoid being really part of the

alisphenoid bone.

The vomer is a large median bone which posteriorly les between

the two pterygoids. In front it forms a vertical plate which

supports the secondary palate exactly as in Mammals and extends

to near the front of the snout.

SKULLS OF CYNODONT REPTILES. ~ 905

The alisphenoid bone is one of the most interesting bones in

the skull. It isa large flat bone which extends from the pterygoid

below to the parietal above. It is well seen in the type skull of

Cynognathus crateronotus, but even better in the Capetown

specimen of C. berryi (P]. XLVI. fig. 1). The upper portion of

the bone is irregularly quadrilateral. The upper side articulates

with the parietal, and the posterior with what I believe to be the

prootic. Between the alisphenoid and the prootic are two large

oval foramina. At the posterior and lower corner the alisphenoid

is continued as a slender bone to the quadrate. At its anterior

and lower corner it meets the pterygoids and clasps the basi-

sphenoid. There appears to be an opening into the brain-cavity

between the base of the alisphenoid and the basisphenoid.

The basisphenoid is clasped by the alisphenoids and meets the

basioccipital posteriorly.

The basioccipital is a small bone lying behind the basisphenoid.

It forms the middle part of the occipital condylar region. On

each side’ there is a large round foramen which is pretty certainly

the foramen for the exit of nerves ix., x., xi., and xii.

Between the outer part of the basioccipital and the quadrate

there stretches a rounded pillow-like bone concerning which there

may be some difference of opinion. I believe it to be the

stapes, for reasons which will be stated later.

The lower jaw is remarkable for the great size of the dentary,

which posteriorly nearly reaches the articulation. Elsewhere I

have dealt at some length with the structure of the jaw. The

splenial is long and slender. The surangular and angular are also

feeble splint-like bones. The articular is fairly well developed but

short. I cannot satisfy myself that there is a distinct coronoid

bone as is stated by Seeley.

The only points in which the Cynognathus skull is nearer to

the mammal than that of Bawria and Nythosaurus are: (a) the

closer approach of the jugal to the articulation, (6) the greater

development of the dentary, (c) the greater reduction of the

angular and surangular, and (d) the more mammal-like occipital

condyle. On the whole it is not so near the mammalian ancestor

as either Bauria ov Nythosaurus.

The dental formula appears to be i. 4, ¢. * 3

7 Col ay) Miler ge

Trirachodon.

(Text-figs. 173 & 174.)

Trirachodon is best known by the type skull which is in the

Albany Museum. Though the skull is immature and much

crushed it is practically perfect. Two or three other known skulls

though imperfect show the uncrushed condition of the greater part

of the adult skull.

The premaxilla is smaller than in Cynognathus, not meeting the

nasal behind the nostril, at least not on the face.

906 : DR. R. BROOM ON THE

The septomaxillary is entirely in the nostril, forming no part of

the face.

The maxilla is long and extends far back below the orbit.

The snout is fairly broad at the root of the canine and along the

upper part of the maxilla, but is much narrowed in the molar

region. There are two foramina for the maxillary branch of

nerve Y.

The nasal is moderately broad in front, narrow in the middle,

and very broad behind.

The lachrymal is small, but forms most of the anterior wall of

the orbit.

Text-fig. 173.

Ju-

Zz_

Side view of the skull of Trirachodon kannemeyeri. The drawing is chiefly founded

on the crushed and immature skull which forms the type. Two mature

and uncrushed but imperfect skulls in the Albany Museum and a good snout

in my own collection have made it possible to correct the crushing of the type

and completely restore the skull in the adult cqndition.

The prefrontal is about twice as Jong as broad, and forms most

of the upper margin of the orbit; it unites, as in Cynognathus,

with the postorbital, completely shutting out the frontal from the

orbital margin.

The frontal is fairly like that of Cynognathus, but it does not

extend so far forward. Posteriorly, as in the former genus, it

tapers away between the postorbitals.

The postorbital is more like that of Vythosaurus. It forms the

upper third of the postorbital arch, uniting with the jugal. It

extends backwards on the side of the parietal a little beyond the

pineal foramen.

The parietal is like that of Cynognathus, but the pineal foramen

is much smaller.

The jugal is, on the whole, like that of Cynognathus. It has,

however, the inferior process much better developed. It forms

the lower and posterior half of the orbital margin.

The squamosal differs from that of Cynognathus in not meeting

the postorbital, but ending in front above the jugal, very much as

SKULLS OF CYNODONT REPTILES. 907

in Vythosawrus. As the back of the skull is relatively narrower

than in Cynognathus, the posterior part of the squamosal differs

considerably in contour.

Text-fig. 174.

) <a Th

‘ BRASS Nes

ANS RSSET

ASS

BSR TA

THT

Lsbeeas

as:

Lif

none,

ere,

NS a we

Upper view of the skull of Tvirachodon kanneneyert.

From the same specimens as fig. 173.

The quadrate, so far as can be seen, is small but not unlike the

better known types.

The stapes is a very slender straight bone about the thickness

908 DR. R. BROOM ON THE

of a pin, and having similar relations to the stapes in Cyno-

gnathus.

The occiput is not well known, but is apparently very similar

to that of Cynognathus.

The lower jaw has a very large coronoid process and a condylar

process which nearly reaches the articulation. The articular,

angular, and surangular are on the whole very similar to those of

Cynognathus.

One of the most striking points of difference from Vythosaurus

and Cynognathus is in the structure of the molar teeth, which

have flattened tops, and the lower molars instead of passing inside

of them, as in these other genera, meet them much in the same

way as do the molars in Mammals.

The dental formula is the same as in Oynognathus, viz.:

» 4 1 9

Tas) © ap 0S 50

Diademodon and Gomphognathus.

(Pl. XLVI. fig. 9, & text-figs. 175-178.)

Diademodon resembles Gomphognathus so closely, differing only

in size and in the number of molars, that there is some reason for

suspecting that Diademodon may be animmature Gomphognathus.

If this turns out to be the case, the genus must take the earlier

name Diademodon. The following description of a skull is based

on a beautifully preserved skull in the British Museum, which

may be called Gomphognathus minor, but which unfortunately has

lost the snout, on a fairly good skull of D. mastacus in the South

African Museum, on two other good skulls of Gomphognathus in

the British Museum, and on the type skull of Gomphognathus

kannemeyert in the Albany Museum. As the result of the

examination of this very fine material, the Gomphognathus skull

is better known even than that of Cynognathus, and almost as

well as that of the living Ornithorhynchus.

The premaxilla is fairly large, but, as in all other Cynodonts,

considerably overlapped by the front of the maxilla, It has a

strong internasal process. and forms the anterior and most of the

lower border of the nostril. It has a very considerable palatal

development, the two bones meeting in the middle line behind the

anterior palatine foramen.

The septomaxillary is well developed, lying along the greater

part of the outer wall of the nostril. Most of it is within the

nostril, but a small part of the upper end appears on the face.

The maxilla is not unlike that of Zrirachodon. The maxillary

branch of nerve v. has two foramina. On the upper part of the

maxilla close to the nasal is a small oval depression, presumably

for the lodgment of a gland.

The nasal is narrow in front but broad behind. The nostrils

look more upwards than in any of the previously described

Cynodonts, and the nasal passes forward between them to a

narrow process.

SKULLS OF CYNODONT REPTILES. 909

The lachrymal is of fair size. In Gomphognathus minor it

forms the front of the orbit and much of the inner wall. In

Gomphognathus polyphagus ib is considerably larger, extending to

part of the upper margin.

Text-fig. 175.

Tiki

Side view of the skull of Gomphognathus minor. With the exception of the front

half of the snout and the lower jaw, the drawing is from the beautifully preserved

type in the British Museum. The front of the snout is restored from a specimen

ot Gomphognathus kannemeyeri 1m the British Museum, and the lower jaw 1s

from the lower jaw of the type of G. kannemeyeri in the Albany Museum.

Both these latter are slightly modified to fit the skull of Gomphognathus

minor.

The prefrontal forms most of the upper margin of the orbit.

In Gomphognathus polyphagus it is considerably smaller than in

G. minor, owing to its being encroached on by the larger nasal and

lachrymal. By uniting with the postorbital it completely shuts

out the frontal from the orbital margin.

The frontal is relatively small, the two together forming

only about one-third of the interorbital space. As in all the

other Cynodonts except Bauwria and Sesamodon, the frontals

posteriorly taper away to a point between the postorbitals.

The postorbital is a fairly large bone with an external limb

which meets the jugal, forming the postorbital arch, and a posterior

process which les along the parietal. In G. polyphagus the

postorbital bar is relatively slenderer than in G. minor.

The jugal is very large. It is essentially similar to that of

Trirachodon. The inferior process is much larger and the

posterior extension much deeper. It passes backwards some

distance behind the plane of the quadrate.

The parietal is small and there is a small pineal foramen.

The squamosal is extremely large. The inner portion

articulates with the parietal. The outer and anterior portion

lies above the jugal, forming with it a powerful zygomatic arch.

Inferiorly the squamosal supports the small quadrate. The

peculiar shape of the bone can best be understood from the figures.

910 DR. R. BROOM ON THE

The occiput is very similar to that of Cynognathus, but the

limits of the various elements are less satisfactorily known. The

condyle is double, but is relatively smaller and less mammal-like

than in Cynognathus.

The palate is beautifully shown in three of the British Museum

specimens,

Text-fig. 176.

Upper view of skull of Gomphognathus minor.

The snout restored from Gomphognathus kannemeyeri.

In the anterior palatine opening there is a pair of narrow bones

showing what I suggested a good many years ago were probably

prevomers. Only one specimen is known in which they are shown.

They are evidently not parts of the premaxille, and as they are in

position exactly corresponding to the prevomers of Ornitho-

rhynchus, | am still of opinion that they correspond to the paired

vomers of most reptiles and the prevomers of Platypus and of

Miniopterus.

SKULLS OF CYNODONYT REPTILES. 911

The secondary palate is almost typically mammalian, the maxillee

and the palatines having the same relations as in the mammal,

The palatine besides forming part of the secondary palate curves

round inside of the maxilla, forming the outer wall and part of

the roof of the posterior nares. There is a posterior palatine

foramen situated exactly as in Mammals.

Text-fig. 177.

IMG XXI, XI.

Under view of skull of Gomphognathus minor.

The snout restored from Gomphognathus kannemeyeri.

The vomer is large and, as in typical Mammals, it forms the

median support of the basicranial axis from the sphenoidal region

to near the front of thenose. It forms about one-third of the roof

of the pharynx, and two small tubercles on its posterior part are

probably for the attachment of pharyngeal muscles. In the

region of the secondary palate the vomer forms for a considerable

distance a median support.

The pterygoid is of moderate size, but much smaller than in any

912 P DR. R. BROOM ON THE

other reptilian group. In front it meets the jngal and the

palatine and lies along the outer side of the vomer. It forms a

large descending pterygoid process. Posteriorly it lies against the

vomer and terminates by meeting the alisphenoid as in

Cynognathus.

Text-fig. 178.

Posterior view of skull of Gomphognathus minor.

The basioccipital, basisphenoid, and exoccipital are all

apparently very similar to those bones in Cynognathus. There

is a large foramen for the exit of nerves ix., x., xi., and xi.

Outside of this foramen, but further from the base, is another

foramen which I believe to be the fenestra ovalis, but this 1s less

certain than the nature of the larger foramen, about which there

is no doubt.

The alisphenoid in all its relations is very similar to that in

Cynognathus. At its lower end near where it meets the pterygoid

is an opening which leads into the brain-cavity, and most probably

it was by this opening that the internal carotid artery entered

‘the cranium.

The quadrate is relatively rather smaller than in Cynognathus,

but is fixed into the squamosal bone in a very similar fashion.

Tt clasps the lower margin of the bone, and posteriorly it has two

processes which fit into grooves. In the type specimen of

Gomphognathus kannemeyert, where the articulars do not fit on to

the quadrates, I thought the quadrates had been displaced, but I

am rather inclined to think it is the articulars, as the London

specimens show that the quadrate is not likely to be readily

disarticulated.

In the median section of the skull (Pl. XLVI. fig. 9) a number

of most interesting features are revealed. In the posterior

cranial region there are seen the foramina for the exit of a

number of the cranial nerves. Close to the occipital condyle

are two small foramina for nerve xii. These after passing a

short distance through the bone open into the large foramen

lacerum posticum. This large foramen is also situated well back

and doubtless transmitted also nerves ix., x., and xi. In front

SKULLS OF CYNODONT REPTILES. 913

of this foramen jugulare is the prootic bone, which appears to

have two foramina in it. The posterior I believe to be for

nerve vill., aud the anterior for vii. Between the prootic and

the large alisphenoid is seen the opening for probably both the

2nd and 3rd branches of nerve v. _ Immediately below this

foramen, the prootic sends a sharp bony process upwards, inwards,

and forwards. Probably it lay on the inner side of the Gasserian

ganglion.

The basisphenoid is a large bone, along the front of which lies

the back part of the vomer. There is no orbito-sphenoid and no

presphenoid bone.

The vomer passes from the basisphenoid to about the middle of

the hard palate. Along its dorsal surface ran the cartilaginous

cranial axis, against which the grooved upper surface of the vomer

fits. At the front of the vomer the median cartilage is ossified,

and the bone is apparently the homologue of the mammalian

mesethmoid. In the figure given of the median section the

prevomer (P.vo.) and the septomaxillary (Sma.) are largely

hypothetical.

Sesamodon and Alelinodon.

(Pl. XLVI. figs. 3, 4, & 5, and text-figs. 179, 180.)

These two allied genera, which are both unfortunately very

imperfectly known, stand at present by themselves some distance

apart from the other Cynodonts. Each genus is known only by

a single specimen, which in the case of Scsaimoden brownt 18 only

fair, and in the case of Melinodon simus ver ypoor. Still the interest

attaching to the specimens is so very great that it is necessary to

figure them as fully as possible.

Though the only known skull of Sesamodon browni is very badly

weathered and considerably crushed, it is fortunately possible to

restore the external appearance with much certainty. In fact the

only points in the external anatomy that remain in doubt are the

articular region, the middle of the occiput, and parts of the jugal,

frontal, parietal, and squamosal bones.

The premaxilla is not very satisfactorily preserved, but presents

no unusual features.

The septomaxillary is apparently fairly similar to that of

Nythosaurus, appearing on the face to a considerable extent.

The maxilla is large and resembles to some extent that of

Trirachodon, while in other respects it is nearer to Vythosaurus.

The canine is relatively smaller than in either Zrirachodon or

Nythosaurus, but is situated, as in Vythosaurus, far back from

the front of the bone. The molars, with the exception of the Ist

which is small, form a uniform series, and the maxillary bones are

much approximated in the molar region, as in Zrirachodon.

The nasal resembles on the whole that of Bauria. The nostril

is directed mainly forward and the nasal to some extent over-

hangs it. The bone is moderately broad throughout its whole

914 DR. R. BROOM ON THE

length, but is chiefly remarkable in being narrower behind than

in front. In all other Cynodonts except Lauria the nasal is, as in

Marsupials, much broader posteriorly.

Text-fig. 179.

Side view of skull of Sesamodon browni. Somewhat restored from the only known

specimen, which forms the type. Ihe whole of the preorbital portion of the

skull except the front of the premaxilla is preserved in the specimen, though the

bone in parts is weathered off, leaving only the impression. Though the teeth

are imperfect, remains or impressions of all are present, so that the full dentition

_can be restored with much certainty, the only doubt being the exact length of

“the incisors and canines. The orbit and the temporal fossa are satisfactorily

preserved, and the squamosal is fairly well preserved on the right side of the

skull. The lower border of the jugal is unknown. Both mandibles are in

position but much weathered. The horizontal ramus ts fully known, but much

of the ascending ramus is lost. As, however, the top of the coronoid process 1s

preserved in position, the greater part of the dentary can be restored with

certainty. The condylar process is badly preserved. A considerable part of

what is believed to be the angular and prokably part of the articular are

preserved. As the position of the glenoid cavity is known, the general shape of

the back of the jaw can be restored with some probability.

The lachrymal is small and completely separated from the nasal

by the prefrontal. In this, Sesamodon again agrees with Lauria

and differs from all other known Cynodonts.

The prefrontal is much larger than the lachrymal and meets the

frontal, nasal, maxilla, and lachrymal as in auria.

The frontal is almost completely lost from the specimen, but

just sufficient of the impression of the bone is left to show that it

reached the orbit as in Bauria. In this also, Sesamodon differs

from almost all other known Cynodonts.

The postorbital is fairly large but slender. It forms with a

small part of the jugal a complete postorbital arch. It only

extends backwards on the parietal a very short distance.

The parietal is completely lost except just sufficient to show

the width of the bone.

The jugal is not well preserved except in the upper part. . It

forms the whole of the lower orbital margin and part of the

SKULLS OF CYNODONT REPTILES, 915

postorbital arch. In the drawing given the lower margin is

entirely hypothetical. The posterior or zygomatic portion of the

bone is very short.

Text-fig. 180.

Upper view of skull of Sesamodon browmi, restored from the type. From the specimen

the whole of the frontals, parietals, and most of the occiput have weathered away,

so that these parts are unknown. "As, however, the prefrontals, postorbitals, and

the margins of the orbits and temporal fosse are preserved, the only points

that are left in doubt are the position of the fronto-parietal suture, whether

there is a parietal foramen, and the nature of the condyles.

The squamosal is sufficiently well preserved to show its main

features, and it is seen to be unlike that of any other Cynodont.

As in Bauria it is much smaller than in the higher types. It

articulates, as in other forms, with the parietal, but on passing

outwards it has not, asin Bawre ‘1a, a poster ior ridge. The zygomatic

portion is much Shomhen than in Bawria or any other known type,

but it is relatively fairly deep.

What appears to be the quadrate is a small flattened bone which

I have shown in the figure, but the parts are crushed and some-

what displaced, and it is impossible to speak with certainty of the

condition.

916 DR. R. BROOM ON THE

The occiput is badly preserved, but is remarkable for the great

lateral extension of the exoccipital, which passes out behind and

below the squamosal. Owing to some degree of crushing it is

difficult to be quite sure of the relations of the external end of

the exoccipital, but it certainly extends much further out than in

any other known Cynodont.

The palate, so far as preserved, differs considerably from that of

the typical Cynodonts. There is asecondary palate of the ordinary

type, but the pterygoid appears to have a larger pterygoid process

than usual. Behind the pterygoid process there isa fan-like bony

expansion which passes backwards and outwards towards the

articular region. It looks as if it might be all pterygoid, but

owing to the crushed and weathered condition of the specimen it

is impossible to be sure. Possibly, as in the typical Cynodonts, it

is part of the alisphenoid.

The structure of the lower jaw cannot be satisfactorily made

out. The dentary has a very large coronoid process ; in fact the

coronoid process is as large as the horizontal ramus. There also

is some evidence of a condylar process. The articulation is

apparently, mainly at least, formed bya rounded articular supported

by possibly an angular and surangular. There is a well-developed

splenial bone.

Pages A:

The dental formula is 1. 3, ¢. 7, m. ;

Melinodon is closely allied to Sesamodon and pretty certainly

belongs to the same family. The teeth are of the same type, but

relatively much smaller. The specimen is so imperfect that it is

impossible to make much of the skull. I have figured it as

preserved (Pl. XLVI. fig. 3).

Sesamodon resembles Bauria and differs from the other

Cynodonts in the following characters :—

Uae

1. The nostril is directed more forwards than outwards.

2. The nasal is not widened posteriorly.

3. The prefrontal is larger than the lachrymal and prevents

the lachrymal from meeting the nasal.

4. The frontal forms part of the orbital margin.

5. The postorbital arch is feeble: incomplete in Bauria.

6. The molars show no sign of cusps.

In the following characters Sesamodon comes nearer the

Mammals than any of the other known Cynodonts :—

1. An articulation for the lower jaw which permits of some

degree of antero-posterior movement.

2. The lower canine lies outside the edge of the maxilla when

the jaw is closed.

In addition to the mammalian characteristics peculiar to

Sesamodon, it combines most of those mammalian characters

seen in Bauria with most of those found in the other higher

Cynodonts.

SKULLS OF CYNODONT REPTILES. 917

Peculiarities of the Mammalian Skull, apparently derived

from a Cynodont Ancestor.

Most of the bones of the mammalian skull have their homo-

logues in the Labyrinthodont skull, but they are also to be found in

the skulls of most reptiles. There is, however, no close resemblance

between the mammalian and the batrachian bones, and in many

cases the differences in cranial structure are so great that the

gap between the mammal and any known batrachian must be

enormous. When we examine the Cotylosaurian skull, we find that

the resemblance to that of the mammal is still remote, but any

little resemblance there was in the Labyrinthodoent is here

increased, while there are many mammal-like characters not seen

in the lower type.

The Pelycosaurians of the Lower Permian are so much more

mammai-like than any of the lower forms that, notwithstanding

their remarkable specialisations, one cannot help feeling, as Cope

felt, that here were forms fairly near to the remote mammalian

ancestor.

The Therocephalians and Anomodonts of the Middle Permian

times are in essentials still more mammal-like. For the first time

we get a dentition clearly divided into incisors, canines, and

molars ; for the first time we get a lower Jaw witha dentary which

has a large coronoid process. We get a zygomatic arch formed on

the mammalian type, and we lose for the first time the quadrato-

jugal. We also get most marked mammalian characters in the

posteranial skeleton.

The Therocephalians survived into Upper Permian times, but

hitherto they have not been found in Triassic beds. In Upper

Triassic times their place was taken by the Cynodonts. Though

the gap between the Therocephalians at present known and the

Cynodonts is very considerable, the primitive Cynodont Bawria is

to some extent a connecting link.

Almost all the characters in which the Cynodont skull differs

from the Therocephalian are characters which are met with in

Mammals. Of these the most noteworthy are :—

1. Formation of a secondary palate.

2. Vomer very large, extending forward as a support to the

secondary palate.

Great reduction or complete loss of prevomers (Lauria).

Loss of the postfrontal bone.

Great reduction or loss of the pineal foramen.

Two occipital condyles.

Reduction of the quadrate.

A large alisphenoid bone instead of the homologous rod-

like ‘‘ epipterygoid” or columella cranii of the Thero-

cephalians and Anomodonts.

9. Pterygoids not extending back to the quadrates, the

posterior extension being replaced by the alisphenoids.

10. Reduction of the angular and surangular, and greater

development of the dentary.

Proc. Zoou. Soc.—1911, No. LXITI. 63

CoN See Se

918 DR. R. BROOM ON THE

The presence of this large number of mammalian characters in

the Cynodont skull, and the absence of any in either skull or

skeleton that might not have been expected in the mammalian

ancestor, make the case very strong for the mammalian ancestor

having been a C ynodont. But the evidence becomes even stronger

when we find that most of the peculiarities, even minor

peculiarities, of the mammalian skull have light thrown on them

by the condition of affairs in the Cynodont skull.

Let us consider some of the more remarkable characters of the

mammalian skull in the light of our knowledge of the Cynodont.

Premaaillary.

One of the most striking peculiarities of the mammalian skull

is that the nostrils are separated only by cartilage, so that if the

cartilage be removed the nostrils are united. In most reptiles, in

birds and amphibians the nostrils are divided by an upward and

backward process of the premaxilla, the internasal process. As it

is present in Cotylosaurs, Dromasaurs, Pelycosaurs, Therocepha-

lians, Anomodonts, and even Cynodonts, one might fancy that

here was evidence against the Cynodont ancestry. But there is

good reason to believe that the early Mammals retained the

internasal process and that it was only lost after the Mammals

were well established.

In both Ornithorhynchus and Hehidna the young animal has an

internasal process developed on the premaxilla almost exactly as

in reptiles. The fact that it is retained as a support to the

caruncle or egg-tooth in no way invalidates the conclusion that it

is the reptilian internasal process that has been retained. For

there cannot have been a time when there was acaruncle without

a support, and thus the internasal process must be as old as the

caruncle. As we may be pretty certain that the mammalian

ancestor was oviparous, we may safely conclude that the internasal

process is not a neomorph, but the reptilian structure handed on,

In Zritylodon there is an imperfect but distinct little internasal

process. The only known specimen is too imperfect to enable me

to say whether it formed a complete though slender process which

joined with the nasals. Even if it did not in the adult, it is

vather probable that it did in the very young animal, since

Tritylodon is so much more primitive than the marsupial that not

improbably it was oviparous.

Tn the skulls of young Diprotodonts (e. g. J/acropus) a rudiment

of the internasal process is usually present. And in the young

Trichosurus at birth the internasal process, as I recently pomted

out, can be traced right round in front of the nose. In the very

young marsupial, the nostrils are entirely lateral and wide apart,

and the nasal cartilages pass round in front of each, leaving a

sulcus in the middle line between tke two. The premaxillaries

send up short processes along the sulcus, but from the ends of the

processes two strands of condensed but unossified cells can be easily

SKULLS OF CYNODONT REPTILES. 919

traced round to the top of the snout. Were these tracts ossified

we would have a condition practically similar to that of the

Cynodont.

Septomaxillary.

Kitchen Parker many years ago recognised this as a distinct

membrane-bone in the Lizards and Snakes, but it is only recently

that much attention has been paid to it, chiefly as the result of the

work of one or two paleontologists in Europe and America. The

bone is not known in Labyrinthodonts and probably does not

occur in the Amphibia (the supposed septomaxillary of Xenopus

being probably not homologous). We find it, however, in the very

earliest true reptiles, and we can trace it on the one hand through

numerous members of the Diapsida, and on the other through

most of the mammal-like Reptiles on to Mammals.

A septomaxillary has been found in Pareiasaurus, Pariotichus,

and Procolophon. In these primitive genera it is mainly within

the nostrils, and probably fulfils its main function as a roof to

Jacobson’s organ.

When we come to the mammal-like Reptiles, we find it in the

Pelycosaurs still mainly within the nostril. In the Dinocephalians

(Delphinognathus, Tapinocephalus) it comes partly on to the face.

In the Dromasaurians (Galepus), it forms a very appreciable

portion of the facial wall; and in the Therocephalians (Seylaco-

saurus, Aloposaurus), it also appears pretty largely on the face.

In the Anomodonts it is absent, probably because they had lost

their organ of Jacobson, as would appear from the loss of the

prevomer.

In the Cynodonts, the septomaxillary is always present. In

the lower types it appears on the face, but in the higher forms it

is almost entirely inside the nostril.

Among Mammals a septomaxillary is known only in some of

the lower forms. In Yvitylodon it appears on the face between

the nasal and premaxillary, in much the same way as in Vytho-

saurus. In Ornithorhynchus and Echidna it would appear from

the researches of Gaupp that what used to be regarded as the

upper part of the premaxillary is really the septomaxillary. If

this be so, as seems pretty certain, then the Monotremes have the

septomaxillaries better developed than in the Cynodonts.

The only higher mammal in which there is a bone to be regarded

as probably the septomaxillary is Dasypus. Here a small bone,

which I described in 1897 asthe “ nasal-floor bone ” and suggested

might be homologous with the upper part of the premaxillary in

the Monotreme, is probably to be regarded as a rudimentary septo-

maxillary.

Vomer and Prevomers.

In 1895, and more fully in 1902, I showed that there was

reason to believe that the so-called reptilian ‘“‘ vomers” were not

homologous with the mammalian vomer, but that being formed as

splints to the paraseptal cartilages in close ae pceianion with the

63°

920 DR. R. BROOM ON THE

organs of Jacobson, they were really homologous with the bones

that unite to form the ‘‘dumb-bell bone” of Ornithorhynchus ;

and that the mammalian vomer had its homologue in the so-called

“‘narasphenoid”’ of the lower forms. As a new name was neces-

sary for the reptilian ‘‘ vomers,” I proposed the name prevomer.

In the Batrachia we find all three bones well developed, the

median true vomer or parasphenoid being especially large to

support the base of the skull. When in the earliest true Reptiles

the pterygoids came together, there was little need for the median

vomer and it became greatly reduced. In the Cotylosaurian

Diadectes the median vomer is still a fairly strong rod, but in the

later Cotylosaurians or primitive Diapsidans Pariotichus and

Procolophon the vomer is a very short pointed process. In most

later Diapsidans the vomer remains a small unimportant element.

It developes to a fair size in the Ophidia and becomes large and

much specialised in the Chelonia. The prevomers, on the other

hand, remain large in most Diapsidans, but where, as in the

Chelonians and Crocodilians, the organs of Jacobson become much

reduced or lost the prevomers likewise tend to disappear.

In the mammal-like Reptiles the vomer shows great variations.

In the Therocephalians it is small as in the primitive Diapsidans,

but with the development of a secondary palate a new function

is given to it, and it becomes large. In the Anomodonts, though

the secondary palate is only imperfectly formed, the vomer is large

and extends well forward. In the Cynodonts, where the secondary

palate is complete, the vomer is very large and extends from the

basisphenoid to near the front of the snout. The front part of

the bone corresponds so exactly in its relations to the mammalian

vomer, that it is impossible to doubt that the bones are homologous.

On the other hand, if the anterior part of the bone were lost it is

probable that every one would agree, from the relations of the

back part, that it was the homologue of the reptilian so-called

“‘parasphenoid.” In Mammals the vomer varies greatly in size.

It is relatively very large in the Cetacea, sometimes extending

from the basioccipital to the front of the rostrum, while in the

Rodentia it is often more or less rudimentary.

The prevomers are large in the Dinocephalians and Thero-

cephalians. In the Anomodonts they have completely disappeared.

In the Cynodonts, with the formation of the secondary palate

they are either greatly reduced (Gomphognathus) or quite absent

(Bauria). In Mammals the prevomers are usually absent, their

function as supports to Jacobson’s cartilages being taken by the

palatine processes of the premaxille. Im only two mammals are

they known for certain to occur as distinct bones, viz. Oriitho-

rhynchus and Miniopterus, and in both of these the pair of bones”

fuse together to form a median bone before the animal is full-

grown.

Some observations have recently been made which at first sight

appear to cast a little doubt on the homology of the parasphenoid

with the mammalian vomer.

SKULLS OF CYNODONT REPTILES. 921

Versluys has discovered what he believes to be a large para-

sphenoid in Dermochelys in addition to the vomer and in no way

connected with it. If this determination be correct, it will

probably turn out that the Chelonian vomer is after all a pair of

prevomers fused. The early development of the Chelonian vomer

has not, so far as I am aware, ever been examined, and in my paper

on the reptilian and mammalian vomerine bones I spoke very

guardedly on the subject. So far as we know, the Chelonian

vomer is always a median unpaired bone. But if it be a true

vomer, what of Versluys’ supposed parasphenoid? Fuchs has

shown that in Chelone the basisphenoid is ossified by a large

irregular exostosis on its under side, and that this exostosis bears

relations to the pterygoids very similar to those which the para-

sphenoid of Versluys does. In the light of the observations of

Fuchs, I think it must be concluded that the supposed parasphenoid

in Dermochelys is entirely a development of the basisphenoid, and

not the homologue of the parasphenoid of other reptiles.

Gaupp and Fuchs have both apparently discovered a rudimentary

ossification behind the vomer in Chelonians which they believe

to be a true parasphenoid, and Fuchs has discovered what he

believes to be a rudimentary parasphenoid in Didelphys. The

situation of these rudimentary ossifications is undoubtedly that of

the parasphenoid, but they are also in the region normally occupied

by the vomer in Mammals. When a bone which occupies one region

in an ancestor comes to occupy a somewhat different region in

a descendant through a portion of the bone becoming aborted, it

is by no means uncommon that rudimentary ossifications can be

detected in the region abandoned. The os caruncule-is undoubt-

edly the internasal process of the premaxilla in Ornithorhynchus

and Hehidna, but though it is quite detached from the pre-

maxilla, it is nevertheless a portion of the premaxilla. In the

case of the vomer, supernumerary ossifications appear to be not

uncommon both in front and behind. In Orycteropus there are

two small ossifications in front, apparently not prevomers, but

detached ossifications of the true vomer. Kitchen Parker seems

to have found them so commonly present that in some groups

he regarded them as the rule. Speaking of the condition in

Marsupials he says: “‘The main vomer is often relatively small ;

there is, nearly always, a pair of antero-lateral vomers....and

large postero-lateral, and other, or postero-medial vomers ; these

are very irregular and unsymmetrical in the young Cuscus

especially, in which I find ¢en vomerine bones.” Parker’s postero-

medial vomers are probably the ossifications regarded by Fuchs as

parasphenoids, and there seems no reason to regard them as of

any more morphological significance than the Wormian bones in

the human skull.

Alisphenoid.

Until recently the alisphenoid bone has been looked upon,

like the orbito-sphenoid, as an ossification of the cranial wall, and

922 DR. R. BROOM ON THE

according to Parker “ the alisphenoids and orbito-sphenoids appear

as chondrifications of the walls of the skull.” In studying the

development of the marsupial skull some years ago, I found that

the alisphenoid has originally nothing to do with the walls of the

skull. It first appears asa short rounded rod lying outside the

trabecula and quite independent of it or of any other skeletal

structure. In its relations it seems exactly to correspond to the

middle part of the cartilaginous bar on which the pterygoid bone

develops in Lizards and Sphenodon. In the majority of Lizards

this middle part gives rise to the epipterygoid or columella cranii.

When, asin Chameleon, the epipterygoid is rudimentary, the short

bar which forms its base is almost exactly similar in structure and

relations to the bar from which the alisphenoid developes in

Mammals. One therefore seems driven to the conclusion that the

epipterygoid and the alisphenoid are different developments of

the same element. And this conclusion seems borne out by

comparative anatomy, for we find that most Reptiles have either an

epipterygoid or an alisphenoid, but never both. In Lizards we

find an epipterygoid, but never an alisphenoid: in Snakes an

alisphenoid, but never an epipterygoid.

There seems little doubt that the epipterygoid is the early type

of development. We find, for example, in the primitive Proco-

lophon a columella cranii almost exactly like that of the lizard.

In the Therocephalians the columelle are long and slender, but

usually flattened. In the Anomodonts they are slender, but

rounded. In the Cynodonts we find no longer the columella

cranii, but in its place a broad fan-shaped alisphenoid. The

Cynodont alisphenoid further differs from the columella cranii

of the earlier forms in having the lower part well developed and

replacing the backward extension of the pterygoid. In the

mammal the alisphenoid differs from that of the Cynodont mainly

in having the 2nd and 3rd branches of nerve v. passing through

it instead of behind it.

[Vote by Kprror.—_In Dr. Broom’s memoir as presented to the

Society there followed here a discussion of the quadrate and

tympanic, illustrated by two diagrams. A recent discovery made

by Dr. Broom has considerably modified his views, and he has

asked leave to withdraw the paragraphs omitted here until he

has time to work out and present to the Society in a fuller

form the bearings of his new facts.— August 11th, 1911.]

Angular.

The angular is found in all mammal-like reptiles. It is large

in the Dinocephalians, Anomodonts, and Therocephalians, but

comparatively small in the Cynodonts. In Mammals there is a

small splint-bone on the lower side of Meckel’s cartilage which is

probably the remains of the angular. In Ornithorhynchus there

appear to be two splint-bones, one being probably the surangular.

SKULLS OF CYNODONT REPTILES. 923

In the following diagram is represented what appears to be the

genetic relationships of the principal known Cynodont genera :—

Therocephalian Ancestor.

Bauria

~~ Alurosuchus

“ie | ~G@alesaurus

Mammalhan xe

ape tor Ne

Ance stor. Sesamodon Nythosaurus

v Melinodon

Trirachodon

Cynognathus

=~ 4 va

Diademodon

Gomphognathus

List of References to the Principal Literature.

Baur, G.—On the Quadrate in the Mammalia. Q.J. Micr. Se. Vol. xxviii, 1886,

p. 169.

Broom, R.—On the Fate of the Quadrate in Mammals. Ann. Mag. N. Hist., Nov.

1890, p. 409.

— On the Occurrence of an apparent distinct Prevomer in Gomphognathus.

Journ. Anat. & Phys., Vol. xxxi, 1897, p. 277.

— On the Structure and Affinities of Udenodon. P.Z.S. 1901, Vol. ii,

p. 162.

— On the Classification of the Theriodontsand their Allies. Rep. 8. Afr. Assoc.

Ady. Se. 1903, p. 286.

—— On the Lower Jaw of a small Mammal from the Karroo Beds of Aliwal

North, South Africa. Geol. Mag., Aug. 19038, p. 345.

— On the Mammalian and Reptilian Vomerine Bones. Proc. Linn. Soc.

N.S. W., 1902, p. 545.

— Onthe Theriodonts in the Albany Museum. Rec. Alb. Mus., Vol. i, 1904, p.82.

— On the Structure of the Theriodont Mandible and on its Mode of Articu-

lation with the Skull. P.Z.S., 1904, Vol. i, p. 90.

— Preliminary Notice of some new Fossil Reptiles collected by Mr. Alfred

Brown at Aliwal North, S. Africa. Rec. Alb. Mus., Vol. i, 1905, p. 269.

(Preliminary notice of Sesamodon browni and Melinodon simus.)

924 DR. R. BROOM ON THE

Broom, R.—Reptiles of the Karroo Formation. In Rogers’ ‘ An Introd. to the

Geol. of Cape Colony.’ London, 1905, p. 228.

On some Points in the Anatomy of the Theriodont Reptile Diademodon.

P. ZS. 1905, Vol. i, p. 96.

—— Ona New Cynodont Reptile (42lurosuchus browni). Tr.S. Afr. Phil. Soc.,

Vol. xvi, p. 376.

—— Some recent Advances in South African Paleontology. Science, Vol. xxvi,

1907, p. 796.

— On the Origin of Mammals. Rep. Brit. & S. Af. Assoc. Adv. Sce., Vol. 1,

Johannesburg, 1907.

—— The Origin of the Mammal-like Reptiles. P.Z.S. 1907, p. 1047.

—— Observations on the Development of the Marsupial Skull. Proc. Linn. Soc.

N.S. W., 1909, p. 195.

—— Notice of some new South African Fossil Amphibians and Reptiles. Ann.

S. Afr. Mus., Vol. vii, 1909, p. 251.

(Description of Bawria cynops.)

A Comparison of the Permian Reptiles of North America with those of

South Africa. Bull. Am. Mus. Nat. Hist., Vol. xxviii, Art. 20, 1910, p. 197.

Doxto, L.—On the Malleus of the Lacertilia, and the Malar and Quadrate Bones

of Mammalia. Q.J.Micr. Sc. Vol. xxiii, 1883, p. 579.

Fucus, H.—Untersuchungen iiber die Entwicklung der Gehérknéchelchen, des

Squamosums und des Niefergelenkes der Siiugetiere, &c. Archiv f.

Anat. u. Phys., Anat. Abth., 1906.

—— Ueber einen Rest des Parasphenoids bei einem rezenten Siugetiere. Anat.

Anz., Bd. xxxii, 1908, p. 584.

-——— bBetrachtungen iiber die Schlafengegend am Schidel der Quadrupeda. Anat.

Anz., Bd. xxxv, 1909, p. 113.

— Ueber Knorpelbildung in Deckknochen, nebst Untersuchungen und Betracht-

ungen tiber Gehorknéchelchen, Kiefer und Kietergelenk der Wirbeltiere.

Archiv f. Anat. u. Phys., Anat. Abt., 1907.

—— Ueber das Pterygoid, Palatinum und Parasphenoid der Quadrupeden, &c.

Anat. Anz., Bd. xxxvi, 1910, p. 33.

Gapow, H.—On the Modifications of the First and Second Visceral Arches, with

especial reference to the Homologies of the Auditory Ossicles. Phil.

__ Trans. (B) Vol. clxxix, 1888, p. 451.

——- The Evolution of the Auditory Ossicles. Anat. Anz., Bd. xix, 1901, p. 396.

—— The Origin of the Mammalia. Zeitschr. f. Morph. u. Anthrop., Bd. iy,

Heft 11, 1902, p. 345.

Gaurr, E.—Die Nicht-Homologie des Unterkiefers in der Wirbeltierreihe. Ver-

handl. anat. Gesellsch. xix. Versamml. in Gent., 1905, p. 125.

— Zur Entwicklungsgeschichte und vergleichenden Morphologie des Schiadels

von Hehidna aculeata var. typica. Abdr. Semon’s Zool. Forschungsreisen

in Australien. Jena, 1908.

GreGory, W. K.—The Orders of Mammals. Pt. II. Genetic Relations of the

Mammalian Orders: with a Discussion of the Origin of the Mammalia

and of the Problem of the Auditory Ossicles. Bull. Am. Mus. Nat.

Hist., Vol. xxvii, 1910, p. 105.

Kinesiey, J. S.—The Ossicula Auditus. Tufts College Studies, No. 6, 1900, p. 203.

The Origin of the Mammals. Science, n. s., Vol. xiv, 1901, p. 193.

Ksniipere, K.—Beitrage zur Entwickelungsgeschichte des Kiefergelenkes. Ge-

genbaur’s Morph. Jahrb., Bd. xxx, 1904, p. 159.

Lusoscu, W.—Ueber das Kietergelenk der Monotremen. Jenaische Zeitsch. f.

Naturw., Bd. xli, 1906, p. 549.

Osporn, H. F.—The Triassic Mammals, Dromatheriwn and Microconodon. Proc.

Am. Phil. Soe., Vol. xxiv, 1887, pp. 109-111,

The Origin of the Mammalia. Am. Nat., Vol. xxxu, 1898, p. 309.

Remarks at the Discussion on the Origin of Mammals. Proc. Internat.

Cong. Zool. Cambridge, 1898, p. 415.

The Origin of Mammals. Am. Journ. Se. (4), Vol. vii, 1899, p. 92.

Origin of the Mammalia, III. Occipital Condyles of Reptilian Tripar-

tite Type. Am. Nat., Vol. xxxiv, 1900, p. 943.

The Reptilian Subclasses Diapsida and Synapsida and the Early History of

the Diaptosauria. Mem. Am. Mus. Nat. Hist., Vol.i, pt. 8, 1903, p. 449.

Owen, R.—On some Reptilian Remains from South Atrica. Edinb. New Phil.

Journ., Vol. x, 1859, p. 289; also Quart. Journ. Geol. Soc. Lond., Vol. xvi,

1860, p. 45.

— Palexontology, or a Systematic Summary of Extinct Animals and their

Geological Relations. Edinburgh, 1860; 2nd Hd. 1861.

— Descriptive and Illustrative Catalogue of the Fossil Reptilia of South

Africa in the Collection of the British Museum. London, 1876.

Bia

SKULLS OF CYNODONT REPTILES. 925

Owen, R.—On the Skull and Dentition of a Triassic Mammal (Tritylodon longevus

Owen) from South Africa. Quart. Journ. Geol. Soc. Lond., Vol. xl,

1884, p. 146.

— On the Skull and Dentition of a Triassic Saurian (Galesaurus planiceps

Ow.). Quart. Journ. Geol. Soc. Lond., Vol. xliii, 1887, p. 1.

SrErLey, H.G.—Researches on the Structure, Organisation, and Classification of

the Fossil Reptilia. WI. On the Anomodont Reptilia and their Allies.

Phil. Trans., Vol. clxxx (1889) B, p. 215.

——. idem IX. Sect. 1. On the Therosuchia. Phil. Trans., Vol. clxxxv (1894)

5 DP. 987.

— Idem. IX. Sect. 2. The Reputed Mammals from the Karroo Formation

of Cape Colony.—Part IX. Sect. 3. On Diademodon. Phil. Trans.,

Vol. elxxxv (1894) B, p. 1019.

—. idem IX. Sect. 4. On the Gemphodontia. Phil. Trans., Vol. clxxxvi (1895)

5 Do Ue

—— Idem. IX. Sect. 5. On the Skeleton in New Cynodontia frem the Karroo

Rocks. Phil. Trans., Vol. clxxxvi (1895) B, p. 59.

— On the Dentition of the Palate in the South African Fossil Reptile Genus

Cynognathus. Geol. Mag., 1908, p. 486.

Tuyne, F. W.—Squamosal Bone in Tetrapodous Vertebrata. Proc. Bost. Soc. Nat.

Hist., Vol. xxxii, No. 11, p. 387, 1906.

Verstuys, J.—Ein grosses Parasphenoid bei Dermochelys coriacea Linn. Zool.

Jahrb., Bd. xxviii, Anat., 1909, p. 283.

Bemerkungen zum Parasphenoid von Dermochelys. Anat. Anz., Bd. xxxvi,

1910, p. 487.

WILLISTON, Sega Temporal Arches of the Reptilia. Biol. Bull., Vol. vii, 1904,

p. 175.

Witson, J. T.—On the Skeleton of the Snout of the Mammary Foetus of Mono-

tremes. Proc. Linn. Soc. N.S.W., 1901, pt. 4, p. 717.

Woopwarp, A. S.—Outlines of Vertebrate Palaeontology for Students of Zoo-

logy. Cambridge, 1898.

Explanation of lettering in the Plate and Text-figures.

Ang. Angular; Art. Articular; Aud.gr. Auditory groove; B.O. Basioccipital ;

B.S. Basisphenoid; Dent. Dentary; EH.O. Exoccipital; Ew.st. Extrastapedial ;

Fr. Frontal; Ju. Jugal; I.P. Interparietal; Ia. Lachrymal; Mail. Malleus ;

Men. Meniscus; Mx. Maxilla; Na. Nasal; O.O. Opisthotic; Pa. Parietal ;

Pal, Palatine; Pmx. Premaxilla; Po.O. Postorbital; Pr... Prefrontal; Pt.

Pterygoid; P.Vo. Prevomer; Qu. Quadrate; S.Ang. Surangular; Sma. Septo-

maxillary ; S.O. Supraoccipital ; Sq. Squamosal; Tym. Tympanic; Vo. Vomer.

EXPLANATION OF PLATE XLVI.

. Side view of the cranial wall of Cynognathus berryi. Half nat. size.

. Occiput of Cynognathus berryi. Half nat. size. :

. Side view ot the skull of Melinodon simus. Nat.size. This represents the

type and only known specimen. It is so badly crushed that it is

impossible to restore it with any confidence. Six molars of the right side

are preserved, and the axes of two molar series make with each other an

angle of about 60°. The frontal region is certainly narrow, and as the teeth

are very similar to those of Sesamodon it is probable that the skull is also

somewhat similar, but probably the snout is shorter in Melinodon and the

skull relatively broader.

. The molar teeth of the lett side of Sesamodon browni as preserved. X 1s.

. Base of the skull of Sesamodon browni as preserved. X 15.

. Quadrate and part of the squamosal of Bawria cynops, as seen from the

front. Nearly twice nat. size.

. Base of skull of Bawria cynops, viewed partly from the side to show the

deep keel of the basisphenoid.

. Occiput and base of skull of Bawria cynops, viewed from behind and partly

from below. Slightly restored. About two-thirds nat. size.

. Median section of skull of Diademodon. All parts in unbroken line are

from the specimen in the South African Museum. The prevomer and septo-

maxillary as restored are founded partly on the British Museum specimen

ot Gomphognathus and are partly hypothetical, the inner part of neither

bone bemg known in any specimen. About half nat. size.

(SU Cl

Wey (2) NN Qoe

926 DR. A. HOPEWELL-SMITH AND DR. H. W. MARETT TIMS ON

39. Tooth-germs in the Wallaby AMacropus billardier.

By A. Hoprwett-Smirn, L.R.C.P., M.R.C.S., and

H. W. Marerr Tims, M.A., M.D., F.1.8., F.Z.S.,

King’s College, Cambridge.

[Received and Read May 23, 1911. |

(Plate XLVII.* and Text-figures 181-189.)

The dentition of the Marsupials offers, in some respects, points

of greater interest than does that of other mammals. Though

much has been already written upon this subject, there are still

problems awaiting solution.

The complete history of the tooth-genesis has been worked out

in few members of this class, due no doubt to the difficulty of

obtaining a sufficiently complete series of embryos of any one

species. Any contribution, therefore, however small, may become

of importance as forming a link in completing the chain of

evidence.

The material upon which these observations are based consisted

of three embryos of Macropus billardieri, obtained from the Seven

Sisters Islands by Mr. Brooke Nicholls of Melbourne, to whom

we are greatly indebted for his kindness in sending them for

examination. These embryos give the following measurements :—

Space | STace | Space

1 oe ee pase TI.

mm. mm. mm.

Tip of snout to occiput (circumferentially) ............) 21 | 85 38

3 35 5 (Gnistrare nteline) eeeeneseeeee 15 27 27

Tip of snout to root of tail (circumferentially) .........) 64 104 110

Vertex of head _,, 3 (amstraielit line) =... 35 | 66 52

CUESTA Ot tack not oly Mian HEN Sabai Rae blee Sabie he Alan THe: kp BL Dif

Total len¢th from tip of snout to tip of tail (cireum- |

ferential.) cetcaa cee aetna cae eaia eect nian 135 137

Serial sections were cut in a vertical transverse direction

though the skulls of Stages I and II and stained with borax

carmine. The jaws of Stage III, which gave measurements

approximately equal to those of Stage IT and presumably therefore

of an embryo of about the same age, were dissected and clarified

in oil of cloves after the method recommended by Huxley, to

show the relative positions of the calcified teeth.

The most detailed accounts of the development of the teeth in

Diprotodont Marsupials are those given by M. F. Woodward (12)

* For explanation of the Plate see p. 942.

Im, GS, US, Il, DAHVAUL.

TOOTH- GERMS IN MACROPUS BILLARDIERI.

TOOTH-GERMS IN A WALLABY. 927

and by Deppendorf (1); the latter, however, does not appear

to have examined any specimen of MJacropus. Woodward has

furnished the results of his examination of 14 embryos belonging

to four species, viz. I. giganteus, M. eugenit, M. bennetti, and

M. brachyurus. From the measurements given by him, allowing

for the differences in size of the adult animals of the different

species, our Stage I would appear to be considerably younger than

any of the embryos which he examined. Before discussing the

general considerations arising from our observations, we will

proceed to a description of the conditions found in our material.

Srace I.

Upper Jaw.—At the anterior extremity of the upper jaw the

oral epithelium sinks into the substance of the jaw, forming

a broad triangular cellular mass the outline of which is very

irregular. There are projections of some size extending laterally

into the premaxillary region. None of these would, by themselves,

suggest tooth vestiges, but the possibility of their being of such a

nature is not entir ely ‘negatived because, as will be shown in the

sequel, similar but more definite structures are to be seen in the

lower jaw. The evidence here is, however, too problematical to

permit of definite conclusions being drawn.

Text-fig. 181.

> e e \

e 2 * @ee ‘ n Nee

89 oS z

ana 733 s0% Fy | ee a q =

ee ROR ee oe

= — Ge ~ = aoe

een OY =~ Neg i) me

Capaty Sen 9*00epccecgege oso

= = ~ = os = = =—_— «<>

Be eee een or!

ie ae oy ee ie

eee fi dew ey) ee Oral

i —~ eee sO 2 -

Fea ee ee arene = epithelium

ee a See heme pens ae

55 =aa

Section showing the tooth-band (X) passing horizontally inwards and forming

the anterior part of the tooth-germ B.

On either side of the middle line close to the nasal septum is a

minute slightly calcified tooth-germ with a dental papilla of

928 DR. A. HOPEWELL-SMITH AND DR. H. W. MARETT TIMS ON

definite shape. This germ will for the present be indicated by

the letter A.

From a point just external to the point of connection of the

neck of the enamel organ of 4 to the oral epithelium, the tooth-

band passes horizontally inwards towards the middle line

(text-fig. 181)*. This rapidly increases in size, forming on section

a conical mass of cells which soon loses all connection with the

oral epithelium. From the histological characters of the cells

and from the fact that in the more posterior sections a portion

of the stellate reticulum of the enamel organ is visible, this

structure must be regarded as a second tooth-germ (£).

It is considerably larger than the preceding germ 4, though

there is not the same amount of differentiation into the more

typical dental tissues.

Text-fig. 182.

Section through the tooth-germ B at its deeper end.

The independent connection of the enamel organ 4 with the

oral epithelium shows that it is not to be regarded as the

morphological successor to A.

To neither A nor Bare there either palatal or labial down-

* Owing to the difficulty of accurate orientation, the anterior sections of a jaw cut

in the vertical-transverse direction will be in a plane parallel with the outer surface

of the jaw. This has been borne in mind in describing the relative positions of the

various structures.

TOOTH-GERMS IN A WALLABY. 929

growths which could serve as an indication as to which dentition

the germs should be referred.

As this tooth is traced backwards it assumes a peculiar shape on

section. Its inner margin is sharp, the outer one rounded, while

on the dorsal surface is a relatively deep fissure (text-fig. 182).

The peculiarity of the adult tooth is thus early indicated.

As B disappears from the sections a new tooth-germ (C) makes

its appearance. Though minute it has a fully formed enamel

organ and dental papilla, but without any trace of calcification.

This germ has a very superficial position in the jaw.

Closely following upon Cis a large uncalcified tooth (D) lying

to the palatal aspect of and slightly posterior to the preceding

tooth-germ. Connected with the neck of the enamel organ is a

minute predecessor slightly invaginated by a rudimentary dental

papilla (text-fig. 183).

Text-fig. 183.

<aye 088 One

Wa Noe x

aay aye,

Section showing the tooth-germ D with its predecessor lying to the labial side

of the neck-band.

Yet another germ (/’) of some size soon makes its appearance,

occupying a superficial position in the jaw. There is a slight

bulging of. the dental lamina to the labial side of the neck of this

enamel organ, which may or may not indicate an abortive attempt

at the formation of a predecessor.

The dental lamina from this point is continued backwards for

some distance as a distinct band without showing any definite

930 DR. A. HOPEWELL-SMITH AND DR. H. W. MARETY TIMS ON

trace of a tooth-germ. After an interval, another tooth (//) of

considerable size appears, lasting through three slides. It has

a dental papilla with blunted apex. As yet there is no definite

evidence of calcification. There are neither palatal nor labial

downgrowths of the dental lamina. ‘This tooth we identify as

the first of the cheek series.

The tooth-band continues backwards in a well-marked condition

giving evidence of a distinct swelling which in Stage II has

developed into an undoubted tooth-germ. Without a knowledge

of the subsequent history of this swelling, one would have

hesitated to attach importance to it.

Posteriorly to this is the large enamel organ of Stage I from the

neck of which springs a definite palatal downgrowth of the dental

lamina. The shape of the dental papilla is definitely molariform,

the outer cusp already attaining to a higher level than the inner.

One more enamel organ (/), the most posterior in the jaw

at this stage, is to be seen following closely upon the preceding

one. It has but reached the flask-shaped stage.

Thus it is seen that there are representatives of eight teeth in the

upper jaw at this stage which, by comparison with those present

in Stage II, we identify as incisors 1, 2, 3, 4 and 6, and premolars

1,3 and4. The reasons which have led us to arrive at this con-

clusion will be detailed subsequently.

Lower Jaw.—Near to the mandibular symphysis is a well-

defined involution of the oral epithelium into the subjacent

mesoblastic tissues. The cells, both peripheral and central,

of this tubular downgrowth have precisely the same characters

as those covering the alveolar margin. From the appearance of

these cells and from the inclination of the downgrowth (down-

wards and towards the middle line) being the same as that of

undoubted tooth-germs situated more posteriorly in the jaw, the

opinion that it is a vestigial tooth-germ (a) seems to be justified.

Very closely following upon this structure and placed at a deeper

level in the jaw, is a small well-calcified tooth (6) showing both

dentine and enamel. The appearance of this tooth suggests

degeneration, and it may safely be asserted that this tooth, though

calcified, is not the one which will ultimately become the functional

incisor.

The difficulty in identifying the exact relationships of tooth-

germs close to the symphysis is always great, due in part to a

certain amount of crowding, accentuated in the Macropodide by

the very large size of the functional incisor and in part to the

difficulty of correct orientation. We at first thought that these

two structures stood to each other in the morphological relation-

ship of predecessor and successor. From a careful consideration

of all the facts we believe that this is not the case, but that they

are the representatives of two separate teeth and that the close

approximation to each other is due to the crowding caused by the

large functional incisor. The appearance of @ suggests a tooth

TOOTH-GERMS IN A WALLABY. 931

which is attempting but failing to develope; it has not the

“concentric ” appearance of a vestigial tooth-germ such as one

so often sees as the last evidence of existence of a predecessor to a

calcified tooth.

Rapidly succeeding the calcified tooth 6 is a well-defined flask-

shaped dental rudiment lying in a very superficial position in the

jaw. This, though close to, is quite independent of any of the

other tooth-germs here present and will in the meanwhile be

referred to by the letter c (text-fig. 184).

Text-fig. 184.

- oe

° 7 OC = He

— Ze. mae 4 2 ose,

Sue Wig > oe oyguant™

23TH 2 @ o%7 ye 0)

= 12 2,2 «© = Ww

as — » 4,009 9a68° ¢

sis ea . ?

= >= POS GSS °

1 oF G wes, 1 - ,

1 TNT GMRRR SS

— ? Le pe O &., 0-7

oe 5 7s eee, 9 0

roy ’ aay, er. - yen

r} eee 1 00 te Des! NDbibin D

A eo? oe 0 ’ Be Ps

’ 6 .

ast, ovoy. ; 0%

Section showing flask-shaped rudiment of tooth-germ c.

At this point the dental lamina runs horizontally inwards, as

in the upper jaw. It forms a thickened band of cells lying some

distance below the tooth-germ c. Tracing this band backwards

it quickly becomes connected with the upper part of the enamel

organ of a large tooth (d) in which the dental tissues are differen-

tiated and which extends through several slides. In relation to

the neck of this enamel organ is a lingually situated downgrowth

of the dental lamina indicating a potential successor (ds) to the

tooth d. From this poimt the tooth-band vanishes for some

distance (text-fig. 185).

As the tooth itself dies out the dental lamina once more comes

into prominence and gives rise to an enlargement indicating in our

DR. A. HOPEWELL-SMITH AND DR. H. W. MARETY TIMS ON

932

Text-fig. 185.

end of tooth-germ ¢ and commencement of enamel

Section through posterior

an of d and rudiment of a successor to d.

org

Text-fig. 186.

Section showing the enlarged downgrowth of the dental lamina indicating the

vestige of the tooth-germ e.

TOOTH-GERMS IN A WALLABY. 933

opinion a tooth vestige (e). It lies close beneath the alveolar

margin (text-fig. 186).

Then occurs a further interval through which the tooth-band

persists without giving rise to any further dental enlargements.

Three further teeth (h, 7, and 7) of premolariform pattern, follow

in the cheek region. Of these the middle one is considerably

the most extensive, the posterior one being quite small. To the

two anterior teeth there are indefinite indications of lingual

downgrowths of the dental lamina.

As in the upper jaw there are indications of eight teeth, five

antemolars and three maxillary teeth,

A general impression of the number and relative positions of

the dental structures in both jaws may perhaps be best realised

by the following diagram, which has been drawn to scale in the

horizontal direction.

Text-fig. 187.

Diagram showing number and relative positions of the tooth-germs in the jaws

of Stage I. Calcified germs more heavily outlined.

Srace IT.

The difference in size between this embryo and that of Stage I

is very considerable, and as a consequence a number of additional

teeth have appeared in both Jaws and those which were present

in the earlier stage have developed very materially. It is un-

fortunate that we have not had at our disposal an embryo of

Proc. Zoou. Soc.—1911, No. LXIV. 64

934 DR. A. HOPEWELL-SMITH AND DR. H. W. MARETT TIMS ON

intermediate size, as this would have rendered the precise

interpretation a matter of less difficulty and of greater certainty.

Without going into details we will summarise as briefly as

possible the conditions found in the later stage.

Upper Jaw.—Anteriorly is a small calcified tooth (4’) oceupying

a position similar to the tooth found in the previous stage and is

undoubtedly the same tooth still persisting. It shows no signs of

being erupted, neither has it undergone any further development.

That it will ultimately become absorbed without attaining fune-

tional activity seems to be certain.

Behind this hes a large incisiform tooth already heavily

calcified. This tooth (b') is the largest of the anterior series.

All traces of the tooth-band between dA’ and 45’ have become

lost, and several sections intervene between the disappearance

of A’ and the commencement of the enamel organ of B. Thus

we are of the decided opinion that these are two morphologically

distinct teeth, a point to which reference will be made later.

This conclusion is the same as that at which we arrived in Stage I.

C" is still a small unealeified tooth lying superficially, and

appears to have been pushed out of the serial line towards the

outer aspect of the jaw.

D' is very similar to 5, but perhaps not quite so large.

Posteriorly to )’ is a small calcified tooth (4’) of irregular shape

and situated close to the alveolar margin of the jaw. ‘This tooth

is obviously vestigial and will never become functionally active.

According to our identification this tooth is unrepresented in the

earlier stage.

After an interval another large incisiform tooth is seen (2”).

It is neither so large nor so heavily calcified as B’ or D’.

G' is a tooth of some size showing but traces of commencing

calcification. From its relation to the premaxillo-maxillary

suture we regard this tooth as the canine. It is a tooth in an

early stage of development which has appeared since the age of

Stage I. There is a ‘“‘ concentric epithelial body” which is to be

regarded as the last trace of a vestigial predecessor.

The first of the true maxillary teeth is elongated and of

premolariform type (H’) without any indications of a predecessor

or successor. The third tooth (/’) is like unto it but of much

larger size, extending through many more sections.

Between these two calcified teeth is a very deeply placed tooth-

germ with a neck of remarkable length (/)) (Pl. XLVII. fig. 1).

‘here is no evidence of calcification. This is evidently the

second maxillary tooth which, crowded between the large calcified

first and third, has been forced into the abnormally deep situation,

besides being pushed from the linear series towards the palatal

side of the jaw.

The fourth maxillary tooth (A’) is large and in it calcification

is but just commencing. From the neck of the enamel organ

springs a marked labial downgrowth of the dental lamina, con-

nected with which is a ‘concentric epithelial body,” the vestige

TOOTH-GERMS IN A WALLABY. 935

of a predecessor (Pl. XLVIT. figs. 2. & 3). ‘The importance of

this will be discussed later. The last tooth of the series (Z’) is

uncalcified and at an earlier stage of development than the

maxillary teeth in front of it. It lies much nearer to the alveolar

margin than do the others, and there is this further point of

interest, that the anterior end of this tooth overlaps superficially

the posterior end of the tooth in front, which is distinctly not the

case with the more anteriorly situated teeth.

According to our identification we recognise at this stage six

incisors, one canine and five maxillary teeth.

Lower Jaw.—The difficulty of interpreting the appearances in the

region of the mandibular symphysis is even greater than in the

earlier stage. The enormous development of the lower functional

incisor, extending as it does through thirty slides*, has disarranged

all the parts, more particularly anteriorly where the tip of the

tooth becomes more superficial and where it now reaches well in

front of the point at which its young enamel organ connected with

the oral epithelium.

Text-fig. 188.

Section showing involution of the oral epithelium

at the mandibular symphysis.

At the symphysis the oral epithelium is turned inwards in the

middle line so as to partially cover the opposing ends of the

mandibular rami. At this point there is a definite involution

of the epithelium into the subjacent tissues. The section being

cut at this point in a plane parallel with the anterior surface, the

* A rough estimate may be obtained by stating that a slide carries an average

of twelve sections, each section being 745 mm. in thickness.

: 64*

936 DR. A. HOPEWELL-SMITH AND DR. H. W. MARETT TIMS ON

direction of this digitiform involution is inwards and _ slightly

upwards. ‘There is an involution on either side of the symphysis

and they are quite symmetrical. The position makes the inter-

pretation somewhat doubtful, otherwise we should have but little

doubt in ranking them as tooth rudiments. Regarding the origin,

connection, and histological characters as ef more importance

than mere position, we shall regard it as such and provisionally

indicate it by the letter a’ (text-fig. 188).

Close to the middle line, but on the alveolar margin of the

mandible, is a more globular involution containing cells commenc-

ing to be arranged as a concentrical epithelial body such as several

authors now recognise as being tooth vestiges. This will be

referred to as 6’. This also is a bilaterally symmetrical structure

and extends through several sections. Immediately above this is

a fissure in the oral epithelium containing some deeply stained

fibrous-looking material. The appearance suggests the possibility

of this marking the situation where the minute calcified tooth,

present in the jaw in this position 1m the earlier stage, may have

been erupted. There is otherwise no trace of the calcified vestige.

Text-fig. 189.

Showing the relative positions of the teeth in Stage IT.

Calcified teeth more heavily outlined.

A broad bulbous involution of the oral epithelium is seen close

to the middle line (c’) which soon loses its connection with the

TOOTH-GERMS IN A WALLABY. 937

alveolar epithelium, remaining as an isolated spherical mass.

It probably is the same structure described in Stage I, and is

naturally of larger size.

Lying ata deeper level but entirely independent of it, is the

anterior extremity of the large calcified functional incisor. That

these two structures have no morphological relationship with each

other is made clear, not only by their relative position but also

by the fact that the connection of the enamel organ with the

surface is definitely visible in a section farther back, together

with a “ concentric” vestige of a predecessor and an indication of

a potential successor. This large functional incisor we shall

indicate as d’,

Then follows an interval of thirteen slides without any trace of

teeth, and throughout the greater part of this interval the dental

lamina is scarcely visible. At length a definite “concentric

epithelial mass” occurs embedded in the dental lamina. It is im-

possible to say whether this is the vestige of a canine or of an outer

incisor. From its position relative to that of the opposing teeth, it

is probably the vestige of the latter tooth. The point in favour of

its being a vestigial canine is its propinquity to the first true post-

canine tooth. However, it is a point of quite minor importance.

The description already given of the true maxillary teeth

applies almost equally well to the corresponding structures in the

lower jaw. The only two points to which attention may be drawn

are (i) the larger size and definite calcification of the 4th tooth of

the lower series; and (ii) the absence of a definite ‘“ concentric”

predecessor to the same tooth, though there is a very minute

structure which may indicate its remains, the position in relation

to the tooth itself being the correct one. That it should have

disappeared is what would be expected from the greater size and

calcification of the successional tooth.

Jil, Hisronoey.

There are certain histological peculiarities revealed by an

examination of these jaws. They are briefly as follows :—

(i) The oral epithelium along the alveolar margins is very thick

and heaped up to an unusual extent. In no other mammal that

we have examined have we ever seen this character carried to such

excess. This is the ‘‘Zahnwall” of the German writers. It is

said to be a marked character in the Ungulates, and in our ex-

perience it is certainly not of common occurrence among manimalia

generally. Of course no morphological significance is implied

by the comparison.

(ii) The somewhat unusual compactness of the stellate reticulum

and the definiteness of the stratum intermedium of the enamel

organ.

(iii) The precocious developinent of the enamel which seems

to calcify as soon as the dentine, and even in some cases to

938 DR. A. HOPEWELL-SMITH AND DR. H. W. MARELT TIMS ON

precede it. Spherules of calco-globulin are clearly seen in many

of the ameloblasts.

(iv) The abundant evidence of blood-vessels within the enamel

organ is a very striking feature (Pl. XLVII. fig. 5). The fact, of

the vascularity of the enamel organ is one upon which there

are contradictory statements. Wedl, Magitot, Legros, Sudduth

and Paul deny the existence of blood-vessels in this situation.

On the other hand, Prof. Poulton and the late Prof. G. B. Howes

have mentioned their presence in the enamel organs of the Rat.

In Macropus there is no doubt whatever. They can be seen

entering the enamel organ apparently at more than one point on

the surface, and are often of sufficient size to clearly recognise the

blood corpuscles within them. They can be traced to a point about

midway between the outer and inner enamel epithelium, but we

have not seen them proceeding farther, neither have we found them

in the stratum intermedium, the position in which they were

described by the authors above mentioned.

The fact of the vascularity of the enamel organ is one of im-

portance in connection with the various theories held as to the

processes of the calcification and enamel formation and of the

functions of the stellate reticulum. It is extr emely probable that

the unusual vascularity of the enamel organs in this animal is

correlated with the precocious development of the enamel, to which

reference has just been made.

(v) Evidence of the fusion of enamel organs. There appears to

be some evidence of the possibility of the occurrence of such fusion

though the evidence is not decisive. Pl. XLVII. fig. 4, shows the

subdivision of an enamel organ into two parts by an epithelial

septum passing from the outer enamel epithelium to the inner

where the latter lies over the apex of the dentinal papilla. This

occurs in more than one cheek-tooth and may be seen on both sides.

We have never met with anything of the kind before. ‘That this

septum is not a small capillary running directly across the stellate

reticulum we are assured. Wecan offer no other suggestion than

that it is a double enamel organ taking part in the formation of a

single tooth.

This is a point of considerable morphological importance as

bearing upon the question of the evolution of the molar patterns.

Fusion is presupposed to have occurred by all writers who believe

in any form of concrescence, but little satistactory ev idence has

been forthcoming in the mammalia and little weight can be given

to the evidence derived from the disintegration ‘of the Cetacean

molars.

Amongst the lower vertebrates the evidences of fusion seem to

be conclusive. In one family of upper Paleozoic Sharks, the

Cochliodontide, there is a fusion into plates not only of adjoining

teeth of one series, but also of successional teeth of several series.

Semon (4) has also shown that a fusion of individual cusps takes

place in Ceratodus, and Rose has seen the cusps in the process of

fusion in the teeth of achameleon (3). Woodward (/oe, cit.) speaks

TOOTH-GERMS IN A WALLABY. 939

of asupernumerary incisor in the upper jaw of anadult Perameles

in the teaching collection of the Royal College of Science in which

there are “ indications of at least three fangs, and is obviously a

fusion of (these) teeth.” The possibility of tooth fusion in the

mammalia is therefore by no means improbable. The matiter

must remain unsettled until more satisfactory evidence is forth-

coming, but the condition figured would seem to point in that

direction and we can only repeat that we can suggest no other

explanation.

LV. GENERAL CONSIDERATIONS.

_ A. Serial homologies of the Teeth.

Upper Incisors.—Assuming our identification of the teeth to be

correct, we have six incisors present in the upper jaw, of which

the 5th is unrepresented in the earlier stage and has attained but

a slight degree of development in Stage IJ. This excessive

number of upper incisors confirms the original statement of M. F.

Woodward (11), as the result of his observations on Petrogale

penicillata and Macropus giganteus. That author is obviously

aware of the difficulty in accepting so large a number, for he wr ites

‘(p. 465): “The discovery of six pairs of incisors, although an abso-

lute fact, is in many respects an unfortunate one, as we know

of no adult mammal with so many, and even amongst Reptiles,

many Lizards and Crocodiles have the number of teeth in each

premaxilla restricted to five.” He then proceeds to discuss various

possible explanations such as the late calcification of the functional

incisors in Jacropus and the greater space in the premaxillary

region in Petrogale as the possible determining causes. A reference

to our text-figures 187 & 189 (pp. 933 & 936) will show that

Woodward has possibly found the correct explanation, both

factors appearing to be present in our specimens.

The difficulty is, however, not fully stated by a mere reference

to recent forms, for Osborn (2) in his paper, “On the Structure

and Classification of Mesozoic Mammalia,” considers that the

dental formula of the primitive heterodont mammal should be

taken as 1.4, c. 1, pm. 4, m. 8.

In a paper published three years later (13) Woodward appears

to have modified his former opinion, for in spite of having pre-

viously spoken of the presence of six upper incisors as ‘fan Alcolite

fact,” he writes “ Tam now disposed to regard the three minute

teeth which I described in the upper jaw of several genera

belonging to this family” (¢.e. Macropodide) “as the premilk

predecessors of the three functional incisors.” He gives no reasons

for the alteration in his opinion, as in his previous paper he dis-

cusses and discards the interpretation which he appears later to

have adopted. He further makes no reference to the alteration

in the identification of the homologies of the functional incisors

which such a change of opinion would involve.

We have naturally therefore paid particular attention to these

940 DR. A. HOPEWELL-SMITH AND DR. H. W. MARETT TIMS ON

‘views, and we agree with Woodward’s first statement that the

presence of six upper incisors, in spite of all the difficulties which

it involves, is ‘‘an absolute fact.” The one point upon which we

join issue with Woodward is his view of the homology of the first

functional incisor which he regards as being i. 1, whilst we hold

that it is i. 2. The small calcified vestige in both our stages

is clearly anterior to and independent of the first large tooth

which is obviously the first functional incisor. Our disagree-

ment is due to differences of interpretation of the conditions

present in a very puzzling region, 7%. e. close to the median

symphysis; and though we hesitate to put ourselves in opposition

to so accurate and experienced an observer, we nevertheless are

compelled to adhere to the opinions already stated. Weare, how-

ever, quite in accord with him in regarding the outer functional

incisors as being the 4th and 6th, the 3rd and 5th disappearing.

We, like Woodward, are therefore at variance with the opinion

of Oldfield Thomas (5), who regards the missing incisors as being

the outermost ones of the series. Though Deppendorf (1) does

not appear to have examined the jaw of MJacropus, he has made

interesting observations on the tooth-genesis in many other mar-

supials. In #pyprymnus he identifies the three functional incisors

as the 2nd, 3rd, and 5th. Thus he agrees with us in the identifi-

cation of the 1st functional teeth, but differs from both Woodward

and ourselves in the identification of the two outermost functional

premaxillary teeth. It is unfortunate that there should thus be

four different versions given on this subject. Possibly the con-

ditions described by Deppendorf may be really different and not

merely a difference in interpretation, as he carried out his

researches in other genera, in which the relative proportions of

the bone may be somewhat different and thus produce different

results.

Lower Incisors—In the lower jaw we believe there are evidences

of five ante-molar teeth. Whether the most posterior one is the

vestige of a canine or of an outer incisor, it is impossible to deter-

mine, Of theremaining four the first three are vestigial, so that

according to our interpretation the functional incisor of the adult

‘is i. 4. This statement is at variance with Woodward’s conclusion

as to the homology of this tooth which he regards as i.2. In his

illustration he figures two vestiges which obviously correspond

with our b' and ¢’, his first incisor in Petrogale being a minute

calcified tooth just as is the germ 6’. The morphological 3rd incisor

of Petrogale is “ more vestigial than the first,” just as c’ is more

vestigial than 6’.

The discrepancies between our interpretations probably depend

on the following facts. (1) Woodward seems to have examined the

jaw on one side only, and to do so would have divided it through at

the symphysis. In doing so he would most certainly destroy the

minute vestige a which we have found lying practicaily within

the symphysis. (2) He regards the posterior vestige as being

ete as to the large functional incisor, whereas we regard it as

eing anterior. We both agree in stating that it lies close to the

TOOTH-GERMS IN A WALLABY. 941

tip of the large tooth, and there is no doubt that the exact mor-

phological localisation of the vestige in relation to the large incisor

is a matter of difficulty. On the whole, therefore, we are in closer

agreement with Woodward's conclusions than with those of any

other writer.

The close approximation which we have found between the

representatives of these four anterior incisors is of considerable

interest when we remember the condition of the lower incisors in

Didelphys.

Canines.—These teeth call for no further observations.

Maxillary teeth—The fourth true maxillary tooth of the upper

jaw, as has already been stated, has a well-marked ‘ concentric

epithelial body” lying superficially to it. The importance of

these structures as representing the last trace of a tooth vestige

has been frequently insisted upon by one of us (H. W.M. T.)

(7,8), and the more recent researches of Wilson & Hill (10) seem

to have placed the matter beyond doubt. So long as the present

distinction as to whether or not there are predecessors is the cri-

terion of distinction between premolars and molars, confusion is

bound to result. Embryology often reveals the presence of these

vestiges which breaks down this conception of a true molar tooth.

It would in our opinion be better to call all post-canine teeth

maxillary teeth. Until such a suggestion is adopted we must

regard the fourth tooth of the series as a premolar (pm. 4).

The relationship of this tooth to ae one immediately behind it

suggests that the fifth tooth, 7. e. m’, is the deciduous tooth which

is replaced by pm’, a suggestion strongly urged by one of us in a

previous paper (8).

_ It seems to be tolerably certain that the 2nd maxillary tooth

will not be able to develop further, thus confirming the opinion of

Oldfield Thomas that pm? is the missing pr emolen,

The same interpretation holds good io the lower teeth.

To which dentition do the teeth of Marsupials belong ?

No very decided answer to this question can be obtained from

the material examined. ‘Traces of a successional dentition are

scanty, while those of a deciduous series are somewhat more

abundant and more distinct. One of us (H.W. M.'T.) has in

more than one paper (6, 7) previously urged, upon general grounds,

the view that the permanent dentition of the marsupials is the

permanent dentition of the Eutheria and that both the deciduous

and successional pm* belong to one and the same dentition. This

conclusion has been independently arrived at by Wilson & Hill

(10) and adopted by C. 8. Tomes (9). The joint authors of the

present paper are in agreement upon this subject, and we have

found nothing in the material which we have examined to cause

us to doubt the correctness of this view.

Note.-—I would desire to express my gratitude to the Odonto-

logical Society of Great Britain, for a grant in aid of researches

942

ON TOOTH-GERMS IN A WALLABY.

upon mammalian dentitions, and which has in part been used for

the purposes of this investigation ——H. W. M. '.

LITERATURE.

1. Deprenporr, T.—‘ Zur Entwickelungsgeschichte des Zahn-

12.

13.

Fig.

systems fete Mar supialier.” Semon’s Zoolog. For schungs-

reisen in Australien und dem malayischen Archipel,” Bd. ill.

1897-1910, pp. 243-402.

. Osporn, H. F.—‘ On the Structure and Classification of the

Mesozoic Mammalia.” Journ. Acad. Nat. Sci. Philadelphia,

vol. ix. p. 186.

Ross, C.— Ueber die Zahnentwickelung vom Chameleon.”

ia Anzeiger, Bd. vil. 1893, pp. 566— B77.

. Semon, R.—‘Die tiussere Hingwiekelun g des Ceratodus forsteri.”

Zoolog. Forschungsreisen in Australien und dem malayischen

Archipel, TBials te 1893.

THomAs, OLDFIELD.—‘‘ On the homologies and succession of

the Teeth in the Dasyuride, ete.” Phil. Trans. Lond.,

vol. 178, 1888, pp. 443-462.

Rint e Ww. Lassa, — Notes on the Dentition of the Dog.”

eae Anzeiger, Bd. xi, 1896, pp. 537-546.

“The evolution of the Teeth inthe Mammalia.

Anat. & Phys. vol. xxxvii. 1903, pp. 132-149.

“On the succession and homologies of the Molar and

Premolar Teeth in the Mammalia.” Jour. Anat. & Phys.

vol. xxxvi, 1902, pp. 321-343.

th)

Journ.

. Tomes, C. S.—“ Dental Anatomy.” London, 1898.

. Witson, J. T., & Hitt, J. P.—‘“ Observations upon the

development and succession of the Teeth in Perameles,

together with a Contribution to the discussion of the homo:

logies of the teeth in Marsupial animals.” Quart. Journ.

Mier. Sci. Lond., vol. xxxix. n.s., 1896-97, pp. 427-584.

“ Observations on tooth development in Orni-

thorhynchus.” Quart. Journ. Micr. Sei. Lond., vol. li. n.s.,

1907, pp. 137-165.

Woopwarp, M. F.—‘‘ On the development of the Teeth in the

Macropodide.” Proc. Zool. Soc. Lond. 1893, pp. 450-471.

“On the Teeth of the Marsupialia, with especial

reference to the pre-milk dentition.” Anat. Anzeiger,

Bd. xii. 1896, pp. 281-291.

EXPLANATION OF PLATE XLVII.

1. Showing the deeply situated bulbous rudiment of pm? with the very long

neck of the enamel organ reaching to the surface of the oral epithelium.

2. The enamel organ of pm with the labial dow ngrowth of the dental lamina

near to the free extremity of which is an enlargement, a “concentric epi-

thelial body ” (c.e.b.).

8. A ‘‘ concentric epithelial body ” more highly magnified.

4, Showing the sub-division of the enamel organ into two parts—?a fusion of

two distinct enamel organs.

5. Showing the presence of bloodevessels w aie the stellate reticulum of an

enamel organ.

P.Z.9.1944. Pl. XLVILL.

G.M.Woodward delet lith. West, Newman imp-

IDION(O BISA EMCI IalO) lee G37

DR. C. W. ANDREWS ON A NEW FOSSIL MAMMAL, 943

40, On a New Species of Dinotherium (Dinotherium hobley:™)

from British Hast Africa. By C. W. AnprEws, D.NSc.,

F.R.S., F.Z.8. (British Museum, Natural History) f.

[Received & Read May 23, 1911.]

(Plate XLVIIT.F)

During the last few years great additions have been made to

our knowledge of the extinct mammalia of Africa, but hitherto

the discoveries of their remains have been confined to the

northern and southern portions of the continent. Now, however,

a new find of mammalian bones in British Hast Africa shows

that great hopes may be entertained that before long light will

be thrown on the history of the group in the central portions of

the continent. Recently Mr. C. W. Hobley, C.M.G., Commissioner

of Mines in British East Africa, sent to the British Museum a

small box of bones from the neighbourhood of Karungu on the

east side of Lake Victoria Nyanza. Most of the specimens are

indeterminable fragments, probably picked up on the surface,

but in addition to these there are some beautifully preserved

teeth with a portion of the mandible (Pl. XLVIII. figs. 1, 1 a) of

a small species of Dinotherium; a small imperfect Proboscidean

caleaneum (fig, 5), a patella, and some other fragments probably

belong to the same animal. The teeth and bone are in a wonder-

fully good state of preservation, being hard and not easily broken,

and there can be no doubt that further collecting in the same

locality will yield results of the very highest importance and

‘interest. '

The teeth preserved all belong to the lower jaw: they are

pm. 4, m. 2, and m. 3 of the left side, and pm. 3 of the right; the

left pm. 4 and m. 2 have been replaced in their sockets in the

portion of the jaw preserved; this also contains the freshly

broken roots of pm. 3 and m. 13 m. 3 is isolated, the portion of

the jaw behind m. 2 being lost. Anteriorly the jaw fragment ceases

-at the level of the front of pm. 2, where its inner border is turning

in towards the symphysis; its ventral portion is missing, the

dental canal being exposed.

The teeth.—'The anterior premolar (figs. 3, 4) has a crown con-

sisting of a high, laterally compressed antero-external cusp, to the

inner face of which a shorter and more rounded antero-internal

cusp is closely united. The hinder half of the tooth consists of a

comparatively low and compressed outer tubercle separated from

‘the antero-external cusp by a notch, and a small rounded inner

* [The complete account of this new species appears here, but as the name and

a preliminary diagnosis were published in the ‘ Abstract’ the species is distinguished

by the name being under'lined.—Eprror. |

+ Published by permission of the Trustees of the British Museum.

t For explanation of the Plate sce p. 945,

944 DR. C. W. ANDREWS ON A NEW FOSSIL MAMMAL,

tubercle which is joined to the outer by a ridge widening from

within outwards,

The last premolar (pm. 4) (figs. 1, 1 @) consists essentially of two

transverse crests which are united towards the outer side by a low

obscure ridge. On the anterior face of the tooth there is a slight

prominence of the cingulum, the surface of wear of which becomes

continuous with that of the outer end of the anterior crest.

There is also a low transverse ridge formed by the cingulum”

on the posterior face of the tooth.

The first molar has unfortunately been lost; the second (m. 2)

consists of a pair of transverse crests, the ends of which, especially

the inner end, are higher than the middle and the anterior face

of which is concave. Qn the posterior end there isa well-developed

ridge of the cingulum extending nearly the whole width of the

tooth. The third molar (m. 3) is similar to the second except

that it has a well-developed triangular talon, the outer side of

which is formed by a prominent tubercle from which a crest

diminishing in height and width runs to the inner edge of the

tooth,

In size and, on the whole, in the pattern of the teeth this

Dinotherium is very similar to ). cwviert Kaup, which is from the

lower and middle Miocene of France, being apparently especially

characteristic of the Burdigalien horizon. Detailed comparison

however shows some differences. In pm. 3 the separation of the

antero-internal cusp is much more distinct than in a specimen of

the same tooth ascribed to D. cwviert ; the last premolar is shorter

in proportion to its width than in that species, and in the talon of

m. 3 the outer tubercle is much more distinctly and independently

developed. These differences, coupled with the remoteness of the

localities in which the two forms are found, seem to justify the

establishment of a new species for this East African animal and |

propose that it shall be called

DINOTHERIUM HOBLEYI Andrews.

Abstract, P. Z.5. 1911, p. 35 (May 30th).

The dimensions (in centimetres) of the teeth in the type-

specimen are :—

Length. Width.

[DIU Se eee: 4-0 32

Ole Ape cance 4°9 4-2

U1 Ob, PAIRS rec 6:2 5:2

MUO ee ee 02 54

An imperfect proboscidean caleaneum (fig. 5) probably belongs

to this species. It belongs to the left side, and a great part of its

outer half is broken away and some of the other faces are abraded.

Its extreme length is 17:2 em., the cast of a caleaneum referred

to Dinotherium giganteum is 32°7 em. or nearly twice as long.

From the same locality there have been obtained a lower molar

ON RATS FROM THE SOCIETY’S GARDENS. 945

and the distal end of a radius of a small species of Rhinoceros

(? Aceratherium), portions of the carapace of a very large

Chelonian, probably a species of Zestudo, fragments of the shell

of Trionyx, and some Crocodilian remains.

From some portions of matrix adherent to some of the bones it

appears that they are preserved in a bed of tough clay with much

calcareous matter and numerous grains of blown sand ; this deposit

is probably of lacustrine origin, but in the absence of any molluscs

or other invertebrates, it is not possible to be certain either as to

its origin or as to its exact age. Judging from the Dinotheriwm

remains, the beds are probably lower or middle Miocene. If they

should fortunately turn out to contain a rich mammalian fauna,

probably this discovery will lead to a great advance in our

knowledge of the history of several groups of Artiodactyls, of

the Hyracoids, and possibly of the Anthropoidea. It is greatly

to be desired that a careful collection should be made as soon

as possible.

EXPLANATION OF PLATE XLVIII.

Fig.1. Dinotherium hableyi, portion of left ramus of mandible with pm. 4, m. 2, m. 3,

trom above; la, ditto, from side. Type specimen.

. Ditto, crown of m.2 of another individual.

. Ditto, pm. 3 of the type specimen trom above ; 3 a, ditto, from side.

Ditto, crown of pm.3 of another individual.

Ditto, left calecaneum from inner side.

Om oo to

eub., facet for cuboid ; ect., ectal facet ; sus., sustentacular facet ; é.c., tuber calcis.

All the figures are ? natural size.

EXHIBITIONS AND NOTICES.

June 13, 1911.

Epwin T. Newton, Esq., F.R.S., in the Chair.

Mr. H. G. Puimmer, F.RB.S., F.Z.S., Pathologist to the Society,

presented a Report on the Pathological Examination of Rats

(Mus decumanus) caught in the Regent’s Park and in the Society’s

Gardens. 500 rats had been examined between the Ist of J anuary

and the 17th of May, 1911, all in a precisely similar manner, The

spleen, lungs, glands, and blood were examined microscopically ;

and from any animal which looked in any way unhealthy cultures

were made.

The results were summarized as follows :—5 rats were caught

in the Park, and 495 in the Gardens: 283 of these were males

and 217 females.

3 rats had tubercle, 10 had tapeworm cysts in the liver, 49 had

Trypanosoma lewisi in their blood, 2 had empyema (not tubercular),

1 had a tumour of the lower jaw (the result of an old injury), and

1 had pleuritis and hydrothorax (not tubercular), ;

946 MR. R. I. POCOCK ON THE CRESTED RAT.

Bacteria were found in 71 rats: in 40 in the lungs, and in 31

in the spleen.

Saccharomycetes were found in the lungs of 16 rats.

Fleas were found on 4 rats, and lice on 3 rats.

The general condition of the rats was very good, and in none

was anything at all suspicious found.

Dr. R. W. Ssaurexpr, C.M.Z.S., sent for exhibition a photograph

he had taken of a living specimen of a male albino Woodchuck,

Arctomys monax, that had been sent to him from Virginia,

UWESsA;

Mr. R. E. Houpine exhibited and made remarks upon the

Horns of a Highland Ram, a Fallow Deer, and a Roebuck, which

were fused at the base, and also the skull of a coursing Greyhound

with abnornal dentition.

Mr. R. I. Pocock, F.R.S., F.Z.S., Superintendent of the Society’s

Gardens, exhibited the skin and skull of a specimen of the

rare Crested Rat, Lophiomys ibeanus Thos,, which had been sent

from Nakuru for the Zoological Gardens by Mr. R. B. Woosnam,

C.M.Z.S., but had unfortunately died on the voyage. After

alluding to the well-known peculiarities of the skull in this Rodent,

Mr. Pocock drew attention to the arrangement and coloration

of the hair (text-fig. 190), and expressed the opinion that the

alternating areas of black and white with which each individual

hair is ornamented, must make the animal conspicuous at dusk,

if confidence be placed in the analogy supplied by such animals

as Poreupines and Zorillas. The coat consists of two kinds of

hair,—a softer close-set under-fur dusky grey at the base, then

white, then dark at the tip, and much longer, coarser hair usually

dark at the base, then white, then black, then white at the tip.

These hairs are so arranged that the dark and white bands of the

nnder-fur coincide exactly with the alternating bands of the same

colour in the coarser hairs, while the long white tips of the latter

oroject clear of the rest of the coat. From this arrangement it

results that when the hair is raised the median white bands

combine to form a continuous white mass thrown into relief by

the dusky base and the distal black area, the whole being su-

mounted by the white tips of the long hairs shining with almost

silvery lustre.

The coat of the upper side, moreover, is divided into three

definite regions—a median dorsal and a lateral on each side—by a

band extending from the shoulder to the hip and consisting of

subspiniform greyish hairs of peculiar spongy texture, and thick

in the middle but narrowest at the base and apex. When the

animal is viewed from above with its coat erect, the white areas

of the region on each side below this dividing band form a

MR. R. I. POCOCK ON THE CRESTED RAT. 947

continuous white lateral stripe which anteriorly merges more or

less, according to the species, with the white area on the summit

of the head; and when seen from the side the same white stripe

is visible as well as the white area of the hairs of the dorsal crest.

Text-fig. 190. -

Dorsal and side views of Lophiomys ibeanus.

In the typical form of Lophiomys, namely LZ. imhausi, the tail

is longer and much whiter and the frontal band Fareeln longer

and more conspicuous than in ZL. ibeanus. ‘These two Fontes

948 THE HON, P, A. METHUEN ON

conjoined must, theoretically, combine to make the former species,

or race, more easily seen at night than the latter.

That the Crested Rat is nocturnal was proved by observations

made upon the specimen of L. wmhausi that was kept in the

Jardin d’Acclimatation, Paris. It was also recorded of this

animal by Milne-Edwards that when irritated it raised its dorsal

crest erect and defended itself by biting vigorously.

Mr. Pocock concluded by remarking that the specimen exhibited,

which before being skinned was perfectly fresh having been kept

in a refrigerator, had a most peculiar but indescribable smell.

From this it might be inferred that Lophiomys was perhaps a

protected self-advertiser. On the other hand, it was considered

possible that it might be a mimic of the Porcupine, since the

coloration of the two was in a general way very similar *.

PAPERS.

41, On an Amphipod from the Transyaal.

By the Hon, Paun A. Muruuen, F.Z.S.

[Received April 27, 1911: Read June 13, 1911.)

(Plates X LIX.—LI.*)

Introduction.

About twelve months ago, Mr. Hewitt, of the Albany Museum,

Grahamstown, who, at that time, was at the Transvaal Museum,

Pretoria, communicated in a letter to the Rev. Noel Roberts and

to me, his discovery of a blind Amphipod together with some

Copepods in a cave at Irene, which lies about 9 miles south of

Pretoria; at the time of writing he considered the species to

belong to the genus Hucrangonyx Stebbing. Owing to other

work he postponed the description of it, and, when I came out

some few months ago, he very kindly gave me the specimens

collected to deal with.

Not long ago I visited the same cave and secured two specimens :

none of the Amphipods taken from this cave were large.

In February, Mr. Austen Roberts and myself went to the

Makapan Caves, which lie in hilly country about 15 miles west of

Potgietersrust, in the Transvaal, for we heard that some of these

caves contained water, and we hoped to obtain a more plentiful

supply of these cave Crustacea.

The first cave we visited—the more famous historically—con-

tained nothing we were in search of, so we directed our steps to

* Tn the discussion that followed this exhibition, Dr. R. E. Drake-Brockman, who

was acquainted with Lophiomys in Somaliland, remarked that the natives of that

country regarded these rats as young Porcupines.

«-+ Foy explanation of the Plates see p. 957.

“ayo URUTMAL '}S3AQ

IsSbaHeod

XANODNVAEONH

Ie ies) dlchiltle Jeu Ti,

West,Newman lith.

BUCRANGONYX ROBERTSI.

PGS) eS) JIL LAL

oe ras

eas

Zi,

West,Newman lith.

BHUCRANGONYX ROBERITSI.

AN AMPHIPOD FROM THE TRANSVAAL, 949

two others on the opposite side of the valley. In both we found

a plentiful supply of water.

As to these two caves, one was considerably deeper than the

other. In the first one we entered, we reached the level floor

fairly soon ; here we found shallow sheets of water supporting a

large number of Gammarids, Copepods, and Ostracods ; the bottom

was muddy and the floor of the cave generally covered by the

droppings of bats. No light entered as far in as this. The

bottom of the second cave we reached after a rather long, steep,

and winding descent: the stalactites and stalagmites showed it

to be a limestone cave. The water here was as clear as crystal,

rich in Gammarids and poor in Copepods; there were no

Ostracods, but a few aquatic worms were taken; some terrestrial

Isopods and some spiders were also found. Though the water

appeared quite still, a fresh supply was without doubt being added

continually. The bottom was rocky, covered by a thin layer of

fine mud, and the floor of the cave, as in the other, was strewn

with bat droppings.

Later in the day a few Gammarids, similar in every respect

to the others taken, were caught under stones at a spring in

the vicinity, but none were ever taken in the spruit which runs

through the valley, where the numerous crabs would make short

work of such fry.

All the Gammarids so far found in the Transvaal represent a

single species, belonging most probably to the genus Hucrangonya

Stebbing, and closely related to Hucrangonyx vejdovskyi Stebbing,

vide (3. p.389 and 4). However, this little cave form has not been

placed in this genus without notice being taken of certain resem-

blances and affinities to the genus Viphargus Schiddte(3. p. 405),

more so to Weoniphargus Stebbing (3. p. 404 and 2, p.73), and

the genera Crangonyx Bate and Paracrangonyx Stebbing (8.

p. 969 and 1. p. 218). In many ways this species appears to be a

generalized Niphargus-Crangonyx type, aS 1s seen mainly in the

nature of the telson and third uropods ; the breadth of the second

joint of pereiopods 3 to 5, and the total length of these appendages

as compared to pereiopods 1 and 2; the shape and size of the

gnathopods ; the number of sete and spines on the inner and

outer plates of the maxillula; the structure of the lips; the size

of the accessory flagellum and total length of the antennule.

It is in the sense of the comparative generalization of its

structure that this creature may be called primitive; primitive as

opposed to the more recent genera Paracrangonyx, Apocrangonyx,

and Crangonyx, but, on the other hand, more recent than the

genus Gammarus, and probably NViphargus.. This conclusion has

been arrived at more by the study of the telson than of any other

part (vide 1. p. 219).

The large size of the outer ramus of the third uropod appears

to be almost peculiar, but, judging from Chilton’s (1. p. 218)

remarks on the variability of this element in Crangonyz, this

character can be given apparently too much attention.

Proc. Zoot, Soc.—1911, No. LXV. 65

950 THE HON. P. A. METHUEN ON

It may be as well to record the outstanding characters of this

new species in order, and mention points of similarity and dif-

ference between it and the various genera and species mentioned

above, as they appear significant in leading us to some conclusion

as to the proper systematic position of the only known fresh-water

Amphipod from South Africa (unless some littoral marine forms,

taken in a stream in the Cape Peninsula and in vleis on the Cape

Flats within two or three miles of the sea, be considered as true

freshwater creatures).

The antennules are longer than the antenne, the accessory

flagellum small and 2-jointed, subequal to the first joint of the

normal flagellum. In Hucrangonya vejdovskyi, the second joint

of the accessory flagellum is half the length of the first. In

Neoniphargus, the accessory flagellum is 2-jointed, longer or

shorter than first flagellular joint.

Upper lip rounded as in Viphargus, Crangonyzx, etc.

Lower lip, inner lobes fairly small; in Paracrangonyx they are

small; in Wiphargus well developed; in Crangonyx flagellula

Benedict, very small. Mandibular processes moderate; in Hucran-

gonyx vejdouvskyi unusually prolonged.

Mandibles normal, dissimilar. Second joint of palp, subequal

to third, is rather broad owing to convexity of inner margin; in

Eucrangonyx vejdovskyi second joint broad, considerably longer

than third; in Paracrangonyx third joint subequal to second,

which is not expanded (in figure) ; in Viphargus third joint longer

than second.

Maxillula : inner plate with two plumose setz, outer with seven

spines serrated on the inner side; two or three sete to inner

plate in Niphargus and Neoniphargus, the former with seven

spines to the inner plate, the latter with six ; in Paracrangonyx

the same as for Hucrangonyx robertsi; inner plate of Crangonyx

with six sete, of Hucrangonyx vejdovskyi with four setee.

Maxilla.as in Hucrangonyx.

Maxillipeds as in Hucrangonyx (most probably), not unlike

those in Paracrangonye.

The two pairs of gnathopods as in Hucrangonyx vejdovskyt, but

palm equal to hind margin; in #. vejdovskyi it is much shorter.

Pereiopods 1 and 2 slender and slightly shorter than succeed-

ing pereiopods; in H. vejdovskyi they are “said to be slightly

longer.” Pereiopod 5 the longest. The second joints of all the

pereiopods much as in Z. vejdovskyi.

Pleopods normal; in Paracrangonyx slight and one-branched.

In uropod 1, peduncle a little longer than the equal rami; in

E. vejdovskyi as long.

In uropod 2, rami slightly unequal, peduncle as long as shortest

ramus; in #. vejdovskyi shorter than rami.

Uropod 3 has peduncle half the length of long outer ramus,

inner ramus minute with single spine; in H. vejdovskyi peduncle

half the length of outer ramus, inner ramus flattened, rudi-

mentary, shorter than peduncle; in Crangonyx and Paracran-

AN AMPHIPOD FROM THE TRANSVAAL. 951

gonyx uropod 3 not elongate, so also Neoniphargus which has

inner ramus minute; Miphargus has outer ramus long, two-

jointed, and the inner small.

