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MM PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

ZOOLOGICAL SOCIETY

OF LONDON.

1904, vol. II.

(MAY—DECEMBER.)

PRINTED FOR THE SOCIETY,

AND SOLD AT THEIR HOUSE IN HANOVER-SQUARE,

LONDON:

MESSRS. LONGMANS, GREEN, AND CO,

PATERNOSTER ROW.

eins ae

OF THE

COUNCIL AND OFFICERS

OF THE

ZOOLOGICAL

SOCIETY OF LONDON.

1904.

COUNCIL.

(Hlected April 29th, 1904.)

His Grace Tut Duke or BeprorD, K.G., President.

Sir ALEXANDER BatrD, Br.

Wittiam T. BuanrorD, Hsq.,

Clk DSR RS. Vice

President.

GrorceE A. Boutencer, Hsq.,

E.R.S., Vice-President.

Tuomas H. Burroucues, Ksq.

Freperic G. D. Drewirt, Esq.,

M.D., F.R.C.P.

Hersert Druce, Ese., F.LS.,

Vice-President.

Cartes Drummond, Hsq.,

Treasurer.

FREDERICK GILLETT, Esq.

F. DuCane Gopman, Ksq.,

D.C.L.,F.R.S., Vice-President.

AuBERT GtntueR, Esq., M.D.,

Pu.D., F.B.S., Vice-President.

Sir Epmunp Gites Lopmr, Br.

K. G. B. Mzapr-Watpo, Hsq.

P. Cuaumers Mircuexy, Esq.,

M.A., D.Sc., Secretary.

THe Hon. L. WaAtrer Rorts-

SCHILD, D.Sc., M.P.

Howarp Saunpers, Hsq., Vice-

President.

DavipSuHarp, Bsq., M.D.,F.R.S.

OLDFIELD Thomas, Hsq., F.R.S.

Surron Timntis, Esq.

CHARLES 8. Tomes, Esq., M.A.,

E.RB.S.

Avueustus F. Wiener, Esq.

PRINCIPAL OFFICERS.

P. Cuaumers Mrrcuett, Esq., M.A., D.Sc., Secretary.

Frank E. Bepparp, Esq., M.A., F.R.S., Prosector.

R. I. Pocock, Esq., Superintendent of the Gardens.

Mr. F. H. Warernouss, Librarian.

> Mr. Joun Barrow, Accountant.

Mr. W. H. Cots, Chief Clerk.

Mr. Georce ArtHur Dousuepay, Clerk of Publications.

Mr. Arruur THomson, Assistant Superintendent of the

Gardens.

S70,6% &

LIST OF CONTENTS.

May 3, 1904.

Page

Dr. Graham Renshaw, F.Z.8S. Exhibition of a drawing of a

youne ’Adrrcamebilepliamtis.sesen.ttiaees seen ces clctina clases 1

The Secretary. Exhibition of a photograph, taken by Mr.

Brankgkiaess of ai@ uaa... ino eaeeneaicns as eebeaenece ees 2

Mr. F. E. Beddard, F.R.S. Exhibition of the brain of a

Troupial (Quiscalus versicolor) infested by worms........-

Mr. R. H. Burne, F.Z.S. Exhibition, on behalf of Prof.

Stewart, of specimens of the female reproductive organs

of certain Marsupials and photographs of a Leathery

Riche (ReMi art en ie a meme mar Renner. Sort: Ammen AAAs a 2

1. On the Osteology and Systematic Position of the rare

Malagasy Bat Myzopoda aurita. By OLDFIELD THoMAs,

HORAS RE ZS. (Ce late oooa et scodes wanes an cscass sc 2

2. Contributions to the Anatomy of the Lacertilia.—(3) On

some Points in the Vascular System of Chameleon and

other Lizards. By Franx HE. Bepparp, M.A., F.R.S.,

IPTOSeChOrRuORE ney SOCIC Yarn eme ence codes ase eee se 6

3. Notes on the Gill-rakers of the Spoonbill Sturgeon, Poly-

odon spathula. By A. D. Imus, B.Sc. (Lond.), Assistant

Demonstrator in Zoology in the University of Birming-

inenanueais(Wellayi JUL.) \ i eek Retain OHSRBA Seen taono Hae sann LaMaMOA Anat e 22.

Page

4, On the Cranial Osteology of the Fishes of the Families

Hlopide and Albulide, with Remarks on the Morphology

of the Skull in the Lower Teleostean Fishes generally.

By W.G. Ripewoop, D.Sc., F.L.8., Lecturer on Biology

at St. Mary’s Hospital Medical School, London ......... 39

May 17, 1904.

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

Menagerie im tApril i904 oo. iii socctot saniomnss is scaieesestaeeee 82

Dr. W. T. Calman, F.Z.S. Exhibition of, and remarks upon,

a blind Crustacean (Munidopsis polymorpha) from the

Hiclandkonwanzarokes@amarcles meveme-peeeeeieacas-eecaraceeeee 82

Mr. F. EK. Beddard, F.R.S. Exhibition of, and remarks upon,

a specimen of the Lizard Chlamydosaurus kingii with

HE MMOLALGVORCS cy dale hictaasnieciae wale siaisost lee tesarats no selbtawnalsbanre naletalneteirs 82

Mr. Oldfield Thomas, F.R.S. Exhibition of, on behalf of the

President, a sketch by a Chinese artist of Pére David’s

[Deere iigonsal |e eWiaE al: “SOBA, odonpnenonscoeéaceAdboaudeeaceceLodsec 83

1. On some Nudibranchs from East Africa and Zanzibar.—

Part V. By Sir C. Extot, K.C.M.G., late H.M. Com-

missioner for the East African Protectorate, F.Z.S.

(Elites TEU GME), ied teehee romance sales iesieteine eseals 83

2. Description of a new Tree-Frog of the Genus Hyla, from

British Guiana, carrying eggs on the back. By G. A.

BOULENGER, HRI). Viena.ss abate we)! o...s.:. cece. 106

3. Notes upon the Anatomy of certain Snakes of the Family

Boide. By Frank K. Bepparp, M.A., F.R.S., Prosector

OPENS SOCICLY scowcesewbce'sr ae coplsies aeciaaet creo MEME sae 107

4, On Entomostraca collected in Natal by Mr. James Gibson.

By G. Srewarpson Brapy, M.D., LL.D., D.S8c., F.RB.S.,

OMEZS: (blates Vil VIITS) Sen cateus sie cee eeemeee 121

June 7, 1904.

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

Menaeerie an) May PO04 oe stows 6.2 «brs eee RCP eeRieeets 129

The Secretary. Note on two specimens of the Orang-utan

he had seen in Paris

Seow wees reese eee seers sere sssseroesscersesescn

Dr. Giinther, F.R.S. Exhibition of, on behalf of the President,

and remarks upon, a series of mounted specimens of

hybrid) Pheasants ne Neha eect e ns once ice

Dr. F. D. Drewitt, F.Z.8._ Exhibition of, and remarks upon,

some antlers of the North-African Red Deer ............

Dr. F. D. Drewitt, F.Z.8. Exhibition of a pair of horns of

Loder’s Gazelle from Southern Algeria

ee cee recesses sesseceee

Dr. Graham Renshaw, F.Z.8. Exhibition of photographs of,

and remarks upon, a pair of Short-horned Buffaloes in

the Antwerp Zoological Gardens

eco ese eee soe esses esas eeesecees

Mr. F. KE. Beddard, F.R.S.: Exhibition of, and remarks upon,

the skull of a Cape Crowned Orane ............2..ececeeees

Mr. R. E. Holding. Exhibition of photographs of, and re-

marks upon, the antlers of the Wapiti in various stages

OEP OLN ae SALE ASE SOLE one

Mr. R. E. Holding. Exhibition of a pair of antlers of the

imishmRed Deere. 4.548. i toi oer ae aaa cea ends coe teas

Mr. R. I. Pocock. Exhibition of, and remarks upon, living

hairless specimens of the House-Mouse and Brown Rat.

Mr. R. L. Pocock. Exhibition of, and remarks upon, young

examples of the Egyptian Fat-tailed Gerbille ............

1. On some New and Little-known Butterflies, mainly from

high elevations in the N.E. Himalayas. By Lt.-Col.

J VEAL COLM Wawcnrn, 1 (Plate TXG)a oss asia. sets. .5- 50

2. On Seasonal Phases in Butterflies. By A. G. Burinr,

1 Bel oa BY 3)0 Sad LS pa ey Zi AIRED a asses iA Ue a ge OT

3. The Prey of the Lion. By Capt. Ricnarp Crawsuay,

BZ SM capa anette vaccine aeeione sh aniinc wa duacwa rcs aa Que aa casan as

4, Note onan apparently Abnormal Position of the “Brephos”

within the Body of a Skink (Chalcides lineatus). By

Frank EH. Bepparp, M.A., F.R.S., Prosector to the

DOGLOUY Fie imran nan eco kaminsen caacin, sr .itican ce ee Lats

Page

129

130

131

133

. Contributions to the Knowledge of the Visceral Anatomy

of the Pelagic Serpents Hydrus platyurus and Platyurus

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

Prosector bo, DHE SOCLELY | 2: -en-)e/02- ele o- =e eee enact str

6. On the Presence of a Parasternum in the Lacertilian

Genus TViliqgua, and on the Poststernal Ribs in that

Genus. By Frank HE. Bepparp, M.A., F.R.S., Prosector

UG, LMENSOCIEbY, vere: coe-eigect- aie «elon ariel pe eer esac

7. On the rare Rodent Dinomys branickii Peters. By Dr.

Eur A. Gorupr, 0.M.Z.S., Director of the Goeldi

Wiavisermin, Jee, (IERHIe: 2G) pas esaspooncosc0asdesasecasasococns

8. The Black Wild Cat of Transcaucasia. By C. SATuNIN,

Gui Uintits), (CLINE Si © enone oc anonaeasesce caesuadduesnnodescgcoC eco

9. On a Buffalo-Skull from East Central Africa. By R.

IUSADI SIGRID So50n scncpsnopaysdcdoee doa oe pba eaaecne so suDsHESOdouSo Od:

10. The Ichang Tufted Deer. By R. LyDEKKER...............

11. On Two New Labyrinthodont Skulls of the Genera

Capitosaurus and Aphaneramma. By A. Smrra Woop-

WARD. GO BARS. RVZSe (Plates x0) cs 2xcnIl) ar. cc.

November 15, 1904.

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

Menagerie in June, July, August, September, and

MEto ber; Mi OO4 Mee oc aesed Hamat seeoeesen eereeeee. dae ree ees

Mr. R. Lydekker. Note on a sketch of a Deer from Hainan.

Mr. F. E. Beddard, F.R.S. Note on the voice of a young

ISBiMEAIKOO caggrasoducotuereosdnorosdunssosundanedoeldoooebesoRdsesd

Mr. Frederick Gillett, F.Z.S. Exhibition of, and remarks

upon, some antlers of the Altai Stag, shed in the

Sociteuys Menagerie ci c.c.cearsen tess ceete ners ane ee eee

Dr. P. L. Sclater, F.R.S8. Note on the specimens of the

Okapi in the Congo Free State Museum ..................

Mr. W. B. Tegetmeier, F.Z.S. Exhibition of a specimen of

an Asiatic King-Crab picked up alive off the Isle of

WaT CNG a chaantiaeagin seins lunar eros sabe baje'se tin cs te see ee eee eee

Prof. J. C. Ewart, F.R.S. Exhibition of skins and de-

scription of a new species of Zebra from Hast Africa ...

147

154

170

178

179

180

181

18]

Vil

1. On Mammals from the Island of Fernando Po, collected

by Mr. E. Seimund. By Ouprrexp 'THomas, F.R.S.,

EZ Se): (Plate Nal eae cere sou. ogden scladtonsz de tclhes

On Hylocherus, the Forest-Pig of Central Africa. By

OxupFietp THomas, F.R.S., F.Z.S. (Plates XIV. & XV.)

3. On the Species of Crowned Cranes. By P. CHALMERS

Mircuet, M.A., D.Sc., Secretary to the Society .........

4, On the Mouse-Hares of the Genus Ochotona. By J. Lewis

IBONHOTE:) MESA\. RUZES i iyitad.c teeny de seieeiehs opie euboeereas

On some Edible and other New Species of Harthworms

from the North Island of New Zealand. By W. B.

Benuam, D.Sc., M.A., F.Z.S., Professor of Biology in

the University of Otago, New Zealand ...................5.

November 29, 1904.

Dr. Walter Kidd, F.Z.S. Exhibition of a drawing of, and

remarks upon, the extensor surface of the hand of a

C@lainmpaanee Re as ae sala Sah aise dale mie ieiae are cus Gade ctaten

1. Some Observations on the Field Natural History of the

Lion, By Captain Ricnarp Crawsnay, F.Z.8.. .........

. On some Nudibranchs from Hast Africa and Zanzibar.—

Part VI. By Sir C. Huror, K.C.M.G., late H.M. Com-

missioner for the Kast African Protectorate, F.Z.S.

(Ga Best NG VIBE SS UV AU. is 2 oe ese ety arura natin ors itnteetae a pace a

8. On a small Collection of Freshwater Entomostraca from

South Africa. By Roserr Gurney, B.A., F.Z:S.

(Plate XVIII.) ...... Se Sra ai MRts ae Rit pup a eect tates Mee DMAP TAvos 1h ol nT

4, On the Morphology and Classification of the Asellota-

Group of Crustaceans, with Descriptions of the Genus

Stenetrium Hasw. and its Species. By H. J. Hansen,

JED, LIDS | (Uelenves OCD oO-14)) Mop sbeocsbaednedee

5. On the Lacerta depressa of Camerano. By G. A.

BouLencer, F.R.S., V.P.Z.S. (Plate XXII.) ............

6. On Old Pictures of Giraffes and Zebras. By R. LyDEKKER.

7. On Two Lorises. By R. Livprxxer. (Plate XXIII.) ... «

Page

183

263

264

268

298

Vill

December 13, 1904.

Page

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

Menasericin November 1904 ss ..5 ee eee 347

Mr. Oldfield Thomas, F.R.S. Exhibition of specimens and

description of a new Gazelle from Palestine ............... 347

1. The Characters and Synonymy of the British Species of

Sponges of the Genus Leucosolenia. By EB. A. Mrncuiy,

HZ.S.); University College) London)... 2.2... ae 349

2. Descriptions of Thirty-two new Species of Halticine

(Phytophagous Coleoptera) from South and Central

America. By Martin JAcosy, F.ELS. ..........0.0.0.cc00e 396

3. Notes on Anthropoid Apes. By the Hon. Watrer

Rorusceitp, Ph.D., F.Z.8. (Plate XXIV.)............... 413

4. Descriptions of Indian and Burmese Land-Shells referred

to the Genera Macrochlamys, Bensonia, Taphrospira

(gen. nov.), Microcystina, Huplecta, and Polita. By

We BEANFORD, WO. He. Iie BRS. VPs:

(ETC XOXO ) Ode, .2): ae ee: so mee oan tgs eee nee 44]

5. On the Cranial Osteology of the Clupeoid Fishes. By

W. G. Ripewoop, D.Sc., F.Z.8., Lecturer on Biology at

St. Mary’s Hospital Medical School, London............... 448

ALPHABETICAL LIST

OF THER

CONTRIBUTORS,

With References to the several Articles contributed by each.

BEDDARD, FRANK E., M.A., F.R.S., F.R.S.E., Prosector to the

Society.

Exhibition of the brain of a Troupial (Quiscalus versi-

CoLommuntested by, WOEMIS. ss... cen. see eaeeeres ees asec ester

Contributions to the Anatomy of the Lacertilia.—(3) On

some Points in the Vascular System of Chameleon and

other) Wizards 5... heat Suh. Ge ee Vie aa eke sea paths facet

. Exhibition of, and remarks upon, a specimen of the

Lizard Chlamydosaurus kingit with femoral pores .........

Notes upon the Anatomy of certain Snakes of the

Wamnihy. Bot lease eeeee hae ae PSone ta atatelscichdaie ato. ots otters

Exhibition of, and remarks upon, the skull of a Cape

Crowned i Camere sos sneer cele la arr a ahas

Note on an apparently Abnormal Position of the

“ Brephos” within the Body of a Skink (Chalcides lineatus).

Page

131

145

BrepparD, Frank E., M.A. &. (Continued.)

Contributions to the Knowledge of the Visceral Anatomy

of the Pelagic Serpents Hydrus platyurus and Platyurus

colubrinus

See eseeereseser esses aesecessccesoese sere esses eeseesesseesese

On the Presence of a Parasternum in the Lacertilian

Genus Z7%liqua, and on the Poststernal Ribs in that Genus.

Note on the voice of a young Kangaroo

eee cece ern ecces eve

Benuam, W. Buaxuanp, D.Sc., M.A., F.Z.S., Professor of

Biology in the University of Otago, New Zealand.

On some Edible and other New Species of Earthworms

from the North Island of New Zealand

BuanForD, WitL1AM THomas, C.I.E., LL.D., F.R.S., V.P.Z.S.

Descriptions of Indian and Burmese Land-Shells re-

ferred to the Genera Macrochlamys, Bensonia, Taphrospira

(gen. nov.), Microcystina, Euplecta, and Polita. (Plate

KOKA)

wee estes asters eo esesoeesreseseseesersesossseeeeest sees esosseene

Bonuotr, J. Lewis, M.A., F.Z.S.

On the Mouse-Hares of the Genus Ochotona ............

BovuLencer, Grorcr Assert, F.R.S., V.P.Z.S.

Description of a new Tree-Frog of the Genus Hyla from

British Guiana, carrying eggs on the back. (Plate V.)...

On the Lacerta depressa of Camerano. (Plate XXII).

Brapvy, G. Stewarpson, M.D., LL.D., D.Sc., F.R.S., C.M.Z.S.

On Entomostraca collected in Natal by My. James

Ciloson., ui (Places Vall — Vela sen ec caci. as see oe eee

Page

147

154

179

44]

205°

Burne, RicuarpD Hiaerns, F.Z.8.

Exhibition, on behalf of Prof. Stewart, of specimens of

the female reproductive organs of certain Marsupials and

photographs of a Leathery Turtle .............:..eeseeeeee seen

Butter, ARTHUR GARDINER, Ph.D., F.LS., F.Z8.

On Seasonal Phases in Butterflies..................es0cese0e

Catman, Witn1AM Tuomas, D.Sc., F.Z.S., of the British

Museum (Natural History).

Exhibition of, and remarks upon, a blind Crustacean

(Munidopsis polymorpha) from the Island of Lanzarote,

COU TaV TEES San REPS SHON git eR mee eae NOY a RUN rn nT Relient

CrawsHay, Capt. Ricuarp, F.Z.8.

Ave Preys of the giv. 26 scn nc epnemastoncceracee aceneceane ces

Some Observations on the Field Natural History of the

DreEwirtt, Freperic GrorcE Dawtrey, M.A., M.D., F.R.C.P.,

E.ZS.

Exhibition of, and remarks upon, some antlers of the

INorth=Atacan’ Red) Weer ii. ss eins senseoeee on saseeecenseeee

Exhibition of a pair of horns of Loder’s Gazelle from

Soubhermn Algeria isosceles eee cates ee sie ects nial

Entor, Sir Cuarues, K.C.M.G., late H.M. Commissioner for

the East African Protectorate, F.Z.S.

On some Nudibranchs from East Africa and Zanzibar.

Parte, (Blabesplliin ds IVE NR: bee. Riot a idee k ast

On some Nudibranchs from Hast Africa and Zanzibar.

arty WUsta (latesmNGV plone EVEL UR sn ee

Page

264

130

xil

Ewart, Prof. James Cossar, M.D., F.R.S., F.R.C.P., F.Z.8.

Exhibition of skins and description of a new species

ol Zebra irom bast vActrica: ssh Welle 2 tye eae eae et melee)

Fawcert, Lt.-Col. J. Mancoum.

On some New and Little-known Butterflies, mainly

from high elevations in the N.H. Himalayas. (Plate IX.)

GiILLtEert, FreperRicK WILLIAM ALFRED HeErpert, F.Z.S.

Exhibition of, and remarks upon, some antlers of the

Altai Stag, shed in the Society’s Menagerie..................

Gorpt, Dr. Emit Aveust, Director of the Museu Goeldi,

Para, C.M.Z.S.

On the rare Rodent Dinomys branickii Peters. (Plate X.)

GintuER, ALBERT CHARLES Louis Gorruitr, M.A., M.D.,

Ph.D., F.R:S., V.P.Z.S.

Exhibition of, on behalf of the President, and remarks

upon, a series of mounted specimens of hybrid Pheasants.

Gurney, Ropert, B.A., F.Z.S.

On a small Collection of Freshwater Entomostraca from

Southeainicass | (Plate XG Vil) e lee ee acne. eeenee eae eaenee

Hansen, H. J., Ph.D., F.M.LS.

On the Morphology and Classification of the A sellota-

Group of Crustaceans, with Descriptions of the Genus

Page

181

134

179

158

129

298

Stenetrium Hasw. and its Species. (Plates XIX.—XXI.) 302

Page

Houpine, Ricuarp E.

Exhibition of photographs of, and remarks upon, the

antlers of the Wapiti in various stages of growth ......... 131.

Exhibition of a pair of antlers of the Irish Red Deer . 133

Imus, A. D., B.Sc. (Lond.), Assistant Demonstrator in

Zoology in the University of Birmingham.

Notes on the Gill-rakers of the Spoonbill Sturgeon,

Halonen, Goadoudia, (Ween IU.) Senkencnosoocesoncoscoacscnas 22

Jacosy, Martin, F.E.S.

Descriptions of Thirty-two new Species of Halticine

(Phytophagous Coleoptera) from South and Central

NTTOVSTENGEY, herd ic Coico ae OER An nO AP MIE dt GASSED BION Me em er ON 396

Kipp, Water, M.D., M.R.C.S., F.Z.8.

Exhibition of a drawing of, and remarks upon, the

extensor surface of the hand of a Chimpanzee ............ 263

LypEKKER, RicHArp, B.A., F.R.S., F.Z.S.

On a Buffalo-Skull from East Central Africa ............ 163

AD nes Tike axes over a binnnineyel IDisteyp) Re cable seosnbabosoudsdecgoceadess ae 166

Note on a sketch of a Deer from Hainan ............... 178

On Old Pictures of Giraffes and Zebras ................5 339

Oni Two Worisess) (Plate XxX ne eee sna te seces seen ne 345

Mincutn, Prof, Epwarp Aurrep, M.A., F.Z.S.

The Characters and Synonymy of the British Species

of Sponges of the Genus Lewcosolenta ......ccescecscesncncens 349

X1V

MircHeELL, Perer Cuatmers, M.A., D.Sc., Secretary to the

Society.

Exhibition of a photograph, taken by Mr. Frank Haes,

Ob a Qua eaig Sih eT ile BW curl aay cate canter ee eee eee ets

Report on the Additions to the Society’s Menagerie in

PASTEL 9 OA iets asc tustles sepaauanicd. ee A cbyashs fk Ceara eee

Report on the Additions to the Society's Menagerie in

IM ctiyafi 9 OA ee RRAG SET, RENE. MOB Selon atk htatoles abet «a Mobste ame

Note on two specimens of the Orang-utan he had seen

sir, J Aaya vi fans af ra eres har a SOR eed ET a SOE NTS 518

Report on the Additions to the Society’s Menagerie in

June, July, August, September, and October, 1904 ......

On the Species @it Orronanee! OveNES ssacaccuacooncascccnscor

Report on the Additions to the Society’s Menagerie in

IW@wenmnloce WOOL coopnssadccoseonacouncos onognosoduoccoatacsbooedded

Pocock, Recinatp Inygs, Superintendent of the Gardens.

Exhibition of, and remarks upon, living hairless speci-

mens of the House-Mouse and Brown Rat ..................

Exhibition of, and remarks upon, young examples of

the Egyptian Fat-tailed Gerbille ...................000eseseeee

Rensuaw, Granam, M.B., M.R.CS., F.Z.8.

Exhibition of a drawing of a young African Elephant.

Exhibition of photographs of, and remarks upon, a pair

of Short-horned Buffaloes in the Antwerp Zoological

Garden Se eee ote gcc BU asthe okibaiie Cai etnie cake ee rears

Page

129

133

Page

RipEewoop, Waurer G., D.Sc., F.L.S., F.Z.8., Lecturer on

Biology at St. Mary’s Hospital Medical School,

London.

On the Cranial Osteology of the Fishes of the Families

Hlopide and Albulide, with Remarks on the Morphology

of the Skull in the Lower Teleostean Fishes generally .... 35

On the Cranial Osteology of the Clupeoid Fishes ...... 448

Roruscuinp, The Hon. Lionen Water, M.P., D.Sc., Ph.D.,

E.Z8.

Notes on Anthropoid Apes. (Plate XXIV.)............ 413

Satunin, Constantin, C.M.Z.S.

The Black Wild Cat of Transcaucasia ..........0+.0;e0000. 162

Sciater, Pain Luriuny, M.A., D.Sec., Ph.D., F.BS.,

WSs, EZ:

Note on the specimens of the Okapi in the Congo Free

Steen Meuse uae sy. aiischisd nc ced tac cule cam etencson clare anne eee 180

TEGEIMEIER, WILLIAM BuRNHARD, F.Z.S.

Exhibition of a specimen of an Asiatic King-Crab

picked up alive off the Isle of Wight ........................ 181

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

On the Osteology and Systematic Position of the rare

Malagasy Bat Myzopoda aurita. (Plate I.) ............... 2

Exhibition of, on behalf of the President, a sketch by

a Chinese artist of Pére David’s Deer from Hainan ...... 83

XV1

: Page

THomas, OLDFIELD, F.R.S., F.Z.8. (Continued.)

On Mammals from the Island of Fernando Po, collected

by, Vir Ee Semaund).)((Plate: XeMi) eens ech eee eeeet ese 183

On Hylocherus, the Forest-Pig of Central Africa.

(Plates DV XV) ne eee ouics senna celts nane eee Ree 193

Exhibition of specimens and description of a new

Gazelle from Palestimes se 2.-ceeae eee eee ee eee eee 347

Woopwarp, ArtHur Smita, LL.D., F.R.S., F.Z.8.

On Two New Labyrinthodont Skulls of the Genera

Capitosaurus and Aphaneramma, (Plates XI. & XII.). 170

Hist OF Mina Es:

1904.—Vor. II.

Plate

ll, Wipes, COUGH eGo uboenoDG0o orcs cc onposocuoaedot

JUL, GRIMS OE IRON OU, 5 Ra aoc coco oop eacaseangedone

Il. 1. Notodoris minor. 2. Trevelyana coccinea. 8. T’. cey- |

Uavaved, 4 I, CROCE weacboobrnovogr sono bcodon ooo \

IV. 1. Trevelyana bicolor. 2. Nembrotha cristata. 3. N. >

affinis. 4. Martonia levis. 5. Teeth of (a) Bornedla |

digitata, (b) B. excepta, and (c) B. simpler ........ a)

\o JEDI COU Acaceugeucvenodoveeevnoobpoudcodncouns

VI.

WAG, \ 1Benserrnastneeony xo INEHENL Sob Ga oagonaoconcvccueonvan

VIII.

1X. Butterflies from the N.E. Himalayas ................

GQ ID UCTONS OREITHO aco eos csc nuongeon soso onucnoDodon

XI. Capitosaurus stantonensis .....-----+ ++ +++ eeee see

XIl. Aphaneramma rostratum 1... cee eee cece

XIII. Scotonycteris bedfordt...... 1 se eee rece ee teen es

XIV. | apa :

Xv. ( Hylocheeris meimert2hagent vesserssancrarsccnccrne é

XVI. 1. Phyllidia nobilis. 2,3. Hervia lineata. 4,5. Facelina }

lineata, 6. Stiliger varians. 7,8. Elysia marginata . |

XVIL 9. Madrella ferruginosa. 10, 11. Fucelina lineata. t.

12. Stiliger irregularis. 13, 13a. Placobranchus |

ocellatus. 14-17. Elysia dubia. 18. LE. marginata . |

Proc. Zoou. Soc.—1904, Vou. II. b

268

Plate

XVIII,

XIX,

XX.

XXI.

XXII.

XXIII.

XXIV.

XXY.

XVill

Page

South-African Freshwater Entomostraca ............ 298

1. Stenetrium armatum Hasw. 2. S. mediterraneum, )

NE SD) BMS CAH ACH Op TION ARON Ilr eo co bike aw ak ade

1. Stenetrium serratum, n. sp. 2. S. occidentale, n. sp,

GUMS RONLVILENSE USD Wiles ss cvenaiatcta here ee eee

1. Steneirium antillense, n. sp. 2. S. stamense, n. sp.

3-6. Various Asedlota

CCC aC eI Ti Ca I Yat Tt CCCI arya Tar)

Lacerta depressa Camerano

ote tea ke vs BieL cht oie eS eae 332

RACES Of JuOVISES:,)-soryeraterara anja wroieie cust tea te eee 345

Sumia vellerosus (Gray) (very old male) .............. 418

Macrochlamys and similar Land-Shells from India .... 441

LIST OF TEXT-FIGURES.

1904.—Vot. II.