Telson with emargination and almost square, as in ZL’. vejdovskyi ;

in Crangonyx entire or partly cleft; in Paracrangonyx entire ;

in Veoniphargus partly cleft ; in Viphargus deeply cleft.

This evidence seems sufficient to warrant the inclusion of

this new species in the genus Hucrangonyx near the species

EL. vejdovskyi.

I have taken the opportunity here offered of naming the species

after the Rev. Noel Roberts, in recognition of his enthusiasm for

this branch of zoology.

Detailed description of the new species.

Family GAMMARIDA® Leach.

Genus Eucranconyx Stebbing.

EUCRANGONYX ROBERTSI, sp.n. (Plate XLIX.)

Female: length of largest specimen taken 11 mm.*; colour

dirty yellow or pink in the shallower cave near Potgietersrust,

semi-transparent white in the deeper cave. No trace of eyes

was discovered.

Body rather compressed, smooth, no carina, rostrum barely

perceptible, a few minute spines on dorsal part of head and third

segments ; the last three or five segments with spines on posterior

margin.

First four coxal plates of pereion deeper than the rest, the first

smallest, the third and fourth deepest ; the fourth plate broader

than the others; side-plates of the fifth, sixth, and seventh

pereion segments small, those of fifth and sixth deeply emarginate

behind. These coxal plates bear marginally small spines.

Antennules (Pl. L. fig. 1) long, longer than the antenne;

flagellum much longer than the peduncle; first joint of the

peduncle a little shorter than second and third joints together ;

second joint rather longer than third ; accessory flagellum (Pl. L.

fig. 1 q@) two-jointed—both joints armed with sete,—minute,

hardly as long as first joint of flagellum; all the joints of the

antennule provided at their distal extremity with moderate sete,

of which one or two on each segment are conspicuously longer

than the rest; the second joint of the peduncle has also two

groups of small setee and one other seta besides about its middle

length ; the arrangement of the setz on each joint of the flagellum

appears to be constant in disposition after the eighth joint, and

to be about seven in number.

Antenne (Pl. L. fig. 2) about four-sevenths the length of the

* The length includes that part from the anterior margin of the head to the

posterior margin of the last pleon segment.

65*

952 THE HON. P. A. METHUEN ON

antennules; peduncle longer than flagellum, the ultimate and

penultimate joints well armed with numerous sete which are

arranged, generally speaking, in succeeding and nearly parallel

semicrescents about all parts of the joints except the proximal

extremities; on the other joints of the peduncle the sete re-

present a much diminished quantity; the distal parts of the

flagellum joints carry about eight sete each, disposed in an un-

varying sequence.

Lips: wpper lip (Pl. L. fig. 4) rounded, with some minute setz

about the extremity, at the centre of which they take an inward

course for a short way on each side. Lower lip (Pl. L. fig. 3):

inner lobes with minute sete on distal convex margins; outer

lobes with longer setze; mandibular process not elongate, bearing

small sete.

Mandibles (P\. L. figs. 5 & 5a): palp rather long when com-

pared with the size of the mandible itself; first joint shortest,

devoid of sete; second joint rather broad and curved, the side

opposite the biting process markedly convex and armed with

two rows of sete (about fifteen in number); the ultimate joint is

about the same length as, or a little longer than, the penultimate

and is provided with terminal sete on both lateral margins. Palp

on each side similar except that basal joints are of slhghtly

unequal lengths. Mandibles themselves of unequal size and of

slightly different structure (vide figs. 5, 5 @).

Mawillula (Pl. L. fig. 6) resembles that of Paracrangonyx com-

pactus Chilton very closely. The first jomt of the palp is short,

the terminal joint bears about nine bristles; the endite of the

third joint of the maxillula is armed with seven stout serrated

bristles, which resemble those found in a similar position in

P. compactus but differ in being more extensively serrated, and,

further, at the base of the innermost of these bristles is found a

group of fine hairs; this endite is about double as broad as the

palp; the endite of the first maxillula joint (accepting Hansen's

interpretation of the endites) appears to be exactly similar to

that of P. compactus, bearing two fairly long plumose sete and a

number of hairs on its inner margin.

Mazilla (Pl. L. fig. 7): the endites are simple, regular, and

well-developed structures, both armed distally with a great number

of setee which are disposed in two rows, the more distal of these

rows being considerably longer than the rest, at any rate in the

case of the endite of the third jomt; the inner side of the endite

of the second joint bears a number of fine hairs, and between

these and the distal rows of setze les one long plumose and another

smaller, slightly plumose seta.

Mazilliped (P1. L. fig. 8): the shape of the joints like those of

Paracrangonyx compactus, the whole limb being, however, some-

what slenderer ; the seventh joint alone is shorter than that of

P. compactus; a great number of closely-set sete and bristles

clothe the inner surface of the two endites and of the fifth, sixth,

and seventh joints. ‘The basipoditic endite or inner plate carries

AN AMPHIPOD FROM THE TRANSVAAL. 953

three plumose sete on its inner margin, and distally three short

stout bristles and about eleven plumose setee. The endite of the

ischiopodite or outer plate carries a number of fairly short stout

bristles distally, and along two-thirds of its inner margin six of

the distal bristles are serrated.

Gnathopods: about the same size and length; the fifth and

sixth joints of gnathopod I. somewhat broader than those of

gnathopod II. In the male, gnathopod I. is a little longer than

gnathopod IT.

Gnathopod I, (Pl. L. fig. 9) well developed, subchelate; the

coxa of the first joint in shape making almost a parallelogram,

sloping forward somewhat, anterior portion deeper than posterior,

diagonal from posterior ventral corner to anterior dorsal corner

equalling greatest length of coxa; anterior margin bearing five

small spines and ventral margin five also. The second joint or

basos is narrow at the base, has anterior side straight, and bears

six feebly serrated sete, the most distal the longest; posterior

margin convex, most markedly about the middle of the joint

which is here seen to be two-fifths in breadth to length; the

posterior margin carries a number of long sete, and following

this arrangement, four setze proximally, equidistant apart, followed

by a group of two, then a group of three, a group of two, another

group of three setz all serrated, and distally a group of four

smaller sete, not serrated. The third joint or ischium is short,

broader than long, and bears on its posterior distal extremity five

sete not serrated. The fourth joint or meros has the posterior

distal portion rounded and well provided with numerous serrated

setee, proximal to which lie a group of four serrated sete; on the

anterior margin four small sete can be noticed. The fifth joint

or carpus, which is more or less triangular, has a dense cluster of

serrated setze on its short posterior portion, which are arranged

into groups, one consisting of a row of marginal sete, another of

apparently two rows placed submarginally ; the anterior margin

of the joint carries four sete. The sixth joint or propodos is

subovate, equal in length to the longest measurement of the basos,

as wide as the greatest vertical depth of the coxa; anterior margin

convex, with four groups of four fairly short sete arranged at

varying intervals from a fifth group of six longer sete situated

near the base of the dactylos; in addition, a small solitary seta

will be noticed external to the third of these groups; the posterior

margin, starting from the proximal end, is straight for a short

distance, to a part where four small well-marked prominences are

encountered ; on each of these are placed two finger-shaped pro-

cesses of unequal length and, excepting on the last prominence,

two sete ; behind the fourth prominence lies a row of sete: the

palm, to which the dactylos fits closely, follows immediately ; it is

provided with a series of very short spines arranged in couples,

each couple equidistant apart; a number of short sete, nine in

number, are also present opposite the border of the most posterior

muscle, disposed more or less in a straight line, and varying

954 THE HON. P. A. METHUEN ON

slightly in individual length ; between the eighth seta of this

series and the middle portion of the palp are placed three fairly

long sete, of which the central is the longest. The dactylos,

which is slightly curved, is provided with two sete on its anterior

and four very short sete on its posterior margin; this inner

margin bears distally a small tooth distinct from the long

curved terminal unguis; between these two lie two small sete.

None of the sete on the ultimate and penultimate joints are

serrated.

Gnathopod II. (Pl. LI. fig. 10) is very like gnathopod I. Only

the chief differences will be mentioned. The sete on the second

joint are somewhat different in disposition and in length. The

third joint is somewhat longer and narrower and possesses more

setee. The fourth joint has fewer sete and only one fairly dense

group of about ten sete. The fifth joint is three-quarters as

broad as long (the same measurements in gnathopod I. show the

breadth to be greater than the length); the anterior and posterior

margins are comparatively long, the former the longest; the sete

on the posterior margin are longer and more numerous and are

arranged in seven or eight groups. The sixth joint has, on the

anterior part, an additional group of fairly long sete, which vary

in number in each group: the whole joint is longer and narrower:

the palm does not differ much, but the proximal half of the pos-

terior margin bears eight groups of long feebly serrated setee,

between which and the palm lie five finger-shaped processes, one

conspicuously longer than the rest with another little one at its

base. The dactylos and terminal unguis spine are both shorter ;

between the latter and the small terminal tooth there are no

sete.

Pereiopod J. (Pl. LI. fig. 11): the coxa of the first joint is

almost rectangular; the anterior and ventral margins are,

however, somewhat concave, the middle part of the posterior

margin slightly convex; the anterior margin has four short sete,

the ventral three. The second joint is long, narrow at the

proximal base, whence it widens rapidly but nowhere greatly,

being never more than a quarter as broad as long; nine spines of

moderate length and seven spines of greater length arm its

anterior and posterior margins respectively ; further, a group of

four sete are placed at the distal extremity of the posterior

margin. The third joint is a little longer than that of the

gnathopods but is considerably narrower; its posterior margin

carries a group of four setz distally. The fourth joint is long,

longer and broader than the fifth, the two joints together about

the same length as the second joint: the anterior margin carries

three setee unequal in length—at the base of the longest a minute

seta can be seen—and distally a group of three sete, one of which

is long; the posterior margin bears three groups of three sete

each, then one short seta, then distally a group of three sete.

The fifth joint, which is equal in length to, but broader than, the

sixth joint, carries on its anterior margin two minute sete, and

AN AMPHIPOD FROM THE TRANSVAAL. 955

distally a group of three small sete; the posterior margin is pro-

vided with four groups of sete, the proximal group consisting of

three sete, the next of two, then another of two, the most distal

group of four medium and one large seta. The sixth joint, which

is slender, bears on its posterior margin six groups of three sete

to each group, and on its anterior margin three minute sete, and

distally a group of four setze.

Pereiopod II. is practically similar in all respects to pereiopod L.,

to which it is subequal; on the second and third joints there are

two or three small additional sete marginally.

Of the remaining pereiopods the fifth is the longest, the third

the shortest, these differences of length being appr eoable but not

great; they are, however, considerably longer than the first and

second. In the third pereiopod, although the second and third

joints are somewhat shorter than those of the first pereiopod, the

great difference in length is made up by the much greater length

of the fourth, fifth, sixth, and seventh joints of the third pereiopod;

at the same time these four joints, though slender, are of the same

breadth as those of the first pereiopod.

Pereiopod II], (Pi. LI. fig. 12): the coxa of the first joint is

somewhat concave anteriorly, emarginate posteriorly ; anteriorly

and ventrally it bears some small spines. The second joint is

rather narrow at the base, broadens very rapidly, and then

narrows gradually towards its distal extremity ; it is a little more

than twice as long as broad; the anterior margin is slightly

convex and is beset witha number of moderate spines—deseribing

from the proximal end, three single ones equidistant apart, six in

pairs subequidistant apart, and terminally a group of three spines:

the posterior margin has fourteen shorter spines equidistant apart

and terminally two closer together; behind these latter a large

spine. The third joint is slightly rounded and carries a group of

four fairly small sete. The fourth joint is broader at the base

than that of the first pereiopod; the anterior margin carries nine

small sete, arranged in pairs all but the most proximal, and

distally a group of four sete, one conspicuously larger than the

others; the posterior margin carries two moderate spines and

three small ones and distally two fairly large and two small spines.

The fifth joint is armed with five groups of spines, the most

proximal group of two, the next three of three, and the following

of two spines to each group, and distally with a sixth group made

up of two larger and two smaller spines; opposite the last group

is a group of one larger and three smaller spines, some distance

behind which is a single spine, and behind this spine again two

groups of three spines, and disposed proximally to these a couple

of small spines close together. The sixth joint is conspicuous on

account of the Jarger size of some of the spines on its anterior

margin ; on this margin four groups of three spines to each are

fixed, and distally a group of eight spines varying in length:

the opposite margin six groups of spines are seen, each ee

possessing two or three spines. The seventh joint, including the

956 THE HON. P. A. METHUEN ON

claw, is one-third as long as that of pereiopod I., and carries on

the posterior margin two small spines.

In pereiopod IV. the details of the arrangement of the sete

differ very slightly.

In pereiopod V. the third and fourth joints are equal in length

to, but broader than, those of the third pereiopod; the seta

arrangement is not exactly similar, but this slight difference is

negligible. In Irene examples the penultimate joint of this

appendage carries much longer spines on one side than is the case

in specimens from the Makapan caves.

The pleopods are normal; the basal portion bears two branches

and two little coupling-hooks. The longest pleopod is the

second.

Pleopod I. (Pl. LI. fig. 13): basal portion oblong, slender, and

devoid of sete, except the two coupling-hooks ; these little

structures bear a few tubercles and one slightly hooked at the

end. The inner ramus is somewhat longer than the outer; each

ramus consists of thirteen joints which bear long plumose sete.

Pleopod II.: the basal part bears distally on its anterior margin

two fairly long spines, which are absent in the other pleopods and

appear to be absent altogether in the male.

Pleopod III.: the number of joints to the rami are two or

three less than those of the first pleopod.

The wropods: the first uropod the longest; the third uropod

not quite so long as the first; the second uropod about two-thirds.

as long as the first. The peduncle of the first uropod long, longer

than either ramus, much longer than peduncle of second; peduncle

of second much longer than peduncle of third. Outer ramus of

third uropod long, much longer than those of first ; inner ramus

minute. Rami of first uropod longer than those of second.

Uropod I. (Pl. Li. fig. 14): peduncle, broadest at the base,

bears five spines on posterior margin, and two others close

together distally. Outer ramus a trifle shorter than inner, which

bears five spines on posterior margin and two larger and four

smaller ones at the extremity. The outer ramus with four rows

of spines, three in each row, and another row basally of two

spines; distally two larger and two smaller spines.

Uropod IT, (Pl. Li. fig. 15): peduncle fairly stout as compared

with that of uropod I., narrower basally, but broader distally than

same; on posterior margin two spines; on median ridge (vide fig.)

five spines, the most distal being the longest. Inner ramus

longer than outer, which carries on posterior margin four spines,

and distally two longer and five shorter spines; anterior margin

has two small spines; the outer ramus bears posteriorly four

spines, two longer and three shorter distally, and eight spines

arranged in couples anteriorly.

Uropod ITI, (Pl. LI. fig. 16): peduncle about the same breadth

as that of uropod IJ.; anterior margin bears distally two small

spines; opposite these is seen the inner ramus bearing one small

spine, not counting spine it is one-fourth the length of the peduncle.

AN AMPHIPOD FROM THE TRANSVAAL. 957

Anterior margin of outer ramus carries six groups of spines, three

to each group except the two most distal, which are made up of

two spines each; distally is a group of fairly long setz: the

posterior margin shows five groups of spines, the first two groups

of three spines, and the other three of four spines each. At

junction of peduncle and outer ramus three spines can be dis-

tinguished. In the male the outer ramus appears to be hardly as

large as in the female.

Telson (Pl. LI. fig. 17) almost square; sides slightly convex :

two small plumose sete on each side bear two larger and two

smaller spines placed in a posterior position. The structure

excavate behind; in the male this excavation not quite so deep

as in the female. In specimens from Irene one of the terminal

spines on each side is much longer than in other specimens.

Literature referred to.

1. Cuinron, C.—‘ The Subterranean Crustacea of New Zealand.”

Trans. Linn. Soc., 2nd ser. Zool. vol. vi. pt. 2, 1894.

2. Smirn, G. W.—“‘The Fresh-water Crustacea of Tasmania.”

Trans. Linn. Soc., 2nd ser. Zool. vol. xi. pt. 4, 1909.

3. Srepsrne, T. R. R.—Amphipoda Gammaridea in Das Tierreich,

21 Lief. Berlin, 1906.

4. Vuspovsky.—Crangonyx subterraneus (non Bate 1859) in SB.

Bohm. Ges., nr. 10, p. 12, t. 1-3.

EXPLANATION OF PLATES XLIX.-LI.

The appendages have been drawn with the aid of a camera lucida.

All the figures are of Hucrangonyx robertsi, 2.

PratE XLIX.

Side view of female, X 11.

Prate L.

Fig. 1. Antennule, x 29.

la. Accessory flagellum, X ca. 160.

2. Antenna, X 29.

3. Lower lip, X ca. 45.

4, Upper lip, X ca. 45.

5 & 5a. Mandibles, X 29.

. Maxillula, X 75.

. Maxilla, X 75.

. Maxilliped, X 40.

. Gnathopod I, X 29.

co O-1

Prats LI,

Fig. 10. Gnathopod IT, X 29.

11. Pereiopod [, x 29.

12. Pereiopod III, X 29.

13. Pleopod I, x 29.

14. Uropod I, X 29.

15. Uvropod II, x 29.

16. Uropod III, x 29.

17. Telson, X 29.

Text-fig. 191.

958 MR. R. LYDEKKER ON

42. An African Rhinoceros, Klipspringer, and Gazelle *.

By R. LyprexKer f.

[Received May 9, 1911: Read June 13, 1911.]

(Text-figures 191-193.)

I. THe Somatt RHINOCEROS.

The presentation to the British Museum by Mr. Drake-

Brockman of two skulls of the Somali Rhinoceros, affords an

opportunity of considering whether that animal is entitled to

rank as a distinct local race of the so-called black species.

Sportsmen, I am told, almost invariably regard it in that light;

* The complete account of the two new forms described in this paper appears

here, but as the names and preliminary diagnoses were published in the ‘Abstract,’

the species are distinguished by the names being underlined.—Ep1Tor.

+ By permission of the Trustees of the British Museum.

Upper aspect of skulls of East African (A) and Somali (B) Rhinoceroses,

THREE AFRICAN MAMMALS. 959

and Mr. Ward informs me that such heads as he has mounted

indicate a relatively small animal, with horns inferior in size to

those of the Eastern and the Southern Rhinoceroses, and a skin

with a somewhat different epidermal structure.

According to modern views in regard to nomenclature, the

Somali Rhinoceros already has a scientific name, since it is re-

ferred to by Count Joseph Potocki on page 82 of his work entitled

‘Sport in Somaliland, London, 1900, as Rhinoceros bicornis

somaliensis; and although no description was published at the

time, the accompanying plates apparently render the name valid.

Text-fig. 192.

Front (A) and side (B) views of skull of Nigerian Klipspringer.

Of the two skulls presented by Mr. Drake-Brockman, one is

that of a subadult animal, with the whole of the permanent

dentition in use, and almost perfect, although the tip of the pre-

maxille is broken off. The other, which is considerably more

imperfect, is that of a younger animal, with the upper premolars

only just coming into wear.

Compared with a skull of Rhinoceros bicornis from East Africa

(B.M. No. 7.2.26.1)* (text-fig.'191), of somewhat greater age than

either of the Somali specimens, the latter are seen at once to

differ by the narrower form of the whole upper surface, both at

the interparietal constriction and at the orbital expansion. The

boss for the front horn is also much less expanded in the Somali

skulls, and there is less convexity in the region immediately behind

* T am fully aware that this is not the type locality of the species.

960 MR. R. LYDEKKER ON

this. Moreover, the palate is more decidedly vaulted in the

Somali than in the East African skull. So faras I can ascertain,

these differences appear to be constant in all the skulls available

for comparison.

The differences in the proportions of the Somali and East

African skulls will be apparent from the following table :—-

E. African, Somali.

iKensbh ofjupper aspectiwmsaeeceenceteee ee eens 227 ins. . 23 ims.

reali hsabhon outs aieee anne see ee anne 11 94

TPRVEMRAIUG VAIN Gbegysoc 200 cccaaansormonssassean eas. 14 203

A ROTMEM HO WANCNE D5 45 sco opgss0gedgo0nsoooobosceoneseS 142 Te

Length of upper tooth-row (excluding p.1)... 102 10¥

AW elit has Sot ain’ Ss cas, Ns hone ate et eee Pac a 24

As these dimensions amply demonstrate the racial distinctness

of the two forms, the Somali animal may stand as &. bicornis

somaliensis Potocki; the specimen here described occupying the

position of type. It may be added that if the East African and

Somali skulls were of the same age, the difference in the lengths

of the upper tooth-row would be greater.

Ii, Tue Nicrrian Kirpsprincer (Oreotragus saltator porteust).

Lydekker, Abstract P. Z.S. 1911, p. 38 (June 20).

Early in May, as I have already stated in the Field news-

paper for that month, Mr. Rowland Ward directed my attention

to the skull and borns of a male Klipspringer from the Duchi ’n

Wai range of the Yola province of Northern Nigeria, lying to

the south-west of Lake Chad. The skull was taken from an

animal shot there by Dr. E. J. Porteus, by whom it was kindly

piaced at my disposal. Klpspringers, it appears, are quite

familiar to the natives working in the Yola tin-mines, by whom

they are known as gaddi-dueki, a term equivalent to Hill Duiker,

The skull (text-fig. 192) differs from that of an East African

Klipspringer by its much greater width ; its diameter across the

orbits being 34 inches, whereas that of the Hast African specimen

is 212 inches, It is further distinguished by the marked bending

down of the margin of the lachrymal bone, which consequently

has a distinct lateral surface in place of a sharp edge, and also

shows only a comparatively small part of its total area from the

frontal aspect.

Klipspringers, so far as [ am aware, appear to be unknown on

the West Coast north of Angola*; while to the north, Abyssinia

is the nearest point to Yola where they are found. As the

country between the last two districts is to a great extent low

desert, the Yola Klipspringer must be quite isolated.

On this ground, and also on account of the peculiarities in the

form of the skull, it is clearly entitled to racial distinction, and

I propose to name it Oreotragus saltator porteusi, in honour of

Dr. Porteus.

* For the information that Klipspringers inhabit Angola, I am indebted to

Mr. E. A. Hamilton.

THREE AFRICAN MAMMALS. 961

The type of the new race of Klipspringer will be the aforesaid

skull, which Dr. Porteus has kindly presented to the British

Museum.

I know of no Antelope with a distribution identical with that

of the Klipspringer, as now extended.

About a fortnight after describing this skull I received a letter,

dated Naragata, Northern Nigeria, from Mr. M. P. Hyatt,

informing me that he had recently killed three Klipspringers—

an adult buck and two does—in that part of the country.

IIT. An ALGERIAN GAZELLE (Gazella hayi).

Lydekker, Abstract P. Z.S. 1911, p. 38 (June 20),

At the close of 1909, Mr. M. V. Hay presented to the British

Museum the skin and skull of an adult male Gazelle, shot by

himself in Algeria between Constantine and Biskra, and sup-

posed to be a Doreas (Gazella dorcas). In due course the specimen

was set up, and placed on exhibition as a representative of that

species. Recently, however, as already mentioned in the Field

newspaper *, it was pointed out to me by the donor that the

specimen differed considerably from the Dorcas, and I was informed

at the same time that its distinctness is reeognised by the Arabs,

who call it rhozal-rim ; rhozal being the name of the dorcas, and

rim that of Loder’s Gazelle (G. leptoceros). On comparison of the

specimen with an undoubted Dorcas from the Biskra district, the

difference between the two became apparent (text-fig. 193).

Mr. Hay’s specimen is about the same size as a Dorcas, but

appears to have rather larger ears,—I say appears to have, because

one 1s never certain whether there may not have been stretching

or shrinking in the mounting. Its most distinctive characteristic

is, however, to be found in the form of the horns, which com-

pletely lack the double, sublyrate curvature of those of the Dorcas,

and incline almost directly upwards and slightly inwards, with a

slight inward and forward turn at the tips. There are also much

fewer rings on the horns, the present specimen having only 12,

whereas a Dorcas may have 24 or 25; possibly a more aged

example of the new form might develop two or three more rings,

but even then the difference would be very considerable.

The face-markings differ considerably from those of a Dorcas,

and are more like those of an Edmi (4. euvieri), the middle stripe

being much darker, with a conspicuous nose-spot, and the light

eye-stripes much less apparent, while the forehead lacks the

chestnut tint of the Dorcas. The body does not show the faint

light stripe above the flank-band which occurs in the Dorcas ; the

knee-tufts are larger and blacker; and the tail is brown instead

of black, with a shorter fawn area at the base.

As the skull of Mr. Hay’s specimen is mounted in the skin, I

cannot give cranial characters; but the foregoing features are

amply sufficient to distinguish the new Gazelle from the Dorcas,

* June 3rd, 1911.

962 ON THREE AFRICAN MAMMALS.

which is found much more abundantly in the district: frequented

by the former. ‘This being so, there is no doubt as to the dis-

tinctness of the new species, which I have named Gazella hayi,

from all other Algerian Gazelles, unless it be G’. kavella and the

so-called Antilope corrina, neither of which can now be identified.

The type of the new species will, of course, be the Museum

specimen.

both from the Constantine—Biskra district,

Heads of Gazella hayi (A) and G, dorcas (B)

P49 Ili, Jl, ILI.

Witherby & Co., Imp.

CAPRA PYRENAICA HISPANICA.

(Summer Pelage)

AS Iii, 2, Li.

Witherby & Co, Imp.

CAPRA PYRENAICA VICTORIA 4é, &.

(Summer Pelage)

IP ZoS> t@iil, Pi, ILM

Witherby & Co., Imp.

CAPRA PYRENAICA VICTORI€.

(Winter Pelage)

ON THE SPANISH IBEX. 963

43. The Subspecies of the Spanish Ibex.

By Prof. Ancen Caprera, O.M.Z.S.

[Received May 13, 1911: Read June 13, 1911. ]}

(Plates LIT.-LIV.* and Text-figures 194-199.)

The first scientific description of a Spanish Ibex was that

published in 1833 by F. Cuvier, in the great iconographic work

‘ Histoire Naturelle des Mainmifeéres,’ pl. 396. It was based on

a young male from the Pyrenees, of which the author gave

a beautiful figure in winter pelage, but was not accompanied by

any technical name. In the index of the work, published in 1842,

the animal is erroneously called Capra ibex, the fact being

apparently ignored that four years before Schinz had described

the same forn m of Goat under the name Capra p yrenaice T ie

In volume xxvi. (1848) of the ‘Comptes Rendus’ of the Paris

Academy of Science, Schimper briefly mentioned the Ibex of the

Andalusian sierras, naming it C. hispanica and considering it as

quite a different species, a view followed by all the authors “of the

time, and sustained still in our own days by F orsyth Major =

and Graells$. Modern zoologists, however, seem to agree in

admitting only one species of Spanish Ibex, though admitting that

there are some differences between the specimens coming from the

Pyrenees and those from other parts of the Peninsula. Sclater ||

considers the latter as a “ slightly altered phase” of the foruione

and ‘Trouessart, in his ‘ Conspectus Mammalium Europe,’

expresses the same idea in a more modern fashion, describing

two different subspecies: Capra pyrenaica, from the ‘“‘ chaine des

Pyrénées,” and C. pyrenaica hispanica, from the “chaines de

montagnes de |’Espagne Centrale et Méridionale.”

I do’ not intend to discuss now the meaning of the terms species

and subspecies. While awaiting a satisfactory and univer. sally

accepted definition of these wor ds, I agree with other authors in

considering all the Spanish Wild Goats as belonging to a single

species ; but as to the number of subspecies, I think there are,

not two, but three at least, the Ibex of Central Spain being quite

different from both the Pyrenean and the Andalusian forms.

This view has been anticipated by Ménégaux in Perrier’s ‘ Vie

des Aniimaux,’ as he says that ‘la forme qui habite les sierras du

centre de la Péninsule fait le passage entre les deux formes

susnommeées ” (pyrenaica and hispanica). This central subspecies

remains, however, unnamed and undescribed as a different form,

and to name and describe it are the chief purposes of the present.

paper.

* For explanation of the Plates see p. 977.

+ Neue Denkschr. Allg. Schweiz. Ges. Nat. ii. 1838, p. 9.

t Atti Soc. Tose. Sc. Nat. iv. 1879, p. 2.

Memorias Acad. Cienc. de Madrid, Xvil, 1897, p. 356.

|| P. Z.S. 1886, p. 315,

964 PROF. A. CABRERA ON

Before doing so, must remark on the geographical distribution

of the species and on the differences between the various races.

In that interesting book ‘ Unexplored Spain,’ MM. Chapman

and Buck have quite recently told the history of the Wild Goats

of Spain and the manner in which they are protected against

imminent extinction, giving some details about their present

distribution. The authors are not exact, however, in saying that

the isolated colonies now formed by the Ibex have been “ separated

from each other during ages.” There are, in fact, strong reasons

for believing that in the past Ibexes inhabited every suitable

point of almost every mountain ridge in Spain. Names recalling its

existence, suchas Las Cabras, Cabrales, Cabrera, Cabreira, Cebreros*,

&e., are quite common in all the mountainous districts. In the

seventeenth century the species was found in all the Sierra Morena

and Sierra de Cazorla. The Sierra de Segura, Sierra de Francia

in the Salamanca Province, and the Toledo Mountains, where it

does not exist to-day,’formed parts of its range sixty years ago,

and it has been found in the Sierra de Bejar, between the Sierras

of Francia and Gredos, so recently as 1897 7; and in 186], the

date of Seoane’s ‘ Fauna mastolégica de Galicia,’ a few individuals

remained in the mountains of that region. That Ibexes inhabit,

or at least inhabited in 18904, the mountains of Gerez, in the

northern border of Portugal, is a well-known fact.

In connection with the existence of Ibexes in the western

extreme of the Cantabrian chain, it must be remembered that

fossil remains of Capra pyrenaica have been found in Santander,

and from this we may surmise that the species reached the north-

western corner of the Peninsula from the Pyrenees, through that

northern ridge. It spread from there southwards into the

central sierras, either through Portugal by the Serra da Estrella,

included in its range by Trouessart, or directly through the

Burgos mountainsand the Guadarrama. Father Saturio Gonzalez,

a noteworthy collector of mammals, tells me. that he has found in

the old monastery of Silos a pair of Ibex horns which have been

preserved there for centuries, and about sixty miles east from

Santo Domingo de Silos there are a Sierra de Cabrejas and

a village named Cabrejas del Pinar. ‘These facts seem to suggest

that the Ibex was once common in the mountains connecting the

northern chain with the Sierra de Gredos, but, since there is not

any evidence of its existence in the Guadarrama, too much

confidence cannot be put in this hypothesis.

As to the Andalusian sierras, it is evident that the Ibex found

* Oebra was frequently used instead of cabra as a name for the Ibex during the

twelfth and thirteenth centuries. It was from this that the old Spanish writer

Father Sarmiento supposed that Zebras formerly existed in Spain. The modern

vernacular naine of the animal is cabra montés (Mountain Goat), or simply montés,

but there are some other local names. In the mountains about the lower Ebro it is

called sawvatge (the wild one); the people of Galicia call it.craba brava or craba fera

(Wild Goat), whereas in the Pyrenees the name bucardo, related to the English buck

and the French bouwe and bowqguetin, is commonly used.

+ Rivas Mateos, Actas Soc. Esp. Hist. Nat. 1897, p. 208.

+ P. de Oliveira and L. Vieira, Annaes Scienc. Naturaes, 111. 1896, p. 91.

Text-fig. 194.

THE SPANISH IBEX. 965

its way to them by the Toledo mountains and the Sierra Morena,

turning afterwards along the mountainous ridges of Hastern

Spain northwards to the lower Ebro basin. Notwithstanding the

proximity of this region to the Pyrenees, I cannot believe that

the Ibex inhabiting it immigrated directly from the Pyrenean

chain, as it is very different from the form found here, and quite

indistinguishable from the southern race.

Baceanic !s.

x q

foc OD 8

(Li 3 xy

S. a

oy cS

é TANS

LISBO

In the geographical range of the Spanish Ibex four perfectly

distinct areas may be distinguished, although now reduced toa

number of small isolated colonies by continued persecution from

Proc. Zoou, Soc.—1911, No. LX VI. 66

Map showing the probable former distribution (dotted areas) and present colonies (vertical lines)

of the Spanish Ibex.

966 PROF. A. CABRERA ON

the Middle Ages, when wild-goat meat was a very favoured dish

at every Spanish table. A different subspecies is found in each

of these areas, as follows :—

(a) Pyrenean area, comprising the Spanish side of the Pyrenees

and, in former times, the eastern part of the Cantabrian chain.

Tts peculiar Ibex is Capra pyrenaica pyrenaica. It may be con-

sidered as practically extinct, being today found only in the

northern extreme of the Huesca Province, about the Mount

Perdido*. Two old bucks, three females, and three or four

half-grown individuals remained there in 19077. In a recent

letter on this subject, the Count of San Juan, who spends a great

part of his time hunting in the Pyrenees, kindly informs me:

“T think that probably no more than ten or twelve Ibexes

remain in all the Pyrenean chain. A pair survived recently in

the Maladeta; somebody shot the female, and the male sought

refuge among a herd of domestic Goats and was subsequently

killed by the goatherd.”

(6) North-western, or Atlantic, area, formed by the mountains

of Galicia and Northern Portugal. The lack of suitable material

prevents correct identification of the Wild Goat found there, but

from the description by Barboza du Bocage t I surmise that it

represents a peculiar subspecies, which I do not care to describe

at present. At all events, it is well-nigh extinct, only a few

specimens, if any, remaining in the Portuguese mountains of

Gerez. Not being a Spanish race, we need not discuss it now.

past, the ridges of El Barco, Bejar and Francia, and the hills

of Toledo. The subspecies inhabiting this area, at present reduced

to a single colony in the highest peaks of Gredos, will be named

and described below. MM. Chapman and Buck have told the

history of this Ibex so accurately that it is unnecessary to repeat

it here. The colony consists of about three hundred and fifty

head, and having been under royal protection since 1905 it is

rapidly improving.

(¢) Mediterranean area, from the mountains forming the

Guadalquivir basin, eastwards and northwards through the

sierras of the Valencia Province to the mouth of the Ebro. It

is inhabited by Capra pyrenaica hispanica (type locality, Sierra

Nevada), a subspecies not so near extinction as the Pyrenean and

Central forms. No less than six colonies, in fact, are known to

exist, the exact number of heads in each of them being unknown.

Three of these colonies are in South Spain, viz. :—one in Sierra

Nevada, another in the two parallel ridges of Sierra Bermeja and

* The Mont Perdu of the French. It is a custom with many English writers to

use French names for localities on the Spanish slope of the Pyrenees, but, in my

opinion, such a course is against commonsense. Since these localities are in Spain,

Spanish names must be preferred in every case in which there is not an English name

for them.

+ Gourdon, Bull. Soc. Se. Nat. de ’Ouest de la France, (2) viii. 1908, p. 12.

+ Mem. Acad. Se. Lisboa, 1857.

THE SPANISH IBEX. 967

Sierra de Ronda from their junction in the Sierra de Tolox, and

the third one in Sierra Morena, near Fuencaliente, under the

protection of the Marquis of Mérito. Another community of

C. p. hispanica is that of Sierra Martés, Valencia Province,

mentioned by Chapman and Buck. It is very possible that

Ibexes exist also in the mountains on the opposite bank of the

river Cabriel, a name itself derived from “cabra”; but if such is

the case, these Goats cannot be considered a different colony, since

they may swim across the river, as a specimen now preserved in

the collection of the Institute of Valencia was actually seen to do.

The two remaining colonies of this race are established on the

lower Ebro, not far from Tortosa. One of these comprises the

Sierra de Card6 and the Tivisa Mountains, where an immature

male, recently received by the Madrid Museum, was obtained.

The other colony is found on the other side of the river, on

Mount Caro.

At first glance, the three subspecies of the Spanish Ibex are

much alike, their chief differences being in the horns and in the

extent of the black markings peculiar to these Goats. The

species, as a whole, may be described as a pale brown animal

with the outer side of the limbs black, a black band on the lower

part of the flanks, and a short black mane, continued along the

back by a narrow stripe. The forehead and the beard are

blackish or very dark brown, and the belly and inner part of

the limbs white. In winter pelage there is a whitish underfur,

quite absent in summer, when the general colour is browner and

the black areas become more abruptly definite. The females lack

in all seasons the mane and the black markings of the head and

body, presenting only a blackish tint on the anterior face of the

limbs *, and it is the same with young males, in which the black

areas appear in the second or third year, becoming larger and

darker as the animal grows older. It is, therefore, very difficult

to ascertain the differences between the various subspecies when

quite adult males are not at hand.

In the typical Capra pyrenaica the dorsal line appears con-

siderably broadened on the withers, frequently forming a large

lozenge-shaped blot which in old specimens spreads laterally

over the shoulders, coming downwards to coalesce with the black

of the fore limbs. The black of the hind limbs extends upwards

over the whole external surface of the thighs and on the hind-

quarters, sometimes reaching the median stripe on the rump.

The Mediterranean C. p. hispanica has these black areas con-

siderably reduced, the dorsal stripe being not broadened anywhere,

and the black of the fore limbs reaching at most the lower part

of the shoulder and the chest, whereas on the thighs it does not

reach the haunches and is narrowed to a mere band connecting

* In one of the illustrations in Chapman and Buck’s ‘ Unexplored Spain,’ repre-

senting the shooting of Ibexes in the Sierra de Gredos, the females are erroneously

depicted with a well-marked dorsal line. In reality,this stripe is,in females and

young males, very faint or quite obsolete.

66*

PROF A, CABRERA ON

Text-fig. 195.

968

Distribution of the black areas on the body of the Pyrenean (A), Gredos (B),

and Mediterranean (C) races of the Spanish Ibex.

THE SPANISH IBEX. 969

the band of the flank with the black of the leg, thus dividing the

white of the inner side of the thigh from the general brown

colour.

The Goat of Central Spain represents a stage intermediate

between typical pyrenaica and p. hispanica by the spreading of

the black areas, this colour invading the lower half of the shoulders

and covering the whole outer side of the thighs, but not reaching

the withers nor the haunches. The dorsal stripe, as in hispanica,

is of practically the same width from neck to tail.

Text-fig. 195 clearly shows the gradual decrease in the amount

of black as the species approaches the Mediterranean coast. The

variation is to some extent parallel to that indicated by Lydekker *

for the subspecies of Capra sibirica, but in the case of the Spanish

Ibex it cannot be attributed to differences in elevation nor to

the presence or absence of snow in the localities frequented by

each race. I think it interesting to note that young males of

C. pyrenaica pyrenaica, when two years old, closely resemble,

in the black markings, adult males of the Central subspecies,

the young of the latter being in turn similar to the adult

C. p. hispanica.

As to the colour of the upper surface of the body, in winter

pelage the three subspecies are much alike, the general tint

being pale brownish grey in C. pyrenaica pyrenaica, dirty

buff, more or less clouded with black towards the lower part

of the flanks, in the Ibex of Central Spain, and a paler and less

blackened buff in C. p. hispanica. The under side of the neck

is black or dark brown in the Pyrenean and Central races, and

slightly clouded with black in the Mediterranean form. I have

never seen a specimen of true pyrenaica in summer pelage,

but Trouessart describes its colour during that season as “ gris

brun foncé.” In both the two other subspecies it is pale brown,

washed with white on the flanks. here is, however, a difference

of tint between them, the Central Ibex being browner and the

Mediterranean one redder. The colour of the Ibexes of Gredos

is near the broccoli-brown of Ridgway, whereas in C. p. hispanica

it is a tint intermediate between fawn and cinnamon. Each

hair is white at the root, after which there is an undulated space

rather curiously coloured, as it has one side white and the other

side brown. This particoloured space is followed by a broad pale

band, and the hair ends in a dark brown point. Now, the

difference in colour between the two subspecies depends on the

pale subterminal space being entirely cream-buff in the Central

Ibex, and white with a broad red ring in hispanica.

Another noteworthy difference between these two Ibexes is

found in the colour of the hind border of the thighs. In the

Central form this part is buff, abruptly contrasting with the

brown of the haunches; whereas in the [bex of the Mediterranean

region it 1s coloured like the rump and the flanks, the tint being

* P.Z.S. 1901, i. p. 91.

970 PROF. A. CABRERA ON

only a little paler. Moreover, the sides of the head, grey in the

Pyrenean and Central races, are pale buff, clouded with brownish

red, in the Mediterranean form.

Text-fig. 196.

aN 4 i

SAW

ont :

Skull and horns of adult male of the Gredos Ibex (x 1).

Madrid Museum, No. 1523.

The skulls of C. pyrenaica pyrenaica and C. p. hispanica have

been described by Forsyth Major, who gives a number of dif-

ferential characteristics, most of them, I think, either merely

individual or due to age. A reliable one appears, however, to be

found in the shape of the nasals, which are more abruptly

narrowed in front in Aispanica than in true pyrenaica. In this

respect the Ibex of Gredos is nearer to the Mediterranean race,

the border of the nasals forming an almost perfect V in their

distal third.

Writing of the horns of Ibexes from different localities, Chapman

THE SPANISH IBEX. 971

and Buck* assert that “‘ examples from the two outside extremes

(Pyrenees and Nevada) most closely assimilate in their flattened

and compressed form of horn.” If by this a lateral compression

is to be understood, my own experience bears out this statement ;

but if we must understand that the individual horn in the

Pyrenean and Mediterranean races is flatter from front to behind

than in the Gredos subspecies, on careful inspection I cannot agree

with the above-quoted authors. The only reliable method for

Text-tes NOTE

Cross-sections of left horns of the Gredos Ibex (x 4).

A. Madrid Museum. Type.

B. Madrid Museum, No. 447.

(In this and the two following figures the sections represent the horn as

being seen from the tip, the front face appearing above and the inner keel to the

right side.)

investigating the true differences, and the one I have followed

with every specimen examined, consists in the taking of a cross-

section of the horn about the middle of its length. This section

is pear-shaped, and somewhat variable even in specimens from the

same locality ; but in each subspecies it is always easily referable

to a peculiar type. Now, in the Gredos Ibex (text-fig. 197) it is

* “Unexplored Spain,’ p. 144.

972 PROF. A. CABRERA ON

invariably flatter and broader than in any one of the other sub-

species, the difference being chiefly due to the great width, in the

former, of the flat upper surface of the inner keel, between its

edge and the round part of the horn. The horn sections

of OC. p. pyrenaica and C. p. hispanica, although much alike

in their more rounded and narrow contour, also differ in form.

In true pyrenaica the portion of the upper or front face imme-

diately above the keel is markedly hollow, whereas in hispanica

Text-fig. 198.

Cross-sections of left horns of the Pyrenean Ibex (X $).

A. Mainz Museum. Type.

B. Toulouse Museum.

it is slightly convex, the Mediterranean Ibex approaching

in this the Gredos subspecies. Of course, these differences can

be appreciated only in adult males, as the horns of the young in

all the races have a somewhat rounded section with a short pro-

jecting tip corresponding to the inner keel. Every specimen

I have compared has horns with at least six annulations *.

As to the curvature and direction, the horns are absolutely alike

* The popular belief that each knot or annulation on the horns means a year in

the age of an Ibex is no more true than the one assigning the same value to the

tines of deer; but, since the number of knots depends on the horn growing, it

becomes evident that many annulations, like many tines in deer horns, always indicate

an old animal.

THE SPANISH IBEX. 973

in the three races, being twisted in a half-turn of spiral, with the

tips sometimes directed inwards and downwards, as in Capra

cylindricornis, but generally pointing upwards Thus, the horns,

when seen from the front, form a very open lyre, not unlike the

lower half of the horns of the Pir Panjal Markhor. In the females

they are very short, somewhat lyrate and quite cylindrical,

differing from those of young males in the complete absence of

keel. The size of the horns in the adult male is decidedly larger

in the Pyrenean Ibex, the two other subspecies being practically

Text-fig. 199.

Cross-sections of left horns of the Mediterranean Ibex (xX #)-

A. Madrid Museum, No. 449.

B. Madrid Museum, No. 1042.

similar in this respecs. I have neither seen nor found mentioned

any specimen from either Central Spain or the Mediterranean

area with horns about one metre in length, such as frequently

occur in Ibexes from the Pyrenees. There followsa table of horn-

measurements of adult males in the three subspecies. The

dimensions of specimens marked [Ch. & B.] after the owner's

name, are converted into millimetres from Chapman and Buck’s

‘Unexplored Spain’; those of specimens marked [W.| from

Rowland Ward’s ‘ Records of Big Game.’

PROF. A. CABRERA ON

Pyrenean Thee }

Length on

outside circum- Tip to tip. Locality. Owner.

curve. ference. |

mm. mm. mm. |

1020 260 | Valibierna. Bagneres de Luchon Museum.

970 a | | North Spain. Paris Museum.

830 be: | Bee Ordesa Valley. Bordeaux Museum.

(Sin 22 675 Ordesa Valley *. Sir Victor Brooke [Ch. & B.]. |

750 | 960 | 450 (about) + Pyrenees. Mainz Museum (Type) f.

730 | 855 585 | Ordesa Valley. Sir Victor Brooke [Ch. & B.]. |

698 | 255 495 | Pyrenees. British Museum [ W.].

630 250 | Benasque. Nantes Museum.

610 | Pyrenees. Yaris Museum.

590 | | ; Querigtiena. Toulouse Museum.

Gredos Ibex.

815° | 252 690 + Gredos. |H.M. the King of Spain.

768 | 240 595 Central Spain. / MM. Chapman & Buck [W.}.

740 | 250 530 Bohoyo. Madrid Museum (No. 447).

730 220 470 + Gredos. | Madrid Museum (No. 1523).

M5 | 245 620 Gredos. | Marquis of Torrecilla.

700 165 520 Gredos. Senor Prado Palacio.

680 245 520 Las Hoyuelas. Marquis of Viana.

650 | 260 550 Madrigal de la Vera. | Madrid Museum (No. 448).

630 260 540 Las Hoyuelas. Marquis of Viana.

613 230 | Gredos. MM. Chapman and Buck

on A [Ch. & B.]. |

Mediterranean Ibex.

850 a a | Sierra Morena. | Marquis of Mérito [Ch. & B.]. |

740 200 | 525 | Sierra Nevada. | Madrid Museum (No. 449).

730 230 585 | Sierra Nevada. | MM. Chapman and Buck

| (Ch. & B.]. |

650 Be .. | Sierra Nevada. | Bordeaux Museum.

590 220 320 South Spain. | Senor E. Cortina.

590 210 470 | Sierra Nevada. | Madrid Museum (No. 921).

580 200 Sierra Nevada. | Madrid Museum (No. 1048).

580 200 sai Sierra Nevada. | Mainz Museum.

570 250 410 Sierra Nevada. | Madrid Museum (No. 1042).

655 200 | 415

Sierra Martes.

| Senor P. Burgoyne [Ch. & B.}.

* MM. Chapman and Buck give merely “ Pyrenees”? as the locality of Sir Victor

Brooke’s specimens, but according to a letter of the owner himself, published by

Count Russell (see Gourdon, Bull. Soc. Sc. Nat. de ?Ouest de la France, viii. 1908,

pp. 6-8), they were obtained in Ordesa Valley, the Val d’Arras of the French.

+ The tip of a horn is slightly broken.

Capra pyrenaica was based by Schinz on specimens in the Mainz Museum,

which are still there, as Dr. Reichenow kindly informs me.

One of these specimens

being an adult, although not old, male in winter coat, and the winter pelage being

described first by Schinz, I think it convenient to choose it as the type.

THE SPANISH IBEX. 975

T must now proceed to the complete description of the Central

Spain Ibex. His Majesty King Alfonso XIII. has specially and

graciously permitted me, at my own request, to dedicate this

subspecies to Her Majesty Queen Victoria of Spain. I have

great pleasure in doing so, in recognition both of her love of

nature and of the fact that Royal protection only has prevented

the total extinction of this splendid ruminant.

CAPRA PYRENAICA VICTORLE, subsp. n.

Diagnosis.— An intermediate form, in size and in the extent of

the black markings, between C. p. pyrenaica and C. p. hispanica,

rather browner than hispanica in the summer coat, and with horns

similar in size to those of that race, but comparatively broader

and flatter.

Colouwr.—Adult male, summer pelage: Upper surface of body

pale broccoli-brown, sometimes inclining to raw umber, and more

or less white-washed on the sides. Neck fawn-coloured, paler, and

frequently whitish, on the under surface. A black stripe, bordered

with white hairs, starts from a large black blot covering the nape

and runs along the upper part of the neck and over the back,

reaching to the tip of the tail. On the neck the hairs of this

stripe are Jong and erect, forming a short mane. On the back

the white hairs form a narrow and not very distinct light area on

both sides of the black median line. Belly and inner side of

thighs white. A broad black band, grizzled with white at the

borders, crosses obliquely the lower part of the flanks, its anterior

point going into the white of the underparts. Thighs black,

except on the hinder border, which is light buff. The black

spreads downwards, encircling the limb above the hock and

covering the front and sides of the leg and the whole foot from

some distance above the false hoofs. The fore feet and the front

and lateral surfaces of the fore limbs are likewise black, this

colour reaching the lower half of the shoulders, the chest, and the

lower part of the ventral aspect of the neck, where it is coarsely

mixed with white. Back of the legs creamy white. Forehead

seal-brown; cheeks brownish grey; the eyes encircled with

ochraceous buff, and the muzzle and the upper lip are of the same

colour. The beard brownish black, this dark tint covering also

the sides of the lower jaw to the rim of the mouth, whereas the

middle of the lower lip is dirty white. The ears fawn-coloured on

their outer aspect, yellowish white within.

Winter pelage: The main colour of the upper parts of the body

and neck turns in winter a dirty buff, densely clouded with black

on the flanks, the hairs being white at their bases and then pale

eream-buff with a brown or blackish tip, and covering a whitish

under-fur. Throat and underside of the neck seal-brown, touched

here and there with chestnut. Black areas distributed as in the

summer coat, but not so abruptly defined, their upper borders

melting into the black clouding on the sides. The colours of the

head are practically the same as in summer, the cheeks only being

slightly paler and somewhat buffy.

976 ON THE SPANISH IBEX.

Adult female: In summer pelage the general colour is an

intermediate tint between cinnamon and fawn, paling to creamy

white on the underparts, the inner side of the limbs, and the

lateral and hinder surface of the legs. ‘The muzzle and the sides

of the face cream-buff. The fore part of the legs, from a short

distance above the knees and hocks, Vandyk-brown. Tail like

the back, with a seal-brown tip. Dorsal stripe and bands of the

flanks quite absent. In winter coat the main colour is a dark

dirty buff.

Young of both sexes, in the first year: Colour like adult females,

but somewhat paler; the markings on the legs pale chestnut.

The males begin to show the dark areas of the body in the third

year, the black tint appearing first on the chest and lower part of

the shoulders.

Skull and horns.—See above for the comparison between this

and the other subspecies.

Measurements (of type, mounted).— Length from nose to root

of tail, along the curves, 1355 mm.; tail, 130; hind foot, with

hoofs, 385; ear, 120; height at shoulder, 700 *.

Skull (of paratype, Madrid Museum, No. 1523): Total length,

264 mm.; interorbital breadth, 110 ; length of nasals along median

suture, 95; greatest width of ditto, 40; upper molar series, 68 ;

lower molar series, 75. For the horn-measurements, see the

foregoing table (p. 974).

Type.—Old male in summer coat, from Madrigal de Ja Vera, on

the southern slope of the Sierra de Gredos. Madrid Museum,

No. 448.

The differences between this subspecies and both the typical

and the Mediterranean forms of C. pyrenaica, fully discussed

above, will be, I hope, clearly shown by the accompanying figures

and plates. I have considered it unnecessary to give a coloured

figure of the Pyrenean form, as there is a tolerably good one in

Lydekker’s ‘ Wild Oxen, Sheep, and Goats,’ besides the portrait of

a young male in Cuvier’s ‘ Histoire Naturelle des Mammiféres.’

Unfortunately, the same cannot be said about C. p. hispanica.

The figure published by Schinz in ‘Monographien der Siugethiere’

is anything but good, and the one in Rosenhauer’s ‘'Thiere

Andalusiens’ is not much better, the rigidity and other defects of

the mounted specimen, after which it was evidently made, being

too faithfully reproduced by the artist. As for the Ibex of

Central Spain, in Graells’ ‘ Fauna Mastodolégica Iberica,’ there

is a plate which appears to be an attempt to represent some of

the specimens in the Madrid Museum ; but they are figured in

quite a grotesque and childish way, and the colour is entirely

false. The Ibexes on the Risco del Fraile in Chapman and Buck’s

‘Unexplored Spain’ are correctly drawn; but the illustration,

being uncoloured, cannot give a complete idea of the animal.

* In an old buck from the Sierra Nevada, in the Madrid Museum, the length

from nose to root of tail is 1190 mm.; the hind foot, 305.

SUSKehEIININstel WH IANONONGE INL: tS

‘SISNHEVaGVH ISdITTMHd VOOOGVN ¢ SISNHNVaGND ISdITIHd VOOOCVIN +

‘OULD UEUIMAN 4Se\

x Hl

FARO

AT id JI6t SAd

AS. USGL PAL ILL,

West, Newman chromo.

2. MADOQUA SWAYNEI.

1. MADOQUA PIACENTINII.

ON THE SOMALI DIK-DIKS. 977

I must not close this paper without expressing my sincere

acknowledgments to Prof. Dr. Reichenow, Mainz; Dr. A.

Ménégaux, Paris; M. A. de Montlezun, Toulouse; and M. Chaine,

Bordeaux, for helping me with measurements and other information

about specimens in their respective museums.

EXPLANATION OF THE PLATES.

Prate LIT.

Capra pyrenaica hispanica.—Adult male from the Sierra Nevada, in summer coat.

Madrid Museum of Natural Science.

Prats LIT.

Capra pyrenaica victorie.—Old male (type) from Madrigal de la Vera, and adult

female trom Bohoyo, both in summer coat. Madrid Museum of Natural Science.

Prate LIV.

Capra pyrenaica victorie.—Old male from Bohoyo, in winter coat.

Madrid Museum of Natural Science.

44. On Antelopes of the Genera Madoqua and Rhynchotragus

found in Somaliland. By R. EH. Drake-Brocxmay,

MERC Selistu.Oaky., HZ.)

[Received and Read June 138, 1911.]

(Plates LV. & LVI.*)

THe DIK-DIKs.

The Somali country might justly be termed the home of the

Dik-diks,' for not only are most of the known species found there

but all the remainder at present known, save Madogua dama-

rensis, are to be found in the adjoining territories.

These little Antelopes, affording but poor sport for the big-

game hunter, have been quite overlooked by the sportsman until

the last fifteen years or so.

Dik-diks are invariably found in what might be termed the

acacia bush country, dry and arid regions where the trees seldom

grow beyond the height of bushes, and where there is sufticient

thick undergrowth to afford them protection, while at the same

time it permits of their running about freely between the bushes

and plants. In the dense aloe and sansevieria patches so

frequently met with in these parched regions, Dik-diks will nearly

always be plentiful, owing to the excellent shelter they afford for

these diminutive creatures.

Most of the small plantsand bushes supply them with food, but

the various varieties of stunted acacias are undoubtedly their

favourites. Personally I cannot recall having met with Dik-diks

in any place where these stunted acacias were not to be found.

* Wor explanation of the Plates see page 984.

978 DR. R. E. DRAKE-BROCKMAN ON

It has been doubted whether they can exist without water ;

Swayne says, talking of all the Dik-diks, that they “like to be near

water, going to drink at midday and just after nightfall,” whereas

the Somalis maintain that if a Dik-dik drinks water it will die.

In the Badminton Library, vol. 1., ‘Big Game Shooting in

East Africa,’ Mr. F. J. Jackson wrote concerning the Paa, the

local name for M/. kirkii, “it is therefore quite evident that the

juices of the vegetation on which it feeds and the dews at night

are sufficient for its requirements.”

My own experience is in entire agreement with the last-

mentioned authority, although I am not prepared to say that

Dik-diks never drink.

Those who may have observed them closely will have noticed

how a single pair will, even though disturbed, be found day after

day in much the same spot, perhaps a square acre in extent, miles

from the nearest water; here they will live all their lives if

unmolested. It is surprising to see how, after shooting the

female, the male will cling to his old haunts, living quite alone

for months together.

On one occasion I shot the female of a pair which used to live

in a small acacia grove where I frequently went to get a Francolin

for the pot ; not wanting the male I left him alone, and shortly

after left the district for seven or eight months, on the lapse of

which I returned, and while after Francolin again saw my old

friend who, not having found another mate, was living quite alone ;

there was no question about his being unaccompanied, as I saw

him every time I visited the spot. This is all the more remark-

able, as Dik-diks were plentiful enough in the locality.

There is one point I should like to emphasise before proceeding

to deal with each of the various species in turn, and that is the

presence, in a large proportion of the Dik-diks, of small white

spots on the muzzle.

Professor Linnberg has given specific rank to a Rhynchotragus

from Lake Baringo because of this peculiarity, but I venture to

think that this feature will not be found to be constant. In

specimens of M. phillips from the same district some will be

seen to possess these white muzzle-spots, while in others they are

absent.

Mapogua pHitirest Thos. (PI. LV. fig. 3.)

Phillips’ Dik-dik presents such striking variations in different

localities that for some time past I have been endeavouring to get

together a series of specimens which will show these variations

clearly and enable me to decide as nearly as possible the limits of

this interesting species.

In point of fact, were it not for the skull- measurements being so

similar, | should have been tempted to give each of the varieties

distinct specific rank, as the localities frequented by each are very

clearly defined.

Phillips’ Dik-dik was first described by Mr, Oldfield Thomas,

THE SOMALI DIK-DIKS. 979

who took as the type a specimen obtained by Mr. Lort Phillips

at Dobwein, 40 miles south of Berbera.

This spot I have been unable to locate, so can only conclude,

after carefully examining the type specimen, that it was some-

where along the Golis foothills or, perhaps, just on the top of

Mirso, which I will show is the southerly limit of the coast

or Guban variety, to which I gave subspecific rank in July 1909 *

and called MW. phillipsi gubanensis.

This subspecies is found all along the coast-belt of British

Somaliland, passing northwards into French Somaliland, certainly

as far north as Djibouti, but how much farther I have been

unable to ascertain, and eastwards towards the Mijertain country,

but how far I have not yet determined, owing to there being no

specimens available from that area.

What I propose to designate as the true I. phillipsi is the

brilliant rufous or dark cinnamon-flanked Dik. dik found through-

out the interior of British Somaliland from, roughly speaking, the

Golis Range away to the south and west into the Haud, where,

especially in the west, the animal’s flanks are so red that at

a short distance it looks rufous all over; this is in marked

contrast with the coast variety, which looks quite grey.

As one passes through the hilly country from Jig-jigga to

Harrar and also to the west and south-west of the former,

one finds another and darker Dik-dik, which in its wild state

looks of a dark red-brown colour. This is the Dik-dik to which

Mr. Neumann gave the name J/. hararensis, and so distinct does

it appear in life from J. phillips: that it fully deserves a name so

that it might not be confounded with M. phillipsi; but in my

opinion it deserves only subspecific rank, and should be called

M. phillipsi hararensis (P|. LV. fig. 2). It is to be expected that

in localities where the soils are so distinct as in the coast-belt

and the Haud one would see some variation in the coloration of

the pelage, and here in this species we have this variation very

distinctly marked; but this is not the only difference, another

equally potent factor, in the shape of altitude with its accom-

panying variations in temperature, also assists in increasing

the difference between the type species and its subspecies, by

rendering the pelage thicker.

For instance, the hair in the Guban variety is scantier and

shorter than in the Harrar and neighbourhood specimens; in

the former the individual hairs from the dorsal region of the

back measure from 13-23 mm., whereas in the latter variety they

are usually 30mm. or more. This difference renders M/. phillipsi

gubanensis a much more sleek-looking animal (EILWe ities),

The pelage of the true W. phillipsi, which is the common Dik-dik

found all over the interior of the “ Horn of Africa,” is inter-

mediate between the two.

The skull-measurements in all three are, as one would expect,

practically identical.

* Ann. & Mag. N. H. ser. 8, Vol. 4. p. 49.

980 DR. R. E. DRAKE-BROCKMAN ON

This Dik-dik is known to the Somalis by the name “ Gol ass,”

owing to its bright ved flanks. It ranges from near Djibouti in

the north and the Ennia Galla country in the west throughout

the Somali country to the east coast and as far south as probably

the 3rd parallel. A large number of the skins of this Dik-dik

are brought down to the Benadir ports for sale, from Central and

East Central Somaliland.

T might here add that I consider that little importance can be

attached to the colour of the crest, as it 1s very variable. In

some it is of a bright fulvous, in others of a dull reddish brown,

while in not a few the hairs are tipped with black. The white

eye-patch also varies in distinctness with age.

Mapboqua swayNEI Thos. (Pl. LVI. fig. 2.)

This is the smallest of the Somali Dik-diks, and is, according to

Swayne, known to the Ogaden Somalis by the name “ Guyu,” but

this name I have never heard it called myself. The word “ Guyu ”

in Somali means any living animal.

There is very little known about the exact habitat of this little

Dik-dik, owing to the fact that sportsmen have usually confounded

it with the ubiquitous MW. phillipsi. The type specimen was

bought by Swayne from a native in the town of Berbera.

IT have been on the look-out for it for some years, but have

failed to come across it either alive or dead in British Somaliland.

I first met it during my journey with the Anglo-Abyssinian

Boundary Commission in 1908, south of Ginir on the river Web,

one of the affluents of the Juba, where it was plentiful practically

all along the left bank of this river up to its junction with the

Ganale. This I took to be its westerly limit.

One of my collectors has lately brought me two specimens of a

Dik-dik from Eastern Somaliland, as far south as Obbia on the

coast ; he obtained these from a place called Gharabwein about

12 miles inland from Obbia, where they were plentiful, and

were the only Dik-diks seen, although at Eil Hur, not more than

10 miles distant, J/. phillipsi abounded and this Dik-dik

resembling JZ. swaynet was absent.

From the above I conclude that J. swaynei and the above-

mentioned species which I am about to describe stretch right

across Central Somaliland from east to west, where they are

locally distributed, and surrounded by the commoner J. phillipsi.

Neither J. swaynei nor the allied form go as far south as

Mogadishu, as my collector was unable to procure a specimen

of either species there or on the Webi Shebeleh, nor was he

able to purchase any of their skins in the market, although the

Somalis bring Dik-dik skins in thousands for sale in the coast

towns in Italian Somaliland. The only two species the skins of

which he was able to obtain were J. phillipsi and R. quentheri.

The Obbia specimens differ from those I obtained on the Web

in Western Somaliland in that there is no yellow suffusion of the

THE SOMALI DIK-DIKS, 981

grizzling on the back in the former, as there is in the latter, and

the bright rufous nose-patch is not continuous with the crest,

which may be either rufous or dull earthy brown.

In his ‘Game Animals of Africa,’ page 191, Mr. Lydekker says,

speaking of J. swaynei, “The bucks weigh but 6 lbs. and the

does even less.” Now it is an invariable rule among these small

antelopes to find the females heavier than the males. There is

usually a difference of 1 lb. between the sexes. Adult males of

Swayne’s Dik-dik weigh about 43 lbs. and the females 5 or 52 lbs.,

whereas the average weight of a buck of MW. phillipsi is 52 Ibs.

and of a doe 63 lbs. The heaviest buck of the latter species I

have ever weighed was 6? lbs., whereas the heaviest doe was

8 lbs.

I should place the range of this Dik-dik between the 5th and

9th parallels of latitude, where, although it intermingles with

M. phillips, it is not nearly so common. _ It is quite possible that

it is not to be found farther north than the 8th parallel, as

Mr. Dodds, a friend of mine, shot a large number of Dik-diks in

the Ogaden Rer Ali country, especially around Daggahbur and

Milmil, and failed to procure a single specimen of IZ. swaynei,

although both J/. phillipsi and R. guentheri were obtained.

MADOQUA PIACENTINIL, sp.n. (Pl. LV1. fig. 1.)

Although this little Dik-dik agrees, as regards its size and

skull-measurements, with J. swaynei, it differs so markedly in

coloration that I have considered it advisable to propose a new

name for it.

In M. swaynei the grizzling is very indistinct, being invariably

suffused with a dull buff or clay colour, whereas in this species

the grey grizzling is so fine and distinct that it at once attracts

attention.

The coloration of the legs in both species is similar. The

neck in M. piacentini is fairly grizzled all round, the pale throat

and buff chin patches being completely cut off from the pinkish

buff of the chest. Apart from the fine grey grizzling, the most

distinctive patches are about the head.

In IZ. piacentinu there is a bright rufous diamond-shaped nose-

patch which stands out conspicuously on the grizzled head, while

the terminal part of the long hairs of the crest in both my

specimens is of a dull creamy buff. The hairs of the crest,

however, vary so much in Dik-diks that little importance can be

attached to this feature. The ears, which are of a dark buff, have

avery distinct black edging on the outside, about 4 mm. in width

anteriorly and dwindling down to a mere edging posteriorly.

This peculiarity I have never seen in any of the other Somali

Dik-diks.

I was successful in getting only two specimens of this very

handsome Dik-dik, both males, but both present exactly similar

features. ‘They were obtained by one of my collectors at a place

Proc. Zoou. Soc.—1911, No. LX VII. 67

982 DR. R. E. DRAKE-BROCKMAN ON

called Gharabwein, within a day’s march of Obbia in the

Mijertain country, Italian Somaliland. They were inhabiting

thick aloe scrub country, and were plentiful im a locality some

little distance from water; they were very local and appeared to

be surrounded by JZ. phillipst.

The following are the measurements in the flesh of the type

specimen—a male in the British Museum, No. 356.

Hieadwandgbodiyaecca-ue oboe crcese aes A475 mm.

Wail cops ene aire Pd are tee Tien cae cae SS

Himd foGbiweamaiasithesacchcael apie. yen

A OE eee eer eee Sr omOGnE On: San ¢ suena CA He

IW ielig iG Terenas ieeunce Greene Tasers 2 lbs

Skull-measurements.

WiNeiseyll lWenaventlin " Qaqeeneusscncnesosoaconscce 92 mm.

Teiniseul Weraverslay 843 occescnaccgac ooacecacoece SOi

Posterior edge of orbit to gnathion 609 ,,

iWippericheek-teebtht..-.sen.c-cedeeas 4 Ser oS

IDeineid a Oi Weslo q- che scecone ococasseocc NGO

iBreaditiwotenasal sehen eee saa 8:9),

In comparing the above skull-measurements with a typical

M. swaynei, it will be found that they are practically identical.

This bright and beautiful Dik-dik I propose to name in honour

of my friend Mr. Piacentini, the Acting Consul-General for Italy

in Aden, through whose kindness and help I was able to send my

collector to the Mijertain country.

RHYNCHOTRAGUS GUENTHERI Thos.

This Dik-dik, known to the Ogaden Somalis by the name

‘“¢Ghussleh” or ‘‘ Gussuleh,” owing to its habit, when startled, of

dashing off in leaps and bounds, giving vent to a peculiar

whistling ery which sounds like “ Ghuss-Ghuss-Ghuss,” is chiefly

found in Western and West Central Somaliland. Swayne tells us

that he first met with it when travelling in a south-westerly

direction in the Rer Amaden country, 7. e., roughly speaking,

between the 7th and 8th parallels of north latitude; this is its

northerly limit, as Mr. J. H. Dodds tells me that during a recent

journey he made in Western Somaliland he never met this Dik-dik

until he got to Daggahbur, a well-known Ogaden watering-place

near Milmil.

Starting from Daggahbur in the north this Dik-dik passes in a

southerly and south-westerly direction for Dolo at the junction of

the Dawa and Ganale rivers, and then crossing the Juba is

ubiquitous throughout north-eastern British East Africa. I do

not think it reaches the coast anywhere, being there replaced by

R. kirkii. In Somaliland proper its easterly and south-easterly

limits are still unknown.

It probably extends for a considerable way down the Webi

THE SOMALI DIK-DIKS. 983

‘Shebeleh river, very nearly reaching to the coast in the Hawiya

country, as thousands of their skins are yearly sold in the market

in Mogadishu by the Hawiya Somalis. Theyare either caught in

native traps or shot with bow and arrow.

RHYNCHOTRAGUS KIRKIL Ginth.

Kirk’s Dik-dik, which was named so far back as 1880, inhabits

only the most southern angle of the Somali country east of the

Juba River; it, however, extends across that river and south-

wards into British East Africa as far as Kilimanjaro. How far

north it extends into Somaliland proper Lam unable to say, as the

material at present available is insufticient.

The type specimen came from Brava on the Benadir coast of

Italian Somaliland, but my collector failed to get me any specimens

from Mogadishu, so [ conclude that it does not extend farther

north than this, being replaced on the coast by JZ. phillipsi and

in Central Somaliland by 2. guentheri, of which hundreds of skins

are offered for sale in the market in Mogadishu. I have only

been able so far to procure the skins without head-skins or skulls

from that locality, so am unable to definitely state whether the

skins belong to A. guentheri or an allied form; they certainly

appear to agree with my specimens of the former.

RHYNCOTRAGUS CORDEAUXI Dr.-Br.

This Dik-dik, which was named by me last year, should really

be included among the Abyssinian Dik-diks, together with

f.erlangert, M. phillipst hararensis, and Rk. guentheri wroughtoni,

but like I. phillipsi hararensis, it will probably be found to

inhabit the north-eastern part of the Esa country.

It was first obtained by me in the bush country to the west and

north-west of Dirre Dawa, and is at present, so far as I am aware,

recorded only from the Danakil country:

I traced it as far west as the Gurgurra River, one of the

tributaries of the Hawash, and it is this latter river that marks

out its westerly limit; the northern edge of the Harrar and

Arussi plateaux mark its southern limits, but how far north

and north-east it strays I am at present unable to definitely

‘state.

This is a fine species, and most resembles in appearance

M. phillipst gubanensis, only it is a much bigger animal.

RHYNCOTRAGUS ERLANGERI Neum.

Erlanger’s Dik-dik hails from Eastern Abyssinia. As one

descends from the great Arussi plateau on the low-lying country,

towards the east, called by the Gallas ‘“‘ Gamogi,” this is the

Dik-dik one finds.

If a line be drawn on the map between Harrar in the north

-and Ginir in the south, this line will cut through an acacia-bush

67*

984 ON THE SOMALI DIK-DIKS.

country, more or less intersected by rivers and streams which drain

the great Arussi plateau; this, the Ennia Galla country, is where.

Erlanger’s Dik-dik is found.

The type specimen was obtained at Sheikh Hussein, which is.

30 or 40 miles or so east of the edge of the plateau, while my own

specimen was obtained some 20 miles to the west of Sheikh

Hussein opposite the foothills of Mt. Abu el Kassim, on the south

-bank of the river Wabi.

So far as I am aware, these are the only two specimens recorded,

so that at present its limits must remain undefined ; but I fancy I

shall be fairly accurate in suggesting that its habitat probably

lies to the west of the Harrar-Ginir line, being replaced to the east:

of it by WZ. phillipsi hararensis and to the south by IZ. swaynei.

RHYNCHOTRAGUS GUENTHERI WROUGHTONI Dr.-Br.

I have now come to a very interesting subspecies, namely

R. quentheri wroughtoni. The only specimen recorded is the type,

which I obtained on the north bank of the Wabi River among the.

foothills of Mt. Abu el Kassim.

The presence of this subspecies in a spot so far removed from

the natural habitat of the species can only be explained by its.

having at some period or other found its way up the river Wabi,

which is one of the main tributaries of the Webi Shebeleh, which

flows through the vast area inhabited by &. guentheri. Its

darker coloration and large ears are probably accounted for by its

environment, namely the dense acacia bush on the banks of the

river; the size and shape of its ears certainly point to this.

I am inclined to think that this subspecies will only be found

close to the river, as both MW. erlangeri and R. swaynei are to be

found within a short distance of it.

EXPLANATION OF THE PLATES.

Puate LV.

Fig. 1. Madoqua phillipsi gubanensis. | Fig. 2. M. phillipsi hararensis.

Fig. 3. Madoqua phillipsi.

Prate LVI.

Fig. 1. Madoqua piacentinii. | Fig. 2. Madoqua swaynei.

t=}

ON WING-CLAWS IN YOUNG BLACK-BACKED PORPHYRIOS. 985

EXHIBITIONS AND NOTICES.

June 27, 1911.

FREDERICK GILLETT, Esq., Vice-President,

in the Chair.

The Secretary read the following report on the additions made

to the Society’s Menagerie during the month of May 1911 :—

The registered additions to the Society’s Menagerie during the

month of May were 445 in number. Of these 163 were acquired

by presentation, 200 by purchase, 41 were received on deposit,

25 in exchange, and 16 were born in the Gardens.

The total number of departures during the month, by death

and by removals, was 204.

Amongst the additions special attention may be called to the

following :—

1 African Rhinoceros (2hinoceros bicornis) ¢, from Nairobi,

received from R. B. Woosnam, Esq., C.M.Z.S., for H.M. THe

Kine’s African Collection, on May 19th.

1 Californian Sea-Lion (Otaria californiana) 2, from the

North Pacific Ocean, purchased on May 10th.

2 Three-coloured Parrot-Finches (Lrythrura trichroa), new to

the Collection, from New Guinea, received in exchange on

May 4th.

A Collection of 38 Fishes, received on May Ist, comprising the

following species, all new to the Collection :—

4 Sword-tails (Xiphophorus helleri) from Mexico, 2 Fighting Fish

(Betta splendens)from Singapore, 2 Fan-tailed Cyprinodons (ivulus

flabellicauda) from Mexico, 2 Elegant Cyprinodons (Haplochilus

elegans) from the Niger River, 2 Chaper’s Cyprinodons (Haplo-

chilus chapert) from Sierra Leone, 2 Timid Cyprinodons (Haplo-

chilus panchax) from Cochin India, presented by P. Arnold, Esq. ;

2 Poey’s Cyprinodons (Rivulus poeyi) from Para, 6 Ocellated

Cyprinodons (fivulus ocellatus) from Santos, 10 Zebra Fish

(Dania rero) from Bengal, presented by G. A. Boulenger, Esq.,

F.Z.S.; 2 Freshwater Flying-Fish (Pantodon buchholzi) and 2

Gular Cyprinodons (fundulus gularis) from the Niger River, and

2 Rainbow Fish (Zrichogaster labius) from Bengal, purchased.

My. D. Seru-Smiru, F.Z.8., the Society’s Curator of Birds, ex-

hibited two immature Black-backed Porphyrios (Porphyrio melano-

notus) which had been bred in the Gardens, and remarked upon

their possession of a well-developed claw (text-fig. 200, p.986) onthe

pollex. Although these wing-claws were said to be functional

only in the Hoatzin amongst living birds, the exhibitor believed

that they were so also in the present species and also probably in

986 DR. W. T. CALMAN ON THE BRINE SHRIMP.

‘the Common Moorhen, these birds using them in climbing

amongst reeds and herbage.

Text-fig. 200.

Wing of young Porphyrio melanonotus with well-developed claw.

Mr. J. Lewis Bonuorse, M.A., F.Z.S., exhibited a pair of

Egyptian Desert-Mice (Weriones crassus) which showed a darker

and more rufous colour than normal examples. This coloration

had been artificially produced by keeping the animals in a moist

atmosphere at 80° Fahr. They were first exposed to these con-

ditions on the 7th of April, and a month later were conspicuously

darker ; after that the darkening process still continued, but more

slowly, and they appeared for some time previous to being killed

to have reached a limit to their darkening. During this same

period, and owing to the fine weather, other examples had been in

a temperature that rose during the day to 90°, falling at night to

60° or even lower. This had apparently produced no change in

their coloration. The change in the examples exhibited was:

therefore probably due rather to the humidity than to the temper-

ature of the atmosphere. Mr. Bonhote was therefore inclined to

think that the pale colour of desert animals was due to the

extreme dryness of the atmosphere rather than to any special

assimilation of their colour to the surroundings.

Dr. W. T. Catman, F.Z.S., exhibited a number of living

specimens of the Brine Shrimp (Artemia salina), bred from

Tidman’s Sea Salt. The Brine Shrimp, a small Crustacean

THE SECRETARY ON THE AFRICAN RHINOCEROS. 987

belonging to the Sub-class Branchiopoda, was found in various

parts of the world, living in salt lakes and in the shallow

ponds in which sea-water is exposed to evaporation for the

manufacture of salt. It formerly occurred in England, but had

probably long been extinct in this country. An accidental

observation recently made at the Natural History Museum

showed, however, that it wasa very easy matter to obtain a supply

of living specimens. ‘“'Tidman’s Sea Salt,” as sold in the shops,

frequently, if not always, contained living eggs of Artemia, and

an 8 °/, solution, allowed to stand for a few days, was found to

contain a swarm of nauplius larve. The first attempt at rearing

these failed owing probably to lack of food-material in the water.

The juice of green leaves pounded in a mortar and strained

through muslin was found to be a suitable food, and the addition

of a few drops of this at intervals of about a week enabled the

specimens exhibited to be raised to maturity. All of them were

females, and swarms of larve of the second (parthenogenetic),

generation had appeared.

The Secrerary remarked that on a recent visit to the Ostrich

Farm of Mr. Carl Hagenbeck at Stellingen, near Hamburg, he

had seen in the incubator fertile eggs of Struthio massaicus from

German East Africa, S. australis from South Africa, and S. mo-

lybdophanes from Somaliland, the eggs all having been laid at

Stellingen. A. Reichenow (‘ Die Vogel Afrikas,’ vol. i. p. 7) had

already described and figured certain specific differences in the

number and arrangement of the pits on the eggs of these species.

He himself had been interested to notice that the eggs of the

Masai Ostrich were much larger than those of the others, more

spherical in shape, and very smooth and porcelanous in texture.

Those of the Cape Ostrich were somewhat similar in shape and

texture, but were much smaller; Mr. Hagenbeck had informed

him that a pair of the Masai Ostrich bred by himself and sent out

to the Cape were regarded by expert ostrich farmers there as

unusually large birds. The eggs of the Somali Ostrich were

larger than those of the Cape Ostrich, but smaller than those of

the Masai species, and were markedly oval in shape witha mor gleee

less polished surface.

The SecrETARY also remarked that on his recent visit to

Mr. Hagenbeck’s Zoological Park at Stellingen, near Hamburg, he

had the pleasure of seeing a fine young pair of the common African

Rhinoceros, obtained from British East Africa, the exact locality

being unknown. The male closely resembled the ordinary figures

and mounted examples of the species, in that the skin appeared

to be smoothly stretched over the sides of the body, but the ears

were fringed with long tufts of hair. The female, on the other

hand, had no hair on the margin of the ears, and the general

external appearance was very different. At first sight it seemed

988 MR. R. I. POCOCK ON HYBRID FOALS,

as 1f it were in very poor condition, the ribs standing out through

the skin, but closer inspection showed that in reality the skin

of the flanks was disposed in thick, permanent folds, arranged

roughly like ribs. Thinking it possible that these differences

might indicate the existence of distinct races of the Rhinoceros,

on returning to London he had at once examined the Society’s

own pair of examples of this species, both of which had come from

British East Africa, probably somewhere near Nairobi. The

female, purchased in 1906, had the ears unfringed with hair, like

those of Mr. Hagenbeck’s female, but the rib-folds on the skin

were no more than indicated, although there were very heavy

permanent folds round the neck. In the male, obtained in the

current year from Nairobi as part of the King’s African Collec-

tion, the ears were fringed with hair as in Mr. Hagenbeck’s male,

whilst the rib-like folds on the skin were extremelystrongly marked,

as in the case of Mr. Hagenbeck’s female. The presence or

absence of the marginal fringe on the ears was therefore probably

either an individual or a sexual character. In the absence of

knowledge of the exact provenance of all the four examples,

nothing could be said as to whether or no the presence of the rib-

like permanent folds on the body were racial. Their existence,

however, as well as the presence of the heavy fold round the

neck, showed that it was not correct to distinguish the Asiatic

Rhinoceroses from those of Africa by the presence in the former

of permanent skin-folds. The neck-fold was almost identical ir

both, whilst, although they were differently arranged, deep body~

folds occurred in both.

Mr. R. I. Pocock, F.R.S., F.Z.S., Superintendent of the

Society’s Gardens, exhibited a photograph (text-fig. 201) of a

foal born in the Gardens on June 21st and bred between a male

Somaliland Wild Ass (Zquus asinus somaliensis) and a female

Mountain Zebra (Hquus zebra) and made the following remarks :—

“The period of gestation, dating from the day of service to the

birth, was 12? months. The general colour of the foal is sandy

fawn, the ground tint of the legs being markedly whiter. The

ears are long, as in both parents, and have a large apical black

patch, fading inferiorly in front to brown, and a brown transverse

basal stripe, running upwards mesially: a corresponding basal

stripe is present on the ear in the dam but not in the sire.

There is no white tip to the ear such as is seen in all Zebras.

The lips and area round the nostrils are black, and there is no

white on the muzzle, such as is seen in all typical Asses. Half-

way between the forehead and the muzzle there is an area

covered with many close-set narrow brown stripes and some very

faint stripes are traceable on the lower edges of the under jaw.

The mane is like that of the sire (text-fig. 202), black in the centre

and sandy fawn externally, the pale external portion showing no

trace of breaking up into evenly spaced tufts, such as are seen in

MR. R. I. POCOCK ON HYBRID FOALS. 989

Text-fig. 201.

er on

Text-fig. 202.

Somaliland Ass, the sire of the hybrid foals.

990 MR. R. I. POCOCK ON HYBRID FOALS.

all Zebras. A narrow black spinal stripe extends from the mane

to the black tip of the tail, but it is indistinct over the hind

quarters and on the upper side of the tail where the hair is long,

and recalls the dorsal and caudal mane or crest seen in the foal

of Grévy’s Zebra. There area very distinct black shoulder-stripe

and a few abbreviated stripes both in front and behind it; there

are also indistinct traces of close-set stripes on the lower border

of the neck and a deep black belly stripe; but for the rest the

body is unstriped and of a tolerably uniform sandy fawn colour

all over, like that of the sire. Both front and hind legs are

marked with strong black stripes, broader and more numerous.

than in the sire but much less numerous and more widely spaced

than in the dam. On the inside of the legs they extend just

above the knees (carpus) and hocks (tarsus), but externally those

of the hind leg reach almost to the stifle-joint (Anee), while those

on the front leg reach to about the same height. The callosities.

on the front leg are of medium size, being much smaller relatively

than in the dam, but actually of about the same size as in the

sire; and, as in the latter, there is no dewlap and the hairs along

the spine project backwards.

In general appearance this foal, which is a female, decidedly

favours the sire on account of the absence of stripes on the body

and the sandy fawn ground-colour. But it may be noted that

in the presence of the spinal stripe, the shoulder-stripe, and the

stripe on the base of the ear, it shows much greater similarity to

the typical form of African Ass, as exemplified by domestic

breeds, than to the race to which its sire belongs.

Whether the stripes will become more numerous as age

advances, remains to be seen.

This appears to be the first record of the birth of a hybrid

between the Somaliland Ass and the Mountain Zebra. Several

crosses between the domestic Ass and this Zebra have, however,

been described. The best extant account was given by F. Cuvier

(Hist. Nat. Mamm. iii. pl. 315, 1824), who accurately described

and figured a hybrid produced by a male black Spanish Ass and

a female Mountain Zebra. This animal agreed very closely with

the one just born in the Gardens, except that the ground colour,

when the animal was fourteen years old, was dark grey even on

the legs and there were distinct spots on the basal half of the tail.

The distribution of stripes was practically the same in the two;

and Cuvier’s figure shows no dewlap on the throat. The

extension of a crest of hair along the spine from the mane to the

tail and the presence of the basal stripe on the ear were noticed by

this author. The difference between this specimen and the one

born in the Gardens in colour of the body and legs is probably to

be explained by the blackness of the coat of the sire; but

St. Hilaire, who saw the foal when newly born, mentioned that

its general colour was yellowish chestnut ; at two years, however,

it was grey and this tint was retained until death. The period

MR. R. I. POCOCK ON HYBRID FOALS. 99}

of gestation was a fortnight over twelve months, being shorter by

one week than in the case of the animal born in our Gardens.

Another hybrid of the same kind is figured on plate 28 of the

volume on Horses in Jardine’s ‘ Naturalist’s Library.’ The

animal seems to have been much more copiously striped than our

specimen. Many strong but abbreviated stripes are shown running

along the saddle behind the withers; the leg stripes extend

farther up the quarters and the body, head and neck are marked

with indistinct wavy and close-set stripes. The croup, however,

seems to have been self-coloured sandy fawn like the rest of the

body. In this case it is not known which of the two species was

sire and which dam.

In the ‘ Knowsley Menagerie, p. 73, two hybrids between

Mountain Zebra mares and Asses of African descent are

described. One sired by a Maltese Ass is represented by the

right figure of the pl. lvii. The ground-colour of the body is

dark grey, that of the belly and legs white; the face below the

eyes is tan, and there is no white on the muzzle; the ears are

large with the tip broadly black and a broad stripe near the base ;

the mane is grey and unstriped. There are no stripes on the

face; but the body and neck are covered with narrow wavy

stripes which break up into small spots upon the hind quarters ;

the shoulder-stripe is very distinct, broader than the others and

forked; the belly is unstriped, but the legs are distinctly striped.

Except for the presence of spots on the croup, this animal is rather

like those figured in the ‘ Naturalist’s Library.’

The second specimen, sired by an ass of unspecified breed, is:

described as grey with an indistinct cross and a few narrow dark

stripes on the shoulder and fore legs [nothing is said about the

hind legs]; the upper side of the tail, which is elongate and

tufted, is stated to be slightly banded ; and the ears are said to

be moderate. Attention is drawn to the presence of scarcely any

stripes on this animal, which is contrasted on that account with

the one sired by the Maltese Ass. Although according to the text:

and the legends of the plates, this animal is unfigured, the

description applies very closely to the specimen represented by

the left-hand figure of pl. lvii., which purports to be a hybrid

between a male Hemione (=Onager) and a female Zebra.

This figure represents an animal sandy fawn in colour with

the legs striped, but somewhat sparsely, only slightly higher than

the level of the belly which is lighter than the flanks; the head

is fawn with some narrow close-set rufous stripes in the middle

about half-way between the eyes and the nostrils; the muzzle is

ashy grey, without any white; the ears are moderately long, with

a black tip; the mane is black in the middle, white externally,

the white hairs showing a decided tendency to break up into tufts

as in all zebras and quaggas; continuous with the black mane is.

a black spinal stripe; there is a distinct black shoulder-stripe,

followed by several abbreviated and less distinct stripes upon and

992 MR. R. I, POCOCK ON HYBRID FOALS.

just behind the withers. In the text this animal is merely

described as having the shoulders and legs banded. Its re-

semblance to the hybrid just born in the Gardens is very close

and, indeed, surprising considering the difference in coloration

and general character between the Onager and the Wild Ass of

Somaliland. Jin our hybrid, however, the ears are decidedly

longer and the mane shows no signs of being striped. Apart from

these differences, there is little to choose between the two. But

these differences are sufficient to make one hesitate in adopting

the suggestion that the animal described as a hybrid between an

Onager and a Mountain Zebra may have been a hybrid between

a Domestic Ass and a Mountain Zebra.”

[Supplementary note added July 4th, 1911.]

Since the above given account was read a hybrid foal (text-

fig. 203) between the same Somaliland Wild Ass and a Chapman’s

Quagga (1. quagga chapmanni) has been born. The period of

gestation was twelve months and four days from the date of

Text-fig. 203.

Chapman’s Quagga and hybrid foal.

service. The foal is very like the Mountain Zebra hybrid. The

ears, however, are smaller, as was to be expected from the

relatively smaller ears of the Quagga dam. ‘The ground colour of

the body, too, is a little paler, while the legs are only slightly

paler than the body. The leg stripes are less distinct and less

numerous and the insides of the legs are scarcely banded. The

MR. R. I. POCOCK ON HYBRID FOALS. 993

shoulder-stripe is shorter and simpler ; the spinal strip is black and

zig-zag over the saddle, and very faint, short, narrow, close-set:

stripes are detectable on each side of it along the back and

elsewhere on the body, the hairs of these ‘ ghost-stripes’ being a

shade darker and glossier than the spaces between them. The

spinal crest is not so pronounced; the apical black patch on the

ear is smaller, and the basal stripe, although much narrower,

is decidedly blacker. Faint narrow stripes are present on the nose

as in the first described hybrid; the lips and area round the

nostrils are black, and the two patches above the nostrils which

are dark tan in the dam are dark greyish brown in the foal,

and somewhat sharply contrasted with the narrow lighter area

that intervenes between them and the general sandy grey hue

of the nose. ;

The left-hand figure of pl. lviii. of the ‘ Knowsley Menagerie’

represents a hybrid between a Domestic Ass and a Burchell’s

Quagga (1. quagga burchelli). This hybrid appears to differ from

the one above described in having a few quite distinct brown

stripes on the body and very few stripes on the legs. The

ground colour of the legs, moreover, is markedly whiter. Since

Burchell’s Quagga differs from Chapman’s in having the legs

whiter and almost stripeless, the difference in the coloration of

the limbs between the two hybrids is not a matter for surprise.

In connection with the two hybrids born in the Gardens and

those that have been bred elsewhere previously between the same

species, namely H. asinus and L. zebra, or LH. quagga, irrespective

of the exact race or breed of the species, the following points

may be noticed. The white muzzle of H. asinus is eliminated.

In other respects asinine characters are dominant over zebrine

and quaggine characters, as is shown by the absence of stripes

on the mane, the disappearance of the white tip to the ear, and the

suppression, partial or complete, of the stripes on the neck, head,

body, and quarters. Even when stripes are visible on these areas

they do not resemble in width and arrangement those of Mountain

Zebras and Quaggas, but are more suggestive of the narrower

stripes of Grévy’s Zebra (L. grevyi), as in the case of some horse-

quagga hybrids bred by Prof. Cossar Ewart. Examples of typical

#H. asinus carry a dark basal patch on the ear, a spinal and

a shoulder stripe, and very frequently distinct or indistinct bars

on the legs. The spinal stripe is also frequent on all Asiatic asses;

and the Mongolian species (4. hemionus hemionus) sometimes

shows traces of a shoulder-stripe and of leg-stripes on the knees

and hocks as well. The Tibetan Wild Ass (Z. kiang) has a large

dark basal patch on the ear, and spinal, shoulder and leg stripes

are commonly visible in many horses. Although the basal ear-

patch, the shoulder stripe, and spinal stripe are absent in typical

examples of H. asinus somaliensis, no one doubts that this race

is descended from asses bearing the marks in question. Inter-

mediate forms indeed, with very narrow spinal and shoulder stripes

and a dusky patch on the ear, connect the Somaliland Ass with

994 MR, F. BE. BEDDARD ON

ordinary domestic varieties; and in all the many foals born

in the Gardens between our Somaliland Ass and domestic asses of

English and Spanish breeds, the ear-patch, shoulder and spinal

stripes were present as in the dams.

The above stated facts suggest that, with the possible exception

of the ear-patch in horses, the shoulder and spinal stripes as well

as the stripes on the legs have been lost comparatively recently

by the species that are without them.

PAPERS.

45, Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea. By Frank HE. Bepparp, M.A.,

F.R.S., F.Z.S., Prosector to the Society.

[Received May 28, 1911: Read June 27, 1911.]

(Text-figures 204-215.)

Il. On two New GENERA OF CESTODES FROM MAMMALS.

The following communication to the Society contains an

account of the anatomy of two species of Tapeworms, contained

in the collection of Cestoidea belonging to the Society. I

feel it necessary to refer them to new genera, and the facts which

I shall presently bring forward will I hope justify my so doing.

The first of these species is certainly near to Zhysanosoma, and is,

as I think, clearly to be referred to the same genus as a species

recently described by me * and temporarily reterred to the genus

Thysanosoma. I pointed out, however, in that paper a number

of differences from other tapeworms referred to the genus

Thysanosoma, and intimated that it probably would be necessary

to institute a new genus for the worm. ‘This expression of

opinion is justified by the discovery of a second species which

is clearly of the same genus. The second species considered in

the present paper was placed by me temporarily, and in a report to

the Prosectorial Committee, in the genus Sertiella. It shows,

however, some aitinities to Tenia (sensu stricto) and seems on

account of various peculiarities, which in due course will be

pointed out, to demand the creation of a separate genus.

T shall commence with an anatomical description of the new

species allied to Thysanosoma; this I place in a new genus which

I propose to call Thysanotenia.

Thysanotenia lemuris, gen. et sp. n.

An example of a Black-headed Lemur (Lemur macaco), which

had lived in the Gardens three years and three months, contained

* See No. I. of this series, P. Z.S. 1911, p. 651.

NEW MAMMALIAN TAPEWORMS. 995

two perfect examples, and a number of large fragments, of a

tapeworm which I refer to the above new genus and species.

There is, as it appears to me, no doubt whatever that these

tapeworms belong to the same genus as a species which I have

recently described in the “ Proceedings” * of this Society as Thy-

sanosoma gambianum from the Gambian Pouched Rat (Cricetomys

gambianus). ‘The external characters are in very near agreement.

The present species agrees with Zhysanosoma gambianum in the

unarmed head, in the fact that the segments increase in length

posteriorly but are never longer than broad, that the genital

apertures are unilateral, and that the eggs are enclosed in a

numerous series of paruterine organs in the fully mature

segments. This set of characters is sufficient to refer the present

species, as well as Thysanosoma gambianum, to the subfamily

Thysanosomine of the Anoplocephalide.

Text-fig. 204.

Thysanotenia lemuris, about twice natural size.

Of the two specimens which possessed a scolex, the larger

(text-fig. 205) measured at least four inches in length, and was

probably longer as it ended abruptly and without a definite termina]

* Above, p. 651.

996 MR. F. E. BEDDARD ON

proglottid, such as was visible in the younger, smaller, example (text-

fig. 204). Furthermore, among the fragments was one measuring

nearly five inches in length, which may, for what I know to the

contrary, have belonged to the first mentioned individual, since it.

consisted entirely of mature proglottids, which were of greater

length individually than the terminal proglottids of the specimen

Text-fig. 205.

Thysanotenia lemuris, a second specimen. Above and to the left is the scolex

more highly magnified; below and to the right, two segments of another

individual showing the papillae which bear the genital pores.

described and figured in this communication. The longest

proglottids of the mature fragment were rather more than 3 mm.

long, and rather less than 3 em. wide at the posterior end of the

proglottid, which is considerably wider than the anterior end.

The proglottids have no fringes posteriorly such as are

present in Thysanosoma and provide the reason for the name

given.

NEW MAMMALIAN TAPEWORMS. 997

The scolex (text-fig. 205) is of moderate dimensions, not very

minute, and quite visible as such by the naked eye. It is

more or less spherical, and is marked by some black pigment

below the level of the suckers. A pigmentation of the

suckers is not uncommon among the Cestoidea, and it seems

occasionally to pervade the scolex also. The suckers have a

circular orifice, and present no marked features on examination

with a lens. ‘They lie some way below the summit of the scolex,

and are directed laterally ; there is no trace, that I could see, of

an upward direction. The smoothly rounded upper end of the

scolex shows no hooks. ‘The suckers lie near to the lower end of

the head, which is followed by a distinct neck of 2 or 3 mm. in

leneth, in which the strobilisation was not apparent. The

strobila are at first much broader than long, and each overlaps

its successor at the edges. In the larger specimen, the dimensions

of which are given above, the proglottids did not attain to an

equality in length and breadth until about three inches or so

from the scolex. ‘They never attain to a length which is greatly

in excess of their breadth, as will be apparent from the measure-

ments which I have given above. It is noteworthy that in the

smaller of the two individuals which are described here, the very

few last segments alone showed a length equal to their breadth.

In this worm (text-fig. 204) the last proglottid had an oval form,

and was of less diameter than those which immediately preceded

it. This form is usual for the last proglottid, and serves to

emphasize the fact that this specimen was complete and had

not begun to cast off proglottids. The ripe proglottids differed

from the anterior ones in their greater thickness, and in the fact

that their surface was marked by a longitudinal furrowing,

caused as I should imagine by unequal distention with embryos.

The shape of these proglottids also was different from that of

those lying in front, in that they were hourglass-shaped, with,

however, as has already been mentioned, a greater diameter

posteriorly than anteriorly. The genital pores are unilateral

without exception from end to end of the body. They are not at

all visible until the segments have begun to increase in length,

and are perhaps less obvious upon the very distended posterior

proglottids. The pores are particularly obvious in this species,

on account of the fact that they are borne upon slender processes

of the Body-wall, as will be more specially described later in

connection with the generative organs. These genital processes

lie near to the anterior border of the segments.

If the above account of the external characters of this tape-

worm be compared with my account of the external characters of

Thysanotenra gambiana, it will be seen that the two species,

though agreeing in many features, nevertheless show several

points of distinction, which enable them to be fully defined

as distinct species from these external characters only. It is

particularly to be noted that the mdividual of the two examples

of Thysanctenia lemuris which I regard as a not fully developed

Proc. Zoou. Soc.—1911, No. LX VIII. 68

998 MR. F, E. BEDDARD ON

worm is like the adult Thysanotenia gambiana in its characters.

Thus the proglottids do not increase at all appreciably in length

until the very end of the body, those anterior to this being

broader than long. It is, however, clearly a difference between

the two species that there are in Thysanotenia lemwris a much

longer series of proglottids of greater length than diameter, and

that the conical projection of the body which bears the generative

pore is more pronounced. Furthermore, while the species described

in the present communication has along neck, there is no neck in

Th. gambiana, the strobilisation commencing immediately after

the scolex. Finally, 7h. lemuris has black pigment in the scolex,

which is wanting in Zh. gambiana. There are thus several

obvious differences between the species, which as I think prevent

any confusion between them.

In transverse sections (text-figs. 206, 207) it was only ‘possible

to see a single water-vascular tube on each side. This is of con-

siderable diameter, though it fluctuates in size from place to place

Text-fig. 206.

Thysanotenia lemuris. Two sections across a proglottid, one of whicn (on the

right) shows the genital papilla. .The half only of each complete section is

shown. R&.S, Receptaculum seminis ; Sy.d., sperm-duct ; Z., testes scattered in

parenchyma; Va., vagina; W., excretory tube.

and is connected with its fellow in each proglottid by a transverse

tube, which hes near to the posterior boundary of the proglottid.