Page

1. Afferent renal system of Chameleon vulgaris ..... 0.00. s eens 8

2. Certain anterior arteries and veins of Chameleon vulgaris...... 12

3. Visceral arteries and veins of Pygopus lepidopus..........++0+ 13

4, Renal venous system of Pygopus lepidopus .........+..+200e% 15

5. Hepatic portal system of Pygopus lepidopus..... Soe HiRes 17

6. Part of hepatic portal system of Pygopus lepidopus .......... 18

7. Some arteries of Phelsuma madaygascariensts .......e++.-000: 19

Sb ORRIN CEI ADSL 6 ong MOMS tole AWOS AO DCEO Dee OdeooOns 38

Sb Joa TE COTES TENG OE OE IMMA mmds sk eomone do bone boooe 40

10. Elops saurus, hyopalatine arch, opercular bones, &c., with

MORNE? ONS) 94 51 Sbig'd BB elo ie ERisecctis Wet chanel ccalee wallse irr taibey che or anemet ee 40

LC Sos: sours tic oramciiial SKE letOM list tke patel ate cle oie oe) auaregeiete 42

12 Cheyanne GEA Calla Eee UR BUMES 4 5 ols he S CoD MOOS Coad oboous 43

13. Megalops cyprinoides, right side of skull .............- 00005 46

14. Megalops cyprinoides, hyopalatine arch, opercular bones, and

TRACT GUA MCE L eee odhostbcceceosdosesoer sod oat 46

Tas CieniOn io eit ook eMss GOROTCOTOUS nis dodacaconeaedeodden sean 48

16. Albula conorhynchus, right side of skull ..................-: 50

17. Albula conorhynchus, hyopalatine arch, opercular bones, and

Maaichible MOtGisTOe macs. oteee eee ses Git rtcwaae ovale eigen alee (ne 50

18. Albula conorhynchus, hesgatneemacel SIKGIGIOMN 565506 886 beccud abn | OH

19, Part of intercostal arterial system of Eryx jaculus .........+.. 109

20. Part of intercostal arterial system of Eryx jaculus............ 110

21. Certam abdominal veins in Eryx conicus .......... 002 eves 114

22, Certain abdommal veins im Bry jaculus 2... eens ne 115

23. Liver and certain adjacent veins in Bryx conicus ......++.00+ 118

DAMAtntlers Gk VWapith irom phioxOoTaplse vale c miei: ool sane ses 132

25, Dissection of Chalcides lineatus..... Sc RRR Wiarton ODOR 146

Page

26. A portion of the internal surface of the lung of Platyurus

COULDNINUS Hare stirs nants ee aise se OE RP 150

27. A portion of the internal surface of the lung of Hydrus

PNOLYUFUS se acai ee ee Mas obs PO EE EE ie 152

28. A portion of the internal surface of the non-vascular part of the

nines CHE Ne) EONS (MUM US “a escdecenesocncc¢ancasas5e 153

Soe Nodommnal xibsot Miligualscuncovdes tyes tseee eerie 156

30. Tihqua seincoides, ventral flap of musculature................ 157

31. Front view of skull of male Buflalo, Bos (Bubalus) caffer

TEOCNEROBY eg oa te ahaa ROT ee Ue 6 FCO CU ec 165

32. Left lateral view of adult male skulis of Elaphodus michianus (A)

andeHAchangenstsi()) warren cecil ean ee eee 167

03. Front view of adult male skulls of Llaphodus michianus (A) and

EE MCCHONGENSIS (1S) 5 age enioutep es ake eukey. Were /hies aie eaten aes 168

34. Capitosaurus stantonensis, hinder view of occiput, restored .... 172

30. Ward’s Zebra, to show long ears and face-stripes ............ 181

36. Ward’s Zebra, to show “ sadheon ” and pao dorsal band .... 182

SiemeleandKor Balear.CamegulOnunune ere meinen. Hic see eee 202

TOMA MolMes QUEM CIGZOUChICE HS ene aa teens ear ie ie einen 202

BY), GLSNGL Ot VAC (ROROMIID A decbegecasgdagngseuagnccun4on 203

Ai miveadvol Baleanica cece a tery can doy is tenn A aon See 203

4). Maoridrilus mauiensis. A spermatheca ................-+++: 223

A2. Maoridrilus mauensis. A penialicheetay.. 5.6... cee doe ee 223

43. Maoridrilus mauiensis. Tip of penial cheta ................ 224

44, Maoridrilus mawiensis. 'Tip of penial cheta ................ 204

45. Octochetus michaelsent. Spermatheca............:00e+-2+0 225

46. Dinodrilus beddardi. View of part of the fore-body.......... 227

47, Dinodrilus beddardi. Spermatheca .............00+0+-+00-- 228

AS; hododralus edulis. Viembrall view tps y= ace «1 eee asisteteysieue= 232

AO hhodod rls cds e) Aymem inion recy isp eieirit ae yearn ieee 233

50. Rhododrilus edulis. The prostate of the left side ............ 233

oleiohododilus edulis. A penialichretay wage isin ieee -waeiekreiek 233

52. Rhododrilus edulis. The tip of a penial cheta .............. 233

53. Rhododrilus edulis. Diagrammatic sketch of a section through

Ge PORNO DOES sa. case ny eR acs: « cay cnt acted eee ene 234

Op LULododi ius Cause, speriathecar ne see ante rae 234

55. Rhododrilus bestt. Ventral view of the clitellar segments .... 236

56. Ehododrilus bestt. An enlarged view of one of the tubercula

OG ALTOS Cle SM eke oes Oh eRe SEL ES a8 0 bt Oo 236

57. Rhododrilus bestx. Portion of prostate..............++.--++- 236

Dow LodonrviusiOeste.. Au penial chester piasacni sree eer eee 237

59. Rhododrilus bestt. Tip of penial cheta ...........-..0+-0-. 237 -

60. Rhododrilus besti. Tip of penial cheta, side view............ 287

Gltwuhododnemsbestt. A ispermabtheca 20. 4. pcre ase een 237

62. Tokea esculenta. Ventral view of clitellar region ...........- 241

33, Tokea esculenta. Enlarged view of the male pore ............ 241

64. Tokea esculenta. A somewhat diagrammatic drawing of a

dissectioniolatehwormie nn eee ee en ere 242

SOK

Page

65. Tokea esculenta. An asymmetrical arrangement of the

POLO SALES |v oleae een mMent te cael der ole ilchelecoce MMe Rede walt muaeh Ste eter = 243

66. Tokea esculenta. A diagram, compiled from serial sections,

showing the course of the sperm-ducts .................. 244

Cie Token esculenta. . Spermat neces cisin-l ejel-yelvows acpedeske 244

68. Tokea sapida, similar view to that in text-fig.62 .......... 245

GOR Uokea sapida.., Spenmuat bie cammeryectn wetetyelleuerlets ort ater 245

70. Tokea urewere, similar view to that in text-fig. 62 .......... 247

71. Tokea urewere. View of the anterior end of the prostate .... 248

ies Lokea wreweres jp Spermarhecaly iieiss.q-ter de ae eR eae rere ot 248

73. Tokea huttont, similar view to that in text-fig.62 .......... 248

74. Tokea hutiont. View of the prostates ................505: 249

103, HokeatputionimpsSpenm atheca tyiestien sao» oc ioniielstals sey alan 249

76. Tvkea suteri, similar view to that in text-fig. 62 ............ 250

(0. Male) oie | Syren, A We sae beioo gon bobcat coop odT 250

(8. Tokea kirkt, similar view to that in text-fig. 62 ............ 251

AOS Loko kirhiy 1 SperMmirxknecay «Asked <hs sekie steicen s, tieiens) eee 251

80. Tokea maorica, similar view to that in text-fig. 62 .......... 253

81. Tokea maorica, the prostates and the gland of the tuberculun

AUG AIL OUTS oes cara cc'c SENN AVS ALN CRY RL CE eM heer Sneha 254

Sh, AOAC) TUBES TS NOPANENANO Gs occ es oscs6 so odme ae De ooKaN 254

85 *, George the Fourth’s Nubian Giraffe at “Windees 5 Re 340

86. Bane the Fourth’s Nubian Giraffe in its native country .... 341

87. Grom of Capen (i) Grirathes) we \ te ratermneaeteie cho ils pec css teueveneers 342

88) Mountain Zebra... 02. ink 6 oe elt 343

89. Queen Charlotte’s Mountain Zebra ........... eee eee ee eee 344

90) Skull and horns\of, Gazella tmenrvlliy 2 sn =). 348

oN , 364

92. | Spicules-of Lewcosolenta complicata.......ecuesceseeees 367

93. 369

fe Spicules of Lewcosolenta variabtlis ....... cece cee eee eee 1 a

96. Spicules of Sycon sp. and Lewcosolenia variabilis ............ 381

97. . - . 388

93. Spicules of Leucosolenia botryoides .................... 390

99. Skull of Gorilla gorilla (Savage & Wyman). (Side view.) .. 416

100. Skull of Gorilla gorilla (Savage & Wyman). (Front view.).. 417

101. Skull of Gortl/a gorilla matschiet Rothsch. (Side view.) .... 418

102. Skull of Gorilla gorilla matschiet Rothsch. (Front view.) .. 419

103. Skull of Gorilla gorilla diehh Matschie ..........5.0....00 420

104) Skull of Gorzlla beringend Matschie <2... 3)... somes ss ec 421

IOS), Jaleo oye Gaus Liha, 7 seb ose sooo osoeeb os usuDGde 423

106. Skull of Sima satyrus Linn. (Fully adult.) .............. 428

107; Skull of Sunvw satus Winn. (Here adult.) i) 2. sane. e ee 424

IOS, Tani Out Seana walierasue (EN) adoccuoc0e¢eancvausocccuds 425

% By an oversight in numbering, text-figures 83 and 84 have been omitted.

XXil

Page

109. Left canine tooth of 1. Stmia satyrus Linn.; 2. Simia vel-

CEG OSUS AU CUBA) aire at asec esiniss'alo olen Ses gi ora Glee aE RTO ce 426

110. Simia satyrus schweinfurtht (Gigl.) cee. Oe ee eee 497

111. Head of Stnia pygmeus raripilosus Rothsch.. 2 oe. . ce eae. 428

112. Head of Simia satyrus marungensis (Noack)......4. wes nag} 431

143° Head. of Simid:pygmeus Schrebero: viaiv. cs liscia. nian, sae) 432

114. Head of Stmea pygmeus chimpanse Matschie .. 0202.0. 0004. 433

J15. Head of Sima koolookamba (Du Chaillu)...... 22.0. e es eee 45

116. Skull of Pongo pygmeus forma agrias (Schreber). vs... oo.8. 437

117. Skull of Pongo pygmaeus forma pygmeus ee) Vise noo) 438

Ja82 Cranium of, Chitocentrusidorabi) at. wealy wliinse . wor node? 449

If. Chirocentrus. dorah, ught side ofigkull 1) 22/2. . snosioi pady) 451

120, Chirocentrus dorab, hyopalatine arch, opercular bones, «and

mandible. ..CQ .gitvizat. af Jadtwod.woly splice busine pou) 452

121. Chirocentrus dorab, hyobranchial skeleton .y5....2)s. 0800. 453

122% Cranium of. Clupen jintao! ni.ladt. of weir celudia. says pedo) 455

128° Clupea finta, right side of skull ....cradisagaga ...Wi poy) 456

124, Clupea finta, hyopalatine arch, opercular bones, and mandible . 457

125. Clupea harengus. A. Hyobranchial skeleton. B. Fourth and

fifth, branchial arches.of, the right side ........ .wisyVoodwe. . 458

126% Cranium.of Chatoéssus erebt ....svadinmeqe. . wasxoont pode} 464

12%. Chatoéssus erebt, melt sidelofiskulliidu VY. elisuol, ol) agian . 465

128. Chatoéssus erebt, hyopalatine arch, opercular bones, and man-

CDN a edoip tare rsyeycisislie,e = eiaiMteds HRB EGREICD (3) AGEs) 10. GOT 466

129. Chatoéssus erebi, hyobranchial skeleton .........00. 6000000. 468

130%; Cranium of. Dussumierta acutailo®, sistinall, eoltalngd. wpen' 469

IGIS Dussumierta, acuta, right side\of skully\ysos) Jo.ccio, bos. Dail: 470

182. Dussunveria acuta, hyopalatine arch, opercular bones, and

mandible .........% eset oor DUDINGUOD, DUTIOAMN AL 10 ZAMAN, : AT]

1835 Cranium of Engrauls encrasicholus.........02550 sce sess ses 478

134. Engraulhs encrasicholus, right side of skull .........5,25555 474

185. Engraulis encrasicholus. A. Hyopalatine arch and mandible.

B. Fourth and fifth branchial arches.» C. Hyoid ...s0... AT5

eGo Cranium of (Covlia asus... sen cmwcneeect te. a caster pa erates 478

em Covlionnasus mich t side) of Slswlly eerie ener ered 479

138. Coilia nasus, hyopalatine arch and mandible oy... so... 0. ee. 481

189) Cotha nasus, hyobranchial skeletow 2). o.. ye ee 48]

1405 Cramunvoi Chanos salnvoneus. s6\. WN. W OR. WT) pute 483

141). Chanos:salmoneus, right side ‘ofiskull ww wsinononoed Je lies 485

142: Chanos salmoneus, hyopalatine arch and mandible .......... 486

143. Chanos salmoneus, hyobranchial skeleton... vee eee 487

LIST OF NEW GENERIC TERMS

PROPOSED IN THE PRESENT VOLUME (1904, vor. I1.).

Agasicles (Coleopt.) .........

Aphaneramma (Rept.)

Dinodriloides (Vermes) ......

Ectocyelops (Entomostr.) ...

Page

5896, 400

170, 1738

226

esses ae

Page

Sophraenella (Coleopt.)...... 396, 405

Taphrospira (Mollusca) ............ 441

Mokeay(Viermes)) s-..ces--0--eee 220, 240

PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

1904, Vol. II. (May to December).

May 3, 1904.

G. A. Boutencsr, Esq., F.R.S., Vice-President,

in the Chair.

The Secretary exhibited, on behalf of Dr. Graham Renshaw,

F.Z.8., an outline-drawing of a young African Elephant showing

an unusual development of the hairy covering of the body.

Dr. Renshaw had supplied the following notes on the specimen

in question :—‘“ The Elephant was a small calf presented by

M. Verreaux to the Jardin des Plantes Museum, Paris, the locality

being somewhat loosely given as ‘the Cape of Good Hope.’ No

date appeared on the label, but asa skeleton of a female LAinoceros

simus (mounted in the adjoining gallery of Comparative Anatomy)

was labelled ‘ Ed. Verreaux, 1846, perhaps it might be assumed

that the Elephant was also presented about that date. In any case,

it had evidently been in the collection for very many years. The

hairy covering showed a remarkable development in many parts of

the body. In this specimen the tusks had not begun to appear at

the time of death. It would be very interesting to ascertain if

these abnormally hairy individuals tend to lose this covering with

advancing years. Similar instances have been noted in the

Indian species, and brought to the notice of the Zoological

Society. Thus, I understand that a photograph of a mother and

calf, taken in Ceylon and showing considerable development of

hair, was exhibited at a former meeting of the Society; that

attention was also drawn to an individual (sold in 1882) which

Proc. Zoot, Soc.—1904, Vor. II. No. I. 1

2 MR. OLDFIELD THOMAS ON THE [May 3,

lived in the Gardens and was remarkable for the hair along its

back and on its head; and that the National Collection contains

a young calf, remarkably hairy, born in the Regent's Park col-

lection in 1903. All these seem to be referable to the Indian

Hlephant, so that, as the first recorded instance of the same

phenomenon in the African species, this note may be of interest.”

The Secretary also exhibited a photograph, presented to the

Society by Mr. Frank Haes, of the female specimen of the Quagga

which had lived in the Society's Gardens.

Mr. F. E. Beddard, F.R.S., exhibited the brain of a Troupial

(Quiscalus versicolor) in the hinder part of both cerebral hemi-

spheres of which was an entangled mass of Nematoid worms

lying below the pia mater. The bird was reported to have

dropped down suddenly from its perch “in a fit.”

Mr. R. H. Burne, F.Z.8., on behalf of Professor Stewart,

exhibited and made remarks upon specimens of the female

reproductive organs of the Marsupials Perameles obesula, Tricho-

surus vulpecula (one specimen from a virgin in which the septum

between the median vaginz was still intact, and another from an

individual that had borne young, in which the septum was

absorbed), and Dasyurus viverrimus, during pregnancy, showing

the increased size of the uteri, due mainly to enormous thickening

of the mucosa. These specimens had been presented to the Royal

College of Surgeons by Mr. James P. Hill, of Sydney University.

Mr. Burne also exhibited photographs of a Leathery Turtle

(Dermochelys coriacea), lately purchased by the Royal College of

Surgeons. The individual, which came from Japan, was a female,

measuring 4 ft. 4 in. in length. ©

The following papers were read :—

1. On the Osteology and Systematic Position of the rare

Malagasy Bat Myzopoda aurita. By Otprinip THomas,

F.R.S., F.Z.8.

[ Received April 9, 1904.]

(Plate 1.*)

In 1899 the British Museum was fortunately able to acquire

a specimen of the exceedingly rare and remarkable Bat Myzopoda

aurita, which had been obtained at Tamatave, Madagascar, by

Dr. Tuchébaud.

This animal was first described by Milne-Edwards and Grandi-

* For explanation of the Plate, see p. 6.

Pa. I QOS, Woll.t elT.

Bule, Sons and Danielsson, Ltd.

MY ZORODA AUIR WTA

1904, |. MALAGASY BAT MYZOPODA AURITA. 3

dier in 1878*, and further descriptions of the type specimen

were given by Dr. Dobson 7, but in neither case was there any

account of the skull or skeleton, nor were any figures published

to illustrate its many peculiarities. It is therefore thought that

some figures of the animal, with an account of its osteological

characters, will be of service to students of the Chiroptera.

Of the external characters little further notice is necessary

beyond drawing attention to the figures given (PI. I.) of the ear

(fig. 1) and of the peculiar mushroom-shaped process (fig. 1 a) at

its outer base, an organ unknown in other Bats.

I would, however, express my opinion that the “ irregularly

square lobe in the usual position of the tragus or slightly in

front of it, continuous above with the keel of the ear-conch”

(Dobson), does actually represent the tragus of other Bats,

coalesced in this case with the inner base of the ear, an arrange-

ment unique within the Order.

The penis is short, slender, pointed, and without a bone; the glans

very slender, styliform, about 2 mm. in length, the opening of

the urethra at its base above; the prepuce wrinkled and quite

naked externally, a very unusual character.

The palate has a single undivided ridge, convex forwards,

running across between the anterior premolars, and six pairs of

divided ridges between the larger cheek-teeth (see Pl. I. fig. 8).

The skull is short, broad, smooth, and rounded, in a superficial

view not unlike that of a Chilonycteris, though shorter-muzzled.

The brain-case is proportionally large, smooth, spherical, unridged,

not abnormally raised above the face-line. Nasal notch shallow,

quite unlike the deep notch characteristic of the Vespertilionide ;

but, unfortunately, the specimen is too old to show the nasal and

other sutures, so that the exact disposition of the bones cannot be

described. Premaxille united between and in front of the rather

irregular palatine foramina, but not bearing teeth in the middle

line; their structure and that of the anterior nares very similar

to what is found in WVatalus. Posterior nares low from above

downwards, much obstructed by the large rounded keel of the

vomer, which is ankylosed to the palatine in the middle line;

opening of nares narrowly U-shaped, the front edge of the opening

level with the front of m.° Pterygoids narrow, divergent, ending

in long, curved, hamular processes. Two large circular basi-

sphenoid pits present, one on each side between the antero-

internal corner of the bulla and the back of the pterygoids ; their

dividing septum T-shaped in section, its buccal surface broad,

smooth, and in level continuation with the mesopterygoid fossa

and the basioccipital, but narrowed upwards, where the floors of

the two pits approach each other closely. Bulle and cochlez of

medium dimensions. Basioccipital smooth and broad, quite filling

up the space between the bulle.

* Bull. Soc. Philom. 1878, p. 220.

+ P.Z.S. 1878, p.371. Report on accessions to Chiroptera during 1878-1880, p. 23 :

British Association Reports, 1880.

4 MR. OLDFIELD THOMAS ON THE [May 3,

Teeth of the normal insectivorous structure ; the reduction in

size of the incisors, two anterior upper and median lower pre-

molars as in Myotis, though the resemblance is clearly quite

superficial.

Dimensions of the skull :—

Greatest length 16 mm. ; upper length in middle line 14-4 ; basal

length 13°5 ; zygomatic breadth 11:8; intertemporal constriction

4s breadth of brain-case 9:6; palate, length 5-8, breadth inside

m.” 3, outside m.° 6:4; front of upper canine to back of m.* 6° 35

front of lower canine to back of iy, (Ofc

The skeleton as a whole, including the structure of the sternum

and pelvis, appears to be remarkably like that of certain Vesper-

tilionide, notably Scotophilus ; but I suppose this to be due to its

being of a primitive nature, without any noteworthy specialisations

inthe way of ankyloses or hypertrophy of parts. Wonderful

differences are to be found in these respects in various genera of

Chiroptera *, but I have not studied the subject sufficiently to

form an opinion as to the systematic value to be attached to these

modifications. Fortunately, Mr. Gerrit Miller is now engaged on

a general revision of the classification of Bats, and is dealing fully

with the skeletons, so that I do not propose to do more than give

a short description of that of J/yzopoda for incorporation in his

work,

The vertebral formula is C. 7, D. 13, L. 5, 8. 4, C. 8; total

37, plus a cartilaginous rudiment of a ninth caudal vertebra.

None of the vertebre are ankylosed together, except of course

those of the sacrum, thus widely differing from the condition in

Natalus. Hight of the ribs are connected with the sternum by

costal cartilages, the ninth, tenth, and eleventh also having car-

tilaginous continuations not reaching to the sternum ; the twelfth

and thirteenth are practically without such continuations.

The sternum is simple and little specialised ; the presternum

normal, with a rather elongate manubrial process (fig. 2 a), which

is but little expanded terminally ; the broad sterno-costal plate T

between the presternum and first rib, so characteristic of Bats, of

average dimensions; mesosternum ‘slender, 6 mm. in lensth,

slightly keeled, the costal cartilages of the 3rd to the 7th ribs

articulating with its sides at subequal distances, that of the 8th

joining it side by side with that of the 7th at its posterior end ;

xiphisternum 2°8 mm. long, slender, scarcely broadened terminally.

Proportions of scapula as shown in the Plate (figs. 2 & 4), its antero-

internal angle produced into a well-marked process projecting

downwards and inwards towards the vertebral column. Coracoid

about 3 mm. in length, therefore not long for a Bat of this size,

simple, slightly falciform, not expanded terminally.

The pelvis is on the whole very similar to that of Scotophilus

* The peculiarly ankylosed vertebral column of Natalus and the highly modified

sternum of Kerivouwla are examples.

+ The homologies and proper name of this bone are not as yet definitely settled.

Tt is the “7” of Winge’s figure on p. 44 of his Chiroptera of Lagoa Santa (E Mus.

Lun@3si, 1892), where its rclation to the epicoracoid of Parker is discussed.

1904.] MALAGASY BAT MYZOPODA AURITA. 5

temmincktwt in its proportions and the angle at which it is set.

Tt is even less expanded across the ilia, which are rounded instead

of being concave above. Its length, dorsally, is 9 mm. ; its breadth

at the acetabulum 5:4, and in front across the ilia 3°8; its depth

from front to back 4:6. The symphysis is more slender, though

still firmly ossified, and the obturator foramen is smaller. The

pectineal process is unusually slightly developed, far less than in

any Bat that I have had the opportunity of examining, and is

indeed hardly to be called a process.

In its entirety, therefore, the skeleton of Myzopoda is remark-

able for the simplicity and non-specialisation of the different

parts. In all other Bats’ skeletons that I have examined (not, I

confess, a very large number, but still representing most of the

groups) there are some specialisations, such as unusual ankyloses or

peculiar development and expansions of processes, but in M/yzopoda

none of these are present, while even normal Chiropterous

characters, such as the elongation of the coracoid and the pectineal

process on the pelvis, are at a minimum.

It will be evident from the above description, from those of

Milne-Edwards, Grandidier, and Dobson, and from the figures

now published, that Myzopoda is a most remarkable and peculiar

kind of Bat, and that it must certainly form a special Family,

the Myzopodide, though the affinities of this Family are by no

means clear. Both Milne-EKdwards and Dobson referred Wyzopoda

to the Vespertilionide, forming of it a group equivalent in value

to the Plecoti, Vespertiliones, and Minioptert. Now, however, that

Mr. Miller has taken Vatalus and Thyroptera out of the Vesper-

tilionide *, the chief reason for leaving it in that Family has been

removed, for it is only to those genera that Myzopoda could be

supposed to be related.

On the whole, it appears to me probable that I/yzopoda is most

nearly related to the Natalide and Mormoopidey, being a

descendant—specialised in some characters and primitive in

others—of the common ancestors of those groups. The occur-

rence of sucking-disks both in Myzopoda and Thyroptera is no

doubt a coincidence, for these organs would appear to be com-

paratively recent specialisations, On the other hand, the

possession of three phalanges in the middle finger in the same

two genera is presumably a primitive character, for both Phyllo-

stomatide and Mormoopide possess three, and the loss of the third

one in Vatalus and Furipterus of the Natalide { is one of the

characters in which that Family shows an approach to the still

more highly-developed Vespertilionide.

* “History and Characters of the Family Natalide,” Am. Nat. xii. p. 245 (1899).

+ As has already been done by various other authors, I would regard the

“ Lobostomine” of Dobson as forming a Family distinct trom the Phyllostomatide.

t Mr. Miller states that the long terminal phalanx of the two-phalanged Natalus

and Furipterus “is divided into two in Thyroptera,’ but without very definite

embryological evidence it would be difficult to accept this homologisation. The

terminal bar of cartilage beyond the second phalanx in the Vespertilionide is a

hint at a different homology, and one more in accordance with the usual ideas on

the subject.

6 MK. F, E. BEDDARD ON THE | May 3,

The interest of the relationship of this Madagascar Bat to

purely American groups is obvious, as adding one more to the

forms of that island which show the same affinity.

But the alliance of the Myzopodide to either the Natalide

or the Mormoopide is by no means close, and there can be no

question that the Family is quite a distinct one. As a diagnostic

point, the coalescence of the tragus with the ear-conch is note-

worthy, for almost every Family of Chiroptera can be diagnosed

by the development and structure of this organ, and its unique

condition in Myzopoda is an index to the special peculiarity of

the group.

Still more interesting will it be, as bearing on the question of

a southern connection between the faunas of the Old and New

Worlds, if the New Zealand Mystacops should prove to be a

member of the same series of forms. Its three-jointed middle

finger and the general structure of its skull and dentition point

in this direction, while its exserted tail and warty lower lip

suggest the Mormoopide quite as much as the Emballonuride, to

which it has been usually referred. Pending the publication of

Mr. Miller’s researches on the classification of Bats, however, I

do not wish to do more than suggest a possibility that M/ystacops,

like Myzopoda, may prove to have American affinities.

EXPLANATION OF PLATE I.

Myzopoda aurita.

Fig.1. Side view of head. +.

la. Mushroom-shaped process of ear, enlarged.

2. Sternum and scapula. +4.

2a. Side view of manubrial process of presternum.

3. Pelvis. +.

4, Scapula and humerus from behind. 2.

5, Skull. 2,

6. Skull, basal view. +.

7. Skull, side view. 3.

8. Palate-ridges. +.

2. Contributions to the Anatomy of the Lacertilia.—(3) On

some Points in the Vascular System of Chamaleon and

other Lizards. By Franx HE. Bepparp, M.A., F.R.S.,

Prosector to the Society.

[Received March 14, 1904.]

(Text-figures 1-7.)

The present communication is a continuation of a paper laid

before the Society in March last * dealing with the venous system

in a few genera of Lacertilia. In the present paper I deal with

the venous and also, to some extent, with the arterial systems of a

* “Contributions to the Anatomy cf the Lacertilia, No.1,” P.Z.S. 1904, vol. i.

p. 436.

1904. | ANATOMY OF THE LACERTILIA. (Sa

number of Lizards, more particularly with Chameleon, of which I

have had the opportunity of dissecting four specimens belonging

to the common species, Ch. vulgaris. The other genera, with which

I deal in a less comprehensive way, are Pygopus and the Geckos

Phelsuma and Tarentola. Of the genus Pygopus we possess, so far

as I am aware, no knowledge of the vascular system. Chameleon

has lately formed the subject of some investigations on the part

of Prof. Hochstetter, to whose memoir due reference will be

made in the course of the following pages. Inasmuch as that

anatomist was unable, through a deficiency in injection, to give

much account of the hepatic portal system, I am able to add

something to the existing knowledge of that aberrant lizard, as

well as to confirm a good many of the facts elucidated by Prof.

Hochstetter with regard to other tracts of the venous system.

CHAMA4LEON VULGARIS.

Of this species I have dissected four individuals, of which one

was a male and the rest females. The male specimen was fully

injected in both venous and arterial systems: one of the female

examples was injected from the anterior abdominal vein, and the

renal vessels were successfully filled as well as the intestinal

portal system. Iam therefore able to offer some facts concerning

both the arterial and venous systems. The latter has been partly

described by Hochstetter *, who has specially studied the renal

afferent and efferent veins, with a description of which I shall

begin. As that anatomist has pointed out, they differ con-

siderably from those of other Lizards; but I shall have occasion

to point out in a subsequent page that Pygopus resembles the

Chameleon in one important particular. J am further able to

note those points in which the veins in question show individual

variation. With Hochstetter’s account I find myself quite in

agreement ; there are, however, a few details to which he does

not refer. The afferent renal vein, which is formed by division

of the caudal into the two afferent renals below the kidney,

receives a number of oviducal veins of which I give a full account

later. It also receives at least one vein from the dorsal parietes

before the ischiadic joins it. The anterior abdominal vein arises

from the afferent renal just at the line of division between the

anterior wider and the posterior narrower region of the kidney.

From it immediately a branch is given off which divides into two,

of which the posterior supplies the dorsal parietes. The anterior

branch is the one referred to by Hochstetter as joining the pos-

terior vertebral in front of the kidney (A in text-fig. 1, p. 8). It

appears to me that this vein may be looked upon as the equivalent

of the lateral abdominal vein in other Lacertilia, for instance in

Iguanay. If this homology be not accepted, then the vein in

question is wanting in Chameleon.

a Peinase zur Entwickelungsgeschichte des Venensystems, &c., ’ Morph. Jahrb.

1X. p. rap

‘ y ‘ On the Venous System in certain Lacertilia,” P. Z.S. 1904, vol. 1. p. 489.

8 MR. F. E, BEDDARD ON THE [May 3,

There are, however, several points of resemblance between the

two veins. In Chameleon, as in Iguana, for instance, the lateral

abdominal vein is connected with the suprarenal bodies on each

side. They both arise from the anterior abdominal vein shortly

after the connection of the latter with the ischiadic vein.

Text-fig. 1.

Afferent renal system of Chameleon vulgaris.

A, vein communicating anteriorly and posteriorly with B, afferent renal. Ant.4bd.,

anterior abdominal; Od., oviducal veins; Z.K., left kidney; S7., suprarenal

body ; S7.v., suprarenal veins,

(N.B.—The branches from the afferent renal vein to the kidney are not indicated.)

The continuation of the afferent renal into the posterior verte-

bral is shown in the accompanying drawing (text-fig. 1). In one

specimen, the injected male, I found the vein only on one side of

the body, the left*. In the others it was present on both sides,

and usually unequally developed on the two sides; in one the

* A similar asymmetry occurs, as will be pointed out shortly, in Pygopus.

1904.] ANATOMY OF THE LACERTILIA. 9

left was rather longer than the right, and the reverse was the

case in another specimen. I presume that these veins are the

posterior cardinals ; they lie exactly in the same straight line as

the two azygos veins anteriorly, which are admitted to be the

anterior section of the posterior cardinals. It receives branches

from between the ribs running superficially over the musculature.

These differ on the two sides of the body in the individual where

they were best shown, a large specimen in which I describe later

the oviducal veins. On the right side, close to the point where

the posterior vertebral loses itself in the parietes, it receives two

thinnish veins from the parietes superficially. Further back, and

close to the anterior end of the suprarenal body, a thicker vein,

fed by two branches which run backwards and forwards respec-

tively along the parietes, enters it. On the left side there is only

one anterior vein debouching into the posterior vertebral. The

corresponding vein to the thicker posterior vein on the right side

opens directly into the suprarenal body, as explained later and as

illustrated in the figure to which reference has already been

made.

I mention later that in Pygopus there is a vein obviously

corresponding to the posterior vertebral. There is also, as I have

described, a similar vein in Jguwana* which arises from the

parietes close to the median side on the left, and forms one of the

afferent suprarenal vessels. In J/guana, as in most Lizards, the

suprarenal body lies a long way in front of the kidney, hence

the absence in that form of a connection between the posterior

vertebral and the kidney.

In a more fully mature individual of large size, which I owe

to the kindness of Mr. J. F. Ochs, F.Z.S., the veins of the kidney

region were very successfully injected, and enable me to describe,

which was impossible in the other specimen, the oviducal veins.

There was a slight difference on the two sides of the body; on

the left side three oviducal veins reach the posterior vertebral

vein in front of the kidney, one crossing the suprarenal body to

do so. In the region of the kidney itself three oviducal veins

reach the afferent renal in front of the anterior abdominal, and

at least two behind it. On the right side, the most anterior

oviducal vein takes up, before reaching the posterior vertebral

vein, a vein from the dorsal parietes anterior to, but in the same

straight line with, the posterior vertebral at the point where it

plunges into the thickness of the parietes.

Some of the oviducal veins take up a branch from the supra-

renal body before joining the posterior vertebral ; in other cases

the efferent suprarenal veins open directly into the posterior

vertebral.

A remarkable fact about the oviducal veins is the anterior

termination of the longitudinal vein running along the entire

oviduct into which they open.. This vein could be readily traced

* Toc. cit. p. 443.

10 MR. F. E. BEDDARD ON THE [May 3,

forwards beyond the funnel of the oviduct along the membrane

which supports the oviduct, and which ends anteriorly in a trans-

verse ligament tying the anterior border of each liver-lobe to the

parietes*. The vein on each side passes along this ligament and

enters the liver-lobe, forming thus a part of the hepatic portal

system. I have not seen a description of a similar state of affairs

in any other lizard, and I have not myself observed anything of

the kind. These vessels may take the place of the dorsal parieto-

hepatic trunk or trunks, which, as Hochstetter correctly notes,

are apparently absent in Chameleon. Physiologically they would

appear to be equivalent, since they convey blood from the dorsal

parietes.

LEpigastric Veins.—These veins were properly injected only in

one out of the four specimens which I have been able to examine.

The principal vein of the series, as in Z%liqua, is the median

epigastric. But the two lateral epigastrics are not absent, though

of diminished importance as compared with their condition in

Iguana and in Varanus. They arise from the anterior abdominal

of each side behind the fat-body, and run along the body-wall

dorsally of the fat-body on each side. They are small vessels,

and seem to end in branches of the much more important median

epigastric. The latter vessel arises from the anterior abdominal

just at the anterior end of the fat-body, to which it gives off a

branch ; whether there is a branch to the other (the left) fat-

body, [ do not know. The epigastric passes forwards near to the

middle line and to the right of the anterior abdominal as far as

the point where the anterior abdominal receives the portal vein.

At this point it opens into the anterior abdominal; the main

trunk, however, diminished in calibre, still continues its forward

course, and seems to be connected with the ventral parietal

affluents of the liver, though I am unable to make an accurate

statement as to the mode of its connection. It gives off in its

course a good many branches, which seem to anastomose with

similar branches of the anterior abdominal.

‘The anterior abdominal vein of each side is, like that of

Varanus (as made known by Hochstetter 7), a direct continuation

of the vein from the hind leg of its own side. Hach receives a

branch from the kidney and dorsal parietes, which has been

already referred to. A peculiarity of the Chameleon, as com-

pared with at least some Lacertilia{, is that the two anterior

abdominal roots unite to form the single median unpaired trunk

before they reach, and receive all the six branches from, the

paired fat-bodies. The anterior abdominal vein pursues a median

course between the two closely approximated fat-bodies; and

here several more or less regularly arranged branches from

each fat-body reach the unpaired region of the anterior abdominal.

* “Qn some Points in the Anatomy of Tupinambis teguexin,” P. Z. 8. 1904,

vol. i. p. 465.

+ Loc. cit. p. 467.

- ft Beddard, “On the Venous System in certain Lizards,” P. Z.S. 1904, vol. 1. p. 436.

1904. ] ANATOMY OF THE LACERTILIA. i

It receives minute branches from the parietes throughout its

whole course to the liver. Contrary to what is found in most

Lizards, the portal vein joins the anterior abdominal at a long

distance from the liver, in fact about halfway between the origin

and termination of the vein. The system of veins of the bladder

is connected with the anterior abdominal. A median vessel

leaves the bladder anteriorly and joins the undivided region of

the anterior abdominal. Another branch (?0n each side) runs

from the hinder part of the bladder to the right anterior abdo-

minal behind the entrance of the fat-body veins.

Suprarenal Portal Veins —As a general rule, there appears to

be no distinct suprarenal portal system: the suprarenal body

receives branches from the posterior vertebral vein given off along

the course of the latter, which have been already referred to. But

on one side of one individual there were separate suprarenal

portals. This is illustrated in the accompanying drawing (text-

fig. 1, p. 8). On the right side of the body two veins collecting

blood from the parietes, instead of joining the posterior vertebral

vein, pour their contents directly into the suprarenal body of that

side. The more usual absence of an independent suprarenal

system is to be explained, as I imagine, by the fact that the

suprarenal bodies are closer to the kidneys than they are in

some other Lizards, such as /guana, where the suprarenal system

is quite independent.

Dorsal Aorta and its Branches.—The subclavian arteries (text-

fig. 2, p. 12) spring from the right aortic arch, and I observed a

difference in two specimens dissected as to their mode of origin. In

the one the two arteries arose independently—that of the right side

being anterior to the left. In the other specimen the twosubclavians

sprang from a common trunk. Both subclavians give off vessels

to the parietes (shown in the drawing referred to). The right-

hand artery gives off a single trunk which immediately divides

into two, one going to the parietes, the other supplying the

cesophagus and running there in company with a branch of the

azygos vein of that side, ‘The left subclavian gives two branches

to the parietes following each other, which are not paired tubes

as is the case with the intercostals given off from the aorta

itself. The right aorta gives off two pairs of intercostals before

it joins the left. The left aorta, on the other hand, gives origin

to gastric trunks, of which I counted three before the junction

of the two aorte. The gastric trunks of the common aorta arise

alternately from either side of that artery, and are disposed in

pairs supplying naturally each side of the stomach. There are

eight of these arranged in four pairs anterior to the origin of

the mesenterics.

The intercostal arteries are for the most part strictly paired,

the two arteries for a given vertebra arising exactly side by side.

Occasionally, however, there is an irregularity, one of the two

arteries arising a little in front of the other.

As is the case with most Lizards, the gastrosplenic artery

12 MR. F, E. BEDDARD ON THE [May 3,

arises behind the superior and inferior mesenterics, and crosses

over them shortly after its origin. In two out of three specimens

the two mesenterics arose from the aorta by a common trunk; in

the third they arose separately. The succeeding arteries I have

dissected only in one individual, where they were asymmetrical.

Text-fig. 2.

Certain anterior arteries and veins of Chameleon vulgaris.

Ao., aorta cut short just after union of right and left aortic arches; a, 4, c, paired

intercostals; Az., azygos vein; J., jugular; @., esophageal artery and vein ;

R.Scl., L.Scl., right and left subclavian arteries.

On the right side there first arise two arteries for the ovary and

oviduct; then comes the renal. On the left side the ovarian

artery is followed by another trunk which divides into renal and

ovarian, and which lies behind the renal of the right side.

PYGOPUS LEPIDOPUS.

I have examined only one example of this species, which was

not injected. The reptile, however, had died in very good con-

dition for my purposes, for the veins were turgid with blood.

1904. | ANATOMY OF THE LACERTILIA,. 13

Arteries.—The aorta gives off eight branches to the stomach

along its whole course. The next branch which arises therefrom

Text-fig. 3.

Visceral arteries and veins of Pygopus lepidopus.

ant.abd., anterior abdominal vein; 4, ¢, d, e, branches of portal vem; g., gastric

affluent ; gast., gastric arteries of last pair; g hp. gastrohepatic ; i.c., Inter-

costal; od., oviducal; ov., ovarian and suprarenal arteries; 7., renal; R.K.,

right kidney.

goes not only to the stomach but to the liver and spleen. It

arises in a rather curious fashion (see text-fig. 3). The vessel

14 MR. F, E. BEDDARD ON THE [May 3,

springs from the ventral surface of the aorta, just in front of a

pair of intercostals; it crosses one of these intercostals, but blends

with that artery at the point of crossing. The next visceral

artery to arise from the aorta is the oviducal of each side; these

are strictly symmetrical, and each arises, not independently from

the aorta, but from the intercostal of its own side. Next arises

the cecocolic artery, which is crossed shortly after its origin in

the usual way * by the subsequently arising celiac trunk, which

is, it will be observed, a single trunk. Upon this trunk follows

the ovarian and suprarenals, of which one arises independently

from the aorta and the other from the intercostal of its own

side. Then occurs a long gap, the next trunk to arise being the

right renal, which springs from an intercostal. So, too, and a

little further back, does another oviducal artery.

On the whole, therefore, the most remarkable feature of the

arterial system of Pygopus appears to be the origin of many of

the arterial branches from the intercostals instead of directly

from the aorta. Noteworthy, too, is the large number of gastric

arteries, which is perhaps to be looked upon as associated with the

snake-like form of this lizard.

There being only rudimentary hind limbs, the system of the vene

renales advehentes is much simplified (Text-fig.4,p.15). The caudal

vein divides into two branches, one for each kidney. Each vein

soon divides into three branches, 7. e. the anterior abdominal,

the lateral abdominal, and the afferent renal. This last vein,

in the case of the right kidney, runs over the gland, giving off

branches, nearly to its anterior extremity. On the left side the

termination of this vein was particularly interesting, as showing

a distinct resemblance to the Chameleon. As in the latter

reptile, the vein does not end upon the kidney, but is prolonged

beyond it for some little distance, and is lost in the parietes to the

left of the middleline. It clearly represents the posterior vertebral

vein, and its presence on one side of the body only is, it will be

remembered, occasionally paralleled in Chameleon. At about the

middle of the kidney a vein from the oviduct (on each side)

reaches the afferent renal.

The efferent renals arise at first as a single trunk very near to

the posterior end of the kidneys, and of course between them.

This trunk divides into two before reaching the middle of the

kidneys. The left efferent renal receives first of all a vein from

the posterior region of the left ovary, and then passes in close

contact to, receiving branches from, the left suprarenal. It forms

the right efferent renal just in front of the left and just behind

the right suprarenal. Before their junction the right efferent

renal receives a branch from the posterior region of the right

ovary. Into the vena cava thus formed by the union of the two

efferent renals opens first of all, and at about the middle of the

right suprarenal, the anterior left ovarian vein, into which opens

* See Hochstetter’s account of visceral arteries in Lacertilia in Morph. Jahrb.

vol. xxvi. p. 213.

1904. ] ANATOMY OF THE LACERTILIA. 15

a branch collecting blood from the parietes and the left oviduct.

Just beyond the right suprarenal a vein from that gland opens

into the vena cava, and immediately beyond that again the

anterior right ovarian vein, which—nearer to the ovary than

in the case of the left ovarian veins—receives a branch from the

oviduct and the membrane connecting this with the lung. The

vena cava then pursues its course, without receiving further

branches, to the liver.

Text-fig. 4.

Renal venous system of Pygopus lepidopus.

ant.abd., anterior abdominal; K, right; KK’, left kidney; Zv., lateral abdominal

vein; od., oviducal; v.r.a., vena renalis advehens; v.c.i., vena cava; ep.,

epigastric.

Suprarenal Portals—These veins are asymmetrical. The right

suprarenal body has two afferent veins and the left only one.

The right-hand vessels pass dorsally of the vena cava; that of the

left suprarenal ventrally of the left vena renalis revehens.

16 MR. F. E. BEDDARD ON THE [| May 3,

The azygos veins are to be found on both sides of the body;

but that of the right side is much the larger, and extends back,

for the space of about eight ribs, some little way beyond the

anterior end of the lungs. On the left side there is a very

slender azygos, which might readily be missed on account of its

delicacy. This latter receives, before entering the left jugular, a

branch from the cesophagus. In Chameleon, as I have already

pointed out, the esophageal vein enters the right side of the

heart via the azygos vein.

Lateral Abdominal Veins.—These veins arise at the junction

of each root of the anterior abdominal with the vena renalis

advehens. They do not arise from the anterior abdominals

themselves as is the case with most Lizards. Their course along

the body-wall is, however, like that of many other Lizards. On

the right side the vein is connected superficially by slender

branches with the dorsal parieto-hepatic veins of that side.

Anteriorly the veins die away.

Hepatic Portal System.—As in most, but not all, Lizards,

nearly the full complement of hepatic portal vessels is present

in Pygopus. It receives blood, that is to say, from the anterior

abdominal, from the epigastric, the stomach, and the dorsal parietes.

There are only wanting independent vessels from the ventral

parietes present in many Lizards and so marked a feature of

Chameleon.

The epigastric vein appears to be single; if lateral epigastrics

are present they are small—unless, indeed, the vessels which I

homologise with the lateral abdominals of other Lacertilia are

really to be looked upon as lateral epigastrics. The single epigas-

tric lies to the left of the middle line. It is connected with the

anterior abdominal by two branches just behind the liver. It

gives off a large number of branches to the liver. I counted

eight of these altogether: they extend along the whole of the

liver, but do not arise at regular intervals, and are of unequal

size. Anterior to the liver is a tract of the epigastric which

gives off, at any rate, one branch to the vena cava before passing

headwards and ending in a way that I have not been able to

ascertain. It is noteworthy that here, asin Phelswma, in both

of which there are no marked lateral epigastrics, the median

epigastric takes up a portion of the duties of the lateral epigas-

trics, and conveys some of the blood from the posterior abdominal

region straight to the heart.

The dorsal parieto-hepatic vessels in this lizard are numerous

and large. Altogether three of them enter the liver, and at

least two of them are connected, as already mentioned, with the

lateral abdominal veins by fine branches running superficially over

the parietes. There are also three on the left side, more anteriorly,

which communicate with the liver via the stomach.

There are four gastrohepatic vessels, which arise separately from

a continuous longitudinal trunk running along the stomach, which

is fed not only from the vessels of the stomach itself, but also by

1904. ] ANATOMY OF THE LACERTILIA. ly

four vessels emerging’ from the parietes on the left side of the

vertebral column: these have just been referred to. This longi-

tudinal trunk, after giving off the four gastrohepatic vessels,

divides posteriorly into two branches (4, in text-fig. 5), which run

Text-fig. 5.

Hepatic portal system of Pygopus lepidopus.

Ant.Abd., anterior abdominal; A, origin of G, gastric branch of portal ;

L, liver; Pv., portal vein; Sé., stomach.

side by side, and are connected by cross-anastomoses. One of

these branches joins the general portal system ; the other enters

the anterior abdominal vein just where it receives the portal

vein.

Proc. Zoou. Soc.—-1904, Vou. IT. No. II. 2

18 MR. F. E. BEDDARD ON THE [May 3,

It is a noteworthy fact about the liver of this lizard that

a good many of the longitudinally running veins passing through

it appear at intervals on its surface, instead of being covered

throughout as is the general rule.

The portal vein has the following affluents (see text-fig. 3, b, c,

d, e, p. 13, & text-fig. 6, Spl.):—a gastric branch which really

belongs to the anterior abdominal system and has been already

referred to; just posterior to the liver a splenic vein; behind

this two veins near to each other, one of which is splenic,

the other gastric; still more posteriorly a large vein which

Text-fig. 6.

Part of hepatic portal system of Pygopus lepidopus.

Ao., aorta with chief visceral branches; G‘b., gall-bladder; Z, liver; P, pancreas:

P.v., portal vein; Sp/., spleen supplied by one artery and two veins; V.c.i.,

vena cava.

is formed of two branches, one of which is gastric, the other

intestinal ; behind this again another vein from the small

intestine joins the common trunk; the last branch of iraportance

is one from a dilated, almost globular, region of the small in-

testine. The main trunk then pursues a straight course along the

rest of the gut.

PHELSUMA MADAGASCARIENSIS.

I have a few notes only to offer upon the vascular system of

this Gecko, of which I have examined only a single male

specimen.

1904. | ANATOMY OF THE LACERTILIA. 19

The two subclavian arteries (text-fig. 7) arise independently

from the right aortic arch, the right subclavian a little in front

of the left. In their course within the body-cavity they are not

Text-fig. 7.

Some arteries of Phelsuma madagascariensis.

Ao., aorta at junction of right and left arches; Az., azygos vein; G, gastric

arteries ; Int., ceecocolic artery; Scl., subclavians; Sz., mesenteric; Spl., spleen.

concealed by musculature as is the case with these arteries in

Tiliqua. here are three gastric and cesophageal arteries of

20 MR. F. E. BEDDARD ON THE [May 3,

secondary importance in front of the main gastrosplenic artery.

Then follow, after a considerable gap, the two mesenteric arteries,

which cross in the usual Lacertilian fashion *, the anterior sup-

plying the cecum and large intestine. The second gives rise to

two main branches immediately after it has passed under the

first.

Hepatic Portal System.—In addition to the anterior abdominal

vein, concerning which I] have nothing to say, and the median

epigastric, which I describe elsewhere, the liver receives a single

gastrohepatic vein and a considerable series from the dorsal

parietes. There appear to be none from the ventral parietes

independent of the epigastric. There are, however, no less than

three dorsal parieto-hepatie vessels, of which two are larger than

the third. This system is thus more conspicuously developed

than in some Lizards.

Epigastric Vein.—I succeeded in discovering only the median

epigastric vein in Phelswma ; whether the others are present or

not, [am unable to say. This vein joins the anterior abdominal

posteriorly, as in most Lizards, and does not seem to be prolonged

further back—to the fat-bodies, for example.

Anteriorly it joins the liver after receiving an important

branch from the ventral parietes. The epigastric system is,

furthermore, represented by two or three veins from the

ventral parietes, arising anteriorly to the entrance of the median -

epigastric into the liver, which do not communicate with the

liver, but open into the vena cava in front of it. I observed

a slight connection between the epigastric and one of these veins.

Suprarenal Portal System.—This system is by no means so

clearly marked off from the general systemic circulation as it

is in many Lizards, In the case of the left suprarenal body,

T noticed two afferent veins arising from the parietes. One of

these arises anteriorly from the parietes near to the middle line,

and may be looked upon as a portion of the posterior cardinal ;

the second vein arises laterally and runs at right angles to the

longitudinal axis of the body. These vessels seem to join a

considerable plexus lying between the gonad and the vas deferens,

which plexus is also continuous with the system of the vena cava

through the spermatic veins.

The azygos vein is present only on the right side, and is not

very extensive.

The anterior abdominal vein gives off anteriorly on each side

a lateral abdominal before it unites with its fellow in the middle

line.

TARENTOLA ANNULARIS.

The vascular system of this Gecko, so far as I have been able

to.examine into its details, does not show wide differences from

that of Phelswuma. There are, nevertheless, a few facts to which

* These arteries in many Geckos are described by Hochstetter in Morph. Jahrb.

vol. xxvi. p. 213. > ;

1904. ] ANATOMY OF THE LACERTILIA. 21

I think it desirable to call attention as tending to emphasise the

peculiarities of the Geckonide as compared with other families of

Lizards. As to the apparent differences between Tarentola and

Phelsuma, I am unable at present to lay much stress upon

them. The arterial system shows one important agreement with

that of Phelswma. There are, in fact, three small gastric arteries

supplying the stomach exclusively which in’ both genera have

precisely the same arrangement; that is to say, there is one

anterior artery, followed after a considerable interval by the two

next which are close together.

But Tarentola has not a gastrosplenic artery arising after these

and before the origin of the cceliac, such as that which is present

in Phelsuma.

I did not observe in Phelswma—but I do not assert that it does

not exist—a lateral artery on either side present in Zarentola,

This artery is in effect a series of anastomoses between the ex-

tremities of the intercostal arteries which arise from the aorta

and run along the ribs towards the ventral middle line. Whether

it is to be compared to the epigastric artery of either side or not,

I do not know.

The hepatic portal system and the single median epigastric vein

are almost exactly like the corresponding veins of Phelswma.

T found, however, only a single parieto-hepatic vessel. The

epigastric is connected anteriorly with the vena cava, as in

Phelsuma.

Suprarenal Portals.—Kach suprarenal body has, as in Phelswma,

two afferent veins, The anterior vein of the left suprarenal body

passes backwards to it, arising from the side of the vertebral

column. This is exactly what I observed in Phelsuma.

The azygos vein is as in the last genus.

The lateral abdominals appear to be rather shorter than in

Pheisuma, plunging at once into the thickness of the parietes,

REsuMn.

It may be useful to state in a few words the chief new facts

contained in this communication.

The most noteworthy new features in the vascular system of

the Chameleon as compared with those of other Lizards are;—

(1) The large number of gastric arteries situated in pairs and

supplying right and left sides of the viscus.

(2) The connection of the longitudinal oviducal vein with the

hepatic portal system, of which it is an affluent apparently not

represented in some other Lizards.

(3) The entire restriction of the dorsoventral oviducal veins to

the efferent renal and to its forward prolongation the anterior

vertebral. In /gwana, for instance, these veins open partly into

the afferent renal.

(4) The absence of aggaepicnens for wanely, running lateral

epigastrics.

22 MR. A. D. IMMS ON THE GILL-RAKERS [ May 3,

_ (5) The short course of the lateral abdominal veins and their

fusion anteriorly with the posterior vertebral.

(6) The tendency towards a disappearance of a special supra-

renal portal system.

As to Pygopus, the following points seem to me to be deserving

of special notice :—

(1) The large number of gastric arteries.

(2) The origin of a considerable number of the visceral arteries

from the intercostals, and not directly from the aorta.

(3) The large number of dorsal parieto-hepatic portals, and the

existence of an equal number of portals arising dorsally on the left

side and reaching the liver via the stomach.

(4) The connection between the branches of the well-developed

lateral abdominals and the dorsal parieto-hepatics.

(5) The absence, or at most small development, of lateral

epigastric veins.

(6) The abundant connection (by 8 trunks) of the median

epigastric with the liver and its connection anteriorly with the

vena cava.

(7) The presence, as in Chameleon, of a posterior vertebral

vein continuous with the afferent renal.

(8) The opening of a single oviducal vein into the afferent

renal posteriorly, and of a single vein into the ovarian and thence

into the caval vein anteriorly.

Concerning ZVarentola and Phelswma, there are fewer general

observations to offer. But I may direct special attention to the

following :—

(1) The absence of at least conspicuous lateral epigastrics.

(2) The connection of the median epigastric with the vena cava

anteriorly.

(3) The restriction of the azygos to the right side.

(4) The shortness of the lateral abdominals.

3. Notes on the Gill-rakers of the Spoonbill Sturgeon,

Polyodon spathula*. By A. D. Imus, B.Sc. (Lond.),

Assistant Demonstrator in Zoology in the University

of Birmingham.

[Received April 19, 1904.]

(Plate II. +)

The gill-rakers of fishes are organs which present a considerable

range of variation in form and structure, but which, as yet, have

been very inadequately studied. In their most familiar form they

* Communicated by Prof. T. W. Barnes, F. RS., F. is

{ For explanation of the Plate, see p. 34.

ID oe Se Oo MIP I Il.

Mintern Bros ump.

A.D. Imms del. M.P Parker. hth.

GIULIS RVUNSEIRS Ol IP OINAOIDON.

1904. ] OF THE SPOONBILL STURGEON. — 23

occur as a single or double row of protuberances disposed along

the concave inner or pharyngeal margins of the branchial arches.

They may be modified, however, so as to become tooth-like, seti-

form, or even filamentous. In some fishes they are completely

absent. Among the Elasmobranchs these structures are found in

a number of species, and, as a typical example, they are well seen

in Acanthias vulgaris. In this fish they occur as lanceolate

projections which are developed principally along the anterior

edges of the pharyngeal margins of the arches. In the “ Basking

Shark,” Cetorhinus (Selache) maximus, they are found as a series

of greatly elongated, coarse, seta-like structures which are disposed

along the inner margins of the branchial arches upon both the

anterior and posterior edges. A somewhat similar apparatus is

described as being present in the South African “Whale Shark,”

Khinodon tymeus. Among the Holocephali the gill-rakers are

small in size and are not conspicuous ; they are seen in Chimera as

rows of small tubercles. By far the greater number of the

Teleostomes possess gill-rakers, and it is in this subclass that they

exhibit their widest range of variation. A contribution towards

a systematic study of them in the freshwater members of the

Teleostomi has been made quite recently by Zander*. He has

examined a considerable number of species, and finds that in

carnivorous types like Hso« and Lucioperca the gill-rakers are in

the form of teeth. In numerous other genera they form a sieve-

like filtering-apparatus (‘‘ Siebfortsiitze”), which is developed in

some cases upon both edges of the branchial arches, as in Perca,

Acerina, Lota, and the Cyprinide; or in others only upon the

anterior edges, as in Clupea alosa, Coregonus fera, C’. albula, and

Osmerus. ‘The relative fineness of the ‘‘Siebfortsitze” is correlated

with the nature of the food of the species where they occur, and

attains its extreme development in this respect amongst those

fishes which subsist upon plankton. Poptat also has studied the

gill-rakers in the Teleostomi, and has given brief descriptions of

their structure and disposition in numerous species. As the

result of his studies, he finds that they are specifically peculiar or

diagnostic in all the forms which he examined, and interprets

their arrangement and development in relation to the form of the

mouth and the nature of the food. All three genera of the

Dipnoi possess gill-rakers. They are largest and stoutest in

Neoceradotus. In Protopterus and Lepidosiren they are present

in the condition of minute, slender, pointed projections, and in

the specimens which [ examined they are slightly more delicate

in the first mentioned of the last two genera.

The object of the present article is to call attention to some

interesting features with regard to the structure and function of

gill-rakers in the case of the “Spoonbill” or ‘“ Paddle Fish,”

Polyodon spathula. Iam indebted to Prof. T. W. Bridge, F.R.S.,

for suggesting to me that I should examine these organs, and

* Zeitsch. fir wiss. Zool. Bd. Ixxv. 1903, pp. 233-258.

+ Ann. Sci. Nat., Zool. t. xii. 1900, pp. 139-216.

24 MR. A. D. IMMS ON THE GILL-RAKERS [May 3,

also for his kindness in placing ample material at my disposal and

for valued assistance rendered to me in various ways.

Anyone who has made even a cursory examination of the gills

of this fish cannot fail to have been struck with the appearance

of the regular comb-like organs which the rows of gill-rakers form

on each face of the branchial arches. Although they are familiar

to most zoologists and are characteristic of Polyodon, no one, so

far as [am aware, has devoted to them more than passing notice.

From among the early accounts of this fish it will serve the present

purpose sufficiently if reference be made to a single source only.

It is in a letter by Dr. 8. P. Hildreth to the editor of the ‘ American

Journal of Science’ that the following mention is made of the gill-

rakers. Heremarks: ‘“‘ The jaws are without teeth; but the fauces

are lined with several tissues of the most beautiful network,

evidently for the purpose of collecting its food from the water, by

straining, or passing it through the ciliary membranes, in the

same manner as practised by the spermaceti whale.”* Another

observer, I, W, Clemens, says that Polyodon “had five pairs of

gills which were double. Each of these duplicatures were thickly

set with teeth, of about the diameter and consistence of best

Russian bristles, and one and a fourth inches long.”+ He

mentions that the particular fish he examined measured 4 ft. 8 in.

in length.

Very little further information is to be gleaned from the works

of any of the later writers. Reference is made to their occurrence

in this fish by Owen £; and Giinther§$ remarks that each branchial

arch has a double series of very long setiform gill-rakers, and that

the two series are separated by a broad membrane. No adequate

figures of them appear to have been published by any author.

A representation of a branchial arch of Polyodon, which also

shows the very characteristic disposition of the gill-rakers, is given

by Prof, Wiedersheim in the 2nd edition of his ‘ Vergleichenden

Anatomie der Wirbelthiere,’ but has been omitted in the later

edition,

Before describing the gill-rakers of Polyodon, it will be necessary

to refer to certain peculiar features in connection with the

branchial arches. Each arch has undergone a remarkable antero-

posterior compression, so that all its segments, and more especially

the epibranchial and ceratobranchial pieces, assume the form of

relatively thin cartilaginous plates, The plates are so obliquely

disposed that the proper anterior and posterior surfaces look

outward and inward respectively, while the concave inner and the

convex outer margins are nearly anterior and posterior respec-

tively. As usual in other fishes, the gill-rakers of the first four

branchial arches form two rows in relation with each arch; and,

* “Notice of the Spoonbill Sturgeon or Paddle Fish of the Ohio (Polyodon feuille

of Lacépéde),”’ Silliman’s Amer. Journ. of Science, vol. xii. (1827) p. 203,

+ Ibid. p. 204.

* ‘Comparative Anatomy,’ vol. i. p. 482.

§ Brit, Mus. Cat. of Fishes, yol. vili. (1870) p. 346.

1904.] OF THE SPOONBILL STURGEON. 25

again following the general rule, they are attached to the anterior

and posterior surfaces, close to the concave Inner margin, in the

form of an anterior and a posterior series, but from the obliquity

of the surfaces of the arch the two series appear as if disposed

along the outer and inner faces of an arch. They will be referred

to in future, however, as the anterior or the posterior series, as

the case may be.

If the gill-rakers are surveyed from the first branchial arch

backwards to the fourth arch, they are seen to become progres-

sively shorter in length, and, furthermore, those of them that

are situated along the outer or anterior aspect of an arch

are somewhat longer than those carried along the inner or

posterior aspect of the same arch (PI. II. fig. 2, 2.9.7. and 0.9.r.).

Hence the series of the longest gill-rakers is carried on the outer

aspect of the first branchial arch, while the row borne on the inner

aspect of the fourth branchial arch is composed of the shortest

gill-rakers. It is also worthy of note that the longest gill-rakers

in either an anterior or a posterior series of a branchial arch are

those situated nearest to the junction of a ceratobranchial with

an epibranchial element (fig. 1, 7.), the gill-rakers gradually

increasing in length from the dorsal and ventral extremities of

an arch until the centre of the concavity is reached, where they

attain their maximum. The fifth branchial arch carries gill-rakers

along its anterior surface only, and they are slightly longer than

those disposed along the posterior aspect of the preceding arch.

The fifth gill-arch is itself much reduced, since it retains only its

ceratobranchial element, and, in correspondence with this, gill-

rakers are not developed on the opposing face of the preceding

arch in relation to its epibranchial cartilage, but only with the

ceratobranchial *.

Owing to the extraordinary compression of the plate-like

branchial arches +, combined with the attachment of the gill-rakers

to their concave inner margins, the greater portion of each arch

practically forms a stout cartilaginous septum, which separates

the anterior from the posterior series of gill-rakers in relation

with each arch, much in the same way that an inter-branchial

septum would separate the double series of gill-filaments on the

opposite or outer margin of a branchial arch (figs. 1 & 2). The

necessity for this curious modification is by no means obvious.

It would seem that, were the septum absent, such delicate

and fragile organs as the gill-rakers would be very lable

to get dislocated or clogged together, and perhaps damaged,

through one series of gill-rakers rubbing against the other.

As it is, each row is kept in a beautifully regular order, and

not a single gill-raker will be noticed to be disarranged from

its proper position, and all of them when not in use are closely

applied to the surface of the septum. The function of this septum

* A full account of the skeleton of the visceral arches of Polyodon is given by

Prof. T. W. Bridge in the Phil. Trans. 1878, vol. 169; vide pp. 702-712 & pl.57. figs.8 &9.