The single water-vascular tube of this species appears to represent

by its position the innermost of the two tubes found in 7hysano-

twnia yambiana. I could find no network of minute tubules arising

from these, such as are obvious in the last mentioned species.

Genital organs.—On the whole, the genital system of the species

dealt with in the present communication is not very like that of

Thysanotena gambiana. ‘There is, however, a very important

NEW MAMMALIAN TAPEWORMS. 999

point of agreement, in the similarity of the enclosure of the eggs

in numerous paruterine organs, which perhaps outweighs the

various dissimilarities which will be dealt with in considering the

testes and the ovaries and their ducts. The testes occupy in

Thysanotenia lemuris a quite different position in the body from

that which they occupy in Zh. gambiana. They lie in the former

species entirely between the water-vascular tubes, and in the

medullary region of course of each proglottid. Like the other

organs of the genital system, they commence to be visible rather

late in the body, thus contrasting very markedly with the

conditions obtaining in the second genus described in the present

paper. The testes lie mainly in the posterior region of each

proglottid, but in front of the transverse water-vascular vessel.

They form continuous rows each only one deep, and not, except

perhaps here and there, at all crowded. The testes are also

extended anteriorly to quite the front end of the proglottid on

either side of the ovaries.

The cirrus sae is divided, but not sharply, into two regions.

The terminal part which encloses the cirrus is narrow, and this

tube expands posteriorly into an oval vesicle of much greater

dimensions. ‘The whole structure has in fact much the shape of a

soda-water bottle. The hinder part of the sac is about on a level

with the receptaculum seminis and the sac itself is gorged with

sperm, the whole of the available space being filled with a mass of

sperm. It is encircled by a thick layer of muscle fibres which

run longitudinally to the longer axis of the sac, and form a

continuous coat passing in this direction to all round the sac

when it is viewed in longitudinal section, that is in transverse

sections of the proglottid. At the posterior end the thick

muscular layer is interrupted for a minute space, to permit of

the entrance of the sperm-duct which narrows greatly at its

entrance into the cirrus sac, or rather into that part of the

cirrus sac which is specialised as a vesicula seminalis. This

narrow region of the vas deferens immediately widens out

into a long tract of duct, which may be regarded as a second

vesicula seminalis. This tube is wide, quite as wide as in species

of Lertiella, and is coiled upon itself as it passes back towards

the opposite extremity of the body. This region of the sperm-

duct would be, if unwrapped from its coiling, of considerable

length; I found it to be gorged with sperm. The cirrus itself

was generally protruding from the genital aperture. It is not

large and has the usual structure of this organ. The difference

which it shows from the sperm-duct is very striking, and

furnishes an argument in favous of those who would hold that

the cirrus is not merely the end of the sperm-duct which is

capable of protrusion. It seems in this species at any rate to be

a structure independent of the sperm-duct. There is a certain

resemblance between the bottle-shaped cirrus sac and the bottle-

shaped receptaculum seminis and the end of the vagina. Both

of these tubes lie to the same side of the iateral excretory vessels —

68*

1000 MR. F. E. BEDDARD ON

if we are to regard the ovary as ventral, they lie to the dorsal

side.

The vaginal pore lies behind the opening of the male duct

into the genital cloaca. The vagina is a perfectly straight, delicate-

walled tube as in so many tapeworms. Posteriorly and close to

the lateral water-vessel, the vagina expands into a somewhat pear-

shaped receptaculum seminis, from the wider, posterior end of

which the vagina emerges again abruptly and runs a curved oblique

course towards the ventral side of the body. This latter region

of the vagina is of the same calibre and appearance as the

terminal section which opens into the genital] cloaca. The

swollen receptaculum seminis (text-fig. 206, p. 998) has on the other

hand thick glandular walls. The ovary and the yolk-gland he

anteriorly in each proglottid and very nearly in the middle of

the proglottid, verging however to the pore side, the position

being therefore quite different from that which characterises

Thysanotenia gambiana. The ovary is not large, neither is the

yolk-gland.

Whether a uterus exists as a definite structure at any period in

the development of the sexual organs, I am not able to state with

certainty. But I am disposed to think that a distinct uterus

does not exist. At the most, it must have a very brief existence,

for I can find nothing intermediate between eggs scattered in the

parenchyma and in the paruterine organs. J found nothing like

what has been described in Zhysanosoma. In this genus there

are stated to be outpocketings of the uterus round each of which

is formed a paruterine organ. I found in the present species a

condensation of the parenchyma round eggs or groups of eggs,

precisely as I have described in Thysanotenia gambiana. These

latter became more marked, and were then to be described as paru-

terine organs. In fact, the state of affairs which characterises the

genus Thysanotenia is to be looked upon as a further stage in

the development of such a genus as Oochoristica with the inter-

mediate formation of a uterus dropped out, or at least rendered

of very little importance. It is also like Davainea

Text-figure 207 represents a transverse section through a fully

mature proglottid of this worm. It may be compared with a

corresponding illustration of the other species of the genus,

Thysanotenia gambiana*, when certain differences will be

apparent coupled naturally with fundamental points of agreement.

In both species, the greater part of the medullary region of the

segment is occupied by the numerous paruterine organs, which

are only one layer deep. They extend between the excretory

vessels and up to those vessels on either side. In Thysanotenia

lemuris, however, the transverse row of paruterine sacs is at most

thirteen to sixteen, while they are much more numerous in

Th. gambiana. Furthermore, ina given section the greater number

by far—very often all—of the paruterine organs are seen to be

* P. ZS. 1911, text-fig. 158, p. 658.

NEW MAMMALIAN TAPEWORMS. 1001

- without a contained embryo or embryos. This does not mean,

of course, that these are paruterine organs which contain no

eges. It is simply an expression of the fact that in the present

Text-fig. 207.

Thysanotenia lemuris, transverse section through ripe proglottid, showing the

longitudinal external furrowing of the proglottid. e, paruterine organs, in only

two of which were eggs to be seen in this particular section; ¢, excretory

tubes.

1002 MR. F. E. BEDDARD ON

species each paruterus contains fewer eggs (or embryos) than

is the case with Thysanotenia gambiana. In no case have I

seen in a given section more than three embryos within

a single paruterine sac in the species which forms the

subject of the present communication. In reference to this par-

ticular I may compare text-figure 207 with the text-figure of

my paper dealing with ‘“ Thysanosoma ” gambianwm*. The

paruterine organs themselves were of about the same size in the

two species. The presence of so few embryos in a single par-

uterine sac produces naturally a very distinctive appearance which

at once distinguishes the two species from each other. In each

paruterine organ of 7h. lemuris it was possible to distinguish

a cortical and a medullary region of different appearance. ‘That

this was not possible in the other species is probably to be

accounted for by the large number of embryos which filled them.

It will, I think, be admitted that this tapeworm presents char-

acters which will not fit in with those of any known genus. It

contradicts indeed the definitions of families as given by Ransom,

at any rate to some extent; for I should be disposed to place

the genus in the neighbourhood of TZhysanosoma in the

family Anoplocephalide ; and yet this family is characterised by

the absence of a neck, present in the genus which is now under

consideration. The only other position in the series which this

worm could occupy, as I think, is in the subfamily Paruterine of

the family Hymenolepidide ; but in this subfamily the paruterine

organs are limited to one or two, and there is nothing like the

numerous organs met with in both of the species which I describe

here under the generic name of Vhysanotenia. This latter

reason as well as the unilateral genital pores prevent the inclusion

of the species in the genus Stilesia.

This new genus, Thysanotenia tT, may be thus defined :—

Thysanotenia, gen. nov.

Moderately large tapeworms, four to six inches in length and

three millimetres in breadth. Scolex unarmed, with laterally placed

suckers. Proglottids posteriorly as long as or slightly longer than broad.

Genital pores unilateral, borne upon a projection of the body near to

the anterior end of proglottids. Hxcretory tubes wide, one or two (and

these lying side by side) on each side, with or without ramified

branches. Reproductive organs a single set in each segment. Testes

numerous, or very numerous. Uterus a narrow transverse sac or

absent. Many paruterine organs in ripe segments. LEygs without

pyriform apparatus. Adults in Mammals.

* Loc. cit. text-fig. 159, p. 659.

+ Inasmuch as there are no fringes to the proglottids posteriorly such as occur in

Thysanosoma, the name selected is rather a misnomer. I have, however, used it

pore for the purpose of fixing what I consider to be the systematic position of

the worm,

NEW MAMMALIAN TAPEWORMS. 1003

Accepting the above as the generic characters of the genus

Thysanotenia, the two species may be thus defined :—

(1) Thysanotenia gambiana F. E. B.

Thysanosoma gambianum Beddard, P. Z. 8. 1911, p. 651.

Length about six inches, greatest diameter 6 millimetres. Segments

never longer than broad and only a few at the posterior end as lon;

as broad. Genital papilla not very conspicuous. Two excretory tubes

on each side, the inner of the two the larger, placed laterally

to each other ; a network of fine tubules connected with these. Testes

in two groups, the larger lying on the side furthest from the

genital pore, occupying the space between the two excretory tubes of

that side and a little beyond on each side. Ovary and yolk-gland on

pore side lying between the two excretory tubes and «a little to the

immer side also. Sperm-duct narrow or coiled, with a small vesicula

seminalis. No receptaculum seminis ; vagina opens into a terminal

muscular sac. Puruterine organs cach with many embryos. Uterus

a transverse sac.

Host, Gambian Pouched Rat (Cricetomys gambianus).

(2) Thysanotenia lemuris, sp. n.

Length four to sia inches with diameter of three millimetres. Seg-

ments at end of body rather longer than broad. Genital papilla very

conspicuous. One ewcretory tube on each side of body corresponding

to the innermost of the two present in Th. gambiana. No network of

tubules connected with this. Testes scatter ed through posterior part of

the body and anteriorly to the sides of ovary. Ovary and yolk-gland

submedian in position, slightly to pore side of segment. Sperm-duct

wide and coiled after issuing from cirrus sac. Lreceptaculum senrinis

present. Paruterine organs with only three or four embryos in each.

A uterus not formed (2). —

Host, Black Lemur (Lemur macaco).

It is obvious from the above definition and from what has been

said in the course of this paper that the two species, which I assign

to this new genus Thysanotenia, differ from each other in a

good many points, and perhaps may be considered to merit generic

separation.

Anoplotenia dasyuri, gen. et sp. n.

Atabout the same time, 7. e. from February 9th—11th, 1911, three

examples of the Tasmanian Devil (Dasyurus ursinus) died in the

Gardens ; only one of them was found to contain tapeworms, and

these occurred in that example in very great numbers. The

specimens belong, as I believe, to a new species, and I am also

disposed to form for it a new genus, which is to some extent allied

to the Anoplocephalide, but also, in the form of the uterus,

suggests Zwinia, sensu stricto. These facts have suggested the

1004 MR. F. E. BEDDARD ON

generic name which I here propose. I considered after a rough

examination of the worm that it might possibly be referred to

the genus Bertiella, and suggested this in a report to the

Prosectorial Committee; I am now convinced that it cannot be

included in that genus and that its characters will not allow of

its inclusion in any known genus. The movements of the living

worm were particularly active. The worm is not a long form and

has a rather unusually large head, as will be gathered from the

Text-fig. 208.

Anoplotenia dasyuri, enlarged about five times.

accompanying figure (text-fig. 208). I could find no trace of any

hooks or of any terminal sucker or other structure in the

rostellar region. The four suckers are unarmed and of large

size.

The scolex is well marked off from the strobila, although there

is no obvious neck—that is to say, the segmentation begins

apparently at onee. This is brought about not only by the

NEW MAMMALIAN TAPEWORMS, 1005

actually large size of the head but by the fact that the first strobila

are much narrower than those immediately following, and thus

an apparent constriction occurs which emphasizes the distinction

of the head. In other cases the conditions are apparently the

same; but an examination with a lens shows that though a

constriction at about the same distance from the rostellum exists,

the formation of strobila exists beyond this point and has invaded

the hinder region of the actual scolex. It is a little difficult

therefore to assert that a neck is or is not characteristic of

Anoplotenia dasyuri. The occasional commencement of

strobilisation immediately behind the scolex is not unsuggestive

of Oochoristica*, with which genus the present has some likeness,

and there is in the same way a kind of hint of a commencing

pseudoscolex.

The body of the worm is about an inch in length or sometimes

rather longer; but it never grows to a great size. The segments

which immediately follow the head are very short; but this

region of the body is not long, perhaps a couple of millimetres, and

contains but few segments, in some cases not more than a dozen,

in others rather more. The segments increase gradually in

length and towards the end of the body come to be three or four

times as long as they are broad. In the hinder segments the

genital pores are sometimes quite obvious when the worm is

examined with a lens, and irregularly alternate in position from

one side of the body to the other. In transverse sections the

body is apt to be hourglass-shaped owing to the greater thickness

of the lateral edges of the body due to the bulging caused by

the unusually large cirrus sacs. ‘This is certainly the case with

the shorter and flatter anterior proglottids.

In transverse sections, through the anterior region of the body

in mature segments, which are, however, not long and distended

with ova, the layers of the cortex can be readily distinguished.

The cortex (text-fig. 209) is of about the same diameter as the

medullary portion. It is distinguishable into a much thicker

outer layer of longitudinal fibres, and a much thinner inner

layer of longitudinal fibres. The two are separated by delicate

transversely-running fibres, of which there are also a set

within the inner layer of longitudinal fibres and thus bordering

upon the medulla. The inner layer of longitudinal fibres is

particularly conspicuous for the reason that several are closely

grouped into a bundle, of which bundles there is only a single

row, as is shown in text-figure 209. This arrangement of the

muscular fibres is only apparent in the more anterior segments.

It ceases to be obvious in ripe proglottids such as that represented

in text-fig. 213 (p. 1012) where the body is gorged with eggs.

In these segments however, where, as will be pointed out at

length presently, the eggs are partly contained in a uterus and

partly scattered singly or in groups through the medullary

* See P. Z. S. 1911, p. 628.

1006 MR. F. E. BEDDARD ON

parenchyma, a new set of muscular fibres becomes apparent which

I have not observed in the anterior segments and which therefore

if really present are not so plain in those segments. The fibres run.

across the medullary parenchyma in a dorgo-ventral direction and

apparently belong to the cortical layer of circular fibres. In

text-fig. 213 some of these fibres are seen to cross the medullary

parenchyma and then to join the longitudinal series of cortical

fibres. Possibly these fibres assist in the rupture of the

proglottids to expel the eggs, and their presence may also account

for the particularly active movements of this species which have

been referred to above.

Text-fig. 209.

Anoplotenia dasyur?, transverse section through part of body-wall. I, two delicate

layers of circular fibres between which is a special layer of longitudinal fibres

grouped into bundles. 7, testis.

The excretory vessels are two on each side and are accurately

superposed, 7. e. dorsal and ventral. Later on in the body only one

vessel is very plainly visible on each side, and this is of greater

calibre than anteriorly.

The gonads and their ducts appear very early in the strobila of

Anoplotenia dasyuri. They commence to be visible within less

than 1 mm. of the head immediately after the anterior set of very

short segments. ‘The segments, when the ducts are first visible,

NEW MAMMALIAN TAPEWORMS, 1007

are absolutely crammed with apparently mature (certainly very

nearly mature) testes. The ducts showed no signs of specialisation.

The vas deferens only expanded slightly and gradually into the

elongated terminal sac, which is so much specialised in the mature

segments to be described later, This immature condition of the

ducts persisted for only six segments. Thereafter the cirrus sacs

were nearly or quite fully developed. In these anterior segments

the ovaries were not so forward in development as the testes.

Text-fig. 210.

Anoplotenia dasyuri, longitudinal section of proglottid. A., shell-gland; Ov., ovary,

below which is seen the cirrus sac ; 7’., testes ; U., uterus, between which and the

cirrus sac are seen the coils of the vas deferens cut transversely ; W., transverse

excretory tube. The posterior part of the proglottid is above.

The ovaries of Anoplotenia dasyuri lie posteriorly in the

segment but anteriorly to the vitelline glands, and when fully

developed are large and distinctly double. They have the very

common bushy form and occupy a good deal of the posterior

region of the segment. I did not observe the ovaries to be fully

developed until the first segment, in which the cirrus sac is also

fully developed and in which the uterus has begun to appear.

When the uterus has become so far developed as to fill the greater

part of the proglottid, the ovaries quite distinctly lie in continuity

with the masses of not fully mature eggs which here fill up the

chambers of the uterus.

The vitelline glands are quite evident close to the posterior

border of the proglottid; they do not extend out so far laterally

as do the ovaries.

The vagina of this worm is, as is so usual in the group, a quite

straight tube for the greater part of its extent, that is to say it

is not coiled. It lies behind the cirrus sac and opens into the

1008 MR. F, E. BEDDARD ON

commencement of the genital cloaca, perforating the muscular

pad. Its walls are not distinctly cellular and they stain deepiy,

both of which features are very common in these animals.

Distally, the vagina may be easily followed until it opens into a

well-marked receptaculum seminis, which is very large and con-

spicuous in Anoplotenia. In young segments its course is straight

across the segment; in older ones it passes straight to the

middle line in an oblique course, then bends back along the middle

line of the segment. This sac lies almost exactly in the middle

of the body and is absolutely circular in transverse sections of the

proglottids. ‘The vagina enters it on the ventral surface and

leaves it again at an exactly corresponding point further towards

the non-pore side of the proglottid. Thus the tube leaving the

receptaculum has to be followed for a short distance in sections

before it can be ascertained whether it is the distal or proximal

part of the vagina. The receptaculum was gorged with sperm. It

is not spherical as might be imagined from its circular contour in

section, but narrows to the more slender tube at one end which is

beyond the exit of the vagina.

Towards its point of opening into the genital cloaca, the vagina

lies parallel to the cirrus sac, in some cases being less oblique

in its course than in other proglottids. It has a very thick

muscular wall, that is to say thick relatively to the size of its very

small lumen. This muscular coat is double, an inner longitudinal

and an outer circular layer being present. Opposite the actual

point of opening into the genital cloaca, the vagina suddenly

widens into a very small sac lying closely adpressed to the

muscular pad which forms part of the wall of the genital cloaca ;

from this sac, a narrow tube passes at right angles to the

rest of the vagina and perforates the muscular pad. The terminal

sac of the vagina is seen to be filled with sperm. It is related

perhaps to the lateral orifice of the cirrus within the invaginated

pouch of the cirrus sac: this when protruded forms a bulbous

extremity, near to which the orifice would pour its contents

into, quite fill and perhaps even somewhat dilate, this terminal

sac of the vagina. This matter is, however, more fully dealt with

under my description of the cirrus sac and penis (on p. 1014).

I am disposed to think that the receptaculum seminis is no

more than a dilatation upon the vagina, for it could easily owe its

shape to mere gorging with sperm, and its walls appear to be like

those of the rest of the vagina and to have lost their cellular

character. It is at any rate greatly disguised in them as in so

many other tapeworms. As Gough has lately pointed out,* this

non-cellular appearance is preceded by a distinct wall of cells. If

the receptaculum seminis be as I suggest merely a local swelling

of the vagina, it is clearly quite different in its nature from the

receptaculum described above in Zhysanotenia lemuris. ‘This

latter is most obviously a distinct and definitely specialised region

* “Tapeworms of the subfamily Avitelline,”’ Quart. Journ. Micr: Sei. vol. lvi.

NEW MAMMALIAN TAPEWORMS. 1009

of the female tubes. For it is apparent in less mature proglottids

and has there walls of a different character from the slender

Text-figc. 211.

Anoplotenia dasyuri, two vipe proglottids viewed as transparent objects.

L., masses of eggs at posterior end of proglottid; R., reticular portion of uterus ;

Sp.d., cirrus sac.

1010 MR. F. E. BEDDARD ON

vagina connected with it. And, moreover, in these younger

proglottids its shape cannot be due to any distention by sperm,

for the sac was quite empty of sperm. It might be con-

venient to restrict the term receptaculum seminis for cases

of this kind, and not perhaps to name specially the distended

region of the vagina in which the sperm is chiefly massed in

other forms.

Text-fig. 212.

Anoplotenia dasyuri, transverse section through ripe proglottid.

O.,ova; U., cavity of uterus.

The uterus of this tapeworm is visible very early in the body

in correlation with the early development of the organs of repro-

NEW MAMMALIAN TAPEWORMS. 1011

duction generally. I found, in fact, that the uterus was quite

recognisable in the first segment which had a fully developed

cirrus pouch, and that segment was one of the earliest to have

attained an appreciable length and lay hardly a millimetre

behind the scolex. In this segment the uterus showed (in a

longitudinal horizontal section through the anterior region of the

body) a rounded form stretched in the direction of the transverse

section of the body and thus rather oval in outline, and it

occupied precisely the median region of the proglottid. In this

particular uterus I found no ova. A segment or two further

back the uterus is already larger, but it still has the form of a

more or less oval sac, extending in these segments towards the

pore-side and having thus become eccentric in position. The

eccentricity, however, is not very strongly marked. In these

segments the uteri were full of ova.

"The uterus in these segments lay near to the posterior boundary

of the proglottid and was transversely elongated in form; it

was distinctly posterior to the strong muscular cirrus sac. The

uterus possessed a distinct epithelial wall that was of sufficient

thickness to show itself in all my sections (cf. text-fig. 210, p. 1007).

Later, the epithelial wall is not obvious, but the cavity has plain

boundaries and can be recognised as a definite cavity and not

merely a system of irregular lacune.

In the posterior ripe proglottids the uterus undergoes some

changes which are not altogether easy to follow and to correlate.

When the elongated, fully ripe proglotiids are examined mounted

in glycerine, the eggs are seen to be arranged throughout them

in a way which differs slightly in different proglottids but

is as arule at any rate on the same plan. ‘The eggs occur in

clusters and strings which give the appearance of a retiform

uterus. In this, at times, a median string of eggs giving off

lateral branches may be recognised. And though these lateral

branches join here and there and thus make a network, the

general appearance given in such segments is that of the uterus

of 7 cenia, Which is characterised ‘rs a median stem and lateral

branches. Very commonly the ova are more thickly clustered

together in the posterior region of each segment. There is, I

think, little doubt that if the worm were examined only in this

way, the uterus would be pronounced to be reticular. A study

of sections, however, leads to a rather different interpretation of

the arrangements visible in solid preparations. In some trans-

verse sections, such as that illustrated in text-fig. 212, the whole of

the interior of the proglottid is occupied by the uterus and the

contained masses of developing eggs. These appear to lie in a

large undivided cavity, which I take to be the uterus. This

region corresponds to the posterior part of the segment, where as

already mentioned the eggs tend to become massed.

In other sections through the same proglottid as that which

has just been referred to, the conditions observable were different.

There are, as is shown in the accompanying figure, eggs and

1012 MR. F. E. BEDDARD ON

groups of eggs which correspond to the thinner strings of eggs

in proglottids which are viewed as solid objects rendered trans-

parent by glycerine. These groups vary in size, but it would

appear that they are imbedded in the parenchyma of the medul-

lary region and are not contained in cavities—that, in fact, there

Text-fig. 213.

Anoplotenia dasyuri, transverse section through ripe proglottid.

O., ova, contained in uterus (U.); O1, eggs scattered through parenchyma.

is no uterus here at all. It may, of course, have been present

and have disappeared. In intermediate proglottids the uterus

forms a series of cavities which apparently intercommunicate and

thus constitute a network. In these cavities the eggs are not

mature—at any rate, the membranes have not yet appeared.

This system of cavities fills up a great deal of the available space

NEW MAMMALIAN TAPEWORMS. 1013

and the testes become much restricted and tend to disappear.

The most noticeable part of the medullary tissue left is a pro-

jection from the region of the cirrus sac lodging the coil of the

vas deferens.

It would appear therefore that the uterus in this genus

Anoplotenia rather combines the characters of that organ in

several other genera than presents us with a new type. That it

passes through a simple saccular stage is not perhaps a fact of

any great moment; for that occurs in many genera. But it 1s

undoubtedly reticular at one time, and, contrary to what is found

in such cases, the reticular stage is not permanent.

Text-fig. 214.

RS.

Anoplotenia dasyuri, longitudinal section through proglottid, showing the branched

and reticular uterus with ovain smaller and larger clumps. B.S., receptaculum

seminis; W., excretory tube.

The ultimate condition of the uterus does not fall definitely

within any of the types used by Ransom in his table of arrange-

ment of the genera of Teenioid Cestodes ; and the condition of this

organ in Anoplotenia dasyuri is really one of the chief reasons

upon which I base its generic distinction from other forms.

The ¢estes of this species are very numerous in those anterior

segments in which they are at their full development. They

are pressed closely together and overlap and appear to fill all of

the available space left between the ovaries and other organs of the

proglottid. As the latter are posterior in the segment, the testes

are mainly anterior. Both in longitudinal and sagittal segments

the testes can be seen to be limited in their occurience only by

Proc. Zoot, Soc.—1911, No. LXIX., eo

1014 MR. F. E. BEDDARD ON

the cortex. In correlation with their large numbers, the testes

are of small size. They cannot, obviously, be said to be either

dorsal or ventral or anterior or posterior in position.

A good deal of the anterior part of each mature proglottid is

occupied by a large coil of the vas deferens which forms a larger

mass than in many tapeworms figured or known to me at first

hand. Although this coil lies anteriorly in the segment on a

level with the lar ge cirrus sac, it does not touch the anterior

boundary of the segment. In front of it there is to be seen a

considerable heap of testes. The coil is generally in close contact

with the cirrus sac. I found no vesicula seminalis in this species,

but the coiled tube gets wider when ripe.

The cirrus sac and its contained structures are rather re-

markable in this species and much more complicated than in

many other species, including the three that I have already

described in my former communication *. As already stated,

the organs of reproduction appear very early in the chain of

proglottids. It is, however, not for some segments that the

cirrus sac is fully developed. The fully developed cirrus sac

coincides with the first appearance of the uterus. As already

mentioned, in considering the external characters of this worm,

the cirrus sac when mature is so large as to cause an appreciable

bulge in the segment. It therefore entirely fills the medullary

region of that part of the segment where it occurs, as seen in

a transverse section. In such sections it may also be seen

that the cirrus sac and the genital cloaca together (of which

a description follows) occupy about one-third of the entire breadth

of a proglottid.

The cirrus sac is nearly if not quite spherical in shape, and

consists of an outer coat and of an inner mass of tissue. The

outer coat is not very thick and is muscular, the fibres no doubt

serving to compress the sac and thus bring about the evagi-

nation of the cirrus. The internal tissue of the cirrus sac is

a tissue in which the cirrus itself is embedded, and it entirely

fills the sac save where it is traversed by the cirrus. It contains

many nuclei scattered fairly closely throughout it, and delicate

fibres which may be occasionally seen to possess a distinctly

retiform arrangement. It seems to me to be a soft tissue which

is an elastic packing material transmitting to the cirrus the

contractions of the external muscular coat of the cirrus sac. In

some sections indeed the nuclei can be seen to be more compressed

and regularly arranged in lines in the immediate neighbourhood

of the eversible cirrus sac. This might seem to argue some

contractility possessed by the tissue forming the core of the

cirrus sac.

The cirrus itself is peculiar and complicated in structure. It

consists of two parts. First of all there is the part which lies

immediately within the cirrus sac and which is perfectly con-

tinuous with the vas deferens, and shows no sudden differences

* P.Z. S. 1911, p. 626.

NEW MAMMALIAN TAPEWORMS. 1015

of structure that I can detect from the vas deferens. It lies in

a loose coil of only two or threeturns. Followed distally, this tube

Text-fig. 215.

Anoplotenia dasyuri. Cirrus sac with penis in various stages of retraction.

In the upper figure the penis is completely retracted, in the middle figure it:

is completely protruded; in the lower figure it is incompletely protruded.

A, muscular pad on anterior side of genital cloaca; B, muscular pad on posterior

side of genital cloaca which is perforated by opening of vagina (Ja.) ; C., cirrus;

P., penis, on the posterior side ot which, as is shown in the middle figure, the

cirrus opens; Sp.d., vas deferens.

(SiS)

1016 MR. F. E. BEDDARD ON

is seen to open into an invaginated sae which lies pushed into the

‘solid core of the cirrus sac, as is shown in the text-figure accom-

panying this description. This sac is irregular in form with

crumpled walls, and it is surrounded by a layer of particularly

stout muscular fibres, which lie therefore within the cirrus sac

and form a differentiated portion of its core. These muscular

fibres lie loosely round the invaginated sac. The wall of the

latter is rather thickened to form a pad lying on the anterior

side, and close to this the cirrus opens into it, their cavities

becoming continuous. The opening into the sac is opposite to

the pad, and therefore on the posterior side of the sac.

The cirrus sac does not open directly on to the exterior, but

through a genital cloaca which is itself much complicated. <A

horizontal section through the whole structure is represented in

text-fig. 215. The genital cloaca may be divided into three or

four regions; the invaginated penis (as we may term the in-

and evaginable sac into which the cirrus opens) is continuous

with a wide but narrow cavity of quite as great a diameter

(antero-posterior) as the cirrus sac itself, but very narrow from

side to side. After this comes a tubular cavity surrounded

by a very thick layer of circular fibres which are perforated, as

already mentioned, on the posterior side by the vagina which

here enters the genital cloaca. In horizontal sections such as that

represented in text-fig. 215, this layer of muscles appears as two

strong muscular pads, that on the posterior side being longer

from side to side than the one opposite. After this the canal

widens a little for the terminal part of its course and its walls

are rather crumpled, there being a particularly deep recess”

immediately after the sphincter region which precedes it. A

considerable portion of the cirrus sac can be evaginated, sometimes

more and sometimes less. These differences also are shown in

text-figure 215. In some cases only the anterior side of the

cirrus sac is protruded, which in such a case barely reaches

the external orifice of the genital canal. In other cases

much more is protruded, and the cirrus itself is drawn down

into this penial protrusion which extends well beyond the

external orifice. In this latter case, however, which represents

the extreme of what I have seen in my sections, the actual opening

of the vas deferens is not at the tip of the protruded penis, but

at the side and within the genital canal. I may not perhaps have

seen instances of extreme protrusion. It is noteworthy that the

orifice of the vas deferens is on the posterior side of the penis and

thus corresponds to the vaginal orifice. Possibly a complete

extrusion of the penis takes place in cases of cross-fertilisation.

The following are the general characters of this species * :—

Head quite unarmed, of rather large size and with four unarmed

suckers, Strobilisation begins at once, there being no “neck”: the

jirst two or three strobila wider than those which immediately follow

* IT do not attempt to discriminate between generic and specific characters.

NEW MAMMALIAN TAPEWORMS. 1017

and of same diameter as or wider than head, thus forming «a

rudimentary pseudoscolex. The proylottids increase very rapidly to

a consideruble length, the posterior beiny longer than broad and

becoming detached. Not more than ten or a dozen anterior short

proylottids. Genital orifices single and irregularly alternate.

Exeretory tubes posteriorly one on each side of body, that of one side

being as a rule wider than that of opposite side; in anterior segments

two on each side. Testes very numerous, filling up the whole space

left by other organs in proylottid. Vas deferens coiled. Cirrus sac very

large and spherical and somewhat peculiar in structure, with an

eversible sac reaching the exterior through a much differentiated

genital cloaca. Ovarres posterior in segment and with vitelline glands

posterior to these. Vagina straiyht and narrow, opening posteriorly

to currus sac; a receptaculum semuus present. Uterus at first a

simple sac, later a reticulum, and later still part of the uterus remains,

while other eggs are imbedded singly or in groups in the medullary

parenchyma. Hggs without V-shaped apparatus.

It will be, as I think, evident from the résume of the characters

of this species just given, that it cannot be referred with any

confidence to any one of the really known genera of the Tetra-

cotylea. The convenient table giving a key to the various genera

used by Ransom in his memoir enables one to refer the species

from Dasyurus ursinus to the neighbourhood of Oochoristica, Tenia

(s.s.) or Bertiella. Of the latter genus several species are known

from Marsupials; but they are not known from the present

genus, and appear to be nearly limited to the herbivorous (at

any rate Diprotodont) genera, i.e. Phalanger, Phalangista, and

Phascolarctos*.

These species, however, are certainly not congeneric with that

which I describe in the present paper. They agree with the

generic definition given by Ransom7, who doubtless took them, as

well as the species of Bertiella from Apes and Rodents and Birds,

into consideration when formulating his definition. The worms

studied by myself show the following important differences from

Bertiella as defined by Ransom :—The strobilisation is different,

the posterior strobila being much longer than broad ; the genital

canals pass between the dorsal and ventral excretory vessels ;

the testes exist throughout the segment save where space is

occupied by the ovaries etc.; the uterus is of a totally different

character; the cirrus sac is also totally different from anything

figured in Bertiella. I do not feel able therefore to refer this

species from Dasyurus ursinus to the genus Bertiella.

I am of opinion that the present genus is nearer to the genus

Oochoristica. The latter genus actually occurs in carnivorous

Marsupials but in Neotropical forms, in fact in Didelphys, and not,

however, so far as I am aware, in Australian Marsupials. The

* See Zschokke in Semon’s ‘ Reise,’ Jena 1898, for B. obesa and B. semoni; and

the same author, “ Neue Studien an Cestoden aplacentaler Siugethiere,” Zeitschr.

wiss. Zool. lxv. 1899, for B. edulis and B. sarasinorum. Also Janicki, “ Die

Cestoden Neu Guinea’s ” in Nova Guinea, Livy. v., 1906, p. 281, for B. rigida.

+ Loe. cit. p. 62.

1018 MISS R. HARRISON AND PROF. S. J. HICKSON ON

general form of the body and the segmentation is not unlike in

the two genera; and especially to be noted is a resemblance in

the scolex. In the present genus as in the Oochoristica from

Tamandua tetradactyla, described by myself * some months since,

the strobila, as it were, invade the scolex. On the other hand, the

early disappearance of the uterus and the imbedding of the ova.

singly in the medullary parenchyma is a character of Oochoristica

which distinguishes it from the genus which I propose to call

Anoplotenia. The peculiar cirrus sac and the very complex

genital cloaca are points in which Anoplotenia differs from all the

genera with which I here compare it.

There now remains the genus Tenia (sensu stricto) to which the

present species shows a certain amount of likeness in the uterus,

which is rather pronounced in certain proglottids. There is in

fact occasionally a quite distinct median stem with branches.

Tenia, however, has an armed rostellum which is sometimes not

armed asin 7’. (eniarhynchus) saginata, where the hooks drop out

early and are replaced by a sucker-like structurey. There is

nothing of this kind in the present species, which moreover bears

no such close likeness to Zenia saginata as would warrant its

inclusion in the same genus or subgenus. Another genus in

which the uterus has a marked median stem and lateral branches

is Catenotenia,: the species of which occur in the mouse and in

the squirrel. in this genus, however, the testes and ovaries have

a different position from that which is met with in the tapeworm

dealt with in the present memoir, and the relation of the genital

duct to the excretory tubes is also different.

46. Some Madreporaria from the Persian Gulf. By Ruts

Harrison, Oxford §. With a Note on the Memoir and

some Further Notes on Pyrophyllia inflata by SyDNEY

J. Hickson, M.A., D.Sc., F.R.S., F.Z.S.

[Received May 19, 1911: Read June 27, 1911. ]

(Plates LVIT. & LVIITI. |! and Text-figures 216-221).

This collection of Madreporarian corais was made by Mr. F. W.

Townsend, and entrusted to me for identification and description

by Professor Hickson. I should like to take this opportunity of

thanking Professor Hickson for putting this interesting piece of

work in my hands. My thanks are also due to Professor Bourne

for allowing me to carry on the work in his laboratory and placing

all its resources at my disposal, and for help and advice during the

* P. Z.S. 1911, p. 627. I ought to have mentioned in that paper that something

of the same kind appvears to occur in O. rostellata (see Zschokke, Zeitschr. wiss.

Zool. vol. Ixxxini. 1905).

i Cf. Bronn’s “'Thierreich,” Vol. iv. Abth. B. p. 1720.

{£ Janicki, Zeitschr. wiss. Zool. 1906, vol. Ixxxi. p. 505.

§ Communicated by Prof. S. J. Hickson, E.R.S., F.Z.S.

|| For explanation of the Plates see p. 1044.

1% Bots) MSIL, PW IOWA,

MADREPORARIA FROM THE PERSIAN GULF.

I, & S, Ill, il. ISVAN,

pis Paina ee roy

poster

MADREPORARIA FROM THE PERSIAN GULF.

Bale & Danielsson Lttim;

CORALS FROM THE PERSIAN GULF. 1019

progress of the work; to Professor Sollas for permission to use

his apparatus, which enabled me to grind sections, photograph and

reconstruct a wax model of Zrematotrochus zelandic ; to Professor

Jeffrey Bell for permission to examine various Madreporaria in

the British Museum; and to Professor Stanley Gardiner for the

loan of a large number of his specimens of Heterocyathus

. equicostatus for comparison with the present collection.

The collection consists of examples of the following species :—

Family GuyNnrp&.

Pyrophyllia inflata Hickson [25].

Family FLABELLIDS.

Flabellum magnificum v. Marenzeller [28].

Family TurBInoLup.

Heterocyathus cequicostatus Milne-EKdwards &

Haime [29].

Heterocyathus heterocostatus, sp. n.

Paracyathus cavatus Alcock [1}.

Trematotrochus zelandie Dunean {14}.

Agelecyathus persicus Duncan |14 |.

Family Funerp.

Fungia patella Milne-Edwards & Haime [29].

Family HupsaAMMUID&.

Heteropsammia aphrodes Alcock |1}.

Dendrophyllia sp. ¢

All these specimens are in the Manchester Museum. The

occurrence of a recent Zrematotrochus is worthy of special

attention. Hitherto, the genus has been known from seven

fossil and one recent species from Australia and Australasian seas,

and it is remarkable to find it appearing in so remote a locality

as the Persian Gulf. More remarkable still is the fact that this

very species has already been described from Cook’s Strait,

New Zealand, by Professor Martin Duncan under the name of

Conocyathus zelandie. 'The resemblance of the coral I have been

examining to the figures and description (so far as it went) of

this Conocyathus was so striking, that it occurred to me that

possibly Duncan had overlooked the perforations of the wall,

characteristic of a Trematotrochus. Owing to the kindness of

Professor Jeffrey Bell I have been able to re-examine the original

type specimens in the British Museum, and the result of this

examination has been to convince me that the corals are absolutely

identical. ‘The British Museum species have not been so care-

fully cleaned and dried as those in the present collection, and in

places the intercostal furrows are somewhat choked up by sand

and grit; but that perforations do exist, exactly similar to those

of the Persian Gulf species, | have no hesitation in stating ; in

parts of the corallum they are clearly visible, but unless one were

1020 MISS R. HARRISON AND PROF. S. J. HICKSON ON

expecting to find them it would be very easy to overlook them,

and it is not altogether surprising that Duncan should have done

so. Had he observed them, I venture to think he would not have

called the coral a Conocyathus. In his ‘ Revision of Families and

Genera’ he places Conocyathus, Trematotrochus, and Turbinolia

together as closely allied forms; indeed the perforations of the

theca of Trematotrochus form the only feature which separates it

from Conocyathus. Now that perforations have been observed in

Conocyathus zelandiw, we must either amend the definition of that

genus, or else remove this species to the genus Zvrematotrochus.

The existence of pores is a character of such importance that the

latter course seems to me advisable, and in future Conocyathus

zelandie should be known as Z'rematotrochus zelandiwe. The

specific name is unfortunate and apt to be misleading, as is

bound to be the case when a specimen is given a name denoting

the locality in which it was originally found, and subsequently

appears in other parts of the world. A further consideration of

the structure and systematic position of this coral will be found

in the systematic part of this paper.

Family GUYNIID&.

PyROPHYLLIA INFLATA Hickson [25]. (Pl. LVII. figs. 8-11;

Pl. LVIII. figs. 18, 19.)

About sixty specimens of this species were obtained on a

gravelly bottom at a depth of 156 fathoms in the Gulf of Oman.

Professor Hickson has added some further notes to his original

description of this species at the end of this memoir (p. 1039).

FLABELLUM MAGNIFICUM v. Marenzeller [28.] (Pl. LVII.

figs. 1-3.)

Corallum fan-shaped, wall and septa very thin and delicate.

Numerous rootlets descend vertically downwards, each such

rootlet communicating with two interseptal chambers on opposite

sides of a septum, and firmly fixed on a mass of mud and

serpulid tubes. Calice oval in outline, but constricted in the

middle of the short diameter; practically semi-circular at the

ends of the long axis. ‘Two specimens, measuring as follows :—

I. II.

1a (ytd Ni Ave peeer eee ral, eh aan se aenre 2Suailicee Hy ees SELON TT Ta 15 mm.

N Rio ar 6) LOOM CVENTACIES socooausocnvooonndve seconn eon 60 . 40

Calice ry Shoitidiamerers ‘ 4

‘ Greatest width ..............000000. 40 ,, 26 ,,

Least width? ees fees. aoO) see Zowes

Numibertofiseptaywen.ea ee eer cee eee rela 96 |

As 6p DUNT SEI, saocodondenocoouocedenaact 24, 24 |

CORALS FROM THE PERSIAN GULF. 1021

Costee faintly indicated throughout entire height of the corallum

which is also marked with fine wavy transverse accretion lines.

Septa in six systems of six cycles of which the sixth cycle is

incomplete, while there are present a few rudimentary septa of

a seventh cycle. Those of the first three cycles equal and reaching

the columella, edges vertical, entire until within about 4 mm. of

the columella where a few coarse denticulations may be present ;

beset with minute blunt spines arranged in transverse and radial

rows. Septa of lower cycles become successively smaller. Columella

parietal, only slightly developed.

Locality. Telegraph Cable, Persian Gulf. Depth not recorded.

In the larger specimen there are 175 septa arranged as

follows *:—

Bhi " | Number of super-

Number of 7 unter of sels of numerary septa of

septa. 1 cycle wanting. 7th cycle.

Chambers] aier.cseereeet 31 | 3 2

Be 2) tec eae 25 7 none

a) Wubeene ecdane 30 2 none

EL. 26 | 6 none

a 5 30 | 2 none

Gite shove ike 33 5 6

otaliaesee scenes 175 25 8

This coral differs from v. Marenzeller’s species in its smaller size

and the presence of numerous rootlets. Gardiner [19] has shown

that the presence or absence of such rootlets is a variable

character, and he has described rootlets in Flabellum rubrum, a

species in which they had not previously been recognized.

v. Marenzeller’s species was made for a single specimen, and as

the agreement between the calicular and septal characters of the

specimens under consideration and those of the type are so

similar, it has been considered advisable not to separate them.

Family TURBINOLIID4.

Genus HETEROCYATHUS.

The genus Heterocyathus has been critically examined by

Gardiner [20], who had at his disposal a very large number

of specimens. He absorbed the species of Semper and Rehberg

in a single variable species originally described by Milne-

* The septa are considered as being divided into six chambers, bounded by

primary septa; the chambers are considered in rotation beginning with one to the

right-hand side of a directive septum.

1022 MISS R. HARRISON AND PROF. S. J. HICKSON ON

Edwards & Haime [22] as Heterocyathus cwquicostatus ; to this

Bourne [6] has since added Stephanoseris rowsseawi. .

In the large collection of corals from 8. Africa, Gardiner

recognized two types with the following diagnostic characters :—

Type I. Coste equal in size, rounded and covered with low

granules (Pl. LVIILI. fig. 12), small intercostal spaces ; base smooth,

low granules, only traces of cost ; theca with thin upper edge,

never more than 1 mm. above columella; septa in four cycles,

thick with narrow interseptal spaces, average exsertness 1°5 mm. ;

cycle i. broader and more exsert than cycle 11., 11. than ii1., iv.

more exsert than ii1., higher on either side of i. than of 11. Sides set

with low ridges, edges not toothed ; commonly certain ones, or all,

coloured black; pali before all cycles, large and conspicuous ;

columella a mass of rods decreasing in size and height from

the pali towards the centre of the axial fossa, densely packed

together.

Tyre 11. Coste of cycles i., ii., and ii. larger than iv., which

consists of a row of separate granules rougher and higher than in

Type I, with broader intercostal spaces (Pl. LVIII. fig. 13); base

roughly granular, costee sometimes extending on to it; theca with

thin upper edge, generally 2 to 3 mm. above the top of the

columella; septa in four cycles with a tendency to have some of

a fifth cycle represented, thin with broad interseptal spaces,

average exsertness 3 mm.; ridges on sides conspicuous, edges

toothed towards centre of calice; all the same colour, white or

some shade of grey; no proper pali or columella; septa iv. fuse

with septa ili., and these again with 11. ; septa 1. generally separate,

but in the centre fuse with the rest forming a mass of trabecule

covered by fine points which run up along the septa of cycles 1. to

ui. for some distance, almost like fine teeth.

In describing the collection of Mr. J. J. Simpson and

Dr. Rudmose-Brown from Burma, Miss Poole [24] recognized

the Gardiner’s two types, and added a third type with a fifth

eycle of septa, four crowns of nodular pali, and a deeper fossa.

I have been able to re-examine the collections described

by Professor Gardiner and Miss Poole, and, so far as the former

collection is concerned, the two types are sufficiently different,

in my opinion, to be regarded as two species. Gardiner himself

has. described them as “ two perfectly distinct modes of growth,

almost two varieties.” I have searched in vain for intermediates,

and although I had no difficulty in picking out the two specimens

which the author describes as doubtful, they are both too much

broken and corroded to afford any clear evidence that they

vepresent a form intermediate between the two types. The

general facies are entirely different. In Type I, the tendency is

for the base to be broader than the calice in correlation with

rounded granular coste, thick septa, practically no fossa, and a

papilliform well-developed columella. In Type I, the tendency

is for the coral to taper away toa point from a circular calice ;

such is the form of the only known example without a commensal

CORALS FROM THE PERSIAN GULF. 1023

Aspidosiphon; the flat base of the normal specimen is never so

large as the calice, and this shape is invariably in correlation with

alternating coste, thin septa, a deep fossa, and an ill-defined

columella.

The characters of the coste of the two types are very definite.

Fig. 12 on Plate LVIII. represents two adjacent cost of Type I,

low broad ridges, covered all over with fine uniform granules; the

intercostal furrows are small and shallow. Fig. 13 on Plate LVIII.

represents two adjacent cost of Type Il: on the right is the

type of costa which corresponds to septal cycles i., 1i., and 111.,

a prominent exsert ridge, on which the granulations tend

to become regular transverse bars; on the left is a costa corres-

ponding to septal cycle iv., slightly less exsert, and beset with

very irregular coarse granulations; the intercostal furrows are

deep. These marked differences, always in correlation with the

differences of the calicular characters to which reference has

already been made, are too consistent to retain both forms in the

same species, True, there are many examples of Type I which

have an alternation of broad and narrow coste, but such coste are

always alike in being lowand uniformly granular; the alternation

of coarsely granular coste and much exsert narrow coste is never

discernible. In some, however, there is a tendency for the coste

of Type I to become more exsert and more coarsely granular at

the calicular margin, but this generally occurs where the whole

growth of the coral has been distorted by being fixed to an

abnormally large shell, and 1t might be thought that this distortion

had influenced the form of growth. Indeed, it suggested the

possibility that the shell on which the coral fixes itself, and the

position which that shell assumes within the actual body of the

coral, might influence the mode of growth to such an extent

that not only the general shape but the character of the septa

and coste might be controlled. This, however, is not the case.

There are well-defined examples of Types I and IL both fixed

on exactly similar shells: in some the Aspidosiphon in corals of

both types is coiled horizontally forming a flat base, in others the

corals are fixed to shells which le sometimes horizontally,

sometimes vertically within the coral zoophyte, and there are

always examples of both types harbouring Aspidosiphons in shells

which assume either position; so that the species of shell or

the position it assumes exercises no influence on the manner of

growth of these two well-defined types. Such types vary about

two distinct modes, with a slight overlapping of the extremes of

variation ; and are not themselves the extremes of a single growth

mode. The relation of the species to each other is represented in

text-fig. 216.

Of some 252 individuals in the South African collection, there

are 225 of Type I and 19 of Type IJ,and 8 which have characters

commonto both types. The characters which separate the two types

cannot be Bees and given numerical values ; consequently

in text-fig. 216 A the point about which the two growth-modes

1024 MISS R. HARRISON AND PROF. S. J. HICKSON ON

vary is purely arbitrary, but the number of individuals which vary

about these points is a real number, and represents the two

distinct growth-modes of two different species. If the two types

were varieties of a single growth-mode, the variation might be

expressed by such a diagram as text-fig. 216 B, but that would

represent a condition of affairs wholly contrary to the numerical

facts; for the intermediates, which in reality are less than a

thirtieth of the whole number, here represent the greater number

of forms.

Text-fig. 216.

() 20 40 60 80 100 420 140 160 180 200 220 240 260

ty) 20 40 60 go 100 120 140 /60 180 200 220 240 260

A.—A diagrammatic representation of two species which vary about two distinct modes,

the extremes of which converge towards one another. The abscisse give numbers

which vary about each mode respectively.

—A diagr: atic representation of a single species varyin 0 i inar ,

B.—A diagrammatic representation of a singl ar about an imaginary mode

Type I is undoubtedly the H. equicostatus of Milne-Edwards

& Haime, although, as I shall show, there are considerable

variations within the type. Type II is probably identical with

Verrill’s [42] H. alternatus, a species which has escaped the

notice of several authors. This species possesses all the

characters which separate Gardiner’s Type II from Type I,—the

base smaller than the disk, with a slight constriction above it, and

then walls spreading obliquely outwards to the edge of the disk,

CORALS FROM THE PERSIAN GULF, 1025

alternating costz, paliform teeth exsert before all septal cycles

except the last, and an ill-developed papillose columella which

scarcely rises above the surface of the broad shallow central fossa.

When Miss Poole stated that the forms in the Burmese

collection belonged to three different Types, two of which were

identical with Gardiner’s Types I and II,she had not the advantage

of seeing the actual specimens, and was in error. All the

Burmese forms belong to Gardiner’s type I, but they show great

variation, and in order to justify their inclusion with H. cequi-

costatus three types must be recognized. These types may be

called A, B and C to avoid confusion, but it must be understood

that they correspond with Miss Poole’s Types I, II & III.

Type A=Type I as defined by Gardiner. There is normally a

single small aperture on the basal surface in addition to the

opening of the Aspidosiphon chamber. In a few, this additional

aperture is absent, whilst in others, two or at most three such

apertures occur.

Type B is a much lower flattened form without a fossa; a

tendency for the coste to alternate; four septal cycles, the

exsertness of the septa of the fourth cycle on either side of the

primaries is very marked, and the joining over the tertiaries

and secondaries gives a characteristic star-like appearance ; the

columella is less well-developed, it is a compact trabecular mass,

and not distinctly papilliform as in Type A; the pores of the

endodermal canals are more numerous, and are not restricted to

the base, but are distributed irregularly round the lower part of

the corallum.

Type C is a taller form, the base tends to equal the disk in

diameter, and the walls are nearly perpendicular ; a fifth cycle of

the septa is present ; there is a distinct fossa, and the centrally

depressed columella is a trabeculate mass asin the previous type;

the pores of the endodermal canals are disposed in a ring a little

below the calicular margin.

In defining these three types, I have retained Gardiner’s Type 1

and: Miss Poole’s Type C in order to avoid confusion as far as

possible ; but it must be borne in mind that Type B as here

defined is very different from Gardiner’s Type IT.

Practically the whole of the South African collection belongs to

Type A, the Burmese collection belongs to Types B and C; the

specimens Miss Poole described as belonging to Type I should be

included in Type B. The Ceylon collection described by Professor

Bourne is somewhat intermediate between Types A and B. These

three types will embrace the species of Semper, Rehberg, and

Alcock*.

H. parasiticus Semper [38]. Intermediate between Types A

and B.

H. philippinensis Semper [38]. Two types. PI. xx. fig. 12 is

Type B. PI. xx. figs. 13 & 14, Type C.

* T have not considered H. sulcatus and H. lamellosus Verrill, and H. cochlea

Gmelin, as I have not been able to obtain first-hand reference to these species.

1026 MISS R. HARRISON AND PROF. S. J. HICKSON ON

H. pulchellus Rehberg [37]. Intermediate between Types B

and @. General facies, fifth septal cycle and depressed

columella, Type C. Alternating costze and distribution of

lateral pores, Type B.

H. oblongatus Rehberg [37]. Type C.

H. wood-masoni Alcock [1]. Type B.

The specimen from the Persian Gulf also belongs to Type B.

The other specimens of Heterocyathus in the Persian Gulf

collection constitute a new species. The tendency to remain

conical is very marked; the coste are distinct from those of other

species ; and the uniformity in size separate it sufficiently until a

good series of intermediates are found which will link it up

with /7. alternatus.

I therefore recognize in the genus the following species :-—

1. H. cequicostatus Milne-Edwards & Haime.

Stephanoseris rousseaui Milne-Kdwards & Haime.

H. parasiticus Semper.

H. philippinensis Semper.

H. oblongatus Rehberg.

H. pulchellus Rehberg.

H., wood-masoni Alcock.

2. H. alternatus Verrill.

H. cwequicostatus (Gardiner’s Type I).

3. H. heterocostatus, sp. 0.

Hrrerocyatuus aquicostatus Milne-Edwards & Haime [29].

A single specimen belonging to Type B. Coste, equal in

number to septa, extend to base, beset with irregular spines.

Base roughly granular. Slight calicular fossa. Septa in six

systems of four complete cycles, very spiny ; those of the fourth

eycle fuse with those of the third, and these in turn fuse with

those of the second. Inner margins of septa pass imperceptibly

into the trabecular columella.

Locality. Telegraph Cable, Persian Gulf. Depth 40 fathoms

HETEROCYATHUS HETEROCOSTATUS, sp. n. (Pl. LVII. fig. 6;

Pl. LVIII. fig. 14.)

Corallum simple, free, variable in shape from a low discoid

form to a conical cornuate form. Upper part of corallum

deeper in colour than base. Height varies between 3mm. and

7 mm., and calice between 3°53 mm. and 6 x 5mm. Coste of

two distinct types. Those corresponding to the septa of the first,

second and third cycles are visible from the base to the tip of the

calice, prominent sharp ridges with a single series of coarse

eranulations ; the alternate coste, corresponding to the septa of

the fourth cycle, extend only about half-way down the corallite

from the lip of the calice; they are less prominent, and beset

with numerous irregular granulations. Calice slightly elliptical,

CORALS FROM THE PERSIAN GULF. 1027

open, deep. Septa in four complete cycles. Primaries large, very

prominently exsert ; quaternaries join over the tertiaries, and again

deep down in the calice over the secondaries; quaternaries on either

side of the primaries much developed, and prominently exsert in

the manner typical of the genus Heterocyathus ; all septa beset with

numerous fine granulations arranged in radial ridges. Pali in the

form of small denticulations, which pass imperceptibly into ¢

parietal, fasciculate, centrally depressed columella. At the base, a

circular aperture leads to an Aspidosiphon chamber containing ¢

small Gastropod shell, in which a Sipunculid lives commensally

with the coral zoophyte.

Nine specimens.

Locality. Karachi. Depth 15-40 fms.; bottom, shell-sand.

Paracyatuus cavarus Alcock [1]. (Pl. LVII. fig. 5.)

Corallum simple, fixed by spreading base and expanding slightly

to lip of calice. Height varies between 15 mm. and 20 mm., but

the shortest has the largest calice. Calice variable in shape,

probably owing to local surroundings ; in some specimens calice is

circular, in others oval, while others have various indentations

and exerescences. Diameters of calices 15 x 15 mm., 16 x 12 mm.,

17x 10mm.,18x15mm., 20x13mm.,20x18mm. Coste visible

from base upwards, low broad ridges covered with minute

granulations, corresponding to each septum; towards the lip of

the calice these costal ridges become more marked, and somewhat

stouter and more exsert coste alternate with slightly smaller

coste. This alternation of larger and smaller costze is apparent

in some specimens at the extreme base of the corallum as well as

at the ealicular margin. Septa close, not markedly exsert ; the

size of the septa and the extent to which they are exsert

diminish in a descending order of magnitude according to the

eyele to which they belong: those of the first cycle are larger

than those of the second, the second than the third, and go on.

They are uniformly covered with minute granulations, which are

arranged in a series of longitudinal and radial rows. he calice

is widely open, and the inner margins of the septa are cut up

into large irregular nodules representing small pali, which pass

imperceptibly into a fasciculate parietal, centrally depressed

columella, ‘There appear to be typically five orders of septa, but

those of the lower orders are not always easy to interpret. ‘The

primaries are always large and extend to the columella; their

paliform nodules are slightly larger and more projecting than those

of the septa of lower orders; the septa of lower orders tend to fuse

together near the columella, and pass into it as an ill-defined mass

of minute pali. The larger size of the primary septa and the

grouping together of those of other orders give a hexagonal

starlike effect to the calice. In none of the seven specimens from

the Persian Gulf are the five cycles complete, although in one

there are 94 septa, only two septa of the fifth cycle being missing.

In the specimen figured (text-fig. 217), however, there are also

1028 MISS R. HARRISON AND PROF. 8S. J. HICKSON ON

94 septa, but here the arrangement is not so simple: in one lateral

chamber on each side between the primary and secondary septa

the fifth cycle is unrepresented, there being only three instead of

the normal seven septa. On the other hand, in both the apical

Text-fig. 217.

Diazram ui tic plan of the sah? of Paracyathus cavatus. The primary and

secondary septa are black, the tertiaries are cross-hatched, the quaternaries are

crossed with diagonal lines, the quinaries are left blank; i, ii, ii, iv, v, septa

of the first, second, third, fourth, and fifth cycles.

chambers at one end of the long axis there is an excess of septa.

In both chambers, between the primary and secondary septa on

the one side, and the primary and secondary septa on the other

CORALS FROM THE PERSIAN GULF. 1029

side, there are more than the normal number of septa. It

appears as if the tertiary septa of both these chambers had been

split by a wholly superfluous quaternary septum, while in one

case a quaternary septum has been further split by a super-

numerary quinary. The same phenomenon is apparent to a

greater or lesser degree in all the remaining five specimens. : In

one specimen there is a sudden outpushing of the wall of the

calice between a primary and a tertiary septum, in which there

are no fewer than nine instead of the normal three septa; in

every case there is a costa to correspond with each septum, and

the alternation of large and small septa and coste is maintained.

Seven specimens fixed on to a mass of mud, shell and serpulid

tubes.

Locality. From Cable 60 miles S.W. of Bushire, Persian Gulf,

Depth 30 fms.

Two specimens are infested by the Cirriped Pyrgoma stokesii,

which forms a bulbous gall within the wall of the coral (Pl.

LVII. fig. 5c). This distortion does not, however, interrupt the

regularity of the coste, which are distinctly visible on the outside

of the parasitic chamber; nor does the presence of this parasite

appear to affect the number of septa; there is no excess or

shortage of septa in the attacked specimens beyond that which is

normally observed in the unattacked individuals.

Genus Trematorrocuus T. Woods [39].

TREMATOTROCHUS ZELANDIZ. (Pl. LVII. fig. 4; Pl. LVIII.

figs. 15-17).

Conocyathus zelandie Duncan [14].

Corallum regularly conico-cylindrical, free, without trace of

attachment. No epitheca. Height of largest specimen 7 mm.,

‘diameter of calice 3°4 mm. Coste in four complete cycles,

‘prominent, smooth, and equal in the upper part of the corallum.

Only those of the first two cycles extend to the base; those of

the third cycle extend downwards for about three-quarters, and

those of the fourth cycle for a distance varying between a third

and three-fifths of the height of the whole corallum. The cost

between those of the second and third cycles are longer than the

costee between those of the first and third cycles; they do not

join those of the preceding cycle, but there is a thickening of the

costz of the first three cycles below the point at which that of

the succeeding cycle ends. Intercostal furrows penetrated by

minute, regularly disposed perforations (Pl. LVII. fig. 4). Calice

circular, no fossa, Septa in six systems of three complete cycles,

all exsert, those of the first cycle more prominently so than those

of the succeeding cycles; very thin and beset with small spinous

granulations. Septa of the third cycle join those of the second

a short distance below the lip of the calice ; they are incomplete

at their inner margins, large fenestrations occurring at the point

Proc. Zoot. Soc.—1911, No. LXX. 70

1030 MISS R. HARRISON AND PROF. S. J. HICKSON ON

where the septa of the third cycle join those of the second, and

those of the first and second join the columella. The lower

fifth of the corallum has been filled up internally by a secondary

deposit of calcareous matter, but the outline of the original struc-

ture is faintly discernible in photographs of sections at this level,

and reveals that the columella is parietal, formed by the union of

the inner ends of the septa.

Three specimens.

Locality. Persian Gulf. Depth not recorded.

The genus 7rematotrochus was proposed by Tenison Woods [39 |

for a fossil from the Miocene of Australia. The coral had all

the characters and appearance of a Turbinoliid, but with the

important difference that the wall was penetiated by large per-

forations between the coste, giving free communication between

the interseptal chambers and the exterior.

This remarkable coral did not receive the attention it deserved,

and subsequent authors received sceptically the statement that

these perforations were really present; for the point called in

question the validity of the division of the Madreporaria into

Perforata and Imperforata. Duncan [15] pointed out the diffi-

culty of placing this genus, but referred it to the Turbinoliide

in close relation with Turbinolia, Stylocyathus, Conocyathus, and

Bistylia*.

- In a series of papers published in the Transactions of the

Royal Society of South Australia, Dennant [7 & 8] has since

described six more fossil forms and one recent form (7. verconis)

from Australia and Australian Seas. In the first of these

papers he describes two species from Eocene and one species

from Miocene deposits. The recent species was found in St.

Vincent’s Gulf and Backstairs Passage at depths of from 15 to

22 fathoms. All have the characteristic perforations, which

fact caused him to remove the genus altogether from the Turbi-

noliide and place it among the Hupsammiide. Later he described

three different species from older Eocene beds than those in

which his previous species were found, and in two of these

the perforations do not pierce the wall, but are merely pore-

like cavities extending half-way through the thickness of

the wall like the intercostal dimples of some Turbinolide.

This made him put the genus back among the Turbinolide,

and he referred to Gregory’s [23] suggestion that the Perforate

type of coral has been derived from the Imperforate. He

further remarked on the wide distribution through time of the

original species 7’. fenestratus, and appended a drawing of a

portion of the wall cf one of them, placing the existence of the

pores beyond all suspicion of doubt. All the previously described

species come from Australia, and Dennant pointed out the

* Duncan uses the phrase “one of the species”’ as if more than one species were

known. At the time ofthe publication of his paper (1885) I am not aware that

any other species had been cescribed.

CORALS FROM THE PERSIAN GULF. 1031

occurrence of a recent form as evidence of the close relationship

between the living and fossil fauna ef that country. He made

no attempt to discuss the significance of these forms in con-

nection with the existing classification of the Madreporaria, but

dismissed the subject with the single statement—“ the broad

distinction usually made between perforate and non-perforate

corals breaks down.”

The occurrence of another recent species of this problem-

atical genus again forces the subject on the attention of the

systematist. That it isa T'rematotrochus is, | think, unquestion-

able. It agrees with Dennant’s latest definition in all points

but one :—He has described the septa as being solid; but in

T. zelandie the septa are incomplete, large fenestrations occurring

at the point at which the tertiaries join the secondaries, and the

primaries and secondaries join the false columella. Realizing

that the existence of these fenestrations and of the pores in the

outer wall might again be doubted (having already admitted

that these latter are minute and difficult to see), I have ground

sections of one specimen, thinking that actual photographs would

be more convincing than external observations. These sections

have been made at intervals of #, mm. and the entire coral has

been ground away; every section was photographed at a magni

fication of 20, and from these photographs a wax model has

been constructed. There is, therefore, a permanent record of the

structure of this most interesting coral, and it would be idle to

deny the existence of either thecal pores or septal fenestrations.

But in spite of this, I think that it is rightly placed among the

Imperforate corals.

In text-fig. 218, A and C are diagrammatic representations

of transverse sections of portions of the wall of a Heterocyathus

and a Heterepsammia (after Bourne [6'), for comparison with B,

a transverse section of 7. zelandiw. The Heterocyathus has a

solid theea embedded in stereoplasm. The Heteropsammia has

a porous spengy wall. A glance at the WVrematotrochus will

show that the extremely thin wall is incomplete, and that here

and there are small communications between the interseptal

chambers and the exterior; but it could not be regarded as a

“porous coenenchyme” such as is diagnostic of the division

Perforata. ‘The condition is much more that of a Fungiid which

has grown conically instead of spreading out with a widely open

oral surface, with that part of the wall between the perforations

representing synapticula. In some of the fossil forms the per-

forations are so large and the intervening solid structure little

more than a narrow bar, that the comparison with synapticula

is more striking than in the present instance. The analogy,

however, must not be pressed too far, but it widens the gulf

between 7rematotrochus and the Perforata, and tends to justify

its inclusion among the Imperforata.

_ Another point to be emphasized is the localities in which the

species are found. Hitherto the eight species of the genus were

CAO}

MISS R, HARRISON AND PROF. S. J. HICKSON ON

1032

ted area of Australia and the seas round

imi

all found in the |

the

but now we find it

that continent and New Zealand

‘saiod [eqsoo1aqut “d 2a $ vusetdooseys “wag f sopoAo YIAMOF pur ‘party} “Spuooes ‘ys1y ay} Jo vydas “At “tir “1 “I

*(aumog «aqye) ru2payorm vrmunsdowazazy JO WN]]R109 ayy YSUOAYY UOTZOOS OLYBULLUBASLIP B JO WOO “{)

‘DIPUNIAZ SNYIOAZOINUALT, JO VANI[BAOD OY} YSNOAYy WOTZOIS O1yVUUUMIBISVIG “q

‘(ananog aeqye) sngnzsooinba snyjvhoowazezy JO WIN][R109 AY} YSNOAY} UOTZaS OTAVLUUTUBALSEIp B JO UOIWO “W

"BIG “8Y-489,

lentity of the last with Conocyathus zelandi

1€

The

Persian Gulf.

has already been pointed out in the introductory remarks of this

CORALS FROM THE PERSIAN GULF. 1033

paper, and the occurrence of the same species in localities so far

apart 1s remarkable.

I had hoped that the sections might throw some further

light on the question of septal sequence, but unfortunately the

lower fifth of the coral has been filled up solidly by a secondary

deposit of calcareous matter, and the interseptal chambers do

not begin to appear distinctly until a level is reached where

twelve septa and twenty-four coste are already present. If it

Text-fig. 219.

eae

A. Diagrammatic plan of the cost of Tremtotrochus zelandie, showing the

difference in length of the cost of the fourth cycle. i, ii, ili, iv, costae of the

first, second, third, and fourth cycles.

B. Diagrammatic plan of the septa of Ti-ematotrochus zelantie, representing the

exsert peripheral portion of the primary septa, and the exsert pali of the

secondary septa. i, li, iii, septa of the first, second, and third cycles; ea.s., exsert

portion of the septa of the first cycle; p., pall.

may be taken that the cost correspond to the septa and that the

base of a coral represents the earliest formation, it might be

objected that Pourtalés’ law for septal sequence has not been

followed in this case; for the cost which extend to the base,

twelve in number, are those corresponding to the first and

third (according to Pourtalés [35]) cycles of septa. That is to

say, if the secondary septa split peripherally in a Y-shape, and

a tertiary septum grew up between the arms of the Y, the

septa generally spoken of as secondary and tertiary ave named

1034 MISS R. HARRISON AND PROF. S. J. HICKSON ON

conversely to the order in which they appear in ontogeny. But

it must be remembered that the stem of the Y is also the

representative of the secondary septum according to either the

popular or Pourtalés’ nomenclature ; and these septa still exist

in their normal position midway between the primary septa,

and project upwards as pali, or paliform lobes: in the lower

portion of the colony, before the secondary septa have begun

to branch, the coste corresponding to them are also unbranched,

and extend to the base. A comparison of the diagrams A, B,

in text-fig. 219, illustrates this point.

Diagram A represents 4 plan of the cost and Ba plan of the

septa; in Aa line “aa” has been drawn across the lowest part

of the corallum, and in B a corresponding circle “aa” has been

drawn round the inner part of the septal plan; in both, the

costee and septa respectively of the first and second cycles are

present; presumably the coral must at one time have passed

through a stage when these cycles only were present. In

T. fenestratus this is the arrangement in the adult. Again,

another line “66” is drawn across A at a higher level and a

corresponding circle “bb” on B; in both, the cost and septa of

three cycles are present. When the secondary septa branched

peripherally, new costs arose in connexion with these branches,

and the original secondary eoste remained in connexion with

the inner unbranched part of the secondary septa which persist

as pali or paliform lobes. Bourne [6, text-fig. I. 2] has given

a coniprehensive diagram iHlustrating the relation of septa and

pali according to Pourtalés’ primciple: diagram B illustrates the

arrangement in 1. zelandie, which is entirely in agreement

with it.

The fact that the costz of the fourth cycle are of different lengths

(see text-fig. 219 A, iv) is in accordance with Duerden’s [11]

account of the septal sequence of Siderastrwa radians, in which

he states that new mesenterial pairs appear in some interseptal

chambers, before those in others of the same eycle.

AGELECYATHUS PERSICUS Duncan [14].

Corallites rising from an encrusting base, expanding slightly

from base to calice. Calice elliptical, depressed at one end of

the long axis. Height of corallum from 5-10 mm. at depressed

end of the long axis, and from 15-20 mm. at the opposite end.

Diameter of calice 9x75 mm. to 11x9 mm. The shape of the

corallites suggests that the colony was fixed on a vertical sur-

face, and the inequality of height in the two extremities of the

long axis is a response to an effort on the part. of the indi-

vidual corallites to assume a vertical position. Coste in four

complete cycles, distinct from base upwards, slightly exsert and

finely granular. Septa in six systems of four cycles of which

the last is incomplete, although all the coste of the fourth

cycle are present; septa of the first two cycles more exsert

than those of the third cycle; all three cycles reach the columella ;

CORALS FROM THE PERSIAN GULF. 1035

septa beset with minute granulations arranged in radial ridges.

An incomplete single crown of pali before the septa of the third

cycle. Columella fascicular.

Text-fig. 220.

Diagrammatic plan of the septaof Agelecyathus persicus. Septal orders represented

as in text-fic. 217. Six detached pali are represented. i. ii, iii, iv, septa of the

first, second, third and fourth cycles; p., pali.

A single specimen consisting of five individual corallites arising

from a common encrusting base.

Locality. From Cable 60 miles 8.W. of Bushire, Persian Gulf.

Depth 30 fms.

Before some of the septa of the third cycle there are distinct

upwardly projecting lobes corresponding to pali. Thus in the

calice figured (text-fig. 220) there are six such pali opposite six

septa of the third cycle, three at each end of the long axis;

there are two less conspicuous lobes before two septa of this

cycle, while the remaining four septa of the third cycle have

no trace of any such Jobe. Milne-Edwards and Haime [29]

divided the family Turbinoliide into two sub-families—the Tur-

binolinz without pali, and the Caryophylline with pali; and this

classification has found more favour with subsequent authors than

Dunecan’s [15] later classification, which divided the family into

Turbinolide simplices, Turbinolide gemmantes, and T'urbinolide

reptantes, according to their habits of growth and reproduction.

This coral affords evidence in favour of Dunean’s classification ;

indeed it would be difficult to know in which of Milne-Edwards’

1036 MISS R. HARRISON AND PROF. 8. J. HICKSON ON

and Haime’s groups to place it, for here in a single calice is a

combination of the diagnostic characters of the two sub-families :

pali are present before some and not before other septa of the

same cycle.

This specimen is considerably larger than Duncan's original

example from the same locality, and differs from his flew of

the type specimen in having the septa in the first two cycles

much exsert; but he calls attention to the variability of the

pali, the size and the number of the septa, as being features

of the individual corallites. The similarity in other details 3s

close.

Family FUNG611D &.

FuNG6IA PATELLA Ellis & Solander [16].

Three specimens from the Persian Gulf which are identical

with Doderlein’s figure [10, pl. i.] of the Cycloseris-form of

ff, patella, Wayland Vaughan’s figure [40, pls. xxv. & xxviii.] of

I’. patella, and Gardinexr’s figure [18, pl. 11x.] of Cycloseris hexa-

gonalis. They afford further evidence of the wide distribution

of the species, and for the justification of absorbing the genus

Cycloseris in the genus Fungia.

Locality. Twe specimens from Shaikh Shuaib Island, Persian

Gulf. Bottom, rock. Depth 10 fms. Exact locality of third

specimen not recorded.

Family HuPSAMMIID&.

HereroPsAMMIA APHRODES Alcock [1].

Eight specimens each with a single calice, The inflated spongy

exsert edges of the septa of the first two cycles are well developed.

Locality. Telegraph Cable, Persian Gult. Depth 49 fms.

DENDROPHYLLIA sp. ? de Blainville [5]. (Pl. LVIL. fig. 7.)

Colony bushy, springing from an encrusting base on which are

numerous small individuals. Budding lateral or basal. Polyps

occasionally joined by horizontal bars, but in every case such bars

are inhabited by a small commensal worm, and have probably

been formed in connexion with this parasite. Polyps tend to

assume a vertical position, but one polyp which appears to

arise from a horizontal bar is directed downwards. Size of the

polyps very variable; some searcely rising from the encrusting

base; some attaining to a height of 25 mm. Calice varies

between 2x2 mm. in the youngest to 7x6 mm. in the oldest

individuals. Coste distinct from base upwards, equal, granular,

not exsert, correspond in number to the septa. Septa in

six systems of four cycles with a few representatives of the

fifth cycle, all irregular. Some septa of the lower cycles are

grouped together round that of the preceding cycle in the typical

dendrophylliid manner, while others extend to the columella, and

are equal in size to those of preceding cycles. All septa are

CORALS FROM THE PERSIAN GULF. 1037

finely granular, the lower orders are perforate. Calicular fossa

very deep. Columella fascicular, parietal; well developed in older

individuals.

Locality. From Cable 60 miles 8.W. of Bushire, Persian Gulf.

Depth 30 fms.

With only a single specimen of a coral such as this, which

exhibits such a wide range of variability, the creation of

a new species has not seemed to me justifiable. I have, therefore,

merely noted its characters and appended a photograph of the

specimen, until such time as it may be found in greater numbers.

Note on Miss Harrison’s memoir on some Madreporaria from

the Persian Gulf, and some further notes on Pyrophylli«

inflata. By Sypney J. Hickson, M.A., F.R.S., F.Z.8.

The manuscript of Miss Harrison’s paper was sent to me shortly

before she left this country for India, with a request that I would

read it and revise it for publication. The number of species in

the collection is small and there is only one that is new to science,

but there are several points in the paper which it seemed to me

required rather fuller consideration than she has given to them,

and I have ventured therefore to write an addendum, leaving her

original memoir intact. Had she remained in England I would

have suggested the inclusion of these remarks in her paper, but

under the circumstances, I think it is better to publish them

under a separate title and thereby take the whole responsibility

for them. I have rearranged the order of the species, furnished

the list on p. 1019, and added the family names, but in other

respects the paper is as it left her hands.

At the present time our knowledge of the fauna of the Persian

Gulf is very limited. None of the great deep-sea exploring

expeditions have visited it, and independent investigators with

sutticient knowledge and energy in this region have been few and

far between.

Mr. F. W. Townsend, of the telegraph staff of the Indian

Government, bas made a large and valuable collection of shells,

and these have been described in a series of papers by Cosmo

Melvill and Standen,* but so far as I can discover, very few

genera and species belonging to other groups of marine animals

have been recorded. The richness and interest of the Molluscan

fauna suggests that many new forms have still to be discovered

in the Gulf, but the subject of special importance that the study of

this fauna would shed light upon, is the relation of the fauna

of the Persian Gulf to that of the Mediterranean Sea. It has

frequently been suggested that in the past there was a connexion

between the Indian Ocean and Atlantic Ocean by way of the

Mediterranean Sea ; and, judging by the present day geographical

features, it is probable that the last connexion between them

* Wor list of papers see Proc. Zool. Soc. 1906, p. 783.

1038 MISS R. HARRISON AND PROF. S. J. HICKSON ON

was broken by the formation of the Isthmus of Suez. But the

last connexion but one was that by way of a strait, of which

the remaining part is now the Persian Gulf; and this gulf with

its narrow outlet into the Indian Ocean, high temperature and

great rivers, might be expected to retain some of the fauna which

had been subject to very similar conditions in the South-east

corner of the Mediterranean Sea. Melvill and Standen call

attention to the considerable generic analogy between the Mollusca

of the two regions, and point out that the species of the

Persian Gulf show close affinities with South-Huropean forms.

It would be quite premature to draw any far-reaching

generalisations as to the distribution of Madreporaria from the

few specimens that are here recorded, but attention may be

called to one or two points of general interest.

The occurrence of Pyrophyllia in deep water in the Persian

Gulf is interesting from the point of view of geographical

distribution. Alcock [3] in his comments on the deep-sea Madre-

poraria of the Indian Ocean, calls attention to the ‘“ many

intimate aftinities of the fauna of moderate depths of the Indian

seas with the North Atlantic fauna,” and considers them ‘to be

sufficient to suggest a direct sea-connexion, in the past, between

the Atlantic and Indian Oceans, and the case of Caryophyllia

communis and Flabellum laciniatum would indicate that the

connexion was by way of the Mediterranean.”

The case of Pyrophyllia and Guynia appears to me to give

even stronger evidence of the truth of this hypothesis, than that of

the two species quoted. The genera Caryophyllia and Plabellum

are both very widely distributed recent corals, and it is possible

that in comparatively recent times these two species may have

had an almost cosmopolitan distribution. Guynia and Pyrophyllha

are, so far as is known at present, very restricted in their distri-

bution and are totally unlike any other recent coral—with the

possible exception of the West Indian genus Haplophyllia.

Although so much alike in important characters, they are

sufficiently distinct for us to believe that they were separated

from one another at a very remote period. It is rather more

difficult to believe that the Indian Ocean and Mediterranean

specimens of Caryophyllia communis and Flabellwm laciniatwm

can have undergone no differential change since the time when

the Mediterranean Sea and the Indian Ocean were in commu-

nication. A second point of interest is, that the only other coral

with which Pyrophyllia shows aflinities, namely Conosmila,

should be found in the Tertiary deposits of Australia. Standing

by itself, this is only one of those facts of geographical distribution

which it is important to note but impossible to explain in a

satisfactory way. But its importance as a fact is emphasized

when it is placed side by side with the facts of the distribution of

Trematotrochus.

As Miss Harrison points out, the specimens of Z’rematotrechus

CORALS FROM THE PERSIAN GULF. 1039

found in the Persian Gulf are closely related to species of corals

from the Tertiary deposits of Australia, and to a recent coral

found at depths of from 15 to 22 fathoms in St. Vineent Gulf,* and

ilentical with a recent coral from “no very great depth” im

Cook’s Strait, New Zealand.

Of the geograpical distribution of the other species very little need

be said. Heterocyathus heterocostatus is new, but a closely related

species H. equicostatus, of which one specimen was found, appears

to be widely distributed in the Indian Ocean. Pwracyathus cavatus

is found in the Indian Ocean and is said by Alcock to he closely

related to the Hocene fossil P. crassus from the London Clay.

Agelecyathus persicus was previously recorded by Duncan from

the Persian Gulf, but was said to oecur also off St. Helena.

Fungia patella occurs in the Indian Ocean. ‘The type specimen

of Flabellum magnificwm was found off Sumatra at a depth of

470 metres. The genera Heteropsamnia and Dendrophylliaappear

to be widely distributed in the Indian Ocean. So far then as

this smail collection of corals is concerned, Pyrophyllia inflata is

the only species that even suggests a former connexion of the

Gulf with the Mediterranean Sea.

The genus Pyrophyllia was briefly described in the ‘‘ Manchester

Memoirs,” 1910; but it may be convenient to take this

opportunity of adding a few general remarks on the genus and

of publishing some further illustrations (P]. LVII. figs. 8-11;

Pl. LVITTI. figs. 18, 19).

Pyrophyllia inflata is a small unattached solitary coral, of

about 4 to 5mm. in length, and 1 mm. in diameter at the margin

of the calyx.

The two most important characters are :—(1) Its very

pronounced and-invariable octoradiate symmetry, and (2) the

presence on the external surface of well-marked accretion ridges

with short but definite costal spines (text-fig. 221).

Pyrophyllia is related to the recent genus Guynia, and

to the extinct Tertiary genus Conosmilia. The number of

septa in Pyrophyllia and in Guynia is sixteen, and of these eight

are larger and may be called the*primary septa, and the remaining

eight are smaller and may be called the secondary septa.

In Guyma one of the eight primary septa is larger than

the others, this large septum being according to Duncan a

““very marked rugose peculiarity.” Moreover, in Guynia four of

the primary septa are sometimes larger than the other four, so

that according to Duncan the system of septa is four primary

septa, four secondary septa, and eight tertiary septa.

In these respects and in others of less importance, Guynia

seems to be distinct from Pyrophyllia. In the genus Conosmilia

there is a more variable arrangement of the septa. In

C. elegans, C. litwolus, and C. anomala there are eight

primary septa, eight secondary septa, and thirty-two tertiary

* The specimens of this coral Z. verconis were much worn and were no doubt

dead corals when collected.

1040 MISS R. HARRISON AND PROF. S. J. HICKSON ON

septa; but in C. striata there are only six primary septa, six

secondary septa, and twelve tertiary septa. The more recently

described species of this genus, C. granulata and C. stylifera

(Dennant 7), agree with C. striata in the hexaradial arrangement

of the septa. In Conosmilia, moreover, the accretion ridges do

not appear to exist unless they are represented by the “ beautiful

herring-bone ornamentation of the surface.”

Diagrammatic sketches of the septal arrangement of Pyrophyllia inflata.

g 12) g Yrop

c., columella; m.s., secondary septa; p.s., primary septa.

A. Arrangement of the septa just below the margin of the calyx. B. At the base

of the calyx. C. In the lower parts of the coral. (From Mem. Manch. Lit.

Phil. Soc. 1910.)

The columella of Conosmilia resembles Pyrophyllia im being

laminate and in this respect differs from Guynia, in which the

columella is cylindrical.

There has been a great deal of hesitation in giving the two genera

Guynia and Conosmilia a definite resting-place in the system of

corals,

Duncan * at first placed the genus Guynia in the Order

Rugosa and in the family Cyathaxoniide, but subsequently

removed it [15] to the family Turbinoliide. Miss Ogilvie placed

it in her new family Amphiastreeidze +.

Dunean $ at first placed the genus Conosmilia in the Order

Rugosa, family Stauride, but subsequently removed it to the family

Astreidee Simplices and placed it close to the genus Zrochosmilia.

Miss Ogilvie placed this genus in the family Turbinoliide.

In my preliminary account of the genus, I remarked that ‘it

cannot be denied that Pyrophyllia has some characters reminiscent

of the extinct Rugosa,” but on reconsideration I do not feel

* Phil. Trans. R. §. 1872. y Phil. Trans. 1896. £ Phil. Trans. 1872.

CORALS FROM THE PERSIAN GULF. -1041

satisfied that these characters—the octoradiate symmetry, the

pronounced accretion ridges and the septal fusions in the lower

part of the coral—really indicate any true affinities with Paleeozoic

corals. Duncan found one specimen of Guynia in which, owing

to the abortion (?) of two septa, the upper part of the corallum

showed a hexaradiate symmetry. This specimen was regarded

by him as ‘‘very suggestive in the matter of the evolution of

the hexameral from the octomeral types, or rather from the

tetrameral.” But the view that was in his mind, that the

modern hexaradiate corals are descended from a previous tetra-

radiate type, is not one which commends itself to more recent

investigators of coral structure. The researches of Duerden*

suggest that the tetraradiate symmetry of Rugose corals is derived

from a more primitive hexaradiate symmetry, and there is no

evidence in the development of modern corals, or satisfactory

evidence in paleontology, that the modern hexaradiate corals are

derived from a previous tetraradiate or octoradiate ancestry.

It is true that many Paleozoic corals exhibit four dominant

primary septa, but there are none with eight dominant septa.

The true octoradiate condition is known only in certain species of

the Tertiary genus Conosmilia and in the recent genera Guynia

and Pyrophyllia. ‘The recent genus Haplophillia has also eight

primary septa and eight secondary septa, but it is suggested by

Gardiner that this genus is only a growth stage of “Duncania.

The close relationship of Conosmilia to the simple modern

hexaradiate corals suggested by the classification of Duncan and

of Ogilvie, seems to point to the conclusion that the Guyniide

represent an early offshoot of the modern hexaradiate line of

descent, and that their true affinities lie rather with the modern

Turbinoliide than with any known Paleozoic coral.

I may refer in conclusion to some remarks made in my

preliminary paper on the presence of endotheca in Pyrophyllia.

Although Duncan’s definition of endotheca is not very explicit,

I have come to the conclusion that there is no structure in

Pyrophyllia that really corresponds with what is usually regarded

as endotheca by the paleontologists. A longitudinal section of a

Pyrophylia (Pl. LVITI. fig. 19) shows three regions: an upper

calicular region (a) in which the septa are free from one another

and from the columella, a middle region (y) in which the septa

are more or less irregularly fused with one another and with the

columella, and a lower region (z) in which the septa are again free

and the thecal wall inflated.

It seems probable from the texture of the well-preserved

specimens that the living tissues were confined to the upper

calicular region. ‘The growth in thickness and the fusion of septa

and columella in the middle region (see text-fig. 221) perform the

same function as the endotheca of Paleozoic corals, as they ‘“ unite

septa, close the loculi, and enable the coral to grow in height

and strength, and limit the growth downwards of mesenteries

* Ann. N. H. xviii. 1906.

1042 MISS R. HARRISON AND PROF. S. J. HICKSON ON

and soft parts,” but there are no thin plate-like structures

corresponding with those that constitute the greater part of the

endotheca of Paleozoic corals.

13.

14.

15.

16.

18.

. Atcock, A.

. DENNANT, J.

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Marine Survey Steamer ‘ Investigator.

Soc. Bengal, Ixii. p. 169.

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Royal Indian Survey Ship ‘ Investigator.’ Calcutta, 1898,

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. Atcock, A.—‘“‘ Report on the Madreporaria of the Siboga

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. Bourne, G. C.—* Solitary Corals collected by Professor

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the Australian Tertiaries.” Trans. Roy. Soc. South

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“Recent Corals from the South Australian

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Anz. xxiv. 1901, p. 353.

. Doperte, L.—‘‘ Die Korallen-Gattung Pungia.” Abh. v.

d. Senckenbergischen naturforschenden Gesellschaft, xxvil.

1905, p. l.

. Durrpen, J. E.—‘‘ The Morphology of the Madreporaria. V.

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e280:

Dean, P. M.—‘“ A Description of the Madreporaria dredged

up during the Expedition of H.M.S. ‘Porcupine.’” Trans.

Zool. Soc. London, viii. 1874, p. 303; x. 1879, p. 235.

Duncan, P. M.—“ Notices of some Deep-Sea and Littoral

Corals from the Atlantic Ocean, Caribbean, Indian, New-

Zealand, Persian Gulf and Japanese &c. Seas.” P. Z.S. 1876,

. 428.

inca P. M.—‘“ A Revision of the Madreporaria.” Journ.

Linn. Soe., Zoology, xviii. 1885, p. 1.

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1786.

GARDINER, J. 8.—‘‘On the Solitary Corals collected by the

author in the South Pacific.” P. Z.S. 1898, p. 525.

GARDINER, J. S.—“ On the Solitary Corals collected by Dr. A.

Willey.” Willey’s Zool. Results, 1899, p. 161.°

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CORALS FROM THE PERSIAN GULF. 1043

Garpiner, J. S.—“South African Corals, of the Genus

Flabellum.” Marine Investigations in 8. Africa, II., Cape

> Town, 1904, p. 117.

Garpiner, J. S.—“The Turbinolid Corals of 8. Africa.”

Marine Investigations in 8. Africa, III., Cape Town, 1904,

2 45,

Gasset J. S—“The Fauna and Geography of the

Maldive and lLaccadive Archipelagoes.” IT. Suppl. i.

Madreporaria, Pt. 111., Fungiide, Pt.iv., Turbinolide, p. 933.

GARDINER, J. S.—‘* The Percy Sladen Trust Expedition to

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B25c

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Kent, W. Savitte.—‘*On some new and little-known

Species of Madrepores, or Stony Corals, in the British

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Ann. des Sci. Nat., 3° sér. x. 1848, p. 65.

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Coralliaires.” 3 vols., Paris 1857.

Mosstey, H. N.—“ Deep-Sea Corals.” ‘Challenger’ Reports,

Zoology, 11. 1881, p. 127.

Ortmann, A. E.—“ Beobachtungen an Steinkorallen von der

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1044 ON CORALS FROM THE PERSIAN GULF.

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Polyzoa.” Trans. Roy. Soc. New South Wales, xii. p. 57.

Vaucuan, T. W.—‘ A Critical Review of the Literature on

the Simple Genera of the Madreporaria Fungida, with a

tentative Classification.” Proc. U.S. Nat. Mus, xxviii.

1905, p. 371.

Vaueuan, T. W.—‘‘ Recent Madreporaria of the Hawaiian

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Verritt, A. K.—‘‘ Corals and Polyps of the North Pacific

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Amer. Journ. Sci. & Arts, 2nd ser. xlix., 1870.

EXPLANATION OF THE PLATES.

Prate LVII.

1, Flabellam magnificum von Marenzeller. Viewed from above. x 3 diam.

2. Flabellum magnificum. Viewed from below to show the tubular rootlets of

attachment.

3. Flabellum magnificum. Side view.

4. Trematotrochus zelandie Duncan. Transverse section of the coral.

x ca. 12 diam. Compare text-fig. 218 3, p. 1032.

5. Paracyathus cavatus Alcock. A. View of avertical fracture. B. View from

above. C. Side view showing on both sides above, galls formed by Pyrgoma.

Nat. size.

6. Hetcrocyathus heterocostatus, sp.n. Nine specimens showing the variations

in height. x 2 diam.

7. Dendrophyllia sp.? x % diam.

8. Pyrophyllia inflata. Side view. x 5 diam.

9. Pyrophyllia inflata. View of the mouth of the calyx showing the columella

and primary septa.

10. Pyrophyllia inflata. View of a specimen that has been broken in half

longitudinally showing the irregular arrangement of the fused septa in

the lower regions of the ecral.

11. Pyrophyllia inflata. View of a specimen showing the fusion of the eight

primary septa and the secondary septa. Compare text-fig. 221 B, p. 1040

Prare LVIII.

ig. 12. Heterocyathus equicostatus M.-E. & H. Two adjacent costee which are

low broad ridges covered all over with uniform fine granules, the

intercostal furrows being small and shallow. ‘his speéimen belcngs to

Gardiner’s Type I.

13. Heterocyathus alternatus Verrill. Two adjacent cost. The costs on

the right (A) is the type of costa corresponding with the septal cycles

i, ii, andiii. The costa on the left corresponds to the septal cycle iv.

The intercostal furrows are deep. This specimen belongs to Gardiner’s

H. e@quicostatus Type 1.

14. Heterocyathus heterocostatus, sp.n. Side view. x ca. 14 diam.

15. Trematotrochus zelandie Duncan. Side view. x ca. 12 diam. Compare

text-fig. 219 A, p. 1033.

16. Trematotrochus zelandie. View of the base.

™ Trematotrochus zelandie. View of the calyx from above. Compare text-

fig. 219 B.

18. Pyrophyllia infiata Hickson. View of the calyx showing the columella,

the eight primary septa, four of the eight secondary septa, and the limes of

accretion.

19. Pyrophyllia inflata. Wongitudinal vertical section showing the fusion of

the primary septa in the lower parts of the coral. «, region of the calyx ;

y, middle region; z, inflated base.

‘P.

pith etiam

Hath

le y4iSy WSL PAL ILO.

(RS ene

SMC E 12

ON VARIATION IN A JELLYFISH. 1045

47. On Variation in the Medusa of Merisia lyonsi. By

Cartes L. Boutuncer, M.A., F.Z.S., Lecturer on

Zoology in the University of Birmingham.

[Received May 23, 1911: Read June 27, 1911.]

(Plate LIX.* & Text-figures 222-228.)

In 1908 I published an account [1]+ of a new lacustrine

Hydromedusan, Marisia lyousi, obtained by Dr. Cunnington and

myself from the brackish waters of Lake Qurun in the Fayim

Province of Egypt.

In my paper, whilst describing the anatomy of this interesting

form, | called attention to the fact that the number of tentacles

and radial canals in the medusa stage was subject to much

variation, and mentioned that in a series of 400 individuals

which I examined, 55, or nearly 14 per cent., differed from the

normal.

My description of the abnormal specimens was very short,

and it has been suggested to me that it would be of interest to

furnish a more detailed account of the variation of this medusa,

as well as to figure some of the more peculiar abnormalities.

This I was all the more prepared to do, as further study of the

collection had revealed additional points of interest in connection

with this phenomenon.

Variation is known to occur frequently in jelly-fishes ¢, and in

some species it has been very carefully studied ; although in many

cases the series dealt with were numerically far greater than the

one at my disposal, I know of no form in which such a variety of

abnormalities occur as in Merisia. The interest of the series I

am about to describe is not diminished by the fact that all its

members were collected in one locality and belong to the same

sex.

As the greater part of my material had already been distributed

when I decided to take up this subject again, it was necessary to

re-examine the specimens in the Natural History Museum,

London, and in the Cambridge Museum of Zoology; for permis-

sion to do this, I have te thank Mr. R. Kirkpatrick and Mr. L.

Doncaster, under whose charge the specimens are preserved in

these institutions.

In my description of Marisia lyonsi, I drew up the following

table to show the number and arrangement of the radial canals

* For explanation of the Plate see p. 1056.

+ The figures in brackets refer to the List of References on p. 1055.

t Cf. List of References on p. 1055.

Proc. Zoou. Soc.—1911, No. LX XI, 71

1046 MR. C. L. BOULENGER ON

and tentacles in 400 meduse taken at random from the material

at my disposal :—

Table showing the Number and Arrangement of the Radial Canals

and Tentacles in 400 individuals.

Number of Tentacles.

Sumter |Syeet of} ——— :

Individuals. Canals. | Perradial. ‘Interradial.|Adradial.! Subradial. | Total.

1 3 3 = = — 3

345 4. 4 — — — 4,

1 4 4 1 — _ 5

1 4 4 1 2 = 7

i 4 4, = 1 — 5

2 4 4 2 4 — 10

10 4 4 4 = = 8

28 4 4 4 8 =m | 16

1 4, 4 4 8 6 | 22

5 5 — — — | 6

1 6 6 — — = 6

Total 400

It will be noticed that the 55 abnormal individuals fall naturally

into two well-marked groups :—(@) which includes those medusze

which deviate from the normal tetramerous symmetry ; and (8),

which includes medusz with the normal number of radial canals

and primary, perradial tentacles, but possessing, in addition,

secondary tentacles situated between the four primary ones and

not connected with the stomach by means of radial canals.

It is my intention to discuss these two groups separately, as

it is obvious that in them we are dealing with two completely

distinct phenomena.

(a) The normal medusa of Merisca lyonsi is provided with

four radial canals at the distal extremities of which the four

perradial tentacles are given off. The gonad-bearing region of

the stomach is produced into four perradially situated pouches

which in the adults extend as finger-shaped diverticula for a

considerable distance along the radial canals.

Eleven specimens out of the 400 medusze examined (7. e. 2°75

percent.) were found to deviate from this general tetramerous

symmetry ; one medusa possessed only three radial canals, three

tentacles and three gonadial diverticula ; nine meduse had five, and

one had six of these structures. This type of variation is known

to occur in many species of meduse which are normally tetra-

merous in symmetry ; among craspedote forms it has been studied

best in Obelia (Hucope) [2], Clytia [3], Sarsia [4, 5], Rathkea

(Lizzia) [7], Podocoryne [8], Gonionemus [8], and Limnocnida [9].

VARIATION IN A JELLYFISH. 1047

Limnocnida being another African lacustrine form is of

particular interest. Giinther found that out of 70 individuals

collected by Dr. Cunnington in Lake Tanganyika, 54 meduse

showed the typical number (four) of radial canals, whilst 16, or

24 per cent., had five or more ; as in the case of Marisia, specimens

with five radial canals were commoner than those with a larger

number. Ginther, however, found no meduse with only three

canals, but such trimerous forms have been described in other

genera, e.g. Podocoryne [8| and Rathkea |7].

Text-fig. 222.

Merisia lyonsi.

Diagrams to show the arrangement of the tentacles im four medusze.

B represeuts the normal tetramerous form.

Similar variations occur in several members of the Scypho-

medusz: perhaps the most complete account of the phenomenon is

that given by KE. T. Browne for Aurelia aurita [11, 12]. This

author examined the variation in the number of tentaculocysts

in the ephyra larve and adults of the jelly-fish, and found that

713

1048 MR. C. L. BOULENGER ON

20°9°/, and 22°/,, respectively, were abnormal, the number of

these sense-organs ranging between 6 and 15, the normal number

being, of course, 8. ‘These figures, however, include a number of

specimens which are undoubtedly teratological monstrosities.

Browne pointed out that variation in the number of tentaculo-

cysts does not necessarily interfere with the other organs of the

body, but that there is a correlated variation between the number of

genital pouches and buccal arms as shewn by eight specimens :—

4 individuals had 3 genital pouches and 3 buccal arms, 1 had 5

pouches and 5 arms, and 3 had 6 of these organs.

In Merisia there must necessarily be a correlation between

the number of primary tentacles and radial canals, since recent

researches on the development of the gonophores of Hydromedusze

[13, 14] have shown that these structures arise together from the

endodermal pouches of the young medusa-bud,

There is also a correlated variation between the number of

radial canals and gonadial pouches, and this is only what we

should expect as the latter structures spread outwards from the

stomach on to the radial canals. ‘The single 6-rayed specimen,

however, had one gonadial pouch very poorly developed although

the other five were quite normal.

It is interesting to note that variation in the number of

primary radii in the medusa does not necessarily affect the

symmetry of the manubrium; this point is well shown in the

sections figured on PI. LIX.