+ Lacépéde (1798) speaks of the branchial arches as cartilage-plates.

26 MR. A. D, IMMS ON THE GILL-RAKERS [May 3,

appears to be to prevent the gill-rakers from becoming damaged

in the way suggested, and to enable them to be stowed away in a

regular order, and within a small compass, so as to admit of the

closing of the operculum in the acts of respiration.

Tn an example of Polyodon the length of which measured 166 cm.

(5 ft. 4in.) from the tip of the rostrum to the extremity of the tail,

the longest gill-rakers measured 45 mm. (1? in.) in their greatest

length; their average length is about 28°38 mm. (1g in.). It will

be seen upon referring to fig. 3 (PI. IT.), that each gill-raker consists

of two parts, viz. a slender shaft, which tapers gradually towards

its free extremity, and a basal portion, which is embedded under

the mucous membrane covering the branchial arch. The basal

portion in the specimen figured measures 8 mm. long and 1°5 mm.

in its greatest breadth; the shaft near to where it joins the base

measures *75 mm. across. Each gill-raker is flattened from side

to side at the base, while the shaft is nearly square in transverse

section. In their natural position, the gill-rakers are disposed

with their flat surfaces at right angles to the septum and

are packed very closely together, the interval between any two

scarcely measuring 25mm. In colour they are pale yellow-brown,

and many of them are somewhat darker at their extreme points.

Their surface is smooth and shining; they are extremely brittle,

and when viewed with transmitted light they have a translucent

appearance. When dried they are bone-coloured and perfectly

opaque.

The method of attachment of the gill-rakers to the branchial arch

is shown in PI. IT. fig. 5, where the lower part of one is represented.

It will be seen that the basal portion is inserted just under the

mucous membrane (m.) covering the branchial arch, and that it

lies parallel with the cartilaginous septum. It is attached to a

branchial arch by numerous elastic fibres (e,f.), which are firmly

inserted into the gill-raker and form a remarkably tough, strong

ligament. Some additional fibres (e./f.') closely ensheath the base

at its lower extremity, and others serve to unite adjacent gill-

rakers to one another. Inserted on the outside of each are some

muscle-fibres (m.f.) which pass downwards and inwards to be

attached to the cartilage of the branchial arch.

A sample consisting of 869 grm. of the shaft portions of gill-

rakers, after having been washed in distilled water and then

thoroughly dried, yielded upon analysis the following chemical

composition * :—

Organic matter ...... 32°81) °/

Mineral matter ...... 67°189 °/,.

The mineral matter consists of calcium phosphate 26°80 °/,,

together with carbonates, fluorides, chlorides, and sulphates of

calcium, with a little magnesium and iron.

On account of the small amount of calcium phosphate they

* I am indebted to Mr. C. J. Thompson, of the Chemical Laboratory of the

Birmingham University, for undertaking this analysis. :

1904.] OF THE SPOONBILL STURGEON, 27

contain, the gill-rakers differ very greatly in their chemical

composition either from bone or dentine.

I found that the gill-rakers, after being decalcified and then boiled

for a few minutes in a strong solution of potassium hydroxide,

retained their general form, but the only sign of structure visible

in them was a coarse fibrous groundwork. After a prolonged

treatment with potash this disappeared, and all that remained

was a small amount of a gelatinous precipitate.

If the shaft of a gill-raker be thinned by rubbing down on the

surface of a fine hone, and then mounted in Canada balsam, a

good deal of its minute structure can be made out. When

viewed under a magnifying-power of 40 diameters, it is seen to

consist of a transparent, faintly yellowish ground-substance,

scattered through which are great numbers of lacune with

canaliculi radiating from them. Running through the matrix or

ground-substance in a longitudinal direction are yellowish-brown

canals which contain blood-capillaries (P1. II. fig. 6). In thin sec-

tions the ground-substance appears quite colourless, and when seen

under a fairly high magnification indications of lamination are

visible in it at the periphery, but they die out towards the centre.

The lacune are for the most part slightly larger than those which

are seen in a section of a human long-bone, and they bear no

definite relation to the blood-channels, so that there are no

indications of Haversian systems. The canaliculi are not

nearly so numerous as those found in typical bone, but they are

more frequently branched, and, moreover, they principally arise

from the two opposite ends of a lacuna, and those belonging to one

lacuna freely anastomose with those of several of the neighbouring

ones (fig. 7). A very marked tendency is exhibited by the lacune

to be disposed with their long axes parallel to the surface of the

shaft with their canaliculi running in a similar direction. The

channels containing the blood-capillaries, which are most numerous

towards the base of the shaft (fig. 6), take the form of longitudinal

canals anastomosing with one another by means of short lateral

connections which are given off at frequent intervals. Traced

higher up the shaft, the number of these canals becomes consider-

ably less, and the anastomoses with neighbouring canals become

fewer. As the extremity is reached they become reduced to two

or three trunks, which eventually join with one another just under

the extreme point (fig. 8). The exact method of the termination

of the channels, however, is not easily to be seen in spirit material,

owing to the small amount of blood that is present in the capillary

vessels which are contained within them. In the basal portion of

the gill-raker vascular channels are entirely absent, but running

down the centre is a long, narrow, apparently empty cavity. The

matrix is homogeneous, and exhibits in places faint traces of

lamination. It is well supplied with lacune, and many of them

differ from those found in the matrix of the shaft in having a

knotted or less regular outline, and in being more profusely

branched.

28 MR. A, D. IMMS ON THE GILL-RAKERS [| May 3,

In a gill-raker which has been decalcified in a 10 per cent.

solution of acetic acid, and afterwards stained with Kleinenberg’s

hematoxylin and cut into longitudinal sections with the microtome,

some additional features may be observed (Pl. II. fig. 4). The

ground-substance exhibits indications of being stratified and its

layers show varied capabilities for absorbing staining-reagents.

Traversing it are numerous canals, which are lined internally bya

definite membrane and contain one or more blood-vessels and some

loose connective tissue. In the region of the shaft the lacunz have

protoplasmic contents which are r eadily stained, together with one

or more deeply staining bodies which are apparently nuclei. Thése

lacunee which are situated nearest to the blood-channels can be

distinctly seen to be in communication with them by means of

their canaliculi. In the basal part of the gill-raker many of the

lacunee are shrunken in their outlines and are very poor in

stainable contents. The lower part of the central cavity, which

is represented at c.c., appears to be nearly empty, containing

only some nucleated tissue in which no definite cells are to be

distinguished. The elastic fibres of the ligament, already men-

tioned, penetrate deeply into the substance of the base in much

the same manner as the perforating fibres of Sharpey, which are

composed partly of bundles of elastic fibres, pierce the circum-

ferential lamelle in bone. Im fig. 4 (ef), where these fibres are

seen in section, they appear as variously shaped dots according to

the angle through which they have been cut, and they are very

evenly distributed through the peripheral ground-substance. The

principal blood-vessel is seen to enter the gill-raker about the

point of junction of the shaft with the basal part. The vessel

then breaks up into several branches, which penetrate the ground-

substance and reach the canals traversing it. The mucous mem-

brane of the branchial arch (m. in figs. 4 & 5), accompanied by

capillaries, is prolonged upwards as a complete and continuous

investment to the outer surface of the gill-raker. Ina decalcified

gill-raker the presence of a membrane covering it is easy to make

out without cutting sections, as it can be stripped off by using

a fine needle under a dissecting microscope.

The only fish which possesses gill-rakers at all comparable with

those of Polyodon is Cetorhinus maximus. In this species the

gill-rakers are of the same general form and, in proportion to the

much greater size of the animal, they are correspondingly larger

andstouter. The investigations of Hannover * and Turner f have

shown, on histological grounds, that there is good reason for

believing them to be very greatly elongated teeth. In common

with those of Polyodon, they consist at their bases of a matrix

permeated with anastomosing canals containing blood-vessels; in

the shaft they differ in that they contain but a single canal which

runs straight up to the tip. There are no lacune, but the matrix

* Kong. Danske Vidensk.-Selskabs Skrifter, 1868, p. 485. A résumé is given in

French in the Ann. Sci. Nat., Zool. t. ix. 1868, p. 373: : :

+ Journ. Anat. & Phys. xiv. 1879, pp. 273- 286, pl. xii.

1904.] | _ OF THE SPOONBILL STURGEON. 29

contains large numbers of coarse dentine-tubes which arise from

the walls of the canals and run outwards to the periphery, where

they form a layer of hard dentine. Turner regards these gill-

rakers as being composed of vaso-dentine, but Tomes in referring

to them adds ‘ (? osteo-dentine).” *

It is worthy of note that in fishes of the genus Chetodon and

their allies the maxillary teeth appear to have been modified along

the same lines as those by which the setiform type of gill-raker

has been produced. As their name implies, the teeth of these

fishes are bristle-like; they resemble the hairs of a fine brush in

- being flexible and elastic, and they are composed of a yellowish,

shining, semi-transparent tissue 7.

I would suggest that possibly the gill-rakers of Polyodon are

morphologically the much modified descendants of exoskeletal

structures which have migrated along with the ectoderm on to the

branchial arches. The fact that the mucous membrane covering

the branchial arches is regarded as being endodermal in origin,

offers considerable difficulty to any idea that such structures could

have developed there independently and in sitw, unless they have

arisen in the underlying mesoblast. Klaatsch, however, from a

study of the placoid scales in Mustelus and some other Hlasmo-

branchs, has arrived at the conclusion that their scleroblasts are

ectodermal in origin and are derived from the same layer as that

which gives rise to the enamel. This layer, which is at first homo-

geneous, becomes divided into a portion which has been usually

considered to be of mesodermal origin, while the rest remains in

connection with the ectodermt. Hence he considers that a

“ dermal” exoskeleton is not mesodermal in its ultimate origin. It

is worthy of note that, with regard to the pharyngeal teeth of

many fishes, several writers are inclined to believe that their

presence is due to a migration of the ectoderm into the cavity of

the pharynx. For this reason, and on account of the difficulty of

reconciling them with the presence of anything except ectoderm,

I would suggest the possibility that the skeletal tissue of the gill-

rakers of Polyodon has arisen from portions of the epiblast

forming the outer portions of the gill-clefts, which have migrated

on to the inner or pharyngeal margins of the branchial arches. At

all events, if any migration of epiblast has taken place, the latter

route seems at least as feasible as a backward migration from the

stomodeeum.

In Cetorhinus the gill-rakers retain many structural features in

common with the teeth of the animal, but in Polyodon they

appear to have undergone a more special modification along lines

of their own. The structure of the teeth in the young Polyodon

has been described by Zograff§, but, after a comparison of the

* “Dental Anatomy,’ p. 220.

+ Vide Owen, ‘Odontography,’ pp. 8 & 105, pl. i. fig. 2.

~ Morph. Jahrb. xxi. 1894, pp. 153-240.

§ “Ueber die Zahne der Knorpel-Ganoiden,” Biol. Centralbl., Bd. vil. 1887-88,

p. 181. Ann. Sci. Nat., Zool. 8 ser. t. 1. p. 203, pl. 4. fies. 3, 4, & 6,

30 MR. A. D, IMMS ON THE GILL-RAKERS [ May 3,

gill-rakers with his account, I have not been able to make out any

salient points of resemblance between the two structures. With

the exception of the rhombic plates and ‘“fulcra” of the tail,

the scales are too degenerate to admit of a similar comparison

being extended to them. The plates and “fulcra” of the tail

are, however, tolerably well-developed structures. The former I

have examined after having thinned them, by rubbing down on

the surface of a fine hone, in the same way in which the gill-

rakers were treated. The matrix of a scale is colourless in thin

slices and is pervaded everywhere by lacune which are similar to

those found in a gill-raker, but it does not contain any blood-

channels. The substance of the plate appears to be deposited

around a longitudinal core-like centre in the matrix. Adjacent

plates are united to one another by means of ligamentous connec-

tions the fibres of which penetrate deeply into their matrix. These

fibres are comparable to what Hertwig calls the ‘ Schuppen-

ligament” of the scales of Lepidosteus.

In structure there is, therefore, a considerable likeness

between a rhombic plate and the basal portion of a gill-raker.

The matrix and its lacune are identical in both cases; the

hollowed core or cavity in the base of the gill-raker might be

compared to the core of one of the plates, and to this may be

added the absence of blood-channels in both cases. The fibres

which connect a gill-raker to the branchial arch, and also which

bind adjacent ones together, are comparable with the ligaments

which unite neighbouring scales. For these reasons I think

that it is not improbable that the basal portion of each

gill-raker is the homologue of a ganoid scale—i. ¢., of one of

the rhombic plates which are found along the sides of the upper

lobe of the tail. The shaft or principal part of a gill-raker may

correspond to a greatly elongated spine, or to one of the evanescent

spines which are found in relation with each scale in the develop-

ing Lepidosteus, and which are regarded as representing the

spinous portions of placoid scales. In Lepidosteus, as Nickerson

has pointed out, the basal plate, which is the essential part of

a ganoid scale, has come to be developed independently of

the vestigial spines, instead of being a continuation of the process

by which the latter are produced; and, in comparison with the

basal plate of the Selachian scale, it has increased greatly in size

and importance and has incorporated within itself fibres from the

dermis*. Ina gill-raker, it would seem that we have a basal

plate which is similarly specialised, though not to so great an

extent, but that there has been no corresponding reduction in the

spinous portion, which, on the contrary, has become greatly

elongated. It has no trace, however, of a hard dentine layer, nor

of a coat of enamel or ganoin, unless the covering of mucous

membrane is to be looked upon as the representative of the latter,

* Bull. Mus. Comp. Zool., Harvard, vol. xxiv. 1893, pp. 115-140,

1904. ] . OF THE SPOONBILL STURGEON. 3l

The absence of such hard parts is to be correlated with their not

being subjected to any use which would involve hard wear or

much friction. In short, it is possible that the gill-rakers are to

be looked upon as exoskeletal parts which were derived from an

ancestral Selachian condition, where they exhibited little or no

differentiation either in form or structure. Subsequently they

became modified along lines of their own in order to fulfil

particular functions, some migrating into the mouth to become

teeth, while others passed on to the branchial arches and have

given rise to the gill-rakers.

From what I have described of their structure, the gill-rakers,

at least their shaft portions, appear to be composed of a substance

which bears a close resemblance to osteo-dentine, if not identical

with it. Osteo-dentine was defined by Owen as that type of

dentine in which the matrix is arranged around the vascular

channels in the form of concentric rings, and in which lacuns

similar to those of bone are found*. Tomes regards osteo-dentine

as a substance which is developed by calcification proceeding

through the interior of a pulp, and not by means of the calcification

of a special layer of cells (odontoblasts) as is the case with other

types of dentine. Consequently, inatooth or structure composed

of osteo-dentine there is no single pulp, but pulp and calcified

tissue are quite inextricably mixed up, the vascular channels

containing masses of pulp-structure as well as blood-vessels. In

vaso-dentine there is a distinct pulp-cavity from which radiate

canals which contain minute blood-vessels only. He further calls

attention to the fact that in some teeth neither of the characteristics

defined by Owen occurs, though, if their manner of development be

taken into account, they are unquestionably made of osteo-dentine?.

Apart from any knowledge of their mode of development, the

substance of the gill-rakers of Polyodon bears a closer likeness to

osteo-dentine than to any other structure, for the following reasons.

It resembles that type of dentine in the absence of a common

pulp-cavity, and in the nature of the anastomosing channels

which contain one or more blood-vessels and some loose con-

nective tissue (pulp-remains?), The presence of bone-lacune is

an additional point of resemblance, though Tomes does not look

upon it as being diagnostic of osteo-dentine, since they, or

spaces very similar to them, are present occasionally in other

kinds of dentine.

In Cetorhinus the teeth are relatively greatly reduced in size,

and its food consists principally of minute surface organisms.

The gill-rakers serve as a straining-apparatus which prevents

the food-particles from passing into the branchial sacs with the

outflowing current of water. As mentioned by Prof. Turner ¢,

* Comp. Anat. vol. i. p 362.

+ “On the Structure and Development of Vaso-dentine,” Phil. Trans. 1878, p. 40.

Also Dental Anat., 2nd edition, pp. 88-92.

t Loe. cit. p. 275.

32 - -MR..A, D. IMMS ON THE GILL-RAKERS [May 3,

Cornish has stated, in an account of a supposed Basking Shark,

that in front of each gill a slight comb-like apparatus extended the

whole length of the ray. As the mouth was opened, the comb

automatically fell back to a right angle with the gill-ray, and

effectually barred the egress through the gills of anything except

water taken in through the mouth*. Although this apparatus has

received attention from numerous zoologists, no one, so far as I

am aware, has offered any suggestion as to the means by which

the gill-rakers are brought to interlock with one another when

they are in use.

The occurrence of minute teeth in Polyodon is a well-known

feature. According to Johannes Miiller, there are found im

young specimens (a foot long) two rows of small teeth in the

upper jaw and one row in the lower jaw. Similar teeth are found

on the two anterior branchial arches where they join the floor of

the mouth, and upon their opposite extremities where they join

the palate. He mentions that examples over 3 feet long are.

edentulous. Ina specimen in the Zoological Museum of the

University of Birmingham which measures 88°4 cm. (2 ft. 10 in.)

long, I find that there are unmistakable teeth arranged on the

jaws, as Miiller states; those in the upper Jaw are worn down a

little more than those in the lower. In another fish measuring

139°1 cm. (4 ft. 53 in.) in length, I have been unable to detect

any trace of teeth.

The nature of the food of Polyodon is correlated with the

vestigial character of the teeth. The fish is described as stirring

up with its spatulate snout the mud at the bottom of the waters

of the “bayous and lowland” streams which it frequents, and

feeding upon the microscopical organisms contained in it ; but the

evidence which supports such a statement appears to be rarely

quoted, and it leads one to believe that it is not so definite as one

would wish. An early writer, already referred to, namely

T. W. Clemens, remarked that the Polyodon which he dissected

“had no food in its intestines—all that was observable was a

small quantity of substance resembling chyle, but of the consistence

of honey.” T. H. Bean §, quoting Prof. 8. A. Forbes, says that

“the long snout is useful in procuring its food, which consists

chiefly of entomostracans, water-worms, aquatic plants, leeches,

beetles, and insect larvee.”

In the hope of being able to furnish some additional obser-

vations, I made a careful microscopical examination of the

contents of the whole course of the alimentary canal in two

specimens of Polyodon. In both cases the food appeared to have

been much acted upon by the digestive secretion and very little

* ‘Zoologist, 1870, p. 2253.

+ © Anatomie der Myxinoiden,’ p. 150.

+ Loe. cit. p. 204.

§ “Cat. of the Fishes of New York,” Bull. 60 of the New York State Museum, 1903,

p. 62.

1904.] OF THE SPOONBILL STURGEON. 33

could be made out with regard to its nature. The specimens

had been in spirit for a long time, which greatly increased

the difficulty of identification. I was able, however, to recognise

among it remains of parts of the exoskeleton of insect larvee

together with portions of the spiral thickenings of the trachez, a

few small Oligochete worms, fragments of plant-remains, and

some earthy matter.

There is every veason to believe that the gill-vakers of Polyodon

ave similay in function to those of Cetorhinus. On account

of their fineness, and the closeness with which they are packed

together, they would form an even more effective straining-

apparatus than they do im the case of the latter fish.

Tn the absence of direct observations on the living fish, as to the

precise method by which the eill-rakers constitute an efticient

filtering-mechanism, recourse must be had to anatomical evidence.

If the gillvakers act as a filter, it is clear that the anterior series

of vakers of one branchial arch must be inclined forwards so as

to meet the posterior series of the preceding arch, which have

become inclined backwards for the purpose, so that the two series

of gill-vakers interlock or interdigitate across the cleft. Under

ordinary circumstances, however, the gill-rakers are closely

applied to the anterior or posterior surfaces of the flattened

branchial arches to which they belong, and they do not in

the least incline across a cleft, or tend to meet those of an

adjacent arch on the opposite side of the cleft. In all probability

the necessary movements of the gill-vakers are brought about by

means of the contractions of the muscle-fibres, represented in PI. I.

fig. 5, which extend downwards and inwards from the outside of

a gill-raker and are attached to the cartilage of the branchial

areh. By the contraction of these muscle-fibres the gill-rakers

would be pulled outwards so as to form an angle of about 60° with

the septum. When the fibres are relaxed, the gill-rakers, on

account of the pull that is exerted upon them by the stretched

elastic fibres, would spring back of their own accord to close against

the septum, and so take up the position they occupy when not in use.

The anterior row carried by the first gill-arch is composed, as

already mentioned, of the largest mdividual gill-rakers. The

yeason for this appears to be that they have to bridge over the

interval between that arch and the hyoid, since the Jatter is devoid

of gill-rakers.

In concluding these few notes it may be worth while to mention

that the gillvakers of the other surviving Chondrostean Ganoids

differ very greatly from those of Polyodon, although all these

Fishes share the common character of having a greatly reduced

dentition. In Psephurus they are comparatively short and are

moderate in number. In Scaphirhynchus they are small fan-

shaped structures each of which terminates in three or four points.

In Acipenser the gill-rakers are small, flattened, and pointed organs

which differ somewhat in relative size among the various species.

Proc. Zoon. Soc.—1904, Vor. IT. No. IT. 3

D4 ON THE GILL-RAKERS OF THE SPOONBILL STURGEON. [May 3,

Doubtless these variations in form ave correlated with differences

in the nature of the food in the different genera.

The most important features with regard to the gill-rakers of

Polyodon may be summarised as follows :—

1. The gill-vakers are setiform structures, and each consists of a

basal portion, which is attached by means of elastic fibres

to the branchial arch, and a long, pare shaft portion which

forms its prmeipal part.

2. The matrix of a gill-vaker contains numerous Jacunze which

are connected with one another by means of canaliculi.

In the shaft portion it contains, in addition, a series of

anastomosing channels in which le blood-capillaries and

some loose connective tissue.

3, Structurally, a gill-raker bears a very close resemblance to

osteo-dentine.

4. The mucous membrane covering a branchial arch is pr molonged

over each gill-raker in the fom of a complete inv esting

coat.

. The necessary movements of the gill-rakers appear to be

brought about by means of the contraction of some muscle-

fibres which are inserted on the outside of each gill-vaker

and which pass downwards and inwards to be attached to

the branchial arch.

6. The food of Polyodon consists of microscopical organisms

and the gill-rakers serve as a straining-mechanism which

effectually bars the entry of such particles into the gill-

cavities.

EXPLANATION OF PLATE TI

Fig. 1. A portion of the 2nd branchial arch of Polyodon spathula showing the

general arrangement of the gill-rakers. (% nat. size.)

2. A semi- diagrammatic section taken across the 2nd branchial arch (through

the ceré atobranchial cartilage), to show the position of the gill-rakers in rela-

tion to the septum. (# larger than nat. size.)

A single gill-raker. (Nat. size.)

A section taken through the lower part of a gill-raker which has been

decalcified and afterwards stained with Kleinenberg’s hematoxylin. The

mucous membrane of the branchial arch is prolonged over the surface of the

oill-raker. The matrix of the latter shows indications of lamination and

numerous lacunz are seen scattered through it. (xX 30.)

5. Figure showing the attachment of a gill- raker by means of elastic fibres to

the cartilage of the branchial arch. “(Xx 5.)

6. A portion of the broadest part of the shaft of a gill-raker which has been

thinned by rubbing down upon a fine hone, the preparation afterwards

mounted in balsam. The matrix is seen to contain large numbers of

lacunze mostly with their axes pomting in one direction. The vascular

channels are seen to anastomose with one another at frequent intervals.

(xX 40.)

. Five typical lacune from the shaft of a gill-raker; they are seen to be in

communication with one another by means of their canaliculi. From a

preparation made in the same way as the one represented in fig. 6.

(X 370.)

. View of the extremity of the shaft of a gill-raker mounted whole in glycerine.

This figure shows the termination of the vascular channels in the tip of the

same. (X 20.)

ice)

1904. | ON THE CRANIAL OSTEOLOGY OF ELOPIDA AND ALBULID, 3D

REFERENCE LETTERS.

b., basal portion of gill-vaker. ef, elastic fibres.

b.a., branchial artery. e/f/, elastic fibres surrounding lower-

bf., branchial filaments. most extremity of gill-raker.

b., branchial vein. h., cut end of branchial arch taken

b./v.’, blood-vessel supplying gill-raker. through the hypobranchial

c., central cavity traversing basal cartilage.

portion of gill-raker. j.b.s., cut edge of interbranchial septum.

cart., cartilage of branchial arch. i.g.7., posterior (or inner) gill-raker.

¢.c., central cavity of basal portion of | m. Mucous membrane.

cill-raker seen in section. m.f., muscle-tibres.

c.h., channels containing capillary | 0.g.7., anterior (or outer) gill-raker.

blood-vessels and loose connec- p-. loose nucleated tissue lying in

tive tissue (pulp-remains ?) central cavity.

d., cut ends of elastic fibres which | y., gill-rakers.

are inserted deeply into matrix s., septum formed by cartilage of

of gill-raker. branchial arch.

e., cut end of branchial arch taken s-h., shaft portion of gill-raker.

through the epibranchial

cartilage.

4: On the Cranial Osteology of the Fishes of the Families

Elopide and Albulide, with Remarks on the Morphology

of the Skull in the Lower Teleostean Fishes generally.

By W.G. Riewoop, D.Sc., F.L.S., Lecturer on Biology

at St. Mary’s Hospital Medical School, London.

| Received April 27, 1904. |

; i J

(Text-figures 8-18.)

An investigation on the structure of the skull of the lower

Teleostean fishes was begun by me some years ago for the purpose

of determining what might and what might not be regarded as

vimitive features in the Teleostean skull, and with the object

also of ascertaining the degrees of relationship existing between

the various genera investigated, so far at least as the cranial

characters might bear upon the subject. The work, however, was

repeatedly interrupted by pressure of other occupation, and the

present paper deals only with a small proportion of the whole

inquiry. Descriptions of the skulls of the Elopide and Albulidz

are here given, and I hope before long to publish similar deserip-

tions of the skulls of Mormyride, Notopteride, Hyodontide,

Osteoglosside, and Clupeidee. In the second part of the paper I

offer some remarks, more or less disjointed, upon the morphology

of the Teleostean skull, based upon an examination of the species

of fishes detailed in the list given at the commencement of that

section. The discussion of the affinities of the genera under

consideration is best deferred for the present.

For the material investigated I am indebted very largely

to Prof. G. B. Howes, of the Royal College of Science, and

Mr. G. A. Boulenger, of the British Museum, and to them I hereby

tender my sincere thanks. I wish also gratefully to acknowledge

the help that I have from: time to time received in the way of

OE ’

)

a6 DR. W. G. RIDEWOOD ON THE CRANIAL | May 3,

suggestions and advice from Dr. A. Smith Woodward, Mr. G. A.

Boulenger, and Mr. C. Tate Regan at the British Museum

(Natural History).

The mode of disarticulation and preparation of the skull adopted

for the purpose of the present inquiry may be recommended for

general use. It is obvious that the skulls of Teleostean fishes,

prepared as one now finds them in museums without the disarticu-

lation of any of the parts, are unsatisfactory, by reason of the

impossibility of studying minutely any but the most superficial

bones. On the other hand, the maceration of the skulls until all

the bones fall apart is equally open to objection, since, if the bones

ave stored loosely in a box, a great deal of time is wasted in

sorting out the parts for study; while if the bones are wired at a

little distance from one another in the manner introduced by

Owen, the mounted skull is unnecessarily bulky, and there is

always the possibility of error in the process of rearticulation ; and

the expense is so great as to prevent the general adoption of the

method. If, however, the skull be disarticulated into four parts

as described below, the whole of the bones of the skull can be

studied closely, the skull when stored occupies no more room than

if no disarticulation had been made; the several parts can be

yapidly placed in position for studying thei mutual relations,

and there is no possibility of erroneous rearticulation.

From the fresh head, or one preserved in spirit, the nasal *,

lachrymal, cireumorbital, premaxillary, and maxillary bones of

the left side are removed in one piece, and are carefully cleaned in

such a way as to prevent their coming apart. The left palatine

and left hyomandibular are then disarticulated from the cranium,

the mandibular symphysis is severed, the left interhyal bone is

disarticulated from the hyomandibular, and the whole hyoman-

dibular-palatine arch of the left side, together with the left ramus

of the mandible and the opercular bones of the left side, are

removed in one piece, and are prepared without further dissocia-

tion. The whole hyobranchial skeleton (with the exception of the

hyomandibular and symplectic bones) is then removed and _pre-

pared in one piece. The remainder of the skull is prepared im

one piece: it exhibits all the bones of the right side in their

undisturbed relations, and at the same time presents a freely

exposed left view of the cranium.

It has been found convenient to consider the constituent parts

of the skull grouped as follows :—

1) Craniwm.—The term Cranium, as applied here, is a con-

venient appellation for that complex of not readily separable bones

disposed around the brain. It includes the vomer and _ para-

sphenoid, belonging strictly to the buccal series, and in some cases

(e. g. Osteoglossum) the nasal bones.

(2) Temporal and Preopercular Series.—Post-temporal, supra-

temporal, subtemporal, preopercular, interopercular. The post-

* Tn such skulls as those of the Osteoglosside the nasal is not removed with this

series, but is left with the cranium.

1904. | OSTEOLOGY OF THE ELOPIDS AND ALBULIDS. 37

temporal bone is properly regarded as a constituent of the

shoulder-girdle; but since Gill and others have laid some stress

on the manner in which this bone is attached to the back of the

cranium, it is expedient in the present connection to treat it as a

bone of the skull. The explanation of the exclusion of the

preopercular and interopercular bones from the operculax series is

given on p. 68.

(3) Cireumorbital Series.—The lachrymal bone is included in

this series of bones set around the eye, but it is considered advis-

able to avoid the use of this name. The bone differs in no

important respect from the others of the series, and it 1s not easy

to identify if there are several sensory-canal bones present at the

side of the snout. The nasal, although shut out from the orbital

margin, belongs to the same category, and is included under the

pr esent head, unless it be rigidly united with the cranium as above

mentioned. The term ‘‘ preor bital” is employed to designate that

bone which forms the anterior margin of the orbit. The word is

uae not used in the same sense as it is by Allis (Journ. Morph.

v. 1898), who, in the case of Ama, has applied it to the lateral

eumordl (endosteal prefrontal).

(4) Maxillary Series —Maxillary, premaxillary, surmaxillary

bones.

(5) Mandibular Series.—Dentary, articular, angular.

(6) Hyopalatine Series.—Hyomandibular, symplectic, quadrate,

metapterygoid, entopterygoid, ectopterygoid, palatine.

(7) Opercular Series.—Opercular, subopercular, branchiostegal

rays, jugular plate.

(8) Hyobranchial Series.—All the bones of the hyoidean and

branchial arches except the hyomandibular and symplectic bones.

ELOPID2.

ELOPS SAURUS.

The only published figure of the skull of H/ops appears to be

that given by Agassiz i 4 his ‘ Recherches sur les Poissons Fossiles,’

Atlas, v. pl. G. fig. 1. The figure shows the superficial bones

well, and most of ae can be readily identified, in spite of the

fact that no attempt has been made to label them in any way.

The following remarks are based upon the examination of four

skulls.

Cranium (text-fig. 8, A, B, & C, p. 38).—The cranium is mode-

rately long and slender as seen from the side; in a dorsal view the

posterior part is of considerable breadth. The parietals are small,

and meet in a mesial suture. They lie over the supraoccipital,

which extends well forward beneath the posterior parts of the

frontals. The character of the family Hlopide given by Boulenger

(‘ Poissons du Bassin du Congo,’ 1901, p. 46), “os pariétaux sépare ant

le susoccipital des frontaux,” while applying correctly enough to

Megalops, does not apply in the case of Hlops. The remark is

repeated without modification in the later diagnosis of the family

appearing in the Ann. & Mag. Nat. Hist. 1904, xill. p. 164. The

38 DR. W. G. RIDEWOOD ON 'THE CRANIAL | May 3,

lateral parts of the supraoccipital are at thei anterior ends in

contact with the posterior edges of the alisphenoids.

The posterior temporal fossa is very extensive. It is roofed

over by the frontal, squamosal, and epiotic, while its floor is

formed by the postfrontal, alisphenoid, pro-otic, squamosal, supra-

occipital, and exoccipital. Its aperture is bounded above by the

Text-tig. 8.

Cranium of Flops sauwrus.

A, dorsal view: B, back view; C, left side. For explanation of lettering, see p. 81.

eplotic and squamosal, externally by the squamosal, mesially by

the epiotic, and ventrally by the squamosal, exoccipital, and

opisthotic. The epiotic is so largely overlapped by the parietal

and squamosal bones, that only its posterior part is visible in a

1904. | OSTEOLOGY OF THE ELOPIDH AND ALBULID#. 39

dorsal view of the aanium. ‘The lateral temporal groove lies over

the postfrontal bone, and is partially roofed over by a lammar

extension of the squamosal.

The subtemporal fossa, beneath the facet for the reception of

the posterior head of the hyomandibular, is deep and extends

inwards and upwards beneath the floor of the posterior temporal

fossa. Its roof is formed by the squamosal, its floor and sides by

the pro-otic and exoccipital, while its internal cecal end is formed

by the supraoccipital. The opisthotic is small, and sends a process

forward, below the subtemporal fossa, to meet a backwardly

directed process of the pro-otie.

The basisphenoid is of fair size. Its body is embraced by the

alisphenoids and pro-otics, and it has a vertical descending plate

which divides the eye-muscle canal and touches the parasphenoid

by its lower edge. An orbitosphenoid of moderate size is present,

but the greater part of the interorbital septum is membranous.

The parasphenoid, which bears a spearhead-shaped patch of

fine teeth, extends back as far as the occipital articulation, but

does not project beyond. The eye-muscle canal opens posteriorly

by an aperture of moderate size. The vomer has fine teeth,

disposed in two patches, right and left. The ethmoid region is

very largely cartilaginous. The prefrontals are purely ectosteal,

while the mesethmoid is clearly of double origm, the upper part

(supra-ethmoid of some authors) beg a membrane-bone, while

the lower part, of diminutive size, is a cartilage-bone, separating

without much difficulty from the former, but firmly united with

the vomer.

Temporal Series. The supratemporal is a large thin lamina of

bone with a sensory canal running along its anterior edge, which

edge is in contact with the posterior edge of the parietal and

squamosal bones. The hinder border of the supratemporal is

deeply notched, which gives at first sight the impression that the

bone is double. The mesial edge of the supratemporal les over

the supraoccipital crest, while the lateral edge overlaps the upper

part of the opercular bone. The meeting of the two supra-

temporals in the middle line of the head is noteworthy, and is

reminiscent of Ania.

The post-temporal has one limb loosely bound by fibrous tissue

to the top of the epiotic, and a shorter and more slender limb

attached to the back of the opisthotic. ‘The third limb, lying

beneath the supratemporal, is very short, and just fails to reach

the outer edge of the posterior temporal fossa; but projecting

forward from it is an ossified tendon, which terminates in a kind

of brush in the middle of the fossa, as in Albula.

Circumorbital Series (text-fig. 9, p. 40).—The nasal aperture is

surrounded by three bones; the orbital ring is complete, and

consists of seven bones.

Maxillary Series (text-fig. 9, p. 40).—The gape is large, bounded

above by the premaxilla and maxilla, both of which bear densely-

set minute teeth extending along the whole of their lower borders.

There are two surmanxillary bones.

40) DR, W. G. RIDEWOOD ON THE CRANIAL [May 3,

Mandibular Series (text-figs. 9 & 10).—The dentary is long,

with numerous minute teeth. The distinction between the

Text-fig. 9.

Hlops sawrus, right side of skull. For explanation of lettering, see p. 81

Text-fig. 10.

Ain

CLOT

d j { !

/ oa I "

pop wp wd

EHlops saurus, hyopalatine arch, opercular bones, &c., with mandible; left side,

mesial aspect. 7, jugular plate, dorsal view. For explanation of other lettering,

see p. 81.

ectosteal and endosteal parts of the articular is clearly marked,

and there is in addition a very small sesamoid articular, situated

1904, | OSTEOLCGY OF THE ELOPIDA AND ALBULIDA. 4i

low down, and far in advance of the endosteal articular, 7. e. not

as in Albula. The articular facet for the head of the quadrate is

formed in its upper part by the endosteal articular, and in its

lower part by the angular bone, which is fused with the ectosteal

articular.

Hyopalatine Series (text-fig. 10, p. 40).—The hyomandibular

articulates with the cranium by two heads, the anterior one being

smaller and standing out more distinctly than the other. The

palatine articulates with the ethmoid region by a single head, so

far as can be seen in the dried skull. The symplectic makes with

the lower limb of the hyomandibular an angle of 115 or 120 degrees.

Minute teeth occur on the palatal surfaces of the palatine, ‘ento-

pterygoid, and ectopterygoid bones.

Opercular Series (text-figs. 9 & 10, p. 40).—The shape, size, and

relations of the opercular bones are so clearly shown in the figures

that ne description is necessary. The branchiostegal rays are 35

in number in one specimen, which may be called “specimen A.”

The first one is attached just beneath the anterior end of the

ceratohyal* ; those that follow form a closely-set series along its

ventral border, while the last fourteen are affixed to the outer

face of the epihyal, and gradually become larger and flatter as one

traces the series backwards. The last three extend farther forward

over the outer face of the epihyal than the others, which has the

effect of making the transition to the subopercular and opercular

bones a very gradual one. In no modern form is it more clearly

seen than in Hlops that the opercular and subopercular are the

two terminal elements of the branchiostegal series. In a larger

specimen (B) the numbers are less—20 on the ceratohyal and 12

on the epihyal. In specimen C there are 29 rays on the right

side and 31 onthe left; in D there are 28 on each side: but in

each of these specimens there are probably some rays missing. A

median jugular plate is present, attached by ligament to the back

of the mandibular symphysis (text-fig. 10, js p- 40).

Hyobranchial Series (text-fig. 11, p. 424).—The interhyal is

ossified. The epihyal is relatively large. The upper and lower

hypohyals are approximately equal in size. The glossohyal is a

flat cartilage, horizontally disposed, with a minute endosteal nodule

in its posterior end, and bearing on its upper surface a finely

dentigerous membrane-bone. The urohyal is long, and extends

back as far as the anterior extremity of the fifth ceratobranchial.

A long, finely dentigerous bone overlies the three basibranchials.

The third basibranchial has degenerated in the anterior five-sixths

of its length into a fatty mass, “but the hinder one-sixth is a small,

fairly compact bone partially covered by the long dentigerous

bone just mentioned, the rest of its upper surface being covered

by small, uncoalesced, readily removable dentigerous plates. The

cartilage-plate representing the undifferentiated fourth and fifth

** The eccurrence of branchiostegal rays along the whole length of the ceratohyal,

up to its anterior extremity, is to be regarded as a primitive feature, even more

primitive than the greatness of the number of the rays. Ama is more specialised

than Hlops in this respect.

49 DR. W. G. RIDEWOOD ON THE CRANIAL [May 3,

basibranchials is roughly hexagonal in shape, elongated in an

antero-posterior direction, and continued posteriorly mto a small

rod of cartilage which lies freely between the fifth cerato-

branchials.

The dentigerous plates on the fifth ceratobranchials are readily

removable, which is not usually the case. here is a true ossified

first pharyngobranchial, projecting slightly upwards by the side of

the pavasphenoid, in addition to the small spicular bone, which in

Teleosteans generally has been erroneously taken to represent the

first pharyngobranchial. The spicular bone stands upright from

the antero-superior extremity of the first epibranchial, and serves

Text-fig. 11.

eb t ch

Hlops saurus, hyobranchial skeleton, dorsal view. The epibranchials and

pharyngobranchials of the right side are not shown. For explanation of

lettering, see p. 81.

to attach the branchial skeleton to the pro-otic bone at the point

where the latter meets the opisthotic bone. The fourth pharyngo-

branchial is cartilaginous, and is flanked on its pharyngeal surface

by a dentigerous membrane-bone exactly similar to those which

occur on the second and third pharyngobranchials. Even the first

pharyngobranchial bears a small dentigerous plate im its posterior

part. The transition from these dentigerous plates to the small

toothed plates that accompany the gill-rakers on the epibranchials

is quite gradual. The whole hyobranchial skeleton gives one the

impression that it is in a very simple and unmodified condition.

1904. | OSTEOLOGY OF THE ELOPIDZ® AND ALBULID#. 43

MEGALOPS CYPRINOIDES.

The account of the skull of Megalops given by Shufeldt (U.S.

Fish. Com. Rep. 1883 (1885), pp. 813-816 and figs. 33 and 34) is

very incomplete. His remarks apply to the cranium only, and his

specimen was defective in the parietal and prefrontal regions.

Hay (Zool. Bull. ii. 1, 1898, p. 28) gives a side view of the hinder

part of the cranium of ‘“ Zarpon atlanticus.’ The tollowimg

remarks are based upon the examination of three specimens of

Megalops cyprinotdes.

Text-fig. 12.

is) j

Cranium of Megalops cyprinoides.

A, dorsal view; B, back view; C, left side. For explanation of lettering, see p. 81.

Cranium (text-fig. 12, A, B, & C)—-The pro-otic and associated

bones ave drawn out ina vertical direction, so that the parasphenoid

takes a sharp bend beneath the pro-otics, instead of running

d4 DR. W. G. RIDEWOOD ON THE CRANIAL [May 3,

nearly horizontally as it does in Hlops. The parasphenoid does

not extend as far back as the occipital articulation, but underlies

the anterior two-thirds of the length of the basioccipital. The

eye-muscle canal in two of the specimens examined opens

posteriorly by an aperture which admits of the passage of a fairly

large sewing-needle, but the canal is blind in the third specimen.

The par asphenoid has a long narrow band of numerous small teeth,

and the vomer bears a heart- shaped patch of similar teeth, com-

pletely or incompletely divided into a vight and left half.

The endosteal mesethmoid is in rigid union with the vomer, and

separates fairly readily from the ectosteal mesethmoid ; this gives

the appearance of the vomer being in part a cartilage-bone. A

lateral process of the ectosteal mesethmoid passes downward and

outward to meet a forward process of the ventro-lateral border of

the prefrontal below the nasal sac. This is not present in Hlops.

The parietal bones touch one another in the median line of the

head; each is nearly square in shape. The supraoccipital does

not spread far beneath the parietal bones, and does not come near

the frontals. In the extinct species it appears to have extended

ae forward (see Smith Woodward, Brit. Mus. Cat. Foss. Fish.

iv. pl. ii. fig. 5; also page vi of the Introduction, in which the

roles to Megalops was probably intended by the author to

refer to the extinct species only).

The posterior temporal fosse are large and extend as fax forward

as the orbitosphenoid bone. They communicate with one another

above the roof of the brain-case, although in Hlops they are a

considerable distance apart. The roof of the brain-case is formed

by the alisphenoids (which meet in the middle line above the

brain *) and by a forward growth of the lower part of the supra-

occipital. The supraoccipital either actually touches the alisphenoids,

or a narrow tract of cartilage intervenes. The roof of the poste-

rior temporal fossa is formed by the parietal, squamosal, epiotic,

and frontal bones. The postfrontal is rather hollow, and forms

part of the external wall and floor of the fossa, and the pro-otic

also forms part of the floor. The lateral temporal groove, above

the articular facet for the head of the hyomandibular, is broad

and shallow, and is not roofed over.

The subtemporal fossa is deep, and extends inwards and upwards.

It is bounded above by the squamosal, below by the exoccipital

and pro-otic, behind by both squamosal and exoccipital bones, and

in front by the pro-otic. The opisthotic is comparatively large.

Its postero-superior part, to which the deep limb of the post-

temporal is attached, is small and wedged in between the main

part of the exoccipital and the part of this bone that forms the

posterior border of the subtemporal fossa. The lower part of the

opisthotic, however, extends forwards so as to form an important

constituent of the side of the cranium. It is somewhat bullate in

shape and touches the pro-otic and basioccipital. This bullate

* Hay (Zool. Bull. ii. 1, 1898, p. 32) states that in Tarpon atlanticus the ali-

sphenoids meet in the mid-line below the brain. This is not the case in the specimens

now under consideration.

1904. | OSTEOLOGY OF THE ELOPIDE AND ALBULIDS. 45

portion of the opisthotic is wanting in Hlops, in which genus the

ventro-posterior border of the subtemporal fossa is formed by the

opisthotic, and not by the exoccipital.

On opening the opisthotic bulla of J/egalops there is presented

a fairly large cavity bounded above by the pro-otic and exoccipital,

posteriorly by the exoccipital and basioccipital, anteriorly by the

pro-otic, internally by the pro-otic, exoccipital, and basioccipital,

ventrally and externally by the opisthotic. The chamber opens

postero-ventrally at the side of the basioccipital, and probably

contains a diverticulum of the swim-bladder ; but on this poimt I

am unable to offer any definite statement.

In the recent Megalops, as in fact im the great majority, if not

the whole, of the Malacopterygian fishes, the right and left pro-

otic bones unite above the eye-muscle canal, and thus separate

the basisphenoid from the basioccipital. I should be disposed,

therefore, to regard as pro-otic that bone which in Wegalops priscus

touches the fr ont of the basioceipital and is marked ‘ basisphenoid ”

by Smith Woodward (Brit. Mus. Cat. Foss, Fish. iv. pl. iii. fig. 5;

also p. 26, specimens P. 556 and P. 1698).

The basisphenoid is rather slender, and is 'T-shaped when seen

from the front. The body of it lies between the two pro-oties and

does not touch the alisphenoids. The orbitosphenoid is similar

to that of Hlops.

Temporal Series—Yhe supratemporal is, as in Hlops, a large

thin seale with a notched posterior border, projecting from the

back of the cranial roof. That section of the sensory canal which

normally runs antero-posteriorly in the post- temporal and supra-

temporal fails in Alegalops to leave any impression on either of

these bones, and may possibly be absent. ‘The two supratemporals

meet on the-dorsum of the head. The post-temporal has an

epiotic limb and an opisthotic Limb, the latter being flattened,

and not rod-like; the third hmb is practically obsolete.

Circumorbital Series (text-fig. 13, p. 46).—This series consists

of a nasal bone and nine bones around the eye; the orbital ring

is Incomplete above. The anterior margin of the preorbital bone

enters into a more or less definite articulation with the back of

that head of the maxilla which engages with the palatine. The

COLE uae articulation in Hops is of a less definite character.

Mauillary Series (text-fig. 13, p. 46).—The gape is large, but not

so large as in Hlops; it is bounded above by the premaxilla and

maxilla. The teeth are minute and densely set on the edges of

both of these bones, and extend nearly to the posterior end of

the maxilla. There are two surmaxillary bones of large size.

Mandibular Series (text-figs. 13 & 14, p. 46). —_Except that it is

much higher in proportion to its length, the mandible of Wegalops

rf esembles that of Hlops.

Hyopalatine Series (text-fig. 14, p. 46).—The hyomandibular

articulates with the cranium by a single moderately broad head.

The palatine has a single head for articulation with the ethmoid

region. The symplectic is longer and more slender than in Elops,

but the lower part of the hyomandibular is broader, and its

AG DR. W. G. RIDEWOOD ON THE CRANIAL [May 3,

opercular head is shorter and less mcised below. The symplectic

does not make a definite angle with the hyomandibular, but lies

Text-fig. 13.

Megalops eyprinoides, right side of skull. For explanation of lettering, see p. 81.

Text-fig. 14.

/ 1 Mi

, Char ee

LO 42) SY g COO”

Megalops cyprinoides, lyopalatine arch, opercular bones, and mandible of. left

side, mesial aspect. 7, jugular plate, dorsal view. For explanation of other

lettering, see p. 81. :

in a continuation of the curved main axis of that bone. The main

axis of the hyomandibular is directed downward and forward,

1904, | OSTHOLOGY OF THE ELOPIDA AND ALBULIDA. AT

whereas in Hlops it slopes downward and backward. Minute teeth

occur on the palatal surfaces of the palatine, entopterygoid, and

ectoptery ygoid bones. Hay (Zool. Bull. ii. 1, 1898, p. 39) states

‘that in Ten ‘pon atlanticus teeth may occur even on the quadirate.

Opercular Series (text-figs. 15 & 14, p. 46).-—The subopercular is

relatively longer and narrower than that of Hops. The branchi-

ostegal rays are from 23 to 25 in number. The first thirteen are

attached to the lower edge of the ceratohyal, the remainder to the

outer face of the epihyal. As in Hlops, the series extends along

the whole length of the edge of the ceratohyal. The last six branchi-

ostegal rays become gradually broader, and the last of all extends

farther forward over the outer face of the epihyal than those that

precede it. There is a median jugular plate (text-fig. 14, 7, p. 46)

attached by ligament to the back of the mandibular “symphysis.

Hyobranchial Series—The hyobranchial skeleton bears a close

resemblance to that of Hlops. The interhyal is ossified. The epi-

hyal is proportionately smaller and the two hypohyals relatively

larger than in Hlops, and the endosteal glossohyal is relatively

ereater. The dentigerous patch on the fifth ceratobranchial is

readily removable, as in Hops. There is a first pharyngobranchial

as well as the spicular bone. It stands more upright than that of

Elops, and is attached to the antero-ventral part of the pro-otie.

ALBULID!.

ALBULA CONORHYNCHUS,

Shufeldt (U.S. Fish. Com. Rep. 1883 (1885), pp. 808-813 and

figs. 28-31) has described the cranium and hyopalatine arch of

Viet but the other bones of the skull were apparently missing

from hie specimen. The following remarks are based upon. the

examination of two skulls.

Cranium (text-fig. 15, A, B,& C, p. 48).—The cranium is rather

long, straight, and thin. The par ietals are small; ; they are 1n contact

in the median line, and lie over the Supraoce ‘pital i mm such a way that

hardly more than the spine of the latter is visible in a dorsal view.

The supraoccipital, however, extends well forward, even beneath

the frontals, and it is therefore incorrect to state, as Boulenger

does (Ann. & Mag. Nat. Hist. (7) xiii. 1904, p. 164), that the

supraoccipital 1 18 separated from the frontals by the parietals. The

posterior temporal fossa is roofed over by the fr ‘ontal, squamosal,

parietal, and epiotic; its floor is formed by the sqtamosall and

pro-otic, its outer wall by the squamosal, and its inner wall by

the pro- otie, supraoccipital, and epiotic.

The back of the cranium, immediately on the two sides of the

supraoccipital crest, 1s so ihalllogved that the term vacuity might

almost be applied to these depressions. At the bottom of ach

depression, surrounded by the supraoccipital, epiotic, and ex-

occipital, a small piece of the squamosal is visible (text-fig. 15, B,

sq.). ‘The subtemporal fossa, lying just below the posterior part of

the articular surface for the head of the hyomandibular, is bounded

entirely by the squamosal and exoccipital in the smaller specimen

48 DR. W. G. RIDEWOCD ON THE CRANIAL | May 3

examined, but in the larger specimen the pro-otic forms a small

portion of the anterior wall. It is a deep fossa, and leads inward

and upward beneath the floor of the posterior temporal fossa.

The lateral temporal fossa, above the anterior part of the articular

Text-fig. 15.

Cranium of Albula conorhynchus.

A, dorsal view ; B, back view; C, left side. For explanation of the lettering, see p. 81.

surface for the head of the hyomandibular, is bounded above by

the squamosal and frontal, internally by the pro-otic and squamosal,

below by the pro-otic, externally by the postfrontal, and termi-

nally, ¢. e. antero-internally, by the alisphenoid. It 1s rather

1904. ], OSTEOLOGY OF THE ELOPIDZ AND ALBULIDS. 49

remarkable that the pro-otie rises so high as to touch the

frontal.

The front part of the side of the basioccipital and the lower

part of the pro-otic are greatly inflated, much as in Osmerus.

Incision into the bulla shows that it is for the accommodation

of the very large sacculus, with otolith of equivalent size, and not

for any diverticulum of the swim-bladder. The opisthotic is

moderately small, and lies equally upon the exoccipital, epiotic, and

squamosal; it sends no process forward to meet the pro-otic.

The orbital surface of the alisphenoid is in a plane nearly

transverse to the axis of the cranium. The basisphenoid has the

form of a Y when seen from the front, but a thin plate of bone

continues forward from the stem of the Y into the interorbital

septum to meet the orbitosphenoid. The upper limb of the Y is

attached on each side to the alisphenoid and pro-otic. The

orbitosphenoid is largely developed, and, with the assistance of

the basisphenoid, forms a complete osseous interorbital septum.

The parasphenoid extends very nearly to the occipital articu-

lation, and the eye-muscle canal opens posteriorly by a small

aperture. The part of the parasphenoid lying below the orbital

region bears numerous hemispherical teeth, disposed in a coftin-

shaped patch. Just where the front of the parasphenoid meets

the back of the vomer is an oval space, which is roofed in by a

forward continuation of the dorsal part of the parasphenoid. The

vomer bears two or three transverse rows of pointed teeth. The

ethmoid region of the cranium is long, the distance from the front

of the mesethmoid to the back of the prefrontal being greater

than the length of the orbit. The mesethmoid projects consider-

ably in front of the vomer; it has a trough on its upper surface

instead of the more usual ridge, and below the trough has a

foramen of elliptical shape, visible in a side view only.

Temporal and Preopercular Series.—The post-temporal has

three limbs. The largest lies over the epiotic, the second or deep

one is more slender, and is attached by ligament to the back of

the .opisthotic, while the third one runs forward beneath the -

supratemporal and spreads out into osseous tendons, to which are

attached fibres of the trapezius muscle, inserted into the posterior

temporal vacuity. The supratemporal has a rather flat external

surface, which is flush with the external surface of the squamosal,

The transverse commissure of the sensory-canal system, after

leaving the upper end of the supratemporal does not run in the

parietal, but superficially to its posterior edge. The preopercular

is bent at an angle of about 108 degrees, and the upper limb is

slightly longer than the horizontal limb.

Circumorbital Series——There are in all twelve bones of this

series (text-fig. 16, p. 50). The most anterior ones are curious,

basket-like bones, not much wider than the sensory canals which

they carry. The canals in this region are particularly large.

Maxillary Series.—The upper part of the premaxilla is sculptured

into a spongy-looking basket-work similar to that of the naso-

lachrymal bones. Unless the mouth is very widely opened, the

Proc. Zoou. Soc.—1904, Vou. II. No. IV. 4

50 __-DR. W. G. RIDEWOOD ON THE CRANIAL | May 3,

premaxilla alone bounds the gape above; the premaxillary teeth

are crowded, small, and sharply pointed. The maxilla bears no

teeth; the anterior extremity which articulates with the mes-

ethmoid is sharply incurved. The surmaxilla is single.

Mandibular Series (text-fig. 17).—The dentary bears teeth similar

to those of the premaxilla. There isa clear distinction between the

Text-fig. 16.

Albula conorhynchus, right side of skull. For explanation of lettering, see p. 81.

Text fig. 17.

Albula conorhynchus, hyopalatine arch, opercular bones, and mandible, left side,

mesial aspect. For explanation of lettering, see p. 81.

endosteal and ectosteal parts of the articular, and, lying anterior

to the former and on the lingual side of the latter, is a sesamoid

articular of unusually large size. The angular is not distinct

1904,] OSTEOLOGY OF THE ELOPIDA AND ALBULIDS. 51

from the ectosteal articular, and the articular facet for the head

of the quadrate is formed in its upper part by the endosteal

articular, and in its lower part by the combined angular and

ectosteal articular.

Hyopalatine Series (text-fig. 17, p. 50).—The hyomandibular arti-

culates with the cranium by a single broad head, and its axis slopes

downward and forward. Owing to the smallness of the gape the

quadrate-articular joint is far in advance of the hyomandibular-

cranial articulation, and the symplectic runs nearly horizontally

forwards. The palatine cartilage ossifies at its anteriorand posterior

ends, but remains unossified for the greater portion of its length.

The anterior head articulates with the mesethmoid, the posterior

with the prefrontal. The ectosteal palatine is of small extent, and

bears crowded, small, long, sharp teeth. It is united with the

anterior of the endosteal palatines, but is remote from the posterior

one. The hinder two-thirds of the palatine cartilage, and its

posterior ossification, lie in a trough formed by the ectopterygoid

and entopterygoid. The entopterygoid bears an elongated patch of

hemispherical teeth similar to those on the parasphenoid; the

ectopterygoid is toothless, or may bear two or three small teeth in

continuation of the patch of teeth on the palatine. There is a

prominent, externally-directed, horizontal process of the ectoptery-

goid, which serves to form part of the floor of the orbit. Its outer

edge rests against the upper edge of two of the suborbital bones.

Opercular Series (text-figs. 16 & 17, p. 50).—The subopercular is

large. The branchiostegal rays are 15 in number; they all arise

from the outer face of the hyoid arch. The first two lie freely.

in the branchiostegal membrane, the next ten are attached to

the posterior two-thirds of the ceratohyal, and the last three are

attached to the epihyal. They form a well-graduated series, those

in front having the form of slender curved needles, the hinder

ones being larger and lamellate.

Hyobranchial Series (text-fig. 18, p. 52).—The upper and lower

hypohyals are equal in size, although in an external view the

lower appears to be the larger. The urohyal extends back a little

beyond the posterior extremity of the third basibranchial. The

gill-rakers are all short and stumpy. The interhyal is ossified.

There is a conspicuous dentigerous membrane-bone covering

the first, second, and third basibranchials and the hinder part of

the glossohyal. The teeth are hemispherical, and resemble those

of the parasphenoid and entopterygoid, with which, in fact, they

engage. The glossohyal is movable beneath this dentigerous

plate, but the basibranchials are not. The body of the glossohyal

consists of cartilage in front and cartilage-bone behind, and an

edentulous thin membrane-bone covering both. There is a very

small dentigerous bone covering that cartilage which represents

the fourth and fifth basibranchials; the teeth of this are small

and pointed, like those of the gill-rakers.

The first pharyngobranchial is: perfectly conformable with the

second, and is set in a line with the first epibranchial. A true

spicular bone appears to be wanting. The third pharyngo-

52 DR. W. G. RIDEWOOD ON THE CRANIAL | May 3,

branchial is peculiar, the upright limb being exaggerated, and

the forward one reduced to such an extent that not only does it

fail to run along the mesial edge of thesecond pharyngobranchial,

but it does not even touch it.

Text-fig. 18.

Albula conorhynchus, hyobranchial skeleton, dorsal view. The epibranchials

and pharyngobranchials of the right side are not shown. For explanation of

lettering, see p. 81.

BATHYTHRISSA DORSALIS (Pterothrissus gisu).

I am pleased to be able to confirm a suggestion made by

Boulenger (‘‘ Revision of the Mormyride,” Proc. Zool. Soc. 1898,

p- 776) that Bathythrissa is not intimately related with the

Salmonide, with which family Giinther had associated it (Ann.

& Mag. Nat. Hist. (4) xx. p. 448; Chall. Rep. Deep-Sea Fishes,

p. 221), nor with the Clupeide, with which family Gill grouped

it (Mem. Nat. Acad. Sci. Wash. vi. 1893, p. 131), but is nearly

allied to Aldula. Smith Woodward is evidently of the same

opinion as Boulenger, for in placing the genus /stiews in the

family Albulide, he remarks that Bathythrissa is not clearly dis-

tinguished from Jstiews (Brit. Mus. Cat. Foss. Fishes, iv. 1901,

p. 67, also Introduction, p. vil).

The posterior temporal fosse are completely roofed over, as in

Albula, but the lateral temporal fossee are much shallower. The

relations of .the opisthotic, parietal, and supraoccipital ‘are as in

Albula; except that the supracccipital crest is smaller.’ The basi-:

1904.] © ogrEoLoGy OF THE ELOPIDA AND ALBULIDS. 53

occipital and the lower parts of the pro-otics are inflated, just as

in Albula, but the subtemporal fossz are larger and shallower.

The parasphenoid and entopterygoid teeth are more pointed

than those of Albwla; the vomer and palatine are edentulous.

The eye-muscle canal has no posterior opening ; the basisphenoid

is relatively smaller; the interorbital septum, below the orbito-

sphenoid, is membranous; the prefrontal is of slighter con-

struction ; the oval space in the front part of the parasphenoid is

very small; the ethmoid region of the cranium is a little shorter

in proportion ; the top of the mesethmoid has no groove, but the

oval foramen seen in a side view of the mesethmoid is present.

The shapes, proportions, and relations of the supratemporal and

post-temporal are the same as in Albula, although I have not

recognised the ossified tendons of the post-temporal projecting

into the posterior temporal fossa. The sensory canals of the head,

judging by the shapes of the superficial bones, are even relatively

larger than in Albula. The circumorbital bones, the premaxilla,

maxilla, and surmaxilla do not differ materially from those of

Albula. Giinther (J. c.) says “maxillary with a marginal row of

very small teeth,” but this I cannot confirm.

The mandible differs in shape in consequence of the coronoid

process being situated farther forward ; it is in the posterior half

of the ramus in Albula, but in the anterior half in bathythrissa.

The sesamoid articular of Bathythrissa is unossified. The hyo-

palatine arch resembles that of Albula, except in the matter of

teeth, noted above; the process of the ectopterygoid which passes

outward and backward in the floor of the orbit to join one of the

suborbital bones is longer and more slender; the nodule of

cartilage which in Albula is ossified to form the posterior endosteal

palatine is unossified in bathythrissa, and is connected by a strong

ligament with a process of the orbitosphenoid which is directed for-

ward and downward, and lies on the mesial side of the prefrontal.

The subopercular is smaller than in Albula, but is of the same

general shape; the ventral edge of the interopercular is notched

at a little behind the middle of its length. The sensory tube,

which runs in the lower edge of the preopercular and beneath the

ramus of the mandible, is evidently larger in Bathythrissa than in

Albula. There are only six branchiostegal rays, instead of fifteen

as in Albula. The first four arise from the outer surface of the

ceratohyal, the next from the junction of the ceratohyal and the

epihyal, and the last from the outer surface of the epihyal.