A normal medusa with four tentacles, radial canals and

gonadial diverticula has the distal part of the manubrium

provided with a similar number of conspicuous endodermal

ridges or teniole, which can be seen in the section figured

Pl. LIX. fig. 1); a similar section through a pentamerous

individual (Pl. LIX. fig. 2) demonstrates the fact that, although

five canals and gonads are well developed and the manubrium

appears almost pentagonal, the number of endodermal ridges is

not affected, although their symmetrical arrangement is disturbed.

The above is in accordance with the observations of other

investigators :—Agassiz and Woodworth [2] examined 4000

specimens of Obelia (Hucope), but noted no variations in the

shape of the digestive cavity or in the number of the actinal lobes

of the manubrium, even in specimens with five or six radial

canals in place of the normal number (four) the actinal lobes

being always found to be four in number.

A. G. Mayer [3] has also made some interesting observations

which bear on this point. This author has made a careful study

of the medusa Pseudoclytia pentata, the only Leptomedusan

which is normally pentamerous, and which he considers to be

derived from some species of Clytia (Hpenthis), e.g. C. folleata.

In Pseudoelytia pentata there is much variation both in the number

of radial canals and in the number of oral lips; but whilst the

former incline towards the production of more than five canals,

the oral lips show a decided tendency to revert to the ancestral

condition of four.

VARIATION IN A JELLYFISH, 1049

As mentioned before, variation in the nwouber of radii is

of common occurrence in meduse, and in the above-described

varieties of Merisia lyonsi we are undoubtedly dealing with

a quite ordinary case of meristic variation. As Bateson remarks

in his well-known book on the Study of Variation [15] :—

“In radial series phenomena analogous to those of the variation

in linear series are seen in their simplest form. Just as in linear

series the number of members may be changed by a reconstitution

of the whole series so that it is impossible to point to any one

member as the one lost or added, so may it be in the meristic

variation of radial series: and again as in linear series, single

members of the series may divide. Between these there is no

clear line of distinction.”

In the abnormal specimens of Mwrisia, I do not think that we

are dealing with cases of division or suppression of one or more

radi ; from what we now know of the development of medusa-buds

[13, 14], it seems that the variation is in the number of radial

pouches formed in the early bud.

Since each pouch develops the radial canal and perradial tentacle

of its own section of the medusa-bell, such variation explains the

correlation of the numbers of these two sets of organs. The fact

that the manubrium is independent of the organs situated in the

umbrella is a point decidedly in favour of this view. I have,

unfortunately, not been able to study the development of an

abnormal specimen, all the medusa-buds I have sectioned proved

to be of normal tetramerous symmetry.

(>) The second group of abnormal individuals includes a

number of tetramerous medusze which bear secondary tentacles

between the four primary perradial ones. These secondary

tentacles may be interradial, adradial or subradial in position,

and differ from the perradial ones in not communicating with the

gastric cavity by means of radial canals; they are developed from

the ectodermal and endodermal cells of the umbrella-margin, their

cavities being in communication with the circular canal.

Altogether 44 of the 400 tabulated individuals exhibited this

kind of “abnormality, which is of a very peculiar type ; I have not

been able to find any record of a similar case in a medusa which

normally bears primary tentacles only. There are, of course,

numerous tetramerous meduse which normally posssess. such

secondary tentacles (e. g., Podocoryne, Turritopsis, Oceania,

Limnocnida), and they are known to start life with the fours

primary ones only, the secondary tentacles being developed as

the animals grow, usually in some definite sequence. Thus, in

the majority of cases, the second set of tentacles to appear is the

interradial one, four of these structures growing simultaneously

from the umbrella-margin between the four perradial tentacles.

Hight adradial tentacles are formed next, two in each quadrant

occupying the interspaces between the interradial and perradial

ones, and their formation may be followed by the appearance of a

varying number of subvadial tentacles.

1050 MR. C. L. BOULENGER ON

In such medusz it is obvious that the number of secondary

tentacles is correlated with the size of the individuals, the older

and larger meduse possessing a greater number of these structures

than the smaller and younger ones. In Merisia we are certainly

Text-fig. 223.

Medusa bearing four interradial tentacles im addition te the fenr perradial ones at

the extremities of the radial canals. In this and in the three following text-

figures A represents the medusa as seen in a side view, B its oral aspect.

Text-fig. 224,

Medusa with four interradial and eight adradial tentacles in addition

to the four primary ones.

not dealing with a phenomenon of this kind, for the larger and

more mature meduse are, with rare exceptions, unprovided with

secondary tentacles. In order to show this point more clearly,

I divided the 400 individuals which I examined into two

VARIATION IN A JELLYFISH, 1051

groups :—(i.) including meduse with an umbrella-diameter

measuring between 3 and 2 mm., and (ii.) with a diameter

measuring from 24 mm upwards,

Group (i.) contained 278 individuals of which 39, or about

14 °/,, possessed supernumerary tentacles between the four

primary ones, whereas of the 122 larger medusze belonging to

group (ii.) only 5 were provided with such structures.

Sexual dimorphism is known to occur in a few species of

medusze,* but I have assured myself that this is not the case

with the specimens I am describing; microscopic examination,

by means of sections and whole mounts, showed that all the

meduse’ of Merisia in my collection belonged to the male sex,

whether bearing secondary tentacles or not.

It must be obvious from the above account that the kind of

aviation I have just described is of a nature totally different from

that treated of in the first section of this note. We are certainly

not dealing with a case of ordinary meristic variation, and the

only conclusion I can arrive at is that these multitentacular

meduse form a distinct variety which has arisen as a mutation

from the normal form : it will be interesting to discover whether

this variety will maintain itself in Lake Qurun.

Altogether, forty-six f multitentacular specimens have been

examined by me; it was found that the number of secondary

tentacles varied considerably. 10 individuals possessed four

interradial tentacles as well as the four primary perradial ones

(text-fig. 223); 28 had twelve secondary tentacles, four inter-

vadial and eight adradial (text-fig. 224); and 3 possessed sub-

vadial tentacles in addition to these, the medusa illustrated in text-

fig, 225 (p. 1052) having as many as eight of these structures, two

in each quadrant, situated between the interradial and adradial

tentacles,

The sketches referred to above, although considerably enlarged,

are drawn to scale; they exhibit the fact that this variety of

Merisia lyonsi follows the general rule for multitentacular

meduse, in that the number of secondary tentacles increases

with the size and age of the individuals, The sequence of

the development of the secondary tentacles is also quite normal

in the specimens just described,

Although the majority of the multitentacular specimens of

Merisia exhibited a perfect radial symmetry and developed

their secondary tentacles in a perfectly normal order, a few

medusz proved abnormal in this respect and showed marked

asymmetry. One specimen had a single interradial tentacle

developed between two perradial ones, whilst another bore

interradial and adradial tentacles also in a single quadrant only.

Again, two meduse had these organs developed in two of the

quadrants (text-fig, 226), and a third had secondary tentacles in

* H.e. Stomotoca dinema Agassiz, and Orchistoma pileus Lesson : see Mayer (10)

pp. 111 and 219.

+ T'wo of these specimens were not included in the Table on p 1046.

MR, C. L. BOULENGER ON

1052

Text-fig. 225.

ary tentacles.

acular medusa with twenty second

Multitent

Asymmetrical medusa bearing secondary tentacles in two quadrants only.

VARIATION IN A JELLYFISH. 1053

three quadrants. This specimen had interradial, advadial and

subradial tentacles in two of these, whilst the third quadrant

carried only a single adradial one *.

Giinther has shown that in Limnocnida [9], tentacles of a

particular order are often fully formed in one quadrant before

there is any trace of them in the others. Browne also has called

attention to the fact that in Podocoryne carnea [7] the young

medusz do not always leave the hydroid colony with the same

number of tentacles, all have four perradial ones but the number

of interradial tentacles varies, some having two or three instead

of four, and one specimen he noticed to have a single one only.

I do not think that in the case of the asymmetrical medusz of

Merisia to which I have just referred, we are dealing with any

retardation in the development of the tentacles in certain

quadrants ; some of these individuals were of considerable size

and the gonadial diverticula well developed (ef. text-fig. 226), yet

certain quadrants showed no signs whatsoever of secondary

tentacles, although other quadrants possessed tentacles of the

third or even fourth order.

These five asymmetrical meduse clearly indicate that each

quadrant is capable of forming secondary tentacles quite

independently of the other quadrants of the bell. That this can.

occur in a radially symmetrical animal is distinctly stated by

Bateson [15] who remarks, that in radial series ‘as in Linear

Series, Variation, whether Meristic or Substantive, may take

place either in single segments (quadrants, sixths, etc.), or

simultaneously in all the segments of the body.” This statement

was based on observations made on two meduse, Clavatella

(Hleutheria) prolifera and Aurelia aurita.

Clavatella is a medusa which normally bears a single ocellus

at the base of each of its six tentacles. Claparéde [16] has called

attention to the fact that these ocelli are sometimes doubled ;

this duplicity may occur at the base of a single tentacle or

occasionally at the base of each tentacle instead of one.

Bateson also quotes the observations made by Romanes [4, 5]

on Aurelia aurita. In this form, in addition to changes symmetri-

cally carried out in the whole disc, one or more quadrants may

vary independently. Thus one specimen is figured in which two

quadrants are normal (7.e., each possesses one generative organ

and a set of radial canals) but the other half-disc is divided into

three. Similarly a particular quadrant may possess two sets of

organs or even three, the other three quadrants being normal or

nearly so.

In addition to the forms described above, three other medusze

showed abnormalities of a quite different type, which, although

known to occur in other genera, I consider worth recording.

In the first of these specimens, which was normal as regards

* This specimen is one of those not included in the Table published in my

original paper on Merisia lyonsi.

1054 MR. C. L. BOULENGER ON

the number of its tentacles, one of the latter organs was branched

in such a way as to present the appearance of a smaller tentacle

growing out laterally from a perradial one.

Text-fig. 227,

Diagrams showing the arrangement of tentacles in a normal and

five abnormal medusee of Merisia lyonsi.

P. Perradial tentacle. J. Interradial tentacle. S. Secondary tentacle in an

adradial position. ‘The subradial tentacles are not lettered.

VARIATION IN A JELLYFISH. 1055

Such branched or bifureated tentacles have been recorded pre-

viously by several authors, among whom Agassiz and Woodworth

[2| have described the phenomenon in medusz of Obelia (Hucope),

and Hargitt [8] came across a similarly abnormal specimen of

Gonionemus.

It seems highly probable that such abnormalities are not

congenital, these bifurcated tentacles having no doubt arisen as

the result of injury to normal ones.

Text-fig. 228,

Abnormal medusa in which the apicai canal is retained.

}

The abnormality presented by the other two medusz is of

greater systematic interest: each of these specimens possesses a

well-developed apical canal which projects from the base of the

stomach into the jelly at the summit of the umbrella (text-

fig. 228). This peculiarity is obviously due to the fact that these

specimens have retained the greater part of the canal which in

early life connected the cavity of the medusa-bud with that of the

parent-hydroid.

The presence of an apical canal was at one time considered to

he a specific character of some importance, but recent systematists *

have shown that such a canal occurs frequently as an individual

variation in many species which normally lose this organ in the

adult stage.

List of References.t

1. Boutencer, C. L.—“ On Meorisia lyonsi, anew Hydromedusan

from Lake Qurun.” Quart. Journ. Mier. Sci., vol. li. 1908,

pp. 357-378.

* (Cf, Mayer's remarks on Sarsia prolifera, and other species of this genus (10).

+ his list includes only those memoirs actually referred to in the text of this

paper; for a more complete bibliography I must refer the rcader to Dr, Mayer’s

recently published monograph (10).

1056 ON VARIATION IN A JELLYFISH.

2. Acassiz, A., & Woopworru, W. M.—‘“Studies from the

Newport Marine Laboratory: XL. Some Variations in the

Genus Hucope.” Bull. Mus. Comp. Zool. Harvard Coll.,

vol, xxx. 1896-7, pp. 121-150.

3. Mayer, A. G.—‘ The Variations of a newly-arisen Species of

Medusa.” Mus. Brooklyn Inst. Sci. Bull., vol. i. 1901,

pp. 3-27.

Romangs, G. J.—‘‘ Varietiesand Monstrous Forms of Medusee.”

Journ. Linn. Soc., Zool. vol. xii. p. 527.

Romangs, G. J.—Loc. cit. vol. xiii. p. 190.

Agassiz, A.— Meristic Variation in Sarsia.” Mem. Amer.

Ac. Sci. vol. iv.

Browne, E. T.—‘“‘ On British Hydroids and Meduse.” Proc.

Zool. Soe. London, 1896, p. 459.

. Harerrr, C. W.—‘* Variation among Hydromeduse.” Biol.

Bull. Wood’s Holl. Mar. Lab., vol. 11, 1901, pp. 221-255.

. GintHER, R. T.—* Zoological Results of the Third Tan-

ganyika Expedition conducted by Dr. W. A. Cunnington,

1904-1905.— Report on Limnocinida tanganice ; with a Note

on the Subspecies from the Victoria Nyanza.” Proc. Zool.

Soe. London, 1907, p. 643.

10. Maver, A. G.—‘‘ Medusz of the World.” Carnegie Inst. of

Washington, Publ. No. 109, 1910, vols. 1.-111.

11. Browne, E. T.—‘ On the Variation of the Tentaculocysts of

Aurelia aurita.” Quart. Journ. Micr. Sci., vol. xxxvii.

1894-5, p. 245.

12. Browne, KE. T.—“ Variation in Auvwrelia.” Biometrika,

London, vol. i. 1901.

13. Gorrz, A.—‘ Vergleichende Entwicklungsgeschichte der

Geschlechtsindividuen der Hydropolypen.” Zeit. f. wiss.

Zool., vol. Ixxxvii. 1907.

14. Bounencer, C. L.—‘ On the Origin and Migration of the

Stinging-Cells in Craspedote Meduse.” Quart. Journ. Micr.

Ste, VO lhe UGIO, yo, (ike

15. Batrreson, W.—Materials for the Study of Variation. 1894,

pp. 422-429.

16. CLiararkpE.—Beobachtungen iiber Anatomie und Entwick-

lungsgeschichte Wirbelloser Thiere, 1863, p. 5.

Oo OM OO

EXPLANATION OF PLATE LIX.

Fig. 1. A transverse section through a normal adult medusa of Merisia lyonsi.

The section shows the four radial canals and gonadial diverticula, as well as

the four endodermal ridges in the manubrium.

end., endodermal ridge. g.p., gonadial diverticulum. /am., endodermal

lamella. man., manubrium. ~7.c., radial canal.

Fig. 2. A similar section through a pentamerous specimen of the same species.

Note that although five radial canals and gonadial diverticula are well

developed, only four endodermal ridges occur in the manubrium,

Cf, p. 1048.

Lettering as in fig. 1.

12) Ay Sy USL IPE AS

Huth, Lith? London.

MARGINAL PROCESSES IN LAMELLIBRANCH SHELLS.

ON MARGINAL PROCESSES IN SHELLS. 1057

48. The Marginal Processes of Lamellibranch Shells.

By Cyrim Crosstanp, F.Z.8.

[Received May 27,1911: Read June 27, 1911.]

(Plate LX.* and Text-figs. 229 & 230.)

The existence of the ornamental projections on the surface of

so many lamellibranch shells is rather puzzling to the student

of Bionomics. Especially to one interested in the rate of growth

of mother-of-pearl sheli does their formation seem a sad waste of

shell-making energy.

The drawings given show examples of these processes in young

and adult shells. Inspection reveals one striking fact, that the pro-

portion of shell-building energy thus expended is very much greater

in quite young shells than in the mature ones of several species.

Compare, for example, the figure of a specimen of Margaritifera

Text-fig. 229.

Shells of Murex ramosus and upper valve of adult Chama foliata.

Half natural size.

The illustration will serve for identification of these common shells by

non-conchological readers.

margaritifera (the large mother-of-pearl oyster) seven months old

and that of the portion of the margin of one 54 years old, also the

young of Chama foliata and Chama sp. of about the same age

with the adult shells, and it is seen that relatively the processes

are gigantic in young shells (Pl. LX. and text-fig. 229). They

must have their use, or they could not be so greatly developed at

this, the second, critical stage of the bivalve’s existence Tf (the first

.* For explanation of the Plate see p. 1061.

+ Utility is suggested also by the fact that such processes cannot be regarded as

ancestral features such as in some other cases are more prominent in the young of a

species.

1058 MR. CYRIL CROSSLAND ON

critical stage is that during which the free-swimming larva must

attach itself to a suitable substratum or perish). At this second

stage the Aviculide at least are in the most active state they

attain to, and most other normally sessile lamellibranchs have

well-developed and actively used feet. Zridacna gigas is very

active at this stage, to mention a form probably not often seen

when so young, and particularly well fixed down when adult.

Their future life depends upon their success in crawling into the

position adapted to the special needs of their species. Also they

are now large enough to be attractive as food to fishes and erabs,

but not strong enough to resist claws and pincers. In the case

of M. margaritifera, among other species, the animal has little

to fear after it is one year old, being then too strong-shelled for

ordinary fishes and crabs.

Text-fig. 230.

Two specimens of Avicula zebra attached to Millepora alcicornis, and in the centre

of the latter a specimen of another genus of the Aviculidee which is more

common on corals. Another species of Avicwda is found on “black coral,”

Antipatharia. Half natural size.

Balistes flavimarginatus and B. viridescens, the largest of a

genus of shell-eating fish, are fairly frequently found here * in the

Red Sea, and occasionally break up ‘“ pearl” oysters two or three

years old, but those dissected were found to have preferred smaller

and weaker-shelled species. The big Rays are not common, and

* Dongonab, Port Sudan.

MARGINAL PROCESSES IN SHELLS. 1059

I have not certainly traced any damage to them. T'ridacna,

Spondylus, and Chama similarly have every chance of reaching

old age once the younger stages are passed, and the majority of

specimens found are thick and heavy ; on the other hand, Mar-

garitifera vulgaris (the Ceylon pearl-oyster), Ostrea sp.%, and

Avicula zebra of a year or two old are common, but above this

age are scarcely ever met with in the Red Sea. In the only

fishery for Margaritifera vulgaris which I have seen here, shells

two ox three years old, and small for that age, were being obtained

in comparatively insignificant numbers.

That energy so valuable elsewhere should be wasted upon un-

necessary appendages is also negatived by the consideration that

in this class, as elsewhere in nature, the struggle for existence is

severe, demanding the nicest adaptations to environment. This

is shown by the existence in this one bay of three distinct species

of Margaritifera, anatomically scarcely distinguishable, apparently

living together in the same habitat and obtaining food in

exactly the same way. Only after careful study does it begin to

be seen what delicate adaptations to special environments keep

the three species distinct, each in its own niche in the world.

Observations of the relations between living examples of the

following species and their common foe, Murex ramosus (text-

fig. 229), shows that these ornamental processes have a simple and

essential use to their possessor; indeed, without them they would

fali such easy victims to predatory Prosobranchs that those species

not otherwise protected must become extinct.

The following species living in exactly the same surroundings,

in the same artificial way, were observed, and their mortality

from attacks by Murex noted :—

Ostrea sp.% ...... No spinous processes. _ Decimated.

Avicula zebra... - oy 55

Margaritifera Processes small and Killed in large

mauritir. weak. numbers.

Margaritifera Processessmall,except Keptalive by frequent

vulgaris. in quite young shells. removal of Jurez,

otherwise numbers

perished.

Margaritifera Processes large and Attacked only excep-

margaritifera. strong, remain well tionally.

developed to at

least 6 years old.

We see that in these five species the liability to attack by

Murex (and probably by other prosobranchs, the behaviour of

which I have not observed) is just in proportion to the develop-

ment of spines. ‘The cases of attack upon J/. margaritifera

which I have seen corroborated, occurred only in stunted shells

in which these processes were ill-developed.

In the Aviculide at least the processes are made from the two

1060 MR. CYRIL CROSSLAND ON

outer shell-layers only, and as both of these are weak and horny,

so are the processes. These consequently get worn off the older

parts of the shell, and as increase in diameter becomes slow so

does the development of these processes* and in full-grown

shells practically ceases altogether.

It is obviously an advantage to the “oyster” to grow up

rapidly, and form a shell large enough to resist teeth and pincers

as quickly as possible. MMargaritifera vulgaris succeeds excellently

in this, attaining nearly its full size in a year; but it does this at

the expense of the sooner losing its shell-processes and so being

open to attack from J/urex, and doubtless other predatory

prosobranchs.

In MW. margaritifera also growth is extremely rapid during

the first year, and at the end of it the shell is strong enough

to resist such crabs and ordinary fishes as infest this bay; but

after the second year it is slow, a shell six years old being far

from full-sized. Consequently the formation of marginal pro-

cesses continues later, and with it partial immunity from attack

by durex. Sexual maturity is reached by all these species alike

in the second year.

From the result of my experiments recorded above, one wonders

how the first three species can possibly survive in Nature. The

explanation is very simple: they choose a habitat out of reach

of their foe. Ostrea sp.? and Margaritifera mauritiit inhabit

crannies among other shells £ or in stones, too narrow for Murex

to enter; while Avicula zebra, like other species of the same

genus and a few other genera of the same family, possesses a

foot that fears not the stings of corals, nor even those of J/ille-

pora, and so attains a habitat inaccessible to all others (text-

fig. 230, p. 1058).

Avicula zebra is very common on Millepora alcicornis in the

Red Sea; another advantage of this habitat is support above the

sea-bottom, clear of all obstructions to its respiratory circulation,

which latter is of greater value than would be supposed.

Given protection from enemies, and from being smothered in

mud, all these species will flourish wherever the experimenter

likes to place them, these two considerations being all that

causes the restriction of their habitats in nature.

* Only in a general sense is this true. In two specimens of M. margaritifera

of about the same age, viz. two years, one has added to its diameter much less

than has the other, owing to difference of conditions in which they grew ; but this

has not hindered development of processes, which are merely nearer.together in

the slower-growing specimens. It is when slow growth is due to age that the

formation of processes becomes imperfect.

+ This species, when over a year old, forms a postero-ventral non-nacreous addition

to the shell, similar to, but smaller than, that found in many other Aviculidz,

and which reaches extreme development in Mallews. In all, the addition is an

adaptation to enable the animal to inhabit narrow crevices, and its function is

similar to that of the siphons of those lamellibranchs which live buried in sand, or

in burrows.

{ H.g., on the under sides of large specimens of mother-of-pearl shell brought in

by the divers.

MARGINAL PROCESSES IN SHELLS. LO61

The way in which these processes protect the lamellibranch

has yet to be explained. From actual and repeated observation

one would expect that the prosobranch could extend its proboscis

for a short distance and insert it amongst the processes, which

are not generally very near together, and so drill its hole without

trouble. It is, however, necessary for the Murex to obtain a

very firm hold with its foot before it can operate its drill, and

one finds cases where a distinct impression of the foot has been

made among the growths on the shell before any damage has

been done at all. The point of special interest to the naturalist

as well as to the practical pearl-fisher, is that one cannot estimate

the damage done by Murex by counting how many of a lot of

dead shells have the characteristic hole drilled through them.

In move than three cases out of four no hole is drilled at all,

Murex having found an easier way. It finds the flexible edge of

the shell, then by contractions of its foot breaks a piece away.

The mucus of the foot-glands is then poured out in quantities,

and this has some poisonous effect, as the animal, while still

untouched, ceases to respond to the stimuli which ordinarily

cause a smart closure of the shell. The abundant flesh of this

species is all devoured before putrefaction sets in.

EXPLANATION OF PLATE LX.

Except for figs. 6 and 7, specimens of about the same age have been drawn to the

same scale, the age being something under six months, the magnification 3.

Fig. 1. Chama foliata. A dead shell, widely gaping, seen from above, i.e. from

anterior end. a, hinge; b, ventral edges of shell.

The shell-processes are seen to be about half the diameter of the shell in

length. At this age they and the shell ave brilliantly coloured with dark

red bands on a lght yellow ground. In the adult shell the processes

remain conspicuous (hence the specific name), but in the young they are

out of all proportion to the size of the animal that forms them.

Fig. 2. Upper valve only of Chama sp.? Shell red, processes white. In adult shells

processes quite inconspicuous, generally absent. | Animal then protected by

the extreme solidity of its shell.

Fig. 3. Spondylus sp.? A specimen which is beginning to take the adult shape

and to form the characteristic strong spatulate processes instead of the

brittle needle-lke serrated processes characteristic of the younger stages.

These serrulations are shown in oniy a few instances in the figure.

Fig. 4. A rather younger specimen of the same, with long slender processes only and

edge of shell serrated. Upper valve only drawn.

Fig. 5. Margaritifera margaritiferu. A specimen 12 mm. across shell, with

processes 7 mm. long. Probably three months old; younger specimens

than this have perfectly smooth shells like the umbonal region of the

present specimen.

Fig. 6. The same, X2, aged about 7 months. Showing development of processes

characteristic of the first two or three years’ growth. Some of the radial

rows of processes are thick and spatulate; others, which are thinner, shrivel

and become pointed, as in the figure, on drying.

Fig. 7. Thesame. Part of the edge of a three-year old shell which has grown slowly.

Shells more rapidly developed are distinguished by the wider spacing

of the processes.

Proc. Zoou. Soc.—1911, No, LX XII,

1062 MR. CYRIL CROSSLAND ON

Warning Coloration in a Nudibranch Molluse and in

a Chameleon. By Cyrit Crossuanp, F.Z.8.

[Received June 12, 1911; Read June 27, 1911. ]

I. WARNING COLORATION IN CHROMODORIS.

Since the discovery of warning coloration there has been a

tendency toattribute protective value to displays of brilliant tints

which further observation has rendered doubtful, and the instances

in which actual protection has been experimentally proved are not

so numerous but that one more has some value.

The Chromodorids are, as their name implies, a family of Nudi-

branchs characterised by the development of colour shown by all

its members. Having collected a large number during the past

ten years, I may say that none of the family has been wanting in

this characteristic, except perhaps Casella atromarginata, which,

though handsomely, is not brilliantly, tinted. In this I merely

corroborate the general experience of marine collectors. Besides

this universality of colour the family is well defined structurally ;

indeed for the class the structure is remarkably uniform. The

usual depressed form of body, more or less ample margin, broad

foot, and feathery, usually tripinnate gills, are, in the more typical

Chromodorids, replaced by the opposite characters. A narrow foot

underlies a high body, the mantle is a mere shelf along the top

of its vertical sides, and the gills are simply pinnate *. Internally

the stomach is entirely embedded in the liver.

here is a smaller section of the family in which these char-

acters are less marked, the body having a wider mantle; in some

species a few of the gills are huaralived), and the Promveli Is partly

free from the liver, ‘aun indicating « connection with the ordinary

forms.

Otherwise structure is so uniform throughout the group, that

without detailed description of the colours of the living animal

identification of the species is impossible. Even with this deserip-

tion in full, determination of species is difficult, variation being

very Conetiemible so that, e.g., what one observer sees as black

bands on a white ground is to another white lines on a_ black

body ft.

* Arve these simply-pinnate gills the primitive form of Nudibranch gill, a modifi-

cation of the prosobranch ctenidium The highly specialised Chromodovids supply

the answer, since it is the less specialised members of the family that have the more

complex gills. A detailed study of the Chromodorid gill would show that its struc-

ture is more complex than the term “ simply pinnate ~ leads one to suppose, and

possibly would show details of higher organisation than do the tripinnate feathery

eills of ordinary Dorids. In the dev elopment of these gills from an originally irregular

vascular flap of skin, the tripinnate arrangement would be the earlier stage to be

reached.

+ Chromodoris quadricolor Eliot, Journ. Linn. Soc., Zool. xxxi. Nov. 1908, p. 107

= 0. elizabethina, P. Z.S. 190 L, p. 392, Pl. xxiv. fig. 4. Some other Chromo-

dorids are here figured including Cn igrostriata, a variety of the C. diardii

referred to later in this paper and a good example of the variation now

discussed,

WARNING COLORATION, 1063

The family is therefore probably monophyletic (so far as any

family can be), and it would seem that the power of colour

production is a single character, which, in a few of the cases in

which it has arisen, was associated with distastefulness or other

protective quality, and so, possessing a great survival value, led to

the foundation of the family. What colour or pattern might be

produced selection cared not, so long as it were conspicuous in the

eyes of nudibranch enemies, and duly correlated with inedibility *.

Hence the multitude of patterns and colours found in this one

class, and the variation above mentioned. Possibly one colour

is much the same as another in the biochemistry of the animal,

though so different to the human eye T

The survival value of the devices is evident from the abundance

of species in tropical waters. In collections, while species are

numerous, Individuals are usually rare, but the peculiar habits of

many nudlibranchs, which suddenly appear in local swarms and

shortly disappear again completely, show that as in, e. g., the case

of the Sharks of St. Andrews Bay %, which were not known to

exist there until new fishing methods were introduced, the pro-

portions in which species occur in collections are not at all those

in which they live in the sea, Of the Chromodorids [ remember

par ticularly hs annulata, C. sykesi, C. hilaris. and Ceratosoma

corniyerum as occurring for a few days in quite local swarms,

then dis sappearing absolutely ; while during the past fortnight

a hundred specimens of C. reticulata | PI. axel (p. 1068)] could

quickly be collected within a few yards of where [ write, but

to-morrow there may be not one.

The collection would be the more easy in that this species like

all Chromodorids crawls about in the open instead of skulking

under stones or in crannies after the fashion of most of the class,

except of course at low tide when it would otherwise be killed by

the sun,

In contrast with this, the general inconspicuousness of most

Opisthobranchs culminates in many cases in adaptations which

result in resemblance to surroundings so striking and so well in

aecord with the observed habitat, that there can be no doubt as to

protection being afforded by them §,

* Amidst this diversity we find that in all Chromodorids the edge of the mantle is

specially coloured. In others the gills and rhinophores are coloured differently trom

the rest of the body, as in the case of C. diardii,

+ That the colour of marine animals is often connected with their essential physi-

ology or apparatus of reproduction is dicated by the many cases in which the eggs

of a species are deeply coloured with the principal body colours of the parents.

This is conspicuously the case with numerous species of Polychwta and Nudibranchs,

The Chromodorid hereafter described under the name of C. reticulata lays eges the

colour of the yellow bands on the mantle. The purple band secretes macus of the

same tint,so that probably the purple colour has its physiological use, as well as

warning colour value.

+ McIntosh, W.C., “ Notes from the Gatty Marine Laboratory,” Ann. Nat. Hist,

yol. x. p, 254, 1902.

§ I have never had the fortune to possessa properly stocked aqnarium in which

experiments on this side of the theory conld be undertaken,

e 7

1064 MR. CYRIL CROSSLAND ON

Two interesting facts are worth recording of the coloration of

several species of Tectibranchs, which besides being close simu-

lacra of their environment in other respects, each occur in green

and brown varieties, simulating green or brown weeds, or at least

in one case, the old and young shoots of Zostera. Living speci-

mens of these species are not now within my reach or further

details might be given, and sketches. The same thing happens in

a Scy llaeid*, the The ager green and the brown forms of which are

found near together i in Chuaka Bay, Zanzibar, whilst greenish to

deep brown varieties occur here (Dongonab) in the Red Sear The

other point about the coloration of these forms (Scyllaeid and

Tectibranchs) is the occurrence in so many species of small brilliant

blue spots, sometimes ringed with yellow, the only conspicuous

thing about otherwise scarcely visible creatures. These may be

a recognition marks ” but are more likely to be glands &.

We have then an excellent case for the theory that the brilliant

colour of these animals has a protective value. They walk abroad

where others must creep and hide.

Distastefulness was proved by trying to feed fish with specimens

(mostly CO. reticulata, twice with CO. diardii) thrown from the

windows of my homsebanti The kitchen refuse, and the shelter

afforded from the sun had made the surrounding water populous

with fish, and on a calm morning every detail of their movements

was clear. The water was shallow, the bottom sandy, with a

covering of short ‘sea grass,” the usual habitat of several fish and

often of considerable numbers of Chromodoris reticulata, more

rarely of CO. diardii.

The Garfish, Belone sp., is the only one of these animals with a

specialised diet. It is ever on the watch floating near the surface,

herding together the shoals of “ sardines” (Zngraulis ? boelama),

* Crosslandia viritis Eliot, P. Z.S. 1902, pp. 64-68; C. fusca Khot, Journ. Lim.

Soc., Zool. xxxi. 1908, p. 90.

+ The green Zanzibar specimen was found on Zostera, which is less abundant in

the Red Sea and from which I have not yet obtained Als species.

t Other brilliantly coloured Nudibranch families are the Polyceride and Aeolide.

OF the former I have seen too few specimens to generalise, but one of the most beau-

tiful, the wine-red Plocamopherus ocellatus, was merely a lighter tint of the colour

of its environment, a deep red polyzoan on which it ted. There is some evidence that

it owes part of its ‘colour to its foodstuff! Another, Treveleyana crocea 2, isa little

slug-shaped beast of a brilliant yellow colour. It periodically occurred in great

swarms in Chuaka Bay, Zanzibar, in the open, not hiding in any way. The Aeolids

are all bright coloured, I believe, and all, or nearly all, gain protection by living

amongst hydroid zoophytes, and even can utilise the stinging- thread cells of the

latter in their own bodies.

Tritonids often live among zoophytes, in Zanzibar often among fleshy Alcyonaria,

and are fairly conspicuously coloured. But Melibe fimbriata, a large animal, in colour

and in the shape of its cerata and the processes it bears, looks extremely likea

fragment of drifting fucus-weed.

1 Biot, Sir C., “Marine Biology of the Sudanese Red Sea,” Journ. Linn. Soe.

xxxi. Nov. 1908, p. 104.

2 Bliot, Sir C., “ Nudibranchs from Hast Africa and Zanzibar,” P. Z. 5.1904, vol. ii.

p. 87.

WARNING COLORATION. 1065

occasionally making dashes into the shallow water and carrying off

stragglers. Yeteven elone dashes up for any bait thrown into the

water, though the portions that sink below the surface are left to

the numerous bottom feeders. Of these we have Chrysophrys bifas-

ciata, Pimelepturus cinerescens, and two other species which I have

not identified, and they will eat, or have become accustomed to

eating, almost anything, including shelled specimens of Margariti-

fera vulgaris which had been kept in formalin for three weeks, and

the guts of Balistes flavimarginatus and B. viridescens which had

been in the same preservative overnight. Pimelepturus eats the

fresh viscera of its own species, or the pure white cooked flesh of

Caranx sp. equally readily. But none of these five fishes will eat

either of these two species of Chromodoris; attracted by the splash

they dashed up to them as to any other bait, but one fish after

another, of each species, as the nudibranch sank, at once turned

away after just touching it. Examples of Chrysophrys took the

animals into their mouths, but at once dropped them undamaged.

The behaviour of Pimelepturus was particularly interesting. If

a specimen of C. reticulata, which is white underneath (but for a

few purple spots under the edge of the mantle), fell on the sand

wrong side up, the fish would at once attempt to take it. If, how-

ever, the slug fell right side up, so displaying its gorgeous mantle,

Pimelepturus, swimming over it a few inches away, took no more

notice of it than of a stone.

What actually prevented these numerous fishes, of five distinct

species, from swallowing the Chromodorids? One can hardly

believe that a living slug could have a flavour more powerful

than that of formalin! Moreover, these two species have no odour

sensible to human organs, as have so many marine organisms,

including the Chromodorid Ceratosoma cornigerwm, and the

Tritonid Melibe fmbriata. The behaviour of these fishes, which

had for some weeks been feeding upon all kinds of kitchen

refuse, and especially the fact that when presenting an wn-usual

appearance Chromodoris momentarily attracted them, disposes

of the objection that they merely refused to eat an object which

was strange to them. Indeed, the natural circumstances of the

case make it impossible that C. reticulata should be unfamiliar

to any of these fishes except Belone.

But that this objection is a weighty one andin all supposititious

cases of protective coloration must be carefully considered, the

following examples will show.

The camels of the southern part of the Red Sea Province get

desert grazing all the year round, and are not fed upon “ dira ”

corn (Sorghum) in the summer. The consequence is that a

southern camel has actually to be taught with much patience (for

he deserves all that Kipling said about him) to eat corn. The

feast may be spread before him, but eat it he will not, until his

owner, seizing him by the nostrils, pours the corn into his mouth

with the other hand, spite of his groaning protests. Indeed some

1066 MR. CYRIL CROSSLAND ON

camels never get beyond this stage, and however hungry hold their

heads in the air above the pile of corn, crying like spoiled children

until the owner comes to feed them.

Contrast with this the behaviour of my two tame gazelles. From

six weeks old to maturity they fed on milk, corn, and occasionally

native leguminous plants, When fully adult they were introduced

to bread, sweet biscuits, sugar, green corn-stalks, and the leaves of

Acacia tortilis, all of which they ate readily without the stimulus

of hunger. I once tried to feed a sick camel on stale bread, by

way of invalid diet, It was impossible.

Another case is rather like that of the pigeons that starved

rather than eat a strange seed. I tried to change the feeding

place of my fowls by throwing their corn a yard or two to one side

of the usual place in the direction of the new one. When called the

fowls rushed up, stopped dead at the accustomed spot, and nothing

would make them move a foot beyond it. We can pair the fowls

with the opposite case, as we did the camels with the gazelles.

The land crab Ocypode makes its burrows a little above high-water

mark and never more than a yard or two higher up. But where

the camels and fowls were regularly fed, the ground is full of the

holes of these crabs ten or twenty yards above their usual habitat.

Also, instead of confining themselves to their accustomed diet of

dead fish, these lively crabs are ready for any experiment, from

sweet biscuits to handkerchiefs, and some must now be living on

camel dung and corn.

There are no oysters in Jerusalem, and consequently a native of

that city cannot be persuaded to try one. He has no religious

prejudice, or other dislike, but “ No, I do not know them ” is his

sufficient reason.

IT. WARNING CoLORATION IN A CHAMELEON.

I havenot seen the fact recorded that the Chameleon (Chameleon

sp.) ean change its colour so quickly as to frightenadog. While

staying in Zanzibar my host’s fox terrier showed hostile interest in

a chameleon someone had brought into the house. The chameleon

invariably tried to run away when attacked, but those who know

the species can imagine the ludicrous ineffectiveness of a chame-

leon’s flight. In a few seconds the impossibility of escape seemed

to reach the animal's brain, when it at once turned round, opened

its great pink mouth in the face of the advancing foe, at the same

time rapidly changing colour, becoming almost black. ‘This ruse

succeeded every time, the dog turning off at once. Among the

natural leafy surroundings of a chameleon the startling effect of its

sudden change of colour would be much greater. Imagine a dog or

cat nosing about suspicious of the presence of a live animal, but

unable to see anything. Until almost touching him the chame-

leon sits close, secure in his mantle of invisibility. Then suddenly,

WARNING COLORATION, 1067

when the dog’s nose is within six inches of him, the sudden change

from invisibility to the conspicuous black body and great red

mouth! I think many human beings would be startled, and in

fact the natives of Zanzibar hold the chameleon in horror, and

cannot be persuaded that its bite is not deadly. I tried the effect

upon some of them on my way home. ‘The ordinary native will

show great caution, but no panic, when a chameleon is handled

by a white man. I presented my specimen wneapectedly to

several parties I met on the road, thus :——

On approaching, ‘‘ Good morning,” which was quietly responded

to.

Just after passing, “See what a nice creature is walking on my

back.” Shrieks, and a stampede of fifty yards.

The specimen showed the ordinary light and irritation reactions

which have been frequently recorded.

1068 SIR CHARLES ELIOT ON

50. Chromodorids from the Red Sea, collected and figured

by Mr. Cyril Crossland. By Sir CHarues Huiov,

JS oils Giese BIB gp de Zatse

[Received June 12, 1911: Read June 27, 1911.]

(Plate LXI.*)

The present paper is a pendant to the one preceding it, and

gives some account of three species of Chromodoris from the Red

Sea which, though not new, are worth notice as being either

varieties of known species or forms hitherto imperfectly described.

The matter which it contains is mainly due to Mr. Crossiand, and

I have contributed merely the identification of the species and

a few notes on their internal anatomy. ‘The first species, Chr.

reticulata, is the one on which Mr. Crossland made his interesting

observations regarding the warning coloration of the genus. My

own experience does not entirely support his statement that

Chromodorids do not hide under stones but show themselves in

the open. I have often found them (as well as Zrevelyana crocea)

under stones on tropieal beaches, where their vivid colours

harmonize wonderfully with the ascidians and sponges found in

the same lecality, so that the nudibranch which when isolated 1s

conspicuous, is almost invisible at home. But, as Mr. Crossland

points out, many species are known to arrive on the shore in

considerable bands for the spawning season, and perhaps all do so.

At this critical period, at all events, warning coloration must be

useful to them.

The following notes on three Red Sea species indicate that

there is considerable variation not only in colour but in the details

of the buccal parts. In what appears to be the same species, the

denticulation of the teeth and the shape of the elements in the

labial armature may vary, and thickenings of the rhachis,

amounting to rudimentary central teeth, may be present or

absent.

CHROMODORIS RETICULATA Pease, var. (PI. LXI. figs. 1-3.)

See Bergh, ‘‘ Neue Nacktschnecken ete.” No. iv., Jour. Mus.

Godeffroy, Heft xiv. p. 9 ff.; and Eliot, P.Z. 8. 1904, pp. 386~7.

My. Crossland’s notes on the living animal are as follows :—

“This species is of the soft flat kind. Rhinophores with rather

high cups. Gulls 9, hinder ones short, set in a simple circle open

iivsthiadl. Snbranchedl simply pinnate, motionless. Foot projects

slightly behind the mantle. Mantle-edge slightly or distinctly

wavy, according to the exteusion of the animal.

“ Colour. Greater part of the back a fine network of chocolate-

brown on a grey-white ground; laterally both network and

* Vor explanation of the Plate see p. 1072.

soe

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E.Wilson, Cambridge i

CisURVOIMUOIDKORUD SW IRCOUME AU Ise, TEU ID) PSylavAN,

CHROMODORIDS FROM THE RED SEA. 1069

ground- colour become yellow, forming an ill-defined yellow band

which passes into white. Round the mantle-edge are two deep

coloured bands, the outer orange-yellow, the inner deep violet.

There is a faint line of purple round the extreme edge of the

orange-yellow border. Foot and whole under side pure white

except that colours of mantle-edge are repeated on the under side.

The inner border of violet on upper surface is thickened in places,

and these points are those where the bending of the mantle-edge

is marked when the animal is half contracted.

“ Rhinophores brown with clear white lines along the per-

foliations ; gills grey with white limes on the edge of the folia

and specks elsewhere.

‘‘One specimen measured 5:0 em.x1°6 cm. when fully ex-

tended.”

Four specimens which have assumed a uniform dull plum-

colour, no markings being visible. In most other respects they

agree with Mr. Crossland’s description of the living animal, but

rlheTeS he gives the gills as only 9, I find that they vary from

12 to 18. It often happens that in Chromodorids with many

gills, the smaller plumes are not protruded from the pocket and

thus escape notice.

The labial armature consists of bent rods, but they are not bifid

as in other specimens which I have examined. In twospecimens

the formula of the radula was 63 x 50.0.50 and 65 x 68.0.68 in

the widest rows. The rhachis, as in the specimens described by

me from East Africa, bears triangular thickenings, in which the

base is not clearly defined. ‘The first laterals are as usual in the

genus, low, flattened, and bearing 4 denticles on either side. The

remaining laterals are rather stout, hamate, and bear 6—8 distinct

denticles. The-outermost laterals are lower and bear two or

three denticles on the apex only.

These specimens are referable to Chr. reticulata, but seem to be

a distinct variety characterized by (1) the shape of the rods in the

labial armature, which are not bifid at the tips; (2) the presence

of thickenings on the rhachis of the radula; (3) aviolet border to

the mantle. As will be seen from my previous descriptions of the

species, the coloration is variable.

CHROMODORIS TINCTCRIA Riippell & Leuckart. (Pl). LXI.

figs. 4 & 5.)

Rippell & Leuckart, Neue Wirbellose Thiere des Rothen

Meeres, p. 32.

A single specimen described by Mr. Crossland from the life as

follows :—

“ Chromodorid in shallow water on Zostera growing in sand.

“ Length $85 mm. Breadth of mantle 46 mm.

‘“‘ Body and foot are narrow, mantle more than ordinarily wide,

margin thrown into folds and very mobile.

“Whole animal soft and smooth-skinned, but there are soft

warts on the back, about § inch high and broad. Gills in a row

1070 SIR CHARLES ELIOT ON

shaped GO ona raised base, 18 in number, generally simple but

some are forked; one quite arborescent, bipinnate. Pinnules

rudimentary. The gills are completely retractile and gill-pocket

can close over them. Rhinophores fully extended; numerous

fine white lines om deep crimson ground, otherwise perfoliations

hard to see; completely retractile. Head distinct, with pro-

minent tentacles.

‘General colour greyish white, but this is plentifully sprinkled

with clear white opaque marks, so that general appearance 1s

white. Over the body this is covered wilh 4 a delicate network of

crimson, the mantle being sprinkled with clear spots of the same

colour. Edge of mantle bordered with a thin clear line of bright

orange- -yellow. A broad crimson line runs up each angle of each

gill rhachis.”

Mr. Crossland adds in a letter: ‘“‘ The margin is much more

ample than in other species known to me. I mars never seen one

before in which the undulations are so deep or keep in such

constant motion.”

Ritppell and Leuckart’s diagnosis is “Colore lacteo, pallii

margine sulfureo-limbato : dorso venis punctisque sanguineis

notato: branchiis 19 circiter, pinmatis.” They also say that the

branchie are ‘“ pyramiden-formig,” which corr esponds to Mr.

Crossland’s observation that the pinnules are rudimentary. The

coloration is sufficiently distinct to make the identity of this

animal with Riippell and Leuckart’s Doris tinctoria certain.

The preserved specimen is somewhat distorted but the breadth

of the mantle margin is still noticeable, and the shape is not that

of an ordinary Chromodoris. The internal characters appear to

be those usual in the genus. Though the outer surface of the

liver is purple wherever it is covered by the hermaphrodite gland,

yet the organ itself is of a deep black and leaves a str ong stain.

The labial armature consists of two dark purple plates.

The rhachis bears thickenings much as in Chr. reticulata, and

the radula is of a type common in the genus. The inner laterals

are low and flattened, the innermost bear 3-4 denticles on either

side; the rest are denticulate on the outer side only and the

number of denticles rises gradually from 5 (on the second and

third laterals) to 15 or more. The teeth at the same time become

tall and elegant in shape. There are about 60 laterals in all.

The marginals are denticulate on the tip only but are not

degraded to mere plates.

Curomoports tNorinata (?) Bergh. (Plate LAI. figs. 6 & 7.)

Bergh, Siboga, pp. 157-159.

A single specimen described by My. Crossland from the life

as follows :—

‘¢? CHROMODORIS sp),

Shallow water say 3 feet deep, sand and “sea grass,” in

Dongonab Harbour (March 1911),

CHROMODORIDS FROM THE RED SEA. Og

“ Full length, from edge of mantle in front to tip of foot behind

82 mm., breadth across mantle 40 mm.; these being measured

when the mollusc is crawling. Turned over on its back, the body

is only 20 mm. wide, mantle remaining much as before ; its margin

is not only thus ample, but also mobile. It has no permanent

undulations.

“Right gills, simply pinnate and unbranched, the posterior two

short, the rest long even in proportion to the body. They are not

held stiffly as in so many Chromodorids but wave about in the

water with its motion. (Iam not sure whether there is any active

motion.) Rhinophores straight and sharply pointed ; distal ones

two-thirds perfoliate. They and gills completely retractile.

“ Colour. Back appears ‘of a deep red-brown with numerous

orange spots, small yet conspicuous to the naked eye. As the

orange shows itself elsewhere, we probably really have an orange

ground covered with a thick netw ork of purple (as seen under a

Jens) with small round meshes. This network ends 3 or 4 mm.

from mantle-edge, the inner half of the remainder is therefore a

deep orange. he outer half is violet, light proximally but almost

black in a narrow band distally. There is a very fine line of

primrose-yellow along the extreme edge of the mantle. The

under side of the animal is pure sihite. only the violet ef the

mantle-edge reappears on the other side.

‘« Besides the meshes of the network the back is marked by a series

of clear cut spaces each defined by a darkened line of purple.

These spaces, 2 mm, or so in diameter, may be oval, kidney-shaped,

or circular. Those which are near the orange band of the mantle

are a clear yellowish white, but the rest are clouded with orange

centrally and some have an ill-defined purple spot in the centre.

* Gills and anal papilla are orange on their inner surfaces (those

facing the centre of the cup they form), the rest white with a

little orange.

‘“* Perfoliate parts of rhinophores brown-red passing into purple

at tips. A yellow-white line is conspicuous along the anterior

side, running vertically, and very fine lines of the same run

horizontally on the posterior halves of the perfoliations.”

Mr. Crossland adds that the colour pattern was so complicated

and varied so much in different lights, that he abandoned the effort

of painting it and only made a black and white sketch.

In a letter he alludes to “the clear bilobed area behind the

rhinophores,” beneath which the eyes can just be seen.

The preserved specimen has become of a uniform plum colour

and shows no markings, but otherwise is as described by Mr.

Crossland. The branchie and rhinophores are both very oe

and both completely protruded. The branchie are only 8, but a

the base of the hindmost is a small tubercle which is ee

the beginning of a growing plume.

As in the last species the liver is of a very deep black, only

partly concealed by the hermaphrodite gland, The labial arm-

ature forms a complete circle, olive in colour, and consisting of

1072 - ON CHROMODORIDS FROM THE RED SEA.

bent rods, not strictly bifid but sometimes bearing asmall tubercle

near the tip. The formula of the radula is 70 x 80.0.80 as a

maximum, but it is considerably narrower in the anterior part,

which is also markedly darker in colour. There are thickenings

on the rhachis as in Chr. reticulata but even more distinct. The

first laterals are of the usual flattish shape and bear about

4 denticles on either side. The other laterals are rather tall, and

erect. They bear 8-12 very faint denticles, visible only under a

high power. The outermost laterals are irregularly shaped plates

with a few denticles at the top.

In most respects this animal closely resembles Bergh’s Chr.

inopinata. ‘The remarkable coloration is almost identical (unless

the light area behind the rhinophores proves to bea characteristic

.feature found in other specimens), but there are two differences.

In Bergh’s specimen (1) the branchiz consisted of five tufts each

subdivided into 3, 5 or 8 plumes, (2) the teeth of the radula bore

—7 distinct denticles. The first difference is hardly of specific

value by itself. Typically, the gills of Chromodoris are simply

pinnate but in some species they show a tendency to subdivide.

Still individuals belonging to such species have often normal

pianate gills. The difference in the teeth is more important. If

the denticles as drawn by Bergh are not exaggerated, we must

either recognize two species (the present animal being charac-

terized by its teeth, sumtplas gills, and perhaps some persistent

differences in wolleention) or else admit that considerable variation

in the vadula are possible. I am inclined to adopt the latter

alternative. It will be observed that though the appearance of

the teeth in the two specimens is different, this difference depends

not on an alteration of shape but in the greater or less develop-

ment of denticles. Thickenings on the rhachis seem to be

indifferently present or absent in several species.

EXPLANATION OF PLATE LXI.

Fig. 1. Chromodoris reticulata from a living specimen.

. Chromodoris reticulata. Dorsal view of the mantle and part of body in the

living animal; enlarged.

Chromodoris reticulata. (a) One of the lateral teeth from the middle of a

half row; (6) thickening on the rhachis.

. Chromodoris tinctoria from a living specimen.

Chromodoris tinctoria. Teeth: (a) third lateral, (6) lateral from the middle

of a half row, (c) outer lateral, (d¢) thickening on the rhachis.

Chromodoris inopinata (2) from a living specimen.

Chromodoris inopinata, a lateral tooth.

toe

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Zo OMe elie

B.. Broom, del.

FOSSIL REPTILES FROM SOUTH AFRICA.

jeyAS USMY, Je abesQnul.

R.. Broom, del,

POSSI BEPTILE SS’ FROM SOUTH AMRIC:®

ON NEW PERMIAN REPTILES FROM SOUTH AFRICA. 1073

51. On some New South African Permian Reptiles *.

By R. Broom, D.Se., C.M.Z.8.

[Received May 23, 1911: Read June 27, 1911. |

(Plates LXII. & LXIIL+)

Of the following new fossil reptiles a number have been

discovered by the Rey. J. H. Whaits of Beaufort West, a most

enthusiastic and successful collector. The others have been found

by myself. Though none of the species described represent any

strikingly new types, the series forms an important addition to our

knowledge of the Permian Fauna.

Suborder DINOCEPHALTIA.

MoscicPs CAPENSIS, gen. et sp.n. (PI. LXIT. fig. 1.)

This important new genus was discovered by Mr. Whaits and

myself on the farm Spitzkop in the Moordenaar’s Karroo.

Remains of a number of skeletons were discovered scattered over

and imbedded in an alluvial deposit about a rood in extent.

Though the remains were for the most part fragmentary and

weathered, and so completely mixed up that it is practically im-

possible to pick out the bones belonging to any one individual,

this matters the less seeing that all the skeletons appear to belong

to one species. There is considerable difference in size of the

bones, but I think there is reason to believe that this may be

accounted for by assuming that the small herd which perished

together comprised males, females, and inimature animals. As

portions of thirteen thigh ‘bones have been discovered, there must

have been at least seven individuals.

Owing to the scattered condition of the remains it is impossible

to reconstruct the manus aud pes and to give the exact number

of the pre-sacrai and caudal vertebrze, but with these exceptions

every detail of the skeleton is known. I hope shortiy to give a

full description of the remains in a monograph on the Dinocephalia.

In the present paper I shali merely give a preliminary desciiption

of the skull.

The skull, which I figure and which may be regarded as the

type, is, I believe, that of a young female. The left side is much

weathered and the whole skull considerably crushed, especially in

the occipital region, but otherwise the skull may be regarded as

nearly perfect. The greatest length of the skull is $45 mm., and

from the occipital condyle to the front of the snout 290 mm.

This new animal resembles Delphinognathus conocephalus Seeley

so closely, that it was only after very deliberate consideration

that I decided to place it in a new genus. That it is specifically

* On p. 1079 Dr. Broom proposes two new generic names, viz., Arcfosuchus and

Arctognathus.—ED1ror.

+ For explanation of the Plates see p. 1082.

1074 DR. R. BROOM ON NEW PERMIAN

distinet is beyond question, and the marked difference in the

temporal region seems to me of sufficient importance to Justify the

formation of a new genus for its reception,

The nasal region makes with the line of the teeth a sharp angle

of about 45°, and the snout 1s moderately broad. The pre-

maxillaries are small and each carried three teeth.

The maxillary is moderately largeand flat and carried 12 teeth,

of which the first four are large and may be regarded as canines.

The septo-maxillary is a small bone whieh ‘errs the lower wall of

the nostril, and passes backwards a short distance, separating the

nasal from the maxilla,

The nasals are fairly large rounded bones which are separated

from each other at the lower part by the internasal processes of

the premaxillaries.

The orbit is large and round, protected above by a marked

thickening of the Pupraorbital border and behind by a strong

postorbital arch. The bones surrounding the orbit appear to he

very similar to those of Delphinoc grits and Tapinocephalus,

though the limits of the prefrontal have not been clearly made

out. The lachvymal foramen is large and opens on the face.

The jugal forms the lower border of ‘the orbit. It isa flat and

relatively slender bone which passes back to meet the squamosal

and the quadrato-jngal.

The quadrato-jugal isasmall bone which rests on the descending

process of the squamosal and on the quadrate. It does not as in

Delphinognathus unite with the jugal to enclose a foramen.

The quadrate is very similar to that of the Pelycosaurs. In

the type it is considerably crushed and displaced.

The squamosal is large. It hasa long slender ascending process

which forms part of the posterior wall of the temporal fossa and

meets the parietal. A descending process supports the quadrate,

and an anterior meets the jugal. ‘The upper part of the squamosal

vests on a flat bone which forms part of the oeciput, and which

I regard as the opisthotic from a comparison with the bone in

Marsupials and Cynodonts.

The frontals are large and broad and enormously thickened.

The parietals are relatively small, but like the frontals extremely

thick. They enclose, as in Delphinognathus and Tapinocephalus,

a large pineal foramen. The edges of the foramen are elevated,

but the pineal region stands out less prominently than in

Delphinognathus.

The occiput is moderately flat, and the condyle large and

rounded. The upper side of the eondyle has a deep “hellow

evoove for the medulla, and in the middle line immediately below

the medullary groove is a small deep pit which passes forward

into the basioeeipital for the notochord.

The palate is imperfectly known, but appears te be fairly

similar to that of 7'apinocephalus.

The lower jaw is like that of Delphinognathus but more fully

REPTILES FROM SOUTH AFRICA. 1075

known. The dentary forms about half of the jaw, but owing to

a slender backward process which rests on the surangular it forms

about ¢ of the upper border.

The ‘splenial is slender. The angular forms the larger part of

the outer side of the posterior part of the jaw, and the slender

surangular the upper third. The articular, which has two con-

cavities for the quadrate, is a powerful bone which in front fits

in between the angular and surangular on the inner side of the

jaw.

The teeth are imperfectly known, but appear to be very similar

to those of Delphinognathus and HKecasaurus. This type closely

resembles the tooth figured by Twelvetrees and Seeley as the

tooth of Deuterosawrus, and if these authors are right in regarding

that tooth as belonging to Veuterosaurus, there can be no doubt

that Deuterosaurus is a Dinocephalian closely allied to South

African forms.

Suborder ANOMODONTIA.

DIALURODON WHAITSI, gen. et sp.n. (Pl. LXITI. figs. 6 & 7.)

This beautiful little Endothiodont skull was discovered by the

Rev. J. H. Whaits on the Beaufort West Commonage. It

is fairly complete, but the matrix is so hard that no development

has beenattempted. Fortunately, the skull was discovered broken

into quite a number of scraps, and, but for this, it would have

been regarded as belonging to a small species of Dicynodon.

The lower jaw was, however, broken across obliquely and the

fracture revealed the presence of a series of molars. A second

very badly weathered specimen appears to me to belong to the

same species.

The total length of the skull is 83 mm. From the beak to the

front of the orbit is only 22 mm., and to the back of the orbit

41 mm.,so that the orbit is entirely in the front half of the skull.

The ereatest width of the skull at the back part is about 50 mm.

The parietal region measures 19 mm. across its narrowest part, and

the frontal region is only 13 mm. across. Owing to the forward

position of the orbits the facial part is relatively small and also

narrow. The tusks are typically Dicynodont, but owing to the

premaxillaries beimg very small, placed more forward “than in

Dicynodon.

The molars are arranged in a row like those of Hndothiodon

uniseries Owen, but a second replacing set is seen developing on

the inner side of the functional teeth. There are probably about

8 teeth in use in each jaw. ‘The crowns are not displayed in any

of the teeth of the type specimen, but in the second specimen a

large part of one crown is seen. It is long and slender and has

coarse serrations on the anterior and posterior borders. In this

the teeth agree with those of Hndothiodon, and differ from those

of Pristerodon and Opisthoctenodon.

1076 DR. R. BROOM ON NEW PERMIAN

TAOGNATHUS MEGALODON, gen. et sp.n. (Pl. UXIT. figs. 2-4.)

This new genus and species is founded on an imperfect and

much crushed snout found by myself on the farm Kuilspoort,

Beaufort West district. Though little more than the orbital

region 1S preserved, this specimen is manifestly very unlike any

type previously known.

The frontal region is flat and moderate'y broad, the interorbital

measurement being 16 mm. ‘The orbit measures about 20 mm.

in length.

The greater part of the dentaries is preserved but the anterior

part of the beak is missing. ‘The jaw differs from that of all

previously known Anomodonts in the much longer portion covered

by horn, and also in that the anterior part of the lower jaw

instead of being narrower than deep, is here about twice as broad

as deep.

The teeth are remarkable in that while there is a tusk it

was probably relatively short, and in the lower Jaw there are two

large teeth each about half the size of the tusk. Not im-

probably there were two molars in the maxillary and two in the

mandible.

The nearest ally of Taognathus at present known is Prodicynodon,

but the affinity is not at all close.

OUDENODON BOLORHINUS, sp.n. (PI. LXIIT. fig. 10.)

This new species of Oudenodon was discovered by myself at

Kuilspoort, Beaufort West district. Unfortunately, the type

consists of only the preorbital portion of the skull, and this is

somewhat crushed and weathered.

The most striking characteristics of the species are (1) the

extreme shortness of the snout which brings the front of the

palate nearly under the orbit, and (2) the thickening of the nasal

bones to form a rounded boss which overhangs the nostrils.

The orbit is large and measures about 35 mm. in diameter.

The borders are thickened and rounded.

The maxillary bone is short but powerful; the caniniform

process being very massive. The upper part of the bone forms

part of the thickened lower margin of the orbit and nearly

sepirates the jugal from the lachrymal.

The premaxillary bone is also short and strong, and was

probably as represented in the restoration.

The nasals are short but greatly thickened, forming a large

median boss the sides of which overhang the nostrils.

The frontal bones are short and comparatively narrow, the

interorbital region measuring about 36 mm.

The only species of Oudenodon which seems to come near the

present one is Oudenodon strigiceps Owen, but this latter differs,

apart from differences that may be due to crushing, in the much

ereater size of the nostril and in the much more backward position

of the caniniform process.

REPTILES FROM SOUTH AFRICA, 1077

Suborder THEROCGEPHALIA.

AALUROSAURUS WHAITSI, sp. n. (Pl. LXITI. fig. 8.)

This new species of lurosaurus was obtained by Mr. Whaits

at Beaufort West. lurosaurus felinus also occurs at Beaufort

West on the same horizon, but the difference between the species

is considerable apart from size.

The specimen consists of the greater part of the left dentary,

much of each maxilla, the left premaxilla, and numerous other

fragments of the skull, besides a large number of. fragments of

the postcranial skeleton. The skeleton is so imperfect that no long

bone is entire, and most are represented by articular ends.

A few imperfect vertebrae are present, and a number of dis-

articulated bones of the manus and pes.

The symphysial part of the jaw makes a less obtuse angle with

the lower border of the ramus than in Wlurosaurus felinus.

There are four lower incisors and a single large canine. In the

type only the deep part of the root of the first incisor is left, so

that an accurate measurement of the space oceupied by these teeth

is impossible. The last is situated very close to the canine. All

the incisors are subequal and rounded. The canine measures

10 mm. x 6°5 mm. at the base; the height is not shown. The

molars are small and degenerate. The exact number is uncertain ;

three remain in the jaw, but apparently two have been shed and

replaced by bony tissue. Probably the young animal had 5 molars

(possibly 6). There is a large diastema of 15 mm. between the

canine and what is probably the occupied position of the Ist

molar, and the five molars havea space of 13mm. From the front

of the jaw to the back of the last molar is a distance of 48 mm.

The upper incisors are badly preserved, but the roots are

preserved and occupy a space of 21 mm. ‘The upper canine is

large and but shghtly curved; its anterior border is smooth and

rounded ; the posterior border is serrated. The antero-posterior

measurement at the base is 11 mm.

It seems not improbable that specimen R855 a in the British

Museum may belong to this species.

AKLUROSAURUS TENUIROSTRIS, sp.n. (Pl. LXITTI. fig. 9.)

This species is founded on a snout collected by myself at

Kuilspoort. It resembles lurosawrus whaitsc very closely, but

differs in that, though it is probably a rather larger form, the

snout 1s more slender and the mandible feebler.

The five upper incisors measure 24 mm.as compared with 21 mm.

in 4, whaitsi, and the whole measurement from 7’ tom’ is 65 mm.

in 4. tenwirostris as compared with about 52 mm. in 4. whaitsi.

Some of the lower incisors are well preserved. The first is a

rounded pointed tooth with a feeble serrated ridge on its outer

and posterior side. On the third incisor there is only a very

slightly marked ridge, and there are no distinct serrations, though

it is possible that they have been worn off by friction against

Proc. Zoon, Soc,—1911, No. LX XIII. Ce

1078 DR. R. BROOM ON NEW PERMIAN

the upper teeth. The molars are better developed than in

A. whaitsi.

The arrangement of the bones round the nostril is similar to

that in Scylacosaurus and Aloposaurus.

IcTIDOGNATHUS PARVIDENS, gen. et sp.n. (Pl. LXIT. fig. 5, &

IPL, TUMILIO US stor TT)

This new genus is founded on a small snout found by me at

Kuilspoort. It is, with the exception of Scaloposaurus constrictus,

the smallest known Therocephalian.

From the snout to the orbit measures 46 mm., and the whole

skull was probably not more than 90 mm. in length.

Owing to the weathering of the fossil, which was found in a

sloot and was thus considerably water-worn, the front of the

snout is badly preserved and the teeth are very imperfect. There

appear to be six incisors, all of small size. There is a fair-sized

canine with a very minute Ist canine in front of it. The molars are

numerous, Hight are preserved and two are probably lost, possibly

three. The dental formula would thus appear to be 7. Se. 2 Gdn =

The dental formula of Scaloposawrus was recently given by me as

5 @ 3 9 A suas ;

a.-,¢.-, m.~, the reason for believing that there are three canines

being that the maxillary series begins with oneminute tooth followed

by two larger than any of the succeeding molars. If we regard the

third tooth as a molar, the dental formula becomes the same as in

Lctidognathus. But though the dental formule may be the same,

there is no question but that the species must be placed in separate

genera, the large canine in the present specimen suflicing to remove

it from Scaloposaurus. ‘The canine measures antero-posteriorly

3mm.

ERIPHOSTOMA MICRODON, gen. et sp. n. (PI. LXIII. fig. 12.)

This small imperfect skull was found by Mr. Whaits at

Fraserburg Road. It is in two portions—a rather badly weathered

snout, and an equally badly weathered occipital portion. Though

the contact is missing, the two fragments can be united with fair

accuracy, and the whole skull as restored measures 110 mm. in

length. The snout is very flat and deep, measuring in the

canine region 27 mm. in greatest width, though the snout with

the lower jaw here measures 55 mm. in depth.

The incisors are long, narrow pointed teeth which are situated

well to the front, and are apparently only 4in number. Following

the last incisor is a long diastema of 13 mm. corresponding to the

position of the large lower canine. The canine measures 4°5 mm.

in antero-posterior length. The number of molars is uncertain.

Two only are preserved, and these are long, pointed, slender teeth.

The number of molars must, however, have been few.

The only genera to which Hriphostoma is nearly related are

Ictidosaurus and Lycosaurus. Ictidosaurus angusticeps, the only

known species, was described by me in 1903 from a specimen in the

South-African Museum. JLycosaurus was founded by Owen on

REPTILES FROM SOUTH AF'RICA. 1079

a badly weathered skull in the British Museum, which was made

the type of LZ. pardalis. Unfortunately, the genus Lycosaurus is

at present in considerable confusion owing partly to the bad

condition of the type, and partly to the fact that two other species,

which are not nearly related have been added to the genus.

In Lycosaurus pardalis, Owen determined the dental formula

BSS de ee : sins

to be t.3, ¢.;,™.z. Lydekker not only believes Z. tigrinus Ow.to be

correctly referred to Lycosaurus, but thinks itisreally indistinguish-

able from Z. pardalis. In my opinion the two do not even belong

to the same family. The dental formula of Lycosaurus pardalis

0 Pa 4 9 ° :

I determine as 7.-, c.-, m.. The minute canine in front of the

Jarge one is a character very frequently met with in the early

Therocephalians of the Pareiasaurus zone. The deep square

symphysis with the incisor teeth carried well to the front, and

the small size of the premaxillary portion are all early characters.

Owen gives the locality of the type as “‘Sneewberg mountain-

range,” but this is evidently a mistake, and Lydekker in his

Catalogue merely states that the specimen is from the ‘‘ Karoo

System of the Cape Colony.” It. is highly probable that the

specimen came from the Gouph.

Lycosaurus tigrinus is a much later type of Therocephalian.

Its dental formula is i, O.7, mo As it belongs to a different

genus, I would suggest the new generic name dArctosuchus to

contain A. tigrinus Owen.

Lycosaurus curvimola belongs to a still higher type of Thero-

cephalian, which is not nearly allied either to Lycosawrus or to

He ee 5

Ailurosaurus. Its dental formula is 7.-, ¢.-, m.-. I would

suggest for it the new generic name Arctognathus to contain

A. curvimola Owen.

Removing then these other genera and reserving Lycosaurus

for the type-species, we get a form resembling in many characters

Eriphostoma, but differing in being larger by a half, in having 5

incisors as against 4,and 2 canines instead of 1 as in Hriphostoma.

Lctidosaurus agrees with Lycosaurus as regards the incisor and

canine formula, but differs in having a large number of molars

and a number of other cranial characters. ‘The dental formule

of the three genera are as follows :—

° . )

Wichidosauiuss ee ae oe ee OS

Lycosaurus : pe ia asion 7 SCI Ba

? : 2

Pircpostonigls Vik CSEES ay eG a pean eas

The Geological Horizons of the Beaufort West Specimens.

Some years ago I endeavoured to subdivide the Beaufort series

into paleontological zones. At that time it was only possible to

do so on very broad lines. The area is so extensive, and except

on the upper part of the series there are no lithological characters

(ae

1089 DR. R. BROOM ON NEW PERMIAN

to correspond to the paleontological. Further, as wide areas of

the Karroo are covered by wind-blown dust which forms what

may be regarded as a sort of loess deposit, and the exposed

portions, except in the case of escarpments, are usually isolated, it

becomes a matter of extreme difficulty to connect up the various

beds. The shales for a thickness of 3000 or 4000 feet present no

distinguishing characters, and at present we can do little more

than collect fossil specimens and note the localities.

I subdivided the Beaufort into six zones which may be grouped

as follows :—

(6. Cynognathus Beds.

| 5. Procolophon Beds.

Middle Beaufort 4. Lystrosaurus Beds.

3. Cistecephalus Beds.

Lower Beaufort < 2. Hndothiodon Beds.

1. Pareiasaurus Beds.

Upper Beaufort

The Lystroswurus zone probably corresponds to the Lower

Triassic ; the Cynognathus zone to the Upper Triassic. The

Pareiasaurus beds are probably Middle Permian ; the Hndothiodon

and Cistecephalus beds are probably Upper Permian. <All work

done since 1905 has gone to confirm the correctness of the con-

clusions then arrived at, but it has shown that we will some day

be able to subdivide the zones into a large number of subsidiary

ZONES.

The Pareiasaurus zone is the oldest one where fossils are

numerous. Pareiasawrus is its most characteristic form, but there

are a host of other known genera, mostly of 'Therocephalians.

Small Anomodonts—Dicynodon and Oudenodon—are also not

uncommon.

Above the Pareiasaurus zone we have deposits which measure

about 2000 feet, principally characterised by the abundance of

Dicynodon and Oudenodon. Of every six fossils obtained, five are

those of Dicynodon or Oudenodon. As, however, these genera

occur at all horizons of the Beaufort, it seems unwise to speak

of this as the Dicynodon zone. I have therefore called it the

Endothiodon zone, as Endothiodonts are met with throughout it

and are known at no other horizon. 'Therocephalians are not

common, but may be met with at any height. Curiously enough,

with the exception of Propappus and Sawrosternon, no genera are

known from the Hndothiodon zone which do not belong to the

Anomodontia or the Therocephalia.

Above the Hndothiodon zone is a zone probably not more than

1000 feet thick, which is characterised by the presence of

Cistecephalus, and the higher types of Therocephalians. ‘The zone

is at present the least known of any part of the Beaufort, mainly

because since Andrew Bain’s time no collector has had an oppor-

tunity of doing much with it.

At Beaufort West, one has an opportunity of studying to

advantage the Hndothiodon zone. The township is situated on

REPTILES FROM SOUTH AFRICA. 1081

an undulating plain which is geologically probably not more than

500 feet above the Pareiasaurus zone. The subzone is charac-

terised by the presence of Hndothiodon uniseries Ow.,—a form

that at present is not known from any other part of South Africa.

In the Beaufort West district also occurs Hndothiodon bathystoma

Ow., but whether the horizon of this is above or below that of

FE. wniseries is at present unknown.

Near Beaufort West is the escarpment of the Nieuweveld

which rises a few miles north of the township to a height of 3000

feet or more, and though it is extremely difficult collecting in the

steep slopes of shale, a number of forms of interest have been

collected at various horizons.

As Dicynodon and Oudenodon occur at all levels, it might be

thought well to subdivide the zones by the species of Dicynodon ;

but there is a serious difficulty. Dicynodon is the most trouble-

some genus we have to deal with. Specimens differ so greatly in

size and shape that one hardly knows what to do unless one does

as was practically done by Owen, make every specimen the type

of a distinct species. For many years to come the genus

Dicynodon must remain in utter confusion, and will be useless for

stratigraphical work. On the other hand, the Therocephalians

will be as reliable guides as Ammonites and Trilobites are in the

marine rocks of Europe.

In the following diagram are represented the horizons of the

Beaufort West types described in this paper, and of others whose

horizons are known. It must, however, be understood that the

heights in feet are only approximate.

| 2000 feet ..........55....... Ictidognathus parvidens Br.: Allurosaurus tenui-

rostris Br.; Oudenodon bolorhinus Br.; Cistecephalus

microrhinus Ow.; Dieynodon sp.

| 1500 feet ..................... Aloposaurus gracilis Br.

| Dicynodon sp.

| 1000 feet ..................... DPaognathus megalodon Br.; Oudenodon sp.;

| Dicynodon sp.

500 feet ...............-..... Dicynodon sp.

Dicynodon sp.

Beaufort West horizon ... Hndothiodon uniseries Ow.; Dielurodon whaitsi Br. ;

Alurosaurus whaitsi Br.; Scymnosaurus sp. ;

Dicynodon sp.; Oudenodon sp. lurosaurus

felinus Ow.

It may be well to regard the Hndothiodon zone as reaching

1600 feet above Beaufort West, and then passing into the

Cistecephalus zone.

1082 MR. E. G. BOULENGER ON A

EXPLANATION OF THE PLATES.

Puate LXII.

Fig. 1. Side view of skull of Moschops capensis Broom. 2nat.size. Ang. Angular;

Art. Articular; Ju. Jugal; O.O. Opisthotic; Qu. Quadrate; Q.J. Quadrato-

jugal; S.Ang. Surangular: Sma. Septomaxillary ; Sq. Squamosal.

2. Front part of skull of Taognathus megalodon Broom. Nat. size.

3. Under view of lower jaw of Taognathus megalodon Broom. Nat. size. The

canine of the right side is seen in oblique section.

4. Section across lower jaw of Taognathus megalodon Broom. Nat. size.

5. Upper view of snout of Letidognathus parvidens Broom, Nat. size.

Prate LXIII.

Fig..6. Side view of skull of Dielurodon whaitsi Broom. Nat. size.

7. Section across lower jaw of Dielurodon whaitsi Broom, at the crack

indicated in figure 6. Nat. size.

8. Upper view of mandible of _4lurosaurus whaitsi Broom. Nat. size.

9. Side view of snout of Alurosaurus tenuirostris Broom. 2 nat. size.

10. Side view of snout of Oudenodon bolorhinus Broom. 3 nat. size.

11. Side view of snout of Ietidognathus parvidens Broom. Nat. size.

12. Side view of snout of Hriphostoma microdon Broom. + nat. size.

52. On anew Tree-Frog from Trinidad, living in the Society’s

Gardens. By Hpwarp G. BouLEencer, Curator of

Reptiles to the Society *.

[Received September 20, 1911: Read October 24, 1911.]

(Plate LXIV.+)

In July last the Zoological Society received from Dr. Lewis

H. Gough an interesting collection of Batrachians and Reptiles

brought back by ‘him from Trinidad. Among these I found

examples of three frogs which had not been previously recorded

from Trinidad, viz.: Hyla venulosa Daud., Hyla rubra Daud., and

one which is evidently undescribed, and for which I propose the

name of Hyla goughi, after its discoverer. This little Hyla, one

of the smallest of the genus, was fortunately represented by

numerous specimens, which have enabled me to observe the wide

and rapid changes of colour which this species undergoes, and of

which an idea can be gained from the annexed coloured plate

made by Mr. J. Green at the Gardens under my direction.

HYLa Goucut, sp.n. (Pl. LXIV.)

Tongue circular, slightly nicked and slightly free behind.

Vomerine teeth in two rounded groups between the choane.

Head slightly broader than long; snout rounded, a little shorter

than the eye, which is large and very prominent. Canthus

rostralis feebly marked. Loreal region very slightly concave.

* Communicated by the Secretary.

+ For explanation of the Vlate see p. 1083.

eZ) LONE wileplexgl Va

ad.Green del.et Chromo lith.

GOUGHS TREE FROG.

(HYLA GOUGHI.)

NEW TREE-FPROG FROM TRINIDAD. 1083

Interorbital region a little broader than the upper eyelid.

Tympanum fairly distinct, 4 diameter of eye, Fingers rather

short, with small discs, outer 4 webbed. Subarticular tubercles

feeble. No projecting rudiment of pollex. Toes 4 webbed.

The hind limb being carried forward along the body, ‘the tibio-

tarsal articulation reaches between the eye and the end of the

snout. Tibia half the length of the head and body. Upper parts

smooth, with minute granular warts on the head and the anterior

part of the back. Male with a feebly developed subgular vocal

sac, forming loose folds.

Coloration.—Vhe vapid changes in colour which this frog

undergoes are probably unparalleled in any other Batrachian.

The same individual may vary dorsally from dark brown, reddish

brown, various shades of yellow, to a very pale greyish white.

When startled the majority became of a bright lemon-yellow.

Jn one specimen I observed the head, fore limbs, and anterior

part of the body to be dark brown, whilst the posterior part of

the body and hind limbs were greyish white. In another speci-

men the right half of the body was brown, the left half greyish

white. A brown or grey marking, often hourglass-shaped,

edged with darker or lighter, extending from between the eyes to

the anterior third of the back, followed by one or two transverse

bars, is frequently present, appearing and disappearing with

great rapidity. In one specimen the warking took the shape of

a eross-bar between the eyes and two parallel longitudinal bands

extending along each side of the entire length of the back.

Faint cross-bars on the hind limbs are occasionally present.

Lower parts white or yellow. Iris golden, much obscured by

brown pigment; a clear golden line borders the pupil, which

when fully contracted becomes perfectly linear.

The largest specimen measures 22 mm. from snout to vent.

This little Tree-Frog is extremely agile in its movements,

making leaps of quite six feet. In the daytime it usualiy kept

quiet, sticking to the leaves in the terrarium. On arrival the

males issued a sharp creaky note, but became perfectly mute

after a day or two at the Gardens,

This species appears to be more nearly related to Hyla strigilata

Spix, from Brazil, and Hyla misera Werner (Zoo), Anz. 1903,

p- 252), from Caracas, Venezuela, than to any others. It differs

from both in the shorter snout ; ‘from the former also in having

more fully webbed toes; from the latter in having a smaller

tympanum, and a shorter web between the fingers.

EXPLANATION OF PLATE LXIV.

Gough’s Tree-Frog (Hyla goughi).

1084 MR. C. E. HELLMAYR ON THE

03. A Contribution to the Ornithology of Western Colombia.

By C. EH. Hetimayr, Curator, Division cf Birds,

Zoological Museum, Munich *.

[Received April 26, 1911; Read June 13, 1911.]

Page

Tc Introduction esas ce ee ee ee ee LOS:

If. Account of Mr. M. G. Palmev’s localities.................. 1085

Iif. Account of the Species .. SREY Ae tee dn AM, een 1086

Wik Coneluvsion sy peice We ree mn hed er ae meee ea toy EATON)

I. INTRODUCTION.

The following account is principally based on a collection of

birds made by Mr. Mervyn G. Palmer, one of Mr. W. F. H.

Rosenberg’s field collectors, in the second half of 1908 and in

1909. Though numbering hardly 700 specimens, the set is of

considerable interest, containing a large percentage of rare species

as well as several novelties which I have already deseribed, partly

in the ‘ Bulletin of the British Ornithologists’ Club,’ partly in the

‘ Revue Frangaise d’Ornithologie.’

The district of Western Colombia in which Mr. Palmer has been

werking had not been visited by many naturalists before, and,

putting aside some seattered notices in scientific periodicals, few

accounts have been published about the birds of this little-known

region. Our first knowledge is due to the exertions of the cele-

brated French traveller, Ad. Delattre, who, prior to 1846, made

considerable collections in the vicinity of Buenaventura (Chocé

Bay), in the Western Cordillera at Juntas, Cali, ete., in the neigh-

bourhood of Popayan, notably on the Puracé, as well as at Pasto,

near the Heuadorian frontier. The new species of Humming-

Birds discovered on this trip were made known by Delattre and

Bourcier in the ‘ Revue Zoologique,’ vol. ix., 1846, pp. 305-312,

while a number of birds belonging to other families are discussed by

M. de Lafresnaye in the same journal, vol. x., 1847, pp. 67-79.

The next important contribution is Cassin’s Catalogue of the Birds

collected by Lieut. N. Michler’s expedition to the Rivers Truando

and Atrato, just south of Darien, published in the ‘ Proceedings of

the Academy of Natural Sciences of Philadelphia’ for the year

1860, pp. 132-144, 188-197. This was followed by Sclater and

Salvin’s account of the late T. K. Salmon’s extensive collections

from the State of Antioquia, in the P.Z.S. 1879, pp. 486-550,

wherein 468 species are dealt with. In 1894, Mr. W. F. H. Rosen-

berg visited the Rio Dagua, working chiefly at Juntas and Cali, in

the Western Cordillera. His birds went to the late Adolphe

Boucard, who published a list of the Humming-Birds in ‘The

* Communicated by Dr. P. L. Scuater, F.R.S., F.Z.S.

BIRLS OF WESTERN COLOMBIA. 1085

Humming Bird,’ vol. v., 1895, pp. 5-7, while the bulk of the col-

lection was never reported upon. Mr. Gustav Hopke, im 1896 and

1897, sent a fair series from the same district to Count Berlepsch,

who described several new species in the ‘ Ornithologische Monats-

berichte,’ vol. v., 1897, pp. 173-176, and in the ‘ Ornis’ xiv., Feb.

1907, pp. 347, 361, 365. Mr. Eugene André, in 1899, forwarded

a large collection of birds, from the environs of Buenaventura

and the western slope of the Andes above that town, to Comte

de Dalmas, of Paris. Unfortunately, the greater part of 1t was

subsequently destroyed by accident, and merely a list of the

Trochilide published by Messrs. Simon and de Dalmas in ‘ Ornis,’

x. 1901, pp. 216-224. Lastly, Mr. Outram Bangs, the well-

known ornithologist of Boston, who had obtained two of Palmer's

earlier consignments, gave short accounts of the more interesting

species in the ‘ Proceedings of the Biological Society of Washing-

ton,’ vol. xxi., 1908, pp. 157-162,* and xxiii., 1910, pp. 71-76,

describing several supposed novelties some of which are undoub-

tedly distinct, while others prove to be identical with species

previously named.

The material discussed in this communication is deposited in the

collection of the Zoological Museum at Munich with the exception

of the few species given in brackets.

II. Account oF Mr. PAatmer’s LocALirtizs.

The whole of Mr. Palmer’s collection in possession of the

Munich Museum was secured in the province of Chocé, though

in two different districts. The bulk is from the hot, tropical

valley of the Rio San Juan and its tributaries, the Tamana,

Sipi, Condoto, Calima, Cajon, and Garrapatas Rivers, while a

smaller number of specimens was collected on the Pacific slope of

the Western Cordillera near the head-waters of the San Juan, at

altitudes of from 2800 to 8000 feet.

IT am indebted to Mr. Rosenberg for information about the

nature and altitude of some of the localities which he had visited

himself in 1894, They may be separated into three divisions :—

(A) Rio Dagua District.

Sad Joaquim, Bahia del Chocé (= Buenaventura of Huropean

maps).

El Paillon, several hours’ journey up the Dagua (HK. André).

San José, 600 feet (visited by André and Hopke).

Los Mangos (=Juntas, Rio Dagua), cirea 1000 ft. Hot, tropical

country.

* The statement that the material came from N.W. Colombia, “just south of

Darien,” is a mistake. I am informed by Mr. Rosenberg that the San Antonio

referred to isa village on the road from Buenaventura to Cali, just over the pass

of the Western Cordillera. M1. Bangs appears to have identified it with the San

Antonio, on the Rio Sucio, a tributary of the Atrato, where, however, Mr. M. G.

Palmer never was.

1086 MR. C. FE. HELLMAYR ON THE

Jiménez, 1600 ft., a morning’s walk from Los Mangos, in a

deep ravine by the side of the pass between Los Mangos

and a place called Ventanas, on the road to San Antonio

and Cali. Forest country. Above Ventanas the country

_ becomes open. .

Naranjo (1900 ft.) and Plano de los Monos (2600 ft.). (Visited

by HK. Andre.)

Naranjito, 3900 ft.

Pavas, 4400 ft; San Luis, Bitaco Valley, 4400 ft.

La Maria, 4700 ft.

Palmar, near La Maria.

San Antonio, R. Cali, 5400 to 5800 ft. On road from Buena-

ventura to Cali, just over the Pass of the Western

Cordillera. Mr. Rosenberg passed through San Antonio

many times, but never collected there. The first lot that

went to Mr. Bangs was chiefly brought together at this

locality.

(B) Rio San Juan District (main stream and tributaries). Hot,

tropical country covered with dense forests.

R. San Juan: Noanama, alt. 100 ft.; Tad6, farther north,

230 ft.

R. Cajon, a small affluent.

R. Calima: Guineo (sea-level).

R. Sipi: Sipi, 150 ft.; Rio Garrapatas, 150 ft.

R. Condoto : Condoto, 150 ft.

R, Tamana: Noévita, 150 ft.; El Tigre, 320 ft.; Juntas, 400 ft.*

San Juan.

Pueblo Rico, San Juan slopes, 5200 ft.

Siaté, Rio Siaté, near Pueblo Rico, 5200 ft.

KR. Jamaraya, one of the headwaters of the R. San Juan: Loma

Hermosa, 4180 ft.; La Selva, 4600 ft.

Tatama Mountain. Mr. Palmer collected at various stations

(2794 to 8000 feet).

III. Account oF THE SPECIES.f

1. TURDUS TRISTIS DAGUA Berl.

[Merula tristis Swainson, Philos. Magaz. (n. ser.) i. p. 869

(1827.—Temascaltepec, Mexico). |

Lurdus dague Berlepsch, Ornith. Monatsber. v. p. 176 (1897.—

San José, Rio Dagua, 8.W. Colombia); Hartert, Nov. Zool. v.

1898, p. 478 (Cachavi, N.W. Ecuador).

No. 1957. g ad. San Joaquim, Bahia del Chocé, 1.viii.08.—

Wing 107; tail 84; bill 18 mm.

* Not to be confounded with Juntas on the R. Dagua.

+ [The absence of brackets round the name of an author to indicate that his

species has been transferred to another genus is not due to Mr. Hellmayr, but is in

accordance with the custom of the Zoological Society —Ep:ror. |

BIRDS OF WESTERN COLOMBIA. ' 1087

Nos. 2078, 2180. ¢ ¢ ad. Sipi, ll.ix., 12.x.08.—Wing 110,

HOS ss tale Ser 95m bull S anne

No. 2299. g ad. Noévita, Rio Tamana, 16.xi.08.— Wing 106;

tail 78; bill 18 mm.

No. 2370. 9 ad. Ndévita, Rio Tamana, 5.xi1i.08—Wing 105;

tail 76; bill 18 mm.

No. 2072. 2 imm. Rio Sipi, 10.1x.08.—Wing 107; tail 81;

bill 17 mm.

“Tris, feet and bill dark brown.”

In spite of the late Dr. Sharpe’s doubts there can be no

question that 7’. t. dague is a perfectly distinct form, though, to

my mind, its relations are more correctly expressed by a trinomial

appellation. Compared with a good series of 7’. tristis cnephosa

Bangs, from Chiriqui and Western Costa Rica (Miravelles), the

Colombian birds have shorter wings and tail, a much shorter,

slenderer and darker (blackish-brown) bill, and differ also in

coloration. The upper parts are deep reddish sepia-brown (instead

of greyish olive or olive-brown), the breast and sides dark

rufescent brown (instead of pale brownish grey), and the sides of

the head deep sepia-brown (instead of sooty black).

Tt. dague is peculiar to Western Colombia and N.W. Ecuador,

whence Miketta and Flemming sent numerous specimens to

Mr. Rosenberg. Hcuadorian skins are practically identical with

the topotypes forwarded by Mr. Palmer. This Thrush is an

iwhabitant of the humid lowlands and does not occur in the

mountainous parts of the interior.

2. HyYLocICHLA USTULATA SWAINSONII Cab.

[Turdus ustulatus Nuttall, Manual Orn. U.S. & Canada, Land

Birds, ed. 2, pp. vi, 830 (1840.—Columbia River). |

Turdus swainsoni Cabanis in Tschudi’s Fauna Peruan., Aves,

p- 188 (1844-46. — “ New-Jersey, im Monat October,” coll.

Cabanis).

No. 2302. Q ad. Novita, 17.xi.08. “Iris dark brown, feet

pink, maxilla black, mandible light brown, tip black.”

Identical with other specimens from Bogota, Ecuador, and

Venezuela (Merida).

3. MYADESTES RALLOIDES D’Orb.

Muscipeta ralloides D'Orbigny, Voyage dans |’Amér. mérid.,

Oiseaux, p. 322 (between 1838 & 1847.—Chulamani, Yungas of

Bolivia).

Myiadestes ralloides Sclater & Salvin, P. Z. 8. 1879, p. 492

(Retiro, Concordia, Medellin, 8. Elena).

Nos. 3760, 3771. $ Q ad. Siaté, 5200 ft.: Sept. 17, 1909.—

Wing 84, 82; tail 75, 72; bill 1] mm.

“Tris dark brown, feet light brown, maxilla black, mandible

light brown.”

The specimens agree perfectly with others from Bogota, while

1088 MR. C. E. HELLMAYR ON THE

skins from Cumbre de Valencia, Venezuela, have decidedly smaller

bills. Typical Bolivian birds are not available for comparison.

M. ralloides is probably divisible into several geographical races.

4. HELEODYTES ALBOBRUNNEUS HARTERTI Berl.

[Heleodytes albo-brunneus Lawrence, Ibis, iv. p. 10 (1862.—

“On the line of the Panama Railroad near the summit of the

Atlantic Slope ”.)|

Heleodytes harterti Bevlepsch, Ornis, xiv. p. 347 (Feb. 1907.—

San José, Rio Dagua, W. Colombia).

No. 2522. g ad. El Tigre, Rio Tamana, 320 ft., 9.11.09.— Wing

92: tail 88; bill 23 mm.

No. 2523. @ ad. Same locality and date.—Wing 83; tail 78 ;

bill 20 mm.

“ Tris reddish brown, feet grey, bill black.”

The specimens agree in coloration with the types kindly lent by

Count Berlepsch, and have the back, wings, and tail very much

darker, more brownish black, than typical albobrunneus from

Panama. The dusky markings of the under tail-coverts are

less regular, more spot-like, and the bill is black instead of

pale brown. The difference in size, however, proves to be not

constant, the female from Chocé beimg scarcely larger than

Panama examples *. It may be mentioned that the female has

the head all round pure white hke the male, while that of albo-

brunneus is said to have the pileum clouded or streaked with

pale brown.

H. a. harterti is as yet known only from the Chocé district in

Western Colombia.

5. LEUCOLEPIS PH ZOCEPHALUS PH#OCEPHALUS Scl.

Cyphorinus (sie) pheocephalus Sclater, P. Z. 8. 1860, p. 291

(end of 1860.—Esmeraldas, N.W. Ecuador); Sclater & Salvin,

P. Z.S. 1879, p. 492 (Remedios, Antioquia).

Cyphorhinus brunnescens Sharpe, Cat. B. Brit. Mus. vi. p. 293

(1881.—“ Cauca Valley,” sc. Remedios).

No. 2555. Q ad. Juntas, R. Tamanda, 24.11.09.—Wing 66;

tail 32; bill 20 mm.

“Tris brown, feet dark brown, bill black.”

This specimen, a perfectly adult bird, agrees exactly with topo-

typical examples from N.W. Ecuador (8. Javier), and others from

Chimbo. Perhaps it is a trifle darker chestnut on the back and

slightly deeper rufous underneath. The type of C. brunnescens

Sharpe, which I have examined in the British Museum, is merely

a young bird of C. pheocephalus. Kcuadorian specimens in cor-

responding plumage are exactly like it. Moreover, it 1s a well-

known fact that in the species of this genus the young birds have

the upper parts lighter, more olivaceous brown, and the throat,

* Two specimens measure; wing 80-82; tail 79; bill 19-195 mm.

BIRDS OF WESTERN COLOMBIA. 1089

foreneck, &c. of a much paler, orange-rufous. This variation is

well shown by our series of the nearly allied L. p. lawrencii

Lawr.* from La Vijagua, Eastern Costa Rica fT.

L. p. pheeocephalus is confined to the lowlands of W. Ecuador

and W. Colombia.

6. THRYOPHILUS NIGRICAPILLUS scHoTri Baird.

[Lhryothorus nigricapillus Sclater, P. Z. 8. 1860, p. 84 (1860.—

Nanegal, W. Ecuador). |

Thryophilus schottii Baird, Review Am. B. 1, p. 133 (1864.—

Truando R., Colombia).

Thryothorus nigricapillus (nec Sclater) Cassin, Proc. Acad. N.

Sei. Philad. 1860, p. 195 (River Truando).

Thryophilus nigricapillus (nec Sclater) Sclater & Salvin, P. Z.5

1879, p. 493 (Remedios, Antioquia).

No. 2585. gd ad. Condoto: 2.iv.09.—Wing 67; tail 51; bill

18 min.

Nos! 2181) 258i 2622) OO "ad! Sip] 2s 08)-* Condoto:

20.111., 22.iv.09.— Wing 63-65; tail 46-48 ; bill 17 mm.

Nos. 2618, 2621. 9 2 juv. Condoto: 21,22.iv.09.— Wing 633,

65; tail 46, 48; bill 15 mm.

“Tris brown, feet dark grey, bill black, mandible grey in adults,

yellow in young birds.”

These specimens differ from a good series of 7. n. nigricapillus

from Western Heuador in having the black cross-bands of the

lower parts much broader, more regular as well as more closely

set, especially along the middle Ag the belly. Moreover, chin

and. throat ere distinctly, though narrowly, banded with. blackish,

while they are uniform white in the Ecuadorian form. Even

young birds can easily be distinguished by these characters. In

the coloration of the lower parts T. n. schottit approaches ~

T. sennibadius Salv., of Chiriqui and Costa Rica, but has the top

of the head black like 7’. n. nigricapillus. Its range is evidently

restricted to Western Colombia, from the R. Tr uando south to

the San Juan district. -Like its southern representative, it

exclusively lives in the hot, low country.

. THRYOPHILUS LEUCOPOGON Salvad. & Festa.

Phnyophilus leucopogon Salvadori & Festa, Boll. Mus. Zool.

Torino, xiv. no. 357, p. 6 (1899.—Rio Peripa, W. er

Hellmayr, Journ. f. Orn. 1903, p. 534 (San Javier, N.W.

Ecuador).

No. 2440. g ad. Névita: 28.xii.08.—Wing 58; tail 38; bill

163 mm.

* Cyphorinus lawrencii Lawrence (ex Sclater, MS.), Ann. Lye. N. H. N.Y. viii. p.5

(May 1863,—based on C. cantans (nec Gmelin) Lawrence, 1. ¢. vil. 1861, p. 293.—

Panama Railroad).

+ The presence of dusky bars on the primary coverts upon which the late

Dr. Sharpe Jaid much stress is a purely individual character. Five birds from

W. Ecuador al/ have distinct blackish bars, as also the female trom Juntas.

1090 MR. CG. E. HELLMAYR ON THE

“Tris dark red, feet light grey, maxilla black, mandible grey.”

This specimen agrees perfectly with others from Western

Keuador. In addition to the one from San Javier in the Vienna

Museum mentioned by me, /. c., I have since examined four more

Specimens from N.W. Ecuador in the Tring Museum; one male

and two females also from S. Javier, and a male from Lita,

3000 ft. The wing measures 54-58, the tail 33-37 mm.

T’. leucopogon, though a very well-marked species, is nearest to

T. thoracicus Salv., of Costa Rica, which it closely resembles on

the upper parts, but differs mim the uniform dull ochreous brown

colour of the lower surface, with the exception of the chin and

upper throat which are white with slight blackish edgings, while

in 7’. thoracicus all the throat and breast feathers are white,

broadly margined with black laterally.

T. leucopogon is known only from the coast-belt of Western

Ecuador and Colombia.

8. HenicorRHINA INORNATA Hellm.

Henicorhina imornata Hellmayr, Journ. f. Ornith. l. p. 528

(1903.—Lita, N.W. Ecuador).

No. 2135. 2 ad. Sipi, 30.ix.08.—Wing 55; tail 2743; bill

16 mm.

‘‘ Tris dark brown, feet and bill black.”

This bird is practically identical with the series, including the

type, from N.W. Ecuador, in the Munich Museum. The upper

parts are of the same bright chestnut-rufous hue, the sides of the

breast deep smoky grey, the flanks dark rufous brown, and the

base of the lower mandible is clear yellowish white. H. inornata

is known only from Western Colombia and N.W. Kcuador, where

it inhabits the forests of the humid lowlands as well as the lower

slopes, up to 3000 feet (Lita).

[In the mountains bordering the Cauca Valley it is repre-

sented by another species which Mr. Bangs * has lately separated

as H. leucosticta eucharis, but which I cannot satisfactorily

distinguish from H. 1. prostheleuca, of Chiriqui and Eastern Costa

Rica. Having before me two fine specimens t, I can positively

state that it has nothing to do with H. lewcosticta of Guiana and

Kast Venezuela. Mr. Bangs was apparently misled by a black-

crowned male, but this character is of very little importance in

view of the fact that specimens of prosthelewca in worn plumage

often have the crown nearly uniform black, the brown tips to the

feathers having disappeared through abrasion. Moreover, the two

skins from Primavera have the feathers of the pileum broadly

tipped with umber-brown and the back ete. dull russet- brown,

exactly as in prostheleuca. While a larger series from Colombia

might ultimately reveal some slight differences, the evidence at

hand is not in favour of the southern form being separable. |

* Proce. Biol. Soc. Wash. xxiii. p. 74 (1910.—Pavas, W. Colombia, 4400 feet).

+ & & trom Primavera, Cauca R., 5100 feet, Raap coll., Tring Museum.

BIRDS OF WESTERN COLOMBIA. 1091

9. SEIURUS NOVEBORACENSIS NOVEBORACENSIS Gin.

Motacilla noveboracensis Gmelin, Syst. Nat. 1, i. p. 958 (1789

—ex Waubenton, Pl. Enl. 752. fig. 2: Louisiana).

Siurus noveboracensis Sclater & Salvin, P. Z. 8. 1879, p. 493

(Concordia, Medellin).

Nos. 2131, 2153, 2171. 3b @ ad., (sex not determined). Sipi:

TOR Box Ole,

“Tris dark brown, feet brown, maxilla black, mandible brown.”

These birds apparently belong to typical noveboracensis. Quite

similar specimens we have from Ecuador and Bogota collections.

A common winter visitor to Colombia.

10. Mwnioriuta varia Linn.

Motacilla varia Linneus, Syst. Nat. xii. 1, p. 333 (1766—ex

Brisson & Sloane: Jamaica and San Domingo).

Miiotilta varia Sclater & Salvin, P. Z. 8. 1879, p. 493 (Con-

cordia etc.); Berlepsch, Journ. f. Orn. 1884, p. 282 (Bucara-

manga).

Nos. 2807, 2855. § 2. Pueblo Rico: 27.x.; Loma Hermosa:

Daexa)o: :

“ Tris and feet dark brown, maxilla black, mandible white.”

A common winter visitor to Colombia.

11. DeNpDROICA CASTANEA Wilson.

Sylvia castanea Wilson, Amer. Orn. il. p. 97, pl. 14. f. 4 (1810.

—Pennsylvania).

Dendreca castanea Sclater & Salvin, P. Z.S. 1879, p. 494

(Remedios); Berlepsch, Journ. f. Orn, 1884, p. 282 (Bucara-

manga). —

Nos. 2317, 2331, 2381. § dg juv. Novita: 20,24.xi., 9.xii.08.

“Tris dark brown, feet grey, maxilla black, mandible grey.”

12. DENDROICA ZSYIVA ZSTIVA Gin.

Motacilla cestiva Gmelin, Syst. Nat. 1, ii. p. 996 (1789—ex

Brisson & Daubenton, Pl. Enl. 58, fig. 2: Canada),

Dendreca estiva, Sclater & Salvin, l. c. p. 494 (Medellin) ;

Berlepsch, |. c. p. 282 (Bucaramanga).

Nos. 2593, 2136. g ad., ¢ juv. Condoto: 12.ix.; Sipi:

30.ix.08.

“Tris dark brown, feet light brown, maxilla black, mandible

grey.”

13. BASILEUTERUS TRISTRIATUS TRISTRIATUS Tsch.

Myiodioctes tristriatus Tschudi in Arch. f. Naturg. 10, 1. p. 283

(1844.—Peru); cfr. Berlepsch & Hellmayr, Journ. f. Orn. 1905,

p. @ (erit.).

Basileuterus tristriatus Berlepsch, Journ, f. Orn, 1884, p. 283

(Bucaramang’).

1092 MR. C. E. HELLMAYR ON THE

SL. auricularis Sharpe, Cat. B. Brit. Mus. x. p. 386 (1885.—

Bogota; Pallatanga, W. Ecuador; Simacu, Bolivia; no type

specified).

B. melanotis dedalus Bangs, Proc. Biol. Soc. Wash, xxi.

p- 160 (1908.—San Antonio, Rio Cali, W. Colombia, 5800 ft.).

No. 3748. g ad. Pueblo Rico, 5200 ft., 10.1x.09.— Wing 65 ;

tail 57; bill 103 mm.

Nos. 3749, 3751. 2 (@)ad., Q ad. Pueblo Rico: 10,11.1x.09.—

Wing 64, 613; tail 60, 56; bill 10 mm.

“ Tris dark brown, feet greyish yellow, maxilla black, mandible

grey.”

These specimens agree with a series from Bogota, Ecuador, and

Peru, some of which I had previously compared and found

identical with Tschudi’s type kindly forwarded by the authorities

of the Neuchatel Museum. Iam unable to discover any constant

character on which to separate the Colombian and Ecuadorian

birds from typical Peruvian skins, 6. auwricularis Sharpe being

based on purely individual variations, such as the colour of

the pale crown stripe etc. When compared with B. tristriatus

melanotis Lawr. *, of Costa Rica and Chiriqui, the three examples

from Pueblo Rico differ exactly as indicated by Mr. Bangs,

viz., much brighter green back, deeper (about maize) yellow

underparts, and large, blackish loral spot. However, in all these

points they are absolutely similar to typical tristriatus, to which

Mr. Bangs, evidently misled by Sharpe’s key, does not make

any reference at all. B. t. tristriatus had already been recorded

from Bucaramanga, N. Colombia, by Count Berlepsch.

While I cannot admit the distinctness of the Colombian birds—

for which, moreover, Sharpe’s term auricularis would be an earlier

name—the inhabitants of the mountains of Venezuela constitute

a fairly well-defined race, B. ¢. meridanus Sharpe 7, recognisable

by the reduction or absence of the black loral and auricular spots.

We have in the Munich Museum, three adults from Merida,

and five males and one female from the Cumbra de Valencia ;

and, at Tring, I have examined two skins from Caripé, State of

Cumana, and one adult from Buearito, Tocuyo (Mocquerys).

14. BASILEUTERUS FULVICAUDA SEMICERVINUS Scl.

[ Muscicapa fulvicauda Spix, Av. Bras. 11. p. 20, pl. xxviii. fig. 2

(1825—no locality) |.

Basileuterus senurcervinus Sclater, P. Z. 5. 1860, p. 84 (1860.—

Nanegal, W. Ecuador); Sclater & Salvin, P. Z. 8. 1879, p. 494

(Remedios, Neche); Berlepsch, Journ. f. Orn. 1884, p. 284

(Bucaramanga).

No. 613. Q ad. Juntas, Rio Dagua, 1000ft.: 1.viii.07.—

Wing 60; tail 49; bill 123 mm.

* B. melanotis Lawrence, Ann. Lyc. N. H. N.Y. ix. p. 95 (1868.—Cervantes, Costa

Rica). :

+ B. meridanus Sharpe, Cat. B. Brit. Mus x. p. 3887 (1885.—Merida, Venezuela).

BIRDS OF WESTERN COLOMBIA. ~ 1093

No. 2170. Q ad. Rio Garrapatas, Sipi, 150 ft.: .8.x.08.—

Wing 60; tail 48; bill 12 mm.

“Tris dark brown, feet light brown, bill black.”

These, as well as several other specimens from W. Colombia

in the Tring Museum (Raap and Palmer coll.), are practically

identical with topotypical skins from Nanegal, W. Ecuador. A

series from N.W. Ecuador is not different either.

15, SETOPHAGA RUTICILLA Linn.

Motacilla ruticilla Linneus, Syst. Nat. x. p. 186 (1758—ex

Catesby : Carolina).

Setophaga ruticilla Sclater & Salvin, 1. c. p. 494 (Concordia,

Medellin) ; Berlepsch, 1. c. p. 284 (Bucaramanga).

No. 2392. gad. Névita: 11.x11.09.

No. 3775, —. g gad. Pueblo Rico: xi.09; Siaté: 25.xi.09.

“Tris dark brown, feet black, maxilla black, mandible

brown.”

A common winter visitor to the north-western States of Bone

America.

16. PRoGNE CHALYBEA CHALYBEA Gm.

Hirundo chalybea Gmelin, Syst. Nat. 1, ii. p. 1026 (1789—

ex Brisson and Buffon: Ca emo

Progne chalybea Sclater & Salvin, |. c. p. 495 (Remedios).

Nos. 2356, 2360. ¢ g ad. Ndévita: 30.x1., 1.xii.08.—Wing 128,

125% tail 69; bill 12, 11 mm.

“Tris, feet, and bill black.”

Agreeing with examples from Venezuela, Bogota, and Cayenne,

17. SrELGrDOPrERYX RUFICOLLIS UROPYGIALIS Lawr.

[Hirundo ruficollis Vieillot, Nouv. Dict. xiv.. p. 523 (1817.—

“ Brésil ”’).|

Cotyle uropygialis Lawrence, Ibis, v. p. 181 (1863.— Panama).

Stelgidopteryx uropygialis Sclater & Salvin, |. c. p. 496

(Remedios); Berlepsch, J. f. Orn. 1884, p. 285 (Bucaramanga).

Nos. 2106, 2107. ¢ Q ad. Sipi: 22.ix.08.—Wing 107, 92;

tail 56, 47 mm.

‘“‘ Tris dark brown, feet and bill black.”

Agreeing with specimens from Chiriqui and Costa. Rica. Cfr.

Nov. Zool. xiii. 1906, p. 13.

18. DIGLOSSOPIS CHRULESCENS CARULESCENS Scl.

Diglossopis cerulescens Sclater, Ann. Mag. N. Hist. (2) xvii.

p- 467 (1856.—Caraceas, in Venezuela) ; Sclater & Salvin, P. Z.S.

1879, p. 496 (Santa Elena).

Dig cerulescens Berlepsch, Journ. f. Orn. 1884, p. 286

(Bucaramanga).

IEROC ACO, SOUR ING: IRON, re!

(1094 MR. C. BE. HELLMAYR ON THE

No. —. gad. Tatama, 2794 ft.: 12.x.09.—Wing (2; tail 58;

bill 13 mm.

‘Tris reddish brown, feet dark grey, bill black.”

Agrees with Bogota skins, whereas specimens from Merida are

a darker and more uniform bluish colour below with scarcely

any greyish admixture on the vent. Topotypical birds from

Caraccas are not available for comparison.

D. c. cerulescens is known only from the mountains of Colombia

and Venezuela. In Peru it is replaced by the nearly allied

D. c. pallida Berl. & Stolzm.*

19. DACNIS VENUSTA FULIGINATA Bangs.

'Dacnis venusta Lawrence, Ann. Lyc. N. H. N.Y. vii. p. 464

(1862.—Panama Railroad) |

D. venusta fuliginata Bangs, Proc. Biol. Soc. Wash. xxi. p. 160

(1908.—Jiménez, W. Colombia).

D. venusta (nec Lawrence), Sclater & Salvin, P. Z. 8. 1879,

p- 497 (Remedios).

No. 2215. gad. Noanama: 22.x.08.—Wing 62; tail 39;

bill 11 mm.

Nos. 2243, 2244. § gad. Novita: 7.x.08.—Wing 643, 65;

tail 403, 41; bill 10, 11 mm.

No. “385. gad. Jiménez, 1600ft.: 12.vi.07.—Wing 63;

tail 40; bill 103 mm.

No. —. d a Rio Dagua: 8.vi.95. Rosenberg coll.— Wing 63 ;

tail 403; bill 10 mm.

No. —. Q ad. Rio Dagua: 10.vi1.95. Rosenberg coll. Wing

64; tail 42; bill 11 mm.

‘“‘ Tris crimson, bill and feet black.”

In addition, I have examined two males from Jiménez (June

14, 28), in Mr. Rosenberg’s possession, D. v. fuliginata is a

perfectly good form. Compared with the series from Panama,

Chiriqui and Costa Rica in the Munich Museum, the males sent

by Mr. Palmer differ in having the underparts much darker,

deep black with hardly a trace of the greenish hue so conspicuous

in the Central American birds. The female is smaller tT, and

rather deeper ochraceous buff on the lower belly and under tail-

coverts than those from more northern localities. The bill, in

the southern race, is constantly shorter. Birds from Costa Rica

and Chiriqui have much longer wings and tail, while those from

Panama, in size, approach D. v. fuliginata.

Hight adult males from Chiriqui and Costa Rica measure :

wing 67-70; tail 43-45; bill 12-13 mm.

Three adult males from Panama (Railroad) measure : wing 633—

66; tail 42-43; bill 12-13 mm.

* P, Z.S. 1896, p. 334 (Chachapoyos, N. Peru ; Garita del Sol, C. Peru).

+ Three adult females SNH! Chiriqui measure: wing 65-67; tail 43-443; bill

—13 mm.

BIRDS OF WESTERN COLOMBIA. 1095

20. DAGNIS CAYANA CG@REBICOLOR Scl.

[ Motacilla cayana Linneus, Syst. Nat. xu. 1, p. 336 (1766—

ex Brisson: Cayenne; excl. Hernandez—Mexico)}.

Dacnis cerebicolor Sclater, Contrib. to Ornith. 1851, pp. 106—

12 (1851 (?)—“ New Granada?” sc. Bogotd, cfr. Sclater, Cat.

Coll, Amer. B., 1862, p. 51); Sclater & Salvin, P.Z.S. 1879,

p- 497 (Remedios— 3 imm., ‘‘not quite so bright as specimens

from Bogota”); Hartert, Nov. Zool. v. p. 481 (Paramba, N.W.

Keuador).

D. cayana (errore) Sclater & Salvin, 1. ec. p. 496 (Remedios).

Nos. 2006, 2205, 2214. ¢ gad. Noanama: 24.viii., 20.22.x.08.

—Wing 61-644; tail 424-434; bill 11-114 mm.

Nos. 2319, 2348, 2187. ¢ gad. Névita: 21,28.xi.; Sipi: 13.x.

08.—Wing 61-63; tail 424-44; bill 11-112 mm.

No. 2657. gad. Tadd: 21.v.09.—Wing 63; tail 42; bill

12 mm.

No. 2007. @ ad. Noanama: 24.viii.08.—Wing 59; tail 42;

bill 12 mm.

Nos. 2320, 2374, 2382. 9 9 ad. Né6vita: 21.xi., 7, 9.xii.08.—

Wing 59-62 ; tail 40-43; bill 104-12 mm.

Nos. 2188, 2073. 9 Q ad. Sipi: 10i1x., 13.x.08.—Wing 63,

61; tail 43, 424; bill 11 mm.

Nos. 2631, 2669. 9 2 imm. Tadé: 3, 27.v.09.—Wing 60, 62;

tail 43, 44; bill 11, 114 mm.

Nos. 507, 508, 518, 541, 564, 570, 647,688. ¢ gad. Jiménez,

1600 ft., 28.vi., 12, 13, 18, 23, 24.vii., 9.vi1i.07.—Wing 634-67;

tail 43-47 ; bill 12-13 mm.

No. —. g ad. Rio Oscuro, W. Cordillera, 3000 ft., June 1898,

Batty coll.— Wing 68; tail 49; bill 123 mm.

-Nos, 384, 400, 401, 486, 577, 642,652. 9 9. Jiménez, June to

Aug.— Wing 60-63; tail 42-44; bill 113-12 mm,

“Tris dark red in males, dark brown in females, feet pink,

bill black.”

This series is rather puzzling. The males show a wide amount

of individual variation, every shade of blue between the ultra-

marine of D. c. napaea and the deep purplish blue of D. c. cwre-

bicolor (of Bogota) being represented. The lightest specimen,

no. 2006, from Noanama has the blue portions of the plumage of

exactly the same tone, and the mantle as well as the gular patch

as dull (greenish black) as topotypical Santa Marta skins, from

which it only differs by its smaller size and shorter bill. Next

come no. 2214 (Noanama), no. 564 (Jiménez), and the male froin

Rio Oscuro, which are just a shade darker, more cobalt, with the

black of the mantle and throat deeper, less obscured by dull

greenish tips. Then follow ten specimens which have the plumage

more or less tinged with purple. Finally, three birds, nos. 2187

(Sipi), 508 and 541 (Jiménez), and a fourth from Paramba, N.W.

Keuador, are quite as bright and deep purplish blue as Bogota

skins.

74*

1096 MR. C. E. HELLMAYR ON THE

In coloration, there is no constant difference between the birds

from the foot-hills (Noanama, Névita, Tad6, Sipi) and those

from the more elevated districts (Jiménez, 1600 to 2900 ft. ; Rio

Oscuro, 3000 ft.). The former are, however, on the average, smaller,

and have a shorter, slenderer bill, while the males from Jiménez,

etc., in dimensions, agree with skins from Bogota and Buca-

ramanga.

The females also are rather variable. Most of them have the

top and sides of the head turquoise-blue, scarcely darker than in

the female of D.c. wltramarina from Panama, while in two or

three these parts are of the same dark ultramarine-blue colour as

in Bogota skins of D. c. cwrebicolor.

From what I have said above, it is evident that the majority

of the West Colombian birds are not typical ewrebicolor ; but in

view of their great individual variation, which completely connects

napaea, of Santa Marta, and cwrebicolor, of the Kastern Cordillera,

it seems inadvisable to separate them subspecifically since the new

“form” would mainly consist of ‘‘ intermediates.” One thing,

however, results from the study of that series, viz., that both

napaea and cerebicolor are merely geographical races of D. cayana,

the passage being formed by D.c. ultramarina. 'Thus, we have

the following forms :—

(a) D.c. cayana Linn. Eastern South America from Southern

Brazil to Trinidad, Guiana and Venezuela.

(b) D. ¢. glaucogularis Berl. & Stolzm.* From the eastern

slopes of the Colombian Andes south through Peru to

Eastern Bolivia and Western Brazil (Mattogrosso).

(Pozo Azul ete.).

(d) D.c. ultramarina Lawr.t Isthmus of Panama, Eastern

Costa Rica, H. Nicaragua.

(e) D. c. napaea Bangs§. Santa Marta District, N. Colombia.

(f) D. c. cerebicolor Scl. Andes of Colombia (Western and

Eastern Cordillera) and N.W. Ecuador. Typical in the

Eastern Cordillera.

On some future occasion I hope to give more details about the

variation and geographical distribution of the various races.

21. CHLOROPHANES SPIZA EXSUL Berl. & Tacz. (2)

[Motacilla spiza Linneus, Syst. Nat. x. p. 188 (1758—ex

Edwards: Surinam). |

Chlorophanes spiza exsul Berlepsch & Taczanowski, P. Z.5S.

1883, p. 543 (1884—Chimbo, 8.W. Keuador).

* P. Z.S. 1896, p. 336 (Central Peru: La Gloria, La Merced).

+ Proc. Biol. Soc. Wash. xviii. p. 154 (1905.—Divala, Chiriqui).

Proc. Acad. N. Sci. Philad. 1864, p. 106 (Isthmus of Panama).

Proc. Biol. Soc. Wash. xii. p. 143 (1898.—Santa Marta),

BIRDS OF WESTERN COLOMBIA. 1097

C. atricapilla (errore) Sclater & Salvin, P. Z. 8. 1879, p. 497

(Concordia, Remedios). -

Nos. 2211, 2248, 2385. ¢ gad. Noanama, 22.x.;- Novita:

8.xi., 10.x11.08,— Wing 69 ; tail 45-48; bill 13-133 mm.

Nos. 2212, 2333. ¢g juv., Qad. Novita: 24.xi.; Noanama :

22.x.08.— Wing 68, 65; tail 45; bill 133 mm.

“Tris crimson, feet green, maxilla black, mandible yellow.”

The adult males agree with typical specimens from West

Ecuador in size and in smallness of the bill; but the coloration

is more bluish, especially below, though much less so than in the

Upper Amazonian race, C. s. cerulescens Cass.* Birds from

Chiriqui and Costa Rica (El General de Terraba, Carrillo, La

Vijagua, ete.) are again more greenish, have longer wings (70—

74 mm.), and a longer, more robust bill (143-163 mm.). They

are unquestionably subspecifically distinct. A larger series might

perhaps enable us to separate the West Colombian birds, but for

the present I leave them with C.s. exsul. According to this view,

C. s. exsul ranges from 8.W. Ecuador north to the Pacific slopes

of Colombia.

22. CYANERPES CHRULEA MICRORHYNCHA Berl.

[Certhia cerulea Linneus, Syst. Nat. x. p. 118 (1758—ex

Edwards: Surinam). |

Cereba cerulea microrhyncha Berlepsch, Journ. f. Orn. xxxil.

p. 287 (1884—part. : type ex Bucaramanga, N. Colombia).

Cereba cerulea (nec Linneus) Sclater & Salvin, P. Z. S.

1879, p. 497 (Remedios, Medellin).

Nos. 2257, 2262. ¢ gad. Novita: 10,11:xi1.08.—Wing 55,

534 ; tail 29, 273; bill 18 mm. 3

No. 1961. g ad. San Joaquim, Bahia del Choeé, 3.vi1.08.—

Wing 54; tail 28; bill 173 mm.

No. 2241. g ad. Rio Cajén: 5.xi.08.—Wing 55; tail 28;

bill 173 mm.

No. 2238. ¢ juv. Cajon: 2:x1.08.

Nos. 2242, 2349. 92 Qad. Rio Cajén: 5.xi.08; Noévita:

28.xi.08.— Wing 55; tail 27; bill 17 mm.

“ Tris dark brown (¢ ¢), black (2), bill black, feet wax-yellow

(3), Seem (2)

The adult males agree in the shortness of the bill, and in the

pale azure-blue colour of the cheeks and anterior portion of the

crown, with a number of Bogota skins, but have slightly shorter

wings and tail. Two adult males from Pozuzo (prov. Huanuco,

Peru) are equally small while the bill is even a little shorter.

The difference in size is, however, not likely to be constant, since

a single male from San Augustin, N. Bolivia (3500 feet), has

wings, tail, and bill fully as long as Colombian examples. There

* Chlorophanes cerulescens Cassin, Proc. Acad. N. Sc. Philad. Nov. 1864, p. 268

(1865.—Yuracares, N.E. Bolivia).

1098 MR. GC. E. HELLMAYR ON THE:

can be no question that C.c. microrhyncha is a perfectly valid

form. Cfr. my remarks in Nov, Zool. xii. 1905, p. 9, and J. ¢. xiv.

1907, pp. 42-43.

Its range extends from the mountains of Colombia south to

Northern Bolivia.

23. C@REBA CHLOROPYGA MEXICANA Scl.

[Certhiola chloropyga Cabanis; Mus. Hein. i. p. 97 (1850.—

Bahia). ]

Certhiola mexicana Sclater, P. Z. S. 1856, p. 286 (Jan. 1857.—

S. Mexico, probably Vera Cruz); Sclater & Salvin, P. Z. 8.

1879, p. 497 (Remedios, Medellin).

No. 2822. 9 ad. Pueblo Rico, 5200 ft. : 2.xi.09.— Wing 523 ;

tail 32; bill 12 mm.

“Tris dark brown, feet dark grey, bill black.”

Identical with skins from Bogota and Western Ecuador which

I am unable to separate from others taken in Mexico and

Costa Rica. ;

Birds from Bucaramanga form the transition to C. c. luteola

Cab. Cfr. Berlepsch, Journ. f. Orn. 1884, p. 288.

24. ORYZOBORUS FUNEREUS £THIOPS Scl.

[Oryzoborus funereus Sclater, P. Z.S. 1859, p. 378 (1860.—

Oaxaca, Mexico). |

Oryzoborus cwthiops Sclater, P. Z. 5. 1860, p. 88 (1860.—

Nanegal, W. Ecuador).

No. 1988. g ad. Mouth of Calima, San Juan River, 13.vii.08.

—Wing 55; tail 51; bill 12 mm.

‘‘Tris dark brown, feet and bill black.”

This bird agrees, in size and colour, with a series from Western

Ecuador in the Munich Museum. Specimens from Guatemala,

which I take to be typical O. f. funereus, have longer wings

(60 mm.) and tail (54 mm.), a slightly larger bill, and the females

are much brighter cinnamon underneath. These slight differences

might disappear on comparison of a larger series of Central

American skins, but, for the present, I am unwilling to unite the

two forms without further evidence. O./. ethiops, as defined

here, ranges over the forest district of Western Colombia and

Ecuador. Specimens from Antioquia * and Santa Marta which

I have not seen may also be referable to the southern race.

25. SPOROPHILA OPHTHALMICA Scl.

Spermophila ophthalmica Sclater, P. Z. 8. 1860, p. 276 (end

of 1860—Babahoyo, W. Ecuador); Sharpe, Cat. Birds, xii.

p- 120 (Chocé Bay).

Nos. 1979, 1991. ¢ gad. Mouth of Calima, 13.vili.08;

* O. funereus Scl, & Salvin, P. Z. S. 1879, p. 506 (Medellin).

BIRDS OF WESTERN COLOMBIA. 1099

Guineo, R. Calima: 7.viii.08.—Wing 54-55; tail 45, 47;

bill 10 mm.

No. 2134. Q ad. Sipi: 30.ix.08.—Wing 53; tail 43;

bill 10 mm.

‘‘ Tris dark brown, feet and bill black.”

One of the males (no. 1979: Guineo) agrees with West

Ecuadorian skins, the throat being white with a small black spot

at the base of the mandibular rami. In the other specimen

(no. 1991) the feathers of the upper throat are black, tipped with

white. Both have the black breast-band rather wider than the

large majority of Ecuador skins, though one or two of the latter

hardly differ in that respect.

The male from the mouth of the Calima (no. 1991) approaches

very closely black-throated males of S. aurita, and can only be

distinguished by the white tips to the feathers of the throat !

S. ophthalmica is no doubt merely a southern race of S. aurita™,

but until the status of this puzzling “ species” is better understood,

it would be unwise to employ a trinomial appellation. Mr.

Ridgway’s remarks (in Bull. U.S. Mus. no. 50, Part 1, 1901,

p. 573) should be carefully consulted.

S. ophthalmica is restricted to W. Ecuador and 8.W. Colombia.

96. TERSINA T VIRIDIS OCCIDENTALIS Scl.

[Hirundo viridis Mliger, Prodr. Syst. Mamm. & Av. p. 229

(1811—based on “ L’Hirondelle verte,’ Temminck, Cat. Syst.,

Mus. N. Y. ii. 1889, p. 70.]

Procnias occidentalis Sclater, P. Z.S. 1854, p. 249 (Apr. 1855.

—-“‘ Nova Grenada ”).

P. tersa (nec Linneus) Sclater & Salvin, P. Z. 8, 1879, p. AQT

(Remedios, Antioquia).

P. cerulea occidentalis Berlepsch, Journ. f. Orn. 1884, p. 288

(Bucaramanga).

No. 2035. gad. Noanama: 29.viii.08.—Wing 87; tail 56 ;

bill 9 mm.

Nos. 2404-6, 2412. gdad. Noévita: 16, 18.xii.08—Wing

81-85 ; tail 52-55: bill 9-10 mm.

‘‘ Tris brown, feet and bill black.”

The series agrees, in size and coloration, with other specimens

from Bogota, Eastern Ecuador, Venezuela (Cumana), etc. Cr.

my remarks in Nov. Zool. xvii. 1910, p.271. Birds from Eastern

Brazil (Bahia, Rio) are much larger, and the males have the

plumage of a decidedly paler, more greenish blue (nile-blue).

* Not having seen white-throated males (=hicksii Lawr.) Iam unable to say

in what way they differ from S. ophthabmica, but, judging from published de-

scriptions, the only character of the northern form appears to consist in the rather

wider black band across the chest.

+ For change of generic name see Ridgway, Bull. U. S. Mus. no. 40, pt. iv. p. €89,

footnote b

1100 ; MR. GC. E. HELLMAYR ON THE

27. EuPHoNIA FruLyrcriIssa Sel. [a subsp. ®].

Euphonia fulvicrissa Sclater, P. Z.S. 1856, p. 276 (Jan.

1857.— “Santa Martha in New Grenada”); Cassin, Proc.

Acad. N. Sci. Philad. 1860, p. 143 (Falls of the Truando) ;

Selater & Salvin, P. Z. S. 1879, p. 498 (Remedios, Neche:

Antioquia); Berlepsch, Journ. f. Orn, 1884, p. 288 (Bucara-

manga).

No. 2036. g ad. Noanama, 29.viii.08.—Wing 51; tail 29;

bill 82 mm.

No. 2530. Q ad. El Tigre, 10.11.09.—Wing 50; tail 28;

bill 8 mm.

“Tris dark brown, feet black, bill black, lower mandible blue.”

The adult male from Noanama and another, picked out from a

set of Bogota skins, in the Munich Museum are exactly inter-

mediate between specimens of H. fulvicrissa, from Panama and

Chiriqui, and #. f. purpurascens Hart.*, from N.W. Ecuador.

The back, upper wing-coverts, sides of the head and throat are

glossy metallic steel-blue in the Colombian skins, dull greenish

‘blue or bottle-green in those from Central America, and bright

purple-blue, inclining to violet, in purpurascens. Two skins of

the last named form? have very little, or no white at all, on the

outermost rectrix, while in the specimens from Chiriqui,

Panama, and Bogota a.large portion of the inner web is white.

As, however, the male from Noanama agrees, in that respect, with

purpurascens, much importance cannot be laid upon this

character.

_ The type of Z. fulvicrissa in the British Museum, said to be

from Santa Marta, where, however, recent travellers did not meet

with the species, is to be examined in order to ascertain whether

it belongs to the Colombian ov Veraguan race.

28. KuPHONIA XANTHOGASTER CHOCOENSIS Hellm.

[Euphone sxanthogaster Sundevall, Vetenskaps Akad. Hand.

1833, p. 310, pl. 10. fig. 1 (= ¢ ad.) (1834.—* Brazil”).

Euphonia wanthogaster chocoensis Hellmayr, Rey. Frang. @Orn.

no. 22, p. 23 (1911.—Rio Cajén, W. Colombia).

No. 2015. gad. Noanama: 26.vili.0@2—Wing 61; tail 354;

bill 74 mm.

Nos. 2235, 22367. g¢ g ad. Rio Cajén: 3.xi.08— Wing 58,

D9; tail 33, 34; bill 8 mm.

Nos. 2086, 2176. 9 9 ad. Sipi: 16.ix., .10.x.08.— Wing 56,

57; tail 30, 31; bill 73-8 mm.

“Tris black (¢), dark brown ( @ ), feet dark grey (3), grey (2),

bill black, mandible blue.”

_ * Novit. Zool. viii. p. 377 (1901.—Pambilar, N.W. Ecuador; type in Tring

Museum).

+ The type from Pambilar, and anotuer from San Javier, both in the Tring

Museum.

t Type of subspecies.

BIRDS OF WESTERN COLOMBIA, 1101

' This new subspecies is most nearly allied to typical E. wantho-

gaster, from: Brazil, Peru and Ecuador, but differs in its generally

smaller size, especially shorter and, smaller bill. The males, too,

have the underparts much clearer chrome-yellow, without any

trace of the orange tinge which is always well pronounced along

the middle line in the eastern race; the females also may be

distinguished by the nearly pure cinereous instead of buffy grey

colour of the throat and breast, and by having the middle of the

abdomen much paler buffy. Forehead and crown, in the males,

are clear chrome-yellow, as in average Spee ines of wanthogaster.

E. xanthogaster brevirostris Bonap.*, from Bogota, is much

larger (wing 64-65; tail 38-40; bill 9 ‘mm. ), has the foreneck

and middle of the belly tinged with orange, and the occiput of a

deep brownish orange colour. 4. «. brunneifrons Chapm. t, from

South-eastern Peru, is of the same size as brevirostris, but the

crown is still darker, dull orange rufous.

The new form has been compared with large series of the

various races. Besides the thirteen specimens in the Munich

Museum, I had before me twelve from the Berlepsch Collection,

and three belonging to the Vienna Museum.

29. CHLOROCHRYSA NITIDISSIMA Scl.

Chlorochrysa nitidissima Sclater, P. Z. 5. 1873, p. 728 (1874. —

Antioquia); idem, Ibis, 1875, p. 466, pl. x.; Sclater & Salvin,

P. ZS. 1879, p. 498 (Jerico, Antioquia).

No. 3757. 3S imm. Siat6, Rio Siaté: 16.1x.09.—Wing 67 ;

tail 44°; bill 12 mm.

No, 3745. 9 imm. Pueblo Rico: 9.1x.09.—Wing 67; tail 43;

bill 13 mm.

“Tris brown, feet grey, bill black.”

This species was discovered by the late T. K. Salmon at Jerico,

south of Medellin, on the left side of the Cauca River, in the

Western Cordillera of Colombia. Dr. Sclater described and

figured the adult male. Specimens transmitted by the late J.

H. Batty from Riolima (Cauca Valley) agree perfectly with the

or iginal description.

The immature male and female differ in lacking the lemon-

yellow interscapulium (which is mainly green with but a faint

yellowish tinge) and in having the middle of the belly dull grey

instead of black. Moreover, the general colour is not so bluish.

C. nitidissima is apparently restricted to the Western

Cordillera of Colombia, and found only at altitudes above

5000 feet.

30. CALOSPIZA JOHANNA Dalmas.

Calliste johanne Dalmas, Bull. B. O.C. xi. p. 36 (1900.—E]

* Huphonia brevirostris Bonaparte, Rev. Mag. Zool. (2) ili. p, 136 (1851; ex

Columbia).

HE Euphonia wanthogastra Bysnanmnesseaawas Chapman, Bull. Amer. Mus, N. H. xiy.

226 (1901.—Inca Mines, 8.E. Peru).

1102 MR. C. E. HELLMAYR ON THE

Paillon, near Buenaventura, Chocé, W. Colombia) ; Sclater, Ibis,

1901, p. 597, pl. xii. fig. 2 (Paramba, N.W. Ecuador).

No. 2698. gad. Tadé, 7.vi.09.—Wing 73; tail 50; bill

105 mm.

No. 2388. 9 ad. Novita: 10.xii.08.—Wing 67; tail 46;

bill 10 mm.

No. 2589. ¢ juv. Condoto: 10.iv.09.—Wing 714; tail 49;

bill 9 mm.

“Tris dark brown, feet dark grey, bill black.”

Two more specimens, a female and an immature bird, sex not

determined, from Tadé, June 1909, are in Mr. Rosenberg’s

possession.

There is very little difference between male and female, except

that the latter is rather smaller and not quite so bright on the

head and breast. The young bird lacks the golden hue on the

green portions of the plumage, the black of the head and throat

ismuch duller, and the beautiful golden yellow rump of the adults

is but faintly indicated by a smail patch of dull yellow.

C. johanne is another species peculiar to the humid forest

region of the Pacific coast. Besides the above specimens, and

the type (now in the Tring Museum), the only other examples

on record are two from Paramba, N.W. Ecuador, procured by

one of Mr, Rosenberg’s correspondents *.

31. CALOSPIZA RUFIGULA Bonap.

Tanagrella rufigula Bonaparte, Compt. Rend. Acad. Sci. Paris,

xxxil. (séance 20. Jan.) p. 77 (1851.—Keuador, Bourcier colli.) ;

idem, Rev. Mag. Zool. (2) iii. p. 130 (mars 1851,—reprint of orig.

descr.).

Calliste rufigularis Sclater, Cat. B. Brit. Mus. xi. p. 107 (Pasto,

Colombia t— Lehmann).

No. 2797. @ad. La Selva: 4600 ft., 15.x.09.—Wing 60;

tail 44; bill 10 mm.

“Tris dark brown, feet blue-grey, maxilla black, mandible

blue.”

This bird is identical with typical specimens from Ecuador,

La Selva is the most northerly locality for C. rufigula yet known,

though an example had already been taken by Mr. Lehmann at

Pasto in Southern Colombia, near the Ecuadorian frontier.

32. CALOSPIZA AURULENTA AURULENTA Lafr,

Tanagra (Aglaia) aurulenta Latfresnaye, Rev. Zool. vi. p. 290

(1843.—“ Colombie,” se. Bogota).

Calliste aurulenta Sclater & Salvin, P. Z. 8. 1879, p. 498 (Con-

cordia, Frontino).

* Dr. Selater’s record (Ibis, 1901, p. 597) from Peru is a mistake, cfr. Hellmayr,

Ibis, 1910, p. 328, footnote fF.

+ Not Ecuador as stated 0. cit. —

BIRDS OF WESTERN COLOMBIA. 1103

No. 3747. 9 ad. Pueblo Rico: 5200 ft., 10.ix.09.— Wing

73; tail 48; bill 10 mm.

“Tris dark brown, feet grey, bill black.”

We have also several specimens from Rio Lima, 6000 ft.,

collected by the late J. H. Batty in August 1898. They agree

perfectly with typical Bogoté skins, while birds from Western

Keuador are generally slightly paler on the head and belly.

C. aurulenta sclatert Lafr. replaces it in the Eastern Cordillera

of Colombia. It is known to me only from Bogota specimens.

33, CALOSPIZA ICTEROCEPHALA Bonap.

Calliste icterocephala Bonaparte, Compt. Rend. Ac. Sci. Paris

xxxii. (séance 20. Jan.) p. 76 (1851.—Ecuador, Bourcier coll.) ;

Sclater, Contrib. Orn. 1851, part ii. April, p. 53, pl. Ixx. fig. 1 (the

type stated to have been obtained in the valley of Punta Playa,

south of Quito); Sclater & Salvin, P. Z. S. 1879, p. 498

(Frontino).

Nos. 2825, 2837. 2 2 ad. Pueblo Rico: 2, 6.xi.09.— Wing 71,

69; tail 45, 46; bill 11 mm.

“Tris dark brown, feet grey, bill black.”

Agreeing with specimens from Ecuador and Chiriqui. C’ zetero-

cephala has a wide range, extending from Costa Rica to Western

Keuador. Jam unable to detect differences, either in size or in

colour, between skins from various localities.

34. CALOSPIZA LAVINIA LAVINIA Cass.

Calliste lavinia Cassin, Proc. Acad. N. Sci. Philad. x. p. 178

(1858.—‘ Isthmus of Darien, New Grenada ”).

C’. lavinice Cassin, Proc. Ne N. Sci. Philad. 1860, p. 142 (“ Camp

Toucey, in the mountains of the Rio Truando”—one ¢ imm.);

idem, |. ec. 1864, p. 286, pl. 1. fig. 1 (figure of type).

C. envilive Dalmast Bull. B. O. C. xi. p. 35 (1900.—San José and

El Paillon, near Buenaventura, W. Colombia); Sclater, Ibis,

1901, p. 596, pl. xii. fig. 1 (S. Javier, Rio Cachabé, N.W. Ecuador) ;

‘ofr. Hellmayr, Rev. Frang. d’Orn. no. 11, 1910, p. 161-162

(crit.).

Nos. 2149, 2159. ¢ g ad: Sipi: 2, 5.x.08.—Wing 67, 68;

tail 43, 44; bill 10 mm.

Nos. 2183, 2190. $ ¢ juv. Sipi: 12, 14.x.08—Wing 65;

tail 44; bill 94 mm.

No. 2191. Q@ ad. Sipi: 14.x.08.—Wing 63; tail 42; bill

93 mm.

“« Tris light brown, feet grey, maxilla dark, mandible lighter

brown.”

In addition, the Munich Zoological Museum possesses one

adult, one young male and two females, from San José, near

Buenaventura, 600 feet alt., which were formerly in Comte de

Dalmas’ collection and formed part of the typical series of

C. emilie Dalm. As pointed out by me in Revue Frang. @Orn.

1104 MR. C!. E. HELLMAYR ON THE

No. 11, p. 161-162, the blue-throated form is, however, the true

C. lavinia of Cassin, while the birds inhabiting Veragua and Costa

Rica will have to stand as C. l. dalmasi Hellm. C. Ll. lavinia

ranges from Darien through Western Colombia to N.W. Ecuador,

whence the Tring Museum received several specimens, obtained

by Mr. G. Flemming at San Javier, on the banks of the Rio

Cachabi. It may be mentioned that the same institution

possesses also an adult male, taken by Mr. W. F. H. Rosenberg

in 1895 at Juntas, Rio Dagua.

35. CALOSPIZA GYROLOIDES GYROLOIDES Lafr.

Aglaia gyroloides Latresnaye, Rev. Zool. x. p. 277 (1847—new

name for Aglaia peruviana (nec Tanagra peruviana Desmarest)

Swainson, Anim. in Menag. 1838, p. 356: ‘‘ Peru”—errore! we

substitute Colombia).

Calospiza gyroloides deleticia Bangs, Proc. Biol. Soc. Wash. xxt.

p- 160 (1908.—San Antonio, W. Colombia, 5800 ft.).

Calliste gyroloides Wyatt, Ibis, 1871, p. 325 (Mountain chain

between Bucaramanga and the Magdalena); Berlepsch. Journ. f.

Orn. 1884, p. 289 (Bucaramanga); Sclater & Salvin, P.Z.8.

1879, p. 499 (Concordia, 6000 ft. ; Remedios, 2360 ft.).

No. 632. ¢ ad. Jiménez, 1600 ft.; 6.vii.07.—Wing 81;

tail 55; bill 113 mm.

Nos. 2838, 3754. ¢¢ ad. Pueblo Rico, 5200 ft.; 6.xi.,

14.ix.09.— Wing 783, 82; tail 54, 57; bill 12 mm.

No. 3766. gad. Siaté, Rio Siato, 5200 ft. ; 22.1x.09.— Wing

79; tail 54; bill 12 mm.

Nos. 3753, 3767. 2 Q ad. Siaté, Pueblo Rico: 13, 22.1x.09.—

Wing 74, 75; tail 52, 51; bill 113 mm.

“ Tris dark brown, feet leaden grey, bill black.”

This series as well as other specimens from Rio Lima (Cauca

Valley, 5000 ft.), Bucaramanga and Bogota bear out the character

claimed by Mr. Bangs for his deleticia, viz., all have the lesser

wing-coverts green like the remainder of the wing. Unfortunately,

however, this form is the true gyroloides, as may be easily seen

on referring to the'original description. Lafresnaye’s name was

proposed as a mere substitute for the preoccupied term perwviana

of Swainson, who expressly says: ‘‘ shoulder-coverts green, instead

of golden yellow.” The habitat assigned to perweiana by

Swainson—Lafresnaye does not trouble himself with localities—

is, of course, erroneous since Peruvian birds, as will be shown

hereafter, possess a very large, golden yellow shoulder patch.

Swainson’s type specimen is more likely to have come from the

highlands of Colombia, to which the green-shouldered race appears.

to be confined. While I cannot, therefore, agree with Mr. Bangs

in the application of the name gyroloides, yet this author is

perfectly right in considering the Colombian and Central American

yaces as distinct. The study of the fine series in the Munich

Museum, together with other specimens lent by Count Berlepsch,

Dr. Hartert, and Dr. yon Lorenz of Vienna, shows that there

BIRDS OF WESTERN COLOMBIA. 1105

are at least three geographical forms of C. gyroloides to be dis-

tinguished. In the following lines I give a short resumé of their

Eicon which, I hope, will enable ornithologists to identify

the various subspecies.

(a) C. GYROLOIDES GYROLOIDES Lafr.

Hab. Mountains of Western and Central Colombia: Jiménez

(1600 ft.), Pueblo Rico, Siato (5200 ft.), San Antonio (5800 ft.),

province of Choco (Palmer) ; Rio Lima (5000 ft.), Cauca (Batty) ;

Concordia (6000 ft.), Remedios (2360 ft.), Antioquia (Salmon) ;

Bucaramanga (Minlos, Wyatt); Fusagasuga (6000 ft.: Wheeler) * ;

also found in Bogota collections.

Adult. Bill large and heavy; rufous chin spot immediately

followed by the cerulean blue colour of the under parts; rufous

crown without any yellow border behind or with but a very

narrow, yellowish line; lesser wpper wing-coverts green like the

median and greater ones, sometimes slightly more yellowish green

than the latter, but never yellow.

Specimens from different localities present the following

measurements :—

Four adult males from Chocé: Wing 783-82; tail 54-57 ; bill

114-12 mm.

One adult male from Rio Lima, Cauca: Wing 82; tail 54;

bill 12 mm.

One adult male from Bucaramanga (Mus. Berlepsch): Wing

78; tail 53; bill 11 mm.

Seven adults from Bogota coll.r: Wing 75-82; tail 52-56 ;

bill. 113=12 mm. *

Two females from Choco: Wing"/4, 75; tail 52, 51; bill 114,

12 mm. -

(b) C. GYROLOIDES BANGSI, subsp. n.

Hab. Costa Rica, Chiriqui, and apparently Western Ecuador.

Type in the Zoological Museum of Munich, No. 09.5340. ¢ ad.

Boquete, Chiriqui, 3500 feet, December 17, 1904. H. J. Watson

coll.

Adult. Like C. g. gyroloides. with no or very little green,

between the rufous chin-spot and the cerulean blue colour of the

under parts; yellowish border to posterior edge of rufous cap

absent or but faintly indicated : bill large and heavy ; ; but differs

in having the anterior lesser wing (or shoulder) coverts pale golden

yellow, this colour forming a conspicuous patch on the wing,

abruptly contrasting with the green of the remaining portion.

Specimens from different localities measure as follows :—

Six adult males from Chiriqui: Wing 78-80; tail 54-55;

bill 12 mm.

* Mr. Rosenberg tells me that there are two specimens of the green-shouldered

race from this locality in the British Museum, and that Salmon’s two skins from

Antioquia belong also here.

+ Four in the Munich Museum, three in the collection of Count Berlepsch.

1106 MR. C. E. HELLMAYR ON THE

One adult male from Costa Rica (Naranjo): Wing 81 ; tail 56;

bill 133 mm.

Hight adult males from Western Ecuador (Intag, 8. Nicolas,

Gualea): Wing 78-80; tail 52-58; bill 12-122 mm.

Five adult males from Paramba, N.W. Ecuador: Wing 75-79 ;

tail 52-55; bill 12 mm.

Remarks. It is a curious fact that the birds from Western

Eeuador should belong to the Central American race, and not to

C. g. gyroloides which is found in the neighbouring republic of

Colombia. However, all of the many specimens—nearly thirty

from Paramba, and eleven from other localities in Western

Kceuador—which I have been able to examine showed the pale

golden yellow shoulder-spot well-developed, thereby differing very

markedly from the Colombian form with its uniform green wings.

Although the majority of the Ecuadorian skins have the blue of

the rump rather lighter, and the top and sides of the head

somewhat clearer rufous, yet many specimens are quite indis-

tinguishable from the Chiriqui ones. I do not, of course, believe

in a discontinuous distribution, and fully expect that C. g. bangsi

will be found to exist in the Colombian coast district, having

thus an uninterrupted range from Western Ecuador to Chiriqui,

while C@. g. gyroloides is most probably confined to the high, open

country of the interior. This view is also shared by Mr. W. F. H.

Rosenberg, who has a considerable knowledge of the local distri-

bution of birds in those western districts.

The single Costa Rica male has a decidedly longer bill than any

other example examined by me.

I have named the new fog Mr. Outram Bangs, of Boston,

in recognition of his numerdUs important contributions to the

ornithology of tropical America.

Hab. Upper Amazonia: from the eastern slopes of the Andes

in Colombia (Rio Meta*) and the banks of the Rio Negro

(Marabitanas, Rio Xié) through Eastern Ecuador and Peru to

Northern Bolivia (Yuracarés).

Type in the Zoological Museum of Munich, No. 11,399 ¢ ad.

Chaquimayo Carabaya, 8.E. Peru, 3000 feet, 29.viii.1910. H. &

C. Watkins coll. no. 327.

Adult. Smaller, with a much weaker, slenderer bill ; rufous cap

bordered behind by a very distinct, golden yellow band, from 2

to 3mm. wide; upper throat dark green, separating the rufous

chin-spot from the blue foreneck ; the whole of the lesser and the

adjoiming median upper wing-coverts deep orange golden, forming

a very large, bright shoulder-patch. ‘'I'his patch is about twice as

large asin OC. g. bangsi, and of a much deeper, more orange golden

tinge.

* According to Mr. Rosenberg, the British Museum has four adult birds of this

form from the Rio Meta (800-1500 ft.; Wheeler coll.) and several others obtained

by Buckley in Eastern Ecuador.

BIRDS OF WESTERN COLOMBIA. 1107

Specimens from different localities measure as follows :—

Three adult males from Chanchamayo (C. Peru): Wing 75-77;

tail 49-53 ; bill 10-11 mm.

One adult male from N. Peru (Huayabamba).—Wing 73;

tail 48; bill 11 mm.

Four adult males from 8.E. Peru (Marcapata): Wing 73-76 ;

tail 52-54; bill 11 mm.

Three adult males from Upper Rio Negro*: Wing 70-72: tail

47-493; bill 10 mm.

Two adult males from the Rio Putumayo, §.E. Colombia

(Mus. Berlepsch): Wing 703, 71: tail 49, 50; bill 10-104 mm.

Two adult males from Bogota collections: Wing 75, 76; tail

53, 54; bill 103, 11 mm.

Remarks.—These fifteen examples differ very conspicuously

from C. g. bangsi in the characters given above. The three from

the upper Rio Negro and one from the Putumayo have the head

of a deep chestnut-brown (instead of cinnamon-rufous) colour, but

the second specimen from the Rio Putumayo and the two Bogota

skins agree with the Peruvian ones, The dimensions are also

rather variable, though apparently not connected with any

particular geographical area.

I have named this well-characterized new form after my wife,

who takes considerable interest in ornithology and has materially

helped me in the preparation of this report.

36. CALOSPIZA PALMERI Hellm.

Calospiza palmeri Hellmayr, Rev. Frang. dOrnith. No. 4, p. 49

(1909.—Sipi, W. Colombia); idem, Ibis, 1910, p. 330, pl. v.

Nos. 2164, 21667, 2167, 2186. ¢ ¢ ad. Sipi, Rio Sipi:

7, 13.x.08.—Wing 803-83 ; tai] 554-60; bill 10-11 mm.

No, 2165. 9 ad. Sipi: 7.x.08.—Wing 763; tail 50; bill

11 mm.

‘“‘ Tris dark brown, feet and bill black.”

This fine new bird was fully described by me J. c. and is faith-

fully depicted on plate v. in ‘ The Ibis,’ 1910.

Mr. Palmer has not succeeded in getting additional specimens

of this Tanager.

37. CALOSPIZA LARVATA FANNY Lafr.

Aglaia Fanny Lafresnaye, Rev. Zool. x. p. 72 (1847—* in Nova

Grenada ” (Delattre)—sc. Buenaventura, W. Colombia.)

Calliste francesce Cassin, Proc. Acad. N. Sci. Philad. 1860,

p. 142 (Turbo, N.W. Colombia).

* Two from the Rio Xié, June 1831, one from Marabitanas, March 1831, obtained

by J. rine Vienna Museum [= Calliste gyroloides Pelzeln, Orn. Bras. iii. 1869,

. 207].

P + Type of species.

{ The type is in the Museum of the Academy of Natural Sciences in Phiiadelphia

(cf. Stone, Proc. Acad. N. Sci. Philad. v. 1899, p.51),and not in Baron de Lafresnaye’s

Collection, as erroneously stated by Ridgway (Bull. U. S. Mus. no. 50, pt. ii. p. 50).

1108 MR. C. E. HELLMAYR ON THE

O. larvata francesce Hartert, Nov. Zool. v. 1898, p. 482

(Cachabi, N.W. Ecuador).

Calliste larvata (errore, nec Dubus) Sclater & Salvin, P. Z.8.

1879, p. 499 (Remedios, Antioquia).

Nos. 2050, 2062. ¢ gd ad. Noanama: 2, 4.ix.08.— Wing 69 ;

tail 474-49 ; bill 10 mm.

19.x.08.—Wing 65-682 ; tail 44-47 ; bill 10 mm.

No. 2408. 2 ad. Ndévita: 17.xii.08.—Wing 653; tail 46;

bill 11 mm.

No. 2032. $ juv. Noanama: 28.vi1i.08.

‘“‘ Tris dark brown, feet and bill black.”

This is the true C. J. fanny which was originally described from

specimens obtained by Delattre in the same district of Western

Colombia. The comparison of these birds with the large

series in the Munich Museum clearly proves that the Central-

American form has been wrongly referred to C. l. fanny by

Sclater, Ridgway, and others. (. 1. fanny as represented by

specimens from Western Colombia and N.W. Ecuador (which

are exactly similar) is an exceedingly well characterized race. The

greater upper wing-coverts and the remiges are either uniform

black or show only the faintest trace of greenish fringes in their

apical half; the lower back and rump are very pale sky-blue ; the

blue frontal patch is very strongly developed, reaching as far as

the posterior edge of the eye; the flanks are washed with pale

sky-blue, without any purple-blue admixture; the bill is small

and slender. Birds from Chiriqui and 8.W. Costa Rica (Guana-

caste, Pozo Azul de Pirris), however, differ at a glance in having

the greater wing-coverts and remiges very distinctly margined

with pale green, and the flanks much more extensively and darker

blue, with a strong purplish blue tingeanteriorly. Moreover, the

blue frontal patch is much more restricted, never reaching further

back than to above the middle of the eye, and the bill is larger

and stouter. In all of these points they agree with C. 1. larvata

Dubus*, of which we have a fair series from Guatemala and

Eastern Costa Rica (Carrillo), but may easily be distinguished

from that form by their coppery-golden, instead of deep coppery

reddish-brown throat, much paler blue cheeks with very little

purplish tinge, and slightly paler blue rump.

CO. 1. fanny is restricted to the forest-belt bordering the Pacific

coast, ranging from N.W. Ecuador (Cachabi, 8. Javier, Pambilar)

to Turbo, on the Gulf of Uraba, N. Colombia.

38. CALOSPIZA RUFICERVIX RUFICERVIX Prév. & Des Murs.

Tanagra ruficervix Prévost & Des Murs, Voyage de la Vénus,

Atlas, Oiseaux, pl. v. fig. 1 (1846—no locality).

T. (Calliste) rufivertex iidem, 1. c., Zool. (text), V, 1, p. 212

* Calliste larvata Dubus, Esquiss. Ornith. pl. 9 (1845(?).—Tabasco, $.K. Mexico).

BIRDS OF WESTERN COLOMBTA. 1109

(1855. —The type stated to be from Guatemala, which is doubtless

a mistake),

Calliste ruficervix Sclater & Salvin, P. Z. 8. 1879, p. 499 (Con-

cordia).

Nos. 2813, 2824. 9 2 ad. Pueblo Rico: 20, 26.x.02.—Wing

72, 70; tail 48; bill 10 mm.

‘“‘ Tris dark brown, feet grey, bill black.”

The Munich Zoological Museum also possesses an adult male

from Rio Lima, 5000 ft., taken in August 1898 by the late

J. H. Batty.

The birds from Western Colombia are in every respect similar

to others from Bogota and Western Heuador in our collection.*

Cr. ruficervix is found only in Colombia (Western Cordillera

and Bogota-coll.) and Western Ecuador (Pallatanga, Cayandeled),

living at rather high altitudes. Im Hastern Hcuador (Machay,

Napo) it is represented by the well-characterized C. r. taylort

Tacz. & Berl. t, while, in Peru and Northern Bolivia, C. r. fulvi-

cervic Scl. & Salv.{ takes its place. Of this last-named

form, the Munich Museum possesses specimens obtained by

Mons. G. A. Baer at Nuevo Loreto, Northern Peru, in June

1900.

39. CALOSPIZA LABRADORIDES Boiss,

Tanagra (Aglaia) labrudorides Boissonneau, Rev. Zool. iti. p, 67

(1840.—Santa-Fe-de- Bogota).

Calliste labradorides Sclater & Salvin, P. Z.8. 1879, p. 499 (Con-

cordia, Santa Elena).

No. 3751. 9 ad. Pueblo Rico: 9.ix.09.— Wing 65; tail 47 ;

bill — mm.

‘“‘ Tris dark brown, feet grey, bill black,”

We have also an adult male procured by J. H. Batty at Rio

Lima, 5000 ft., in August 1898. The birds from the Western

Cordillera agree perfectly with a series of Bogota skins.

C. labradorides is peculiar to the mountains of Colombia.

40. BuTHRAUPIS MELANOCHLAMYS Hellm.

B. melanochlamys Hellmayr, Bull. B. O. C. xxv. no. elxi.

p- 112 (June 1910.— La Selva, W. Colombia),

No. 2796. Q ad. La Selva, Rio Jamaraya, 4600 ft., 15.x.09.

Type of species. No. 10.2378, Coll. Zoological Museum, Munich.

Head all round, back and scapulars deep black, with a faint

silky gloss; lower rump and upper tail-coverts rather dull indigo-

* None of the many specimens I have examined showed any trace of the white

auricular spot or any white on the lesser wing-coverts, characters given by

Dr. Sclater tor his Calliste lewcotis (Contrib. to Ornith. 1851, pt. ii. April, p. 58.—

Kenador).

+ Calliste taylori Taczanowski & Berlepsch, P. Z.S. 1885, p. 75 (1885.—-Machay,

E. Ecuador).

* Calliste fulvicervix Sclater & Salvin, P.Z.S, 1876, p. 354, pl. xxx. fig. 1

(Aug. 1876.—Tilotilo, N. Bolivia.)

Proc, Zoot, Soc.—1911, No. LXAXY. (6)

1110 MR. C. E. HELLMAYR ON THE

blue (Valette’s Code des Couleurs, no, 417); lesser and median

upper wing-coverts uniform bright indigo-blue (im shade between

nos. 411 and 412 of Valette’s Code), forming a large pale blue

shoulder-patch ; greater series dusky, on the outer web washed

with dull bluish; primary coverts, remiges and rectrices blackish,

narrowly fringed with dull bluish along outer margin. Below:

throat and sides of breast deep black, like the back; flanks duller

and inclining to blackish slate-colour; rest of ander parts bright

saffron-yellow, passing into a deeper, more orange tinge on the

middle of the chest. Axillaries pale yellow, under wing-coverts

yellowish white.

Bill uniform black. ‘“ Iris brown, feet black.”

Wing 90; tail 53; bill 154 mm.

Mr. Palmer obtained a single specimen of this distinct species

on the San Juan slopes of the Western Cordillera. It belongs to

that section embracing B. arcei Scl. & Salv.* and B. rothschildi

Berl.t, but differs conspicuously in coloration. The most im-

portant characters of B. melanochlamys are the bright, pale blue

shoulder-patch, altogether absent in both of its allies, and the

deep black colour of the head, mantle, and scapulars, without any

bluish tinge. B. arcwi resembles it in having the breast and belly

bright yellow, but the sides and flanks, instead of being exten-

sively uniform deep black, are merely clouded with dusky, and

the upper parts as well as the throat are strongly glossed with

dark purplish blue. 3B. rothschildi, from the, foot-hills of North-

Western Ecuador, has, like the new species, the head and throat

deep black; however, the back, scapulars, upper wing- and tail-

coverts are conspicuously glossed with purplish blue, while the

under parts, with the exception of an orange-yellow patch on the

foreneck, and the lemon-yellow anal region and under tail-coverts,

are black, with a more or less distinct purple-blue sheen.

The bill is deep black in all three species which eventually

will prove to be geographical representatives. Their range is

singularly restricted :—

(a) B. arcei Sel. & Salv. inhabits the Cordillera del Chucu,

Veragua.

(b) B. melanochlamys Hellm. is found in the Western Cordillera

of Colombia, on the sources of the San Juan River.

Western Ecuador: Cachabi (450 ft.), Rio Pichiyacu (500 ft.), ete.

41. BuTHRAUPIS AUREOCINCTA Hellm.

B. aureocincta Hellmayr, Bull. B.O.C. xxv. no. elxi. p. 111

(June 1910.—Tatama Mountain, W. Colombia).

No. 3487. gad. Tatama Mountain, Chocé, 6700 feet, 8.x.09.

Type of species. No. 10.2377. Coll. Zoological Museum, Munich.

3 ad. Top and sides of the head, and nape deep black, with the

* P.Z.S. 1869, p. 439, pl. xxxi. (Cordillera del Chuct, Veragua).

+ Bull. B. O.C. vii. p. ui (Oct. 1897.—Cachabi, N.W. Ecuador) ; Hartert, Nov.

Zool. v. 1898, p. 482, pl. u. fig. 2.

BIRDS OF WESTERN COLOMBIA. WL

exception of a broad, bright saffron-yellow band which starts from

above the eye and descends the sides of the neck where it joins

the yellow malar stripe. Back bright olive-green ; upper wing-

coverts dull indigo-blue, those of the greater series dusky on the

inner web; primary coverts and remiges black, edged with dull

indigo-blue, the innermost secondaries washed with the same

colour on both webs; rectrices blackish, exteriorly broadly mav-

gined with bright olive-green. Cheeks and malar region bright

saffron-yellow ; large chin-spot pale yellow, some of the feathers

with slight dusky edges; throat and sides of the foreneck black,

the feathers of the former with half-concealed, subterminal spots

of pale yellow; a large, bright golden-yellow patch in the middle

of the foreneck; remainder of under parts olive-green, lighter

and more yellowish than the back; under tail-coverts lemon-

yellow, with the basal half olive-green. Avxillaries dusky, tipped

with greenish ; under wing-coverts greyish white.

“Tris brown, feet dark brown, maxilla black, mandible yellow.”

Wing 93; tail 60; bill 14 mm.

This species, of which Mr. Palmer sent only a single adult bird, is

most nearly related to B. edwardsi Elliot.* The two species agree

in the colour of the back, wings and tail, in having the lbpanets and

abdomen olive-green, with a large, golden- yellow patch on the

foreneck, also in rhe shape and coloration of the bill; but

BL. edwardsi may be distinguished at a glance by having the sides

of the head uniform pale blue, without any yellow or black.

B. aureocincta is known only from the Tatama Mountain, on

the sources of the San Juan River, in the Western Cordillera of

Colombia.

B. edwardsi is an inhabitant of the high mountains of Southern

Colombia and Northern Ecuador. The type, in the Paris Museum.

was brought by M. Triana from an uncertain locality in ‘‘ New

Grenada.” A single specimen was obtained by Prof, Orton at

Chillo, on the western slope of the voleano Antisana, 10,000 ft.

alt.7, Ecuador, and the British Museum received three examples

from Pasto, 8. Colombia ¢, through Mr. Lehmann. Count Ber-

Jepsch possesses _a female picked out from a lot of Quito-skins.

There is, in the United States Museum at Washington +, a skin

said to be from Esmeraldas, N.W. Ecuador, but this locality is,

no doubt, incorrect.

42, CoMPSOCOMA NOTABILIS Jard.

Tanagara (sic) notabilis Jardine, New Edinburgh Philos. Journ.

(n. s.) 1. p. 119 (July 1855.—KHastern Cordillera, Ecuador.

W. Jameson coll.).

Nos. 3785, 3786. $$ ad. Tatamda Mountain, 6700 ft.,

8.x.09.— Wing 99, 100; tail 82, 83; bill 19 mm.

«Tris brown, feet black, maxilla black, mandible blue.”

* Nouv. Arch. Mus. Paris, i. Bull. p. 77, pl. iv. fig. 2 (1865.—“ Nouvelle Grenade , ).

t Salvin, Ibis, 1874, p. 307.

{£ Not Ecuador, as stated in the Cat. B. Brit. Mus. xi. p. 150.

WW MR. C. E. HELLMAYR ON THE

The specimens correspond to the original description, and to

the figure in P. Z.S. 1855, plate xc.

So far as I know, this beautiful species has not been recorded

before from Colombia, and was reported only as an inhabitant of

the highlands of Northern Ecuador.

43, TANAGRA PALMARUM MELANOPTERA Scl,

[Tanagra palmarum Wied, Reise nach Brasil. 11. p. 76 (1821.—

Canavieras, Bahia. |}

T. melanoptera (Hartlaub MS.) Sclater, P. Z. 8. 1856, p. 235

(Jan. 1857.—Kastern Peru, etc.).

T. palmarum (nec Wied) Sclater & Salvin, P.Z.S5. 1879,

p- 500 (Remedios).

T. p. melanoptera Berlepsch, Journ. f. Orn. 1884, p. 291

(Bucaramanga).

Nos. 2499, 2650, 2682. g gad. Ndévita: 281.09; Tadeo:

18.v., 1.vi.09.—Wing 92; tail 70-72; bill 13 mm.

No. 2805. g ad. Loma Hermosa, 4180 ft., 23.x.09.—Wing

94; tail 73; bill 13 mm.

Nos. 2424, 2649, 2716. Qad., Qjr., ¢ juv. Névita: 22.xi1.08;

Tad6: 18.v., 18.vi.09.—Wing 89-90; tail 72, 70, 67; bill 123—

13 mm.

“Tris dark brown, feet grey, bill black.”

The adult birds agree with topotypical skins from Northern

Peru and others from Eastern Ecuador (Napo), Venezuela, Trini-

dad, ete., except that the top of the head is of a paler, duller

yellowish olive. In this respect they resemble a series of Bogota

skins. It must be mentioned, however, that two or three from

Bogota have the eap quite as bright olive-yellow as the Peruvian

ones. More information about the geographical distribution of

these forms is required before any further separation can be

attempted.

T. p. violilavata Berl. & Tacz.,* from Western Ecuador, is very

different from the Colombian birds. It has both the upper and

lower parts much more strongly suffused with violet, and lacks

the olive-yellowish cap, the pileum being bluish like the back.

44, RAMPHOCELUS ICTERONOTUS Bonap.

Ramphocelus icteronotus Bonaparte, P. Z.8, 1837, p.121 (1838,

June.—loc. ign., type in Paris Museum); Cassin, Proc. Acad.

N. Sci. Philad. 1860, p. 141 (Turbo, Rs. Atrato and Truando).

Rhamphocelus icteronotus Sclater & Salvin, P. Z.S. 1879, p. 501

(Remedios, Neche).

Nos. 1969, 2207, 2407. § g ad. Guineo: 5.viii.07; Novita:

17.x11.08; Noanama: 21.x.08.— Wing 85-87 ; tail 72-78; bill 15—

16 mm.

No. 2433. § vixad. Novita: 24.xii.08.—Wing 85; tail 75;

bill 15 mm.

* P.Z.S. 1883, ». 546 (1883.—Chimbo, 8.W. Ecuador).

BIRDS OF WESTERN COLOMBIA. AS

No. 2110. g imm. Sipi: 23.1x.08— Wing 85; tail 77;

bill 16 mm.

Nos. 1956, 2228, 2240. 9 9. San Joaquim, 1.viii.08; Rio

Cajon: 2,5.xi. 08.—Wing 79-82; tail 71-76; bill 15-16 mm.

“Tris red, dark brown (no. 2433), feet blue-grey or blue,

bill blue.”

The Munich Museum also possesses specimens obtained by the

late J. H. Batty on the Rio Guapi, 200-300 feet alt.

R. icteronotus ranges from Veragua to S.W. Ecuador (district

of Guayaquil) *, frequenting the forest-region, from sea-level up

to about 3000 feet.

45, PIRANGA RUBRA RUBRA Linn.

Fringilla rubra Linneus, Syst. Nat. x. p. 181 (1758,— based

on Catesby, Nat. Hist. Carolina ete. i. p. 56, pl. 56; Carolina,

Virginia).

Pyranga estiva Cassin, Proe. Acad. N. Sci. Philad. 1860, p. 140

(Turbo); Sclater & Salvin, P.Z.S. 1879, p. 502 (Concordia,

Frontino).

Nos. 2203, 2213. ¢ g¢ ad. Noanama, 100 ft., 20, 22.x.08.—

Wing 94, 95; tail 74, 76 mm.

No. 2223. gad. Rio Cajon : 30.x.08.—Wing 96; tail 73 mm.

Nos. 2817, 2823, 2830. g¢ gad. Pueblo Rico, 5200 ft., 28, 30.x.,

3.x1.09.— Wing 93, 96, 99; tail 72, 74, 76 mm.

No. 2204. g juv. Noanama: 20.x.08.

Nos.2224, 2251, 2390. 9 9. Novita: 9,11.xii.08; Rio Cajon:

30.x.08.— Wing 92-93 mim.

“ Tris dark brown, feet grey or purplish grey, maxilla yellowish

brown or horn-colour, mandible yellowish brown, yellow, or grey.”

A common winter visitor to South America.

46. PH@NICOTHRAUPIS CRISTATA Lawr.

Phenicothraupis cristata Lawrence, Ann. Lye. N. H. N. Y. x1.

‘p. 70 (Feb. 1875.— Bogota).

Phenicothraupis cristata Sclater & Salvin, P. Z.S. 1879, p. 50

(Frontino, Antioquia); Bangs, Proc. Biol. Soc. Wash. KERSTIN 0

(Naranjito, R. Dagua).

Nos. 3778, —. 6 dad. La Selva, 4600 ft., Oct. ?09.— Wing

97, 98; tail 85, 89; bill 18, 19 mm.

No. — (not numbered). dg ad. Pueblo Rico, 5200 ft.,

Noy. 09.—Wing 98; tail 89: bill 173 mm.

No. — (not numbered). g ad. Loma Hermosa, 4150 ft.,

Oct. 20, 09.—Wing 100; tail 90; bill (damaged) mm.

Nos. 2739, 2829. 9 9 ad. Pueblo Rico: viii, 3.x1.09.—Wing

94,95; tail 86; bill 18, 20 mm.

* Ridgway (Bull. U.S. Mus. no. 50, pt. ii. p. 114) erroneously included “ Central

Peru” in the habitat of this species. Palmal whence Taczanowski (P.Z.S. 1877,

p. 332) recorded a couple, is not in Peru, but in the district of Santa Rosa, province

of Guayaquil, S.W. Ecuador. . icteronotus does not occur anywhere in Peru.

1114 MR. C. E. HELLMAYR ON THE

No. 3752. 2 juv. Pueblo Rico: 13.ix.09.—Wing 88; tail 81;

bill 17 mm.

‘Tris brown (adult), grey (juv.), feet brown, bill black.”

The adult males have the back deep vinous red, the upper

wing-coverts and outer webs of the quills scarcely duller. The

females are smaller; the crest is duller and more of a scarlet red ;

the throat rather lighter red; the back and wings are much

duller, brownish vinaceous ; breast and abdomen decidedly paler

pink, with the greyish bases of the feathers more plainly showing

through, etc.

The young bird has the upper parts dusky brown, with the tail-

coverts and rectrices dull testaceous, the lower surface dingy

cinnamomeous passing into dark brown on throat and foreneck.

There is no trace of the red crest, only a few of the crown-feathers

are narrowly edged with dull reddish.

P. cristata is peculiar to the mountainous districts of Western

Colombia. It is only occasionally met with in Bogota collections.

47. CHLOROTHRAUPIS OLIVACEA Cass.

Orthogonys olivaceus Cassin, Proc. Acad. N. Sci. Philad. 1860,

p- 140 (1860.—* Cordilleras Mountains, on the River Truando,

New Grenada”); Sclater & Salvin, P.Z.S. 1879, p. 502 (Reme-

dios, Neche).

Chlorothraupis olivacea Hartert, Nov. Zool. v. 1898, p. 483

(Cachabi, N.W. Ecuador).

Nos. 2137, 2232: $ g ad. Sipi: 11-x.; Rio Cajon: 2.x1.08.—

Wing 92, 94; tail 633, 65; bill 183, 193 mm.

No. 2379. ¢g imvm. Novita: 9.211.08—Wing 89; tail 63;

bill 183 mm.

Nos. 1962, 2101, 2111, 2123. 9 Gad. & imm. 8. Joaquim:

3.vili. ; Sipi: 21, 23, 26.1x.08.—Wing 85, 86, 86, 90; tail 59-60,

bill 18-20 mm.

“Tris dark brown, feet blue or dark grey, maxilla black,

mandible blue.”

Ndult males are dark brownish green above and have the

feathers of the pileum broadly centred with blackish; the lower

parts are nearly uniform olive-green except the upper throat,

which is light yellow. The females are much smaller, lighter

green above, and more yellowish green below with the whole

throat and middle of foreneck pale yellow; the pileum is uniform

olive-green like the back.

C. olivacea is peculiar to the humid forest-district of Western

Colombia and N.W. Ecuador.

48, HerprosPincus xAnTHopyctus Sel,

Tachyphonus wanthopygius Sclater, P. Z. 8. 1854, p. 158,

pl. lxix. (Apr. 1855—“in Nova Grenada”; descr. 2); idem, le.

1855, pl. xc. (deser. ¢ ; Bogota) ; Cassin, Proc. Acad, N. Sci. Philad.

BIRDS OF WESTERN COLOMBIA. WW

1860, p. 142 (Truando; one ¢ adult); Sclater & Salvin, P. Z. 8.

1879, p. 503 (Remedios).

Nos. 2357, 2709. $ gad. Novita: 1.xii.08; Tado: 16.v1.09.—

Wing 92, 89; tail 68, 65; bill 18 mm.

Nos. 2253, 2358, 2610. 9 @ ad. Novita: 9.xi., 1.11.08;

Condoto : 17.iv.09.—Wing 88-90; tail 65-67; bill 16-18 mm.

“Tris dark red, feet dark grey, bill black.”

These birds are identical with Bogotd-skins in the collection of

the Munich Museum. Like the latter, the males have a broad

scarlet supra-auricular stripe, and the lesser upper wing-coverts

bright lemon-colour,

H. wanthopygius ranges from N.W. Colombia (Truando) south-

wards to S.W. Ecuador, where a specimen was obtained by

Dr. Siemiradzki in the vicinity of Chimbo*. In Panama and

Eastern Costa Rica it is represented by the nearly allied, and

perhaps only subspecifically distinct H. rubrifrons Lawy.t

Although considered by Salvin¢ to be inseparable from //. xan-

thopygius of Colombia, two birds from Veragua and Chiriqui

are so much smaller (wing 78, 80; tail 58 mm.) than any of

the numerous (about fifteen) specimens of the southern form, that

there can be scarcely any doubt as to their distinctness. More-

over, in H. rubrifrons the sexes are alike, the male lacking the

scarlet supra-auricular stripe as well as the yellow patch on the

lesser wing-coverts. Cfr. Ridgway, Bull. U.S. Nat. Mus. no. 50,

pt. 11. pp. 104-105.

49, TACHYPHONUS DELATRII Lafr.

Tachyphonus Delatrit Lafresnaye, Rev. Zool. x. p. 72 (1847.—

Buenaventura, W. Colombia).

T. De Lattrei Cassin, Proc. Acad. N. Sci. Philad. 1860, p. 142

(Falls of the Truando).

T. delattrii Sclater & Salvin, P. Z.S. 1879, p. 503 (Remedios,

Neche).

No. 1959. g ad. San Joaquim (Buenaventura): 1.viii.08.—

Wing 74; tail 64; bill 14 mm.

Nos. 2027, 2064. ¢ gad. Noanama: 28.viii., 4.ix.08.— Wing

73, (2; tail 65, 63; bill 13, 14 mm.

INos. 2222, 2238, 3 oad) Rio Cajon: 30.x., 3.x1.08.— Wing

TA. (38> tall Gos G3 sbullil5. 14s mmo:

Nos. 2094, 2109, 2117, 2139.9 G ad. Sipi: 19, 23, 241x.;

1.x.08.—Wing 71-73; tail 62-64, bill 13-14 mm.

Nos 2090M 2180.) 2Gsam eae ClO 8 sip 18.) BOs ails

13.x.08.— Wing 63-67; tail 55-58; bill 12-13 mm.

“Tris dark red, feet and bill black.”

These examples are topotypical, Lafresnaye’s type having been

* Berlepsch & Taczanowski, P. Z.S. 1883, p. 547: Tachyphonus xanthopygius.

+ Tachyphonus rubrifrons Lawrence, Proc. Acad. N. Sci. Philad. xvii. p. 106

(1865.—Line of Panama Railroad).

+ Ibis, 1870, pp. 109-110: 7. propinguus.

1116 MR. C. EK. HELLMAYR ON THE

secured by Delattre at Buenaventura, in the Bay of Choco.

Birds from Western Ecuador are absolutely similar.

Skins from Costa Rica (Carrillo etc.) are larger, with a longer

and more slender bill; the males are of a duller, more brownish

black, and have the vertical crest somewhat deeper orange.

They are perhaps subspecifically separable. The locality ‘ Pasto’ *

is, no doubt, incorrect, for 7. delatrii is found only in the

lowlands and foot-hills.

50. Mirrosprneus cassinit Lawr.

Tachyphonus Cassinti Lawrence, Ann. Lye. N. H. N.Y. vu.

p- 297 (Jan. 1861.—Panama, Railroad).

Tachyphonus ¢ — Cassin, Proc. Acad. N. Sci. Philad. 1860,

p. 142 (Falls of the Rio Truando).

Eucometis cassini Sclater & Salvin, P.Z.S8. 1879, p. 503 (Neche);

Hartert, Nov. Zool. v. 1898, p. 483 (Cachabi, N.W. Ecuador).

Nos. 2612, 2613. g g ad. Condoto: 20.iv.07.— Wing 86, 84;

tail 77, 74; bill 18 mm.

“Tris grey, feet black, bill black, mandible grey.”

These birds agree, in coloration and size, with others from

Costa Rica. Topotypical Panama specimens are not available for

comparison.

M. cassinii ranges from Costa Rica through Western Colombia

southwards to N.W. Ecuador, whence Mr. Rosenberg’s collectors

have sent many specimens*. It is an inhabitant of the humid

forests of the lowlands and foot-hills, up to about 2000 feet.

51. Hem1rHRaAvPis SALMONI Scl.

Dacnis salmoni Sclater, Cat. B. Brit. Mus. xi. p. 27, pl. i.

fig. 2 (1886.—Remedios, Antioquia; = Q); efr. Hellmayr, Nov.

Zool. xiii. 1906, p. 317 (crit.).

Nemosia rosenbergi Rothschild, Bull. B. O. C. vii. p. vi (Oct.

1897.—Cachabi, N.W. Ecuador; = dad.); Hartert, Nov. Zool.

v. 1898, p. 483, pl. 1. fig. 1.

Not DRG OPUS) BOL WR i gf acl Iowan 9) 75 Zlleaxi..

11.xi1.01.— Wing 6 REA 76 ¢ tail 50-53 ; bill 123-123 mm.

No. 2096. ¢ ad. Near ‘Sipi : 19.ix.08.— Wing 68; tail 50 mm.

No. 2261. ¢ vix ad. Novita: 11.x1.08.—Wing 66; tail 49;

bill 185 mm.

Nos. 2324, 2331. ¢ ¢ juv. Novita: 21, 24.x1.08— Wing 65;

tail 49, 50; bill 123, 133 mm.

Nos. 2323, 2337. 9 9 ad. Novita: 21, 25.x1.08 —Wing 957;

tail 42, 43; bill 11-12 mm.

“ Tris dark brown, feet dark brown (d ad.), grey (dg juv. and 2 );

maxilla black, rmaenible light brown or yellowish pense ad.),

grey (do juv. and Qe

* Cat. B. xi. p. 215.

+ Ridgway (Bull. U.S. Mus. no. 50, pt. ii. p. 169) mentions, as existing in tie

U.S. National Museum, a female from“ Guayaquil, S.W. Eenador, but this locality

requires confirmation.

BIRDS OF WESTERN COLOMBIA. iplele7s

[Mus. Brit. 2 imm. Remedios, Antioquia. Type of D. salmon.

—Wing 58;, tail 42; bill 12 mm.]

The splendid series forwarded by Mz. Palmer leaves no longer

any doubt that Dacnis salmoni is the female of WV. rosenbergi.

The females from Novita agree perfectly with the type in the

British Museum, with which they were kindly compared by my

friend Dr. Hartert. Moreover, they correspond exactly to a

detailed description drawn up by me when in London some years

ago. The type of WV. rosenbergi was obtained at Cachabi, N.W.

Keuador, 450 feet, in November 1896, by Mr. W. F. H. Rosenberg,

and is faithfully depicted on the plate quoted above.

The female may be described as follows :—Above dull yellowish

olive, pileum rather darker, hind crown somewhat shaded with

cinereous; upper wing-coverts like the back; quills dusky,

exteriorly edged with yellowish olive ; rectrices dusky, washed

with pale yellowish olive; sides of the head dull olive; throat

and foreneck dingy buff; remainder of under parts bufty white,

niore purely white along the middle of the abdomen ; under tail-

coverts light buff; axillaries and under wing-coverts white ;

distinct margin along the inner web of the remiges olive-yellow.

Wi ings and tail are much shorter than in the adult male, and the

bill is hkewise shorter, with the lower mandible pale hoa brown

instead of whitish.

The young males (nos. 2324, 2331) in coloration resemble the

female, but are very nearly as large as the adult males. They

show a few orange-yellow feathers on the sides of the head. The

mandible is pale brown as in the females.

In structure, H. salmoni agrees with H. chrysomelas Scl. &

Salv. *, from Southern Central America. The females also

present the same style of coloration, that of H. salmoni being,

however, easily recognizable by having the throat and foreneck

buff (instead of olive- -yellow), the belly buffy white (not pale

yellowish), and the upper parts of a considerably darker tinge.

While H. salmon and H. chrysomelas ave strictly congeneric,

T am a little doubtful whether they can be kept in the genus

Tlemithraupis, for the typical species, H. ruficapilla Vieill., and

its allies have a much shorter, ibaendler, less constricted bill.

H. salmoni is peculiar to the humid forest-districts of Western

Colombia and North-western Ecuador.

52. BUARREMON BRUNNEINUCHA Lafr.

Embernagra brunneinucha Lafresnaye, Rev. Zool. 11. p. 97

(1839.—‘“ Mexico”).

Buarremon brunneinuchus Sclater & Salvin, P.Z.S. 1879,

p- 504 (Concordia, Medellin, Santa Elena).

No. 3743. gad. Pueblo Rico: 9.1x.08.—Wing 81; tail 83;

bill 173 mm.

* Tachyphonus chrysomelas Sclater & Salyin, P. Z.S. 1869, p. 440, pl. xxxii.

(Cordillera del Chuet, Veragua)..

1118 MR. C. E. HELLMAYR ON THE

“Tris and feet dark brown, bill black.”

Identical with specimens from Bogota and Venezuela (Cumbre

de Valencia, Mérida). Typical Mexican birds are not available.

Cfr. Ridgway, Bull. U.S. Mus. no. 50, pt. i. p. 466.

53. ARREMON AURANTIIROSTRIS OCCIDENTALIS, subsp. n.

No. 2557. S$ vixad. Juntas, Rio Tamana (405 ft.), 26.11.09.—

Wing 73; tail 56; bill 153 mm.

No. 2573. gad. Condoto, R. Condoto (150 ft.), 27.11.09.-—

Wing 73; tail 58; bill 144 mm.

‘“‘ Tris dark brown, feet pink, bill bright orange-red.”

Messrs. Salvadori and Festa * have most correctly pointed out

the differences between specimens from Western and Eastern

Eeuador. The large series in the Munich Museum, however,

proves that the birds from the western slopes of the Andes are

clearly distinct from A. erythrorhynchus of Bogota, and constitute

a third, hitherto overlooked race. <A. spectabilis, A. erythro-

rhynchus, and the new form are evidently the southern repre-

sentatives of the Central American A. auraniiirostris Lafr. 7 and

should, according to my view, be ranked as subspecies of the

latter. The northern bird may, however, readily be distinguished

by its much larger, heavier bill, much broader, black jugular band,

and pure olive-green black.

Characters, range and synonymy of the three races are as

follows :—

(a) A. AURANTITROSTRIS SPECTABILIS Scl.

Arremon spectabilis Sclater, P. Z. 8. 1854, p. 114, pl. Ixvi.

(Apr. 1855.—Quijos, Hast Ecuador); idem, P. Z. 5. 1856, p. 82

(Quixos); idem, P. Z. 8S. 1858, p. 72 (Rio Napo, E. Ecuador) ;

idem, Cat. B. Brit. Mus. x1. 1886, p. 275 (part.: e-g, Rio Napo;

h, Sarayacu; p, Eastern Peru); Salvadori & Festa, Boll. Mus. Zool.

Torino, xiv. no. 857, 1899, p. 20 (Rio Santiago and R. Zamora,

E. Ecuador).

A. erythrorhynchus (nec Sclater) Berlepsch & Taczanowski,

P. Z.S. 1883, p. 548 (Huambo, E. Peru; in text of A. spectabilis

no. 49); Taczanowski, P. Z.S8. 1882, p. 16 (Huambo); idem,

Orn. Pérou, ii. 1884, p. 535 (Huambo).

Habitat. Easrern Ecuapor: Quijos (fide Gould), Rio Napo

(fide Verreaux), La Concepcion (Petit), Sarayacu (Buckley),

R. Santiago and R. Zamora (esta); Norra Peru: Huambo

(Stolzmann).

Characters. Back, larger upper wing-coverts, and outer webs of

remiges deep brownish or rufescent olive; bend of the wing

bright orange; bill small and stout. Wing 70-73; tail 58-60;

bill 123-133 mm.

Material. A series from La Concepcion, Rio Napo, East

Keuador, obtained by Louis Petit, in the Munich Museum.

* Boll. Mus. Zool. Torino, xiv. no. 357, 1899, p. 21.

+ Rey. Zool. x. p. 72 (1847.—Panama).

BIRDS OF WESLERN COLOMBIA. 1119

Observations. As already stated by Dr. Sclater and more

recently by Salvadori and Festa, birds from Eastern Ecuador

are easily recognizable by the reddish-orange shoulder-spot and

the dark brownish-olive colour of the upper parts. From the

remarks of Berlepsch and Taczanowski it is evident that the

specimens from Huambo, N. Peru, belong to the same race.

(b) A. AURANTITROSTRIS ERYTHRORHYNCHUS Scl.

Arremon erythrorhynchus Sclater, P. Z.S. 1855, p. 83, pl. lxxxix.

(June 1855—~‘in Nova Grenada, Bogota”); idem, 1c. p. 154

(Bogota) ; idem, ].c. 1858, p. 83 (Bogota).

A. spectabilis Sclater, Cat. B. Brit. Mus. xi. p. 276 (part. a, b:

Bogota).

Habitat. Eastern Corompi4: known only from Bogota

collections.

Characters. Back, larger upper wing-coverts, and outer webs

of remiges clear yellowish olive-green; bend of the wing light

lemon-yellow ; bill small and stout (like the preceding form).

Wing: ¢ 74, 2 70; tail: ¢ 60-64, 2 56-58; bill 13 mm.

Material. A series from Bogota in the Munich Museum.

Observations. Bogota skins are quite uniform in their characters

and always readily distinguishable.

A. erythrorhynchus (nec Sclater) Sclater, P.Z.S. 1860, p. 85

(Nanegal, W. Ecuador); idem, ].c. p. 274 (Babahoyo); idem,

]. c. p. 293 (Esmeraldas); Salvadori & Festa, Boll. Mus. Zool.

Torino, xiv. no. 357, 1899, p. 21 (Rio Peripa, Gualea, W. Ecuador ;

crit., synon.).

A. spectabilis (nec Sclater) Taczanowski, P.Z.8. 1877, p. 332

(Palmal, distr. of Guayaquil, S.W. Ecuador); Sclater & Salvin,

P.Z.S. 1879, p. 505 (Remedios, W. Colombia): Berlepsch &

Taczanowski, P.Z.S. 1883, p. 548 (Chimbo, S.W. Ecuador) ;

Sclater, Cat. B. Brit. Mus. xi. p. 275 (part.: ec, d, Remedios ; i—n,

Sta. Rita, Nanegal, Babahoyo, W. Ecuador); Hartert, Nov. Zool.

v. 1898, p. 483 (Cachabi, Chimbo); Goodfellow, Ibis, 1901,

p. 471 (Santo Domingo, 8. Nicolas); Ménégaux in: Mission

Are Merid. Equat. ix. 1, p. B 81 (S. Domingo, Pachijal).

Habitat. Western Ecuapor: Palmal (Jelski), Babahoyo

(Fraser), Chimbo (Stolzmann, Rosenberg), Santa Rita (Buckley),

Gualea, Rio Peripa (Festa), Santo Domingo (Goodfellow, Rivet),

Nanegal (Fraser), Esmeraldas, (Fraser), Pachijal (Rivet), Cachabi

(Rosenberg). etc. WESTERN CoLompra: Juntas, Condoto, Rio

San Juan, Chocé (Palmer), Remedios, Antioquia (Salmon).

Characters. Colour of back, larger wing-coverts, and outer webs

of quills intermediate between A. a. spectabilis and A. a. erythro-

rhynchus, neither dark brownish olive nor clear yellowish green,

but intense olive-green, somewhat shaded with golden: bend of

the wing lemon- or gamboge-yellow, but never orange; bill

1120 MR. C, E. HELLMAYR ON THE

considerably larger, longer as well as slenderer, than in both of its

allies—Wing : ‘S 702 75, 2 67-70; tail: g 58-63, 2 53-58;

bill 14-153 mm.

Type in the Zoological Museum, Munich, no. 09.5862. ¢ ad.

Condoto, Rio Condoto, W. Colombia, 150 feet, 27 March, 1909.

M. G. Palmer coll. no. 2573.

Material. Two adult males from Western Colombia, eight

adults (of both sexes) and young from Western Hcuador (Quito,

Paramba, San Javier, Lita, Carondelet, etc.).

Observations. I cannot perceive any difference between Co-

lombian and Ecuadorian specimens. The characters indicated

above are perfectly constant in the large series (15) examined

by me.

A. a. occidentalis ranges from near sea-level up to about

3500 feet (Paramba, Rio Mira).

54, PsrrTosPIzA RIEFFERII RIEFFERII Boiss.

Tanagra Riefferit Boissonneau, Rey. Zool. 11. p. 4 (1840.—

Santa-Heé-de- Bogota, Colombia).

Psittospiza riefferi Sclater & Salvin, P. Z.8. 1879, p. 505

(Envigado, Concordia, Medellin, Remedios, Santa Elena).

Nos. 2790, 2791. ¢ Q@ ad. Tatama Mountain, 7600 ft, 9.x.09.

Wing 110, 106; tail 81, 79; bill 18 mm.

‘“‘ tris brown, feet and bill scarlet.”

Agreeing with specimens from Bogota and Western Ecuador,

but bill shghtly larger.

P.r. riefferii is well-known as an inhabitant of the mountains

of Colombia and Keuador. In Peru and Bolivia it is replaced

by P. riefferii elegans Tsch.*, of which we have a good series

from various localities in the Munich Museum.

55. SALTATOR ATRIPENNIS Scl.

Saltator atripennis Sclater, Proc. Acad. N. Sei. Philad. vii.

p. 261 (1856.—Popayan, Colombia); Sclater & Salvin, P. Z.5.

1879, p. 505 (Medellin).

No. 2821. ¢ ad. Pueblo Rico, 5200 ft., 29.x.09.— Wing 108;

tail 102; bill 19 mm.

“Tris dark brown, feet and bill black.”

This bird, which can be considered as typical, agrees perfectly

with Sclater’s original description, having the top of the head black

etc. ‘Two specimens from the neighbourhood of Quito (Kcuador)

in our Museum have a somewhat shorter, stouter bill, and the

crown dark cinereous, merely mottled with blackish on the forehead

and above the white eyebrow. However, they may be not

quite adult.

S. atripennis is known only from the highlands of Colombia

and Heuador.

* Saltator elegans Tschudi, Arch. f. Nature. 10; i. D. 288 (1844.—Peru).

BIRDS OF WESTERN COLOMBIA. 21

56. Savraror MAximus P. L. S. Miill.

Tanagra maxima P. L. 8. Miller, Natursyst. Suppl. p. 159

(1776—ex Daubenton, Pl. Enl. 205: Cayenne).

Saltator magnus Sclater & Salvin, P. Z.S. 1879, p. 505 (Reme-

dios, Medellin, Neche); Berlepsch, Journ. f. Orn. 1884, p. 293

(Bucaramanga).

Nos. 2432, 2434. $9 9 ad. Novita: 24.x11.08—Wing 100,

98; tail 89,91; bill 18, 19 mm.

Nos. 2195, 2021. ¢ 2 ad. Noanama: 27.viii., 17.x.08.—Wing

OS 9a- tal) 92. So) bill) 19 mm:

Nos. 2383, 2047. 9 2 imm. Noéovita: 9.xi1.09; Noanama:

2.1x.08.

“Tris dark brown, feet dark grey, bill black, mandible blue-

grey.”

The majority of these skins have the back decidedly purer and

lighter green than our large series from Eastern South America

(Cayenne, Venezuela, Para, E. Ecuador, etc.), though one or two

are scarcely distinguishable on this score.

57. Piryitus erossus Linn.

Loxia grossa Linneus, Syst. Nat. xu. 1, p. 307 (1766.—

“ America”: ex Brisson; we substitute Cayenne as type locality).

Pitylus grossus Cassin, Proc. Acad, Philad. 1860, p. 140 (Falls

of the River Truando); Sclater & Salvin, P.Z.S. 1879, p. 505

(Remedios, Neche).

Nos. 2114, 2157, 2158. g gad. Sipi: 26.ix

93-97 ; tail 82-85; bill 18-19 mm.

No. Q115. 2. Sipi: 24.ix.08.—Wing 93; tail 83; bill 174 mm.

« Tris dark brown, feet grey, bill bright scarlet.”

Not different from specimens obtained in Venezuela, Brazil,

and Eastern Ecuador.

., 0.x.08.— Wing

58. CACICUS UROPYGIALIS Lafr.

Cassicus uropygiali s Lafresnaye, Rev. Zool. vi. p. 290 (1843.—

** Colombie,” sc. Santa Fe de Bogots) ; Sclater & Salvin, P. Z.S.

1879, p. 509 (Jerico).

C. uropygialis ¢ Cassin, Proc. Acad. N. Sci. Philad. 1860, p. 139

(Falls of the R. Truando).

Nos. 1995, 2009, 2088. ¢ g ad. Mouth of Hate Viviane

Noanama: 25.vill.; near Sipi: 17.1x.08.—Wing 133-138; oil

96-98 ; bill 28-30 mm.

Nos. 1996, 2092. 9 9 ad. Mouth of Calima: 17.viil.; Sipi:

19.ix.08.—Wing 122, 123; tail 90; bill 26, 28 mm.

“Tris light (turquoise) blue, feet black, bill light yellow.”

The series agrees with Bogots skins and others from N.W.

Ecuador. C. macrorhynchus Scl. & Salv.*, from Central America,

* Cassiculus microrhynchus Sclater & Salvin, P. Z.S. 1864, p. 353 (Lion

Hill, Panama).

WARY MR. C. E. HELLMAYR ON THE

is much smaller and has a considerably shorter, slenderer bill, but

is most probably only the northern representative of C. uro-

pygialis. The specimens obtained by Lieut. Michler’s party

on the Falls of the Rio Truando, although referred by

Mr. Ridgway * to C. microrhynchus, are more likely to belong

to the present form (¢/7. Cassin, Proc. Ac. N. Sci. Philad. 1867,

p. 64).

59, AMBLYCERCUS HOLOSERICEUS Licht.

Sturnus holosericeus Lichtenstein, Preis-Verz. Mex. Vog. p. |

(1831.—Mexico) ; Journ. f. Ornith. xi. 1863, p. 55 (veprint).

No. 1982. (2). Guineo, Rio Calima: 8.vii.08.—Wing 92;

tail 86; bill 273 mm.

‘Tris white, feet black, bill light yellow.”

This specimen agrees with others from Western Ecuador, in

size and colour. I have not seen typical Mexican birds, which

may be different.

60. CASsIDIX ORYZIVORA VIOLEA Bangs.

[Oriolus oryzivorus Gmelin, Syst. Nat. 1, 1. p. 386 (1788—ex

Latham : Cayenne) ].

Cassidix oryzivora violea Bangs, Proc. New Engl. Zool. Cl. u.

p- 11 (1900.—La Concepcion, Santa Marta, Colombia).

Cassidix oryzivora (errore) Sclater & Salvin, P. Z.S. 1879,

p. 510 (Concordia, Antioquia, Remedios) ; Sclater, P. ZS. 1860,

p- 140 (Pallatanga, W. Ecuador) ; Salvadori & Festa, Boll. Mus.

Zool. Torino, xiv. no. 357, p. 30 (Rio Peripa, R. Daule,

W. Ecuador); Hartert, Nov. Zool. v. p. 485 (Paramba, Chimbo,

W. Ecuador).

No. 2429. g ad. Noévita, 150 ft., 23.x11.08.—Wing 200;

tail 152; bill 37 mm.

No. 2823. g ad. Pueblo Rico, 5200 ft., 30.x.08—Wing 210;

tail 162; bill 393 mm.

No. 2413. 9 ad. Noévita: 18.x11.08. Wing 153; tail 120;

bill 34 mm.

“Tris light yellow, bill and feet black.”

C. o. violea is an excellent form, easily recognizable by the

strong violet-purple gloss of the plumage. In typical (. o.

oryzivora, from Cayenne, N. Brazil, Venezuela, etc., the general

colour is dull black with bronzy reflection, and very little, if any,

violet admixture on pileum and chest. Specimens from Western

Eeuador aiso belong to C. 0. violea.

61. XANTHORNUS MESOMELAS SALVINII Cass.

[ Psarocoliuse mesomelas Wagner, Isis, 1829, p. 755 (1829.—

Mexico.)|

Teierus Salvinii Cassin, Proc. Acad. N. Sci. Philad. 1867, p. 51

* Bull. U.S. Mas. no. 40, pt. ii. p. 190.

BIRDS OF WESTERN COLOMBIA. 1723

(1867.—Costa Rica, Nicaragua, New Granada (Atrato River,

Bogot), ete.; type from Costa Rica, ¢fr. Stone, tc. vol. li.

1899, p. 34).

I. msomelas Cassin, Proce. Ac. Philad. 1860, p. 140 (River

Atrato) ; Sclater & Salvin, P. ZS. 1879, p. 509 (Neche).

No. 2614. 2 ad. Condoto, R. Condoto: 20.iv.09.—Wing 94 ;

tail 100; bill 20 mm.

No. 1971. S$ juv. Guineo, R. Calima : 5.vii1.08.

“Tris dark brown, feet blue, bill black.”

The adult bird agrees with Bogota skins. The length of the

wing, in the latter, varies from 93 to 96 mm. None of my

Colombian specimens showing any trace of white edging to the

inner secondaries, I consider XY. m. salviniias a well-characterized

race.

In 8.W. Ecuador (district of Guayaquil) and N.W. Peru

(Tumbez, Lechugal, Callacate, etc.) another distinct form is met

with. It has been named _Y. m. taczanowskii Ridgw.*, and may

be recognized by its inferior size (wing 85-88 mm.), lesser extent

of black at the base of the outer rectrices, and by having con-

spicuous yellowish-white edges to the innermost secondaries.

Cfr. Berl. & Tacz. P. Z.S. 1883, p. 552 (s. n. 7. mesomelas).

62. HypopyRRHUS PYROHYPOGASTER Tarragon.

Cassicus pyrohypogaster Tarragon, Rev. Zool. x. p. 252 (1847.—-

“ Nouvelle Grenade ”).

Hypopyrrhus pyrrhogaster Sclater & Salvin, P.Z.S. 1879,

p- 510 (Envigado, Santa Elena).

Nos. 2832, 2833. 9 2 (in moult). Pueblo Rico, 5200 ft.,

4.xi.09.