The only differences to be noted in the hyobranchial skeleton

are that the glossohyal has its own teeth, confined to the posterior

fourth of its surface, and is not overlapped by the dentigerous

membrane-bone that belongs to the three basibranchial bones.

The basibranchial teeth stand higher, and are less hemispherical

than in Albula.

Comments on the Skull of the Elopide and Albulidee.

In reviewing the characters which are common to the skull of

the Hlopide and the Albulidee, it is perhaps natural that we should

54 DR. W. G. RIDEWOOD ON THE CRANIAL [ May 3,

be restricted in the main to primitive features. There is no doubt

that the Elopide are the most archaic of existing Teleosteans, and

that the Albulide are in few respects more highly specialised ; but

the study of the skull does not show any direct affinity between

the two families. There is no specialised character common to

both. Such resemblances as exist between them are explicable

by the fact that neither has departed to any great extent from

the ancestral group from which all the Teleostean fishes sprang ;

and there is no evidence (from the study of the skull) that the

divergence of the two families from a single one occurred at any

considerable height above the root of the phylogenetic tree.

In the presence of conal valves of the heart other than those at

the junction of the conus with the ventricle (see Boas, Morph.

Jahrb. vi. 1880, p. 528), Albula exhibits a character common in

Ganoid and still lower fishes, but not possessed by any other

living Teleostean ; while in the possession of a median jugular

or gular plate Hlops and Megalops exhibit a resemblance to the

Ganoid Amia which is not shared by any other Teleostean fish.

The Elopide were abundant in Cretaceous times, and some of the

extinct forms would seem to be more specialised than the living

Elops and Megalops, since they exhibit a reduction in the size of

the jugular plate (Zhrissopater and Spaniodon) and a separation

of the two parietal bones (2hacolepis, Thrissopater, Spaniodon,

and others). (See Smith Woodward, Brit. Mus. Cat. Foss. Fish.

iv. Introduction, p. vi.)

There is no instance among Teleosteans of a paired vomer such

as occurs in Ama and its allies, but it is worthy of note, perhaps,

that in Hlops, and to a lesser extent in J/egalops, the vomerine

teeth are disposed in two patches, right and left.

Of the characters common to the skulls of the Elopide and the

Albulidee may be mentioned the small size of the parietal bones

and their meeting in the median line; the roofing of the posterior

temporal fosse ; the presence of subtemporal fosse ; the presence

of opisthotic, basisphenoid, and orbitosphenoid bones, an upper

and lower hypohyal on each side, and an ossified first pharyngo-

branchial; the distinctness of the endosteal articular from the

ectosteal articular, and the fusion of the angular bone with the

latter ; and the presence of teeth on the dentary, premaxillary,

entopterygoid, and parasphenoid bones.

Teeth occur on the vomer and palatine in Hlops, Megalops, and

Albula, but not in Bathythrissa. Minute denticles occur on the

surface of the ectopterygoid in EHlops and Megalops, but there are

none in Bathythrissa and only two or three small teeth at the

front of the ectopterygoid in Aljula. The mouth in the Albulide

is reduced in size, and its upper border is formed by the pre-

maxille alone, the maxille being toothless, whereas in the Hlopidee

the upper border is supported by both premaxille and maxille,

and both bear teeth. There are two surmaxille on each side in the

Elopidee, and one in the Albulide.

The mandibular suspensorium (hyomandibular and quadrate) is -

slightly tilted forwards in £lops; in Albula it is strongly rotated

1904. ] OSTEOLOGY OF THE ELOPIDZ AND ALBULID. 55

in a forward direction; and in Megalops it has an inclination

intermediate between that of Hlops on the one hand and that of

Albula on the other. The laterally-directed process of the ecto-

pterygoid, which in Albula runs below the eyeball to meet the

upper edge of the suborbital bones, is feebly suggested, in both

Hlops and Megalops, by a narrow, laterally projecting ledge of the

ectopterygoid.

The lateral temporal fossa is roofed over in Hlops, Albula, and

Bathythrissa, but in Megalops the lateral temporal groove is broad

and shallow, and has no roof. The interorbital septum is com-

pletely bony in Albula, but it is largely membranous in the other

genera under consideration. The eye-muscle canal has a posterior

opening in Hlops and Albula, but ends blindly in Bathythrissa,

and is blind in one specimen of Megalops, although opening in the

other two. The ethmoidal region is elongated in the Albulide,

but not in the Hlopidz ; and the palatine has two articulations,

one in advance of the other, whereas the articular head is single

in the Elopide. While in the Albulide the ethmoidal rostrum

projects in front of the mouth, in Hops, and more particularly in

Megalops, the lower jaw projects in advance of the upper.

The supratemporals in the Elopide are large, thin scales of bone,

which meet in the dorsal median line of the head, without forming

a suture; in the Albulide the supratemporal is more laterally

placed, is smaller, and has more the triradiate character so common

among Malacopterygian fishes generally. The branchiostegal rays

are 32-35 in Hlops, 23-25 in Megalops, 15 in Albula, and 6 in

Bathythrissa. The spicular bone present in addition to the true

first pharyngobranchial in the Elopide, is wanting in the

Albulidee.

Remarks on the Morphology of the Skull in the Lower

Teleostean Fishes generally.

In this section of the paper are offered some remarks, generali-

sations, and criticisms of published accounts of the skulls of fishes,

based mainly upon an examination of the following species,

Most of the skulls were specially prepared for the purpose of this

research, it being found that the skulls of fishes as ordinarily

prepared do not lend themselves satisfactorily to such a detailed

examination as that to which it was deemed desirable to submit

them in the present inquiry.

Hlops saurus.

Megalops cyprinoides,

Albula conorhynchus.

Bathythrissa dorsalis.

Mormyrops deliciosus.

Petrocephalus bane.

Gymnarchus niloticus.

Notopterus kapirat.

Hyodon alosoides.

Osteoglossum lerchardti.

Arapaima gigas.

Heterotis niloticus.

Phractolemus ansorgit.

Chanos salmoneus.

Chirocentrus dorab.

Chatoéssus erebt.

Clupea finta.

Dussumieria acuta.

Engraulis encrasicholus.

Cotlia nasus.

56 DR. W. 6. RIDEWOOD ON THE CRANIAL. [May 3,

Ectosteal and Endosteal Bones.—With regard to the relation

between ectosteal and endosteal ossifications in the skull of bony

fishes, there is reason for believing that the ectosteal is the more

primitive, and that even in the few cases in which related ecto-

steal and endosteal bones remain distinct, as in the postfrontal

and sphenotic of Amia*, the endosteal and ectosteal parts of the

articular bone in Amia, Lepidosteust, Arapaima, Albula, Elops,

Megalops, Hyodon, and Gymnarchus, the endosteal and ectosteal

parts of the mesethmoid of Megalops, and those of the glossohyal

in a great variety of forms, the endosteal ossification has been set

up im sympathy with the ossification taking place in the dermal

tissues. The process of ossification is infectious, if one may employ

such a term in this connection, and the increase of blood-supply,

and the redistribution and alteration in the character of the cells

and matrix in the one part is shared by the subjacent parts to a

greater or less degree. As other examples of such related ossifi-

cations, there may be mentioned the squamosal and pterotic, the

postfrontal and sphenotic in Teleosteans, the prefrontal and par-

ethmoid and the ectosteal and endosteal parts of the angular in

such forms as Lepidosteus and Arapaima, in which the upper part

of the bone articulating with the quadrate, is endosteal, while the

ventral surface of the bone is sculptured, and has all the appear-

ances of a dermal bone. Having regard, therefore, to the superior

antiquity of the dermal constituents of the combined bones,

whether these have arisen phylogenetically around sensory canals

or by coalescence of integumentary denticles—a matter ably dis-

cussed by Allis (Journ. Morph. xiv. 1898, pp. 426-431)—it is

preferable to adopt the names that belong to such superficial

ossifications, e.g. squamosal{, prefrontal, postfrontal §. And

* An occasional feature only, although Traquair (‘Ganoid Fishes Brit. Carb.

Form.,’ Palzont. Soc. 1877, p. 16) and Allis (Journ. Morph. ii. 3, 1889, p. 479) seem

to regard it as constant.

+ Van Wijhe (Nied. Arch. f. Zool. v. 3, 1882, pp. 268 & 281) regards the coronoid

also of Lepidosteus and Amia as consisting of separable endosteal and ectosteal

elements, which he calls the autocoronale and the dermocoronale or suprangulare.

The autocoronale of Lepidosteus, however, is either a bone which is to be identified

with the sesamoid articular (see p. 72), or is merely a thickened part of the splenial.

In neither Lepidosteus osseus nor L. viridis have I been able to separate it from the

splenial, yet Van Wijhe (J. e. pl. 16. fig. 9, a.c.) figures it as a separate bone. The

autocoronale of Ama, on the other hand, the bone which is marked d by Bridge

(Journ. Anat. & Phys. xi. 4, 1877), is a special nodule of bone developed in Meckel’s

cartilage in relation with the articulation between the symplectic and the mandible.

(The bone a of Bridge is the angular; 6 and c, which I have never seen as two

separate bones, are the endosteal articular; while the “angular” of Bridge is the

ectosteal articular.)

{ In his description of the skull of Grammicolepis, Shuteldt (Journ. Morph. ii. 2,

1889, p. 280 & fig. 2) discriminates between the squamosal and the pterotic. He

says: “ At the distal extremity of the squamosal I detect a small, flake-like piece of

bone, thoroughly attached, though individualised by sutural traces, which I take to

be the representatives of the pterotic.” Since, however, he also letters the squamosal

and pterotic separately in his figure of such a well-known skull as that of Carana

(fig. 6, p. 285), it would appear that no great importance need be attached to the

distinction.

§ Cole & Johnstone (Proc. & Trans. Liverp. Biol. Soc. xvi. 1902, pp. 160-161),

while admitting the propriety of using the term squamosal for the dermal bone lying

1904. ] OSTEOLOGY OF THE ELOPIDZ AND ALBULID&. 57

this will apply even in cases where the combined bone has sunk

inwards and no longer presents itself as a superficial bone of the

skull ; as, for instance, the prefrontals and postfrontals of Arapaima

and Osteoglosswm, in which genera so many of the dermal bones

are still dermal in position.

Even if in these latter cases it be proved by histological investi-

gation that the bone is a pure cartilage-bone, the argument is not

the less sound. ‘The ossification in the cartilage owed its origin

phylogenetically to a predisposing dermal ossification which

now no longer appears in ontogeny. The terms parethmoid,

sphenotic, and pterotic are, therefore, redundant; and if the

occasion arises for discriminating the two parts of the prefrontal,

postfrontal, and squamosal, they should be distinguished by the

terms ectosteal and endosteal. This practice has hitherto been

followed, more or less, in the case of the articular and palatine

bones. Thetwo constituents of the palatine of Ama, for instance,

are called exosteal and endosteal by Bridge (Journ. Anat. and

Phys. xi. 4, 1877, p. 616), while Allis (Journ. Morph. xii. 3, 1897)

alludes to them as the dermopalatine and autopalatine, employing

the prefixes introduced by van Wijhe (Nied. Arch. f. Zool. v. 3,

1882). The question of ectosteal and endosteal ossification in

fishes has already been discussed at some length by Vrolik (Nied.

Arch. f. Zool. i. 3, 1873), Gegenbaur (Morph. Jahrb. iv. Suppl.

1878), Pouchet (Journ. Anat. et Phys. xiv. 1878), van Wijhe

(J. c. 1882, p. 210 e¢ seqg.), and McMurrich (Proc. Can. Inst. 1

3, 1884, p. 280 ct seg.); and Schmid-Monnard (Zeitschr. f. wiss.

Zool. xxxix. 1883) has recorded some valuable observations on the

mode of origin of such bones as the epiotic and squamosal in

Teleostean fishes.

More recently, Swinnerton (Quart. Journ. Micro. Sci. 1902,

p. 531, footnote) has suggested the prefixes dermo- and chondro-,

“the former being used for bones which are quite free of the

cartilage, and the latter for those which involve cartilage, 1rre-

spective of the degree of ossification in this, or of the retention of

dermal characters.” This does not seem, however, to be a very

satisfactory solution of the difficulty.

Oranial Bones.—The meeting of the two parietal bones in the

median line is, upon paleontological grounds, a more primitive

condition than the separation of these bones by the supraoccipital,

no separation of the parietals occurring in pre-Cretaceous Isospon-

dylous fishes (Smith Woodward, Vert. Paleontology, 1898, p. 113).

We have thus, to all appearances, one sound character by which

to test the relative tendency towards specialisation among Teleo-

stean fishes. But the possibility of a secondary approximation of

superticial to the pterotic and inseparable from it, do not treat the terms postfrontal

and sphenotic in the same way, on the ground that the term postfrontal “ cannot be

correctly applied to a membrane-boue in Fishes.” Why it. cannot, they do not

explain. |

08 DR. W. G. RIDEWOOD ON THE CRANIAL [ May 3,

the parietal bones during evolution must be borne in mind ; and if

the Berycoid fishes are to be regarded as the parental stock of the

Acanthopterygians generally, a view which is supported by both

comparative anatomy and paleontology, all instances of contact

between the parietal bones among the Acanthopterygian fishes

(e. g. Cyttidee, Scorpenide, Triglide) are to be looked upon as

secondary. In support of the plausibility of the hypothesis may

be mentioned the parallel instance of union of epiotic bones in the

middle line in such forms as Lophius, Regalecus, and Luvarus,

this union being without doubt of secondary origin.

In Chanos there exists a condition which is calculated to make

one pause before concluding that even within the limits of the

Malacopterygian fishes the meeting of the parietals necessarily

indicates the retention of the primitive condition ; for in this genus

the parietals are widely separated in the young, but by subse-

quently fusing with the scales of the commissural section of the

sensory-canal system, they come to meet above the supraoccipital

bone*. They do not meet in an extensive suture; indeed, they

leave a considerable portion of the supraoccipital exposed both in

front and behind; but the condition is just sufficient to make it

advisable to trace the development of the roofing-bones of the

cranium in those forms in which the meeting parietal bones are

thin, and beneath which bones the supraoccipital extends a con-

siderable distance forward. This last relation, it may be observed,

is the rule rather than the exception. Boulenger has already

alluded to it in the case of the Salmonidze (Proc. Zool. Soc. 1895,

p. 300), and I gather that he regards the union of the two

parietal bones as secondary if the supraoccipital bone can be

shown to extend beneath them so as to touch the frontal bones.

But if this be so, the condition found in those primitive genera

Hlops and Albula must be secondary, for in them the supra-

occipital touches the frontals beneath the united parietal bones ;

a fact evidently overlooked by Boulenger when he drew up his

synopsis of the families of Teleostean fishes (Ann. & Mag. Nat.

Hist. 1904, vol. xiii. p. 164).

In Clupea, Chatoéssus, and Chirocentrus the cranial roof is

deficient towards the front of its upper surface, and a fontanelle

occurs between the frontal bones and the mesethmoid. Perhaps

this tendency for the frontals to remain apart points to some

affinity with the Characinide and Cyprinide. In Citharinus such

a fenestra extends the whole length of the frontal bones and

involves also the parietals, while in Alestes a fenestra is found

between the parietals and the hinder part of the frontals. A

condition similar to the last occurs in such Cyprinoids as Cato-

stomus, Cyprinus, and Cobitis (see Sagemehl, Morph. Jahrb. x.

and xvii.).

The interfrontal suture is obliterated in Gonorhynchus, but it

does not appear that much weight need be attached to this

* Cope (Trans. Amer. Phil. Soc. n. s. xiv. 1871, p. 455) mentions “ parietals

united” as one of the primary distinctive features of the family Lutodiridz.

1904. ] OSTEOLOGY OF THE ELOPIDE AND ALBULID&. 59

particular character. The persistence of the interfrontal suture

in the different genera of the Murenide, for instance, is very

inconstant.

In Osteoglossum the mesethmoid is separated from the frontals

by the meeting of the two nasal bones in the middle line. In

Megalops the endosteal mesethmoid is more firmly united with the

vomer than with the ectosteal mesethmoid; in Arapaima the

endostea! mesethmoid alone is present, and this does not present

itself on the roof of the cranium; while in Chanos and the

Salmonide an-ectosteal mesethmoid is present, and the cartilage

is unossified.

It is the rule among the Malacopterygian fishes fer the two

exoccipital bones to meet above the foramen magnum, and for the

basioccipital to be excluded from the floor of the brain-cavity by

the union of the two exoccipitals below the brain. The two pro-

otics also unite beneath the brain, and form, with or without

the co-operation of the basisphenoid, the roof of the eye-muscle

canal.

The supraoccipital crest is evidently to be regarded as an osseous

sheet developed in relation with the great trunk-muscles, and not

a backward extension of the supraoccipital bone itself. Chanos

is very interesting in this respect, in that it shows a condition

intermediate between the usual vertically disposed sheet of bone

and the separable brush-like tendon-bone or intermuscular bone

that projects back from the supraoccipital proper in Chatoéssus.

In the latter genus similar and separable osseous brushes project

back from the epiotic bones; and in Sphyrena and Mugil, Starks

(Proc. U.S. Nat. Mus. xxii. 1900) has described and figured

similar brushes, but not separable, continuous with the back of

the epiotic bones. Osseous brushes on the back of the exoccipital

bones are of much more common occurrence, and these are dis-

cussed on p. 65.

The extent to which the supraoccipital, epiotic, and squamosal

crests project backwards, and to which the hinder surface of the

cranium is excavated for the better attachment of the trunk-

muscles, may be taken more or less as a measure of the

specialisation of the skull, since in the pre-Cretaceous bony fishes

the back of the cranium is nearly flat. The excavation of the

back of the cranium has doubtless originated independently in

different groups ; and Allis has pointed out (Zool. Bull. ii. 2, 1898,

p. 92) that this must certainly have been the case in Amia and

Scomber, for in the former the trunk-muscles have extended

forward beneath the parietal bones, whereas in the latter they

lie externally to the parietals. The degree of irregularity of the

back of the skull and the dimensions of the posterior temporal

fossee appear to be in direct proportion to the muscularity of the

front portion of the trunk of the fish.

The orbitosphenoid is a bone which is very variable in its form

and occurrence; it 1s wanting in Osteoglossum, Gonorhynchus, and

Chanos. In the majority of the lower Malacopterygian fishes it

60 DR. W. G. RIDEWOOD ON THE CRANIAL [ May 3,

constitutes an ossification in the upper part of the interorbital

Septum; but in the Mormyride and in Albula and Arapaima it

extends the full height from parasphenoid to frontal. In the

latter genus it is a paired bone—a very unusual feature in

Teleostean fishes, and reminiscent of Amia. Asa rule the bone

is single, being formed by the confluence (although possibly not

during ontogeny) of right and left constituents, the union being

greatest ventrally, so that the bone in transverse section has the

appearance of a U, ora Y, ora T,oranI. Shufeldt, it is to be

noted, speaks of orbitosphenoids in the plural in his description of

the skull of Grammicolepis (Journ. Morph. ii. 2, 1889, p. 280),

but it does not necessarily follow that in this statement he intends

to convey the idea that the right and left parts are really separ-

able; Hay, for instance, speaks of -‘ orbitosphenoids ankylosed in

the mid-line as in the salmon ” (Zool. Bull. ii. 1, 1898, p. 32). It

is to be borne in mind, further, in dealing with the orbitosphenoid

that Boulenger still uses the term in the sense in which Owen

employed it, 7. e. as a designation for the bone which is now more

commonly termed the alisphenoid (see Brit. Mus. Cat. Fishes,

ed. 2, 1.1895, p. 113, fig. C, Percichthys; and ‘ Poissons du Bassin

du Congo,’ 1901, p. li, Lates).

The alisphenoid bones are usually separated, but in Votopterus

they meet one another behind the orbitosphenoid. In JJegalops

the alisphenoids unite above the brain.

A basisphenoid is very generally present in the lower Teleostean

fishes, and has the form of a Y or a IT’ when viewed from the

front; but it appears to be wanting in Arapaima, Heterotis,

Osteoglossum, Gonorhynchus, Chanos, Mormyrus, and Mormyrops.

Itis large in Albula, and assists the orbitosphenoid in forming

a complete bony interorbital septum (text-fig. 15, C, p. 48).

The eye-muscle canal (myodome of American writers) opens

posteriorly by a relatively large aperture, bounded right and left

by the posterior laminz of the parasphenoid, in Clupea, Dussu-

nueria, Chirocentrus, and Hngraulis, and in a somewhat similar

manner in Chatoéssus, although the free wings of the parasphenoid

are here wanting. ‘The canal also opens posteriorly in Hyodon,:

Albula, Hlops, and Megalops; but in the Osteoglosside, the

Mormyride, and in Notopieruws and Chanos it terminates blindly.

The canal is open in the Salmonide ; but owing to the large

amount of cartilage present in the cranium, and the consequent.

shrinkage on drying, the appearances presented by the dried skulls

are apt to be misleading.

While some importance may perhaps be attached to the fact

that the eye-muscle canal either opens posteriorly or terminates

blindly—the facts stated in the last sentence are rather against

this conclusion—no value can be ascribed, so far as I can see, to

a feature upon which Cope has laid some stress (Trans. Amer.

Phil. Soc. n.s. xiv. 1871, pp. 454 & 455), namely, the double

or simple nature of the basis cranii. This refers, so far as I

understand his writings, to the separation of the parasphenoid:

(1904. | OSTEOLOGY OF THE ELOPIDE AND ALBULID®. 61

from the pro-otic floor of the cranium by the eye-muscle vacuity.

The character is one which is very difficult of application ; and it

is a matter of individual opinion whether such a form as Clupea

is to be regarded as having a simple or double basis cranii, for

here the parasphenoid is produced backward into a pair of large

lateral wings, the space between which is freely open below; and

again, to attempt to discriminate, as Cope does, between Woto-

pterus and Osteoglossum by the former having a “ double basis

cvanil” and the latter a “ simple basis cranii” is futile.

Since the terms posterior and lateral temporal grooves (or fosse,

as the case may be) have not always been employed in the same

sense In connection with the cranium of Teleostean fishes, it may

be well to explain that in this paper the prefix “ posterior tem-

poral” is applied to that groove or fossa which lies immediately

external to the epiotic bone, and the ‘“ lateral temporal” to that

depression which lies posterior to, and sometimes also above, the

postfrontal bone, just above the anterior part, or the whole, of the

articulation between the hyomandibular and the cranium. This,

I believe, is the most generally accepted usage of the expressions.

The former space is occupied by the trapezius and trunk-muscles

(Vetter, Jena. Zeitschr. xii. 1878), the latter by the dilatator

operculi and other muscles. The posterior temporal groove is

roofed over to form a posterior temporal fossa in Arapaima,

Osteoglossum, Albula, EHlops, Megalops, and Chanos.

In Clupea, Dussumieria, Chatoéssus, Chirocentrus, and Hugrau-

lis there is an aperture—the “‘ temporal foramen ”—in the side of

the cranium, bounded by the parietal and frontal bones. This

in life is occupied by a fatty mass, and in the dried skull leads

directly from the posterior temporal groove to the cavum cranii.

A short distance behind this is a lateral depression—the “ pre-

epiotic fossa ”—situated immediately in front of the epiotic bone,

and bounded by the parietal, squamosal, and epiotic. In Coilia

there is a small aperture immediately above the most dorsal part

of the upper of the two swim-bladder vesicles, which may possibly

correspond with the temporal foramen of Clwpea and its allies, but

the relations of the parts are rather aberrant. Even in Hngraulis

the pre-epiotic fossa is largely obliterated by the bulging of the

squamosal vesicle. The bottom of the depression is composed of

cartilage in Dusswnueria and in Clupea harengus, as also in

Osmerus, where, too, a shallow pre-epiotic fossa is recognisable.

In Hyodon and in Coregonus pollan, on the other hand, there is

no fossa, but a similar tract of cartilage is present, bounded by

the parietal, squamosal, and epiotic bones.

The large aperture—the “‘ lateral cranial foramen ”—in the side

of the cranium of Votopterus and the Mormyride is bounded by

the squamosal, epiotic, and exoccipital*, and may possibly be

* It is not bounded in Notopterus by the postfrontal and squamosal as stated

by Boulenger (Poissons du Bassin du Congo, 1901, p.115, and Ann. Mag. Nat. Hist.

1904, vol. xiii. p. 164), nor in the Mormyride by the opisthotic and parietal (Poiss.

Bass. Congo, p. 49).

62 DR. W. G. RIDEWOOD ON THE CRANIAL [May 3,

homologous with the pre-epiotic fossa. If the pre-epiotic fossa of

Clupea were to become larger and deeper, the exoccipital would

probably form part of its margin. The relation of the supra-

temporal bone to the lateral cranial foramen in the one case and

to the pre-epiotic fossa in the other, suggests a morphological

equivalence.

The lateral temporal groove is fairly well roofed over in Albula,

OChanos, and Heterotis, and partially so in Hlops, Arapaima,

Osteoglossum, Hyodon, and Notopterus.

In Osteoglossum, Albula, Hlops, and Megalops there is a

depression of considerable size in the side of the cranium, situated

beneath the posterior part of the articular surface for the head of

the hyomandibular. This is the ‘“subtemporal fossa”; it is

bounded by the pro-otic, squamosal, exoccipital, and opisthotic

bones. The fossa is recognisable in Arapaima, but it is only a

shallow depression. According to Hay (Zool. Bull. i. 1, 1898,

p- 30) it is well developed in the Cretaceous genus Xiphactinus,

in which it was erroneously described by Cope as. a foramen.

As already shown by Sagemehl (Morph. Jahrb. xvii.) the sub-

temporal fossa attains to great dimensions in such Cyprinoid

fishes as Cyprinus, Leuciscus, Labeo, and Abranis—it is practically

wanting, however, in Cobitis and Botia. In the former Cypri-

noids it is situated more posteriorly with respect to the articular

facet for the hyomandibular than is the case in the genera

mentioned at the beginning of the preceding paragraph, and it

opens vertically downward instead of downward and outward.

It is so large and deep as to be separated from the dorsal surface

of the skull by but a single layer of bone (epiotic bone). In these

fishes the subtemporal fossze serve for the lodgment of the great

muscles which, by pulling up the inferior pharyngeal bones (fifth

ceratobranchials), bring the teeth upon those bones forcibly against

the callous pad that is carried by the under surface of the basi-

occipital bone.

The subtemporal fossa is not present in the Salmonide. It

appears in the form of a wide but shallow depression in Pro-

chilodus, Alestes, and Hydrocyon*, but the fossa is unrecognisable

in the majority of the Characinide.

The “auditory fenestra,” bounded by the pro-otic, exoccipital,

and basioccipital, and leading into the perilymphatic cavity, is

present in Chatoéssus, Chirocentrus, Clupea, Dussunueria, Hn-

graulis, and Hyodon. In the last-named genus it is traversed by

a vertical bar of the pro-otic.

There are cecal diverticula of the swim-bladder contained in

the squamosal and pro-otic bones respectively in Chatoéssus,

Chirocentrus, Clupeat, Dussuwmieria, Hngraulis, and Coilia. Mega-

lops has a cavity in its opisthotic, which is probably occupied by

a similar diverticulum, although I have not had an opportunity of

tracing the connection with the swim-bladder ; but in Hops there

* See, for instance, Sagemehl, Morph. Jahrb. x. 1, 1884, pl. 2. figs, 13 & 14. -

+ In Clupea sprattus the pro-otic vesicle alone is present.

1904. | OSTEOLOGY OF THE ELOPID AND ALBULIDA. 63

is no such cavity. Theswim-bladder diverticulum in Hyodon and

Notopterus is of an entirely different character. It is large in

size, its outer wall is of fibrous tissue, and its inner wall is con-

stituted by the exoccipital and basioccipital in the former, and

the opisthotic and basioccipital in the latter genus. The want of

uniformity in the relations of the air-vesicles in the above-

mentioned fishes, coupled with the occurrence of such vesicles

in Mormyroids, in which the relations are yet again different,

points to the conclusion that such adaptive features cannot be

relied upon to any large extent in determining whether any two

fishes are closely or distantly related.

The opisthotice or intercalary bone does not appear to be at all

constant in bony fishes, as has already been pointed out by Vrolik

(Nied. Arch. f. Zool. i. 3, 1873), Klein (Jahresh. Ver. vaterl.

Naturk. Wiirtt. xxxv. 1879), and Sagemehl (Morphl. Jahrb. ix.

2, 1883). It must here be borne in mind, however, that what

Gill calls the opisthotic is not the bone that is now generally

known as the opisthotic, but the squamosal, which is of invariable

occurrence. (See Proc. U.S. Nat. Mus. xiii. 1890 (1891), pl. 30.

figs. 2-4, and pl. 31. figs. 2,4, & 5.) In this matter he appears

to be following the now obsolete terminology of Huxley (see

Giinther, ‘Study of Fishes,’ 1880, p. 60). Except in the Gadoid

fishes, the opisthotic is never of very large size. It is well

developed in Hyodon and Megalops, and is moderately large in

Notopterus, Gonorhynchus, Osteoglossum, Arapaima, and Heterotis ;

in most of the Clupeide it is small; it is absent in Hngraulis and

Coilia, in the Mormyride and in Alepocephalus.

As a general rule, in the forms under consideration, the lateral

wing of the parasphenoid that passes up along the anterior

edge of the pro-otic is of very small extent. In Osteoglosswm

leichardti, however, it rises so high as to meet the alisphenoid, and

in Osteoglossum bicirrhosum, Osteoglossuin formosum, and in

Gonorhynchus it enters into relation with both alisphenoid and

postfrontal. This is not exactly comparable with what occurs in

Amia, for in that genus there is one long process of the para-

sphenoid to the endosteal postfrontal (sphenotic), and a separate

shorter one to the alisphenoid. It may here be noted that

Swinnerton (Quart. Journ. Micro. Sci. xlv. 4, 1902, p. 532), in

mentioning the union of processes of the parasphenoid and frontal

immediately in front of the postfrontal bone in Gastrosteus, quotes

Klein as having recorded a similar relation obtaining in the case

of Lophius. ‘The junction, however, in Lophius occurs in front of

the optic foramen, and is in no way comparable with the above.

Although the view of Leydig (Zeitschr. f. wiss. Zool. v. 1854),

Hertwig (Arch. f. mikr. Anat. xi., Suppl. 1874; also Morph.

Jahrb. ii. 1876), Sagemehl (Morph. Jahrb. ix. 2, 1883), and

Klaatsch (Morph. Jahrb. xxi. 2, 1894), that such bones as the

vomer and parasphenoid have arisen by the coalescence of tooth-

bases, is not shared by Walther (Jen. Zeitschr. xvi. 1882) and

Carlsson (Zool. Jahrb. vill. 2, 1894), who claim that tooth-bearing

64 DR. W. G. RIDEWOOD ON THE CRANIAL [ May 3,

bones arise independently of the teeth, it is not unreasonable to

assume that the presence of teeth on these bones indicates a more

primitive condition than the absence of teeth—unless, of course,

the teeth are of secondary, and not primary origin. <A general

review of the skull in Teleostean fishes certainly goes to show that,

although no broad conclusions can be drawn from the presence or

absence of teeth on the vomer, a dentigerous parasphenoid is

habitually associated with other features which may be regarded

as archaic. Of the genera examined, Hlops, Megalops, Albula,

Arapaima, Osteoglossum, Notopterus, Hyodon, and the Mormy-

ride * have teeth on the parasphenoid.