No. 3768. ¢ ad. Siaté, 5200 ft., 23.ix.09.—Wing 142;

tail 140; bill 31 mm.

Nos. 3758, 3761, 3762, 3770. 29 9. Siaté: 17,18, 23.1x.09.—

Wing 127-130; tail 128-133; bill 27-29 mm.

“Tris light yellow, feet and bill black.”

The specimens agree with a number of Bogota skins in the

Munich Museum. As will be seen from the above measurements

the females have much shorter wings and tail.

H. pyrohypogaster is vestricted to the highlands of Colombia. It

is not uncommonly met with in Bogota collections, and Salmon

procured several examples in Antioquia. |

63. CYANOCORAX AFFINIS AFFINIS Pelz.

Cyanocorax affinis Pelzeln, Sitzungsber. Ak. Wiss. Wien,

math.-naturw. Cl. xx. 1, p. 164 (1856.— Bogota); Sclater &

Salvin, P. Z. 8. 1879, p. 510 (Cauca, Remedios).

* Teterus mesomelas taczanowskii Ridgway, Proc. Wash. Acad. Sci. iii. p. 153

(1901.—Guayaquil, 8. W. Ecuador).

WA Bat MR. C. E. HELUMAYR ON THE

C. pileatus (nec Temminck) Cassin, Proc. Acad. N. Se. Philad.

1860, p. 138 (Rs. Truando and Nercua, N.W. Colombia *).

Nos. 2053, 2459. g¢ 2 ad. Noanama: 3.1x.08, 11.1.09.—

Wing 170, 165 ; ; tail 175, 168; bill 27, 29 mm.

No. 2537, So ‘afl El Tigre, R. Tamana, 13.11.09.— Wing 165 ;

tail 166; bill 29 mm.

No. 3746. g imm. Pueblo Rico, 5200 ft., 10.1x.09.—Wing

170; tail 160; bill 29 mm.

‘Tris lemon-yellow, feet and bill black.”

In the pale yellowish-white colour of the under parts and tail-

end these birds agree with Bogota skins, but are slightly smaller 7

In size they approach C. a. zeledoni, from Chiriqui and Costa me

the latter form may, however , easily be distinguished by the bright

creamy-yellow belly, ete.

A good series from Carthagena (the type locality of C. sclateri

Heine {) is required to show whether the shght difference in size

Hee birds from Bogota and Western Colombia is constant.

C. a. affinis is confined to Colombia, ranging from near sea-

level a | to about 5000 feet altitude.

64. CYANOoLYcA PULCHRA Lawr.

Cyanocitta pulchra Lawrence, Ann. Lye. N. H. N.Y. xi. 1875,

p. 163 (Feb. 1876.—Quito, Ecuador).

No. 2789. gad. Tatama Mountain, 7600 ft., 9.x.08.— Wing

130; tail (Gmeulting) about 135; bill 830 mm.

“Tris, feet, and bill black.”

So far as I am aware this is the second known specimen of

this fine species, which was described by Mr. Lawrence from a

single Quito skin in his collection. The bird corresponds exactly

to the original deseription with the sole exception that the smoky-

blackish chest shows a faint cobalt-bluish tinge. C. pulchra cannot

be confused with any other species. It differs from all the forms

of the C. armillata group by the’ beautiful silvery bluish-

white colour of the crown and nape, terminated behind by a

distinct cross-band of bright ultramarine blue; by the dull

brownish-black upper back, without any blue tinge; by the much

duller cobalt-blue throat-patch; by lacking the black jugular

crescent, so conspicuous a feature in all of its allies; by the smoky-

blackish chest, and much duller bluish belly, etc. Moreover, the

bill is much stouter and larger, while the tail, on the other hand,

is much shorter.

There can be no doubt that C. pulchra is an excellent species,

not much like any other member of the genus.

* Ridgway (Bull. U.S. Mus. no. 50, pt. ili. p. 305), with a query, refers the above

quotation to the Central American race C. a. zeledoni Ridgw. (Auk, xvi. 1899,

p. 255.—Talamanea, Costa Rica). But as specimens from Carthazena, N. Colombia.

agree with C. a. affinis from Bogota ete., those from Truando and Nercua most

certainly belong also to the latter, inasmuch as the fauna of those districts is

essentially the same as that of the San Juan River.

+ Bogoté skins measure: wing 175-180; tail 175-180 mm.

¢ Journ. f. Ormith. vii. p. 114 (1860. — Cartagena (an in Guatemala),”.

BIRDS OF WESTERN COLOMBIA. 1125

65. SAYORNIS CINERACEA CINERACEA Lafr.

Tyrannula cineracea Lafresnaye, Rev. Zool. xi. p. 7 (1848.—

“ Caracas, in Venezuela”).

Sayornis cineracea Sclater & Salvin, P.Z.S8. 1879, p. 511

(Medellin, Frontino).

S. ardosiacus (nec Tyrannula ardosiaca Lafresnaye, 1844)

Cassin, Proc. Acad. N. Sc. Philad. 1860, p. 144 (Falls of the Rio

Truando); efr. Ridgway, Bull. U. 8. Mus. no. 50, pt. iv. pp. 512,

(footnote c), 594 (note a).

No. 2831. @ imm. Pueblo Rico, 5200 ft., 4.xi1.09.— Wing 81 ;

tail 73; bill 145 mm.

“Tris dark brown, feet and bill black.”

Tdentical with examples from Merida and Bogota.

66. Copurus LEUCONOTUS Lafr.

Copurus leuconotus Lafresnaye, Rev. Zool. vy. p. 835 (1842—

“in Bolivia,” errore ; we substitute Bogota, Colombia *); Sclater

& Salvin, P. Z.S. 1879, p. 511 (Remedios).

Nos. 1983, 2054, 2068, 2280, 2346. 5 gad. Gnineo; 10.vi1i.08 ;

Noanama: 3, 5.ix.08 ; Novita: 13, 28.xi.08.—Wing 72-75; tail

170-183 ; bill 8-9 mm.

Nos. 1984, 2345, 2347. 9 9. Guineo: 10.viii.08.; Novita;

27, 28.xi.08.— Wing 70; tail 125-135; bill 8mm.

“ Tris dark brown, feet and bill black.”

The females are smaller, with the median rectrices less elongated,

and the pileum is uniform dark sooty without hoary margins.

There is apparently no difference between specimens from so

widely separated localities as Costa Rica and Colombia, nor do

skins from Western Heuador differ.

67. PLATYTRICCUS MYSTACEUS ALBOGULARIS Scl.

[Platyrhynchos mystaceus Vieillot, Nouv. Dict. xxvul. p. 14

(1818—ex Azara: Paraguay). |

Platyrhynchus albogularis Sclater, P. Z.S. 1860, p. 68 (1860.—

Pallatanga, W. Ecuador).

P. cancroma (nec Temminek) Cassin, Proc. Acad. N. Sci. Philad.

1860, p. 144 (Truando).

No. 2811. g ad. Pueblo Rico: 25.x.09.—Wing 62; tail 36 ;

bill 12 mm.

“Tris dark brown, feet pink, bill black.”

This bird is quite typical. Skins from Chiriqui and Costa

Rica agree in coloration, but are slightly smaller (wing 5/—

59 mm.). I have not seen specimens from Eastern Ecuador 7 and

Northern Peru { which may, or may not, be strictly referable

* “ Bolivia” is an obvious pen-slip for “Colombia,” the paper having as title

“ Oiseaux nouveaux de Colombie.”

+ Tacz. & Berl. P. Z.S. 1885, p. 88 (Machay).

* Taczanowski, P. Z.S. 1879, p. 233 (Tambillo).

Proc. Zoot. Soc.—1911, No. LX XVI. 76

1126 MR. C E. HELLMAYR ON THE

to P. m. albogularis. Birds from Northern Venezuela (San

Esteban, Tocuyo, etc.), however, are very different, being much

paler and more greenish brown above, having the throat tinged

with pale yellowish, the lower mandible whitish instead of

mainly blackish, etc., ete. ‘They agree perfectly with topotypes of

P. insularis Allen* from Tobago, and other examples from

Trinidad and Bermudez (Cumana), while a series secured by the

late H. Whitely on Mount Roraima, Guiana, is intermediate

between P. m. mystaceus (of 8.E. Brazil and Paraguay) and

P.m. insularis. Cfr. Hellmayr, Nov. Zool. xiii. 1906, p. 22.

According to my views, the range of the various races of the

group is as follows :—

(a) P. mystaceus mystaceus Vieill. South-eastern Brazil

from Bahia and Minas southwards to Rio Grande do Sul,

Paraguay, and the adjoining province Misiones, Argentine.

(b) P. mystaceus insularis Allen. Tobago, Trinidad, and North

coast of Venezuela t, from Cumand (Bermudez) west to

Tocuyo. [Specimens from Mt. Roraima, British Guiana,

and from the Orinoco R. (Caicara), may form yet another

race. |

Colombia, Panama, Chiriqui, and parts of Costa Rica.

[East Ecuador and North Peru 2]

68. RHYNCHOCYCLUS CINEREICEPS FLAVoTECTUS Hart.

[Cyclorhynchus cinereiceps Sclater, Ibis, i. p. 443 (1859.—

Oaxaca, Mexico). |

Rhynchocyclus megacephala flavotectus Hartert, Nov. Zool. 1x.

p. 608 (1902.—San Javier, N.W. Ecuador).

Nos. 2250, 2396. 3 g ad. Novita: 8.xi., 12.xii.08.— Wing 63 ;

tail 503, 51; bill 12, 13 mm.

No. 2046. dg imm. Noanama: 2.ix.08.—Wing 62; tail 51;

bill 12 mm.

“Tris brown, feet dark grey, maxilla black, mandible horn-

coloured.”

The specimens agree In every respect with the types from San

Javier and two other examples from Paramba, N.W. Ecuador,

kindly lent by the authorities of the Tring Museum. ‘The species

is for the first time recorded from Colombia. 2. ¢. flavotectus,

although described as a subspecies of 2. megacephala auct. nec

Swains.{, is much more nearly allied to 2. cinereiceps, of Central

America, in fact its southern representative. In the coloration

of the under parts the two birds are perfectly alike; but

R. c. flavotectus may be recognized by its much duller and darkec

* Platyrhynchus insularis Allen, Bull. Amer. Mus. N. Y. ii. p. 143 (1889.—

Tobago).

+ P. albogularis Sclater, Cat. B. xiv. p. 67 (part., specimens d, e: S. Esteban,

Venezuela).

{ Cfr. Hellmayr, Verhandl. zool.-bot. Ges. Wien, liii. 1903, pp. 206-8.

BIRDS OF WESTERN COLOMBIA, WN D7/

green (instead of yellowish-green) back, much brighter yellow edges

to the remiges, and by having the median and greater wing-coverts

broadly edged with bright yellow. As a rule, the cinereous

colour of the pileum is somewhat darker, and the tail on the

average shorter.

Specimens from N.W. Ecuador measure as follows :—

(S)ad. §S. Javier: wing 66; tail 53; bill 12mm.

(2)ad. 8. Javier (type): wing 62; tail 51; bill 12 mm.

Two adults (“¢, 2”), Paramba: wing 62, 63: tail 50, 513;

bill 124, 13 mm.

_ Lat first thought that 2. sulphurescens asemus Bangs * might be

the same as the present species. However, Mr. Bangs gives much

larger dimensions (wing 70, tail 59 mm.) and compares his new

form with R&. sulphurescens, to which R. c. flavotectus is but very

distantly related.

R.c. flavotectus is another characteristic form’of the forest district

of the Pacific Coast, ranging from sea-level up to about 3500

feet (Paramba).

69. CRASPEDOPRION ZQUINOCTIALIS Scl.

Cyclorhynchus ceguinoctials Sclater, P. Z. 8. 1858, p. 70

(1858.—Rio Napo, HE. Keuador).

C. brevirestris (nec Cabanis) Cassin, Proc. Acad, N. Sci. Philad.

1860, p. 144 (Rio Truando).

Rhynchocyclus equinoctialis Hartert, Nov. Zool. v. 1898, p. 487

(Cachabi, N.W. Ecuador).

No. 2422. gad. Novita: 21.xii.08—Wing 78; tail 65; bill

16 mm.

No. 2485. gd ad. Noanama: 16.109.—Wing 80; tail 69;

bill 15 mm.

“Tris dark brown, feet dark blue, maxilla black, mandible

horny-white.”

These skins are practically identical with an adult male from

N.W. Ecuador. All have the greater upper wing-coverts and the

inner secondaries conspicuously edged with ochreous-buff, exactly

like C. olivaceus Temm.,f from Eastern Brazil; they differ,

however, from the last named-species in their larger size, much

larger bill, deeper green throat and chest, etc.

Tam not sure whether they are really referable to C. equinoctialis,

not having examined specimens from Eastern Ecuador (the type

locality). According to Sclater’s original description and later

remarks, the true C. wquinoctialis appears to lack the fulvous

margins to the wing-coverts and inner secondaries.

C. cequinoctialis, as understood here, ranges from Ecuador

north to the Isthmus of Panama.

* Proc. Biol. Soc. Wash. xxiii. p. 73 (1910.—Pavas, Western Colombia, 4400 ft.).

+ Platyrhynchos olivaceus Temminek, Rec. Pl. Col., livr. 11. pl. 12. fig. 1

(1820.—* Brésil ”).

t Cat. B. Brit. Mus. xiv. p. 166.

76*

1128 MR. C. E. HELLMAYR ON THE

70. TopIROSTRUM CINEREUM CINEREUM Linn.

Todus cinereus Linneus, Syst.. Nat. x11. 1, p. 178 (1766—ex

Edwards, Glean. N. H. ii. p. 110, pl. 262. fig. inf.: Surinam).

Todirostrum cinereum Cassin, Proc. Acad. N. Sci. Philad. 1860,

p. 144 (Carthagena); Sclater & Salvin, P.Z.S. 1879, p. 512

(Medellin, Santa Elena, Remedios).

No. 1973. ¢ ad. Guineo, Rio Calima: 6.vii.08— Wing 44;

tail 35; bill 14 mm.

‘“‘ Tris white, feet blue-grey, bill black.”

Perfectly identical with specimens from Bogota, Venezuela

(San Esteban), Cayenne, etc. Chin and throat are yellow lke the’

remaining under parts.

In Western Ecuador this form is represented by Jodirostrum

cinereum sclateri Cab. & Heine*, at once known by having the

chin and upper throat white. It extends southwards into the

district of Tumbez, N.W. Peru.

71. CNIPODECTES SUBBRUNNEUS SUBBRUNNEUS Scl.

Cyclorhynchus subbrunneus Sclater, P. Z. 8. 1860, p. 282

(1860.—Babahoyo, 8.W. Ecuador).

Cnipodectes subbrunneus Sclater & Salvin, P. Z. 8. 1879,

p- 514 (Remedios, Antioquia).

No. 2509. SG ad. Névita: 30.1.09.—Wing 94; tail 87; bill

17 mm. .

No. 2539. g ad. Hl Tigre: 13.11.09—Wing 95; tail 92;

bill 17 mm.

Nos. 2605, 2606. ¢ g ad. Condoto: 16, 16.iv.09.—Wing 95,

92; tail 89, 86; bill 164, 17 mm.

No. 2591. S ad. Condoto: 10.iv.09.—Wing 87; tail 83;

bill 163 mm. :

No. 2569. 3 imm. Névita: 15.1.09.—Wing 75; tail 69; bill

163 mm.

“Tris pink (in adults), brown (in imm. male, no. 2569),

feet grey, maxilla black, mandible pink (adults), horn-coloured

(young).”

This series is most interesting, proving as it does that the so-

called “ minor” from Panama and Western Kcuador are nothing

but the young of the larger swbbrunnews. This had already

been suspected by me when, some years ago, I carefully studied and

measured the entire material in the British Museum, but in the

absence of a good series from any locality, it was then impossible

to arrive at a definite conclusion. The occurrence side by side of

these two ‘‘ species” differing only in size always appeared to

me a phenomenon hardly reconcilable with modern views of

zoogeographical distribution.

No. 2569, a young bird, as manifested by the fluffy texture of

the plumage, differs from the other specimens in its much shorter

* Triccus Sclateri Cabanis & Heine, Mus. Hein. ii. p. 50 (1859.—Peru).

BIRDS OF WESTERN COLOMBIA. 1129

wings and tail, less developed crest, more rufescent brown colour

of the upper parts, and especially in having distinct, ochraceous-

buff apical edges to the greater wing-coverts. An unsexed bird

obtained at Santa Rita, W. Ecuador, by Buckley’s collector

Villagomez, and two skins, one marked “ ¢,” from Panama,

“ McLeannan leg.,” in the British Museum, are also immature.

They were erroneously referred by Dr. Sclater to the Peruvian

race C'. s. minor, the type of which is likewise a young bird !

However, adult males from Chamicuros (the type locality of

C. minor) differ slightly from those of Western Ecuador and

Colombia (Remedios) in having both upper and under surfaces

vather paler brown. An adult male obtained on the Rio Puris,

W. Brazil, shows the same variation when compared with our

Chocd series. Hence the name C. s. minor may well be

retained for the Amazonian race, though its (rather trifling)

characters should be confirmed by a better series of fresh skins.

According to my views there are two very closely allied races :—

(1) 0. s. subbrunneus Scl., ranging from 8.W. Ecuador

through Western Colombia to Panama.

(2) OC. s. minor Scl., ranging from Eastern Peru (Chami-

curos) to W. Brazil (Rio Purts).

Their synonymy and ranges are as follows :—

(a) C. SUBBRUNNEUS SUBBRUNNEUS Scl.

Cyclorhynchus subbrunneus Sclater, P. Z. 8. 1860, p. 282

(1860.—Babahoyo, 8.W. Ecuador) ; idem, 1. c. p. 295 (Hsmeraldas,

N.W. Ecuador); Lawrence, Ann. Lyc. N. H. N.Y. vii. 1862,

p. 473 (Panama).

Myiochanes subbrumneus Sclater, Cat. Amer. B. 1862, p. 232

(Babahoyo, Esmeraldas).

Myiochanes sp., Sclater & Salvin, P. Z.S. 1864, p. 360 (Panama).

Cnipodectes subbrunneus Sclater & Salvin, P. Z.S8. 1879, p. 514

(Remedios, Antioquia); Sclater, Cat. Birds Brit. Mus. xiv.

p. 197, pl. xvi. (part.: Babahoyo, Esmeraldas, Balzar: Ecuador ;

Remedios, Colombia); Salvin & Godman, Biol. Centr.-Amer., Aves,

ii. 1889, p. 55 (part.: Colombia, Panama, Ecuador); Salvadori &

Festa, Boll. Mus. Torino, xiv. no. 362, 1899, p. 10 (Rio Peripa,

W. Ecuador); Ridgway, Bull. U. 8. Mus. no. 50, pt. iv. 1907,

p- 484 (Panama, Colombia, W. Ecuador).

C. minor (nec Sclater) Sclater, Cat. B. Brit. Mus. xiv. p. 197

(part. : d, Santa Rita, W. Ecuador; e,/ Panama); Ridgway, Bull.

U.S. Mus. no. 50, pt. iv. p. 485 (part.: Panama, W. Ecuador).

Type locality. Babahoyo, 8.W. Ecuador.

Range. Western Ecuador: Babahoyo, Esmeraldas (Fraser), Rio

Peripa (Festa), Balzar (Jilingworth), Santa Rita (Villagomez).

Western Colombia: Novita, El Tigre, Condoto, Choco distr.

(Palmer); Remedios, Antioquia (Salmon). Panama: Lion Hill

(MacLeannan), Cascajal Coclée (7eyde).

1130 MR. GC. BE. WELLMAYR ON THE

Material.

Wing. Tail. Bill.

Brit. Mus.—One adult from Babahoyo (type)............. 86; 82; 17° mm.

» Aj One adult from Esmeraldas, Ecuador ... 92; 82; Wee op

55 - One adult from Balzar, Ecuador ............ 853 86; LGin ites

Munich Mus.—Five ¢ g ad. from Choeo, W.Colombia. 87-95; 83-92 ; 165-17

Brit. Mus.—One ¢ ad. from Remedios, W. Colombia. 88; 83 ; 17

3s a One juv. from Sta. Rita, Ecuador ......... 74; 72; 16

Pr Two juv. from Panama ......................... 71,735 65,68; 15, 16

Manic Mus.—One ¢ juy. trom Chocd, ive (Caltrain. 75; 69; Goes

Adults from W. Ecuador and Colombia are practically

identical. From Panama I have seen only young birds.

(6) C. SUBBRUNNEUS MINOR Scl.

Cnipodectes ninor Sclater, P. Z.S. 1883, p. 654 (1884.—

Chamicuros, E. Peru); Taczanowski, Orn. Pérou, ii. 1884,

p. 294 (Chamicuros); Sclater, Cat. B. Brit. Mus. xiv. p. 197

(part.: a—c, Chamicuros); Ridgway, Bull. U.S. Mus. no. 50,

pt. iv. 1907, p. 485 (part.: Eastern Peru).

Cnipodectes subbrunneus minor Suethlage, Journ. f. Orn. 1908,

p- 12 (Rio Purts, Brazil),

C. subbrunneus (nec Sclater) Sclater & Salvin, P. Z.S8. 1873,

p. 281 (Chamicuros); Salvin & Godman, Biol. Centr.-Amer., Aves,

1. 1889, p. 55 (part.: Peru); Berlepsch & Leverkuhn, Ornis vi,

1890, p. 16 (Chamicuros).

Type locality. Chamicuros, Eastern Peru.

Range. Kastern Peru: Chamicuros (Bartlett), W. Brazil:

upper R. Purus (Zuber).

Material.

: Wing. Tail. Bill.

Mus. Brit.—Two adult males from Chamicuros ......... 92; 84,83; 153mm.

» s One ¢ juv. from Chamisuros (type) ...... 72; 64; NWS | sy

Mus. Paré.—One adult male from Rio Purvis ............ 86; q7 Gies

72, SERPOPHAGA CINEREA CANA Bangs.

[Huscarthmus cinereus Styickland, Ann. Mag. Nat. Hist. xiii.

p. 414 (1844.—* Chili”’).]

Serpophaga cinerea cana Bangs, Proce. Biol. Soc. Wash. xvii.

p- 113 (1904.—Chirua, Sierra de Santa Marta, Colombia).

S. cinerea (nec Strickland) Sclater & Salvin, P. Z. 8. 1879,

p. 512 (Envigado, Frontino).

No. 2741. g ad. Pueblo Rico, 5200 ft., Sept. 1909.— Wing 564 ;

tail 47; bill 9 mm.

“Tris dark brown, feet and bill black.”

Identical with specimens from Bogotéand Mérida (Venezuela).

Birds from Southern Peru (Andes of Carabaya) which I take

to be true cinerea are darker grey above, and have the breast

more strongly shaded with cinereous, while those from Costa

Rica and Chiriqui, S. cinerea grisea Lawr.*, are smaller, with the

crown more deeply black.

* Serpophaga grisea Lawrence, Ann. Lyc. N. H. N.Y. x. p. 189 (1871.—San

José, Costa Rica).

BIRDS OF WESTERN COLOMBIA. 1131

73. MIONECTES OLIVACEUS HEDERACEUS Bangs.

[ Mionectes olivaceus Lawrence, Ann. Lye. N. H. N. Y. ix.

p- 111 (1868.—Barranea and Dota, Costa Rica). ]

Mionectes oleagineus hederaceus Bangs, Proc. Biol. Soc. Wash.

xxii. p. 73 (1910.— Pavas, Western Cordillera, Colombia,

4400 ft.).

M. striaticollis (evrore) Selater, P. Z. 8. 1859, p. 144 (Palla-

tanga, W. Ecuador); idem, P. Z. 8. 1860, p. 93 (Nanegal, W.

Ecuador); Berlepsch & ‘Taczanowski, P. Z. S. 1884, p. 296

(Surupata, Chaguarpata, W. Ecuador; jide H. v. Berlepsch

in litt.).

M. olivaceus (errore) Hartert, Nov. Zool. v. 1898, p. 487

(Chimbo); Salvadori & Festa, Boll. Mus. Zool. Torino, xiv.

no. 362, 1899, p. 6 (Niebli, Rio Peripa, W. Ecuador) ; Goodfellow,

Ibis, 1901, p. 704 (San Nicolas, Gualea, Canzacota, W. Ecuador) ;

Hartert, Nov. Zool. ix. 1902, p. 607 (Lita, Paramba, N.W.

Keuador).

Nos. 1960, 1998. og. San Joaquim, Bahia del Chocé,

l.viii.08; Boca de Colima, R. San Juan, 18.viii.08.—Wing 68,

69; tail 54; bill 123, 13 mm.

Nos. 922565 2263, 2322) 6 dads Novita: LO 21.08-—

Wing 67-69; tail 58; bill 12-13 mm.

No. 2252. Q ad. Novita: 9.xi.08—Wing 62; tail 48; bill

12 mm.

“Tris brown, feet light grey, maxilla black, base of mandible

pale brown.”

Palmer’s series as well as several examples from Western

Keuador differ at a glance from true JZ, o. olivaceus, of Costa Rica

and Chiriqui, in having the back much duller, less yellowish

green, the pileum more greyish green, and the olive streaks on

the anterior under parts much darker and more strongly marked.

The Colombian and Venezuelan races, WV. o. galbinus Bangs* and

M. o. venezuelensis Ridgw.t, are much brighter green above, the

pileum being scarcely darker than the back, and the general colour

of the lower surface is much deeper yellow, with the dark streaks

more decidedly green.

M. 0. hederaceus replaces the typical form in the lowlands and

on the Pacific slopes of the Western Cordillera of Colombia and

Ecuador.

Whether its ally Jf. striaticollis D’Orb. & Lafr. also occurs

in Western Colombia ¢ I am unable to ascertain, but it should be

mentioned that the numerous skins I have examined from

Western Ecuador were all referable to J/. 0. hederaceus.

* Mionectes olivaceus galbinus Bangs, Proc. New Eng. Zool. Cl. iii. p. 85

(1902.—La Concepcion, Santa Marta, N. Colombia).

+ M. olivaceus venezuelensis Ridgway, Proc. Biol. Soc. Wash. xix. p. 116

(1906.—Guacharo, State of Cumana, N. Venezuela).

{ M. striaticollis Sclater & Salvin, P. Z.S. 1879, p. 512 (Santa Elena).

: IL. s. poliocephalus Bangs, Proc. Biol. Soc. Wash. xxiii. 1910, p. 74 (San Antonio,

io Cali).

WS MR. C. E. HELLMAYR ON THE

74, LEPTOPOGON SUPERCILIARIS POLIOCEPHALUS Cab. & Heine.

| Leptopogon superciliaris Tschudi, Arch. f. Naturg. 10, i. p. 275

(1844.—Peru, 7. e. “ Vorwiilder” of Central Peru; ¢/7. Tschudi,

Faun. Peru., Aves, p. 162.|

L. poliocephalus Cabanis & Heine, Mus, Hein. ii. p. 55 (1859.—

** New Granada,” sc. Bogota).

L. superciliaris (nee Tschudi) Hartert, Nov. Zool. ix. 1902,

p. 607 (Lita, Paramba, N.W. Heuador).

Yo. 2376. dad. Nodovita, 150 ft., 7.x11.08.— Wing 69 ; tail 62;

bill 15 mm.

“Tris dark brown, feet and bill black.”

A second specimen from this region I have seen in Count

Berlepsch’s collection. It is an adult female, and was obtained

by the late Gustav Hopke at San Pablo, 4500 ft. alt., January 21,

1897. (Wing 65; tail 583; bill 143 mm.)

The West Colombian birds agree with a series from Bogota

and San Esteban, Venezuela, in size and coloration. Specimens

from N.W. Ecuador (Paramba) are also similar.

All these examples differ obviously, however, from ZL. s. swper-

celiaris, of which I have examined a large series from Central and

Southern Peru and Northern Bolivia, in the much deeper yellow

abdomen, and in having the throat and chest strongly tinged with

olive-green. The typical race has the under parts very much

paler, about primrose-yellow, with but a slight greenish wash on

breast and sides.

The colour of the apical spots to the wing-coverts is of no diag-

nostic value, for it varies from primrose-yellow to dull ochreous

in individuals taken at the same place,

While there canbe no question as to the distinctness of Z. s

superciliaris and L. s. poliocephalus, the status of the third

geographical race, ZL. s. transandinus Berl. & Tacz.*, from

South-western Ecuador, is not yet definitely settled. Two females

in Count Berlepsch’s collection are, indeed, much smaller than

LL. s. poliocephalus ; but the dimensions given J. c. for a male are

scarcely inferior to the average measurements of adult males

from Colombia and Venezuela, . So far as coloration is concerned

the two forms are practically identical.

Another uncertainty exists with regard to the Central Ameri-

ean birds, which Ridgway and other American naturalists have

referred to ZL. s. superciliaries of Peru. This, however, can

hardly be correct. Hither they belong to JZ. s. poliocephalus,

ov they may represent yet another race.

75. KLaNIA CINEREA PARAMB& Hellm.

| Elainea cinerea Pelzeln, Zur Orn. Bras. ii. pp. 108, 180 (Sept.

1860.— Marabitanas). |

* P. Z. 8. 1883, p. 553 (1884.—Chimho, S.W. Ecuador).

BIRDS OF WESTERN COLOMBIA. 4133

Serpophaga parambe Hellmayr. Bull. B. O. C. xiv. p. 54 (1904.—

Paramba, N.W. Ecuador).

[Tring Museum. ¢ ad. Paramba, N.W. Ecuador, 3500 ft.

23.vii. 1899. Miketta coll. no. 473, Type of species.— Wing 565 ;

tail 48; bill 10 mm.|

No. 2494. d imm. Névita: 26.1.08.—Wing 534; tail 44 ;

bill 95 mm.

No, 2196. ¢ juv. Noanama: 17.x.08.—Wing 56; tail 49;

bill 10 mm.

“ Tris dark brown, feet and bill black, base of mandible brown.”

On receipt of these specimens it at once occurred to me that

they might represent some phase of S. paramba, originally

described from an adult male in the Tring Museum; and the

careful examination of the typical example not only confirmed

the correctness of my surmise, but, furthermore, clearly showed

that S.parambe is merely a western, smaller subspecies of Hlania

cinerea.

In fact, on comparing the types of the two “species” I find

them identical in structural characters as well as in coloration.

However, /. c. parambe may be recognized by its much shorter

wings and tail, and by its rather slenderer, shorter bill.

The immature males obtained by Mr. Palmer differ from the

type in the following points:—The back is bright olive-green

instead of bluish grey; the tips to the upper wing-coverts and

edges to the secondaries are pale yellow, instead of white, the

edges to the rectrices olive-green, not cinereous. Moreover,

the throat only is white, while the remaining under parts,

including axillaries and under wing-coverts, are pale yellow, with

obsolete, greyish-white flammulations on the chest. |The vertical

patch, in no. 2494, is white with a hardly perceptible yellowish

hue in its posterior portion, the rest of the pileum ashy grey, as in

the adult male, but slightly darker.

The immature birds are not unlike the corresponding stage of

F. c. cinerea*, but can easily be distinguished by having the

pileum ashy-grey (not olive-green like the back), the coronal patch

white (instead of pale yellow), and the under parts much paler

yellow, with the throat conspicuously white. The bill is also

narrower and shorter.

FE. c. parambe: evidently replaces H.c. cinerea on the western

sides of the Andes in Ecuador and Colombia.

For comparison, the dimensions of seven skins of /. c. cinerea.

are herewith given :—

Five adult males +... . Wing 60-63 ; tail 55-57; bill 11-12 mm.

Two immature birds ¢ OE O0 40-03 5 0) ye te mtn

* Described by me in Nov. Zool. xv. 1908, p. 46.

+ (a) Marabitanas, Rio Negro: type of species; (4) Bogoté, Mus. H. v. B.;

FE. Bartlett coll.; (¢) Suapure, Caura, Tring Museum.

t (a) ? jr. Suapure, Caura, Tring Museum; (4) Bogota, Mus. Berlepsch.

1134 MR. C. E. HELLMAYR ON THE

76. Luaarus AL3ICOLLIS ALBICOLLIS Vieill.

Tyrannus albicollis Vieillot, Nouv. Dict. xxxv. p. 89 (1819—

ex Azava: Paraguay).

Nos. 2648, 2658. ¢ g ad. Tad6, 230 ft.: 17, 21.v.03—Wing

82, 81; tail 58, 59; bill 11, 12mm.

Nos. 2654 (not numbered). 2 2 juv. Tado: 18, 19.v.09.—

Wing 78; tail 58, 59; bill 11 mm.

“Tris brown, feet and bill black.”

Not different from Brazilian specimens.

77. MYIoZErETES CAYANENSIS CAYANENSIS Linn.

Muscicapa cayanensis Linneeus, Syst. Nat. xii. 1, p. 327 (1766—

ev Brisson: Cayenne).

Hlenia cayennensis Cassin, Proc. Acad. N. Sci. Philad. 1860,

p. 144 (Turbo).

Myiozetetes texensis (errore) Sclater & Salvin, P.Z.S. 1879,

p-. 513 (Hnvigado, Medellin).

No. 2384. Q ad. Novita: 9.xi11.08.—Wing 83; tail (@ncom-

plete); bill 135 mm.

No. 1992. ¢ jr. Bocade Calima: 4.viii.08.—-Wing 85; tail 73 ;

bill 14 mm.

“Tris dark brown, feet and bill black.”

The adult bird agrees with other specimens from Bogota,

Cayenne, and Pard. About the geographical races of this species

cfr. Hellmayr, Abhandl. Bay. Akad. Wiss. Miinchen, i. Cl. xxi. 3,

1906, pp. 649-650.

M. c. rufipennis Lawr. *, synonymised with the typical form by

Mr. Ridgway‘, is easily recognizable by the greater extent of

rufous on the wings. It inhabits the northern littoral of Vene-

zuela (Valencia, Puerto Cabello, ete.) and is not to be confounded

with MM. c¢. erythroptera Latr.¢ from South-eastern Brazil

(Rio, Southern Minas Geracs), which is much larger and has

still more rufous on the wings.

78. CoryPHOrRICCcUS ALBOyITTATUS Lawr.

Pitangus albovittatus Lawrence, Ibis, iv. p. 11 (1862.—Isthmus

of Panama).

Coryphotriccus albovittatus Ridgway, Bull. U.S. Mus. no. 50,

pt. iv. 1907, p. 669 (Panama).

No. 2421. g ad. Novita, Rio Tamana: 21.x11.08.—Wing 81 ;

tail 69; bill 17 mm.

“Tris brown, feet and bill black.”

C. albovittatus, one of the rarest of Neotropical Tyrants, is

apparently recorded for the first time from South America, it

* Myiozetetes rufipennis Lawrence, Ann. Lyc. N. H. N. Y. ix. p. 267 (1869.—

Valencia, Venezuela).

y7 Bull. U.S. Mus. no 50, pt. iv. p. 446.!

£ Lyrannula erythroptera Latresnaye, Rey. Mag. Zool. (2) y. p. 56 (1853.—Brazil).

BIRDS OF WESTERN COLOMBIA. 1135

having previously been known only as an inhabitant of Teeter

Panama.*

The bird agrees very welll with Ridgway’s description except

for its slightly larger size 7 and the colour of the pileum, which

I should call sooty- -blackish rather than sooty-brown. From

C. parvus Pelz. = it differs in its duller, less greenish back and

white throat G@nstead of yellow like the remaining under parts).

In structural characters the two species are per fectly alike.

So far as I know, the above is the only specimen existing in any

European Museum.

79. MyioB1lus SULPHUREIPYGIUS VILLOSUS Scl.

[Tyrannula sulphureipygia Sclater, P. Z.S. 1856, p. 296 (Jan.

1857.—Cordova, Mexico). |

Myiobius villosus Sclater, P. Z. S. 1860, P 93 (1860.—

Nanegal, W. Ecuador) ; Selater & Salvin, P. 7.8 . 1879, p. 514

(Fr nen).

M. sulphureipygius (errore) Cassin, Proc. Acad. N. Sci. Philad.

1860, p. 144 (Rio Truando).

No. 2427. g ad. Névita: 22.xii.08.—Wing 63; tail 54; bill

11 mm.

‘‘Tris dark brown, feet grey, maxilla black, mandible pink.”

This bird, like others from Western Ecuador, differs from

M. s. aureatus Bangs §, of Chiriqui and Costa Rica (Pozo Azul),

in its darker, more brownish-green back, deeper ochraceous chest

and sides, and more brownish under tail-coverts. The differences

between the two races, though slight, appear to be quite constant.

Mr. Ridgway || makes J/. sulphureipygius a subspecies of MZ. aan-

thopygus | =mastacalis Wied |, but this is certainly wrong.

M. b. mastacalis is merely the southern form of J/. barbatus,

the two being completely connected by intermediates in North-

eastern and Northern Brazil (Piauhy,- Maranhao, Para, Rio

Madeira). The representative of the barbatus group in Western

Kceuador, Colombia, Panama, and Costa Rica is J. 6. atricaudus

Lawr., which is clearly specifically distinct from JZ. s. villosus,

with which it occurs in many localities. Mr. Rosenberg’s cor-

respondents sent numbers of both forms from the same places

in N.W. Ecuador.

* Whether the newly described C. a. distinctus Ridgw. (Proc. Biol. Soc. Wash.

xxi. 1908, p. 191—Rio Reventazén, near Guaydbo, E. Costa Rica) is really distinct

can, of course, only be decided when series trom Panama and Eastern Costa Rica

become available.

+ Ridgway gives for Panama specimens the following measurements : wing, ¢

79s, 2 Wi tails Gas, 2 663 bill) 16; 162, mm:

f£ Pitangus. parvus Pelzeln, Zur Ornith. Bras. ii. pp. 111, 181 (1868.—Marabi-

tanas, Rio Negro).

§ WM. xanthopygus aureatus Bangs, Proc. New Engl. Zool. Cl. iv. p. 27 (1908.—

Chiriqui).

Il Bull. U.S. Mus. no. 50, pt. iv. p. 490. *

* Cfr. Hellmayr, Abhandl. Meat Wiss. Miinchen, ii. Cl., xxii. 3, 1906, pp. 641-3.

1136 MR. C. E. HELLMAYR ON THE

Therefore, the natural affinities of these species are correctly

expressed by the following scheme :—

(a) WM. barbatus barbatus: Guiana and Northern Brazil (Rio

Negro, Borba, Para, etc.).

(b) I. barbatus atricaudus; Western Ecuador and Colombia,

Panama, Costa Rica.

west to the Rio Madeira (cfr. Nov. Zool. xvii. 1910,

p- 299).

(d) MW. sulphureipygius sulphureipygius : Southern Mexico ;

2 Guatemala, British Honduras.

(ce) M. sulphureipygius aureatus: From Honduras south to

Panama.

(f) M. sulphureipygius villosus : Western Colombia and Ecua-

dor; ? Peru, N. Bolivia.

80. MyIoBIUs ERYTHRURUS FULYIGULARIS Saly. & Godm.

[Myiobius erythrurus Cabanis, Arch. f. Naturg. 13) ap 249°

pl. 5. fig. 1 (1847.—Guiana, Cayenne). |

M. fulvigularis Salvin & Godman, Biol. Centr.-Amer., Aves,

ii. p. 58 (1889.—Santa Fe, Veragua).

No. 2554. 9 ad. Juntas, Rio Tamana, 23.11.09.—Wing 48 ;

tail 36; bill 84 mm.

“Tris brown, feet yellow, maxilla black, mandible pink.”

This is an extreme example of I. e. fulvigularis, agreeing with

skins from Costa Rica. ‘The forehead is strongly washed with

buff, the under parts are bright buffy ochreous, the throat hghter,

more tawny, etc.

Ofr. my remarks in Nov. Zool. xiv. 1907, p. 48.

81. EMPIDONAX VIRESCENS Vieill.

Platyrhynchos virescens Vieillot, Nouv. Dict. xxvii. p. 22

(1818—based on Muscicapa querula (nee Vieillot) Wilson, Amer.

Orn. ii. 1810, p. 77, pl. 13. fig. 2—Eastern North America).

Empidonax acadicus auct. (nec Gmelin).

No. 2826. (¢)ad. Pueblo Rico: 5200 ft., 2.xi.09.—Wing 74 ;

tail 62; bill 13 mim.

No. 2174. @ ad. Sipi: 9.x.08.—Wing 70; tail 59; bill

12 mm.

“ Tris brown, feet dark grey, maxilla black, mandible yellow.”

A common winter visitor to the Western States of South

America. It has several times been taken in Hcuador.

82. MyI0CHANES RICHARDSONII RICHARDSONII Swains.

Tyrannula richardsonit Swainson, Fauna Bor.-Amer. 11. p. 146,

pl. 46, fig.inf. (1831.—Cumberland House, Saskatchewan, Canada).

No. —. @ ad. Loma Hermosa, 4150 ft.: 22.x.09.— Wing 80;

tail ‘62 ; bill 12 mm.

BIRDS OF WESTERN COLOMBIA. WN S7

“Tris dark brown, feet and bill black.”

This species is also a regular winter visitor to Western South

America, The specimen sent by Mr. Palmer agrees with others

from Costa Rica (San José), Quito, etc.

83. MyzArcuus crinitus Linn.

Lurdus crinitus Linneus, Syst. Nat. x. p. 170 (1758—ex

Catesby, Carolina, i. p. 52, pl. 52: Carolina).

Myiarchus crivitus Berlepsch, Journ. f. Ornith. 1884, p. 303

(Bucaramanga).

Nos. 2409, 2420. 6 ¢ ad. Novita: 17, 21.x11.08.— Wing 101,

107 ; tail 92, 96; bill 21, 22 mm.

No. 2394. 9 (2) ad. Novita: 12.xi1.08.— Wing 1062 ; tail 98 ;

bill 21 mm.

No. 2469. g¢ ad. Noanama: 13.1.09.—Wing 110; tail 102;

bill 22 mm.

“Tris dark brown, feet black, maxilla black, mandible horn-

brown.”

These birds are absolutely identical with skins from the Eastern

United States. From MW. tyrannulus (P. L. 8. Mull.) they differ

in being decidedly greenish instead of greyish brown above, in

the darker cinereous throat and chest, much deeper yellow belly,

and in having the whole of the inner web of the outer rectrices

rufous.

The present record extends the range of MM. crinilus in its

winter quarters considerably to the south. The most southerly

locality yet known was Bucaramanga, whence Lorentz had sent

a specimen to the Bremen Museum (cfr. Berlepsch, 1. c. p. 303).

84. MyYIARCHUS NIGRICEPS Scl.

Myiarchus nigriceps Sclater, P. Z.S. 1860, p. 68 (1860.

tanga, W. Ecuador).

No. 2809.(¢)ad. Pueblo Rico, 5200 ft.: 25.x.09.—Wing 82 ;

tail 75; bill 18 mm.

No. 2208. ¢ imm. Noanama: 21.x.08.—Wing 77; tail 73;

bill 18 mm.

“Tris dark brown, feet and bill black.”

The two skins agree with topotypical examples from Western

Keuador. The pileum is uniform deep black.

Mr. Ridgway, in his great work *, has again confounded two

distinct species under the name J/. nigriceps, overlooking my

remarks in Nov. Zool. xiii. 1906, p. 23 & pp. 323-4, I think I

have conclusively shown that the birds from Venezuela, Amazonia,

and Guiana belong to J. tuberculifer, while M. nigriceps is

restricted to Panama, W. Colombia, W. Ecuador, and Northern

Peru.

The range of the two species is given 7m eatenso in Noy. Zool.

xiii. p. 26.

Palla-

* Bull. U.S. Mus, no. 50, pt. iv. p. 650.

1138 MR. C. E. HELLMAYR ON THE

85. TYRANNUS MELANCHOLICUS SATRAPA Cab. & Heine.

[Tyrannus melancholicus Vieillot, Nouv. Dict. xxxv. p. 84

(1819.—ex Azara: Paraguay). |

Laphyctes satrapa Cabanis & Heine, Mus. Hein. ii. p. 77

(1859.—* Guiana and Caracas”).

Tyrannus melancholicus (nec Vieillot) Sclater & Salvin, P. Z.S.

1879, p. 516 (Retiro, Medellin) ; Cassin, Proc. Acad. N. Sci.

Philad. 1860, p. 143 (Turbo, Carthagena, R. Truando).

No. 2712. g ad. Tadé6, 230 ft., 17.vi.09.—Wing 116; tail

101; bill 21 mm.

“ Tris dark brown, feet and bill black.”

85. CHLOROPIPO HOLOCHLORA Lira Hellm.

[Chloropipo holochlora Sclater, Cat. B. Brit. Mus. xiv. p. 287

(1888.— Bogoti). |

Chloropipo holochlora lite Hellmayr, Nov. Zool. xiii. p. 325

(1906.—Lita, N.W. Ecuador).

CO. holochlora (nec Sclater) Sclater, Cat. B. Brit. Mus. xiv.

p- 287 (part.: specimen 6, “ Pasto,” Colombia); Hartert, Nov.

Zool. v. 1898, p. 488 (Cachabi, Paramba, N.W. Ecuador).

No. 2081. g ad. Sipi: 12.x.08.—Wing 69; tail 44; bill

103 mm.

No. 2352. (3) ad. Novita: 30.%1.08:—Wing 71; tail 46;

bill 113 mm.

No. 2112. 9 ad. Sipi: 23.1x.08.—Wing 663; tail 43; bill

11 mm.

“Iris brown, feet grey or blackish, maxilla black, mandible

grey.”

Identical with topotypical skins from North-western Eeuador.

The upper parts are light olivaceous green, the throat, chest, and

sides somewhat duller. On the eastern (Amazonian) slope of

the Colombian and Ecuadorian Andes this form is replaced by

typical C. h. holochlora.

The locality ‘ Pasto” requires confirmation. The specimen is

typical of C. h. lite.

O. h. lite is restricted to the lowlands and foot-hills of

W. Colombia and N.W. Ecuador.

87. MAsIUS CHRYSOPTERUS BELLUS Hart. & Hellm.

[Pipra chrysoptera Lafresnaye, Rev. Zool. vi. p. 97 (1843.—

Bogota). |

Masius chrysopterus bellus Hartert & MHellmayr, Orn.

Monatsber. xi. p. 35 (1903.—Riolima, Cauca Valley, 4000 ft.).

M. coronulatus (nec Sclater) Sclater & Salvin, P.Z.S. 1879,

pp- 516, 549 (Antioquia ; no exact locality); Pelzeln & Madarasz,

Monogr. Pipr. pl. v. (figure of the Antioquia specimen).

No.—. ¢ ad. Pueblo Rico, 5200 ft., 28.x.08.— Wing 59; tail

43; bill 7 mm.

BIRDS OF WESTERN COLOMBIA. 1139

“Tiis brown, feet light red, maxilla light grey, mandible pale

red.”

This specimen confirms the characters of the subspecies origin-

ally based upon a male obtained by the late J. H. Batty in the

same region. ‘The feathers of the coronal crest, which have

exactly the same shape as in Jf. ¢. coronulatus of Western

Eeuador, being thickened at the end into a horny substance,

are dark brownish red instead of light tobacco-brown.

M.c. bellus is well figured in Pelzeln and Madarasz’s Monograph

of the Pipride, on plate v. In their joint communication *

Hartert and Hellmayr have pointed out that Salmon’s Antioquia

specimen belongs to this form, calling attention to a misleading

statement in Sclater and Salvin’s report in P. Z.8. 1879. AV. .

bellus is as yet known only from the Western Cordillera of

Colombia.

88. PIPRA MENTALIS MINOR Hart.

[Pipra mentalis Sclater, P. Z.S. 1856, p. 299, pl. 121 (1857.

— Cordova, Vera Cruz, Mexico). |

Pipra mentalis minor Hartert, Nov. Zool. v. p. 489 (1898.—

Cachabi, N.W. Ecuador). |

No, 2225. g ad. Rio Cajén: 31.x.08.—Wing 60; tail 29;

bill 8 mm.

No. 2098. d ad. Sipi: 21.1x.08.—Wing 60; tail 29; bill

85 mm.

Nos. 2002, 2201. ¢ g ad. Noanama: 22.viii, 19.x.08.—Wing

58, 57; tail 28, 29; bill 8, 8 mm.

Nos; 220%, 2308, 23805 2371. 3g Sg ad. Novita: 185 Oia;

5, 9.xi1.08.—Wing 57-58 ; tail 28-29; bill 8 mm.

Nos. 2030, 2192, 2430. g S juv. (plumage of female).

Noanama: 27.vill.; Sipi: 14.x.; Novita: 23.x11.08—Wing 60;

tail 29-303 ; bill 84-9 mm.

Nos. 2172, 2226.2 Qad. Sipi: 9.x.08; RioCajon: 31.x.08.—

Wing 59; tail 28, 303; bill 83-9 mm.

“Tris white (in adult male), dark brown (in females and young

males), feet light brown or grey, maxilla dark brown, mandible

pale brown or pink.”

The adult males agree, so far as coloration is concerned, with

typical birds from Northern Ecuador. The head is of the same

intense crimson hue, the gonydeal angle is but narrowly edged

with yellowish, and the imner secondaries only have slight

yellowish-white margins along the inner web. While the bill is

as small and short as in Ecuador specimens, in the length of the

wing and tail the Colombian birds approach P. m. ignifera,

from Chiriqui and Costa Rica. The difference in size, however,

is very slight and not constant enough to warrant their separation

from P. m. minor. Ten adult males from Western Ecuador

* Ornithologische Monatsberichte, xi. pp. 33-25.

1140 MR. C. E. HELLMAYR ON THE

(including type) measure as follows :—wing 55 (twice) to 58; tail

26-28 ; bill 8-9 mm.

P.m. minor thus ranges from Ecuador northwards over the

Pacific lowlands of Western Colombia.

89. PrpRA CORONATA VELUTINA Berl.

| Pipra coronata Spix, Av. Bras. 11. p. 5, pl. vii. fig. 1 (1825.—

S. Paulo d’Olivenca). |

Pipra velutina Berlepsch, Ibis, 1883, p. 492 (1883.—Veragua).

P. cyaneocapilla (nec Hahn) Sclater & Salvin, P.Z.S, 1879,

p- 517 (Medellin, Remedios, R. Neche, Antioquia).

Nos. 2301, 2316, 2340, 2365, 2401. ¢ g ad. Névita: 17, 20,

26.x1., 3, 15.xn-08.

Nos. 2118, 2105, 2146, 2156, 2182, 2241. g g ad. Sipi: 22,

Mae sscegy Ila he 8). 14 oe ONS)

No. 2030. ¢ ad. Noanama: 26.x1.08.

No. 2239. g ad. Rio Cajon: 4.xi.08.

Nos. 2031, 2040, 2058, 2216. g g juv. Noanama: 28, 31.viii.,

Ae OO.

No. 1964. g juv. 8S. Joaquim: 4.viii.08. No. 2160. 3 juv.

Sipi: 6.x.08.

Nos. 2311, 2334, 2361, 2362, 2366. 9 9 ad. Néovita: 19

24.x1., 1, 3.x11.08.

Nos. 2121), 2124, 2173. @ 2 ad. (Sipi: 25, 28.1x, 9.x.08:

No. 1965. 9 ad. 8S. Joaquim: 4.v11.08.

“Tris dark red (in adult males), dark brown (in females and

young males), feet black, maxilla black, mandible blue.”

3d ad.: wing 554-58; tail 24-26; bill 7-8 mm.

@ 2 ad.: wing. 55-58; tail 25-28; bill 74-85 mm.

In ‘The Ibis’ for 1906, pp. 31-32, I have already alluded to

the smaller size of specimens from N.W. Ecuador and Western

Colombia. All of the many skins procured by Mr. Palmer have

much smaller bills and decidedly shorter wings than a series from

Chiriqui and Costa Rica. In the adult males the black of the

plumage is also somewhat deeper. Although I feel pretty sure

that the southern birds constitute a separable race, yet I should

like to examine a series from Veragua, the type locality, before

making any formal separation.

Specimens from different localities measure as follows :—

Males. Wing. Tail. Bill.

mm. mm mm.

Oe nt GUN I acs cee oe aa staiee 62-64 27-29 9-92

ALSO NW 5. CORE) ARO incoodounsgdooAes 62-64 27-28 9-93

3: Panama (Railway) ....:...... 58-59 26-27 1-74

N4bo WY, Gollommlon® soacascscceacsoscc 509-08 243-26 7-8

TS INN 5 WGC GoaShoaeanaddne 5d-57 =. 25-27 = 74-8

Females.

4: Chiriqui and Costa Rica...... 614-63 28-30 93-10

LO Wis Colom nia i eeerereeeneeeetee 544-57 25-27 74-8

SING IBCURNTOIE: Luscusscendsae HG BYE 7-8

BIRDS OF WESTERN COLOMBIA. 114)

90. CHIROMACHERIS VITELLINA Gould.

Pipra vitellina Gould, P. Z. 8. 1843, p. 103 (Dec. 1843,—

Panama).

Chiromacheris vitellina Sclater & Salvin, P..Z.8. 1879, p. 516

(Cauca, Remedios).

Nos. 2025, 2097, 2364, 2494. ¢ g¢ ad. Noanama: 28.viii;

Sipi: 2l.ix; Novita: 3.xii.08, 27.1.09.— Wing 53-54; tail 28-30 ;

bill 10 mm.

Nos. 2067, 2185. 3 gjuv. Noanama: 5.1x.08; Sipi: 13.x.08.

Nos. 2352, 2439. 9 9 ad. Noéovita: 30.x1., 28.x1.08.— Wing

54-55 = taal 29; bill 10 mm.

“Tris dark brown, feet orange, bill black ”

A single male from Panama is rather more yellowish green on

the belly, but does not differ otherwise. C. awrantiaca Salv. is

apparently the geographical representative of C. vitellina in

Veragua and 8.W. Costa Rica.

C. vitellina ranges from the Isthmus of Panama south to

W. Colombia.

91. SapAYoA =NIGMA Hart.

Sapayoa enigma Hartert, Nov. Zool. x. p. 117 (1903.—Rio

Sapayo, N.W. Ecuador); Hellmayr in Wytsman’s Genera Avium,

part ix. 1910, p. 28, pl. 1. figs. 5, 8.

No. 2418. gad. Noévita: 19.xi1.08—Wing 82; tail 66;

bill 15 mm.

No. 2070. g juv. Noanama: 1.1x.08.—Wing 81; tail 61;

bill 135 mm.

Nos. 2071, 2457. 9 imm., 9 ad. Noanama: 5.ix.08, 9.1.09.—

Wing 82, 79; tail 59; bill 15 mm.

“Tris brown, feet black, bill black, mandible grey in adult male

and females, yellow in the young male” (no. 2070).

This curious bird was hitherto represented only by a single

female, the type, from N.W. Keuador, in the Tring Museum.

Tt has been fully described by Hartert, and I have nothing to add

to his excellent account. Sapayoa is certainly related to

Scotothorus, though the shape of the bill, the feathering of the

upper portion of the metatarsus ete., serve to distinguish it at a

glance. ;

The females agree exactly with the type, while the adult male

differs in having the middle of the crown bright golden yellow.

The tail is longer, with the rectrices decidedly narrower, and

the central pair slightly elongated, exceeding the submedian one

by about 3mm. The immature female has the upper part of

the head more yellowish green, and the throat slightly paler

yellowish than the adults. The young male (without any trace of

the golden yellow crown-patch) has a much shorter, smaller bill,

with the lower mandible yellowish white instead of brownish

horn-colour. The range of S. enigma is restricted to the humid

lowlands of the Pacific coast of N.W. Ecuador and W. Colombia.

Proc. Zoo, Soc.—1911, No. LX XVII. ca

1142 MR, C. E. HELLMAYR ON THE

92. 'TITYRA SEMIFASCIATA COLUMBIANA Ridgw.

| Pachyrhynchus semifasciatus Spix, Av. Bras. 11. p. 32, pl. xliv.

fig. 2 (1825.—Para). |

Tityra semifasciata columbiana Ridgway, Proc. Biol. Soc.

Wash. xix. p. 119 (1906.—La Concepcion, Santa Marta).

T. personata (nec Jardine & Selby) Sclater & Salvin, P. Z. 8.

1879, p. 517 (Remedios, Neche).

No. 2502. g ad. Noévita: 29.1.09—Wing 122; tail 77;

bill 24 mm.

No. 2314. 9 ad. Névita: 19.x1.08—Wing 114; tail 69;

bill 24 mm.

“Tris red-brown (¢ ), dark-brown (Q ), feet dark grey, bill red,

tip black.”

As correctly pointed out by Mr. Ridgway, the adult male differs

from typical 7’. semifasciata of Amazonia * by the much

narrower black frontal area, the posterior margin of which ends

in a line with the middle of the eye, instead of extending as far as

its posterior edge. Moreover, the black band across the inner

web of the outer rectrices is much broader, and reaches nearly or

quite to the shaft. The female is much browner on the upper

parts, with the pileum considerably darker.

By the coloration of the female, and the greater extent of

black on the lateral rectrices, this form approaches 7’. s. costaricensis

Ridew. + from Panama and Costa Rica, which, however, is

smaller and has even more black in the tail.

T. s. columbiana ranges from Santa Marta through Western

Colombia to the province of Hsmeraldas, N.W. Ecuador. Speci-

mens from the latter locality agree in every respect with the

Colombian ones.

93, TrryRA ALBITORQUES ALBITORQUES Dubus.

Vityra albitorques Dubus, Bull. Acad. Roy. Belg. xiv. pt. 11.

p. 104 (1847.—Peru); Sclater & Salvin, P. Z. 8. 1879, p. 517

(Remedios).

Nos. 2278, 2452. gg ad. Novita: 13.xi1.08; Noanama:

7.1.08.—Wing 107, 105; tail 67; bill 214, 225 mm.

No. 2525. S ad. El Tigre, R. Tamana: 10.11.09.— Wing 103 ;

tail 64; bill 22 mm.

No. 2524. 9 ad. El Tigre: 9.11.09.—Wing 100; tail 64;

bill 22 mm.

‘“‘ Tris dark brown, feet grey, bill black.”

The males do not appear to be different from those obtained

by Natterer at Manaos, N. Brazil, or from an adult male from

* Mr. Ridgway (Bull. U. S. Mus. no. 50, pt. iv. p. 868) gives as its range

“*Southern Brazil and Bolivia to Central Colombia.” ‘This is not quite correct,

for T. s. semifasciata is strictly an Amazonian species, ranging from Paré west to

the Eastern slopes of the Andes, south to H. Bolivia and W. Mattogrosso.

+ Proc. Biol. Soe, Wash. xix. p, 119 (1906,—Bonilla, Costa ica).

BIRDS OF WESTERN COLOMBIA. iat;

Chyavetas, N. Peru. The back, upper wing- and tail-coverts are

pale pearl-grey, the basal half of the rectrices greyish white, the

remaining portion black with a distinct white apical margin, the

ear-coverts white, etc., exactly as in the skins from Manaos and

Peru.

T. buckleyi Salv. & Godm.* from Eastern Ecuador, the type

of which I have examined in the British Museum, may be

distinguished by the paler, nearly white colour of the upper wing-

coverts and rump, by having the white in the tail confined

to the extreme base of the outer web of the external rectrices,

etc. ete.

94, PLATYPSARIS HOMOCHROUS Sel.

Pachyrhamphus homochrous Sclater, P. Z. S. 1859, p. 142

(1859.—Pallatanga, W. Ecuador).

Hadrostomus homochrous Sclater & Salvin, P. Z. 8. 1879, p. 517

(Remedios).

Nos. 2260, 2600, 2685. ¢ ¢ ad. Névita: 10.xi.; Condoto :

14.iv.09; Tadd: 2.vi.09.—Wing 90; tail 63-65; bill 16—-

17 mm.

No. 2415. $ juv. Novita: 19.x1.08.— Wing 86; tail 61;

bill 17 mm.

No. 2399. 9 ad. Novita: 17.xi1.08.—Wing 85; tail 61;

bill 17 mm.

“Tris dark brown, feet blue-grey, bill black, mandible grey in

the young male and female.”

Identical with skins from N.W. Ecuador, province of

Esmeraldas.

This species ranges from Panama (Railroad) and Santa Marta

in the north through Western Colombia and Western Hcuador

to N.W. Peru (Lechugal, district of Tumbez) 7.

95, PACHYRHAMPHUS DORSALIS Sel,

Pachyrhamphus dorsalis Selater, Cat. Amer. B. p. 245 (1862.—

““ Bogota ?” t); Ridgway, Bull. U. 8. Mus. no. 50, pt. iv. 1907,

p. 832 (Rio Cali, R. Barratoro, Castilla, W. Colombia); Bangs,

Proc. Biol. Soc. Wash. xxi. 1908, p. 157 (“ N.W. Colombia,

just south of Darien,” errore; see footnote above p. 1085),

P. cinerewentris (nec Sclater) Sclater & Salvin, P. Z. 8. 1879,

p. 518 (Santa Elena).

No. 3750. @. Pueblo Rico, 5200 ft., 11.ix.09.—Wing 75;

tail 57; bill 133 mm.,

‘“‘ Tris brown, feet dark grey, bill black.”

* Biol.-Centr, Amer., Aves, ii. p. 120 (Dec. 1890.—Yanayacu, Ecuador).

+ The locality “Sarayacu,” Hast Ecuador, is undoubtedly incorrect. I have, on

several occasions, alluded to the fact that Buckley’s localities cannot be relied upon,

collections from the western and eastern slopes of the Andes having been mixed

up and almost invariably labelled “Sarayacu.”

{ I have examined the type in the British Museum. [It is an undoubted Bogoté

skin, i

77*

1144 MR. C. E. HELLMAYR ON THE

I quite agree with Mr. Ridgway in considering this species

totally different from P. niger cinereiventris Scl., with which

Dr. Sclater later united it. In fact, P. dorsalis appears to be

the Western representative of P. marginatus Licht.* It may,

however, readily be distinguished in the male sex by the uniform

black upper back, separated from the glossy black pileum by a

very broad, light cinereous nape-band. The wings and tail are

longer, and the bill decidedly larger.

The female, as yet undescribed, is very much like that of

P. marginatus, but has a larger, broader bill, and the pilewm is

dark olive, each feather with an apical spot of dull greenish

black.

In addition to the type and another specimen in the British

Museum, I have examined four more adult males from Bogotd in

the collections of the Munich Museum and of Count Berlepsch.

The Tring Museum possesses specimens from Paramba, N.W.

Ecuador, which are perfectly similar to Bogota skins.

P. dorsalis thus ranges from Panama through Western

Colombia to N.W. Ecuador.

96. PACHYRHAMPHUS CINNAMOMEUS Lawr.

Pachyramphus cinnamomeus Lawrence, Ann. Lyc. N. H. N.Y,

vil. p. 295 (Jan. 1861.—Lion Hill Station, Panama. Railway).

Pachyrhamphus cinnamomeus Sclater & Salvin, P. Z. S$. 1879,

p. 518 (Remedios).

P. rufescens® Cassin, Proc. Acad. N. Sci. Philad. 1860, p. 189

(Turbo). ;

Nos. 1980, 2535. ¢ gd ad. Guineo: 8.vi1.08; El Tigre, 320 ft. :

12.11.09.—Wing 78, 79; tail 57, 55; bill 144 mm.

Nos. 1968, 2034. ¢ ¢ imm. (without the rudimentary second

primary). 8. Joaquim, Bahia del Choco, 4.viii; Noanama:

29.viii.08.— Wing 75, 79: tail 57; bill 14 mm.

Nos. 2026, 2350. 992 ad. Noanama: 28.vii.; Ndévita:

28.xi.08.— Wing 75, 76; tail 55, 58; bill 14 mm.

“Tris dark brown, feet blue-grey, maxilla black, mandible

blue.”

The series is exactly like specimens from Western Ecuador,

while skins from Central America (Costa Rica, Guatemala), as

a rule, are of a deeper rufous-tawny on the upper parts.

P. cinnamomeus ranges from Southern Mexico (Tabasco) and

Guatemala southwards to Western Hecuador ‘7.

97. LATHRIA UNIRUFA CASTANEOTINCTA Hart.

[|Lipaugus unirufus Selater, P. Z. 8. 1859, p. 385 (1860.—

Oaxaca, S. Mexico, and Guatemala). |

* =P. atricapillus auct. (nec Merrem); cf. Berlepsch, Nov. Zool. xv. 1908,

p. 141.

+ In Upper Amazonia (Eastern Ecuador; Ucayali, Peru) its place is taken by

another form, closely allied to, or perhaps even identical with, P. castaneus Jard.

& Selby. [= P. rufus auct, nec Boddaert !]

BIRDS OF WESTERN COLOMBIA. 1145

Lathria unirufus castaneotinctus Hartert, Nov. Zool. ix. p. 610

(1902.—Rio Durango, N.W. Ecuador).

Lathria unirufa (nec Sclater) Hartert, Nov. Zool. v. 1898,

p. +90 (Cachabi, N.W. Ecuador); Sclater & Salvin, P. Z.8. 1879,

p-. 918 (Remedios, Neche).

Lipangus unirufus (nec Sclater) Cassin, Proc. Acad. N. Sci.

Philad. 1860, p. 143 (Turbo, R. Truando).

Nos. 2306, 2315, 2045, 2175. ¢ gd ad. Novita: 18, 20.xi. ;

Noanama: 1.ix.08; Sipi, Rio Sipi: 10.x.08.—Wing 130-136 ;

tail 102-108; bill 20-21 mm.

Nos. 2917, 2342, 2343, 2423. 2 2 ad. | Cajon, Ry Cajon ;

28.x.08 ; Novita: All Sep 29, xii.08.— Wing 123-128; tail 101-105;

bill 20-21 mm.

“Tris dark brown, feet grey, maxilla brown, mandible pale

brown or grey.” !

These specimens agree with topotypical birds from N.W.

Ecuador and differ from Z. w. clara Ridgw.* of southern

Central America, in having the back decidedly brighter, more

cinnamon-rufous, and the under parts also darker, deep tawny-

ochraceous. The bill also is light enon instead of blackish

horn-colour.

The specimens from Antioquia (aomedion Neche) in the

British Museum likewise belong to this form; and so probably

also do those obtained by Lieut. Michler’s expedition to the Rio

Truando (although they are referred by Mr. Ridgway t to

Lu. clara), since the fauna of the Atrato region is essentially

identical with that of the Choc6 district.

LL. u. castaneotincta, therefore, appears to range from the coast

of the gulf of Uraba through Western Colombia to the province

of Esineraldas, N.W. Ecuador.

98. LIPAUGUS HOLERYTHRUS ROSENBERGI Hart.

[Lipaugus holerythrus Sclater & Salvin, P. Z. 8. 1860, p. 300

(1860.—Vera Paz, Guatemala). |

Lipaugus holerythr us rosenbergi Hartert, Bull. B. O. C. xvi. p. 12

(1905.—Rio Dagua, W. Colombia).

Lipaugus holerythrus (errore) Hartert, Nov. Zool. v. 1898, p. 489

(Cachabi, N.W. Ecuador).

Nos. 2355, 2375. 3 d ad. Néovita: 30.x1., 7 xii,08.—Wing 104,

NOD sian! 90, 895 bill 19, 20 mm.

No. 2100. 9 ad. Sipi, Rio Sipi: 21.ix.08.—Wing 102;

tail 90; bill 19 mm. ;

“Tris dark brown, feet dark grey or black, maxilla blackish,

mandible brown.”

These birds, which are to be regarded as topotypical of Z wu.

rosenbergi, differ from Central American skins in their darker

* Proc. Biol. Soc. Wash. xix. p. 120 (1906.—Lion Hill, Panama).

y+ Bull, U.S. Mus. no. 50, pt. iv. p. 824.

1146 MR. C. E. HELLMAYR ON THE

tawny russet upper, and deeper tawny under parts, as Sonal

pointed out by Hartert.

Although I have not seen the examples from Antioquia (Neche,

Remedios *), I have little doubt that they also are referable to

the present form rather than to typical L. h. holerythrus.

L. h. rosenbergi is another of the many species peculiar to

Western Colombia and N.W. Ecuador (province Esmeraldas).

99. CorinGA NATTERERID Boiss.

Ampelis nattererii Boissonneau, Rey. Zool. iii. p. 2 (1840.—

Santa-Fé-de- Bogota).

Cotinga nattererii Ridgway, Bull. U.S. Mus. no. 50, pt. iv.

1907, p. 785 (Hastern Panama (Railway line) to Colombia ;

crit.).

Cotinga simont Berlepsch, Ornis, xiv. p. 361 (1907.—San Jose,

Rio Dagua, W. Colombia).

Nos. 2267, 2268, 2269, 2281, 2283, 2289, 2290, 2291, 2292.

6 dad. Névita, Rio Tamang: 12 i 16.xi.08..-Wing 107—

111; tail 66-69; bill 143-153 mm.

No. 2270. gS imm. Novita: 12 ix.08.—Wing 108; tail 67;

bill 153 mm.

Nos. 2282, 2293. 6 g juv. Névita: 14, 16.xi.08.—Wing 105;

tail 68, 70; bill 14, 15 mm.

Nos. 2271, 2275, 2284, 2294, 2295, 2296. 9 9 ad. Novita:

12, 18, 14, 16.xi.08.—Wing 106-110; tail 71-75; bill 143-

15 mm.

“Tris black, feet dark grey, maxilla black, mandible blue.”

The series of adult males is fairly uniform, the variation

being chiefly confined to the intensity of the purple colour

of the throat and abdominal patch. Two specimens, especially

nos. 2281, 2291, have the throat darker than the others. All,

however, show the characteristic blackish cross-bars (whieh

sometimes have a slight metallic greenish sheen) to the feathers

of the throat, already noticed by M. Boissonneau and more fully

described by Mr. Ridgway, and the pale cerulean blue border to

the gonydeal angle; the pileum is constantly deeper blue than

the back; the second primary is very nearly as long as the third,

and never attenuated on its apical portion, so striking a feature

in the allied C. ridgwayi Ridgw., from Chiriqui and 8.W. Costa

Rica.

I am sorry to say that C. simoni is clearly a synonym of

C. nattererii, for the description of Boissonneau, which Count

Berlepsch appears to have overlooked, corresponds exactly to the

Chocé birds.

I do not understand the Count’s statement as regards the

coloration of the female which, he says, eens that of

* TL, holerythrus Sclater & Salvin, P. Z. S. 1879, p. 519 (Neche); Sc a er, Cat. B.

xiv. 1888, p. 357 (part.: Remedios, Neche).

BIRDS OF WESTERN COLOMBIA. 1147

C. cayana. As a matter of fact, the six females obtained by

Mr. Palmer are exactly like C. ridgwayi except in being slightly

deeper buff on the anterior under parts. In the main features,

viz., crown and back sooty edged with buffy white, larger

upper wing coverts broadly margined with cinnamon-buff, dusky

markings of lower surface etc., the females of the two species are

practically identical.

C. nattererii ranges from Eastern Panama (Railway line) south

to Buenaventura, Western Colombia. Occasionally it is also met

with in Bogota collections.

100. CARPODECTES HOPKE! Berl.

Carpodectes hopkei Berlepsch, Orn. Monatsber. v. p. 174 (1897.—

San José, R. Dagua, W. Colombia); Hartert, Nov. Zool. ix, 1902,

p-. 611 (Ventana, N.W. Ecuador).

Nos. 2205, 2272, 2285, 2286, 2287, 2288. ¢ ¢ ad. & imm.

Novita: 12, 14, 18.x1. 08.— Wing 158-170; tail 95-102; bill 20-

213 mm.

No. 2273. (¢) juv. Novita: 12.xi.08.—Wing 158; tail 92;

bill 20 mm.

Nos. 2303, 2325, 2326, 2335, 2339. 9 9 ad. &imm. Névita:

17, 23, 25, 26.x1.08—Wing 135-140; tail 84-87; bill 18-

19 mm.

No. 2274. (¢) juv. Novita: 12.x1.08.—Wing 154; tail 100;

bill 18 mm.

‘Tris orange (or orange-yellow), feet and bill black.”

“J have seen, but been unable to shoot this bird in all parts

of the Chocé from Malaquita (S. T.) up to Sipi” (M. G. Palmer).

The series of males illustrates beautifully the variation accord-

ing toage. In perfectly adult birds the six outer primaries and

the central pair of rectrices bear only a small, rounded black spot

at the tip, while the plumage otherwise is pure white, some-

times with a scarcely perceptible greyish tinge on the pileum.

Immature males have much more black on the wings, and most,

if not all, of the rectrices tipped with black. The younger the

bird, the more extended is the black colour. No. 2287 has not

only the black tips to the three outer primaries much larger than

tht specimens in more advanced stage, but the greater part of the

' inner web of the remaining primaries isalso black. The shafts of

the primaries are pure white in adult, black in immature males.

The latter, besides, have the bastard-quills and primary coverts

edged or tipped with black.

No. 2273 is changing from the grey juvenile plumage into the

white of the adult. Another, still younger, male, No. 2274,

resembles the females, but may be distinguished by its larger size,

much paler cinereous upper parts and breast, whiter belly, ete.

A specimen in the Tring Museum from Ventana, N.W. Ecuador,

is exactly like it.

C. hopket is peculiar to the hot, forest-covered lowlands of

1148 MR. C. E. HELLMAYR ON THE

Western Colombia and the adj joining parts of Ecuador (province

Esmeraldas).

101. QuERuLA puRPURATA P. L. S. Mull.

Muscicapa purpurata P. lL. 8. Miller, Natursyst. Suppl.

p. 169 (1776—ex Daubenton, Pl. Enl. 381 : Cayenne).

Querula cruenta Cassin, Proc. Acad. Philad. 1860, p. 143

(Turbo); Sclater & Salvin, IPS a Se USD, jon" BAO (Pocune,

Remedios).

Nos. 2001, 2151, 2265. ¢ gd ad. Noanama: 22.vii.; Sipi:

3.x.; Névita: 12.xi.08.—Wing 178-182; tail 114-117; bill

24 mm. ;

Nos. 2044, 2152, 2266. 9 2 Noanama: J.ix.; Sipi: 3.x. ;

Novita: 12.x1.08.—Wing 166-168; tail 108-112; bill 22—

233 mm.

* Tris dark brown, feet black, bill blue.”

These birds have generally smaller bills than those from

Guiana, Venezuela (Caura) and Parad, but the difference is not

quite constant. ‘This is a wide- spread species, ranging from

Costa Rica to Peru, and the mouth of the Amazons.

102. SYNALLAXIS PUDICA Scl.

Synallaxis pudica Sclater, P. Z. 8. 1859, p. 191 (1859.—

Bogota) ; Sclater & Salvin, P. Z 8. 1879, p.-521 (Remedios,

Antioquia).

No. 2116. 9 ad: Sipi: 26.1x.08—Wing 60; tail 73; bill

15 mm.

No. 2730. 2 juv. Pueblo Rico, 5200 ft.: 16.viii.09.

“ Tris brown, feet blue-grey (no. 2116), dark brown (no. 2730),

bill black, mandible grey.”

Identical with specimens from Bogota collections.

103. SYNALLAXIS UNIRUFA Lafr.

Synnalaxis (sic) unirufa Lafresnaye, Rev. Zool. vi. p. 290

(1843.—* Colombia ”).

Synallaxis unirufa Sclater & Salvin, P. Z.S. 1879, p. 521

(Antioquia).

No. —. g ad. Tatamaé Mountain, OU ft., 9.x.09.—Wing

61; tail 70 mm. ; bill damaged.

«i Iris brown, lees dark grey, bill black.”

Agreeing with a Bogotdé skin. [he plumage is uniform clear

chestnut rufous, vather paler underneath, except, of course, the

blackish inner webs of the remiges. There is no black cular

patch, so characteristic a feature in the allied S. castanea Sel. iy

from the Silla de Caraccas, Venezuela.

S. unirufa is peculiar to the high mountains of Colombia.

* Ann. Mag. Nat. Hist. (2) xvii. p. 466 (1856.—Caraccas).

BIRDS OF WESTERN COLOMBIA. 1149

104. SIproRNIs ERYTHROPS GRISEIGULARIS Ridgw.

[Synallaxis erythrops Sclater, P. Z. S. 1860, p. 66 (1860.—

Pallatanga, W. Ecuador). |

Acrorchilus erythrops griseigularis Ridgway, Proc. Biol. Soc.

Wash. xxi. p. 72 (1909.—San Antonio, Rio Cali, ““N.W.”

Colombia; M. G. Palmer coll.).

Synallaxis erythrops (nec Sclater) Sclater & Salvin, P. Z.S8. 1879,

p. 521 (Frontino, W. Cordillera).

Nos. 3740, —. 6 ¢ ad. Pueblo Rico (5200 ft.), 7.1x.; Siat6

(5200 ft.), ix.09.— Wing 69; tail 63, 65; bill 13 mm:

No. 2799. 2 juv. Loma Hermosa, R. Jamaraya (4150 ft.),

18.x.09.— Wing 63: tail 68; bill 13 mm.

“ Tris brown, feet greyish green, maxilla black, mandible light

brown.”

The adult birds fully bear out the characters given by Mr.

Ridgway for his subspecies which, moreover, was based upon

one of Palmer’s skins. SS. ¢. griseigularis may easily be recog-

nized from its allies: S. e. erythrops Scl., of Western Heuador,

and 8. e. rufigenis Lawr.*, of Chiriqui and Costa Rica. by its

mouse-grey (instead of buffy brown) foreneck and breast, and

brighter cinnamon-rufous median rectrices.

S. e. griseigularis is peculiar to the elevated districts of the

Western Cordillera of Colombia.

[105. THriPADECTES SCLATERI Berl.

Thripadectes sclateri Berlepsch, Ornis, xiv. p. 365 (Feb. 1907.—

S. Pablo, W. Colombia, 4500 ft. ; G. Hopke coll.).

Rhopoctites alogus Bangs, Proc. Biol. Soc. Wash. xxiii. p. 72

(1910.—Pavas, W. Colombia, 4400 ft.; M.G. Palmer coll.).

Count Berlepsch having kindly submitted to my inspection the

type specimen, 1 can positively state that f. alogus is merely a

synonym of 7’. sclateri, the description of which appears to have

been overlooked by Mr. Bangs. The two types were obtained in

the same district, viz., in the Western Cordillera above Buena-

ventura, Chocé Bay. Although somewhat resembling Automolus

(Rhopoctites) rufobrunneus in general coloration, 7’. sclateri isa very

distinct species, and I agree with its describer that it finds its

natural place in the genus Thripadectes. The structural characters,

especially the shape and size of the bill, are exactly the same as

in 7. flammulatus, while A. rufobrunneus has a much shorter,

slenderer, more depressed bill, and much shorter, weaker tarsi.

The pileum has distinct, though narrow, pale shaft-lines, whereas

it is uniform dusky olive in A. rufobrunneus ; the back is darker,

less reddish ; the throat much paler, light ochraceous instead of

deep orange; the sides of the neck are olive-brown instead of

orange-ochraceous ; the breast and abdomen are darker ochreous

* Synallaxis rufigenis Lawrence, Ann. Lyc. N. H. N.Y. ix. p. 105 (1868.— Costa

Riva).

1150 MR. GC. E. HELLMAYR ON THE

and lack the pale shaft-stripes, so conspicuous in A. rufo-

brunneus, ete. etc.

The type, an adult female, obtained by the late Gustav

Hopke, March 6, 1897, measures—wing 99; tail 95; bill

27 mm. |

106. AUTOMOLUS NIGRICAUDA Hart,

Automolus nigricauda Hartert, Bull. B. O. C. vil. p. xxx,

(1898.—Cachabi, N.W. Ecuador); idem, Nov. Zool. v. 1898,

p. 491 (Cachabi); idem, 1. ¢. ix. 1902, p. 616 (Paramba, 3500 ft. ;

Rio Sapayo, 450 ft.: N.W. Ecuador).

No. 2414. g ad. Noévita, 150 ft., 18.x11.08.— Wing 83 ; tail 71;

bill 23 mm.

“ Tris light brown, feet and bill black.”

This bird is practically identical with the specimens in the

Tring Museum. ‘The species was hitherto known only from

N.W. Ecuador, where Mr. Rosenberg and his collectors obtained

three examples at Cachabi (500 ft.), Paramba (3500 ft.), and Rio

Sapayo (450 ft.).

A. nigricauda is nearly allied to the group of A. rubiginosus

from Central America, but may easily be distinguished by its dull

blackish (instead of bright chestnut rufous) tail, less rufous wings,

darker ferruginous foreneck, dull brownish olive belly, etc., ete.

107. HytoctistEs SUBULATUS ASSIMILIS Ber]. & Tacz.

[Sphenera subulata Spix, Av. Bras. i. p. 82, pl. Ixxxii. fig. 1

(1824—‘* in sylvis fl. Amazonum ”). |

Automolus assimilis Berlepsch & Taczanowski, P. Z. 8S. 1883,

p. 561 (1883.—Chimbo, 8.W. Ecuador); Hartert, Nov. Zool. v.

1898, p. 491 (Chimbo, Cachabi, W. Ecuador).

No. 2080. $ (2) ad. Sipi, Rio Sipi: 12.ix.08.—Wing 82;

tail 71; bill 203 mm.

No. 2467. 3 (%) ad. Noanama: 13.1.09.—Wing 80; tail 69;

bill 20 mm.

Nos. 2725, 2701. 6 Qad. Tadé: 8, 28.vi.09.—Wing 87, 86;

tail 70; bill 21, 22 mm.

No. 2093. @ ad. Sipi: 19.ix.08—Wing 83; tail 67; bill

20 mm.

“ Tris dark brown, feet greyish green or black brown, maxilla

black, mandible light brown.”

This series agrees perfectly with two specimens from Western

Keuador (Carondelet, Paramba) in the Munich Museum, The

upper part of the head is nearly uniform olive-brown with

obsolete dusky edges to the feathers; the back wholly unstriped,

rufescent brown; the upper wing-coverts and outer webs of the

remiges are deep russet-brown; the breast and abdomen dull

light olivaceous, the chest but indistinctly mottled with buffy,

etc. etc.

BIRDS OF WESTERN COLOMBIA, ILI

The Costa Rica form, H. s. virgatus Lawr.*, may be distin-

guished from H. s. assinvilis by its more elongated as well as

slenderer bill (234 to 24 mm.); bright chestnut-brown colour

of the wing-coverts and remiges; blackish ground-colour of the

pileum and nape, with sharply defined, buff shaft-streaks; by

having the upper back distinctly streaked with buff, ete.

Mr. Ridgway has rightly separated P. virgatus under the new

generic term Hyloctistes ~, but it is certainly only subspecifically

distinct from H. s. subulatus and H. s. assimilis, the three races

replacing each other geographically. With regard to the differ-

ences between the two last-named races see my remarks in Noy.

Zool. xvii. 1910, p. 320.

108. XENICOPSIS SUBALARIS SUBALARIS Scl.

Anabates subalaris Sclater, P. Z. S. 1859, p. 141 (1859.

tanga, W. Ecuador).

Anabazenops subalaris Sclater, Cat. B. Brit. Mus. xv. p. 108

(part. : a-c, Pallatanga; d, Quito).

No.—. ¢ ad. Loma Hermosa, R. Jamaraya, 4150 ft.,

19.x.09.— Wing 91; tail 84; bill 19 mm.

“Tris dark grey, feet grey-green, maxilla black, mandible

yellow.”

This bird is somewhat difficult to place. In coloration, notably

in the dark brown ground-colour of the pileum and back, it agrees

with topotypical West Ecuadorian specimens; but it is fully as

large as the Central American form, X. s. lineatws Lawr. f,

which, however, has the head above and the back of a distinctly

paler, more olivaceous brown tinge. Until more specimens from

Western Colombia come to hand its identification can be regarded

only as provisional.

X. s. subalaris inhabits the elevated districts of the Western

Cordillera in Ecuador and Colombia, extending down the slope to

about 2000 feet.

X. mentalis Tacz. & Berl.§, united by Dr. Sclater to 1. s.

subalaris, is a very distinct species, though it may eventually

prove to be the Hastern representative. It differs particularly in

the following characters: the top of the head is much darker,

blackish, and covered with broad, buff shaft-stripes, while in

A. s. subalaris there are but a few narrow hair-like streaks to be

seen ; the upper back also shows much broader as well as more

numerous buff stripes; the whole under surface, posterior to

foreneck, is likewise broadly striped all over with yellowish

Paila-

* Philydor virgatus Lawrence, Ann. Lyc. N. H. N.Y. viii. p. 468 (1867.—Ango-

stura. Costa Rica).

+ Proc. Biol. Soc. Wash. xxii. p. 72 (1909.—Type: Philydor virgatus Lawr.).

t Anabazenops lineatus Lawrence, Ann. Lyc. N. H: N.Y. viii. p. 127 (1865.—

ncoctarss Costa Rica).

§ Anabazenops mentalis Taczanowski & Berlepsch, P. Z.S. 1885, p. 96 (1885.—

Machay, Eastern Ecuador).

1152 MR. C. E. HELLMAYR ON THE

white, etc., etc. The pattern of the upper parts reminds one

rather of Automolus holostictus Scl. & Salv., yet it need not be

emphasized that this species is generically distinct.

Of X. mentalis, I have examined two adult males and an

immature female from Machay, E. Ecuador, and two Bogota

skins, one in the British Museum, the other in Count Berlepsch’s

collection.

109. XENOPS GENIBARBIS LITTORALIS Scl.

[Xenops genibarbis Mliger, Prodr. Syst. Mamm. et Av. p. 213

(1811.—Cameta, Lower Amazons). |

Xenops littoralis Sclater, P. Z. S. 1861, p. 379 (1861.—Esme-

raldas, N.W. Ecuador).

Xenops ruficauda (nec Vieillot) Cassin, Proc. Ac. N, Sci. Philad.

1860, p. 193 (Turbo, N. Colombia).

Xenops genibarbis (nec Lliger) Sclater & Salvin, P. Z.8. 1879,

p. 523 (Remedios).

No. 2039: ¢ imm. Noanama: 31.viii.08.—Wing 61; tail 46;

bill 123 mm.

No. 2479. 9 ad. Noanama: 15.1.09.—Wing 61; tail 463;

bill 15 mm.

No. 2646. g ad. Tadé, R. San Juan: 17.v.09.—Wing 623;

tail 48 ; bill 13 mm.

“ Tris dark brown, feet blue-grey, bill black.”

These birds agree with topotypical examples from the Ecuadorian

province of Esmeraldas. _X. g. mexicanus Scl.*, which ranges

from E. Mexico to Panama, is more rufescent, less olive, under-

neath, and has the back of a brighter, clearer cinnamomeous-

brown.

X. q. littoralis inhabits the lowlands and foothills of Western

Eeuador and Western Colombia.

110. GuyPHORHYNCHUS CUNEATUS CASTELNAUDIT Des Murs.

[Dendrocolaptes cuneatus Lichtenstein, Abhand], Akad. Berlin

a.d. Jahren 1818-19, p. 204, pl. un. fig. 2 (1820—“‘in Brasilice

provincia Bahia”; cfr. idem, |. c. a.d. Jahren 1820-21, p. 264. |

Glyphorhynchus castelnaudii Des Murs, in Castelnau’s Voyage,

Oiseaux, p. 47, pl. xv. fig. 2 (1856.—Santa Maria, Hastern Peru).

Glyphorhynchus cuneatus (evrore) Sclater & Salvin, P. Z.8. 1879,

p- 523 (Remedios).

Nos: 2019 2d: o mm. Orad.) peiprs) L0set Neamamanr

26.vili.08.— Wing 69, 67; tail 70, 62; bill 12 mm.

“Tris dark brown, feet dark grey, maxilla black, mandible

rey.”

Slightly smaller than Upper Amazonian skins, but similar in

coloration.

_ %* Xenops “ mexicanus Sclater, P. Z. S. 1856, p. 289 (Jan. 1857.—Cordova;

E. Mexico).

BIRDS OF WESTERN COLOMBIA. 1158

111. DENDRORNIS LACHRYMOSA ROSTRATA Ridew.

[Dendrornis lachrymosus Lawrence, Ann. Lye. N. ET INE ONES ‘val

p. 467 (1862.— Panama). |

Xiphorhynchus lacrymosus rostratus Ridgway, Proc. Biol.

Soc. Wash. xxii. p. 73 (1909:—Rio Dagua, ‘¢ North-western ”

Colombia).

Dendrornis lacrymosa (errore) Sclater & Salvin, P. Z.8. 1879,

p. 523 (Remedios) ; Hartert, Nov. Zool. ix. 1902, p. 616 (N.W.

Keuador).

Dendrornis — sp. Cassin, Proc. Acad. N. Sci. Philad. 1860,

p. 194 (R. Truando).

Nos. 2587, 2615. g¢ gad. Condoto, R. Condoto, NBO) siaey 3

6,20.iv.09.—Wing 126, 122; tail 103, 98; bill 837 mm.

Nos. 2004, 2005. dg imm., 2 imm. Noanama, 1000 fia: 24,

26.viii.08.— Wing 126, 120; tail 105, 102; bill 35, 32 mm.

“Tris dark brown, feet green, maxilla black, mandible grey.”

These specimens, as well as several others from North-western

Ecuador (Bultin, Carondelet), have the bill slightly deeper and

stouter than typical birds from Panama. The culmen, how-

ever, is not more strongly arched. I must confess I am not

very confident as to the distinctness of this race and should not

be surprised if a larger series would show it to be inseparable

from D. 1. lachrymosa. D.1. rostrata, if really distinguishable,

ranges from N.W. Colombia (Rio Truando) along the Pacific coast

southwards to the north-western portion of Ecuador. It is found

only in the lowlands and on the hill-slopes.

112. DENDRORNIS TRIANGULARIS ZQUATORIALIS Berl. & Tacz.

[Dendrocolaptes triangularis Latresnaye, Rev. Zool. v. p. 134

(1842.—“ Bolivia,” * errore; the type came from Santa-Fé-de-

Bogota, Colombia). |

Dendrornis erythropygia equatorialis Berlepsch & Taczanowski,

P. Z. 8. 1883, p. 563 (1883.— Chimbo, W. Keuador) ; Hartert,

Nov. Zool. v. 1898; p. 490 (Chimbo, Paramba, W. Eeuador).

D. cequatorialis Salvadori & Festa, Boll. Mus. Torino, XIV.

no. 362, 1899, p. 25 (R. Peripa, W. Ecuador).

D. punctigula (errore) Goodfellow, Ibis, 1902, p. 63 (Nanegal,

Intac, Gualea, W. Ecuador; specimens examined).

No. 2378. g ad. Névita: 7.x11.08—Wing 117; tail 103;

bill 34 mm.

* The locality “ Bolivia” is certainly a mistake, for in Mag. Zool. 1843, Oiseaux,

pl. 32, Lafresnaye expressly says: ““ vient de Santa-Fé-de-Bogoté.” This statement

is repeated in Rev. Mag. Zool. (2) ii. 1850, p. 419, in the French text, while the

Latin diagnosis (a verbatim reprint of the original account) terminates with the

indication: “habitat in Bolivia.” Moreover, at that time, Bogoté skins reached

Paris in large numbers, whereas material from Bolivia was very scarce in European

Museums. In fact, D'Orbigny was the only naturalist to send collections from that

country, but he did not obtain the species in question.

1154 MR. CG. E. HELLMAYR ON TIE

No. 2529. gad. El Tigre, 320 ft.: 10.11.09.—Wing 116;

tail 103; bill 834 mm.

“Tris dark brown, feet grey-green or blue, maxilla black, tip

light brown, mandible grey.”

The two skins agree perfectly with a series from Western

Ecuador in the Munich Museum. JD. ¢. cequatorialis is an

excellent form, differing in many important details from the

typical race of Central Colombia, W. Venezuela, etc. The rump

and the wings are much darker chestnut rufous; the ground-

colour of the throat is decidedly buff, instead of whitish ; the dark

olive markings on the throat are restricted to small, rounded

apical spots, while in the allied D. ¢. triangularis the feathers

show a continuous, marginal edge, giving a squamate appearance ;

the top of the head is wholly unstreaked, plain dark olive, only

sometimes with a few buff shaft-lines on the forehead, whereas

the typical form has the entire crown conspicuously streaked or

spotted with pale buff, ete.

D. t. punctigula Ridgw. * from Costa Rica and Chiriqui, is

most closely related to D. t. equatorialis, agreeing in the pattern

of the throat and in the uniform pileum, but differs in the clearer

rufous rump and wings, the deeper olive-green ground-colour, and

the smaller, less rounded spots of the lower parts. Single

examples are, however, not always distinguishable.

D. t. triangularis shows a distinct, whitish lateral stripe near

the lower edge of the maxilla, while in D. t. equatorialis and

D. t. punctigula the upper bill is uniform dark horn-brown or

blackish. This difference is quite constant in the enormous series

which I have examined.

D. ¢. equatorialis is strictly confined to the western slopes of

the Andes in Ecuador and Southern Colombia. In Ecuador it

ranges from sea-level up toabout 3500 feet, for we have specimens

from Ventana (90 feet) and Bultn (160 feet) as well as from Lita

(3200 feet) and Paramba (3500 feet). From Colombia, west of

the Coast Cordillera, I have seen only Palmer’s two skins, one

taken at 150, the other at 320 feet elevation 7.

D. t. triangularis inhabits the Central and Eastern Cordillera

of Colombia, the mountains of Western Venezuela (Merida,

Cumbre de Valencia), the eastern slopes of the Ecuadorian Andes,

and Northern Peru. ‘Two specimens from the Cauca slope of the

Coast Cordillera—one, from Santa Hlena, in the British Museum,

the other, from Riolima, at Tring—also belong to this form, being

in every respect typical. Birds from Hastern HEeuador (Machay,

* Dendrornis punctigula Ridgway, Proc. U.S. Mus. xi. p. 544 (1889.—Naranjo,

Costa Rica).

+ My material of D. ¢. equatorialis consisted of the following specimens :—

6 from Chimbo, including the type (Mus. H.v. Berlepsch & Tring); 2 from Cachabi

(Tring); 7 Paramba, 6 Lita, 1 Ventana, 5 Bultin (Mus. Tring, Munich, Vienna, etc.) ;

3 Nanegal, 2 Nono, 1 Intag (Goodfellow coll—TIring), 2 Chocéd, W. Colombia

(Munich): 385 examples in all.

BIRDS OF WESTERN COLOMBIA. 1155

Baeza *) and North Peru (Chachapoyas) are absolutely identical

with those from Bogoté and Venezuela 7.

Ona future occasion [I hope to give some information respecting

several other species of the genus Dendrornis which, through

lack of proper material, are very imperfectly known.

113. DENDROCINCLA MERULOIDES LAFRESNAYEL Ridgw.

| Dendrosops meruloides Lafresnaye, Rev. Mag. Zool. (2) iii.

p. 467 (1851.—‘“ Cote ferme,” sc. Cumana, Venezuela). |

Dendrocincla lafresnaye. Ridgway, Proc. U.S. Mus. x. 1887,

p. 492 (1888.—‘“ Upper Amazons ?”—errore; we substitute

Colombia as type locality).

D. olivacea latresnayei Oberholser, Proc. Acad. N. Sci. Philad.

1904, p. 457 (crit.).

No. 2460. 9 (?) ad. Noanama: 11.1.09.—Wing 102; tail

88; bill 26 mm.

No. 2113. 9 imm. Sipi: 23.1x.08.—Wing 95; tail 81; bill

25 mm.

‘“‘ Tris dark brown, feet and bill black.”

These specimens agree well with others from Western Ecuador,

and Riolima, Cauca Valley. Bogota skins are slightly more

rufescent both above and below, thus pointing towards JD. J.

pheochroa Berl. & Hart.

D. m. lafresnayert, D. m. pheochroa, and D, m. meruloides are

clearly races of one and the same species, representing each other

geographically. The first-named form may always readily be

distinguished from the two others in having the outer webs of the

quills conspicuously washed with olivaceous, and the dusky

portions of the bill much darker, black instead of horn-coloured.

On the other hand, D. m. meruloides differs from D. i pheochroa

in its much brighter, more cinnamon or ochraceous-brown

coloration.

Range and principal synonyms of these forms are as follows :—

(a) D. MERULOIDES MERULOIDES Lafr.

Dendrocops meruloides Lafresnaye, Rev. Mag. Zool. (2) iii.

p- 467 [1851.—“ Cote ferme,” sc. Cumand; cfr. Ménégaux

& Hellmayr, Mém. Soc. Hist. Nat. Autun, xix. 1906, p. 121

crit.) |.

: Dee meruloides Cabanis & Heine, Mus. Hein. ii. 1859,

p. 34 (Caraccas); Sclater & Salvin, P. Z.8. 1868, p. 167 (Pilar,

Venezuela); Chapman, Bull. Amer. Mus. N.Y. vi. 1894, p. 48

(Trinidad) ; Berlepsch & Hartert, Nov. Zool. ix. 1902, p. 67

* D. erythropygia(!) Goodfellow, Ibis, 1902, p. 63 (Baeza, EH. Ecuador).

+ My material of D. t. triangularis was composed as follows :—9 Bogota (Tring,

Munich, H.& B.), 1 Santa Elena, 1 Riolima,Cauca (Brit. Mus., Tring), 10 Merida

(fring & Munich), 3 Cumbre de Valencia (Munich), 4 East Ecuador (2 Machay,

H. & B.; 2 Baeza, Tring), 2 N. Peru, Chachapoyas (Tring) ; 30 examples in all.

{ =D. olivacea auct. ex parte: America merid.

1156 MR. C. E. HELLMAYR ON THE

(Cumand, San Hsteban) ; Hellmayr, Nov. Zool. xiii. 1906, p. 29

(Trinidad).

D. meruloides aphanta Oberholser, Proc. Acad. N. Sei, Philad.

1904, p. 460 (Lobago).

Type locality : Cumana, Venezuela.

Hab. Tobago ; Trinidad ; North Coast of Venezuela: from

Cumanid, to Puerto Cabello.

Examined: 16 Trinidad, 2 Tobago, 6 Cumana, 1 Caraccas,

12 San Esteban (near Puerto Cabello).

(b) D. meruLorEs PHmocHROA Berl. & Hart.

Dendrocincla® pheochroa Berlepsch & Hartert, Nov. Zool. 1x.

p. 67 (1902.—Munduapo (type), Maipures, Nericagua, Orvinoco ;

Suapure, La Pricion, Caura R.) ; Thering, Rev. Mus. Paul. vi.

1905, p. 437 (Rio Jurud); Snethlage, Jou. f. Orn. 1908, p. 15

(Rio Purts).

D. olivacea pheochroa Oberholser, Proc. Acad. N. Sci. Philad.

1904, p. 458 (Suapure, La Union, Caura).

D. fumigata (errore) Pelzeln, Zur Orn. Bras. 1. 1867, p. 42

(part. Rio Branco, spec. examined) ; Taczanowski, P.Z.S. 1882,

p- 27; idem, Orn. Pérou, il. 1884, p. 168 (Huambo, N. Peru).

D. olivacea (nee Lawrence) Sclater, Cat. B. xv. p. 166 (part. :

I, m, Sarayacu, E. Ecuador; », Rio Napo ; q, Iquitos, INGuletere: 2

spec. examined),

Dendrocolaptes atrirostris (nec Lafr. & D’Orb.) Selater, P. ZS,

1858, p. 63 (Rio Napo; spec. examined).

Type locality: Munduapo, Orinoco R.

Hab. Amazonian region: from the Rio Branco, N. Brazil, and

the Caura Valley, Eastern Venezuela, westwards to N. Peru and

the eastern slopes of the Colombian and Ecuadorian Andes.

Examined : 7 Munduapo (inel. type), 2 Maipures, 2 Nericagua,

6 Caura, 1 Rio Branco, 1 R. Jurua, 2 N. Peru (Iquitos, R. Tigre),

3 Rio Napo, E. Ecuador, 3 Bogotsa.

D. lafresnayei Ridgway, Proc. U.S. Mus. x. 1887, p. 492

(1888.—“ Upper Amazons,” errore).

D. olivacea lafresnayei Allen, Bull. Amer. Mus. xii. 1900, p. 156

(Santa Marta); Oberholser, Proc. Acad. N. Sci. Philad. 1904,

p. 457 (crit.).

Dendrocops atrirostris (errore) Sclater, P.Z.S. 1860, p. 66

(Pallatanga, W. Ecuador) ; idem, l.c. p. 278 (Babahoyo) ; idem,

lc. p. 293 (Esmeraldas).

Dendrocincla atrirostris Taczanowski, P. Z.8. 1877, p. 332

(Palmal, 8.W. Heuador); Berlepsch & Taczanowski, P. Z.8. 1883,

p. 563 (Chimbo, 8.W. Eeuador).

D. olivacea Sclater, Cat. B. Brit. Mus. xv. p. 166 (part. :

Colombia (Bogoti, Manaure), W. Ecuador); Hartert, Nov. Zool.

* Spelt Dendrocinda.

BIRDS OF WESTERN COLOMBIA. eae

v. 1898, p. 491 (Paramba, N.W. Ecuador); Goodfellow, Ibis, 1902,

p- 63 (Sto. Domingo, W. Ecuador); Salvadori & Festa, Boll. Mus,

Zool. Torino, xiv. no. 362, 1899, p. 27 (Vinces, W. Ecuador).

D. 0. anguina Bangs, Proc. Biol. Soc. Wash. xx. p. 138

(1898.—Santa Marta); idem, l.c. xi. 1899, p. 100 (Sierra

Nevada de Santa Marta).

Hab. Colombia (Bogoti, Santa Marta, Riolima, Chocé, ete.) and

Western Ecuador (Paramba, 8. Javier, Esmeraldas, Pallatanga,

Babahoyo, Vinces, Chimbo, Santo Domingo, Palmal, etc.), from

sea-level up to 6000 feet.

Examined: 15 Western Ecuador, 1 Santa Marta (Manaure),

1 Riolima, Cauca, 2 Chocé, 5 Bogota.

N.B.—The original locality ‘‘ Upper Amazons” is doubtless

erroneous, the species being strictly confined to the western slopes

of the Andes. In the Amazonian district it is replaced by the

preceding form.

114. CYMBILANIUS LINEATUS FASCIATUS Ridgw.

|Lanius lineatus Leach, Zool. Misc. i. p. 20, pl. vi. (1814.—

Berbice, British Guiana). |

Cymbilanius lineatus fasciatus Ridgway, Proc. U.S. Mus. vi.

1883, p. 415 (1884.—Rio Sucio, Costa Rica).

C. lineatus (nee Leach) Sclater & Salvin, P. Z.8. 1879, p. 524

(Remedios, Neche).

No. 2571. ¢ imm. Condoto: 27.11.09.—Wing 72; tail 69;

bill 21 mm.

No. 2572. Q ad. Condoto: 27.11.09.—Wing 72; tail 68;

bill 21 mm.

“ Tris red-brown, feet blue-grey, maxilla black, mandible grey.”

Specimens from W. Colombia and W. Ecuador have the black

barring of the lower parts slightly broader than those from

Cayenne and Amazonia. Although topotypical Costa Rican

birds are not available for comparison, | think they belong to

Ridgway’s subspecies.

Cl. fasciatus is confined to the lowlands and foot-hills.

115. 'THAMNOPHILUS MAJOR TRANSANDEANUS Sel.

[Thamnophilus major Vieillot, Nouv. Dict. 11. p. 313 (1816.—

ex Azara: Paraguay). |

T. transandeanus Sclater, P. Z. 8. 1855, p. 18 (1855.—Guay-

aquil, S.W. Ecuador) ; Cassin, Proc. Acad. N. Sci. Philad. 1860,

p- 188 (Ttirbo).

No. 1972. g ad. Guineo, Rio Calima, 6.vii.08.—Wing 92;

tail 70; bill 27 mm.

No. 2528. 9 ad. El Tigre, R. Tamana, 320 ft., 10.11.09.—

Wing 91; tail 69; bill 27 mm.

No. 3777. g ad. la Selva, R. Jamaraya, 4600 ft., 1.x.09.—

Wing 94; tail 70; bill 28 mm.

‘“‘ Tris vermilion, feet blue, bill black.”

Proc. Zoou. Soc.—1911, No. LX XVIII. 78

1158 MR. C. E. HELLMAYR ON THE

The Tring Museum possesses an adult male obtained by

Mr. Rosenberg at Rio Dagua (Juntas), May 27,1895. The males

from W. Colombia agree, in coloration and size, with a series from

Western Ecuador and Chiriqui, the under tail-coverts being

black with distinct white apical margins.

T. m. transandeanus is, otherwise, known only as an inhabitant

of the lowlands (from sea-level up to 1000 feet), and its occurrence

at La Selva (4600 feet alt.) must be regarded as exceptional.

In the Central and Eastern Cordilleras it is replaced by a nearly

allied, but easily recognizable race, 7. m. granadensis Cab. ™*,

which forms the passage to 7. m. melanurus Gould, of Upper

Amazonia. It differs, in the male sex, from 7. m. transandeanus

in having the under tail-coverts cinereous with a white apical

edge preceded by a distinct blackish subterminal band, and in

its much slenderer, weaker bill. I have examined several Bogota

skins in the collections of the Paris and Munich Museums, an

adult male from Antioquia (Salmon), and one from near

Mérida (Briceno) at Paris, and four males from N.W. Venezuela

(3d ad, g¢ imm., Mt. Bucarito, Tocuyo; ¢ ad. Hjido, near

Mérida; ¢ ad. San Esteban—Mocquerys coll.) in the Tring

Museum. All of them very clearly show the characters indicated

above. Cfr. also Ménégaux & Hellmayr, Bull. Soc. Philom.

Paris (9) viii. 1906, pp. 25-26.

116. THAMNOPHILUS NEVIUS ATRINUCHA Salv. & Godm.

[Lanius nevius Gmelin, Syst. Nat. 1, 1. p. 308 (1788—

ex Latham : Cayenne). |

Thamnophilus atrinucha Salvin & Godman, Biol. Centr.-Amer.,

Aves, ii. p. 200 (1892.—“‘ Central America,” no type specified ; we

fix Panama as terra typica).

T. nevius (nec Gmelin) Sclater & Salvin, P. Z.S. 1879, p. 524

(Neche, Antioquia); Berlepsch, Journ. f. Orn. 1884, p. 307

(Bucaramanga); Cassin, Proc. Acad. N. Sci. Philad. 1860, p. 188

(Turbo).

No. 1967. ¢ imm. San Joaquim: 4.viii.08.—Wing 71; tail

61; bill 184 mm.

No. 2049. ¢ imm. Noanama: 2.1x.08.—Wing 70; tail 56;

bill 18 mm.

No. 2127. gad. Sipi: 28.ix.08.—Wing 70; tail 54; bill

183 mm.

Nos. 2411, 2425, 2435. § gd ad. Ndéovita: 18, 22, 24.xii.08.—

Wing 70-74; tail 56-59; bill 183 mm.

Nos. 2395, 2428, 2431, 2436. 9 2. No6vita: 12,23, 24,

26.xii.08.— Wing 68-69 ; tail 54-58; bill 17-183 mm.

‘“‘ Tris brown or red-brown in male, grey in female, feet blue or

blue-grey, maxilla black, mandible grey.”

Specimens from Colombia and Western Ecuador are practically

* Diallactes granadensis Cabanis, Journ. f. Orn. 1872, p. 234 (Bogota coll.).

+ 7. transandeanus (errore) Sclater & Salvin, P.Z.S. 1879, p. 524 (Remedios,

Neche: Antioquia).

BIRDS OF WESTERN COLOMBIA. 1159

identical with others from Costa Rica, 7. . atrinucha replaces

7’. n. nevius on the western sides of the Andes, while the typical

form is found in the Guianas, the Orinoco Valley and _ its

tributaries.

For the distinguishing characters and geographical distribution

of the various races of this group see my remarks in Abhandl.

Bayer. Akad. Wissensch. II. Cl., vol. xxii. 3, 1906, p. 658 ff.

117. MyRMOTHERULA SURINAMENSIS PACIFICA, subsp. n.

No, 2132. gd ad. Sipi: 30.1x.08.—Wing 53; tail 30; bill

16 mm.

Nos. 1974, 2133. 9 9 ad. Guineo, R. Calima: 6.viii.08 ;

Sipi: 30.1x.08.— Wing 52, 50; tail 30; bill 143, 15 mm.

[No. 09.788, g ad. Buenaventura, 23.ii1.99—Wing 522;

tail 29; bill 15 mm.

No. 09.787. g ad. Plano de los Monos, near Naranjo, 2800 ft.,

3.iv.99.— Wing 53; tail 313; bill 15 mm.

No. 09.789-791. 9 9 ad. Buenaventura: 11,18, 23.iii1.99.—

Wing 51-52; tail 28-30; bill 144-15 mm.

These five specimens were collected by Hugéne André. |

“ Tris dark brown, feet blue-grey, bill black, mandible grey.”

Adult. Differs from J/. s. surinamensis Gm., in its longer

tail, much stronger as well as decidedly longer bill, wider white

tips to the outer rectrices (about 3 to 4 mm. on outermost pair),

and in having the upper back less variegated with black, while

the white interscapular blotch is much smaller in the male and

wholly absent in the female. The latter sex, too, has the top of

the head conspicuously clearer, about “tawny ochraceous” (Ridgw.

Nomencl. v. fig. 4) instead of “deep tawny ” (l.c. v. fig. 1), and

the dark stripes on the hind crown and nape are much duller and

less pronounced, being pale dusky olive instead of deep black.

Type in the Zoological Museum, Munich: no. 09.789. 9 ad.

Buenaventura, Chocd, W. Colombia, March 11, 1899. E, André

coll.

Besides the above, I have examined in the Tring Museum

three males and three females obtained at Juntas, Rio Dagua, by

Messrs. Rosenberg and Raap, and in the British Museum

a single female from Remedios, collected by T. K. Salmon.

Dr. Hartert * has already alluded to this form, and, after studying

large series from various localities, I have no hesitation in

separating the Pacific birds from true MZ, s. surinamensis.

The typical race is confined to Surinam, French and British

Guiana, extending westwards to the Caura Valley, Eastern

Venezuela, while JM. s. pacifica ranges from Western Ecuador

through Colombia north to Veragua. The females of both forms

have the sides of the head, the foreneck and chest uniform

ochraceous, thereby differing very conspicuously from the

Amazonian race J. s. multostriata Scl.t.

** Nov. Zool. ix. 1902, p. 612.

+ Cfr. Hellmayr, Noy. Zool. xvii, 1910, p. 345. :

1160 MR. C. E, HELLMAYR. ON THE,

Characters and range of the two races are as follows :—

(a) M. SURINAMENSIS SURINAMENSIS Gm.

Sitta surinamensis Gmelin, Syst. Nat. 1, i. p. 442 (1788—

based on: “ Surinam Nuthatch,” Latham, Gen. Syn. 1, 11. p. 654,

pl. 28 (= @ ): Surinam).

Formicivora quadrivittata (ex Lichtenstem MS.) Cabanis,

Arch. f. Naturg. 13, i. p. 227 [1847.—Guiana (Schomburgk)}.

Myrmotherula surinamensis Sclater, Cat. B. Brit. Mus. xv.

p. 231 (part.: m-g, Brit. Guiana; 2, ata ee Salvin, Ibis,

1885, p. 425 (Camacusa) ; Berlepsch & Hartert , Nov. Zool. ix.

1902, p. 73 (Caura); Berlepsch, Nov. Zool. xy. p. 154 (Cayenne).

Adult. Bill short and slender; white tips to external rectrices.

restricted, not more than 14 to 3 mm. wide on outermost pair.

Upper back, in both sexes, mostly black, variegated with narrow

white edges, base of feathers extensively white, forming a large,

concealed dorsal patch. Female with upper part of the head deep

tawny, crown and nape broadly streaked with deep black; sides

of the head, throat, foreneck and chest bright ochraceous, middle

of throat sometimes paler, ochraceous-buff or buff: rest of lower

parts buff, flanks shaded with greyish. On the sides of the neck

and chest there are sometimes a few faint dusky shaft-lines, but

in most specimens these parts are quite uniform.

Hab. Surinam: Paramaribo (Chunkoo: Tring Museum) ;

French Guiana: Approuague, Ipousin (Cherrie: Tring Museum),

Saint-Jean-du Maroni (Ze Moult: Munich Mus.) ; British

Guiana : Camacusa, Takutu R., Carimang R. (Whitely). Hast

Venezuela, Caura Valley: Nicare, La Pricion, Suapuré, Mato R.

(André, Klages: Tring and Munich Mus.) *

Material. 22 specimens. Birds from various localities measure

as follows :—

Two ¢ gad. Camacusa, British Guiana.— Wing 50, 52; tail 25,

26; bill 14, 15 mm.

Two ¢ ¢ ad. Paramaribo, Surinam.—Wing 50, 52; tail 25,

253; bill 14 mm.

Three ¢ g ad. French Guiana.—Wing 49-51; tail 24-25 ;

bill 14-15 mm.

Five ¢ gd ad. Caura R., Venezuela.—Wing 52; tail 25- a

bill 14-15 mm.

Two 9? 9 ad. Surinam.—Wing 49; tail 243-25; bill 14 mm.

One 2 ad. Carimang R.., Bulbine Ghote, —Wing 502; > tal 26%

bill 143 mm.

Two 9 9 ad, French Guiana.—Wing 49, 50; tail 24, 25 ;

bill 14 mm.

Five 9 9 ad. Caura R., Venezuela— Wing 49-50; tail 25—

27; bill 143-15 mm.

* Tt must remain doubtful, in the absence of females, whether the single male from

Munduapo, Upper Orinoco (ef. Nev. Zool. ix. p. 73) is referable to MW. s. surinamensis -

or to M. s. multostriata.

BIRDS OF WESTERN COLOMBIA, 1161

(6) M. suRINAMENSIS PaciFIcA Hellm.

WM. surinamensis (nee Gmelin) Cassin, Proc. Acad. N. Sci. Philad.

1860, p. 190 (R. Truando, N.Colombia); Lawrence, Ann. Lyc. N. H.

N.Y. vii. 1861, p. 293 (Isthmus of Panama); Sclater & Salvin,

P.Z.S. 1864, p. 356 (Lion Hill, Panama); Sclater & Salvin,

P.Z.S. 1879, p. 525 (Remedios); Berlepsch & Taczanowski,

P. Z.S. 1883, p. 564 (Chimbo, 8.W. Ecuador); Sclater, Cat. B.

Brit. Mus. xv. p. 231 (part.: a-g,j, k, 1); Salvadori & Festa,

Boll. Mus. Zool. Torino, xiv. no. 362, 1899, p. 29. (part.: Peripa,

W. Ecuador) ; Hartert, Nov. Zool. ix. 1902, p. 612° (San Javier,

Pambiltr, N.W. Ecuador); Goodfellow, Ibis, 1902, p. 64

(S. Nicolas, Gualea, W. Ecuador).

“ Myrmotherula ?” Sclater, P.Z.S. 1860, p. 294 (Esmeraldas,

N.W. Keuador).

Adult. Bill strong and heavy ; white tips to external rectrices

larger, varying from three to five millimetres in width on the

outermost pair. Upper back, in the male, much less variegated

with black, the white dorsal patch much less extended. Female

with upper part of the head much lighter, tawny ochraceous, crown

and hind neck with narrow, rather indistinct, dusky olive shaft-

streaks; upper back mainly pale grey, rather sparingly spotted

with black and edged with white, but without any trace of the

white dorsal blotch. Sides of the head and lower parts exactly as

in the female of MW. s. surinamensis.

Hab. Western Ecuador: Peripa (Festa), Intac (Buckley),

San Nicolas, Gualea (Goodfellow d& Hamilton), Chimbo (Siemi-

radzki), Esmeraldas (raser), San Javier, Pambilar (Plemming &

Miketta). Western Colombia: Buenaventura (André), Guineo,

Sipi (Palmer), Juntas (Rosenberg, Raap), Naranjo (André),

Remedios (Salmon), R. Truando (Wood). Isthmus of Panama:

Lion Hill (McLeannan), San Pablo (Salvin), Veragua (Arcé).

| Also in Bogoté coll. |

Material. 41 specimens. Birds from various localities average

as follows :—

Hight dg gd ad. W. Ecuador.—Wing 51-53; tail 28-30 ;

bill 16 mm.

Six ¢ dad. W. Colombia, Chocé district—Wing 523-54 ;

tail 29-314; bill 153-164 mm.

Two § 6 ad. Panama (Lion Hill)—Wing 51; tail 27, 28;

bill 153, 16 mm.

Thirteen 9 @ ad. W. Heuador.—Wing 49-52; tail 28-302 ;

bill 15-16 mm.

Hight ¢ 9. W. Colombia, Chocé district—Wing 50-52;

tail 28-303 ; bill 143-16 mm.

One @. Remedios, N.W. Colombia.—Wing 523; tail 304;

bill 15 mm.

-Two 2 Q@ ad. Bogoté.—Wing 52,53; tail 30,31; bill 15 mm.

One Q ad. Panama (Lion Hill)—Wing 50; tail 273; bill

15 mm.

1162 MR. C. E. HELLMAYR ON THE

118. MyrMornEeRULA FULVIVENTRIS VIDUATA Hart.

[Myrmotherula fulviventris Lawrence, Ann. Lyc. N. H. N.Y.

vil. p. 468 (1862.— Panama). |

Myrmotherula viduata Hartert, Nov. Zool. v. p. 492 (1898.—

Cachabi, N.W. Ecuador ; @ ).

M. fulviventris viduata idem, 1. c. ix. p. 612 (erit.).

M. ornata® Cassin, Proc. Acad. N. Sci. Philad. 1860, p. 191

(Rio Truando).

COE ?” Sclater, P. Z.S. 1860, p. 294 (Esmeraldas, N.W.

Keuador).

M. fulwiventris (nec Lawrence) Sclater & Salvin, P. Z.S. 1879,

p- 525 (Remedios, Antioquia) ; Wyatt, Ibis, 1871, p. 331

(Naranjo, near Bucaramanga); Salvadori & Festa, Boll. Mus.

Zool. Torino, xiv. no. 362, 1899, p. 29 (Peripa, W. Ecuador).

No. 2512. ¢ ad. El Tigre, 4.11.09.—Wing 53; tail 34;

bill 143 mm.

No.. 2426. ¢ juv. Noévita, 22.xi.08.—Wing 514; tail 38;

bill 14 mm.

No. 2471. 2 ad. Noanama, 13.1.09.—Wing 53; tail 38;

bill 143 mm.

“Tris dark brown, feet blackish, bill black, mandible grey.”

These specimens as well as several Bogota skins agree perfectly

with a series from W. Kcuador, including the type, kindly for-

warded by Dr. Hartert from the Tring Museum. JZ, f. viduata

differs from typical fulviventris in having the upper parts

decidedly more brownish or rufescent brown, this being especially

noticeable on the crown and upper tail-coverts; the edges to the

quills and rectrices are also more reddish, and the females have

the belly of a darker fulvous colour. J. f. fulviventris is known

to me from seven skins from Nicaragua (Rio Grande) and

Eastern Costa Rica (Carrillo, Reventazén, Matina R.), but Ihave

not yet seen any from the type locality.

M. f. viduata apparently ranges all over the Pacific lowlands,

up to about 2000 feet altitude, from the Rio Truando in the

north to the neighbourhood of Guayaquil, S.W. Ecuador, in

the south. There is an adult male in the Tring Museum from

Ana Maria (not far from Guayaquil), procured by Herr G. von

Buchwald in December 1905.

119. MyRMOTHERULA AXILLARIS MEZLNA Sel.

[Myrmothera axillaris Vieillot, Nouv. Dict. xii. p. 113 (1817.—

“ Guyane”).

Formicivora melena Sclater, P. Z. 8. 1857, p. 130 (Oct. 1857.—

Bogota).

Myrmotherula melena Sclater & Salvin, P. Z.S8. 1879, p. 525

(Neche, Antioquia); Cassin, Proc. Acad. N. Sci. Philad. 1860,

p- 191 (R. Truando).

No. 2416. ¢ ad. Névita: 19.xii.08.—Wing 53; tail 35;

bill 14 mm.

“Tris brown, feet and bill black.”

Agreeing with Bogota skins.

BIRDS OF WESTERN COLOMBIA. 1163

120. MyYRMOTHERULA SCHISTICOLOR SCHISTICOLOR Lawyr.

Formicivora schisticolor Lawrence, Ann. Lyc. N. H. N. Y. viii.

p- 173 (1865.—Turrialba, Costa Rica).

Myrmotherula schisticolor Hellmayr, Verhandl. zool.-bot. Ges.

Wien, lii. 1903, p. 210 (crit.).

No. 3765. 2. Siaté, Rio Siaté: 21.ix.08.—Wing 57; tail

39; bill 12 mm.

“Tris dark brown, feet plumbeous, bill black, mandible horn-

coloured.”

This bird agrees well with females from Costa Rica (Rio

Naranjo, Boruca), Chiriqui (Boquete), and Western Ecuador,

except in being slightly more greyish, less brownish, on the upper

parts. I have not seen male examples from W. Colombia, but

several from W. Ecuador, which I have before me, I am unable

to separate from typical Costa Rican skins.*

On the other hand, in the Sierra Nevada de Santa Marta,

N. Colombia, and in the mountains of Northern Venezuela

(Cumbre de Valencia, Cuman4), a well-marked geographical race,

M. schisticolor sancte-marte Allen, takes its place. The male

may be immediately recognized in having the black colour

restricted to the throat and middle of the foreneck, while in

M. s. schisticolor it extends over the breast down to the upper

abdomen. The females are not always distinguishable, though

as a rule those of sancte-marte have the back of a purer olive-

grey, without any brownish tinge. The Munich Museum

possesses a good series of this rare form from the Cumbre de

Valencia and the mountains of Cumana, Venezuela. The two

males from 8. Esteban, Venezuela, in the British Museum, Goering

coll.f, are likewise referable to W/. s. sancte-marte.

121. FoRMICIVORA QUIXENSIS CONSOBRINA Scl.

[| Thamnophilus quixensis Cornalia, Vertebr. Syn. Osculati Coll.

p. 12 (1849.—Quixos, Eastern Ecuador). |

Formicivora consobrina Sclater, P. Z. 8. 1860, p. 279 (1860.—

Babahoyo, S.W. Ecuador); Sclater & Salvin, P. Z. 5. 1879, p. 525

(Pocune).

F, quixensis (nee Cornalia) Cassin, Proc. Acad. N. Sci. Philad.

1860, p. 190 (R. Truando).

No. 1963. gd ad. San Joaquim, Bahia del Chocé, 3.viii.08.—

Wing 48; bill 12 mm.

No. 2142. S$ ad. Sipi: 1.x.08.—Wing 48; tail 45; bill

13 mm.

* Carriker (Ann. Carnegie Mus. vi. nos 2-4, 1910, p. 609), following Mr. Ridgway’s

lead, calls the Costa-Rica form MW. menetriesi schisticolor. As a matter of fact,

however, the bird long known under the name MW. menetriesii has nothing to do with

menetriesii of D’Orbigny, as I have clearly demonstrated in the paper quoted above.

M. menetriesii, D’Orb. belongs to quite a different section, being a close ally of

M. cinereiventris Scl. & Salv. Cfr. Bull. Soc. Philom. Paris (9) viii. 1906, pp. 51-2;

Nov. Zool. xiv. 1907, pp. 69-70.

+ M. sancte-marte Allen, Bull. Amer. Mus. xiii. p. 160 (1900.—Valparaiso,

Santa Marta).

t M. menetriesi (errore) Sclater, Cat. B. xv. p. 240 (part.; 5!, c’).

1164 MR. C. E. HELLMAYR ON THE

Nos. 2277, 2327. ¢@ gad. Novita: 13, 23.x1.08.— Wing 48,

49; tail 47, 46; bill 132 mm.

Nos. 2276, 2310. 9 Q ad. Névita: 13, 19.x1.08—Wing 48;

tail 45; bill 13 mm.

No. 2143. @ juv. Sipi: 1.x.08—Wing 47; tail 47; bill

12 mm.

‘“‘ Tris dark brown, feet and bill black.”

The series agrees with specimens from Western Ecuador. One

of the females (no. 2143) approaches /. g. bowcardi Scl., from

Central America, in the paleness of the under parts, yet it can

easily be distinguished by its much smaller, narrower bill, and

much longer white tips to the outer rectrices.

F. q. consobrina is peculiar to W. Ecuador and W. Colombia,

where it inhabits the hot, forest-covered low country from sea-

level up to about 1500 feet.

122. RAMPHOCENUS CINEREIVENTRIS CINEREIVENTRIS Sel.

Rhamphocenus cinereiveniris Sclater, P. Z.S. 1855, p. 76,

pl. Ixxxvi. (June 1855.—‘ Pasto,” 8. Colombia, Delattre coll.).

No. 2506. ¢ ad. Novita: 29.1.09—Wing 53; tail 31;

bul 18 mm.

No. 2102. 9 ad. Sipi: 21.1x.08— Wing 52; tail 32; bill

19 mm.

No. 1976. ¢g imm. Guineo, Rio Calima: 7.viii.08.—Wing 54;

tail 35; bill 18 mm.

“Tris dark brown, feet blue, maxilla black, mandible grey.” __

In addition we have an adult male obtained by Mr.Hugéne

André at El Paillon, near Buenaventura, May 6, 1899.

The specimens from Western Colombia and fourteen others

from various localities in Western Ecuador (Chimbo, 8. Javier,

Cachyjacu, Lita, Rio Verde) agree perfectly with Sclater’s

original description, having a very distinct, dusky brown post-

ocular streak, and the under parts of a rather clear cinereous

with whitish admixture along the middle line. The type-speci-

men is said to have been obtained at Pasto, 8. Colombia, alt.

8000 feet, which is certainly erroneous. JW. c. cinereiventris 1s

exclusively restricted to the low country and slopes of the

Western Cordillera between sea-level and about 3000 feet.

RR. c. semitorquatus Lawr,*, of which | have examined a good

series from Eastern Costa Rica (Carrillo) and Chiriqui (Boquete),

may easily be distinguished in lacking the brown postocular

stripe and in having the belly rather darker cinereous with very

little, if any, whitish suffusion in the middle,

A third geographical race of this group inhabits Hastern

Ecuadory. Jt resembles the Central American form in the

absence of the dusky postocular streak, but the upper parts are

darker and the sides of the head much deeper coloured. I do not,

* Rhamphocenus semitorquatus Lawrence, Ann. Lyc. N. H. N.Y. vii. p, 469

(1862.—Panama).

+ R. cinereiventris (nec Sclater, 1855) Sclater, Cat. B. Brit. Mus, xv. p, 262

(Sarayacu, East. Ecuador).

BIRDS OF WESTERN COLOMBIA. 1165

however, feel justified in bestowing a name upon this subspecies,

having seen but a few skins in not very good condition.

123. CERCOMACRA TYRANNINA RUFIVENTRIS Lawr.

| Pyriglena tyrannina Sclater, P. Z.S. 1855, p. 90, pl. xeviil.

(1855.—Bogota coll.). |

Disythanunus (sic) rufiventris Lawrence, Ann. Lyc. N. H. N.Y.

vill. p. 131 (1865.—New Granada, line of Panama Railroad—

descr. g juv.); cfr. Salvin, Ibis, 1874, p. 316.

Cercomacra crepera Bangs, Auk, xviii. p. 365 (1901.—Divala,

Chiriqui).

No. 1997.. g ad. Boca de Calima, Rio San Juan, 18.viii1.08.—

Wing 67; tail 58; bill 163 mm.

No. 1981. 9 imm. Guineo, R. Calima: 8.vii1.08.— Wing 60 ;

bill 16 mm.

No. 2387. 9 ad. Noévita: 10.xi1.08.—Wing 59; bill 163 mm.

No. 2742. 9 ad. Pueblo Rico, 5200 ft., 8.1x.08.—Wing 62;

tail 67 (!); bill 165 mm.

“Tris dark brown, feet blue-grey, bill black, mandible horn-

coloured in females.”

These birds strictly belong to the dark western form separated by

Mr. Bangs as C. crepera. Mr. Carriker * has already alluded to

the variability of its characters, basing his conclusions upon Costa

Rica skins. On comparing the series in the Munich Museum,

it becomes at once evident that two fairly marked races can be

distinguished. Adult males from Guatemala, Costa Rica, Chiriqui,

W. Colombia and N.W. Ecuador are, notwithstanding some

individual variation, much darker, more slaty blackish, both on

upper and lower parts than specimens from Bogota (topotypes),

the upper Orinoco, British Guiana, and North Brazil (Para, Rio

Negro). The females also are somewhat darker ferruginous

underneath. The single male from Calima is slightly paler

grey on the belly, but similar specimens I have seen from

S.W. Costa Rica, while several from N.W. Ecuador (San Javier)

in intensity of coloration are practically identical with topo-

typical Chiriqui-skins. Mr. Bangs (J. c. p. 366) refers the birds

from Loma del Leon (= Lion Hill), Panama, to typical tyrannina.

I doubt, however, the correctness of this view, since two males

from Monte Oscuro, near Panama City, though not extreme

examples of crepera, are certainly much nearer that form than to

tyrannina. ‘Therefore, it seems to me that Lawrence’s term

rufiventris, founded upon an immature male from Lion Hill, must

take precedence over crepera.

According to my views, the range of the two subspecies would

be as follows :—

(a) C. tyrannina tyrannina (east of the Andes): E. Colombia

(Bogota, Bucaramanga, etc.); Wenezuela: Orinoco (Munduapo)

and Caura Valley; British Guiana; Northern Brazil: Rio Negro

down to Manaos; Obidos, north bank of Amazons; Para district.

* Ann, Carnegie Mus. v, 1908, p. 8; 7.c. vi. 2-4, 1910, p. 618.

1166 MR. C. E. HELLMAYR ON THE

(b) C. tyrannina rufiventris Lawr. Central America from

Guatemala southwards, Western Colombia and Western Kcuador,

south to Chimbo.

124, CercoMACRA NIGRICANS NScl.

Cercomacra nigricans Sclater, P. Z. 8. 1858, p. 245 (Nov. 1858.—

Santa Marta * (type) and Bogots-coll., Colombia ; descr. ¢ imm.) ;

Sclater & Salvin, P. Z. 8. 1879, p. 526 (Remedios, Antioquia) ;

Berlepsch, Journ. f. Orn. 1884, p. 308 (Bucaramanga).

Pyriglena naculicaudis Sclater, op. at. p. 247 (Nov. 1858.—

“Trinidad,” errore; descr. ¢ ad.).

No. 1989. ¢S ad. Mouth of Calima, Rio San Juan, 13.viii.08.—

Wing 67; tail 67; bill 18 mm.

“ Tris dark brown, feet blue, bill black.”

This bird is in the “ maculicaudis” plumage, viz. glossy black

all over, with the campterium, a large dorsal patch, distinct edges

to the wing-coverts, and long apical spots to the outer rectrices

white, and agrees with other adult males from N.W. Ecuador,

and several Bogota skins. Berlepsch & Hartert have shown

that C. maculicaudis and C. nigricans are merely different sexes

of the same species, and the examination of a large series of

sexed specimens has convinced me that this view is perfectly

right. It should also be borne in mind that the supposed two

species have exactly the same range, extending from the Isthmus

of Panama south to Western Ecuador. Mr. Cherrie took three

specimens at Altagracia, Orinoco, two of which, marked “ g ” by

the collector, are in the ‘“ maculicaudis” phase, while the third,

sexed as “92,” is even greyer, both above and below, than the

type of nigricans. . The locality “Trinidad” given for P. maculi-

caudis is certainly erroneous. I have examined the type and

found it practically identical with adult males from Panama and

Bogota. The preparation furnishes no clue as to its origin. The

specimen appears to have been remade, though it looks somewhat

like the skins imported from Northern Colombia (Baranquilla,

Cartagena, etc.).

C. carbonaria Scl. & Salv.t, from the Upper Rio Branco in

Northern Brazil, is quite distinct, although nearly allied. The

male differs in having the upper parts sooty grey ; the throat and

breast much duller black, shading into sooty grey on the flanks ;

in the shorter white tips of the rectrices, as well as in having

distinct, though narrow, white apical edges to the secondaries.

The female, too, is quite different, having the throat white freckled

with dusky, and the belly bright ochraceous with the middle line

whitish. C.carbonaria also has a much narrower and some-

what shorter bill. So far it has been met with only by Natterer,

* Although not taken in recent years by any collector, C. nigricans may yet be

found in the Santa Marta district, for there is a specimen from Calamar, lower

Magdalena (August 5, 1898) in the collection of H.R.H. The Princess Therese of

Bavaria.

+ Nomencl. Av. Neotrop. p. 161 (1873.—Rio Branco, N. Brazil).

BIRDS OF WESTERN COLOMBIA. 1167

who obtained a large series at Forte do Joaquim, on the confines

of British Guiana.

125. CERCOMACRA BERLEPScHI Hart.

Pyriglena berlepschi Hartert, Bull. B. O. C. vii. p. xxix (1898.—

Cachabi, N.W. Ecuador; = ¢ ad.).

Thamnophilus cachabiensis Hartert, 1. c. p. xxix (1898.—

Cachabi, N.W. Ecuador; = @).

Cercomacra berlepschi Hartert, Nov. Zool. ix. 1902, p. 612

(crit.).

No. 1958. g ad. 8. Joaquim, Bahia del Chocd, 1.vi1.08.—

Wing 68: tail 46; bill 194 mm.

No. 2089. gad. Near Sipi, 200 feet alt., 18.1x.08.— Wing 69 ; |

tail 474; bill 19 mm.

No. 1966. Q@ ad. 8S. Joaquim, 4.vii.08.— Wing 67; tail 43 ;

bill 19 mm.

‘‘ Tris dark red, feet and bill black.”

The males are uniform deep black, with a large, concealed

white interscapular patch. The female is rather duller black,

the abdomen slate-blackish, and the feathers of the throat, fore-

neck, middle of the breast, as well as the lesser and median upper

wing-coverts, show very distinct, though sometimes slight, white

apical spots or edges. Dr. Hartert has already pointed out that

these two types of coloration, originally described as different

species, were merely male and female of one and the same species.

From the evidence at hand, there can be no doubt that this view is

correct.

The specimens from Chocé agree in every respect with the

series from N.W. Ecuador in the Tring Museum*. (C. berlepschi

is somewhat difficult to place. In general form it closely re-

sembles some of the short-tailed species, e. g. C. tyrannina,

but the tail is much shorter and slightly rounded instead of

being graduated, the wing relatively longer, and the bill longer

as well as heavier. Although it might some day be found

necessary to create a new genus for its reception, it would be

unwise to do so without a thorough study of all the related

groups.

126. HypocnemMis N&ZVIOIDES Lafr.

Conopophaga nevioides Lafresnaye, Rev. Zool. x. p. 69 (1847—

no locality given; type in Mus. Acad. Philadelphia, coll.

Delattre—we fix as type locality Panama whence the Derby

Museum (Liverpool) possesses two specimens obtained by De-

lattre ; cfr. Sclater, P. Z.S. 1858, p. 254).

Hypocnemis nevioides Cassin, Proc. Acad. N. Sci. Philad. 1860,

p. 190 (Falls of the Truando).

* In this connection a misleading error may be corrected. Hartert (7. c.) gives

the length of the wing for P. berlepschi, as 44 to 46 mm. However, five adult males

from N.W. Ecuador, including the type, in the Tring Museum present the following

dimensions ; wing 67-683 ; tail 48-47; bill 19-20 mm. The females (5) are slightly

smaller; wing 64-68 ; tail 41-46; bill 18-19 mm.

1168 MR. GC. E. HELLMAYR ON THE

No. 2603. g ad. Condoto: 16.iv.09.—Wing 638; tail 38;

bill 16 mm. ;

Nos, 2341, 2491. 9 Q ad. Novita: 26.x1.08, 25.1.09.— Wing

62, 61; tail 37, 34; bill 16 mm.

Nos. 1977, 1999. 2 9 imm. Guineo: 7.viii; mouth of Caiima ;

18.vi1i.08.— Wing 64, 62; tail 37, 35; bill 16 min:

“Tris brown, feet blue-grey, bill Tage, lower mandible of

females grey.”

These specimens as well as others from N.W. Eeuador (San

Javier) agree perfectly with typical Panama examples. Skins from

Miravelles, Costa Rica, have, as a rule, the wing a trifle larger,

and have the back of a clearer, brighter chestnut, but these

variations are so iInconstant that it appears to me highly im-

probable that the northern form, H. n. capnitis Bangs* can be

maintained. This view is shared by Mr. Carriker 7, who collected

a large series of this species in Costa Rica.

Birds from different localities measure as follows :—

Five adult males from Miravelles, Costa Rica: wing 66-67;

tail 34-37 mm.

Two adult males from Panama: wing 65, 653; tail 36, 37 mm.

One adult male from W. Colombia: wing 63; tail 38 mm.

Three adult males from N.W. Ecuador (8. Javier): wing

63-65; tail 35-374 mm.

Four adult females from Miravelles, Costa Rica: wing

63-65 ; tail 33-36 mm.

Two adult females from W. Colombia: wing 62, 61; tail 37,

o4 mm.

One adult female from N.W. Ecuador (S. Javier): wing 61 ;

tail 36 mm.

Both Mr. Bangs and Mr. Carriker give ‘“‘ Pasto, Cauca valley,

Colombia” as the type locahty, but this is an obvious error.

First of all, Lafresnaye does not say where his type came from,

and secondly the occurrence of H. nevioides in the vicinity

of Pasto (which hes high up in the mountains at an elevation

of nearly 8000 feet) is utterly impossible, the species being

restricted to the humid, tropical lowlands, from sea-level to about

2000 feet. Delattre most probably obtained the type at

Panama, whence there are two of his skins in the Derby Museum

(see above).

127. MyYRMELASTES IMMACULATUS IMMACULATUS Lafyr.

Thamnophilus immaculatus Lafresnaye, Rev. Zool. viii. p. 340

(1845—“‘ad Bogotam,” descr. ¢ 2).

_ Myrmeciza berlepschi (nec Ridgway) Bangs, Proc. Biol. Soc.

Wash. xxiii. p. 73 (Palmar, Pavas, La Maria, W. Colombia).

No. 2818. g ad. Pueblo Rico, San Juan slopes, 28.x.09.—

Wing 84; tail 77; bill 22 mm.

* Proc. Biol. Soc. Wash. xix. p. 107 (1906.—Miravelles, Costa Rica).

7 Aun. Carnegie Mus. vi. 1910, p. 619.

BIRDS OF WESTERN COLOMBIA. 1169

[Tring Museum. @ ad. . Primavera, Cordillera occidental,

1904. Raap coll. No. 390.—Wing 79; tail 79; bill 20 mm. }

Mr. Ridgway * has correctly pointed out the distinctness of

M. immaculatus, from Bogota, and IM. 2. berlepschi Ridgw., from

Western Ecuador, though some of his conclusions prove to be

not well-founded. First of all, d/. 2. berlepschi does not occur

anywhere in Colombia, the specimens from Bogota mentioned by

Berlepsch & Taczanowski t, which were kindly forwarded to me

by the Count, being clearly referable to true IZ. immaculatus. The

female from Primavera, W. Colombia, is practically identical with

several Bogota skins in the Berlepsch Collection, In all these

females the lower parts (except the blackish upper throat) are

rufescent brown, mottled with dull smoky grey on the foreneck

and in the middle of the breast, while there is only a narrow

white stripe on the bend of the wing.

Sixteen females from Western Ecuador (Chimbo, Paramba,

Lita, Gualea, Bulan, ete.) have a much larger, heavier bill, the

forehead is more scantily feathered, the bend of the wing shows

a large white patch, and the lower parts are much brighter

rufous brown. The chin as well as the sides of the head are

blackish, exactly as in J/. immaculatus from Colombia. Seventeen

adult males from Western Kcuador also differ from the Colombian

ones in their larger bill, more scantily feathered forehead, and

in having much more white on the shoulders. The general

plumage is sometimes, though not always, deeper black.

M. immaculatus zeledoni Ridgw. t, lately united to the Colom-

bian form by Mr. Carriker§, appears to me to be much more

closely allied to IM. 7. berlepschi. In fact, on comparing three

adult males and two females from Costa Rica (Cariblanco de

Sarapiqui) with the large series from Keuador, I find the differences

not very pronounced. All that can be said is that the northern

birds have the white shoulder-patch slightly smaller. The females

otherwise agree with those from Keuador. More material from

Costa Rica may even show the two forms to be inseparable.

128. MyYRMELASTES EXSUL MACULIFER Hellm.

[Myrmeciza exsul Sclater, P. Z. 8. 1858, p.540 (1859.—Panama:

Delattre). }

Myrmelastes exsul maculifer Hellmayr, Nov. Zool. xiii. p. 340

(1906.—Paramba, N. Ecuador).

Myrmeciza exsul (nec Sclater) Cassin, Proc. Acad. N. Sci. Philad.

1860, p. 191, no. 98 (= 3), 99 (= 2) (Turbo); Sclater & Salvin,

P.Z.8. 1879, p. 526 (Rio Neche, Antioquia).

Myrmelastes cassini Ridgway, Proc. Biol. Soc. Wash. xxi. p.194

(1908.—Turbo, N. Colombia).

* Proc. Biol. Soc. Wash. xxii. 1909, p. 74.

+ P. Z. S. 1883, p. 565.

£ Myrmeciza zeledoni Ridgway, Proc. Biol. Soc. Wash. xxii. p. 74 (1909.—

Guayabo, E. Costa Rica).

§ Ann. Carnegie Mus. vi. 1910, pp. 618-9.

1170 MR. C. E. HELLMAYR ON THE

Nos. 2082, 2145, 2178. g$ dad. Sipi: 12.ix, 2, 12.x.08.—

Wing 67-69; tail 444-47; bill 18 mm.

No. 2234. gad. Rio Cajén: 3.xi.08.—Wing 68; tail 45;

bill 18 mm.

No. 2488. gad. Névita: 28.x11.08.—Wing 69; tail 433 ; bill

19 mm.

Nos. 2017, 2018, 2038. 9 2. Noanama: 26, 31.viii.08.—

Wing 63; tail 40-423; bill 173-18 mm.

Nos. 2179, 2353) 9) adi, od) juv.. )\Sipiy 12x08); INovatare

30.xi.08.— Wing 63, 66; tail 42, 44; bill 18, 19 mm.

“Tris dark red, feet grey, bill black.”

When discussing the status of this species and its northern

limits I alluded to the paler. less rufescent brown coloration of the

specimens from Neche, Northern Colombia, in the British Museum

(cfr. Nov. Zool. xiii. p. 342), and two years later Mr. Ridgway

separated two similar skins from Turbo under the name of

M, cassini*. The present series, however, shows considerable

variation in the colour of the back. Six of the specimens have

the back of exactly the same deep rufous or vandyke- brown shade

as a large series from N.W. Ecuador, while the four remaining

ones are somewhat lighter, more mars-brown above, like the two

examples from Neche, Antioquia. Two adult males and one

female obtained by Mr. Rosenberg on the Rio Dagua in 1895,

agree again with the types from Paramba.

Neither is there any constant difference in the coloration of the

head and underparts. Four males—two from Paramba, one from

Sipi, one from Névita—are decidedly slaty blackish, while the

majority of the skins, both from W. Ecuador and from Choco,

have the chest and belly paler, slate-grey.

Therefore, I cannot admit d/. cassini as a valid form unless

the Turbo birds be distinguished by other characters than those

given in the original description. In addition to the above ten

specimens, the Munich Museum possesses seven males and four

females from N.W. Ecuador; and I have examined ten more

from W. Ecuador, three from the Rio Dagua, in the Tring

Museum, and the couple from Neche in London.

M. e. maculifer, as understoood by me, is another Pacific

type, its range being limited to the forest districts of Western

Ecuador and Colombia.

129. ANOPLOPS BICOLOR DAGU& Hellm.

[Pithys bicolor Lawrence, Ann. Lyc. N. H.N. Y. vii. p. 6

(May 1863.—Panama Railway). |

Gymnopithys bicolor dague Hellmayr, Bull. B.O. C. xvi. p. 83

{1906.—EI Paillon near Buenaventura, Chocé, W. Colombia ; type

in Tring Mus.).

* Mr. Ridgway also mentions a female from Cascajal, Coclé (Panama), as

belonging to this form. There must, however, be some mistake with regard to the

locality, since three adult males from that place in the Tring Museum are clearly

referable to M. easul exsul. Heyde’s localities are not always trustworthy, and, as

he also got many specimens from Western Colombia (Ndévita etc.), an error in

labelling the bird might have occurred.

BIRDS OF WESTERN COLOMBIA. 1171

No. 2336. gad. Noévita: 25.xi.08.—Wing 78; tail 51; bill

20 mm.

Nos. 2564, 2565. ¢ 2 ad. Juntas, R. Tamand: 9.111.09.—

Wing 77, 76; tail 48, 49; bill 20, 193 mm.

“Tris brown, feet blue-grey, maxilla black, mandible grey.”

The specimens are topotypical. The female differs from the

males in having the dark grey colour of the forehead and sides of

occiput less extended and in lacking the blackish border to the

white chest.

A. b. dague is nearest to A. b. bicolor, from Panama, but may

be easily distinguished by the much darker, deep vandyke (rufous)

brown instead of light russet-brown colour of the upper parts,

much darker rufous-brown flanks, larger, stronger bill, and by

the presence of a distinct blackish border to the white chest in

the male sex.

A. b. equatorialis Hellm., from Western Ecuador, agrees with

A. b. daguwe in the dark coloration of the back and flanks, but the

forehead and crown are bright cinnamon rufous, the sides of the

occiput alone being dark cinereous, etc.

The various races of A. bicolor not having been properly

understood hitherto, it is hoped that the subjoined key will aid

ornithologists in identifying specimens of the group. A. leucaspis

Scl. is a near ally of A. bicolor and very likely its Amazonian

(eastern ) representative ; it differs, however, in having the cheeks,

malar region, and pntecion ear-coverts white (instead of black), and

in the possession of a large, concealed, pale cinnamomeous inter-

scapular patch in the female sex.

Key to the species and subspecies.

1, Cheeks, malar region, and anterior ear-coverts white.

Female with a distinct, light cinnamomeous inter-

scapular blotch. {Nasal plumes and whole pileum

bright ferruginous. Lores and narrow sabeenaty

stripe black]... A. leucaspis.

— Cheeks. malar region, “and whole of the ear-coverts

black. Female ‘without a pale interscapular blotch 2.

2. Sides of the occiput dark cinereous, Av Sie con-

trasted with colour of crown ...... 3.

— Sides of the occiput concolor with crown, never

cinereous. [White chest laterally not bordered aby

a blackish stripe]... 5.

3. Forehead and a broad ‘superciliary stripe dark cine-

reous like the sides of the occiput ...... 4.

— Forehead and crown ferruginous, only sides of ‘occiput

dark cinereous ...... : A. bicolor equatorialis.

4. Crown and back light yusset- brown ; ‘sides of body

mars-brown; bill “smaller : 17-183 mm. 58 A. bicolor bicolor.

— Crown and back deep vandyke or dea rufous brow n;

sides of body vandyke brown; bill larger;

ICO arab eee boSenR a eRetpeocuede sameogoncs saecaeend manceds A. bicolor dague.

5. Head above light russet-brown; back umber-brown

or very slightly tinged with rufescent; sides of

body light mars-brown .............0....eee cee cee cece eee A. bicolor olivascens.

— Head above bright ferruginous ; back decidedly cinna-

mon-brown; sides of body rufescent or russet-brown 4. bicolor ruficeps.

1172 MR. C. E. HELLMAYR ON THE

(1) ANopLops LEUCASPIS Scl.

Myrmeciza leucaspis Sclater, P. Z. 8. 1854, p. 253, pl. Ixx.

(1855.—Bogoté, Colombia (type); Chamicuros, Hast. Peru ;

Cobati, Rio Negro).

Pithys leucaspis Sclater & Salvin, P. Z. 8. 1867, p. 576 (Cobati) ;

lidem, I. c. p. 751; iidem, |. c. 1873, p. 276 (Xeberos, Chyavetas,

E. Peru) ; Pelzeln, Zur Orn. Bras. ii. p. 89 (Barcellos, R. Iganna,

R. Uaupés: R. Negro); Taczanowski, Orn. Pér. ii. p. 74 (Tarapoto,

N. Peru).

Gymnopithys leucaspis Salvin & Godman, Biologia, Aves, 11.

p. 222 (Rio Meta, Hast. Colombia).

This species is exclusively found in the forests of Upper

Amazonia, from the eastern slopes of the Colombian Andes

south to Northern Peru. Both Natterer and Wallace obtained it

on the banks of the Rio Negro, while Wheeler transmitted specimens

from the Rio Meta, E. Colombia, to the British Museum. Haux-

well met with it at Chamicuros, Eastern Peru, and KE. Bartlett

procured a series at Chyavetas, Xeberos, and Chamicuros, now in

the Tring Museum. A. leucaspis is also occasionally found in

the trade-collections sent from Bogota to Europe.

Against the statement in the ‘ Cat. of Birds,’ xv. p. 295, I wish

once more to emphasize that it is the female which possesses the

cinnamomeous interscapular blotch. Five sexed males from

Northern Peru (Chamicuros, Chyavetas, Xeberos ; Bartlett coll.)

in Mr. Rothschild’s collection, five from the Rio Negro, obtained

by Natterer, in the Vienna Museum, and one male from Cobati,

R. Negro, Wallace coll. in the British Museum, show no trace of

it; while in five sexed females from the Rio Negro (Natterer ;

Vienna Museum) there is a large conspicuous patch at the base

of the dorsal feathers. Besides the above, I have examined two

males and four females from “ Bogota-coll.” in the collections at

Tring, Munich, and Paris.

(2) ANOPLOPS BICOLOR #QUATORIALIS Hellm.

Pithys bicolor equatorialis Hellmayr, Orn. Monatsber. x. p. 33

(1902.—Lita, N.W. Ecuador).

P. leucaspis (nec Sclater) Hartert, Nov. Zool. v. 1898, p. 495

(Chimbo, W. Ecuador); Goodfellow, Ibis, 1902, p. 65 (Sto.

Domingo, W. Ecuador).

Gymnopithys ruficeps (nec Salvin & Godman) Salvadori & Festa,

Boll. Mus. Zool. Torino, xiv. no. 362, 1899, p. 32 (Santiago,

W. Ecuador); Hartert, Nov. Zool. ix. 1902, p. 613 (Chimbo,

Paramba, 8. Javier, W. Eeuador).

This form, which is confined to Western Ecuador, shares with

A. leucaspis the bright ferruginous colour of the forehead and

crown, but is much darker rufous brown on the back, has the

sides of the occiput dark cinereous, etc., ete. I have examined

twenty specimens from W. Ecuador in the Tring, Vienna, and

Munich Museums. There is never any trace of a pale inter-

scapular patch.

BIRDS OF WESTERN COLOMBIA. 1a

(3) ANOPLOPS BICOLOR RUFICEPS Salv. & Godm.

Gymnopithys ruficeps Salvin & Godman, Biologia, Aves, ii. p. 222

(1892.—* Cauca Valley,” the types are from Neche and Remedios,

Antioquia).

Pithys leucaspis (nec Sclater) Sclater & Salvin, P. Z. 8. 1879,

p- 526 (Remedios, Neche).

This race is evidently confined to the Cauca Valley in the

province of Antioquia, N. Colombia. Besides three of Salmon’s

skins in the British Museum (2) and at Tring (1), I have examined

five Bogota skins * referable to the same form. A. 0. ruficeps

differs at a glance from the preceding subspecies in lacking the

cinereous colour on the sides of the occiput and in the white of

the chest being laterally not bordered with black. It is much

more closely related to A. b. olivascens, but of a more rufescent

colour throughout.

(4) ANopLops BICoLoR DAGUz Hellm.

This well-marked form also has a very limited range, being

known only from the valleys of the Dagua and San Juan Rivers

in Western Colombia. The Tring Museum possesses an adult

male obtained by E. André near Buenaventura, and another from

Juntas, Raap coll. Palmer’s three specimens are the only others

on record. For characters see above.

(5) ANOPLOPS BICOLOR BICOLOR Lawr.

Pithys bicolor Lawrence, Ann. Lye. N. H. N.Y. viii. p. 6 oe

—Panama Railway).

The “typical” subspecies also has a rather limited distribution,

being restricted to Panama and Veragua. I have examined four

skins from the Panama Railroad (McLeannan), one from Chepo,

and one from Santa Fé, Veragua (Arcé), all in the British

Museum.

(6) ANOPLOPS BICOLOR OLIVASCENS Ridgw.

Pithys bicolor olivascens Ridgway, Proc. U. S. Nat. Mus. xiv.

1891, p. 469 (1892.—Santa Ana, Honduras).

This form is more widely distributed, ranging from Hastern

Honduras through Nicaragua, and Costa Rica to Chiri iqui. There

are large series from various localities in the Tring and Munich

Museums.

130. FoRMICARIUS ANALIS DESTRUCTUS Hart.

[Myothera analis Lafresnaye & D’Orbigny, Syn. Av. 1., in Mag.

Zool. el. ii. p. 14 (1837.— Yuracarés, N.E. Bolivia). |

Formicarius analis destructus Hartert, Nov. Zool. v. p. 493

(1898.—Paramba, N.W. Ecuador); idem, 1. c. ix. 1902, p. 614

(N.W. Ecuador; crit.).

F. analis (nec Lafr. & D’Orb.) Sclater, P.Z.S. 1860, p. 294

* One in Tring, two in the Paris Museum, two in the British Museum.

Proc. Zoot. Soc.—1911, No. LX XIX. T®

1174 MR. C; E. HELLMAYR ON THE

(Esmeraldas, N.W. Ecuador); idem, Cat. B. Brit. Mus. XV. p- oe

(part. : Balzar, Santa Rita ; Esmeraldas: W. Keuador).

I. nigr icaypillus (nec Ridgway) Ridgway, Proc. U.S. Mus. xvi.

p- 675 (part.: sp. ew Santa Rita).

F.. destructus Salvadori & Festa, Boll. Mus. Zool. Torino, xiv.

no. 362, 1899, p. 32 (Peripa, W. Ecuador).

Nos. 2508, 2393. ¢ g ad. Névita: 11.xi1.08, 30.1.09.— Wing

90, 86; tail 55, 52; bill 20, 21 mm.

No. 2372. 9 juv. Névita: 5.xi1.08.—Wing 86; bill 18 mm.

‘Tris dark brown, feet and bill black.”

Perfectly identical with topotypical specimens from Paramba,

N.W. Ecuador, the head all round and the hind-neck being sooty

black, abruptly contrasting with the dark rufous brown of the

back.

F. a, destructus is most nearly allied to /. a. nigricapillus Ridgw.

of Eastern Costa Rica. Dr. Hartert * having clearly pointed out

the differences between the two races, I need not dwell any more

upon the subject.

This Yormicarius is confined to the lowlands and foot-hills of

the Pacific coast of Colombia and Ecuador. On the other side of

the Western Cordillera, in the Cauca Valley, another species

or subspecies, /. satwratus Ridgw. tT, takes its place. I have

examined the example from Remedios (Salmon) in the British

Museum, and an adult male obtained by the late J. H. Batty

at Los Jambos (Cauca Valley) in Mr. Rothschild’s Museum at

Tring, which I cannot distinguish from typical Trinidad specimens.

F. satwratus is much paler grey underneath, has the sides of the

neck distinctly cinnamomeous, and the top of the head light

olivaceous brown like the back, besides some minor differences.

131. FoRMICARIUS RUFIPECTUS RUFIPECTUS Salv.

fF. rufipectus Salvin, P. Z. 8. 1866, 73, pl. viii. (1866.—

Santiago de Veragua) ; Salvadori & ines, Boll. Mus. Zool.

Torino, xiv. no. 362 1899, p. 33 (Gualea, W. Keuador) ;

Ménégaux & Hellmayr, Bull. Soe.) Philom: Paris, (9) viii. 1906,

p- 52 (between Ksmeraldas and Pachijal, Oyacachi, W. Keuador) ;

Carriker, Ann. Carnegie Mus. vi. 1910, p. 626 (Rio Cali, W.

Colombia).

No. —. d ad. Pueblo Rico: 16.viii.09.—Wing 99; tail 63;

bill 20 mm. :

No. 2729. 9 ad. Pueblo Rico: 16.viii.09.—Wing 94; tail 59;

bill 214 mm.

“Tris and feet dark brown, bill black.”

These specimens agree well with others from Chiriqui (A. Lara

coll.). The female differs from the male, in addition to its

smaller size, by having the breast somewhat clearer rufous, and

* Nov. Zool. ix. p. 614.

+ Proc. U.S. Mus. xvi. p. 677 (1894.—Trinidad (type) ; Venezuela: S. Esteban ;

N.E. Colombia: Remedios).

BIRDS OF WESTERN COLOMBIA. 1175

the crown dull brownish black, the hind neck only bemg washed

with chestnut, while in the male all the top of the head, from

the forehead to the nape, is dark chestnut-maroon. The sexual

differences noticed by Mr. Carriker, however, do not hold good.

Both specimens have the middle of the belly extensively ochraceous

buff and the flanks dark smoky grey.

This scarce species ranges from Central Costa Rica (Juan

Vinas) to Western Ecuador *, On the eastern (Amazonian)

slopes of the Ecuadorian Andes and in Peru, /. r. rufipectus is

replaced by /. r. thoracicus Tacz. & Berl. f, at once recognizable

by its deep black pileum. I have examined one of the typical

specimens and an adult male from Huaynapata, both in Count

Berlepsch’s collection.

132. PrrrAsoMA ROSENBERGI Hellm.

P. rosenbergi Hellmayr, Rev. Frane¢. d’Orn. ii. p. 51 (April

1911.—Sipi, Rio Sipi).

3 ad. Pileum and nape deep cinnamon-rufous;_ back and

scapulars dull olive, more brownish on the rump, each feather

broadly bordered with black on either side, giving the upper parts

a striped appearance; some of the interscapular feathers white at

the base; upper wing-coverts sepia-brown, washed with russet,

especially on the outer webs, each feather with a small, but

distinct, whitish apical spot, and with a distinct black margin

round the tip; primary coverts uniform blackish brown; remiges

blackish, with the outer web dull russet-brown ; rectrices dusky,

the central ones washed with russet. Lores and broad super-

ciliary stripe, reaching to the sides of the neck, uniform deep

black ; cheeks, subocular and malar region, ear-coverts, chin and

throat ochraceous, rather deeper on the sides of the head; fore-

neck and sides of the breast and flanks dull olive-brownish ;

middle of the breast and abdomen uniform buffy white, this

colour being separated from the ochraceous throat by the dull

brown crescent of the foreneck; under tail-coverts hair-brown,

edged with whitish; axillaries and under wing-coverts dusky

brown, the under primary-coverts tipped with white.

“ Tris dark brown, feet and bill black.”

Wing 97; tail 34; tarsus 45; bill 242 mm.

Type in the Zoological Museum, Munich, No. 09.5759. ¢ ad.

Sipi, Rio Sipi, alt, 150 feet, Sept. 25, 1908. Mervyn G. Palmer

coll. no. 2120.

This species, which is named in honour of Mr. W. F. H.

Rosenberg, of London, who organized Mr. Palmer’s expedition to

Western Colombia, is unfortunately represented by a single spe-

cimen only. ‘The type is a perfectly adult male, as shown by the

* The specimen in the British Museum said to be from Baeza (Eastern Ecuador)

either belongs to F’. x. thoracicus, or else the locality is wrong (Buckley !).

+ F. thoracicus Taczanowski & Berlepsch, P. Z. S, 1885, p. 101 (Machay,

Eastern Ecuador); Salvadori & Festa, Boll. Mus. Zool. Torino, xiv. no. 362, p.33 (S,

José, E. Kcuador); Berlepsch & Stolzmann, Ornis, xiii. 2. 1906, p. 118 (Huavnapata,

S.E. Peru),

Oke

1176 MR. C, E. HELLMAYR ON THE

uniform blackish primary coverts, the whitish apical spots to the

wing-coverts, the black superciliary stripe, etc. Although closely

allied to P. rufopileatum Hart.*, from North-western Ecuador,

it differs from both sexes of that species ft in having no dusky

eross-bands whatever on the lower parts. In P. rufopileatuwm the

male has the under surface of the body, with the exception of

the flanks, regularly barred with black and white, the black

markings of the throat being often broken and less pronounced,

though always present, while only the sides of the head are

ochraceous. The superciliary stripe (uniform black), back and

upper wing-coverts are exactly asin P. rosenbergi. The female

ot P. rufopileatum agrees with the latter in the ochraceous throat,

but can, of course, be easily distinguished by having the feathers

of the superciliary region white, edged with black, and the lower

parts (posterior to throat) bright ochraceous narrowly banded

across with dusky olive. Besides, the back is much more decidedly

brown, and the apical spots to the wing-coverts are bright: buff

instead of white.

133. CoNOPOPHAGA CASTANEICEPS CASTANEICEPS Scl.

Conopophaga castaneiceps Sclater, P. Z. 8. xxiv. p. 47 (1857—

part., type from Bogota, Colombia; = ¢).

C. gutturalis idem, |. c. 1868, p. 574 (Bogota; = @).

No. 3779. @ ad. La Selva, Rio Jamaraya, 4600 ft., 1.x.09.—

Wing 72; tail 39; bill 143 mm,

‘“‘Tris dark brown, feet grey, maxilla black, mandible grey.”

This bird corresponds with Sclater’s description of C. gutturalis.

Tt has the back rather more brownish than a Bogota skin, but

does not differ otherwise. This is, so far as I know, the first

record of C. castaneiceps from the Western Cordillera of Colombia.

In Peru it is replaced by a closely allied form, C. c. b#uwnneinucha

Berl. & Stolazm.f The female of C. c. castaneiceps, in general

coloration, is much hike Grallaricula cucullata Scl., but may

easily be recognized by its white postocular stripe (‘‘ pencil”).

134, ANDRODON ZQUATORIALIS Gould.

Androdon equatorialis Gould, Ann. Mag. N. H. (3) xii. p. 247

(1863. — Ecuador) ; Sclater & Salvin, P. Z. 8. 1879, p. 528

(Remedios, Antioquia).

No. 2129. g ad. Sipi, Rio Sipi: 29.ix.08.—Wing 67; tail

42; bill 40 mm.

Nos. 2074, 2644. 6 d imm. Sipi: 10.1x.08; Tadd: 8.v.09.—

Wing 68, 69; tail 42, 44; bill 37, 40 mm.

“Tris black, feet pink, maxilla black, mandible light yellow.”

The first specimen (No, 2129) has the forehead and crown

* Novit. Zool. viii. p. 370 (1901.—Bultin, N.W. Kcuador).

+ The bird figured in Nov. Zool. ix. 1902, pl. viii. fig. 2 as “juv.” is really

the adult female. This is quite evident from the large series, partly im the Tring

Museum, partly in the possession of Mr. Rosenberg, which I have examined.

t P. Z.S. 1896, p. 385 (Garita del Sol, La Gloria, Chanchamayo, C. Peru).

BIRDS OF WESTERN COLOMBIA. ILI

shining coppery red, passing into golden green on the occiput; in

the two other, apparently immature, birds the forehead is dull

greyish, while the feathers of the crown are dusky, tipped with

bronze-green and edged with reddish bronze. No. 2074 shows a

strong bluish tinge on the nape, completely absent in the other

examples,

With regard to the variation of this species, cfr. Simon, Bull.

Mus. Paris, 1907, no. 1, p. 17.

A. equatorialis is strictly confined to the lowlands and hill-

ranges of Western Ecuador and Colombia, ranging from near

sea-level up to about 2400 feet.

135, THRENETES RUCKERI FRASERI Gould.

[Trochilus Ruckert Bourcier, P.Z.8. 1847, p. 46 (May 1847—

loc. ign.) *.]

Giaucis frasert Gould, Monogr. Trochil. pt. xxiv. pl. 12 (1861.

—KHEsmeraldas, N.W. Ecuador).

Threnetes frasert Hartert, Nov. Zool. v. 1898, p. 493 (Cachabi,

N.W. Ecuador) ; Bangs, Proc. Biol. Soc. Wash. xxiii. 1910, p. 72

(Naranjito, Rio Dagua ; Palmer coll.).

Threnetes ruchert (sic) Boucard, ‘ Humming Bird,’ v. 1895, p. 7

(Rio Dagua).

No. 2417. $ fere ad. Noévita, 150 ft., 19.xii.08.— Wing 60;

tail 38; bill 31 mm.

“ Tris black, feet pink, maxilla black, mandible yellow.”

The differences separating 7’. r. ruckert and T. 7. fraseri are so

slight that their relationship is better expressed by trinomials.

One of the principal characters given by authors for the southern

bird, namely, the lesser extent of the cimnamon-rufous patch on the

foreneck, is of no value at all. Adulé males of fraseri have quite

as much, sometimes even more, rufous on the jugulum than

Central American skins. The only constant points by which to

tell frasert are the rather darker smoke-grey belly, the pure

metallic green upper parts, without any bronzy or golden gloss,

and the much deeper, decidedly bluish-black colour of the median

portion of the lateral rectrices.

Specimens from Western Hcuador have the two central

rectrices dull green, passing to nearly blackish green towards the

tip, while in the male from Névita they are much brighter, uniform

metallic green. In that respect the Choco bird slightly points

towards ruckert from Central America, but otherwise it is

perfectly typical of fraseri.

T. r. fraseri inhabits the forest-covered lowlands of Western

Colombia and Western Ecuador‘, from sea-level up to about

700 feet.

* Bourcier’s rather vague description is possibly not referable to the present species.

The words “gorge et dessous du corps gris-noir bronzé ”’ can scarcely be applied to

T. ruckeri auct., and a re-examination of the type in the Loddiges Collection

seems desirable.

+ Its reported occurrence in Eastern Ecuador (Sarayacu: cfr. Cat. B. xvi. p. 266)

rests on two of Buckley’s skins, no doubt incorrectly labelled.

1178 MR. C. E. HELLMAYR ON THE

136. GLAUCIS HIRSUTA ZNEA Lawr.

| Trochilus hirsutus Gmelin, Syst. Nat. 1, i. p. 490 (1788—ex

Marcgrave: East Brazil). |

Glaucis eneus Lawrence, Proc. Ac. N. Sci. Philad. xix. p. 232

ee fie Vor Rica).

G. columbiana Boucard, Gen. Hum. Birds, p. 402 (end of 1895.

eau Dagua, W. Colombia) ; ; idem, ‘The Humming Bird,’ v. p. 7

(Rio Dagua).

G. hirsuta (nec Gmelin) Sclater & Salvin, P. Z. 8. 1879, p. 528

(Santa Elena).

No. 1985. ¢ (2) ad. Guineo, Rio Calima: 10.viii.08.— Wing

d4L; tail 34; bill 29mm.

a ie is black, feet yellow, maxilla black, mandible Bllevk

grey.”

I agree with Mons. Simon* that the earliest available name

for the small, dark-coloured form of the Pacific coast is the one

given above. Jam unable to find any constant differences, either

in size or in colour, between examples from N.W. Ecuador and

W. Colombia, on the one hand, or those from Central America on

the other.. The uniform cinnamon-brown colour of the under

parts is by no means a constant character of the Western birds,

as already pointed out by Mr. Oberholser ‘.

G. h. cwnea ranges from Costa Rica and Panama south to

N.W. Hcuador (province of Esmeraldas).

137. PH#THORNIS YARUQUI SANCTI-JOHANNIS Hellm.

| Trochilus Yaruqui Bourcier, Compt. Rend. Ac. Sc. Paris, xxxil.

No. 6 (séance 10 Febr. ), p: 187 (1851.—*‘ Les bois tres-chauds

des environs d’Yaruqui,’ W. Kcuador). |

Phaéthornis yaruqui sancti-johannis Hellmayr, Bull. B. O. C.

xxvii. p. 92 (1911.—Condoto).

~ Phethornis yaruqui (nec Bourcier) Cassin, Proc. Acad. N. Sci.

Philad. 1860, p. 194 (R. Truando); Simon & Dalmas, Ornis, x1.

p. 218 (Buenaventura).

Nos. 2128, 2197, 2344, 2598 +. -$ gd ad., ¢ nearly ad. Sipi:

929.ix.; Noanama: 17.x.; Névita: 27.x1.08.; Condoto: 14.iv.09.

—Wing 61-62; tail 55-61; bill 41-43 mm.

Nos. 1987, 2042, 2095. 92 ad, @ 2 imm. Guineo: 11.viil.;

Noanama: 3l.vili.; near Sipi: 19.1x.08.—Wing 58-59; tail

59-63 ; bill 37-40 mm.

“Tris black, feet brown, maxilla black, mandible scarlet.”

Type of subspecies in Zoological Museum, Munich: no. 09.

5807. dad. Condoto, R. Condoto, 150 feet, April 14, 1909.

M. G. Palmer coll. no. 2598.

P. y. sancti-johannis is a very distinct form, differing in many

* Rey. Frang. d’Orn. i. 1910, p. 260.

- + Proc. U.S. Mus. xxiv. 1902, p. 311.

{ Type of subspecies.

BIRDS OF WESTERN COLOMBIA. 1179

important points from P. y. yaruqui of Western Ecuador. The

males may be distinguished by their shorter bill; much broader

and darker, deep ochreous-buff postocular and malar streaks ; and

by having the pale smoke-grey median stripe of the throat and

foreneck much wider and confluent with the smoke-grey abdomen,

which is but slightly spotted with metallic green on the flanks ;

whereas in P. y. yaruqui the under surface is mainly dark metallic

green, with the abdomen and a short stripe along the middle of

the throat sooty blackish. The Chocé birds have, too, the base

of the outer web of the rectrices decidedly washed with greenish.

The females are even more different from the Keuadorian yaruqut.

The under surface is light mouse-grey, the sides of the foreneck

and breast being but sparingly mottled with metallic green, and

the pale median stripe along the throat, foreneck, and chest is

conspicuously mixed with whitish. The deep ochreous-buff post-

ocular stripe is much broader, the white tip to the central rectrices

more extended, but less sharply defined than in the females of

P.y. yaruqui. In the females of both races, the malar streak 1s

white, anteriorly tinged with buff.

Mons. Simon, of Paris, to whom I have submitted our series,

agrees to the distinctness of the Colombian race.

P. y. sancti-johannis replaces the typical form in the tropical

lowlands of West Colombia (from the Truando to Buenaventura),

138. PH#THORNIS SYRMATOPHORUS SYRMATOPHORUS Gould.

Phaéthornis syrmatophorus Gould, Contrib. to Ornith. 1851,

p. 139 (1851.—‘“ Interior of Quito, in Ecuador,” W. Jameson

coll.).

P. syrmatophorus Sclater & Salvin, P. Z. 8. 1879, p. 528

(Santa Elena, Medellin, Antioquia); Simon & Dalmas, Ornis,

xi. 1901, p. 217 (La Tigra, Las Cruces, Western Cordillera).

P. berlepschi K. & Cl. Hartert, Nov. Zool. i. p. 56 (1894.—Rio

Pescado, W. Ecuador).

No. 3782. @imm. La Selva, Rio Jamaraya, 4600 ft., 2.x.09.—

Wing 62; tail 75; bill 41 mm.

“Tris black, feet brown, maxilla black, mandible scarlet with

black tip.”

This bird, as well as Salmon’s skins from Medellin and Santa

Elena in the British Museum, agrees with examples from Western

Ecuador upon which Hartert’s P. berlepschi was based. I regret to

say, however, that this name is a synonym of P. s. syrmatophorus

while the eastern form has to bear the subspecific term columbianus

Boucard *, as already pointed out by Salvadori & Festa rf. Gould

established the species upon specimens obtained by Prof. Jameson.

There are three skins from this source in the Gould Collection

(now in the British Museum): one, marked “ Napo—Jameson,”

* Phaéthornis columbianus Boucard, ‘The Humming Bird, i. p. 17 (1891.—

Bogota; type in Paris Museum examined).

+ Phaéthornis syrmatophorus Salvadori & Festa, Boll. Mus. Zool. Toro, xv.

1180 MR. C. E. HELLMAYR ON THE

belongs to the eastern race, P. s. columbianus, having the malar

streak and a broad stripe down the middle of the lower surface,

from the chin to the anal region, clear white, and the rump as

well as the upper tail-coverts uniform ochraceous buff; the two

others, labelled ‘“‘ Ecuador—Jameson,” agree with the so-called

“< berlepschi” from W. Kcuador, having the malar streak bright

buff, the rump feathers green with buff edges, and the whole of

the lower parts, with exception of a white stripe along the middle

of the throat, ochraceous buff. As will be easily seen from Gould’s

account, his description is a mixtum composituwm, the characters

of the lower parts (“‘ chest, abdomen, and under tail-coverts rich

buff”) being taken from the western form (berlepschi), while the

words “rump and upper tail-coverts rich buff” apply only to

the eastern colwmbianus. Since no marked type of P. syrmato-

phorus exists, M. Boucard was perfectly justified—according to

the rule of the first reviser—in restricting the name to one of

the components, and his name columbianus has to stand for the

eastern race. Thus we have :—

(a) P. s. syrmatophorus (Gould) (syn. berlepschi Hart.).

Western Heuador (Gualea, Rio Pescado, etc.) and Western

Cordillera of Colombia (La Selva, La Tigra, Las Cruces,

Medellin, Santa Elena, etc.).

(b) P. s. columbianus Boucard. Eastern Ecuador (Rio Pastaza

etc.), and Eastern Cordillera of Colombia (Bogota coll.).

139. EUTOXERES AQUILA SALVINI Gould.

[Trochilus Aquila Bourcier, P. Z. 8. 1847, p. 42 (1847—

“Jes environs de Bogota”). |

Hutoweres salvint Gould, Ann. Mag. N. H. (4) i. p. 456

(1868.—Veragua).

E. aquila (nec Bourcier) Simon & Dalmas, Ornis, xi. p. 218

(Plano de los Monos, Western Cordillera).

No. 2445. gad. Noanama, 100 ft., 5.1.08.—Wing 76; tail

54; bill 303 mm.

“Tris black, feet black, plantar surface yellow, maxilla black,

mandible yellow.”

This bird belongs to ZL. a. heterwra, as defined by Hartert &

Hartert*, having much less white in the tail than Bogota-skins

(=true LH. a. aquila). Messrs Salvadori & Festa ft cast some

doubts on the distinctness of H. baroni Hart. & Hart. t, and

take this “ species ” for the young of the ordinary heterura (sensu

Hartert). JI have neither material nor time to enter into that

question now: but whatever the specimens with an olive-grey tail

and minute, if any, white spots to the rectrices may be, there

can be no doubt that it isto this form that Gould’s term JL.

heterura has to be restricted. Although Gould, in the introductory

* Nov. Zool. i. 1894, pp, 53-54.

+ Boll. Mus. Zool. Torino, xv. no. 268, 1900, p. 2.

t Nov. Zool, i. p. 54 (1894.—Rio Pescado, Naranjal, Ecuador).

BIRDS OF WESTERN COLOMBIA, 1181

remarks to his paper, alludes to the variability of the tail-markings,

yet the formal description (J. ¢. p. 456) of H. heterura clearly

refers to the birds afterwards separated as H. baroni, cfr.

especially the words: “ tail olive-grey, 7m some instances tipped

with sullied white.”

The next available name for ZH. heterwra Hart. (nec Gould)

seems to be ZL. salvini Gould, based upon specimens from Veragua

which I cannot satisfactorily distinguish from those taken in

W. Colombia and W. Ecuador.

140. FLORISUGA MELLIVORA MELLIVORA Linn.

Trochilus mellivorus Linneus, Syst. Nat. x. p. 121 (1758—ex

Edwards: Surinam).

Florisuga mellivora Simon & Dalmas, Ornis, xi. p. 218 (Las

Cruces, West. Cordillera) ; Sclater & Salvin, P. Z. 5. 1879, p. 529

(Remedios).

Nos. 2055, 2238. Gad, gimm. Noanama: 3.ix.; Cajon,

R. Cajén : 3.x1.08.— Wing 66, 65; tail 38, 35; bill 18, 19mm.

‘Tris, feet, and bill black.”

We have also a number of skins obtained by the late J. H.

Batty on the Rio Zapota, in August and September 1898.

I’, mellivora is a wide-spread species.

14]. PoLYERATA ROSENBERGI Bouc.

Polyerata roserbergi Boucard, Genera of Humming Birds, p. 399

(end of 1895.—Rio Dagua, W. Colombia): idem, ‘ The Humming

Bird,’ v. Dec. 1895, p. 6.

Nos. 2247, 2351. ¢ Sg ad. Novita: 7, 28.xi. 08.—Wing 55,

563; tail 30, 34; bill 22, 203. mm.

No. 2551. dg ad. Juntas (Rio Tamana) : 20.11.09.— Wing 57 ;

tail 32; bill 20 mm.

No. 2570. dad. Rio Condoto: 26.111.09.—Wing 55; tail

314; bill 21 mm.

No. 2373. ¢ imm. Novita: 5.xii.08.—Wing 55; tail 32;

bill 20 mm.

No. 2255. @Q ad. Novita: 9.x1.08.— Wing 503; tail 29; bill

21 mm.

“ Tris, bill, and feet black.”

The original examples, obtained by Mr. Rosenberg in the same

district in 1894, were all more or less immature. ‘The fine series

now forwarded by Mr. Palmer enables me to state that none of

the characters which served to distinguish the Hcuadorian race

P.r. reint* holds good, and I am afraid the latter name will have

to be placed as a synonym of P. rosenbergi. Thanks to the kind-

ness of Dr. von Lorenz, of the Vienna Museum, I have been able

to examine a series of seven specimens of the so-called reinz:

266 ad,2 6 5 imm., 3 2 9, all from N.W. Ecuadort. The

* Polyerata reina Berlepsch, Orn. Monatsber. v. p. 58 (1897.—Western Ecuador).

+ They are from the following localities : Carondelet (60 ft.), Bult (60 ft.), San

Javier (60 ft.), Pambilar (60 ft.), Charco Redondo (160 ft.).

1182 MR. C, E. HELLMAYR ON THE

adult males from W. Colombia have the throat and foreneck of

exactly the some golden green hue as the Ecuadorian ones, while

the violet-blue pectoral area, in the latter, is by no means less

extended. There is no difference in the development of the

bronzy base to the outer rectrices between the two series, and the

dimensions are practically the same. The female from Novita

differs from the three “‘ reint” (ew Ecuador) by having the throat:

less spotted with shining green, though this is very insignificant.

Measurements :—

P. rosenbergi.

Wing. Tail. Bill.

mm. min. mm.

Four adult males from W. Colombia... 55-57 30-34 204-22

One immature male __,, 5 Jase DS) 32 20

One female - x as 0S 29 21

OE. ERAONOUN

Two adult males from N.W. Eeuador... 53,55 30,32 22

Two immature males __,, <s Sag 5a Oeahe Bi 21

Three adult females Be “A ... 90-52 ° 27,28 21-22

P. rosenbergi is thus found to range over the humid, tropical

coast-district of N.W. Ecuador and W. Colombia.

142. URANOMITRA FRANCLE Bourc. & Muls.

Trochilus Francie Bourcier & Mulsant, Ann. Sci. phys. et nat.,

d’ Agric. et d’Industr. Lyon, ix. p. 324 (1846.—Bogota ; descr.

¢ ad.)

Cyanomyia francie Sclater & Salvin, P. Z. 8. 1879, p. 530

(Antioquia).

Agyrtria Francie Simon & Dalmas, Ornis, xi. p. 218 (La Tigra,

Western Cordillera).

Uranomitra francie Boucard, ‘'The Humming Bird,’ vy. p. 6 (Rio

Dagua).

Nos. 2733, 2736, 3756. Gg ad. Pueblo Rico, 5200 ft., 16,

18.viii.; Siaté: 15.ix.09.—Wing 55; tail 35, 36; bill. 21-24 mm.

No. 2735. (¢)juv. Pueblo Rico: 18.viii.09.—Wing 56; tail

34 mm.

“Tris and feet black, maxilla black, mandible brown, tip black.”

Similar to Bogota-skins.

U. francie is as yet known only from the mountains of

Colombia.

143. AMAZILIA TZACATL JUCUNDA Heine.

[Zrochilus Tzacatl De la Llave, Registro Trimestre, 11. No. 5,

p. 48 (1833.—Mexico) *.]

Eranna jucunda Heine, Journ. f. Orn. xi. p. 188 (1863.— Baba-

hoyo & Esmeraldas, W. Ecuador).

* Ofr. Richmond, Auk, xvi. 1899, p. 324.

BIRDS OF WESTERN COLOMBIA. 1183

Amazilia fuscicaudata (nec Fraser) Boucard, ‘The Humming

Bird, v. p. 7 (Rio Dagua).

A. riefferi Sclater & Salvin, P. Z. S. 1879, p. 530 (Antioquia) ;

Simon & Dalmas, Ornis, xi. p. 221 (Naranjo).

Nos. 2056, 2057, 2075, 2488. ¢S gad. Noanama: 3.1x.08.;

Novita: 23.1.09; Rio Sipi: 10.ix.08— Wing 57-59; tail 35-36 ;

bill 213—22 mm.

No. 2410. @ juv. Noévita: 18.xii.08.—Wing 57; tail 34;

bill 22 mm.

“Tris and feet black, bill brown, tip black ; maxilla black in the

young bird.”

Identical in colour and dimensions with specimens from Western

Kceuador.

A. t. jucunda, which inhabits the low countries of Western

Ecuador and Western Colombia, agrees with the typical Central

American race in shape and length of the bill, but may be distin-

guished by having the upper mandible fleshy brown (instead of

blackish), and the abdomen conspicuously darker smoke-grey.

A, t. fuscicaudata, from Eastern Colombia (Bogota) and the

mountains of Merida, is much smaller, and has a considerably

shorter, stouter bill.

144, CHLOROSTILBON PUMILUS Gould.

Chlorostilbon pumilus Gould, Ann. Mag. N. H. (4) ix. p. 199

(1872.—Citado and Pallatanga, W. Ecuador); Simon & Dalmas,

Ornis, xi. p. 221 (Naranjo, La Tigra, Western Cordillera of

Colombia).

C. angustipennis (errore) Sclater & Salvin, P. Z. 8. 1879, p. 531

(Medellin).

No. 2732. oad. Pueblo Rico: 16.viii.09.—Wing and tail

in moult; bill 133 mm.

“Tris, feet, and bill black.”

This specimen is typical of C. puwmilws, which had already been

obtained in the same region by Eugéne André, as well as further

north, in Antioquia, by the late '[. K. Salmon. Topotypical birds

from Ecuador I have not seen.

(. pumilus inhabits the mountainous ranges of Ecuador and

Western Colombia.

145. THALURANIA FANNYI Del. & Boutce.

Trochilus Fannyi Delattre et Bourcier, Rev. Zool. ix. p. 310

[1846—“les bords de la rivitre de Dagua, pres Saint-Bonaven-

ture (Nouvelle-Grenade)” |.

Thalurania Fannie Simon & Dalmas, Ornis, xi. p. 221 (Buena-

ventura, El Paillon, Naranjo); Boucard, ‘The Humming Bird,’ v.

p- 6 (Rio Dagua).

No. 2814. oad. Pueblo Rico, 5200 ft., 26.x.09. Wing 55;

tail 39; bill 20 mm.

1184 MR. C. E. HELLMAYR ON THE

Nos. 2443, 3773. ¢ ¢ imm. Noanama, 100 ft., 4.1.5 Siaté,

5200 ft., 24.ix.09.—Wing 54; tail 39, 36; bill 20, 18 mm.

Nos. 2051, 2447. @ 9 ad. Noanama: 2.ix. 08, 5.1.09.—Wing

50, 51; tail 31, 322; bill 20 mm.

“ Tris and bill black; feet in males black, in females grey.”

The differences between 7’. fannyi and 7’. verticeps Gould*,

from Western Ecuador, have been well pointed out by Simon &

Dalmas. The adult male shows all the characters assigned to

Jannyi, viz , the very dark, almost black colour of the occiput and

mantle, the nearly complete, violet-blue interscapular band, the

deeply forked tail, etc. The blue margin bordering posteriorly the

glittering green crown? is not a constant feature, it being well

developed in No. 3773, scarcely indicated in No. 2443, and com-

pletely absent in No. 2814(¢ ad.). Both of the females have the

shining bluish shoulder-patch as described by Simon & Dalmas.

T. fannyi is restricted to Western Colombia, ranging from

sea-level up to about 5000 feet in the western slope of the Coast

Cordillera ¢. In Western Ecuador it is replaced by 7’. verticeps

Gould, which may be only subspecifically separable.

146. CHALYBURA UROCHRYSA Gould.

Hypuroptila urochrysa Gould, P. Z. 8. 1861, p. 198 (1861.—

«« Panama ”’).

Chalybura buffoni (errore) Sclater & Salvin, P. Z. 8. 1879,

p. 529 (Remedios, Santa Elena).

H. urochrysea Salvin, Cat. B. Brit. Mus. xvi. p. 89 (Medellin,

Remedios).

Chalybura urochrysea Hartert, Nov. Zool. v. 1898, p. 494

(Cachabi, N.W. Ecuador).

Nos. 2083, 2329, 2581. ¢ gd ad. Sipi: 12.ix.08; Ndévita:

23.x1.08 ; Condoto: 1.iv.09.—Wing 67-704; tail 42-46; bill

233-24 mm.

No. 2218. dimm. Rio Cajon: 28.x.08.— Wing 684 ; tail 42 ;

bill 24 mm.

Noss eZ 091 20905) 2b 22.2). OVO wads Ye SipitlSre 2 pipe.

6.x.08; Rio Cajon : 29.x.08.— Wing 62-66 ; tail 38-42; bill 214-

24 mm.

Tris black, feet pink, maxilla black, mandible pink (@ ), light

brown (¢).”

This rare species is easily recognizable by its bronze-green tail,

combined with the pale mandible. The females have the six

outer tail-feathers distinctly tipped with greyish white, and

the under tail-coverts pale mouse-grey (brownish grey). The

specimens obtained by Salmon in Antioquia I have examined

* In Jardine’s Contrib. to Ornith. 1851, pt. ii. p. 79, pl. xxi. (Apr. 1851.—Quito,

Ecuador),

+ Cfr. Hartert, Tierreich, livr. 9, 1900, p. 85.

i The reported occurrence at Popayan and Pasto (!) does not rest upon reliable

authority.

BIRDS OF WESTERN COLOMBIA. 1185

in the British Museum, and found them to belong to the present

species. Strangely enough, they had been determined by Salvin as

C. buffoni.

C. urochrysa is, according to our actual knowledge, confined to

the dense forests along the Pacific coast of N.W. Ecuador and

W. Colombia *. It is particularly common in the lowlands,

though Salmon also procured examples near Remedios, at 2300

feet elevation.

At higher altitudes, however, in the Western Cordillera another

species of Chalybura occurs, specimens of which were obtained

at Rio Dagua t by Rosenberg, and at Naranjét by E. André.

It agrees with C. buffoni from Bogota, in the coloration of the

tail (rectrices bluish black, more or less edged with bronze green)

and lower parts, but has the lower mandible mostly pale

brownish, like C’. wrochrysa. With only a single example before

me I refrain, however, from naming this apparently undescribed

form.

147. BoISSONNEAUA FLAVESCENS FLAVESCENS Lodd.

Trochilus flavescens Loddiges, P. Z.8. 1832, p. 7 (1832.—

Popayan, Colombia).

Panoplites flavescens Sclater & Salvin, P. Z. S. 1879, p. 529

(Medellin).

No. 2793. gad. Tatamd Mountain, 8000 ft., 12.x.08.—Wing 80;

tail 52; bill 174 mm.

“ Tris, feet, and bill black.”

This bird fully agrees with Bogota skins except that the green

tip to the inner web of the outermost rectrix is a trifle longer.

Keuadorian specimens have much broader green tips to the lateral

rectrices, and the buff portions of the tail as well as the under

tail-coverts are deeper in tint. They have been separated by Mr.

Oberholser as L. f. tinochlora §.

148. HRIocNEMIS AURELIZ Bourc. & Muls.

Trochilus Aurelie Bourcier & Mulsant, Ann. Sci. phys. et

nat., d’Agric. et d’Industr. Lyon, ix. p. 315 (1846.—Bogoti).

Eriocnemis aurelie Sclater & Salvin, P. Z. S. 1879, p. 530

(Santa Elena, Medellin).

No. 2737. 3 juv. Pueblo Rico, 5200 ft.: 6.ix.08.—Wing 63 ;

tail 40; bill 19 mm.

“Tris black, feet grey, bill black.”

Notwithstanding its rather long bill, this specimen appears

* The original locality “ Panama” is open to doubt. Warscewicz was very

careless about labelling, and, moreover, not one of the many later travellers met

with the species on the Isthmus.

+ Hypuroptila buffoni Boucard, ‘The Humming Bird,’ v. 1895, p. 8 (specimen

examined by me).

t Chalybura buffoni Simon & Dalmas, Ornis, xi. p. 222 (Naranjé, 1900 ft.).

§ Proc. U.S. Mus. xxiv. p. 329 (1902.—West side of Corazdn, Ecuador).

1186 MR. C. E. HELLMAYR ON ‘THE

to belong to the present species, for in coloration it closely

resembles several Bogota skins in corresponding stage.

E. aurelic is well-known as an inhabitant of the mountains of

Colombia.

149. UrostrictE BENJAMINI Boure.

Trochilus Benjamini Bourcier, Compt. Rend. Ac. Sci. Paris,

xxxi. no. 6, Febr. p. 187 (1851—“les régions chauds des

environs de Gualea,” W. Ecuador).

No. —(¢) ad. la Selva, Rio Jamaraya, 4600 ft., Oct. 09.—

Wing 52; tail 35; bill 23 mm.

Compared with specimens from Western Ecuador this bird has

a decidedly longer bill and more white along the middle of the

belly. From a single skin it is difficult to say whether these

divergencies are of any importance.

Like ordinary benjamini the specimen from La Selva has a

large lilac prepectoral spot.

So far as I am aware this is the first Colombian record of

U. benjamini, although it is fairly common in Ecuador on both

sides of the Andes.

150. HELIANGELUS ExoRTIS Fras.

Trochilus exortis Fraser, P. Z. S. 1840, p. 14 (1840.—Guaduas,

Colombia).

Heliotrypha parzudakii Sclater & Salvin, P. Z. S. 1879, p. 529

(Santa Elena). aed

No. 2792. g ad. Tatama Mountain, 8000 ft., 12.x.09.—

Wing 69; tail 49; bill 17 mm.

“ Tris, feet, and bill black.”

A common bird in the mountains of Colombia and northern

Ecuador. Palmer’s specimen agrees well with Bogoti skins.

151. Heuiorurix BArroti Boure. & Muls.

Trochilus Barroti Bourcier et Mulsant, Ann. Sci. phys. et nat.,

d’Agric, etc. Lyon, vi. p. 48 (1843.—* Carthagéne,” Colombia).

Heliothri« barroti Sclater & Salvin, P. Z. 8. 1879, p. 529

(Remedios).

No. 2198. g ad. Noanama; 17.x.08.—Wing 66; tail 42;

bill 16 mm. —

No, 2329. g¢ juv. Névita; 23.xi.08—Wing 65; tail 45 <

bill 154 mm.

“Tris, feet, and bill black.”

Not apprecially different from Costa Rican skins. Z. barroti,

which is very likely only a geographical race of H. auritus,

ranges southwards as far as Western Ecuador,

BIRDS OF WESTERN COLOMBIA. 1187

152. CYANOLESBIA KINGII, subsp.

[Ornismya king Lesson, Hist. Nat. Trochil. p. 107, pl. 38

(1832. —“* Jamaique,” errore — description and plate evidently

apply to the common blue-tailed Bogota form, with a large violet-

blue gular patch)].

No. 3784. gad. Tatama Mountain, 4600 ft. 7.x.09.— Wing 74;

tail (slightly moulting) 93; bill 14 mm.

“Tris, feet, and bill black.”

This bird cannot be identified with any of the known forms.

It is somewhat intermediate between OC. k. kingii from Bogota,

and C. k. cwlestis * from Ecuador, though apparently different

from either. With the latter, it shares the long, robust beak, the

broad, rufous-buff edges to the under tail-coverts, and the bronzy-

green general tone of the lower parts; but the rufescent margins

to the feathers of the breast and abdomen are completely absent,

and the colour of the tail is more like Bogoté specimens. It

should be mentioned, however, that the under surface of the

rectrices is much duller, more blackish than in any other example

examined by me. More material will probably show the West

Colombian birds to constitute a distinct race. Curiously enough,

both Hartert 7, and Simon & Dalmas{ record the green-tailed

C. k.emme Berl., without gular spot, from the Western Cordillera,

while the Tatama bird has the throat extensively violet-blue.

153. CHLORONERPES RUBIGINOSUS GULARIS Harg.

[Picus rubiginosus Swainson, Zool. Ilustr. i. pl. xiv. (1820-1.—

“Spanish Main,” 7. e. Venezuela, environs of Cumans) |].

Chloronerpes gularis Hargitt, Ibis, (6) i. p. 230 (1889.—Santa

EKlena, Antioquia).

C. rubiginosus (errore) Sclater & Salvin, P. Z. S. 1879, p. 533

(Retiro, Concordia, Santa Elena).

_ No. 2798. g ad. Loma Hermosa, Rio Jamaraya, 4150 ft.,

18.x.09.— Wing 122; tail 84; bili 263 mm.

“ Tris dark brown, feet plumbeous, bill black.”

This bird agrees perfectly with Hargitt’s description. It differs

from C. 7. rubiginosus, from N.E. Venezuela (Cumana) and

Trinidad, in having the rump and tail-coverts pale yellow in

decided contrast to the deep tawny olive of the upper back, and

the under parts more richly yellow with the blackish bars

conspicuously narrower and confined to the foreneck and breast:

The throat is much less variegated with whitish, and the feathers

of the pileum, from the forehead to the nape, are tipped with

crimson.

C. r. gularis is limited to Western Colombia, inhabiting both

the Coast and the Central Cordillera.

* Cynanthus coelestis Gould, Introd. Humming-Birds p, 102 (1861.—Ecuador).

+ Tierreich, Lief. 9, 1900, p. 176.

{ Ornis, xi. 1901, p. 223 (Las Cruces, W. Cordillera),

1188 MR. C. E. HELLMAYR ON THE

154. MELANERPES PUCHERANI PUCHERANI Malh.

Zebrapicus pucherani Malherbe, Rev. Mag. Zool. (2) i. p. 542

(1849.—“ Tabago,” errore.*).

Nos. 2258, 2279. ¢g dad. Névita: 10, 13.xi.08.— Wing 115,

113; tail 67, 65; bill 26, 25 mm.

Nos. 1970, 2259. 9 2 Guineo: 5.vui.; Névita: 10.xi.08.—

Wing 108; tail 60, 62; bill 21 mm.

‘“‘ Tris brown, feet grey or greyish green, bill black.”

The series agrees with specimens from Western Ecuador.

M. p. pucherani ranges from Nicaragua southwards through

Western Colombia to Guayaquil, S.W. Ecuador. Birds from

Central America are perhaps slightly different, but I have not

sufficient material to decide the question. The form occurring

from Honduras north to 8. Mexico has lately been separated as

M. p. perileucus Todd 7.

155. VENILIORNIS KIRKII CECILIIT Malh.

[Picus (Chloropicus) Kirkii Malherbe, Rev. Zool. viii. p. 400

(1845.—Tobago)].

Mesopicos Cecilii Malherbe, Rev. Mag. Zool. (2) i. p. 538

(1849.—Colombia).

Chloronerpes cecilie Sclater & Salvin, P. Z. 8. 1879, p. 533

(Antioquia, Remedios, Neche).

No. 1986.( 3.) ad. Guineo, Rio Calima : 10.viii.08.— Wing 84 ;

tail 55; bill 201 mm.

ub Th brown, feet pinkish grey, maxilla dark grey, mandible

paler.”

This bird, in abraded, bleached summer plumage, is not

appreciably different from Bogotaskins. Specimens from Western

Ecuador also agree well with the latter.

V. k. cecilii is peculiar to Colombia (Coast, Central and Kastern

Cordillera £) and Western Ecuador. In Panama it is replaced

by the doubtfully separable V. k. darienensis Ridgw. §

156. CrLeus LoricAtus Reichb.

Meiglyptes loricatus Reichenbach, Scansorie, p. 405, pl. delxxxi.

figs. 4495-96 (1854.—“ Peru”; descr. 9 ).

Celeus loricatus Sclater & Salvin, P. Z. S. 1879, p. 533

(Remedios & Neche).