There is probably in a considerable number of the lower

Teleosteans the half of a vertebral centrum fused with the basi-

occipital and exoccipitals to form the hollow-cone articular surface

at the back of the cranium, so that the cranial articulation really

takes place between this half-centrum and the first free centrum.

Such a condition has long been known to exist in Amia, in which

the corresponding neural arch persists; and Shufeldt (Rep. U.S.

Com. Fish. 1883 (1885), p. 816) has shown that in MJegalops the

suture between the half-centrum and the exoccipitals and basi-

occipital is readily recognisable. When this half-centrum is

removed, the transverse end of the basicranial axis presents a

rough surface, with a suture of the shape of an inverted T or Y

separating the basioccipital below and the right and left ex-

occipitals above. In all of the forms at present under considera-

tion, the wpper part of the suture between the half-centrum and

the exoccipitals is recognisable in the floor of the foramen magnum,

although the suture may no longer be visible down the side of the

half-centrum ; and in these cases the half-centrum can only be

removed by the application of some force, and the inverted T

suture above mentioned be brought into view.

Tn the Salmonide, in Hyodon and in Heterotis, although there

is not a very definite articulation between the first centrum and

the basioccipital and exocciprtals, and only a very sight range of

movement is possible between these, the centrum separates quite

readily, and exposes the trivadiate or inverted T suture, as already

figured by Gegenbaur (Festschr. A. v. Kolliker, Leipzig, 1887,

pl.i. fig. 9) and Parker (Phil. Trans. vol. 163. 1873 (1874), pl. viii.

fig. 8). The condition thus approximates to that which is so

commonly met with in the Acanthopterygian fishes, where the

eranial articulation is definitely between the basioccipital and

exoccipitals in front and the first vertebra behind.

In Arapaima the occipital articulation is further complicated

by the fact that the first complete centrum, although free from

the half-centrum in front of it, is not capable of free play, since

its lower portion sends forward a pair of long, stout processes

which are rigidly united by oblique, jagged sutures with the back

of the parasphenoid.

* The parasphenoidal teeth are rudimentary in Mormyrops and absent in

Gymnarchus.

1904. | OSTEOLOGY OF THE ELOPIDE AND ALBULID&. 65

In Gonorhynchus the back of the basioccipital region of the

skull is hemispherical. This convexity of the occipital articulation

is not, however, peculiar to this genus, for Owen and Klein have

recorded the occurrence of such a convex articular surface in

Fistularia (Anat. of Vert. i. 1866, p. 107; and Jahresh. Wiirtt.

1881, p. 325), and Klein in Syngnathus, Phyllopteryx, Gastero-

tokeus, and Ostracion (Jahresh. Wiirtt. 1885, p. 108).

Temporal and Preopercular Series.—The post-temporal is a con-

stituent of the pectoral girdle rather than of the skull, but a

description of the skull can hardly be considered complete unless

mention be made of the manner in which the attachment with

the shoulder-girdle is effected, and this, very naturally, leads to a

consideration of the post-temporal boneitself. The post-temporal

bone is no doubt primarily a scale-bone of the sensory- canal series.

Tt carries the lateral line forward to the supratemporal bone, and

the forward limb of the post-temporal is fairly well developed in

Hngraulis and Chirocentrus. The other two limbs, to the epiotic

and opisthotic respectively, are usually much larger and better

defined than the forward limb that passes towards the supra-

temporal and squamosal. The extremity of the epiotic limb is

usually loosely attached above the epiotic prominence by a broad,

short ligament, but in Chatoéssws the connection is much more

intimate and the freedom of play is greatly restricted. In Woto-

pterus the epiotic limb is wanting.

The opisthotic limb of the post-temporal is absent in Alepo-

cephalus, Coilia, and the Mormyride ; in Gynumarchus both epiotic

and opisthotic limbs are wanting. In Hngraulis the opisthotic

limb is attached to the back of the exoccipital in the absence of a

distinct opisthotic bone. The opisthotic limb lies deeper than the

epiotic limb, and appears not to be part of the dermal bone at all,

but rather one of the numerous intermuscular bones, at the back

of the cranium, which has acquired a secondary connection with

the post-temporal. The limb is always more or less rod-like

(except in Megalops), and unlike the two more superficial limbs.

In Clupea finta and in Chanos it forms the outermost element of

a series of three nearly parallel intermuscular bones, which

gradually increase in size from within outwards. The innermost

of the three is attached to that part of the exoccipital which forms

the lateral boundary of the foramen magnum; the second is

attached to the exoccipital bone midway between this point and

that process of the opisthotic to which the deep limb of the post-

temporal is attached. The posterior ends of the inner and middle

rods spread out like stiff brushes among the trunk-muscles. In

Chirocentrus there are three brushes of tendon-bones on each side

of the back of the cranium; one arises from the exoccipital

immediately dorso-laterally to the vertebral articulation, the

second from the exoccipital a little more laterally, and the third

from that part of the exoccipital that touches the mesial edge of

the opisthotic. This last tendon-bone lies immediately mesial to

Proc. Zoo. Soc.—1904, Vou. II. No. V. D

66 DR. W. G. RIDEWOOD ON THE CRANIAL [May 3,

the opisthotic limb of the post-temporal. In Osteoglosswm also

there are two important intermuscular bony brushes on each side.

These osseous brushes are probably far more common in their

occurrence than is generally suspected, and the reason that they

have not attracted more notice in the past is probably due partly

to the fact that anatomists have been disposed to discount their

value as constituents of the skeleton (although they are just as

important as epipleural bones, which are not disregarded when

dealing with the vertebral column), and partly also because the

preparateur dissects them away from the back of the cranium

when removing the skull from the vertebral column, and they

thus become thrown away with the muscles of the trunk. They

have been noted, however, in the Black Bass and the Tunny by

Shufeldt, who calls them “occipital ribs” (Rep. U.S. Fish. Com.

1883 (1885), p. 805). Hyrtl (Denkschr. Akad. Wiss. Wien, xxi.

1863, p. 3), with considerable acumen, has likened those at the

back of the skull of Chanos to the ossified tendons of birds.

Such intermuscular brushes are not confined to the exoccipital

and opisthotic bones. In Chatoéssws they occur on the supra-

occipital and epiotic prominences, and in Chanos the supraoccipital

spine is produced back into a bony brush exactly similar to that

of Chatoéssus, except in that it is not a separate structure. In

Gonorhynchus the posterior end of the outer intermuscular bone

is connected with the post-temporal by means of fibrous tissue.

There is no opisthotic limb of the post-temporal besides this,

which is clearly the opisthotic limb which has failed to establish

the usual osseous connection with the post-temporal. In Albula,

in addition to a fully developed opisthotic limb, there is an ossified

tendon which projects forward from the inner surface of the post-

temporal bone into the posterior temporal fossa, where it branches

among the fibres of the trapezius muscle.

The supratemporal may in a general way be described as that

dermal bone which receives the lateral line from the post-temporal

and transmits it to the squamosal, and gives off a branch to the

parietal (e. g. Clupea), or to one or more tubular scales of the

transverse commissure of the sensory-canal system separable from

the parietal (e. g. Albula. Salmo). The form of the supratemporal

is thus triradiate. In Gonorhynchus, however, in which also the

commissural scales are separable from the parietals and supra-

occipital, the forking of the canal-system occurs just in front of

the supratemporal, so that this bone is a plain tubular scale.

The supratemporal is a bone which lies either above or posterior

to the squamosal ridge, and either below or external to the epiotic

ridge; and Collinge (Proc. Zool. Soc. 1895, p. 291) is undoubtedly

right in questioning the correctness of Parker’s application of the

term supratemporal to that bone of the lateral-line system which

in the Salmon lies above the preopercular and below the squamosal

ridge (Phil. Trans. vol. 163. 1873, p. 99, and pl. 6. fig. 1, sé).

Parker’s “‘supratemporal” appears to correspond exactly with

that bone which in Chanos lies immediately above the opercular

1904. | OSTEOLOGY OF THE ELOPID® AND ALBULID. 67

bone and carries the sensory canal from the antero-ventral corner

of the supratemporal downwards to the upper end of the pre-

opercular. This bone I propose to term the swbtemporal. The

real supratemporal of the Salmon was overlooked by Parker alto-

gether, although it is a larger bone than the subtemporal. It

lies between the post-temporal and the back of the squamosal,

and bears the usual triradiate sensory canal. In Bruch’s ‘ Ver-

gleichende Osteologie des Rheinlachses’ (Mainz, 1861), a work to

which, curiously enough, Parker refers (¢. ¢. p. 142), both supra-

temporal and subtemporal are correctly shown, the former being

marked «'" (pl. 2. fig. 1) and the latter a.

The subtemporal attains its greatest development, so far as lam

aware, in the Characinid genus Sarcodaces, in which it appears

as a kind of supraopercular bone. Smith Woodward (Proc. Zool.

Soc. 1887, p. 536) has recorded the occurrence of a similar bone

in Fhacolepis, a Cretaceous Teleostean from Brazil which he is

disposed to associate with the Elopidee.

In dArapaima and Osteoglossum the supratemporal is a stout,

partially sculptured bone, firmly united with the cranium

(squamosal and parietal bones). In the Mormyride it is a large,

thin scale of bone forming a loose lateral cover to the lateral

cranial foramen, and in Wotopterus the relations are the same,

although the supratemporal bone itself is much smaller. In

Hyodon the supratemporal is a large curved scale of triangular

shape, which unites in the dorsal median line with its fellow of

the opposite side, and covers the whole of the parietal and a small

part of the frontal as well. In his “Synopsis of the Families of

Teleostean Fishes” (Ann. & Mag. Nat. Hist. (7) xiii. 1904, p. 164)

Boulenger groups the Mormyridz with the Hyodontide on the

ground that they both have the “supratemporal large, plate-like,

covering the greater part of the parietal bone.” As a matter of

fact the supratemporal of the Mormyridz covers very little of

the parietal. In Petrocephalus bane it just overlaps the lateral

edge of the parietal, in Mormyrus oxyrhynchus it just touches

the postero-lateral corner, while in Mormyrops deliciosus and

Gymnarchus niloticus it does not reach the parietal at all.

In Hlops and Megalops the supratemporal is a scale extending

inwards as far as the median plane of the head, overlapping the

top of the opercular bone, and attached by its anterior edge to

the transverse parieto-squamosal ridge. The transverse com-

missure of the sensory-canal system thus does not run in the

parietal, but above it. Each supratemporal is notched behind,

giving the impression of a transverse row of four bones. In

Dussumieria the notch completely divides the bone, giving a series

of four, the laterals of which, according to the foregoing definition,

alone can claim to be regarded as supratemporals. In Chanos

there is a series of eight such bones. Those nearest the median

plane are narrow tubular bones, the next are fused early with the

parietals (the former fuse later with the latter), then come the

supratemporals, and outermost of all the subtemporals.

68 DR. W. G. RIDEWOOD ON THE CRANIAL [May 3,

A very definite relation exists between the supratemporal and

preopercular bones. Disregarding a few exceptional cases, the

supratemporal is a bone which is distinguished by a triradiate

sensory canal, as already pointed out. The posterior tube is con-

tinued back into the post-temporal, the upper tube runs over the

cranium, usually in or over the parietal bones, as the transverse

commissure, while the antero-ventral tube, after traversing the

smpenieul part of the squamosal, passes down into the pre-

opercular*, either directly or through a subtemporal bone. The

opercular and subopercular bones and the branchiostegal rays, on

the other hand, carry no sensory canal, but are ossifications in

the movable flap or gill-cover that projects back from the hyoid

arch, and are thus bones of a different category. This exclusion

of the preopercular from the opercular series is not a new idea,

although independently arrived at, for it is to be noted that Cole

and Johnstone (Proc. & Trans. Liverpool Biol. Soc. xvi. 1902,

p. 175), in describing the osteology of the Plaice, have pointed

out that the preopercular i is a bone developed primarily around a

portion of the lateral-line system, and is therefore of a different

nature from the other opercular bones. On page 177 they classify

the preopercular with the lachrymal, nasal, suborbital and supra-

temporal bones, a step which I am fully prepared to endorse.

The determination of the morphological value of the inter-

opercular bone is not an easy matter. On examining the skull of

Amia it is difficult to believe that the interopercular does not

belong to the same series as the opercular and subopercular bones,

and yet in Lepidosteus the interopercular is quite removed from

the subopercular, and is situated at the front of the lower or

horizontal limb of the preopercular bone. Possibly the name

interopercular has been appled in different fishes to bones of

different morphological value. However, on the separation of the

preopercular from the opercular and branchiostegal series of bones

and the introduction of it into the temporal series, it becomes

necessary to decide whether the interopercular belongs to the one

or the other of these groups, and on the whole I am disposed to

regard it as more closely related to the preopercular than to the

subopercular and opercular bones.

This is not the view taken by Cole and Johnstone, who consider

(Proc. & Trans. Liverp. Biol. Soc. xvi. 1902, p. 175) that the not

infrequent articulation between the interopercular and the epihyal

confirms the view that the interopercular, like the opercular, is a

modified branchiostegal ray.

The ligamentous connection so commonly met with between

the front of the interopercular and the back of the mandible is

probably merely adaptive, and does not point to any morpho-

logical relation existing between this bone and the mandibular

arch. The question is discussed by Rau? (Morph. Jahrb. iv.

Suppl. 1878, pp. 15 & 16, footnote).

* To be strictly logical, such bones as the squamosal should also be classed

as sensory-canal bones of the temporal series; but it is clearly; more expedient to

regard them as component parts of the “cranium.”

1904.] -- OsTHOLOGY OF THE ELOPIDA AND ALBULID. 69

It might reasonably be urged that the interopercular is a bone

of the true opercular series which has been squeezed out of its

primitive position between the subopercular and the last branchi-

ostegal ray, in much the same way as the penultimate member of

the branchiostegal-opercular series has been reduced and forced

out of its position in the Paleoniscid fishes. The bone in question

was originally regarded by Traquair (“‘Ganoid Fishes Brit. Carb.

Form.,” Paleontogr. Soc. 1877, p. 20) as the subopercular, but

subsequently (ibid. 1901, p. 62) as an accessory plate. Smith

Woodward appears to regard it as the interopercular (Brit. Mus.

Cat. Foss. Fish. ii. 1891, p. 487). But, whatever may be the

homology of the bone, the evidence that it has been excalated

and finally lost is tolerably clear. In Teleosteans, however, there

is no evidence available to show that any such displacement of

the interopercular has taken place; even in Pholidophorus, the

lowest of the Teleostean series, the interopercular already occupies

its definitive position in advance of the subopercular.

In that very aberrant genus Phractolemus, the preopercular is:

small and the interopercular remarkably large. The latter bone

is situated below and anterior to the preopercular, and receives

from it the sensory canal that descends from the squamosal..

The interopercular is thus here performing the function of the

missing horizontal limb of the preopercular. In no other case

have I seen the interopercular conveying a sensory canal; in the

vast majority of cases the interopercular is a thin lamina of bone

which is almost entirely concealed by the lower part of the pre-

opercular. The evidence for putting the interopercular with the

preopercular is unsatisfactory, but owing to the perfect manner

in which the branchiostegal rays grade off into the subopercular

and opercular, there is no justification for including it with these

last as a constituent of the skeleton of the gill-cover.

The interopercular is of very regular occurrence, but it is said

to be wanting in Pantodon. As regards the preopercular, it may

be taken as a general rule, not however without exception, that

the horizontal limb of this bone is most developed in those forms

with a greatly reduced mouth, e. g. Gonorhynchus and Chanos,

and absent in those with a very large gape, e. g. Hngraulis and

Coilia. In Albula and Alepocephalus the vertical and horizontal

limbs are nearly equal.

Oireumorbital and Nasal Series—No results of any great

morphological importance are to be expected from a comparative

study of the bones of the cireumorbital series. They are probably

more subject to variation—generic, specific, and individual—than

any other bones of the skull; and the number of the bones

surrounding the orbit is not infrequently found to differ on the

right and left sides of the same skull.

In Arapaima, Heterotis, and Osteoglosswum the enlargement of

the nasal bones, their meeting in a median suture, and their rigid

union with the cranial bones, are features which are not en-

countered in any of the other genera under examination. The

70 DR. W. G. RIDEWOOD ON THE CRANIAL [May 3,

nearest approach to such a condition is found in Wotopterus, in

which the two nasals are fairly large, and just meet in the middle

plane of the head, and are with some difficulty removable from

the underlying mesethmoid and prefrontal bones. A very similar

condition obtains in Petrocephalus, in which the nasal bones are

exceptionally large.

The postorbital cheek-plates are large in Arapaima, Osteo-

glossum, Hyodon, Elops, and Megalops. Both postorbital and

suborbital bones are large in Adbula. In Chirocentrus the bone

lying antero-ventral to the orbit is large. There is one large

suborbital, extending also behind the eye, in Chanos, Hngraulis,

and Olupea harengus, but in Clupea jfinta the postorbital part of

the cheek is protected by two large plates. In some forms, such

as Coilia, Albula, and Notopterws, in which the postorbital and

suborbital sensory canals are large, the bones have a scroll-like

form; but in the other genera the sensory canal lies farther from the

orbital edge of the circumorbital bones (e. g., Clupea, Chatoéssus),

or is more deeply embedded in the bone (e. g., Arapaima, Chanos).

In Notopterus there are processes of the two hindermost of the

suborbital bones directed inwards below the eyeball towards the

ectopterygoid and entopterygoid, with which they are united by

ligament. There is also a process passing inwards from the

anterior end of the front suborbital bone, and entering into close

fibrous union with the under surface of the prefrontal. Boulenger

finds that in some families of the Perciform fishes the “ subocular

shelf” is sufficiently constant to be of taxonomic value (Ann. &

Mag. Nat. Hist. (7) xi. 1904, p. 179).

In this connection it is interesting to note that in Albala and

in Dussumieria there is an outwardly-directed process of the

ectopterygoid, underlying the eyeball, which meets the edge of

the suborbital bones. This process, like the subocular shelf,

serves the purpose either of supporting the eyeball, or of limiting

the lateral play of the series of suborbital bones.

In Osteoglossum there is a close fibrous connection between the

maxilla and the preorbital and suborbital bones, but this is not

the case in Heterotis and Arapaima.

Maxillary Series.—The evidence of paleontology goes to show

that the most primitive form of mouth is that bounded above by

a small premaxilla and a comparatively long maxilla; at all

events, an enlarged premaxilla has not yet been noted in Iso-

spondylous fishes below the Cretaceous strata (Smith Woodward,

Vert. Paleontology, 1898, p. 113). Most of the forms now under

examination have a premaxilla-maxillary gape; but in Albula,

in which, according to Jordan and Gilbert (Bull. U.S. Nat.

Mus. No. 16, 1882, p. 258) and Smith Woodward (Brit. Mus.

Cat. Foss. Fish. iv. p. 59), the lateral margin of the upper jaw

is formed by the maxilla, the maxilla can be of service only

when the mouth is widely opened, and indeed, since it bears no

teeth, while the premaxilla does, it probably does not function in

1904.] OSTEOLOGY OF THE ELOPIDH AND ALBULID&. 71

prehension at all. A similarly constituted mouth occurs in the

Mormyride. The right and left premaxille of Mormyroid fishes

are fused, and the suture obliterated (except in Gymnarchus).

A similar condition is said to exist in Pantodon. In Chanos,

Chatoéssus, and Gonorhynchus the premaxilla alone bounds the

gape above, and is devoid of teeth.

The exceptional, and apparently useless, backward prolongation

of the maxilla of Coilia is a well-known feature of that genus, and

claims but a passing mention.

There are two surmaxille above each maxilla in Coilia, Hn-

graulis, Clupea, Dussumieria, Chirocentrus, Elops, Megalops, and

Alepocephalus, one in Chatoéssus and Albula, and none in Chanos,

Arapaima, Heterotis, Osteoglosswm, the Mormyride, Hyodon,

Notopterus, and Gonorhynchus. Two surmaxille are present in

the Mesozoic families Pholidophoride. Leptolepide, and Oligo-

pleuride.

Mandibular Series.—The angular bone is distinct in Arapaima,

Heterotis, Osteoglossum, Notopterus, and most Clupeoids, but not

in Coilia and Engraulis, nor in the Mormyride, Hyodon, Albula,

Elops, and Megalops. In Notopterus the angular is a much larger

bone than usual. The endosteal and ectosteal parts of the articular

bone are distinct and separable in Arapaima; and, according to

Hay (Zool. Bull. ii. 1, 1898, p. 37), they are also distinct in the

Cretaceous genus Xiphactinus. He shows that Cope’s interpre-

tation of the mandibular bones of this form was entirely erroneous.

Since the remark of Owen’s (Anat. of Vert. i. 1866, p. 123), that

in Arapaima there is a superadded bony piece answering to the

surangular of Reptiles, is credited by so recent a writer as Smith

Woodward (Brit. Mus. Cat. Foss. Fish. ui. 1895, p. xix), it may

be well to point out that this superadded bone, marked 29 @ in

Owen’s fig. 88, is but the endosteal articular displaced. In

Gymnarchus, Hyodon, Albula, Hlops, and Megalops the endosteal

and ectosteal components of the articular would probably separate

with prolonged maceration, for the suture between them is clearly

visible, but I was unable to submit the material in hand to such

treatment. As a rule the suture is not visible in Teleostean

fishes.

In some forms the posterior part of the surface for the articu-

lation with the condyle of the quadrate is formed by the angular,

the part of the angular bone concerned having a distinctly end-

osteal appearance. Such is the case in Gonorhynchus, Arapauma,

Albula, Megalops, Hlops, Gymnarchus, Hyodon, and Heterotis, but

in Osteoglossum, Notopterus, and the Clupeidee the articular facet

is formed by the endosteal articular alone. In Wotopierus and

the Clupeide the angular bone has the appearance of an ectosteal

bone.

Except in a few forms (e.g., Arapaima, Petrocephalus, Hngraulis,

Chatoéssus, Dussumieria) there is a distinct bone lying usually in

front of the endosteal articular and on the inner (lingual) surface

(@ - DR. W. G. RIDEWOOD ON THE CRANIAL [ May 3,

of the ectosteal articular, and developed above or-around Meckel’s

cartilage. This gives attachment to the tendon of a part of the

levator muscle of the mandible, and may be called the ‘sesamoid

articular,” thus making a third component of the articular bone.

Vetter (Jena. Zeitschr. xii. 1878, pl. 13. fig. 8, Ss.) has called

attention to this bone in the Pike as a “ Sesamoidverknécherung

an der Insertion der Hndsehne von A,—A,* an Meckel’schen

Knorpel,” A, being his third or deep portion of the adductor

mandibulz, and A,‘ its tendon.

This sesamoid articular is remarkably large in A/bula (text-fig. 17,

p. 50), Gymnarchus, and Hyodon, in which it is set on the antero-

superior side of the endosteal articular. It is of fair size in

Notopterus, Osteoglossum, and Gonorhynchus; but as a rule it is

small, and is situated just in front of the endosteal articular

(e. g., Heterotis, Clupea), or at a short distance in advance of it

(e. g., Hlops, Megalops).

The sesamoid articular was described in 1878 in the mandible

of Belone as a “ coronoid bone” by Cope (Proc. Amer. Phil. Soe.

xvi. p. 695), who was of opinion that the bone occurred in no

family of Teleostean fishes except the Belonidee. Guill, writing in

1895, described the bone in question as “lying mostly inside of

the upper portion of the dentary” (Proc. U.S. Nat. Mus. xviii.

1895, p. 173), and, admitting that it was not in any way homo-

logous with the coronoid of Lepidosteus, proposed to call it the

“addentary” (J. c. p. 174). The relation of this bone to the

dentary was, however, based upon an error on the part of Dr. Gill.

He writes (/. c. p. 173) :—‘‘ This element appears to have been

unnoticed by most naturalists, and to have been first observed by

Dr. B.C. Brithl. In 1847 Briihl (Anfangsgriinde der vergl. Anat.

aller Thierklassen, Atlas, pl. xi. fig. 17) published a figure of the

disintegrated right mandible in which the supplementary bone is

marked ZK. I have, however, been unable to find any reference

to it in the text.” Asa matter of fact, only two of the four bones

figured by Briihl are those of the mandible, the other two are

parts of the upper jaw, and are marked ZK and OK, which

abbreviations are explained on p. 88 of the text as standing for

‘ Zwischenkiefer’ and ‘ Oberkiefer’ respectively.

The sesamoid articular has been minutely studied by Prof.

Starks, who in a letter to me dated March 15, 1904, states that it

is a small ossicle situated on the inner surface of the articular,

just above Meckel’s cartilage, that it occurs at the lower end of

the ligament (?tendon) which is attached to the articular, and

that it occurs much more commonly than is generally supposed.

In 1900 he published (Proc. U.S. Nat. Mus. xxii. 1900, p. 2,

footnote) a list of eighteen genera of Teleostean fishes in which

he had detected the presence of this “‘ coronoid bone.”

Although teeth occupying the position of splenial teeth, and

referred to as such by Owen (Anat. of Vert. i. 1866, p. 123),

occur in the mandible of Arapaima, no separate splenial is to be

found in any of the fishes examined, which, being the lowest of.

1904. ] OSTEOLOGY OF THE ELOPIDZ AND ALBULIDA. (i)

the Teleostean series (excluding extinct forms such as the Pholi-

dophoridee, in which “it is not yet quite clear that the mandible

was destitute of splenial and coronoid elements” (Smith Wood-

ward, Vert. Paleontol. 1898, p. 114), might more than any others

be expected to possess traces of this constituent.

The size and shape of the dentary bone vary very considerably.

The bone is usually large; but it is much reduced, especially in

its anterior parts, in Chanos. The coronoid process is usually

formed entirely by the dentary, but the ectosteal articular occa-

sionally forms the hinder part of it. In Chatoéssus the process is

situated unusually far forward, in a position recalling that of the

remarkable coronoid process of Gonorhynchus.

In the extinct family Saurodontide the teeth are lodged in

sockets; and Boulenger, in his recent synopsis of the families of

Teleostean fishes (Ann. & Mag. Nat. Hist. xiii, 1904, p. 164),

places the Chirocentride next the Saurodontide, and separates

them from the Clupeidze because they have ‘“ teeth in sockets.”

The teeth of the three specimens of Chirocentrus dorab examined

for the purposes of this investigation are, however, certainly not

lodged in sockets; they are anchylosed to the edge of the bone,

and are flanked by a slight ledge on the external side, exactly as

in Coilia, for instance.

Hyopalatine Series.—The presence of one or two articular heads

for the front of the hyopalatine arch, a matter upon which Swin-

nerton (Quart. Journ. Micro. Sci. xlv. 4, 1902, pp. 551, 556-557,

& 584) lays considerable stress, may possibly be an adaptive

feature related to the length of the ethmoid region of the skull,

for in Hlops and Megalops the ethmoid region is short, and the

front of the hyopalatine arch has but a single head, whereas in

Albula the ethmoid region is long and there are two heads to the

palatine. The difference does not strike one at the outset as

likely to be of fundamental importance: extended investigations

upon dissected, 2. e. not dried, skulls alone can decide the point.

An unusual condition occurs in Cotlia, i which the ethmoid

region of the cranium is short and the palatine has two heads;

but these are right and left, and not anterior and posterior. This

peculiarity is not shared by the allied genus Hngraulis.

The palatine bone is fused with the ectopterygoid in Arapaima,

Osteoglossum, and Notopterus, and in these genera ‘and in Hyodon

there appears to be no endosteal part of the palatine. In the

Mormyroid fishes there is no separate entopterygoid, and the

palatine bone is fused with the side of the vomer.

The metapterygoid attains its maximum size relatively to the

adjacent bones in Coilia; in Gonorhynchus the other extreme is

reached, for here it is reduced to a fine needle of bone. The

articulation between the metapterygoid and entopterygoid region

of the hyopalatine arch of Osteoglosswm and the lateral process of

the parasphenoid described by Bridge in 1895 (Proc. Zool. Soe.

1895) is considered by Swinnerton (/. c. p. 572). on embryological -

74 DR. W. G. RIDEWOOD ON THE CRANIAL | May 3,

grounds to be a primitive articulation persisting in this case, but

lost in the great majority of Teleostean fishes. Such an articula-

tion also exists in Arapaima and Heterotis. In the Mormyroid

fishes the whole of the upper edge of the hyopalatine arch enters

into extensive relation with the under surface of the cranium.

The symplectic is usually rod-like, but in Arapaima, and to a

less extent in Osteoglossum and Notopterus, it spreads in a

squamous manner over the surrounding bones. It is wanting in

the Mormyride. The slope of the hyomandibular varies con-

siderably in the different genera, as also does the angle which

the symplectic makes with the axis of the hyomandibular.

The ectopterygoid is in some cases straight (e. g. Hyodon), in

others moderately bent (e. g. Clupea) or sharply angulate (e. g.

Chatoéssus).

The hyomandibular articulates with the cranium by either one

or two heads; and since two species of the same genus may vary

in this respect, one may conclude that no great importance need

be attached to the difference. The hyomandibular of Clupea

jinta differs from that of Clupea harengus in possessing two

separate heads for articulation with the cranium, whereas the

latter has a single head, broad antero-posteriorly. Since Swin-

nerton (J. c. pp. 541, 544, & 549) has shown that in the de-

velopment of Gustrosteus the double head of the hyomandibular

results from the enlargement of the anterior and posterior ends,

and the reduction of the middle part, of a single broad head,

Clupea harengus may possibly be more primitive than Clupea finta

in this respect.

One would imagine on general grounds that a diminution in

the size of the mouth would be associated with a forward slope of

the hyomandibular, a marked angulation between the symplectic

and the hyomandibular, and a sharp bend in the middle of the

ectopterygoid ; and this is certainly what one finds in Chatoéssus,

and to a iesser extent in Clupea harengus as compared with

Clupea finta. Yet, while Megalops has a smaller mouth than

Hlops, ts symplectic is more nearly in a line with the hyoman-

dibular than in the latter genus; that is to say, the angle, as

measured in degrees, is greater (cf. text-figs. 14 & 10, pp. 46 & 40).

The angulation of the ectopterygoid is approximately the same in

both, and the advancement of the quadrate articulation in Megalops

is brought about by the forward slope of the hyomandibular only.

fyodon has a fairly large mouth, associated with a perfectly straight

ectopterygoid and a backwardly sloping hyomandibular, yet, as in

Elops, the symplectic is strongly bent upon the hyomandibular.

Somewhat similar relations—the symplectic angle, however, being

more open—obtain in the large-mouthed genera Hngraulis and

Coilia. On the other hand, it is interesting to note that in the

small-mouthed forms Gonorhynchus and Chanos the hyomandibular

does not slope forward. In the latter genus the quadrate has

shifted forward along the ectopterygoid, and has parted from the

symplectic entirely.