* The species does not occur on “ Tabago”’ (sc. Tobago), the original locality being

no doubt erroneous. Later, in the Monogr. des Picid. ii. p. 227, Malherbesays that

the specimens he saw in the British Museum came from “ Tabago ” while those

existing in his own collection were from “ Nouvelle-Grenade.” We may therefore

accept Colombia as the type locality since the birds from this country agree very

well with Malherbe’s description and figure (pl. ciii.).

+ Proc. Biol. Soc. Wash. xxiii. p. 154 (1910.—Manatee, British Honduras).

~ Count Berlepsch (J. f. Orn. 1884, p. 314: Chloronerpes cecilie) notices that

examples from Bucaramanga are considerably larger than those from Bogota.

§ Proc. Biol. Soc. Wash. xxiv. p. 33 (1911.—E] Réal, Darien).

BIRDS OF WESLERN COLOMBIA. 1189

C’. mentalis Cassin, Proc. Acad. N. Sci. Philad. 1860 p. 137

(1860.—Turbo & R. Atrato; deser. 3).

No. 2254. (¢) ad. Noévita: 9.x1.08.—Wing 117; tail 70;

bill 21 mm.

‘Tris crimson, feet grey, maxilla black, mandible yellow.”

This specimen, a fully adult male with cheeks, malar region

and upper throat crimson, corresponds exactly with Cassin’s

description of C. mentalis. ‘The types of the latter were obtained

in the same general region, but farther to the north, on the Rio

Atrato.

It appears, however, that J/. loricatus was based upon a female

of the same species,

C. loricatus ranges from Costa Rica to Western Ecuador.

I have not seen specimens from Costa Rica or Panama. If

separable, they will have to bear the name squamatus Lawr. *

157. CEOPHLG@US LINEATUS LINEATUS Linn.

Picus lineatus Linneus, Syst. Nat. 12, 1. p. 174 (1766—ex

Brisson : Cayana).

Dryocopus lineatus Sclater & Salvin, P. Z. 8. 1879, p. 532

(Santa Elena).

No. 2014. 9 ad. Noanama: 25.viu1.08.— Wing 180; tail‘115;

bill 36 mm.

“‘ Tris white, feet blue-grey, bill black.”

[158. PicUMNUS OLIVACEUS GRANADENSIS Lafr.

[Picumnus olivaceus Lafresnaye, Rey. Zool. vill. p. 7 (1845—

“ad. Bogotam, in Colombia ”). |

PR: granadensis Lafresnay Cli nice x. ype 1S. iC S4i ad:

Caly, in Nova Granada ”—coll. Delattre: @ ad., type in Mus.

Acad. Philad., cfr. Stone, Proc. Acad. N. Sci. Philad. li. 1899,

p- 52) ; Hargitt, Cat. B. Brit. Mus. xviii. p. 549 (part.: fA g.

¢ 2 Medellin, Antioquia, examined by me).

P. olivaceus (nec Lafr.) Sclater & Salvin, P. Z. 8. 1879, p. 532

(Medellin, Colombia).

P. canus Bangs, Proc. Biol. Soc. Wash. xxi. p. 72 (1910.—

Naranjito, R. Dagua; one @ ad.).

No. 387. g ad. Primavera, W. Colombia, 5200 ft. Raap coll.—

Wing 60; tail 30; bill 133 mm.

No, 246. ¢ ad. San Isidro, 2700 ft. Raap coll.—Wing 59;

tail 29% bill 12) mm.

Nos. 257, 271.

Raap coll. Wing 58, 59; tail 29, 30; bill 12 mm.

One 2 ad. San Antonio, Western Cordillera, 5400 ft. 13.vi.09.

Fassl coll.— Wing 57; tail 30; bill 12 mm.

Two 6 g ad. Rio Dagua: 12, 16.vi.95. Rosenberg coll.—

Wing 58; tail 292, 30; bill 12-13 mm.

2700 ft.

* Celeus squamatus Lawrence, Ibis, v. p. 184 (1863.—Panama Railroad).

Proc. Zoou. Soc.— 1911, No. LX XX. 80

1190 MR. C. E. HELLMAYR ON THE

This series, which belongs to the Tring Museum, enables us

at last to make out the status of P. granadensis, originally based

upon an adult female from the same district (Cali). I am sorry

to say that Mr. Bangs under P. canus simply redescribed the

true granadensis. This is quite evident on comparing the two

descriptions, which agree almost word for word. In a note

appended to the diagnosis of P. o. harterti *, I have already

alluded to the distinguishing characters of P. 0. granadensis, but

a few additional remarks may not be out of place here. The

geographical variation of this group was quite unsatisfactorily

understood until Mr. Hartert ~ pointed out the differences

existing between the three races then known to him. MHartert

was, however, not acquainted with granadensis Lafr., for the

birds from Western Ecuador to which he applied that name proved

to be distinct and were subsequently separated by me as P. o.

hartertt.

According to our present knowledge, the following subspecies

of P. olivaceus ave easily recognizable :—

(a) P. otrvacnus oxivAceus Lafr.

Type from Bogota.

Adult. Back warm greenish brown or olivaceus brown; chest

strongly tinged with deep olivaceous; breast and abdomen

yellowish with well-defined, though narrow, dusky stripes,

particularly on the lower breast and flanks; quills edged with

olivaceous. Adult males with tips to feathers of forehead

“flame scarlet” (vather duller than pl. vu. fig. 14 of Ridgway’s

Nomenclature).

Hab. Appavently the Eastern Cordillera of Colombia. All the

specimens I have seen, eleven at Tring, and nine in the Munich

Museum, are of the well-known Bogota form.

(6) P. oLIvaceus FLAVorINcTUS Ridgw.t

Type from Pozo Azul, Western Costa Rica.

Adwt. Back not unlike P. 0. olivaceus, but mostly somewhat

darker ; edges of remiges brighter and deeper olivaceous yellow ;

chest decidedly darker, more brownish, less greenish; dusky

stripes of breast and abdomen wider and less sharply defined.

Adult males with tips to feathers of forehead dull “cadmium

orange” (Ridgw. Nomencel. pl. vi. fig. 2.)

iah South-western Costa Rica (south of Pozo Azul de Pirris)

and Chiriqui §.

* Bull. B. O. C. xxiii. March 1909, p. 67.

+ Novit. Zool. ix. 1902, pp. 606-7.

t P. flavotinctus Ridgway, Proc. U.S. Mus. xi. 1888, p. 543 (1889.—Pozo Azul,

vw pe Rica).

§ An additional. subspecies, P. clivaceus panamensis Ridgw. (Proc. Biol. Soc.

W Hae xxiv. 1911, p. 44: Lion Hill Station, Panama) inhabits Panama (Lion Hill,

Tocoumé). There is a single adult male trem the latter locality, obtained

by E. André, in the Tring Museum. This form is nearest to P. o. flavotinctus, but

has the chest buffy yellow (instead of brownish) and the back much paler, light

earthy-brown.

BIRDS OF WESTERN COLOMBIA. 1191

Examined: Two males and two females, Costa Rica (Tring

Museum), four males and one female, Chiriqui (Tring and

Munich Museums).

N.B.—An adult male in bad condition, said to be from San

Esteban, Venezuela, in the British Museum differs merely in

having the breast more strongly tinged with brown. The locality

is most probably erroneous.

P. o. harterti Hellmayr, Bull. B.O.C. xxiii, p. 67 (March

1909.—Paramba, N.W. Ecuador).

P. granadensis (nec Lafr.) Sclater, P. ZS. 1860, p. 95

(Nanegal) ; idem, l.c. p. 287 (Babahoyo); Berlepsch & Taczan-

owski, 1. c. 1883, p. 570 (Chimbo, Yaguachi); iidem, |. c. 1885,

p. 106 (Yaguachi) ; Hartert, Nov. Zool. v. 1898, p. 497 (Chimbo) ;

Salvadori & Festa, Boll. Mus. Zool. Torino, xv. no. 368, 1900,

p. 16 (Vinces) ; Goodfellow, Ibis, 1902, p. 209 (Santo Domingo) ;

Hargitt, Cat. B. Brit. Mus. xviii. p. 549 (part.: 7, k, Babahoyo ;

l, Monj1).

P. olivaceus granadensis Hartert, Nov. Zool. ix. 1902, p. 606

(Chimbo, Paramba, 8. Domingo).

Type from Paramba, N.W. Kcuador, 3500 ft. alt.

Adult. Much like P. o. flavotincéus, but chest olivaceous as in

P. o. olivaceus, and adult males with tips of feathers on forehead

much paler, cadmium yellow (Ridgw, pl. vi. fig. 6).

Hab. Western Ecuador, from sea-level up to 3500 feet.

Exanuned: Kleven males and four females in British, Tring,

and Munich Museums.

(d) P. oLIVACEUS GRANADENSIS Lafr.

Synonymy, see above.

Type from Cali, W. Colembia.

Adult. Differs at once from the three preceding races in having

the back light olive-grey or pale smoky-grey (without greenish

or brownish suffusion) and the under parts creamy-white, nearly

uniform, with but a few obsolete dusky streaks on the flanks, and

the foreneck very slightly tinged with dull greyish. Edges of the

quills creamy white, instead of olivaceous or olive-yellow. Tips

of feathers on forehead in adult males even paler than in

P. 0. harterti, “ chrome-yellow ” (Ridgw. pl. vi. fig. 8.

Hab. Western Cordillera of Colombia: Cali (Delatire), Rio

Dagua (Rosenberg), Naranjito (Palmer); Primavera, San Isidro,

Media Luna (aap); San Antonio (fassl); Medellin, Antioquia

(Salmon).

Obs. In addition to the specimens enumerated above I

examined a couple from Medellin (Salmon) in the British Museum

and found them toagree. One of the Dagua examples (Rosenberg)

is rather more brownish grey above than the others, thereby

forming the passage to P. 0. harterti.

80*

1192 MR. C. E. HELLMAYR O THE

(e) P. oLtvAcEUS DIMotUS Bangs.

Picumnus dimotus Bangs, Bull. Mus. Harvard Coll. xxxix.

p. 146 (1903.—Ceiba, Honduras).

P. olivaceus (nec Lafr.) Hargitt, Cat. B. Brit. Mus. xviii. p. 548

(part.: A, 7, Julian, S. Pedro: Honduras).

Adult. Much like P. o. olivaceus, and agreeing in the adult males

in having the tips of the feathers on the forehead scarlet, but with

back and chest much more greenish, and dusky streaks on belly

less distinct.

Hab. Honduras: Ceiba, Julian, San Pedro; Nicaragua.

Kxamined : Two adult males from Honduras (Whitely) in the

British Museum. |

159. CERYLE TORQUATA TORQUATA Linn.

Alcedo torquata Linneeus, Syst. Nat. 12, i. p. 180 (1766—ex

Brisson: Mexico & Martinique).

Ceryle torquata Cassin, Proc. Acad. N. Sci. Philad. 1860, p. 133

(R. Atrato & Truando); Sclater & Salvin, P.Z.S. 1879, p. 534

(Neche). !

No. 3759. Q ad. Siaté, R. Siaté, 5200 ft., 17.1x.09.—Wing

195; tail 122; bill 65 mm.

“Tris brown, feet greyish green, bill black, base yellow.”

160. CERYLE AMAZONA Lath.

Alcedo amazona Latham, Ind. Ornith. i. p. 257 (1790.—

“ Cayana ”).

Ceryle amazona Cassin, Proc. Acad. N. Sci. Philad. 1860, p. 133

(R. Nercua); Sclater & Salvin, P. Z.8. 1879, p. 534 (Neche).

No. 2397. gad. Ndéovita: 14.x11.08.—Wing 131; tail 85;

bill 66 mm.

‘“‘ Tris dark brown, feet and bill black.”

161. CeryLe inpA Linn.

Alcedo inda Linneus, Syst. Nat. 12, 1 p. 179 (1766—

ex Edwards: ‘“ India occid.”, errore; we substitute Surinam

as type locality).

Ceryle inda Cassin, |. ce. p. 133 (Turbo).

No. 2723. Q ad. ‘Tadd, R. San Juan: 28.11.09.—Wing 97;

tail 62; bill 43 mm.

“Tris dark brown, feet black, maxilla black mandible dark

brown.”

162. CERYLE AMERICANA AMERICANA Gm. (?)

Alcedo americana Gmelin, Syst. Nat. 1, 1. p. 451 (1788—

ex Daubenton, Pl. Knl. 591. figs. 1, 2: Cayenne).

Ceryle cabanisi (nec Tschudi) Sclater & Salvin, P. Z.S. 1879,

p. 534 (Retiro, Concordia, Medellin).

Nos. 1990, 2077, 2209. g g ad. Mouth of Calima: 13.viii. ;

Sipi: 1l.ix.08; Noanama: 21.x.08.—Wing 75-77; tail 54-55 ;

bill 36-39 mim.

BIRDS OF WESTERN COLOMBIA. 1193

“Tris dark brown, feet and bill black.”

These three as well as another adult male from the Cauca

River (Batty coll.) in the Munich Museum differ from a very

large series of skins obtained in Cayenne, Venezuela (Caura),

Brazil, etc., in having the under tail- coverts whoily unspotted

white or very faintly spotted with bronze-green. It should be

mentioned, however, that this character is likewise found in two

specimens from the island of Toba go.

In the absence of typical examples from Western Peru, 1

cannot say whether the birds from Western Colombia should not

be referred to C. a. cabanisii Tsch.* I notice, however, that

the white bars on the quills are by no means broader than in

typical C. a. americana, which is said to be the case in the

Peruvian form.

163. TROGON MASSENA Gould.

Trogon massena Gould, Monogr. Trogon. Ist ed. pl. 16 (1838.—

Guatemala); Cassin, Proc. Acad. N. Sci. Philad. 1860, p. 135

(Truando ; delta of the Atrato).

No. 2008. g ad. Noanama, 100 ft., 26.viii.08.—Wing 170;

tail 168; bill 24 mm.

“Tris dark brown, feet brownish yellow, bill orange-yellow.”

Compared with others from Central America, this bird is

smaller, and has the middle pair of rectrices washed with dull

bluish instead of clear bronze-green. With a single specimen at

hand it is, of course, impossible to judge the value of these

variations. In other respects the male from Noanama is a

typical massena, there being not the least trace of a white

breast-band, ete.

Noanama is the most southerly locality yet known for 7’.

massena. Curiously enough, the late T. K. Salmon obtained

the nearly allied 7. melanurus macrowrus Gould in the State

of Antioquia T, provided that his specimens have been correctly

determined.

164. UrosPaArHA MARTI SEMIRUFA Ncl.

[Prionites martw Spix, Av. Bras. i. p. 64, pl. Ix. (1824—“ in

sylvis Parae”).|

Momotus semirufa Sclater, Rev. Mag. Zool. (2) v. p. 489

(1853—part.: Santa Marta, New Grenada {).

Momotus Martii (nec Spix) Cassin, Proc. Acad. N. Sci. Philad.

1860, p. 1386 (River Nercua).

Urospatha mariwu Sclater & Salvin, P. ZS. 1879, p. 534

(Remedios, Neche).

* Alcedo Cabanisii Tschudi, Fauna Peru., Aves, p. 253 (1844-45.—“ environs of

Lima’”’).

+ T. macrurus Sclater & Salvin, P. ZS. 1879, p. 535 (Remedios, Neche).

{ Although one of the localities, Rio Javarri (W. Brazil), refers to typical U. m.

inane, Sclater’s description Is evidently taken from the Santa Marta specimen.

Cfr. the words: “ cauda spatulata.”

1194 MR. C. E. HELLMAYR ON THE

Nos. 2108, 2419. ¢ gad. Sipi: 23.ix.; Névita: 20.xii.08.—

Wing 147, 150; tail 250, 255; bill45, 47 mm.

No. 2013. 9 ad. Noanama: 25.viii.08.—Wing 148; tail 250;

bill 46 mm.

“Tris dark brown, feet and bill black.”

All three specimens have the median rectrices spatulated and

decidedly bluish. Cfr. also my remarks in Abhandl. Bayer.

Akad. Wiss., II. Cl. vol. xxii. 3, 1906, p. 611, and in Nov. Zool.

xiv. 1907, p. 403.

U. marti semirufa replaces the typical race in Western

Keuador and Colombia, ranging northwards to Costa Rica and

Nicaragua.

165. Momotus 2QUATORIALIS ZQUATORIALIS Gould.

Momotus equatorialis Gould, P. Z. 8. 1857, p. 223 (Jan. 1858.—

‘“‘ Archidona, near the Equatorial line, on a branch of the Rio

Napo”); Sclater & Salvin, P.Z.S. 1879, p. 534 (Envigado,

Retiro, Concordia, Frontino).

No. 2820. g ad. Pueblo Rico, 5200 ft., 29.x.09.—Wing 154;

tail 315; bill 422 mm.

‘* Tris vermilion, feet dark grey, bill black.”

We have also a fine pair procured by the late J. H. Batty

at Rio Lima, 4500 feet, Western Cordillera, in August 1898.

The Colombian skins agree exactly with others from Ambato,

HE. Ecuador, which can be regarded as topotypical.

AM. equatorialis is an inhabitant of the highlands of Ecuador

and Western Colombia. It may prove to be merely the southern

form of the Central American M, lessoni Less. In Peru it is

replaced by a nearly allied race, J/. wquatorialis chlorolemus

Berl. & Stolzm.*.

166. GALBULA MELANOGENTA Scl.

Galbula melanogenia Sclater, Contrib. to Ornith. for 1852, p. 61

(1852—loe. ign.).

G. ruficouda (nee Cuvier) Cassin, Proc. Acad. N. Sci. Philad.

1860, p. 134 (River Nercua).

Nos. 2219, 2513. gg ad, Rio Cajén, 125 ft., 29.x.08 ;

El Tigre, 320 ft., 5.11.09.— Wing 79, 82; tail 93, 94; bill 47 mm.

Nos. 2220, 2567. 9 9 ad. Rio Cajén: 29.x.08; Juntas,

400 ft., 11.1.09.-Wing 79, 81; tail 93; bill 44 mm.

“Tris dark brown, feet yellowish green, bill black.”

The specimens are identical with others from Chiriqui and

Western Ecuador.

167. Bucco prcrorauis Gray.

Bucco pectoralis Gray, Genera Birds, i, pl. xxvi. (Dec. 1846—

no locality given); Wyatt, Ibis, 1871, p. 374 (Magdalena Valley,

between Naranjo and the river); Sclater & Salvin, P. Z.S. 1879,

* P.Z.S. 1902, ii. p. 35 (1902.—Ocobamba near Cuzco, S.E. Peru).

BIRDS OF WESTERN COLOMBIA. 1195

p- 536 (Neche); Berlepsch, Journ. f. Ornith. 1884, p. 315

(Bucaramanga).

Nos. 2464, 2626. ¢ ad, d imm. Noévita: 4.xi1.08; Tado:

29.iv.09.—Wing 100; tail 83; bill 323, 32 mm.

Nos. 2318, 2359, 2464. 9 9 ad., 1 Q imm. Novita: 20.x1.,

1.xii.08; Noanama: 12.1.09.—Wing 95-100; tail 80-84; bill

30-33 mm.

“Tris dark red, feet grey, bill black.”

Adults have the edges of the scapulars, wing-coverts, rump, etc.,

pure white, while they are more or less tinged with buff in

immature birds; the latter also show a slight buffish wash on the

lower flanks and under tail-coverts.

Two skins from Chiriqui have the black portions of the

plumage faintly glossed with oil-green, while our Chocé series,

as also a couple from N.W. Hcuador, show a strong metallic blue

loss.

B. pectoralis ranges from Western Panama (Chiriqui) south-

wards to N.W. Ecuador (province Hsmeraldas *). It is exclusively

found in the hot, dense forests of the lowlands.

168. Bucco TEcTUS SUBTECTUS Scl.

[Bueco tectus Boddaert, Tabl. Pl. Enl. p. 43 (1783—ex

Daubenton, Pl. Enl. 688. fig. 2: Cayenne). |

Bucco subtectus Sclater, P.Z.S8. 1860, p. 296 (1860.—Esme-

raldas, N.W. Ecuador); Sclater & Salvin, P.Z.S. 1879, p. 536

(Neche).

Nos. 2634, 2703. 9 9 ad. Tadoé (230 ft.): 3.v., 12.vi.08.—

Wing 69; tail 56; bill 21, 22 mm.

‘“‘ Tris red, feet and bill black.”

This well-characterized race differs from B. ¢. tectus, of Cayenne,

Para, and Hastern Venezuela (Caura), in several important points :

The upper parts are of a much deeper, more glossy black; the

white spots on the head are confined to the forehead and

anterior crown; the white cross-bar on the inner web of the

submedian pair of rectrices is either wholly absent or but faintly

indicated; the black crescent of the foreneck is much narrower ;

the bill more slender.

Birds from Panama have even less white spotting about the

forehead than those from more southern localities, but otherwise

they do not differ.

B. t. subtectus has exactly the same range as the preceding

species, occurring from Western Panama (Chiriqui) south to

N.W. Hcuador (province Esmeraldas).

169. Bucco NoANAM# Hellm.

Bucco noaname Hellmayr, Bull. B.O.C. xxv. p. 21 (1909.—

Noanama, W. Colombia).

* There are two specimens in the Tring Museum: gad. from Carondelet (60 ft.),

Oct. 16, 1900; 9 ad. Bultin (160 ft.), Dec. 23, 1900, both secured by Mr. G.

Flemming, one of Mr, Rosenberg’s correspondents.

1196 MR. C. E. HELLMAYR ON THE

No. 2033. g ad. Noanama (100 ft.), 28.vili.08. Type of

species.— Wing 81; tail 66; bill 28 mm.

Nos, 2708, 2711. § Q ad. Tado (230 ft.), 8, 16.vi.09.— Wing

81, 80; tail 66; bill 28, 27 mm.

“Tris red or brown, feet grey, bill black.”

Adult (sexes alike). Upper part of the head and nape dark

ashy; back and upper wing-coverts dark sepia-brown, each

feather with a distinct, pale rufescent apical margin ; upper tail-

coverts with rather brighter rufescent cross-bands and edges ;

remiges and rectrices dusky, narrowly fringed with rufescent-buff

along the outer web. Narrow frontal edge and broad super-

ciliaries, reaching as far as the posterior angle of the eye, soiled

white; Jores ashy black; cheeks, ear-coverts, and sides of the

neck dark ashy, some of the feathers edged with pale greyish.

Malar region, chin, aud throat white; chest dull sooty black,

most of the feathers showing, on their concealed basal portion,

a white mesial spot or streak, only to be seen when the feathers

are raised; rest of the belly white, washed with ochreous-buff

and marked with coarse blackish spots or transverse bands; under

tail-coverts uniform buff. Axillaries and under wing-coverts deep

buff, mixed with blackish ; inner web of remiges broadly edged

with buff on its basal half.

This new Puff-bird is not very nearly related to any other

member of the genus. In proportions and style of coloration

(such as the rufescent cross-bands on the upper parts, buff

axillaries, under wing-coverts and quill-lining, ete.) it is not un-

like B. macrodactylus Spix from Amazonia; but the ashy cap,

the much broader blackish breast-band, the coarse spotting of the

belly, ete., as well.as the much larger size, serve to distinguish it

at a glance.

The three specimens sent by Mr. Palmer are perfectly alike,

the female being but very slightly smaller.

B. nodname is, as yet, known only from the valley of the San

Juan River in Western Colombia.

170. MALACOPTILA PANAMENSIS POLIOPIS Scl.

[Malacopiila Panamensis Lafresnaye, Rev. Zool. x. p. 79

(1847.— Panama). |

Malacoptila poliopis Sclater, P. Z. 8. 1862, p. 86, pl. viii.

(1862.—Esmeraldas, N.W. Ecuador, descr. 9 ).

M. panamensis (nec Lafr.) Cassin, Proc. Acad. N. Sci. Philad.

1860, p. 134 (Rio Truando).

Nos. 2085, 2437, 2520. g¢ g ad. Sipi (150 ft.), 12.ix.; Névita

(150 ft.), 26.x11.08 ; El Tigre (320 ft.), 9.11.09.— Wing’ 88-90 ;

tail 75-78; bill 29 mm.

Nos. 2084, 2593. 9 @ ad. Sipi (150 ft.), 12.ix.08 ; Condoto

(150 ft.), 12.iv.09.— Wing 86-88 ; tail 75, 79; bill 29 mm.

“Iris red, feet grey, maxilla black, mandible yellowish grey.”

The series agrees perfectly with topotypical examples from the

BIRDS OF WESTERN COLGMBIA. 1197

province of Esmeraldas. Jf. p. poliopis appears to be a well-

marked race, differing from J. p. panamensis, of Panama,

Chiriqui, and 8.W. Costa Rica, in its darker coloration. The

males are much darker rufous above and have the throat

and foreneck of a deeper cinnamon-rufous shade, whilst the

females may also be distinguished by their fuscous (less brownish)

back, more blackish cheeks, darker ochraceous foreneck, etc.

M. p. poliopis is apparently confined to the lowlands and foot-

hills of the Pacific coast district of Colombia (from the Truando

southwards) and W. Ecuador (south to Guayaquil).

171. MALAcoprinaA MysTACGALIS Lafr.

Monasa mystacalis Lafresnaye, Rev. Mag. Zool. (2) 11. p. 215

(1850.—Colom bia).

Malacoptila panamensis (errore) Sclater & Salvin, P. Z.5.

1879, p. 5386 (Remedios).

M. mystacalis Berlepsch, Journ. f. Ornith. 1884, p. 315

(Bucaramanga ; crit.).

Nos. 2803, 3764. g 2 ad. Loma Hermosa (4150 ft.), 22.x. ;

Siat6, near Pueblo Rico (5200 ft.), 21.1x.09.— Wing 98, 95; tail

96, 94; bill 30, 28 mm.

“ Tris red, feet grey, maxilla black, mandible yellow.”

These two specimens as well as several others from Bogota are

undoubtedly different from J/. p. poliopis. They are much larger,

with a stouter, stronger bill; the forehead is broadly white,

bordered posteriorly by a very distinct, black band ; the cinnamon-

rufous colour below extends down over the breast, leaving only

the middle of the abdomen white, the flanks being mixed with

dull earthy brown, while in all the races of M/. panamensis the

breast and sides are sharply striped with blackish; the upper

parts are of quite another shade of brown, ete. The sexes are

alike, whereas there is a marked sexual difference in the forms

of M. panamensis.

M. mystacalis inhabits the mountains of Colombia, being found

in the Western as well asin the Eastern Cordillera (Bucaramanga,

Bogota).

172. MoNASA PALLESCENS Cass.

Monasa pallescens Cassin, Proc. Ac. N. Sci. Philad. 1860, p. 134

(1860.—Cordilleras of the Rio Truando) ; idem, |. c. 1864, p. 287,

pl. iv.; Wyatt, Ibis, 1871, p. 374 (Paturia, Magdalena Valley) ;

Sclater & Salvin, P. Z.S. 1879, p. 5386 (Remedios, Neche).

No. 2562. Q ad. Juntas, Rio Tamana (405 ft.), 5.111.09.—

Wing 142; tail 132; bill 39 mm.

“Tris dark brown, feet black, bill scarlet.”

A fine example of this scarce species, which is at once known,

among the other white-fronted forms, by its black throat and

smaller, nearly white upper wing-coverts. It corresponds exactly

with Cassin’s description and figure.

1198 MR. C. E. HELLMAYR ON TILE

M. pallescens is peculiar to the forests of Western Colombia,

Juntas being the most southerly locality on record. Specimens

from the Central Cordillera (Remedios, Neche, Paturia) I have

not seen, but from Sclater’s remarks they would seem to be

somewhat different.

173. Capito QUINTICOLOR Elliot.

Capito quinticolor Elliot, Nouv. Arch. Mus. Paris, i. Bull.

p. 76, pl. iv. fig. 1 (1865.—“ Nouvelle Grenade,” Triana coll.) ;

Dalmas, Bull. Soc. Zool. France, xxv. 1900, p. 176 (¢ 9: El

Paillon, near Buenaventura).

No. 2689. gad. ‘Tadé, 230 ft., 2.vi.09.—Wing 82; tail 453;

bill 21 mm.

“ Trides reddish brown, feet dark grey, maxilla black, mandible

blue-grey.”

This bird agrees well with the figure of the type except

in having a distinct orange tinge along the middle of the lower

breast and abdomen. It is an adult male in perfect plumage.

C’. quinticolor is the rarest of the Barbets of South America, only

four specimens being on record. The type, in the Paris Museum,

was brought by Mons. Triana from Colombia, without any

further locality. Count Dalmas recorded a couple obtained by

Eugéne André near Buenaventura, province of Chocé, viz., in the

same general district whence our specimen also comes.

C. quinticolor is apparently confined to the humid, forest-

covered lowlands of Western Colombia.

174. Capito MACULICORONATUS Lawr.

Capito maculicoronatus Lawrence, Ann. Lye. N. H. N. Y. vu.

p. 300 (Jan. 1861.—Lion Hill, Panama); Sclater & Salvin,

P.Z.8. 1879, p. 537 (Remedios, Neche: Antioquia).

Nos. 2297, 2312. § gd ad. Noévita: 16, 19.xi.08.— Wing 88, 85 ;

tail 56, 54; bill 213 mm.

No. 2227. g§ nearlyad. Rio Cajén: 31.x.08.—Wing 85; tail

56; bill 22 mm.

Nos. 2298, 2313. 9 9 ad. Noévita: 16, 19.xi.08.—Wing 84;

tail 53, 54; bill 212, 22 mm.

No. 2028. 9 imm. Noanama: 28.vili.08.—Wing 78; tail 54;

bill 21 mm.

“Tris black (¢), dark brown (2), feet blue, bill blue, tip

black.”

Compared with two specimens from Panama in the Munich

Museum, the males have the yellow tinge across the lower

breast rather paler, but in other respects they are similar.

C. maculicoronatus ranges from Veragua and Panama south-

wards to Western Colombia, and inhabits exclusively the hot,

forest-clad lowlands and foot-hills.

BIRDS OF WESTERN COLOMBIA. 1199

In Western Ecuador, its place is taken by C. squamatus Salv.*,

which differs, in both sexes, in lacking the scarlet patch on the

flanks, in possessing a Jarge white patch on the outer web of the

tertiaries, and in having the forehead orange-red instead of

black. The female, too, has the feathers of the back and the

upper wing-coverts narrowly edged with white. The hitherto

undescribed male has—lke that of C. maculicoronatws—the

cheeks, throat and foreneck white, the chest light saffron-yellow,

and the upper parts uniform glossy black, but there are hardly

any black streaks on the flanks. J have examined a good series

from Carondelet (province HEsmeraldas) and Santo Domingo,

W. Ecuador.

175. CAPITO BOURCIERII SALVINI Shelley.

[| Micropogon Bourcierit Lafresnaye, Rev. Zool. viii. p. 179

(1845—ad Bogotam ; deser. ¢).|

Capito salvint Shelley, Cat. B. Brit. Mus. xix. p. 119 (1891.—

“Central America, from Panama to Costa Rica,” no type

specified !); Dalmas, Bull. Soc. Zool. France, xxv. 1900, p. 180,

note (Las Cruces, Western Cordillera above Buenaventura,

6000 ft.). ;

C. bourciert (nec Lafresnaye) Sclater & Salvin, P.Z.S. 1879,

p- 538 (Frontino, Antioquia).

No. 2806. g ad. Loma Hermosa, Rio Jamaraya, 4150 ft.,

23.x.09.— Wing 74; tail 52; bill 20 mm.

INoseZmoo 250 G Quads. Pueblo) Rice, 52005 fhe sda.

25.x.09.— Wing 76, 73; tail 52; bill 20, 193 mm.

Nos. 2739, —. 2 ad.,? imm. Pueblo Rico: 7.ix., Nov. 09.—

Wing 74; tail 50, 51; bill 20, 19 mm.

Nos. 2808, 3774. 9 Q ad. Loma Hermosa: 23.x.09; Siato,

5200 ft., 25.1x.09.— Wing 71; tail 49, 47; bill 19 mm.

“Tris dark red, feet greyish green (¢) or blue-grey (2), bill

greenish yellow or yellow.”

As already pointed out by Count Dalmas (J.c.) the birds from

W. Colombia are referable to C. 6. salvini. Compared with

a good number from Costa Rica and Chiriqui the specimens from

Chocé are slightly larger on the average, but in coloration they are

nearly similar. The males have the throat dark crimson which

gradually passes into the deep orange of the chest, while, in the

females, the broad black frontal band is immediately followed by

the golden olive of the crown, without the least trace of the pale

blue cross-band, so conspicuous a feature in C. 6. bowrcierit 2, of

the Hastern Cordillera (Bogoti).

Count Dalmas (/.¢. pp. 179-180) most correctly explained the

distinguishing characters of the three races of this group, and I

quite agree with him that the West Ecuadorian birds constitute

* This, 1876, p. 494, pl. xiv. (= 9; Santa Rita); Goodfellow, Ibis, 1902, p. 218

(Santo Domingo).

1200 — MR. C. E. HELLMAYR ON THE

a quite distinct form, immediately recognizable by the crimson

of the throat and foreneck being abruptly contrasted with the

clear sulphur-yellow belly without any orange tinge. His name

CU’. shelleyi* is, however, antedated by C. equatorialis Salvad. &

Festa t by several months. Of this distinct form I have

examined, besides the type actually at Tring, two fine adult

males from Quito and Gualea, in the Munich Museum, and two

males in poor condition, from Nanegal and Esmeraldas, respectively,

in the British Museum collection. The original locality “‘ Rio

Napo” is erroneous, the type being a skin of the well-known

Quito form.

The range of the three races known to me is as follows :—

(a) C. 6. bourcieriti Lafy. Eastern Cordillera of Colombia

( Bogotz coll.), and Eastern Ecuador.

(b) C. 6. salvini Shelley. Costa Rica, Chiriqui and Western

Cordillera of Colombia.

Intac, Chimbo, Pallatanga, Nanegal, Esmeraldas, ete.

[176. SEMNORNIS RAMPHASTINUS Jard.

Tetragonops ramphastinus Jardine, New Edinburgh Philos.

Journ. (n.s.) i. p. 404 (Oct. 1855.—‘ Hastern Cordillera between

Quito and the mountain Cayambe,” N. Ecuador).

No. 87. g ad. La Tigra, W. Cordillera, 5700 ft., 13.iv.,1899.

EH. André coll.— Wing 110; tail 81; bill 22 mm.

“Tris red, feet olive-yellow, bill. yellow, with a dusky spot

towards the tip.”

This bird agrees. perfectly with specimens in the Munich

Museum from the neighbourhood of Quito.

So far as I know the species has not previously been recorded

from Colombia. |

177. RamMpHastos swaAtnsonit Gould.

Rhamphastos Swainson Gould, P. Z.S. 1833, p. 69 (Sept.

1833—“ in montosis Columbie.”)

Ramphastos tocard (nec Vieillot) Sclater & Salvin, P. Z.S. 1879,

p. 537 (Concordia, Medellin, Remedios).

Rk. tocardus Cassin, Proc, Acad. N. Sci. Philad. 1860, p. 136

(River Nercua).

No. 2000. Q ad. Noanama {100 ft.), 22.viii.08.—Wing 230 ;

tail 170; bill 166 mm.

‘“‘Tris dark green; feet blue; base of maxilla below blackish,

diagonal line dark red, remainder yellow; mandible dark red,

apical third black with an oblong yellow patch near the tip.”

This bird is typical swainsonii, having the basal portion of both

mandibles (in skin) rich salmon colour, while in the allied

* Bull. Soc. Zool. France, xxv. p. 179 (Noy. 1900.—* Rio Napo ”’—errore !)

+ Boll. Mus. Zool. Torino, xy. no. 368, p. 22 (Feb. 1900.—Intac, W. Ecuador).

BIRDS OF WESTERN COLOMBIA. 1201

R. ambiguus Swains.* the same parts are dull black. C/r. Hartert’s

observations in Noy. Zool. v. 1898, p. 498.

I am at a loss to understand how the name R. focard Vieill.t

could have ever been applied to the present species. Levaillant’s

figure represents a bird with white throat and chest, and crimson

upper tail-coverts, whereas R. swainsoniz has the latter pure white

and the throat deep yellow. The next available name is the one

used above.

R. swainsonii (tocard auct.) ranges from southern Central

America through Colombia to Western Ecuador.

178. PrEROGLOSSUS ERYTHROPYGIUS SANGUINEUS Gould.

[Pteroglossus erythropygius Gould, P. Z. 8. 1843, p. 15 (July

1843.—Voyage of the ‘Sulphur’; the type locality, later given

as Realejo, Nicaragua, is doubtless erroneous). |

P. sanguineus Gould, Monogr. Ramph., 2nd edit., text to pl. 21

(1854—loe. ign.) ; Cassin, Proc. Acad. N. Sci. Philad. 1867, p. 109

(R. Truando, N. Colombia); Hartert, Nov. Zool. v. 1898,

p. 498 (Cachabi, Paramba, N. W. Ec uador).

P. erythropygius (nec Gould) Cassin, Proc. Acad. N. Sci. Philad.

1860, p. 136 (R. Truando).

No. 2022. gad. Noanama: 27.viii.08.—Wing 151; tail 175

bill (measured with chord) 130 mm.

Nos. 2023, 2024, 2163, 2189. 9 9 ad. Noanama: 27.viu.08 ;

Sipi: 6,14.x.08.—Wing 147-152; tail 170-182; bill 112-120

mm.

“Tris yellow; feet greyish green; maxilla greyish yellow, tip

yellow, broad culminal band and streak along cutting-edge black ;

mandible black.”

The series proves beyond doubt that P. e. sanguineus is a

perfectly valid form, the coloration of the bill being quite

constant in all five specimens. I have also examined ten skins

from various localities in N.W. Ecuador, belonging partly to the

Tring and partly to the Munich Museums.

These fifteen examples invariably have the lower mandible

black, while the maxilla, in addition to the black stripe along the

cutting-edge, shows a broad, black culminal stripe, extending over

more than the two basal thirds of its length. Cassin (/. e. pp. 109—

110) gives the same coloration for six specimens obtained on the

River Truando by Lieutenant Michler’s Expedition. Sometimes

the extreme base of the mandible is shightly clouded or marbled

with dull greyish yellow, and a small spot at the extreme tip is

soiled yellowish.

In P. e. erythropygius Gould, on the other hand, the bill is pale

yellow except a black stripe along the cutting-edges of the

maxilla, and a slight dusky tip to the lower mandible. The

culminal stripe is generally absent, though occasionally specimens

* Zool. Ilustr. iii. pl. 168 (no locality).

tone Dict. xxxiv. p. 281 (1819—based on Leyvaillant, Ois. de Paradis, etc., 11.

pl. 9).

1202 MR. GC. E. HELLMAYR ON THE

may be found with an indistinct, narrow streak down the centre

of the culmen.

P. é. sanguineus inhabits the lowlands and _hill-ranges of

Western Colombia (from the Truando southwards) and N.W.

Keuador (province Ksmeraldas : Cachabi, Paramba, Bulin, etc.)*.

P. ¢é. erythropygius replaces it in the more southern districts of

Western Ecuador: Babahoyo, Pallatanga, Chimbo, Rio Peripa,

Santa Rita, Santo Domingo, ete. The Tring and Munich Museums

possess good series of this form, showing its characters to be

quite constant f.

179. Coceyzus MELACORYPHUS Vieill.

Coccyzus melacoryphus Vieill. Nouv. Dict. viii. p. 271 (1817—

ex Azara: Paraguay).

No, 2104. g ad! Sip: /224%08— “Winey 110 tant 128)

bill 25 mm.

“Tris black, feet grey, bill black.”

Rather small, but not otherwise different from Brazilian

examples.

180. Pronopsirra puLcHRA Berl.

Pionopsitta pulchra Berlepsch, Ornith. Monatsber. v. p. 175

(1897.—San José, Rio Dagua, W. Colombia); Hartert, Nov.

Zool. v. 1898, p. 500 (Cachabi, N.W. Ecuador).

No, 2016. 2 ad. Noanama: 26.vii1.08.— Wing 146; tail 64;

bill (with chord) 22 mm.

“ Tris blue-grey, feet dirty yellow, bill white.”

A fine specimen of this rare Parrot, which was discovered

on the Rio Dagua by the late Gustav Hopke, and was afterwards

met with by Mr. Rosenberg near Cachabi, in the Hcuadorian

province of Esmeraldas.

The species is evidently restricted to the hot lowlands of

W. Colombia and N.W. Ecuador. It is allied to, but quite

distinct from, the Central American P. hamatotis Scl. & Salv.

181. Prionus mMEenstruvus Linn.

Psitiacus menstruus Linneus, Syst. Nat. 12, i. p. 148 (1766—

ex Edwards—hab. ign.—et Brisson: “ Guiane,” sc. Cayenne).

Pionus menstruus Sclater & Salvin, P. Z. 8. 1879, p. 538

(Remedios).

Nos. 2010, 2168, 2169. dg 6 @ ad. Noanama: 25.viii.; Rio

Garrapatas: 8.x.08.—Wing 180-175; tail 75-80; bill 27-

28 mm.

* Tawrence’s record of P. erythropygius from Chiriqui (Ann. Lye. N. H.N. Y. viii.

1865, p. 178) is certainly referable to some other species, perhaps P. torquatus, of

which the Munich Museum has a large series from Boquete.

+ Salvadori & Festa (Boll. Mus. Zool. Torino, xv. no. 368, 1900, p. 23) describe

a “young” bird of P. erythropygius (from Intac) as having the lower mandible and

a distinct culminal stripe black. I cannot help thinking that the specimen in

question really belongs to P. e. sanguineus, though its occurrence so far south would

be remarkable.

BIRDS OF WESTERN COLOMBIA. 1203

“Tris brown, feet whitish, maxilla black with base red, mandible

black.”

Colombian examples of this wide-spread species are apparently

not different from others taken in Cayenne, Brazil, and Venezuela.

There is perhaps generally more rosy suffusion on the foreneck,

though this is not quite constant.

182. MicrAsTUR GUERILLA INTERSTES Bangs.

| Micrastur guerilla Cassin, Proce. Acad. N. Sei. Philad. iv. p. 87

(1848.—Jalapa, Mexico). |

Micrastur interstes Bangs, Auk, xxiv. p. 289 (1907.—La

Estrella, Cartago, Costa Rica).

No. 2330. 9 ad. Noévita, Rio Tamandé (150 ft.), 24.xi.08.—

Wing 175; tail 165; bill (from cere measured with chord)

18 mm.

“Tris and feet yellow, bill black.”

This specimen, a perfectly adult bird in the “ plumbeous phase,”

except in being slightly larger, agrees with an adult from Miramar

(Chiriqui) and a couple from Cartago, Costa Rica (Underwood).

All the under surface, from the foreneck to the tail-coverts, is

closely barred black and white, as described by Mr. Bangs. The

sides of the head and the throat are pale smoky grey (the former

somewhat darker than the latter), exactly as in the Central

American skins; but the scapulars and, to a lesser degree, the

wings are more strongly tinged with chocolate-brown.

M. g. zonothorax Cab.* from Venezuela, of which I have

examined six specimens procured in the mountains around

Merida, differs from the Colombian and Costa Rican birds in

having the sides of the head and the throat dull rufescent brown,

and the quills more or less rufous-brown. Moreover, the mantle

is generally washed with chocolate-brown, whereby its close

relation to MW. rujficollis Vieill. is supported. That zonothorax

is merely a race of M. guerilla is clearly shown by a specimen

from Paramba, N.W. Ecuador, in the Tring Museum, which

combines the dull rufescent brown cheeks and throat with the

blackish wings.

M. g. interstes ranges from Costa Rica and Chiriqui southwards

to Western Kcuador (Nanegal, Paramba, Surupata, etc.).

183. ACCIPITER SUPERCILIOSUS Linn.

‘alco superciliosus Linneus, Syst. Nat. 12, i. p. 128 (1766.—

Surinam : juv.).

Accipiter tinus Sclater & Salvin, P. Z. 8. 1879, p. 541

(Remedios).

No. 2628. 2 imm. Tad6 (230 ft.), 30.iv.09.—Wing 149;

tail 105; bill 13 mm.

“Tris orange, feet yellow, bill black, base yellow.”

* Climacocercus zonothorax Cabanis, Journ, f. Orn, xiii, p. 406 (1865.—Puerto

Cabello, N. Venezuela).

1204 MR. CG. E. HELLMAYR ON THE

184, LEucoPreRNIS PLUMBEA Saly.

Leucopternis plumbea Salvin, Ibis (3) 1. p. 240, pl. vill. (1872.—

Ecuador); Hartert, Nov. Zool. ix. 1902, p. 605 (Paramba,

S. Javier, N.W. Ecuador).

No. 2103. g ad. Sipi, 150 ft., 29. ix.08.— Wing 220; tail 130;

bill 24 mm.

“Tris crimson, feet orange-red, bill black, base orange.”

This bird éoerontonee with Salvin’s description and figure with

the exception that the thighs are but obsoletely barred with

white. The other character istics, viz. the white under wing-

coverts and quill-lining, and the broad white bar across the middle

of the tail, are very well pronounced,

L. plumbea, although nearly allied to LZ. schistacea Sund, *

of Amazonia, may easily be distinguished by its much smaller size,

by lacking the white apical band £0 the rectrices, etc., ete. It was

hitherto known only as an inhabitant of Western Eeuador 7, and

is very rare in collections.

185. LeucoprerNis SEMIPLUMBEA Lawr.

Leucopternis semiplumbeus Lawrence, Ann. Lyc. N. H. N. Y.

vii. p. 288 (1861.—Panama Railroad).

LL. semiplumbea Sclater & Salvin, P. Z. 8. 1879, p. 540

(Remedios); Hartert, Nov. Zool. ix. 1902, p. 605 (Paramba, °

S. Javier, N.W. Ecuador).

No. 2566. 9 ad. Juntas, R. Tamand (405 ft.), 10.1ii1.09.—

Wing 188; tail 130; bill 22 mm.

“Tris yellow, feet orange, maxilla black, base orange, mandible

yellow.”

The skin is in every respect typical. L. semiplumbea ranges

from Nicaragua and Costa Rica through Western Colombia to the

province of Hsmeraldas, N.W. Ecuador.

186. HeERPETOTHERES CAGHINNANS Linn.

Falco cachinnans Linneus, Syst. Nat. x. p. 90 (1758—

ex Rolander: ‘‘ America meridionalis,” hab. subst. Surinam, auct.

Berlepsch, Nov. Zool. xv. p. 290).

Herpetotheres cachinnans Sclater & Salvin, P. Z. S. 1879,

p. 541 (Cauca, Remedios).

ING, ZO o AGl Vévita (150 ft.), 4.xii.08.—Wing 250;

tail 195; bill 26 mm.

‘Tris brown, feet yellow, bill black, base yellow.”

187. Lepropon PALLIATUS Temm.

Falco palliatus (Wied MS.) Temminck, Pl. Cel. livr. 23, pl. 204

(1823.—- “ Brésil et Guiane”; juv.).

* Asturina schistacea Sundevall, Ofvers. Vetensk.-Akad. Forhandl. vil. No. 5,

p- 132, note 3 (1850.—* Brasilia”).

+ Sharpe’s (Cat. B. i. p. 216) record from “ Panama” requires comfirmation.

BIRDS OF WESTERN COLOMBIA 1205

Leptodon cayennensis auct. *

No. 2721. 9 ad. Tado ea ft.), 26.vi.09.—Wing 270; tail 170

bill 29 mm.

“Tris dark grey, feet blue, maxilla black, mandible dark

blue.”

An adult bird of this wide-spread species.

188. CoLuMBA sPECcTOSA Gm.

Columba speciosa Gmelin, Syst. Nat. 1, ii 783 ete

Daubenton, Pl. Enl. 213: Cay enne) ; ches & Salvi Tit aA:

1879, p. 543 (Remedios).

No. 2476. g vixad. Noanama: 14.1.09.—Wing 187; tail 112;

bill 203 mm.

‘Tris and feet dark red, bill scarlet.”

A wide-ranging species.

189. CoLUMBA ALBILINEA ALBILINEA Bonap.

Columba albilinea (ew Gray MS.) Bonaparte, Consp Av. u

p- ol (Nov. 1854.—*“ Nova Granada ”’).

C’. albilineata Sclater & Salvin, P. Z. 8. 1879, p. 543 (Retiro).

No.2801. gad. Loma Hermosa, 4150 ft., 19.x.09.—Wing 195;

tail 140; bill 193 mm.

“Tris light brown, feet and bill lemon-yellow.”

This bird is exactly like specimens from Ecuador. North of

the Isthmus of Panama C. a. albilinea is represented by the

nearly allied C. albilinea crissalis Salvad. +, which has the under-

parts much paler and the top of the head more reddish.

190. CotumBA Goopsont Hart.

Columba goodsoni Hartert, Bull. B. O. C. xii. p. 42 (1902.—

5. Javier, Pambilar, and Carondelet, N.W. Ecuador) ; idem, Nov.

Zool. ix. 1902, p. 602 (Pambilar, 8. Javier, Carondelet, Rio

Sapayo t, Cayapas, N.W. Hcuador).

No. 2041. ¢ ad. Noanama (100 ft.), 31.viu.08.—Wing 150;

tail 113; bill 14 mm.

> tres pink, feet crimson, bill black.”

This specimen, a male in full plumage, agrees Rerieetly with two

topotypes from N.W. Heuador. As pointed out by Hartert, the

Species is a very distinct one and wey be distinguished inguin

C. nigrirostris brunneicauda Carriker $, of Costa Rica and Panama,

in having the top and sides of the head clear plumbeous grey and

* The specific name cayennensis cannot be used for this species, Falco

cayennensis Gmelin (Syst. Nat. 1, 1.1788, p. 269) being preoccupied by Falco

cayennensis of the same author (I. ¢. p. 263).

+ Columba crissalis Salvadori, Cat. B. Brit. Mus. xxi. p. 294 (1893.—Costa

Rica, etc.).

3p Erroneously spelt “ Japayo.”

§ Ann. Carnegie Mus. vi. Nos. 2-4, p. 395 (1910.—Guapiles, Costa Rica).

Prec. Zoot. Soc.—1911, No. LXX XI, dl

1206 MR. C. E. HELLMAYR ON THE

the throat light pearl-grey instead of vinous, the foreneck and

breast much paler, plumbeous washed with lilac, and the abdomen

dull vinous-brown. ‘The bright cinnamon-rufous colour of the

under wing-coverts and inner webs of the remiges serves to dis-

tinguish it at a glance from C. pluinbea bogotensis Berl. & Lev.

and C. subvinacea berlepscht Hart. *, found in the same districts.

C. goodsont is peculiar to the forest-covered lowlands (from sea-

level to about 500 feet) of N.W. Ecuador (province Hsmeraldas)

and Western Colombia.

191.: GEOTRYGON VERAGUENSIS CACHABIENSIS Hart.

[Geotrygon veraguensis Lawrence, Ann. Lyc. N. H. N. Y. viii.

p. 349 (1866.—Veragua). |

G. veraguensis cachabiensis Hartert, Nov. Zool. v. p. 504

(1898.—Cachabi, N.W. Ecuador).

Gi. v. cachaviensis Hartert, |. c. x. 1902, p. 603 (S. Javier, Rio

Sapayo, N.W. Ecuador ; crit..),

No. 2444. g ad. Noanama (100 ft.), 5.1.08—Wing 134;

tail 76; bill 182 mm.

‘“‘ Tris yellow, feet crimson, bill black.”

This specimen is decidedly darker sepia-brown on the back and

wings, and has a stronger violet-purple gloss on the upper mantle

than a single Costa Rican skin of true G. v. veraguensis. The

same differences were noticed by Hartert in a series from

N.W. Ecuador.

G. v. cachabiensis replaces the typical race in the Pacific

lowlands of W. Colombia and the adjoining parts of Ecuador

(province Esmeraldas).

192. GHorrRYGoN BOURCIERI Bonap.

Geotrygon bourcieri Bonaparte, Consp. Av. ii. p. 71 (Nov.

1854.— Lloa, Ecuador).

No. 2816. 2 ad. Pueblo Rico (5200 ft.), 27.x.09.—Wing 155;

tail 98; bill 17 mm.

“Tris yellow, feet light red, bill black.”

Compared with two fine adults from Hcuador (Aguapum) this

bird has the flanks slightly deeper rufous, and the forehead

less rosy, but these variations are very trifling.

This rare Pigeon was hitherto known only from Western

Ecuador (2 and N. Peru) and the present Hewat! extends its

range considerably to the north.

193. Rayncnortyx crnctus Salv.

Odontophorus cinctus Salvin, Ibis, (3) vi. p. 379 (1876.—Veragua;

= 6),

QO. spodiostethus Salvin, Ibis, (4) 1. p. 447 (1878.—Veragua ;

=o ye

* Noy. Zool. v. p. 504 (1908.—Paramba, N.W. Ecuador).

BIRDS OF WESTERN COLOMBIA. 1207

Rhynchoriyx cinctus Hartert, Nov. Zool. ix. 1902, p. 600 (Bultn,

Rio Bogot:i, Pambilar, N.W. Ecuador; crit.).

Nos. 2119, 2125. ¢ $§ ad. Sipi (150 ft.): 25, 28.1x.08.—

Wing 116, 120; tail 45; bill 17 mm.

No. 2126. 9 ad. Sipi: 28.1x.08.—Wing 116; tail 44;

bill 17 mm.

‘* Tris brown, feet blue, maxilla black, mandible grey.”

These specimens fully bear out the conclusions arrived at by

Hartert from the study of a series from N.W. Heuador, viz., that

O. cinctus and O. spodiostethus have been established upon phases

of one and the same species. The two skins, marked as “ ¢ ” by

the collector, agree substantially with the description ot

O. spodiostethus, having the sides of the head and throat bright

ferruginous, the chest dark cinereous, and the rest of the belly deep

ochraceous buff, the flanks finely vermiculated with dusky, and

the under tail-coverts only distinctly barred with black, ete.

The third specimen, sexed as “female,” tallies well with the

description of O. cinctus. It has the head above, nape, and chest

deep rufous-brown, while the remaining under paris are white,

broadly banded with black, except down the middle of the

abdomen; chin and upper throat are white; there is but an

indistinet, buffy whitish superciliary stripe, ete.

While there cannot be any further question as to O. cinctus

being the female, and O. spodiostethus the male of the same

species, yet the comparison of a good series from Veragua might

show the birds from the Pacific coast district of Colombia and

N.W. Heuador to be slightly different. In fact, the Sipi female

differs from Salvin’s and Grant’s descriptions in having the lower

back and rump bright rufescent brown, finely vermiculated with

dusky and spotted with black, especially down the centre, instead

of ‘‘dark grey or vinaceous, mottled with whitish.”

R. cinctus, wherever it occurs, appears to be arare bird. It has

been recorded from the Escondido River (Nicaragua), Panama,

Veragua, the Choco district (W. Colombia), and from the province

of Esmeraldas (N.W. Ecuador).

194. CHAMAPETES GOUDOTII GouDOTII Less.

Ortalida Goudotii Lesson, Man. d’Ornith. i. p. 217 (1828—

“les montagnes du Quindiu,” Colombia).

Chamepetes goudoti Sclater & Salvin, P. Z. 8. 1879, p. 544

(Retiro).

No. 2795. $ ad. Tatamdé Mountain, 4600 ft., 14.x.09.—

Wing 255; tail 245; bill 33 mm.

*“ Tris crimson, feet vermilion, bill black, bare space round the

eye dark blue.”

Identical with a specimen from Bogota. Birds from Ambato,

E. Keuador, are much brighter rufous underneath. They are

certainly subspecifically distinct and may belong to C. g. ruji-

ventris Tsch., from Peru, which I have not yet met with.

1208 MR, C. BE. HELLMAYR ON THE

195. NorHocercus INTERCEDENS Salvad..

Nothocercus intercedens Salvadori, Cat. B. Brit. Mus. xxvii.

p- 513 (1895.—Frontino, W. Colombia).

NV. bonapartii (nec Gray) Sclater & Salvin, P. Z.S. 1879, p. 548

(Frontino, Concordia).

No.2728. Qad. Pueblo Rico (5200 ft.), 16.viii.09.— Wing 203;

tail 70; bill 32 mm.

“Tris dark brown, feet grey, maxilla black, mandible grey.”

The specimen agrees with Salvadori’s description. The throat

is white, washed with pale rufescent buff on its lower portion.

The rump. upper tail-coverts, wings, as well as the abdomen, are

marked with numerous white or buffish dots.

NV. intercedens is peculiar to the Western Cordillera of Colombia

(Frontino, Concordia, Pueblo Rico). In southern Central America

it is represented by WV. frantzii Lawr.*, in the astern Cordillera

and in the mountains of Western Venezuela by 1. bonapartet

Gray ‘.

196. CRECISCUS ALBIGULARIS Lawr.

Corethrura albigularis Lawrence, Ann. Lyc. N. H. N.Y. vi.

p- 302 (Jan. 1861.—Panama Railroad).

Porzana albigularis Sclater & Salvin, P. Z. 5. 1879, p. 546

(Remedios).

Creciscus albigularis Hartert, Nov. Zool. ix. 1902, p. 604

(S. Javier, N.W. Ecuador).

No. 2144. $ vixad. Sipi, Rio Sipi, 1.x.08.—Wing 75; tail 30;

bill 18 mm.

“ Tris orange-yellow, feet dark greenish brown, bill black.”

This bird agrees with typical Panama specimens, the head above

being russet-brown, the chin and upper throat conspicuously

white, etc. There are a few cinnamon-rufous spots, but no white

bars, on the upper wing-coverts.

(San Javier, province Esmeraldas).

197. JONORNIS MARTINICA Linn.

Fulica martinicu Linneeus, Syst. Nat. 12, i. p. 259 (1766—* in

Martinice inundatis ”).

Porphyrio martinicus Sclater & Salvin, 1. ¢. p. 546 (Medellin).

No. 2713. @ ad. Tadé (230 ft.), 18.v1.09.

“Tris light brown, feet yellowish brown, bill red, tip yellow.

198. ARAmIDESs wort Berl. & Tacz.

Aramides wolfi Berlepsch & Taczanowski, P. Z. 8. 1883, p. 576

®* Tinamus frantzii Lawrence, Ann. Lyc. N. H.N. Y. ix. p. 140 (1868.—Cervantes,

Costa Riea).

+ _Tinamus bonapartei G. R. Gray, List Spec, B. Brit. Mus., y. Gallinz,

p. 97 (1867.—Valley of Aragua, W. Venezuela).

BIRDS OF WESTERN COLOMBIA. 1209

(1884.—Chimbo, S.W. Ecuador); Salvadori & Festa, Boll.

Mus. Torino, xv. no. 368, p. 40 (Rio Peripa, W. Kcuador):

Hartert, Nov. Zool. ix. 1902, p. 604 (Pambilar, Carondelet,

N.W. Ecuador).

No. 2264. 9 ad. Névita: 11.xi.08.—Wing 160; tail 70; bill

55 mm.

“Tris scarlet, feet crimson, bill green, tip paler.”

This skin agrees perfectly with others from N.W. Ecuador,

notably with an adult female from Carondelet. Salvadori and

Festa (J. ¢.) have alveady pointed out that the description in the

Cat. B. Brit. Mus. xxiii. p. 55, is not quite exact.

A. wolfi is nearest to A. mangle Spix * and agrees with it in

the clear cinereous colour of the head and neck above, and in the

whitish throat, but may at once be distinguished by its much

larger bill, deep olivaceous-brown back, this colour passing into

rufous-brown on the mantle, black (instead of greyish) lower

abdomen and thighs, much darker, ruddy brown breast and upper

belly, ete.

A. wolfi is new to the fauna of Colombia. It was only reported

as an inhabitant of Western Keuador.

199. DENDROCYGNA DIscoLoR Scl. & Saly.

Dendrocygna discolor Sclater & Salvin, Nomencl. Av. Neotrop.

p- 161 (1873.— Venezuela, Guiana, et Brasilia,” type from

Surinam, cfr. Cat. B. Brit. Mus. xxvii. p. 162).

D. autwmnatis (nec Linneus) Cassin, Proc. Acad. N, Sci. Philad.

1860, p. 197 (R. Truando).

No. 2193. 2 ad. Mouth of Rio Sipi, near Noanama, 16.x.

08.—-Wing 225; tail 62; bill 50 mm.—“ Iris black, feet and bill

blood-red.”

Typical of D. discolor, the lower hind neck being buffy grey,

abruptly contrasted with the chestnut-brown back.

D. discolor is widely distributed in South America south of the

Panama Isthmus.

200. QUERQUEDULA CYANOPTERA Vieill.

Anas cyanoptera Vieillot, Nouv. Dict. v. p. 104 (1816—

“‘ yiviére de La Plata et 4 Buenos Aires”).

No. 2150. 2 ad. Sipi: 3.x.08.—Wing 175; tail 70; bill

40 mm.

“Tris black, feet putty brown, maxilla blue-grey, mandible ~

flesh-coloured.”

201. Carso vicua Vieill.

Hydrocorax vigua Vieillot, Nouv. Dict. viii. p. 90 (1817—ex

Azara: Paraguay).

* Gallinula mangle Spix, Av. Bras. ii. p. 74, pl. 97 (1825,—Brazil).

1210 MR. C. E. HELLMAYR ON THE

Carbo brasilianus? Cassin, Proc. Acad. N. &ci. Philad. 1860,

p: 197 (R. Truando).

~ No. 2045. 9 imm. Noanama: 1.ix.08.—“ Iris emerald-green,

feet black, maxilla grey, mandible yellow.”

IV. Conclusions.

Although our knowledge of Western Colombia is still very far

from complete, yet the researches of the naturalists mentioned

in the first part furnish sufficient evidence on which to base

some conclusions as to the general character and affinities of the

avifauna of this remote district. As might be expected from its

geographical situation, the fauna of Western Colombia shows a

mixed character, being composed partly of Ecuadorian and partly of

Central American species, the former, however, by far preponder-

ating. The most interesting result derived from Mr. Palmer's

collection is evidence of close similarity of the avifauna of the San

Juan Valley to that of the province of Esmeraldas, N.W. Ecuador.

In fact quite a number of remarkable species are apparently peculiar

to S.W. Colombia and the adjoining parts of N. Ecuador, while in

the more southern districts of Western Keuador other more or

less allied forms take their place. In several instances, however,

the Chocé region has a species or subspecies of its own, while

the province of Esmeraldas is inhabited by a representative form

generally distributed over Western Ecuador.

A few species range from Panama to N.W. Heuador, but do

not go further south. Several others extend from Panama to the

Dagua district, whereas in N.W. Ecuador another species or race

is found.

The subjoined tabular lists + may serve to illustrate these facts,

which are of some importance to the student of zoogeographical

problems.

(A) Species peculiar to Western Colombia (Chocé district) and

N.W. Ecuador (province Esmeraldas).

Turdus tristis dague. Dendrornis lachrymosa rostrata,

Henicorhina inornata. Cercomacra berlepschi.

Calospiza johanne. *Caprimulgus rosenbergi.

Hlenia cinerea parambe. Polyerata rosenbergi.

Rhynchocyelus cinereiceps flavotectus. *Hucephala hwmboldti.

Mionectes olivaceus hederaceus. Pionopsitta pulchra.

Chloropipo holochlora lite. Columba goodsont.

Sapayoa enigma. Geotrygon veraguensis cachabiensis.

Carpodectes hopkei. Leucopternis plumbea.

Automolus nigricauda. etc., etc.

+ The species not represented in Mr. Palmer’s collection, but procured by previous

travellers, are marked with an asterisk (*).

BIRDS OF WESTE

N COLOMBIA. IAL

(B) Species peculiar to Western Colombia.

Heleodytes albobrunneus.

Thryophilus nigricapillus schottii.

Huphonia xanthogaster chocoensis.

i. f. fulvicrissa.

Chlorochrysa nitidissima.

Calospiza palmeri.

Buthraupis melanochlamys.

B. aureocincta.

*Ostinops salmoni.

Masius chrysopterus bellus.

Siptornis erythrops griseigularis.

Thripadectes sclateri.

Anoplops bicolor dague.

Pittasoma rosenbergi.

Phethornis yaruqui sancti-johannis.

Thalurania fannyi.

* Adelomyia cervina.

Cyanolesbia kingii subsp.

Bucco noaname.

Capito quinticolor.

C. maculicoronatus (north to Panama).

Monasa pallescens.

Chloronerpes rubiginosus gularis.

* Picumnus olivaceus granadensis.

Pteroglossus e. sanguineus

(north to the Truando, south to N.W.

Eeuador).

Odontophorus parambe baliolus.

Nothocercus intercedens.

Represented in W. Ecuador by

T. n. nigricapillus.

HH. x. xanthogaster (°)

LE. fulvicrissa purpurascens (N.W.).

C. pheenicotis.

B. rothschildi (N.W.).

B. edwardsi (also in §. Colom)ia:

Pasto).

O. atrocastaneus.

M. c. coronulatus.

S. e. erythrops.

A. b. equatorialis.

P. rufopileatwn (N.W.).

P.y. yaruqui.

T. verticeps.

A. melanogenys maculata.

C. k. celestis.

squamatus.

P. 0. harterti.

P. erythropugius erythropygius

(C. & S.W. Ecuador).

O. parambe paranbe.

known to occur elsewhere.

Chiromacheris vitellina.

Pachyrhamphus dorsalis.

Bucco pectoralis.

B. tectus subtectus.

(D) Species ranging from Panama to the Chocd district,

W. Colombia.

Cotinga nattererii.

Capito maculicoronatus.

1212

MR. C. E, HELLMAYR ON THE

(1) Species of Western Colombia represented in southern Central

America (Panama, Costa Rica) by nearly allied forms.

Western Colombia.

*Turdus tristis daque.

Heleodytes albobrunneus harterti.

TLeucolepis p. pheocephalus.

Thryophilus nigricapillus schottii.

+L. leucopogon.

* Henicorhina inornata.

+ Basileuterus t. tristriatus.

*Dacnis venusta fuliginata.

D. cayana corebicolor.

+Sporophila ophthalnica.

*Calospiza lavinia lavinia.

C. gyroloides gyroloides.

*O. johanne.

*O. larvata fanny.

Buthraupis melanoch lamys.

* Heterospingus wanthopygius.

*Hemithraupis salmoni.

}+Cacicus uropygialis.

Cyanocorax affinis affinis.

*Rhynchocyclus cinereiceps flavotectus.

+ Mionectes olivaceus hederaccus.

Myiobius sulphureipygius villosus.

+Pipra mentalis minor.

Chiromacheris vitellina.

+Tityra semifasciata columbiana.

*Lathria wnirufa castaneotincta.

*Lipaugus holerythrus rosenbergi.

*Carpodectes hopkei.

Siptornis erythrops griseiqularis.

+Hyloctistes subulatus assimilis.

+Xenops genibarbis littoralis.

* Dendrornis lachrymosa rostrata.

TD. triangularis equatorialis.

+Uyrmotherula fulviventris viduata.

+ Formicivora quixensis consobrina.

+Ramphocenus c. cinereiventris.

+Myrmelastes exsul maculifer.

Anoplops hicolor dague.

}Hormicarius analis destructus.

Pittasoma rosenbergi.

+Threnetes ruckeri fraseri.

*Chalybura urochrysa.

Chloronerpes rubiginosus gularis.

Picumnus olivaceus granadensis.

}Malacoptila panamensis poliopis.

Central America.

T. tristis cnephosa.

Hi. a. albobrunneus.

Lup. lawrencii.

T. semibadius.

T. thoracicus.

HL, leucosticta prostheleuca.

B. tristriatus melanotis.

D. v. venusta.

D. c. ultramarina.

S. aurita.

C. lavinia dalnasi.

tC. gyroloides bangsi.

C. florida florida and C. florida arcei.

C. larvata subsp.

B. arcei.

H.. rubrirfons.

HL. chrysomelas.

C. microrhynchus.

C. a. zeledoni.

R. c. cinereiceps.

MM. o. olivaceus.

IM. s. aureatus.

P. mentalis ignifera.

C. aurantiaca.

T. s. costaricensis.

Lu. clara.

Lh. holerythrus.

C. nitidus.

S. e. rufigenis.

A, s. virgatus.

X. g. mexicanus.

D. 1. lachrymosa.

D. t. punctigula.

M. f. fulviventris.

FE. q. boucardi.

R. c. semitorquatus.

MM. e. exsul.

A. b. bicolor.

EF. a. nigricapillus (Kastern Costa

Rica).

P. michleri (Panama).

T. r. ruckeri.

C. isawre (Veragua).

C. 1. uropygialis. :

P. 0. panamensis (Panama).

M. p. panamensis.

~ When also oceurring in the province of Esmeraldas (N.W. Ecuador), they are

marked with an asterisk (*): when more generally distributed in Western Ecuador,

with a dagger (t).

BIRDS OF WESTERN COLOMBIA. UALS

Western Colombia (cont.). Central America (cont.).

Aulacorhomphus albivitta pheolemus§. A. ceruleigularis.

*Pionopsitta pulchra. P. hematotis.

*Columba goodsoni. C. nigrirostris brunneicauda.

*Geotrygon veraguensis cachabiensis. G. v. veraguensis.

Nothocercus intercedens. NV. frantzii.

The evidence at hand is hardly sufficient to allow an exact

comparison of the fauna of Western Colombia with that of the

Central Cordillera, yet from what we know it appears that the

Pacific lowlands and foot-hills possess a considerable number of

peculiar elements, e. g. Supayou enigma, Cnipodectes subbrunneus,

Carpodectes hopkei, etc. Some of the species recorded only from

the Western Cordillera may turn up in the Central Cordillera ;

but, on the other hand, it is quite certain that a good many forms

that inhabit the central mountain-chain are altogether absent

from the Pacific slopes.

A thorough exploration of Southern and Central Colombia

cannot be too warmly recommended to the attention of

ornithologists, as offering a wide field for the solution of various

zoogeographical problems.

§ Aulacorhamphus pheolemus Gould, Ann. Mag. N.H. (4) xiv. p. 184 (1874.—

Concordia, Western Cordillera of Colombia (and Merida, in Venezuela—errore !)).

A. petax Bangs, Proc. Biol. Soc. Wash. xxi. p. 158 (1908.—San Antonio, R. Cali,

Western Cordillera of Colombia).

There can be no doubt that the blue-throated Awlacorhamphus, of the Western

Cordillera, has to bear the name A. a. pheolemus, which Mr. Bangs appears to have

overlooked. Although in the Cat. B. Brit. Mus. xix. p. 158, a specimen from

Merida, Venezuela, is given as type, this cannot be correct, for Gould (J. c.) clearly

descrined the Concordia bird (efr. “throat deep greyish blue”). It may be

mentioned that the Merida form was erroneously referred to pheolemus by Gould,

since a series in the Munich Museum undoubtedly belongs to A. a. albivitta, all the

examples having the throat white.

Proc. Zoou. Soc.—1911, No. LX XXII, 82

No. 100.

ABSTRACT OF THE PROCEEDINGS

oF THE

ZOOLOGICAL SOCIETY OF LONDON.*

October 24th, 1911.

Sir Joun Rose BrRAprForp, K.C.M.G., M.D., F.B.S8.,

Vice-President, in the Chair,

The Minutes of the last Scientific Meeting were confirmed.

The Sacrerary read a Report on the Additions that had been

made to the Society’s Menagerie during the months of June,

July, August, and September 1911.

Mr. JAMEs DunBAr-Brunton sent for exhibition two skins and

a mounted skull, with horns, of Bushbuck shot by him in North-

East Rhodesia.

Mr. W. B. Corton, of the Indian Civil Service, exhibited a

number of heads and horns of various species of Gazelles which

he had obtained in the Eastern Sudan, and gave a brief account

of their habits and distribution.

Mr. R. I. Pocock, F.R.S., F.L.S., F.Z.S., Superintendent of

the Gardens, exhibited two photographs of a living male specimen

of Red-fronted Gazelle (Gazella rujfifrons) which was presented to

the Society by Col. Julian Hasler in May 1908, and came from

Kano in Northern Nigeria. This specimen he proposed to make

the type of a new subspecies to be named G. rujifrons hasleri,

since it differed apparently from all previously recorded races of

that species in having the upper side of the nose pure white from

the nostrils back to the preorbital glands.

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent's Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications ; but it may be obtained on the

day of publication at the price of Stapence, or, if desired, sent post-free for

the sum of Sia Shélings per annum, payable in adyance,

My, J. Lewis Bonnorrg, M.A., F.Z.S., exhibited two “ waltzing ”

Rats (Mus ratius) which he had bred in the course of his experi-

ments. They appeared in the F, generation, and it was noted —

that the strain had become weak and degenerate, other individuals

of that generation being born blind or otherwise defective. As

the “ waltzing” character had been proved to have a Mendelian

inheritance in Mice, it would be an interesting fact that, should

this character prove to have a Mendelian inheritance in Rats, it

would be a case of the genesis of a Mendelian character brought

about by artificial conditions, e.g. environment. ‘The original

stock were normal wild individuals. Mr. Bonhote also pointed

out that the two varieties of J. rattus found in Egypt, viz.

M.r. tectorwm with white underparts and IZ. r. alecandrinus with

dark underparts, had a Mendelian inheritance, the former white-

bellied form being dominant to the dark-bellied form. These

experiments also included the study of the inheritance of a fawn-

coloured variety, hitherto unknown, which appeared as a ‘‘ sport”

from wild-caught parents. ‘This variety had also a Mendelian

inheritance, the fawn-coloured ones being recessive to both the

normal wild forms.

Mr. D. Sera-Smuiru, F.Z.S., Curator of Birds, exhibited a

spirit-specimen of a nestling Australian Regent-Bird (Serzeulus

melinus) which had been hatched in the aviary of Mr. Reginald

Phillipps, of 26 Cromwell Grove, West Kensington, during the

past summer.

Mr. E.G. Boutencrr, Curator of Reptiles, gave the description

of anew Tree-Frog from Trinidad, living in the Society’s Gardens.

The Frog, which was brought back by Dr. Lewis H. Gough in

July last, was one of the smallest of the genus Hyla, and was

vemarkable for the rapid changes in colour and markings which

it displayed.

Mr. Bruce F. Cummines. read a paper, communicated. by

Mr. T. A. Cowarp, F.Z.S., on “ Distant Orientation in Batrachia,”

based on observations and experiments made by the author in

North Devon. ‘Two species of Newts had been used for the

experiments, and the results obtained lent support to the hypo-

thesis that these batrachians possessed a homing faculty, but no

very definite instinct for detecting water, even from a short

distance. Of the factors discussed in connection with amphibian

migration, it was suggested that in regard to Newts, a com-

bination of their homing faculty and their marked tendency to

walk downhill was chiefly of assistance to them in finding water

in which to breed.

Mr. OuprieLp Tomas, F.R.S., F.Z.8., read a paper on Mammals

collected in the Provinces of Sze-chwan and Yunnan, W. China,

by Mr. Malcolm Anderson, for the Duke of Bedford’s Exploration

of Kastern Asia. The paper formed No. XV. of the series, and

would be the last on Mr. Anderson’s specimens, as he was now

returning finally to America. During his work on the exploration

he had obtained 2700 specimens, besides many birds, and had

quite revolutionized our knowledge of the area explored.

The present collection, given, as before, to the National Museum

by the Society’s President, consisted of 160 specimens, belonging

to 33 species. ‘The following were described as new :—

RHYNCHONAX ANDERSON, gen. et sp. nn.

Allied to Uropsilus sor VS, but with p’ and i, present ‘no p,.

Head and body 70 mm.; tail 67; hind foot 15-5 5; skull 21-7.

Hab. Omi-san. Type. Male. No. 11.2.1.25. M.P.A. 2504.

NASILLUS GRACILIS, gen. et sp. nn.

Allied to the above, but with p* and p, present ; no i,.

Head and body 66 mm.; tail 55; hind foot 13°5; skull 20-5.

flab. Chin-fu-san. Type. Female. No. 2566.

SOREX WARDI FUMEOLUS, subsp. n.

Larger and darker-coloured than true wardi; the brain-case

broader.

Head and body 60 mm.; tail 60; hind foot 13; skull 18-1.

flab. Wei-choe, Si-ho R. Type. Male. No. 2627.

SORICULUS IRENE, sp. n.

Allied to macrurus, but brain-case lower.

Head and body 60 mm. ; tail 90; hind foot 16; skull 17-2.

Hab. Yuen-ching, Sze-chwan. Type. Female. No. 2673.

CHODSIGOA LARVARUM, sp. n.

Near Ch. hypsibia, but brain-case narrower and higher.

Head and body 68 mm. ; tail 50; hind foot 14; skull 18°8.

Hab. Imperial tombs HE. of Peking. TJype. Female. B.M.

No. 8.8.7.21.

APODEMUS SPECIOSUS ORESTES, subsp. n.

_ Size medium. Tail long; ears medium. Colour near sepia.

Head and body 93 mm.; tail 125; hind foot 24; ear 16.

Hab. Omi-san. Type. Male. B. M. No. 11.2.1. 170.

APODEMUS SPECIOSUS LATRONUM, subsp. n.

Size large. Tail short; ears long. Colour brown.

Head and body 107 mm. ; tail 101; hind foot 25; ear 20.

Hab. 'Ta-tsien-lu. Type. Male. B.M. No. 11.2.1.156.

MiIcROTUS MILLICENS, sp. n.

Skull flattened; an extra angle on m’*; tail long.

Head and body 90 mm. - tail 53 5 hind foot 18° 5 skull 24°3.

Hab. Wei-choe, Si-ho R. Type. Male. No. 2615.

Microrus (HoTHENOMYS) MELANOGASTER BLEUSIS, subsp. n.

Tail comparatively long; m° with four inner angles,

Head and body 98 mm.; tail 55; hind foot 17; skull 24-7.

Hab. BH. of Chao-tung-fu, N. Yunnan. ype. Male. No. 2696.

Microrus (EoTHENOMYS) OLITOR, sp. n.

Near MW. melanogaster, but no extra internal angle on m’.

Head and body 82 mm.; tail 34; hind foot 16; skull 24.

Tab. Chao-tung-fu. Zype. Female. No. 2714.

Microrus (CARYOMYS) ALCINOUS, sp. n.

Like Jf. (C.) eva, but colour very much darker.

Head and body 90 mm.; tail 56; hind foot 17; skull 24.

fab. Wei-choe, Si-ho R. Type. Male. No. 2631.

_ The Sscrerary presented a paper by Mr. EH. P. Srepsine,

E.LS., F.Z.S., entitled “‘ Game Sanctuaries and Game Protection

in India,” in which the author discussed the question of the

formation of Game Sanctuaries and what had been already done

in this direction in various parts of the country. Suggested

additions to the proposed New Indian Game Act were given,

and ‘close seasons’’ for certain species recommended as being

necessary for the preservation of the game of the country.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, November 7th, 1911, at half-past Hight

o'clock P.M., when the following communications will be made :—

1. R. 1. Pococs, F.B.S., F.L.S., F.Z.S.

Lantern exhibition on the Moulting of the Arctic Fox.

fos SHES SINT Meas LEEUW

_ On the Moulting of the King Penguin (Aptenodytes pennanti)

in the Society’s Gardens. (With lantern illustrations.)

3. 7. E. Gunn, F.LS.

On the Presence of Two Ovaries in certain British Birds,

more especially the Faleonide. (With lantern illustrations.)

4, Prof. P. P. Susuxin, C.M.Z.S.

Ontogenetical Transformations of the Bill in Ardea cinerea.

5. A. D. Imus, D.Sc., B.A.

On some Collembola from India, Burma, and Ceylon, with a

Catalogue of the Oriental Species of the Order.

The following papers have been received :—

1. H. B. Preston, F.Z.8.

Diagnoses of New Species of Terrestrial and Fluviatile

Shells from British and German East Africa.

. R. LYDEKKER.

On the Milk-Dentition of the Ratel.

Structure of the Alimentary Canal of the Stick Insect

(Bacillus rosstvi Fabr.), with a Note on the Parthenogenesis of

this Species.

4. Ropert SHELFORD, M.A., B.Z.S.

Ginter, amongst the Blattide ; with a Revision of the

Genus Pr osoplecta Sauss,

5. The Rev. O. Pickarp-Campripen, F.R.S., C.M.Z.S8.

Contributions to the Knowledge of the Spiders and other

Arachnids of Switzerland.

A further Collection of Mammals from Egypt and Sinai.

7. J. T. Cunnincuam, M.A., F.ZS.

Mendelian Experiments on Fowls.

8. Prof. Grorrrey Surrn, M.A.

The Freshwater Crayfishes of Australia,

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Society oF Lonpon,

Recrnt’s Park, Lonpon, N.W.

October 31st, 1911.

No. 101.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

November 7th, 1911.

FrepDErick Ginuert, Esy., Vice-President, in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

Mr. F. Menterra Ocitvis, F.Z.S., communicated a paper by

Mr. T. E. Gunn, F.L.S., ‘‘On the Presence of Two Ovaries in

certain British Birds, more especially the Falconide.” The

author outlined the views held by the majority of English mor-

phologists on the reproductive organs of adult female birds, and

enumerated examples which he had collected during a number of

years where the right as well as the left ovary was present, and,

so far as could be ascertained, in the two cases of which sections

had been made, where the right ovary was functional. He

pointed out the extraordinary preponderance in his examples of

paired ovaries occurring in the Falconide, as compared with those

derived from any other source, and remarked that in that family

the ovaries were usually placed symmetrically one on either side

of the vertebral column and at about the same level. In examples

other than the Falconide this symmetrical arrangement was the

exception rather than the rule, the right ovary generally occupy-

ing a position almost directly below the left, in the left half of

the body-cavity, which, in the author’s opinion, suggested a half-

way home on the road leading to the final disappearance of the

right ovary.

Mr. R. I. Pocock, F.RB.S., F.LS., F.Z.S., Superintendent

of the Gardens, exhibited a series of lantern-slides prepared from

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent's Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications ; but it may be obtained on the

day of publication at the price of Szarpence, or, if desired, sent post-free for

the sum of Six Shillings per annum, payable in adyance.

photographs taken by Mr. P. W. Farmborough, F.Z.S., illustrat-

ing the colour-change and phases in the moult of an Arctic Fox

now living in the Society’s Gardens,

Mr. D. Seru-Smirs, F.Z.S., Curator of Birds, read a paper,

illustrated by lantern-slides, on the Moulting of the King Penguin

(Aptenodytes pennanti) in the Society’s Gardens. He referred

to Mr. de Winton’s paper on the same subject which appeared

in the ‘ Proceedings’ in 1898. The specimen observed by

Mr. de Winton did not moult until it had lived sixteen months

in the Gardens, whereas the specimen now in the menagerie

had moulted twice in six months.

The author stated that the new feathers were almost fully

grown before the old ones were shed, and that the latter had to

be rubbed off by the bird’s beak or feet, as they were firmly

attached to the sheaths of the new feathers. The paper was

illustrated by photographs, which showed the bird in various

stages of the moult, as well as by specimens of the shed feathers.

Dr. S. F. Harmer, M.A., F.R.S., V.P.Z.S., read a paper by

Prof. A. D. Imus, D.Sc., B.A., entitled, ‘‘Some Collembola from

India, Burma, and Ceylon, with a Catalogue of the Oriental

Species of the Order.” Four genera and twenty-eight species

were described as new, amongst the latter the most remarkable

being a form unique among Collembola in possessing a median

cercus to the fifth abdominal segment, and for the reception of

which a new subfamily was formed. The total number of Col-

lembola known from the Oriental region was stated to amount to

53 species comprised within 27 genera.

A paper on the “‘ Ontogenetical Transformations of the Bill in

Ardea cinerea,” by Prof. P. P. Susuxin, C.M.Z.S., was read by

Mr. D.Sera-Smiru, F.Z.8. The author gave a description of the

gradual development of the bill in a series of embryos and young

specimens of the Heron upon which he had made observations.

The simple rhamphotheca proved to be only a late stage of the

compound one, and the form of the Ardeine bill he regarded as

a derivative one, and discussed its resemblance to those of allied

forms.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, November 21st, 1911, at half-past Hight

o'clock P.m., when the following communications will be made :—

1. Dr. Grorrrey Suiru, M.A.

The Freshwater Crayfishes of Australia.

2. Frank E. Bepparp, M.A., F.R.S., F.Z.S8.

Contributions to the Anatomy and Systematic Arrangement

of the Cestoidea.—III. On a New Genus of Tapeworms from

the Bustard (Hupodotis kori).

(SU)

. ALFRED HE. Cameron, M.A., B.Sc.

Structure of the Alimentary Canal of the Stick Insect

(Bacillus ross Fabr.), with a Note on the Parthenogenesis of

this Species.

. G. A. Boutencer, F.R.S., F.Z.S.

Some remarks on the Habits of British Frogs and Toads,

with reference to Mr. Cummings’s recent communication on

“« Distant Orientation in Amphibia.”

5. H. B. Preston, F.Z.S.

Diagnoses of New Species of Terrestrial and Fluviatile

Shells from British and German East Africa.

6. R. LyDEKKER.

On the Milk-Dentition of the Ratel.

The following papers have been received ;—~

1. Ropert SHetrorp, M.A., F.Z.S.

Mimicry amongst the Blattide; with a Revision of the

Genus Prosoplecta Sauss.

2. The Rev. O. PrckArp-CamsBrinax, F.R.S., C.M.Z.S.

Contributions to the Knowledge of the Spiders and other

Arachnids of Switzerland.

3. J. Lewis Bonunors, M.A., F.L.S., F.Z.8.

A further Collection of Mammals from Egypt and Sinai,

4. J. T. Cunnineuam, M.A., F.Z8.

Mendelian Experiments on Fowls.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL, |

Secretary.

ZooLocicaL Society or Lonpon,

ReceEnNtT’s Park, Lonpon, N.W.

November 14th, 1911.

No. 102.

ABSTRACT OF THE PROCHEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

November 21st, 1911.

S. F. Harmer, Esq., M.A., Sc.D., F.R.S., Vice-President,

in the Chair,

The Minutes of the last, Scientific Meeting were confirmed.

The Secretary read a Report on the Additions that had been

made to the Society’s Menagerie during the month of October

issih:

Mr. R. I. Pocock, F.R.S., F.Z.S., Curator of Mammals,

exhibited two living examples of an Elephant-Shrew (JJacro-

scelides sp.) from Bechuanaland, which had been presented to the

Society by Capt. H. O. F. Littledale and forwarded by Dr. L.

Péringuey, C.M.Z.S., and remarked that these specimens, together

with another in the same consignment, but belonging to a different

species, were apparently the first representatives of the genus

Macroscelides the Society had ever possessed. After drawing

attention to some interesting structural features shown by the

animals, Mr. Pocock said that the Society was to be congratulated

upon the safe arrival of these Elephant-Shrews, because, apart

from members of the Hedgehog-family, which from being omni-

vorous are more easy to preserve in captivity, exotic species of

Insectivora are proverbially difficult to keep alive for any length

of time and are therefore seldom exhibited in the Gardens,

Dr. Grorrrey Smiru, M.A., read a paper, communicated by

the SECRETARY, entitled “‘ The Freshwater Crayfishes of Australia.”

The object of this paper was to revise the classification and

—

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent’s Park, N.W., on the Tuesday following the date of Meeting to which

it refers.. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications; but it may be obtained on the

day of publication at the price of Stxpence, or, if desired, sent post-free for

the sum of Stix Shillings per annum, payable in advance.

nomenclature of the Australian and Tasmanian Crayfishes, and to

give diagnoses of the genera and species with their limits of

distribution. The work was based on a large material obtained

from all parts of the continent and from Tasmania. Many of

the specimens had been collected by the author, but the majority

formed a very large collection brought together during the past

twenty years by Prof. Baldwin Spencer.

Four genera were recognised, Astacopsis, Cherops, Parache-

rops, and Hngeus: the first three genera included the Freshwater

Crayfishes proper, and the last-named genus contained the Land-

Crayfishes, which were not dealt with in this paper.

The geographical distribution of these genera and its bearing

upon the geographical problems of Australia was discussed, and

the view was supported that the Bassian Subregion represented

the home of the Australian Crayfish, and that Astacopsis was

nearest the ancestral form.

Mr. F. KE. Bepparp, M.A., F.R.S., F.Z.S., Prosector to the

Society, presented a paper on “A new Genus of Tapeworms from

the Bustard (Hupodotis kori).” Four complete specimens and

some fragments of this Cestode had been obtained from a

S. African Bustard in the Society’s Gardens, and the author

regarded it as a member of the group Tetracotylea, but could not

reconcile its characters with those of any other genus of that

group at present known. He briefly described its anatomical

characters and discussed its systematic position, and proposed a

new genus and species for its reception.

The SECRETARY presented a memoir by Mr. A. E. Cameron,

M.A., B.Sc., entitled “‘ The Structure of the Alimentary Canal of

the Stick-Insect, Bacillus rossii, Fabr., with a Note on the

Parthenogenesis of this Species.”

The author stated that this insect had a rather limited dis-

tribution, occurring in the south of Europe and in the north of

Africa, and that in the wild state it was not found north of

Orleans. Certain peculiarities of the alimentary canal were dealt:

with which were regarded as adaptations to the mode of life of

the species. Attention was drawn to the fact that the male was

only rarely found in the wild state, and that parthenogenetic

reproduction of B. rossii had been verified, for during four

generations the specimens kept by the author had showed no

males. The fact that the males were disappearing suggested that

parthenogenesis was not the primitive method of reproduction,

but that the species had become adapted to it through the failure

of sexual reproduction.

Mr. H. B. Preston, F.Z.S., communicated a paper based on a

collection of Terrestrial and Fluviatile Shells made by Mr. Robin

Kemp in British and German East Africa. One new genus and

thirty-four new species were described, which represented only a

very small portion of the large number of species collected

Mr. EH. G. Bounencer, F.Z.S., presented a short paper by

Mr. G. A. Boutencsr, F.R.S., F.Z.S., containing some remarks

on the habits of British Frogs and Toads, for the information of

those who might feel inclined to carry out further observations on

the subject of the migrations of Amphibia as dealt with in a paper

recently read before the Society. The Common Toad was strongly

recommended as the most suitable Batrachian on which to

institute series of experiments on Distant Orientation.

A paper on the “ Milk-Dentition of the Ratel” was received

from Mr. R. LyprexKsr, in which he described an instance of

primitive features present in the milk-dentition being entirely

lost in the teeth of the permanent series. So far as he was aware,

no such atavistic feature had been hitherto recorded in the case

of any existing mammals.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, February 6th, 1912, at half-past Hight

o’clock p.M., when the following communications will be made :—

1. Mrs. R. Harte THomas, F.Z.S.

On Experimental Pheasant Breeding.

2. J. T. Cunninenam, M.A., F.Z.S.

Mendelian Experiments on Fowls.

3. H. G. Primer, F.R.S., F.L.S., F.Z.S.

Report on the Deaths which occurred in the Zoological

Gardens during 1911,

4, J. Lewis Bonnorte, M.A., F.L.S., F.Z.5.

A further Collection of Mammals from Egypt and Sinai.

The following papers have been received :—

1. Rosert SHetrorD, M.A., F.Z.S.

Mimicry amongst the Blattide; with a Revision of the

Genus Prosoplecta Sauss.

2. The Rev: O. Pickarp-Camprince#, F.R.S., C.M.Z.S.

Contributions to the Knowledge of the Spiders and other

_ Arachnids of Switzerland.

3. Herpert L. Hawnins, M.Sc., F.G.S.

The Classification, Morphology, and Evolution of the Echi-

noidea Holectypoida.

4. H. Lysrer Jameson, M.A., D.Sc., Ph.D., F.Z.S.

Studies on Pearl-Oysters and Pearls.—I. The Structure of

the Shell and Pearls of Margaritifera vulgaris Schumacher :

with an examination of the Cestode Theory of Pearl Pro-

duction. .

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Society oF Lonpon,

RecENt’s Park, Lonpon, N.W.

November 28th, 1911.

Papers (continued).

Page

40. On a new Species of Dinotherium (Dinotherium hobleyi) from British East Africa.

By C. W. Anprews, D.Se., F.R.S., F.Z.S. (British Museum, Natural History).

(Ge TE NGTS VATA) oops ain eet we eee anee Rar eeMeney FURS ABD AISS sw sow o:ni 2 4 cto heal cote ang OcheD

41. On an Amphipod from the Transvaal. By the Hon. Paun A. Mernuen, F.Z.8.

(O15. NUTONE STENT) aa sc otis fote te NS onan etal eA CREE Seg eles AVen NE eae’ a) a's.s een Sata : 948

42. An African Rhinoceros, Klipspringer, and Gazelle. By R. Lyprxker. (Text-figs.

AQT T G38) Wee Maar co age ics aptreretae te chances Lottie ay QUIRED EC 2c Oe aR eRe 958

43. The Subspecies of the Spanish Ibex. By Prof. Ange Caprmra, O.M.Z.S. (Pls. LIL—

DONE errata eset IS AI) Se oles ieee Sree Sra Sc cloit 32 e 35 oir eo OA ee SO EamMeMmE

44. On Antelopes of the Genera Madogua and Rhynchotragus found in Somaliland. By

R. HE. Draxe-Brockman, M.R.C.S., L.R.C.P., F.Z.S8. (Pls. LV. & LVI.) ........-- O77

45, Contributions to the Anatomy and Systematic Arrangement of the Cestoidea.

II. On. Two new Genera of Cestodes from Mammals. By Franx EB. Brppanp, >

M.A., F.R.S., F.Z.S., Prosector to the Society. (Text-figs. 204-215.) ............ 994

46. Some Madreporaria from the Persian Gulf. By Rurn Harrison, Oxford. Witha

Note on the Memoir and some Further Notes on Pyrophyllia inflata by Sypney J.

Hickson, M.A., D.Se., F.R.S., F.Z.S. (Pls. LVII. & LVIII. and Text-figs. 216-221.) 1018

47. On Variation in the Medusa of Merisia lyonst. By Cuaruns L. Boutencer, M.A.,

F.Z.8., Lecturer on Zoology in the University of Birmingham. (PI. LIX. and

Mexteie se 2oBe) ay .tevers -f lsrede iste ieletors SNE LiG Cod OCR TIG ROOD ASIC SP PEA aOR OOO Be, do 1045

48. The Marginal Processes of Lamellibranch Shells. By Cyrim Crosszanp, ¥.Z.8,

(Pl. LX. and Text-figs. 229 & 230.) ....-. RES CARH Cars Soar n ean clas 1057

49. Warning Coloration in a Nudibranch Molluse and in a Chameleon. By Cynrin

ROSSINI REZESS Sete Saya eer eraies ecperetaratelchsitae Cede SR aR tant BRE CGA O ONS 1062

50. Chromodorids from the Red Sea, collected and figured by Mr. Cyril Crossland. By

Sim Crags sbmom ake CAVE Gear C Be BZN. (RIM PCIEys erseietacc are selina tienes 1068

51. On some new South African Permian Reptiles. By R. Broom, D.Sc., C.M.Z.S.

Calg EO OUR MG UNI) SG so Sock eal TERA heed aN ec A ra ne ee Lr

52, On a new Tree-Free from Trinidad, living in the Society’s Gardens. By Epwarp G.

Bovunenerr, Curator of Reptiles. (Pl. LXIV.).......... eT a ii has Serccecrs ecru 1082

Division of Birds, Zoological Museum, Munich ............ SO EEOC REC One eb ae 1084

STIG ere Ia Gein tl (a8 Oboe She O San Oeenh cio seD USS ch Ociph vont Senos lane Bee ade eMW oc i

INE ole Camron each) oa ee cols gp vasaed ee eRe O canoe BA oboe AEA Sood Jeo il

List of Contents............. ener ess NEN shea hater aol iran gdh gcabde (eee Xe oth.s CR iti

Alphabetical List of Contributors ............-- Hei aceasta MaSteR NR ay aie Senay eas ix

News Genericul erinsirnts sielecisutihsavers (atta ater sloasrsi cia esehawa ce eet lesearalsy chen seeseie Seeteent Ne te XV1il

TndextomSerentiticwNames’ \voustrrtccncts acactor tena: sheer as chsh nr) <selipia oa pepc sce) pale ats ia xix

Index of Illustrations

Tiara Micmn eee a evateh niatectetoie roles XXXVI

LIST ‘OF PLATES. —

1911, Parr IV. (pp. 869-1213).

Plate Page

XLIII.

XLIV. Aleyonaria of the Cape of Good Hope ...........-..... - 870

XLV.

XLVI. Skulls of Cynodont Reptiles ................000.....0. 893

XLVII. Tooth-germs in Macropus billardierd ........0-..0ee+s000 926

SCUMLLL. emenhencu men ayant, saistctatcieisetciale\ a lacie eats 943

XULIX.

L. ! HLWwenGng ONY xe TOUTED. es sarees cin voi iale'vm «wie ele e tyem ss eel e : 948

LI.

LIL. Capra pyrenaica hispanica. (Summer Pelage.) .......-..

LI. Capra pyrenaica victorie, § 2. (Summer Pelage.) ...... |

LIV. Capra pyrenaica victorig. (Winter Pelage.) ............

LY. 1. Madoqua phiillipsi gubanensis. 2. Madoqua phillipsi

hararensis. 8. Madoqua phillipsi ......++ BSP oe OT7

LVI. 1. Madoqua piacentinti. 2. Madogqua swaynei .......+++

ae \ Madreporaria from the Persian Gulf .........+2+..--.. 1018

MINS Variation) Merisia Wyomst 6 ice. «sees ovine <2 <a\* =) </- 1045

LX. Marginal processes in Lamellibranch Shells.............. 1057

LXI. Chromodorids from the Red Sea ............00...-: -.. 1068

enna) Fossil Reptiles from South Africa,............. 0. 0eceee 1073

LXIII. J

LXIV. Gough’s Tree-Frog (Hyla gought) 0... ..eceescceccceenes 1082

NOTICE. .

The ‘ Proceedings’ for the year are issued in four parts, paged consecutively,

so that the complete reference is now P. Z. 8.1911, p.... The Distribution

is as follows :—

Part I. issued in March.

~~, Pp ical Ul Re eae June.

eae 3) Ee ate September.

ae me Wier December.

\) ‘ Proceedings,’ 1911, Part III. (pp. 557-868), were published on

aN" September 15th, 1911.

‘The Abstracts of the Proceedings,’ Nos. 100-102, are

contained in this Part. “.

*p 970