1904.] OSTEOLOGY OF THE ELOPIDE AND ALBULIDA. 75

In Chirocentrus, although the mouth is fairly large, the quad-

rate is advanced by a forward rotation of the hyomandibular.

The symplectic in this case is in a direct line with the axis of

the hyomandibular. In Arapaima the mouth can hardly be

described as small, yet the forward slope of the hyomandibular is

excessive. The explanation of this is most probably to be sought

in the great length of the postorbital portion of the head.

In Engraulis and Coilia alone of the forms under consideration

does the quadrate slope backward.

Opercular Series—The reasons for excluding the preopercular

and interopercular bones from this series are given on p. 68.

The bones considered under the present heading are the opercular

and subopercular bones, the branchiostegal rays and the jugular

late.

: Ag is well known, the subopercular bone is wanting in Woto-

pterus. This condition is unparalleled among the other forms

under consideration, although the subopercular is distinctly small

in Osteoglossum and Arapaima, and very small in Heterotis. It

is comparatively large in Albula and Gonorhynchus; it is said to

be wanting in Pantodon.

The lower Malacopterygian fishes are well adapted for demon-

strating the continuity of the opercular and branchiostegal

systems. The opercular and branchiostegal bones are functionally

similar, serving to support the gill-cover, and there seems to be

good reason for regarding them as morphologically similar also.

The view is by no means a new one, for Traquair (“‘ Ganoid

Fishes Brit. Carb. Form.,” Palzeontogr. Soc.) demonstrated it in the

case of the Paleoniscide in 1877, and Shufeldt (Rep. U.S. Com.

Fish. 1883 (85), pp. 818 & 820) mentioned it in 1883; and

although the latter does not in his paper give references to

previous expressions of such opinion, he does not himself claim

the idea as original. The only recent reference to such a view

appears to be that by Cole and Johnstone (Proc. & Trans. Liverp.

Biol. Soc. xvi. 1902, p. 175), who mention the opercular bones

(excepting the preopercular) as “ modified branchiostegal rays.”

In Chanos, Albula, and Hlops the transition from the upper-

most branchiostegal rays to the subopercular and opercular bones

is very evenly graduated, and in Osteoglossum it becomes a matter

of some difficulty to decide whether the bone lying antero-

ventrally to the opercular is a reduced and displaced subopercular,

or the uppermost branchiostegal ray which has lost its connection

with the epihyal. The difficulty of deciding whether a bone lying

below the opercular bone is a subopercular or a free branchi-

ostegal ray, the true subopercular being wanting, must frequently

have occurred to the systematic ichthyologist. Boulenger dis-

tinctly admits the difficulty when dealing with the Mormyroid

fishes (‘ Poissons du Bassin du Congo,’ 1901, p. 50, footnote). In

preparing the skull of Hngraulis the subopercular comes away

readily with the epihyal, and is very liable to be mistaken for a

76 DR. W. G. RIDEWOOD ON THE CRANIAL [May 3,

branchiostegal ray, a fact already pointed out by Valenciennes

(Hist. Nat. Poiss. xxi. 1848, p. 11).

Incidentally it may be pointed out as an item of evidence in

favour of regarding the hyomandibular as an element of the

hyoid arch, a view which Pollard has contested*, that there is

invariably a definite articulation between the front of the oper-

cular bone and a special process from the back of the hyoman-

dibular, comparable with the less definite articulation between

the antero-superior extremities of the branchiostegal rays and the

epihyal and ceratohyal.

It is generally assumed, and the results of the present haces

gation tend to show that the assumption is justifiable, that a

large number of branchiostegal rays is a primitive character.

The greatest number of rays occurs in Hlops, which has from 30

to 35 on each side; Megalops has about 24; and Hngraulis, Coilia,

and Dussumieria from 10 t013. <Albula has 15, but the allied deep-

sea genus Dathythrissa only possesses 6. The commonest numbers

are from 6 to 9: Chanos and Gymmnarchus have as few as 4.

The jugular plate present in Hlops and Megalops is not defi-

nitely related to the hyoid arch, but, on the contrary, is united by

ligament with the mandibular s symphysis, Functionally, at all

events, the jugular plate belongs to the branchiostegal series, and

it is more convenient to treat it under this heading than else-

where. Parker’s recognition of the urohyal as a basibranchiostegal

is altogether erroneous. The urohyal is never superficial and does

not support the gill-cover; it is an ossified tendon of the lingual

retractor muscles (see ‘‘ Hyobranchial Series” below).

Hyobranchial Series —The unfortunate application by Parker

(Phil. Trans. vol. 163. 1873 (1874), p. 101) of the name “ basibran-

chiostegal” to the urohyal bone has occasioned much confusion,

and in spite of the unsuitability of the term it continues to be

employed by certain writers, and appears in a paper published as

recently as 1901 (Supino, Ric. Lab. Anat. Univ. Roma, vii. 3,

1901, p. 18). EHvenif the urohyal could be shown to belong to

the branchiostegal series, the prefix basi- would be misleading as

implying homology with the component parts of the copular

skeleton, for the branchiostegal rays are dermal, not visceral

bones. Asa matter of fact, the urohyal is an ossified tendon, or

an ossification of the mtermuscular septa, as Parker himself

admitted, lying between the two sternohyoid muscles, and is

* Anat. Anz. x. 1895. By his describing the Teleostean skull as metautostylic

(p. 25) I conclude he regards the hyomandibular as a part of the mandibular arch ;

at all events he considers the hyomandibular of the Teleostean to be homologous

with the prespiracular cartilage of Sharks, which, from its position in front of the

spiracle, is undoubtedly a constituent of the mandibular and not of the hyoid arch.

+ Gegenbaur (Morph. Jahrb. iv., Suppl. 1878, p. 17) writes :—“ Er dient, wie sonst,

zur Insertion der subbranchialen Muskulatur, und hat weder zur Membrana br anchi-

ostega noch zu den Kiemenbogen irgend eine Beziehung.” See also Vetter, Jena.

Zeitschr. xii. 1878, pl. xin. fig. 10, Sth. Ginther (‘ Study ‘of Fishes,’ 1880, p. 91) says

that the urohyal “separates the musculi sternohyoidei, and serves for an increased

surface of their insertion.”

1904. ] OSTEOLOGY OF THE ELOPIDH AND ALBULIDS. ne

attached by two ligaments to the right and left hypohyal bones,

except in the Mormyroid fishes, in which it is rigidly fixed beneath

the anterior part of the copular skeleton.

_ Shufeldt (Rep. U.S. Com. Fish. 1883 (85), p. 820) writes that

the urohyal “lies between the sternohyoid muscles, and is not

always present where a glossohyal exists.” The latter part of the

quotation is, I think, open to serious question; it is probably

always present in Teleostean fishes. Brooks (Proc. Roy. Dubl.

Soc. n. s. iv. 4, 1884, pp. 180-183), in his description of the skull

of the Haddock, while adopting the name basibranchiostegal for

this tendon-bone, misapplies the term urohyal to that cartilage

which lies between the last two pairs of ceratobranchials, and

represents the fourth and fifth basibranchials.

In connection with the urohyal, it is of interest to observe that

in Votopterus there are, in addition to the urohyal proper, a pair

of tendon-bones of similar character to it, but of smaller size,

projecting backwards from the posterior end of the ventral surface

of the second basibranchial. Such tendon-bones are also present

in Osteoglossum, Heterotis, and the Mormyride; but in these

fishes they are confluent with the reduced second hypobranchials.

In dealing with the homologies of tendon-bones, one must ever be

prepared to admit the possibility of convergence; thus, while in

Notopterus there are separate tendon-bones related to the second

hypobranchials and the second basibranchial, in Diodon (Dicotyl-

ichthys) a pair of exactly similar bones project down from the

mesial ends of the third ceratobranchials, and represent either the

downwardly directed and greatly elongated third hypobranchials,

or a pair of tendon-bones confluent with the third hypobranchials,

or simply a pair of tendon-bones, the third hypobranchials being

absent. In Polypterus, again, the urohyal as a median bone is

wanting, but a pair of tendon-bones project downward and

backward from the ventral surface of the anterior end of the

ceratohyal.

The hypohyal of each side of the head is usually double, con-

sisting of distinct upper and lower ossifications. The right and

left upper hypohyals are separated by the first basibranchial ox

the glossohyal, or both, but the two lower hypohyals are articu-

lated together in the median plane. The lower hypohyal in most

cases 18 larger than the upper, but in Hyodon, Hlops, Megalops,

and Albula the upper and lower hypohyals are approximately

equalin size. In Arapaima, Heterotis, Osteoglossum, Petrocephalus,

and Notopterus there is but one hypohyal on each side, and this

would appear to represent the upper one. This is rather odd

when considered in relation with the probability that the

single hypohyal of Amita represents the lower of the two. In

Mormyroids other than Petrocephalus no hypohyals are recog-

nisable.

It is difficult to understand why Supino, in his recent work on

the skull of deep-sea Teleosteans (Ric. Lab. Anat. Univ. Roma,

vill.ix. 1901-2), and Starks, in his description of the Serranoid

78 DR. W. G. RIDEWOOD ON THE CRANIAL [May 3,

Roceus (Proc. Wash. Acad. Sci. iii. 1901, pl. lxiv.), allude to the

hypohyal as the basihyal. Owen (Anat. of Vert. i. 1866, pp. 106

& 124), it is true, used the word basihyal in this sense, and was

followed by Giinther (‘Study of Fishes,’ 1880, p. 58) and others;

but the homology between the glossohyal of the Teleostean and

the basihyal of the Elasmobranch is now so firmly established that

there is no justification for reviving an erroneous terminology.

What is more incomprehensible than the retention of an obsolete

application of the term basihyal is the fact that Starks, while

calling the hypohyals the basihyals in 1901 (/.¢.), designates them

hypohyals in 1898 and 1904 (Proce. Calif. Acad. Sci. (3) i. 2, 1898,

pl. xxiii. fig. 8, Sebastolobus; and Proc. U.S. Nat. Mus. xxvii.

1904, p. 603, Berycoid fishes).

Most writers are in agreement as to the application of the

terms epihyal and ceratohyal, but Allis has adopted an unusual

nomenclature which appears to have very little to recommend it

beyond the fact that it serves to locate the main jointing of the

hyoid arch between the epi- and cerato-elements, as occurs in the

branchial arches; he regards the epihyal as a part of the cerato-

hyal (which is, according to his view, a double ossification), and

ealls the interhyal the epihyal (Journ. Morph. xii. 3, 1897).

The glossohyal or basihyal varies greatly in size, being largest

in Hyodon, and extremely reduced in Hngrauhs and Coilia.

There is no separate glossohyal in the Mormyridx; it is either

wanting, or is fused with the first basibranchial. An endosteal

glossohyal is frequently present in addition to the ectosteal and

usually dentigerous bone (e. g., Albula, Chirocentrus, Megalops),

but in such forms as Heterotis, Osteoglossum, Chatoéssus, Clupea,

and Alepocephalus the cartilage remains unossified.

The dentigerous plate which covers the first, second, and third

basibranchials is readily removable in Arapaima, but in most

cases it is fused with the second, and overlaps the hinder part of

the first basibranchial and the anterior part of the third basi-

branchial (e. g., Clupea, Chirocentrus, Engraulis). In Hyodon ib

is fused with the third as well as with the second basibranchial,

and in Albula it is fused with all three basibranchials, and over-

laps, but is not fused with, the posterior half of the glossohyal.

Tt is much reduced in size in Chatoéssus, in which it is edentulous ;

and in Gonorhynchus it is confined to the second basibranchial.

In Dussumieria each of the three basibranchials has its own

dentigerous investing bone. There is in most cases also a much

smaller dentigerous bone covering the plate of cartilage that

represents the fourth and fifth basibranchials.

The first basibranchial is unossified in Heterotis, Notopterus, and

Gonorhynchus. The second basibranchial is remarkably long in

Engraulis, and the parallelism of the first and second cerato-

branchials is much disturbed in consequence. In Chanos the

anterior ends of the fourth and fifth ceratobranchials are separated

by a narrow, elongated tract of cartilage; in Alepocephalus and

Chatoéssus the anterior ends of the third and fourth cerato-

1904. | OSTEOLOGY OF THE ELOPIDA AND ALBULIDA, 79

branchials are set close together, without the usual interval. The

fourth and fifth basibranchials are represented by an undivided

plate of cartilage, as is customary in Teleostean fishes. No trace

of ossification is to be detected in this plate, a fact which is of

interest In relation to the discovery by Swinnerton of separate

ossified fourth and fifth basibranchials in that aberrant and

specialised genus Cromeria (Zool. Jahrvb., Abth. f. Anat. xviii.

1903, p. 66).

The hypobranchials are much reduced in the Mormyroids, and,

as already pointed out, the second hypobranchials are confluent

with prominent tendon-bones. In Coilia and Kngraulis the

second hypobranchials are fused with the sides of the second

basibranchial, and in Chanos the third hypobranchials are fused

with the third basibranchial, but in each case the suture remains

visible.

The “spicular bone” is generally taken to represent the

modified first pharyngobranchial. In Hyodon the first pharyngo-

branchial bone is large, and is certainly upturned and rod-like,

resembling a spicular bone; but in all the forms examined which

have a true spicular bone this arises from the upper surface of the

first epibranchial, and not from its anterior end as it does in

Hyodon. In Chirocentrus the first pharyngobranchial is a conical

bone projecting forward and upward. In Albulait is a bone which

runs forward with its axis in a line with that of the first epi-

branchial, and is clearly serially homologous with the second and

third pharyngobranchials. In Mormyrus and Mormyrops, also,

there is a small ossified first pharyngobranchial, but as a rule this

element of the visceral skeleton is cartilaginous.

The above-mentioned genera appear to have no true spicular

bone, but in Hngraulis, Clupea, Chatoéssus, and Chanos there is a

cartilaginous first pharyngobranchial and a true spicular bone im

addition, while in Klops, Megalops, and Alepocephalus there

is a well-defined and fully ossified first pharyngobranchial as

well as a slender spicular bone. The simultaneous presence of a

spicular bone and an unmistakable first pharyngobranchial bone

effectually disposes of the argument that the spicular bone repre-

sents a modified first pharyngobranchial. The spicular bone has

the value of an ossified ligament for the attachment of the

branchial skeleton to the side of the pro-otic, and is not a primary

constituent of the visceral skeleton. The confusion which has

arisen with respect to it, is probably due to the fact that in some

cases—it is not yet possible to say whether they are many or few—

the first pharyngobranchial takes upon itself the suspensory

function of the real spicular bone. The long bone which in the

Berycide and Cyttidz is attached to the anterior end of the first

epibranchial appears to be a true first pharyngobranchial, but

further observations upon the relations and the development of

this bone are much to be desired.

It is frequently difficult to say whether a bone of the branchial

skeleton is really toothed or not, since in cases where teeth are

80 _ DR. W. G. RIDEWOOD ON THE CRANIAL [ May 3,

present they may be fused to the bone, or may strip off with the

mucous membrane, and leave no scar upon the surface of the

bone. The matter is of some little importance because the mucous

membrane is customarily allowed to dry on the pharyngeal

skeleton to prevent the constituent parts from separating, and

loose teeth thus appear as if they were rigidly attached. The

dentigerous plates on the fifth ceratobranchialare readily removable

in Hlops and Megalops, which is rather remarkable, for the teeth

of the fifth ceratobranchial are usually firmly fixed to the bone

even in those cases in which there is considerable reduction in the

hyobranchial dentition as a whole. It is worthy of remark in

connection with the probable origin of the ectosteal constituent of

the glossohyal by the coalescence of tooth-bases (cf. vomer, &c.,

p- 63), that in the Herring the glossohyal is a cartilage overlaid

by the ectosteal lamina; and the teeth, although they may leave

scars on the bone when the mucous membrane is stripped off, are

not intimately attached to the bone. This evidently indicates a

process of degeneration in the lingual dentition, the first stages in

the transition to an edentulous state being marked by a reduction

of the basal parts of the teeth.

The dentigerous plates lying on the pharyngeal surface of the

second, third, and fourth pharyngobranchials are not collected

together to form an epipharyngeal apparatus as, for instance, in

the Cod, but they remain distinct. The cartilage of the fourth

pharyngobranchial remains unossified, but in some forms, such as

Albula and Chirocentrus, the cartilage in drying shrinks upon the

underlying dentigerous plate in such a way as to give the

impression that itself is ossified. Im Chatoéssus the ectosteal bone,

here toothless, spreads over the mesial and dorsal surfaces of the

cartilage, and gives the effect of a completely ossified fourth

pharyngobranchial. Swinnerton, it may be mentioned, has re-

corded the presence of a truly ossified fourth pharyngobranchial

in Cromeria (Zool. Jahrb., Abth. f. Anat. xvi. 1903, p. 65,

figs. H & K).

Engraulis, Gonorhynchus, and Alepocephalus have a distinct

fifth epibranchial cartilage. The presence of this element in

Alepocephalus was pointed out by Gegenbaur, who has also figured

one in Clupea alosa (Morph. Jahrb. iv. Suppl. p. 24, and pl. 2.

fig. 13, Alosa vulgaris or Clupea vulgaris). In Clupea harengus

and C. finéa the position of the fifth epibranchial is occupied by a

ligament ; in most genera there is no representative of this element

of the visceral skeleton.

The epibranchial accessory organ of respiration produces, in

those Malacopterygians in which it is at all largely developed,

important modifications in the shape and size of the elements of

the fourth and fifth branchial arches. In Chatoéssus, and to a

lesser extent in Chanos, the fifth ceratobranchial and the fourth

epibranchial are considerably increased in width, but in Gono-

rhynchus these bones are only lengthened. For our knowledge of

the structure of the epibranchial organ of Clupeoid fishes we are

1904.] OSTEOLOGY OF THE ELOPIDH AND ALBULID&.

mainly indebted to Hyrtl (Denkschr. Akad. Wiss. Wien, x. |

pp. 47-57, and xxi. 1863, pp. 1-10).

The epibranchial organ is by no means confined to these

but occurs, in one form or another, in MHyodon, in Het

(Hyrtl, Denkschr. Akad. Wiss. Wien, viii. 1854, pp. 73-8

certain Characinoid fishes (Hyrtl, Denkschr. Akad. Wiss.

xxi. 1863; Kner, Verh. zool.-bot. Ges. Wien, xi. 1861, pp.

192; Sagemehl, Morph. Jahrb. x. 1885, p. 114), in Osphrom

Anabas, &e. (Cuv. et Val., Hist. Nat. Poiss. vil. 1831, p. 328

pl. 206; Peters, Miill. Arch. Anat. Phys. 1853, pp. 427-430),

probably in several other forms not related to the above-nai

genera. In the last instances given, the whole of the epibrane

and pharyngobranchial skeleton is modified, while the fifth cet

branchials remain unaltered.

Abbreviations employed in the Figures.

al, alisphenoid. mpée, metapterygoid.

an, angular. max, maxilla,

bb, dentigerous plate covering n, nasal.

the basibranchials. op, opisthotic.

bo, basioccipital. ope, opercular.

br, branchiostegal rays. or, orbitosphenoid.

bs, basisphenoid. p, parietal.

cb, ceratobranchial. pb, pharyngobranchial.

ch, ceratohyal. pl, palatine.

cor, circumorbital bones. pm, premaxilla.

ct, cartilage. pof, postfrontal.

d, dentary. pop, preopercular,

eb, epibranchial. prf, prefrontal.

ecar, ectosteal articular. pro, pro-otic.

ecp, ectopterygoid. ps, parasphenoid.

eh, epihyal. pet, post-temporal.

enar, endosteal articular. ptf, posterior temporal fossa.

enp, entopterygoid. q, quadrate.

eo, exoccipital. sar, sesamoid articular.

ep, eplotic. sm, surmaxilla.

SJ; frontal. soe, supraoccipital.

gh, glossohyal: ~ sop, subopercular.

hb, hypobranchial. sp, spicular bone.

hh, hypohyal. sq, squamosal.

hm, hyomandibular. st, supratem poral.

th, wnterhyal. stf, subtemporal fossa.

iop, teropercular. sy, symplectic.

me, mesethmoid. v, vomer.

Proc. Zoo. Soc.—1904, Von. IT. No, VI. 6

ON THE PORES IN CHLAMYDOSAURUS KINGII. [May 17,

May 17, 1904.

Howarp Saunpers, Esq., F.L.S., Vice-President,

in the Chair.

m The Secretary read the following report on the additions to the

eee Besccicty’s Menagerie during the month of April 1904 :—

ue q The number ae register ad additions to the Society’s Menagerie

during the month ‘of April was 111, of which 46 were by

_ presentation and 20 by purchase, 20 were born in the Gardens,

and 25 were received on deposit. The number of departures

_ during the same period, by death and removals, was 96. _

; Amongst the additions attention may be called to :—

- 1. A fine specimen of the Boatbill (Canchroma cochlearia),

purchased on April Ist.

_ 2. Two Cheetahs (Cynelurus jubatus), from the Soudan,

_ presented by Col. B. Mahon, C.B., F.Z.8., on April 18th.

3. Two Keas (Nestor notabilis), presented by Mr. T. E. Doune,

on April 26th.

Dr. W. T. Calman, F.Z.8., exhibited a specimen of a blind

crustacean (Munidopsis polymorpha), from a subterranean lake in

the Island of Lanzarote, Canaries, collected by Mr. Fairfax

Prevost. The species was described in 1892 by Dr. Koelbel, of

Vienna, from specimens collected by Prof. Simony, but appears

to have been overlooked by recent writers on the group to which

it belongs. The lake in which it is found is salt and communicates

with the sea, the water rising and falling with the tide. All the

other species of the genus, over 100 in number, are inhabitants

of the deep sea, none being recorded from a depth of less than

about 100 fathoms.

Mr. F. E. Beddard, F.R.S., exhibited the body of an example

of the Lizard Chlamydosaurus kingiti which had died in the

Menagerie some years ago, and made the following remarks upon

the absence of femoral pores :—

Mr. Boulenger has pointed out (Cat. Lizards Brit. Mus.) that

while the Australian Agamidee with one exception possess femoral

pores, these structures, are as a rule, wanting in that family.

Among those in which the femoral pores are described, or at

least asserted to be present and in definite numbers, is the genus

Chlamydosaurus. The original describer of the genus, and of the

species, Dr. Gray (in King’s ‘Survey of Australia,’ vol. i1. p. 424

et seq.), however, observed that it was “‘ destitute of femoral pores.”

Duméril and Bibron (‘ Erpétologie générale,’ p. 440) make use of

the phrase “ Des pores fémoraux” as part of their definition of

the genus. As this absolute contradiction occurs, I have thought

it worth while to exhibit to the Society a male example in which

there are certainly no femoral pores fully comparable to those of

PZ.S.1904,vol IL Pl. IIT.

{

C.Crossland, del. Huth,sc.et imp.

lL NOTODORIS MINOR. 2.TREVELYANA COCCINEA.

3.7. CEYLONICA. 47 CROCEA.

PZ.5.1904,vol I. Pl. IV.

C.Crossiand, del Huth, sc. et imp

1. TREVELYANA BICOLOR. 2 NEMBROTHA CRISTATA.

5.N. AFFINIS. 4 MARIONIA LEVIS. 5 TEETH OF (2) BORNELLA DIGITATA

(b)B. EXCEPTA AND (c) B. SIMPLEX.

1904. ] ON NUDIBRANCHS FROM EAST AFRICA AND ZANZIBAR. 83

such other Agamids as, for example, Amphibolurus barbatus (also

exhibited). If the specimen exhibited by me be not a mere

variation, I can understand how it is that the presence of femoral

pores has been asserted by some and denied by Gray. For there

are along the thigh a few larger scales which might lead to the

inference that they were perforated scales. They are not so,

however; and when the skin is reflected no glands can be seen ;

these always accompany the femoral pores, which are, of course,

the external apertures of their ducts.

Mr. Oldfield Thomas, F.R.S., F.Z.S., exhibited, on behalf of the

President, a sketch by a Chinese artist of a hind and fawn of

Pére David’s Deer (Hlaphurus davidianus) from Hainan. Unfor-

tunately the artist had added the antlers of a Peking Stag (Cervus

hortulorum). The sketch was taken from specimens presented to

Mr. E. T. C. Werner, H.B.M. Consul at Hainan. It served to

prove that Pére David’s Deer still survived in Hainan. Whether,

however, it was indigenous or imported, remained to be deter-

mined. Mr, Lydekker was in communication with Mr. Werner on

the subject,

The following papers were read :—

1. On some Nudibranchs from East Africa and Zanzibar.

Part V.* By Sir C. Extot, K.C.M.G., late H.M. Com-

missioner for the Hast African Protectorate, F.Z.S.

[Received March 10, 1904. ]

(Plates IIT. & IV. +)

In my last two papers I treated of the Doridide Crypto-

branchiate as a group, but no systematic importance is to be

attached to the order in which the species now to be described are

arranged.

PTERAEOLIDIA SEMPERI.—Since writing my description of this

species in my second paper (P. Z.S. March 17, 1903, p. 255), I have

read Prof. Bergh’s account cf Vossis, characterised by a lateral ridge

similar to that found in some of my specimens (‘ Opisthobranchiata

of Danish Expedition to Siam,’ 1899-1900, p. 52), and accordingly

carefully re-examined them to see if they should not be referred

to this new genus. It appears that they should not. The radula

is uniseriate, consistently of 18 teeth, and the same in the

specimens which have and those which have not the lateral ridge.

It therefore seems clear that the ridge is found in the genera

where the radula is uniseriate as well as in those where it is

triseriate, and, further, that in alcoholic specimens, at any rate, it

may be present or absent in the same species.

* For Part IV. see P. Z. S. 1904, vol. i. p. 380.

+ For explanation of the Plates, see p. 105.

De.

84 . SIR C. ELIOT ON NUDIBRANCHS [May 17,

Nortoporis Bergh.

[Bergh, “‘ Neue Nacktschnecken d. Siidsee,” p. 111, in Jour.

d. Mus. Godeffroy, viii. 1875; Eliot, Nudibranchiata in J. 8S.

Gardiner’s Fauna and Geography of the Maldive and Laccadive

Archipelagoes, vol. 11. part 1.|

This genus, which is recorded from three parts of the Indo-

Pacific, seems allied to Mgires and the little-known Triopella,

with which it forms a small group of phanerobranchiate Dorids

characterised by a hard texture, valves or other appendages pro-

tecting the gills, and undifferentiated teeth. Both Mgires and

Notodoris have simple unperfoliate rhinophores.

The body of Wotodoris is hard and rough, often marked with

prominent ridges. The frontal veil is large. Tbe branchie, and

sometimes the rhinophores, are protected by valves. There is no

labial armature, and the teeth are hamate with indications of an

accessory denticle. Three species have been described, each from

a single specimen—J. citrina B., WV. gardineri Eliot, and the

present V. minor. They are all yellow, differing chiefly in size,

shape, and the form of the branchial valves. It is just possible

that V. minor may be a young and undeveloped form. It is

smaller than the others, and superficially resembles a Phyllidia.

Tt has no distinct tail, no rhinophore valves, and no longitudinal

ridges. The branchial valve is three-lobed and not much sub-

divided. Possibly the gill is constructed differently from those

of other species. Both JV. citrina and gardineri have rhinophorial

valves and a body tapering off into a tail: the former has a single

dorsal ridge running from the rhinophores to the branchial valve,

which is eight-lobed: the latter has four dorsal ridges and a

branchial valve three-lobed, with elaborate subdivisions.

Noroporis MINOR, sp. n. (Plate III. figs. 1 a—1 g.)

One specimen from Chuaka, east coast of Zanzibar.

The living animal was 13 millimetres long, 5 broad and 4 high.

It was light lemon-yellow in colour, with sharply-marked trans-

verse black lines. The flat sole occupied the whole ventral

surface. The back was not quite smooth, the yellow parts being

really low broad lumps between black depressions. The whole

body was very stiff and rigid, superficially resembling a Phyllidia.

The animal was never seen to move.

In the preserved specimen the yellow has become whitish, but

otherwise the shape and markings of the living animal are pre-

served. The integuments are very hard and full of spicules.

There is no trace of any mantle-edge, and the body slopes straight

down to the sides of the foot. Over the mouth-parts is a strong

rounded frontal veil (figs. 1a & 1e), also descending right down to

the sides of the foot, and extending laterally about as far as the

rhinophores. At the beginning of the posterior third of the body

are the three gill-valves (figs. 1a@—lc). They are not noticeable

except in profile, as they lie rather flat, and are not much sub-

divided. Beneath them lie the gills (fig. 1d), which appear to

1904. | FROM EAST AFRICA AND ZANZIBAR. 85

consist of about 27 small tufts, pinnate, bipinnate, or tripinnate

according to their size, and spread over three areas corresponding

to the valves. Possibly each area represents a separate axis, and

the gills should be described as three tripinnate or quadripinnate

plumes. But this arrangement cannot be demonstrated with

certainty in the preserved specimen, and the living animal never

raised the valves at all. The rhinophores are thick, conical, and

without a trace of perfoliations; they are retracted into simple

holes, provided with neither valves nor raised edges. No oral

tentacles and no groove in the anterior margin of the foot could

be discovered (fig. 1 e).

There is no trace of armature on the labial cuticle. The radula

consists of 33 rows, the largest of which contain about 25 teeth

on each side of the rhachis. The teeth are transparent and

crowded: the innermost are smaller and close over the rhachis;

the outermost are longer and show no trace of irregularity. The

shape of all is much the same, hamate with a rudimentary denticle

under the tip of the hook. They much resemble the teeth of

Notodoris citrina (Bergh, l. c. pl. ix. figs. 39, 40), but are somewhat

more erect and hardly ever show indications of more than one

denticle (fig. 1g). The glans penis spreads out somewhat as in

Nembrotha, and appears to be trifid. The lower part is armed

with a thick mass of minute blunt spines (fig. 1/).

TREVELYANA Kelaart.

[Kelaart, in Ann. Mag. Nat. Hist. 3rd ser. vol. 1. p. 257, 1858;

Bergh, in Semper’s Reisen, Heft xi. p. 441, & xvi. 2, p. 850.]

This genus is recorded only from the Indo-Pacific, where it

seems to be the commonest representative of the Polycerade,

being frequent under stones between tides. The animals are

limaciform, but some specimens at any rate show indications

of a division between the back and sides. The body is smooth,

bears no appendages, and is usually of a light bright colour

varying from red to white. The branchie are rarely less than

ten, often numerous, and generally small. There is no labial

armature or central tooth. The radula is fairly wide, and com-

posed of hamate or awl-shaped teeth, which are often wregular.

The hermaphrodite gland, instead of being spread over the liver,

is collected into two globular masses.

Several of the species, e. g. the 7’. ceylonica and 7’. bicolor given

below, are very imperfectly described by the original authorities,

and hence identification is uncertain. It is clear that the whitish

forms with yellow lines and spots show considerable variety, but

it is hard to say how many of these varieties are specific,

TREVELYANA COCCINEA, sp.n. (Plate III. figs. 2 a-2f.)

One specimen, dredged between Shimoni and Wasin at 6-8

fathoms.

The notes on the living animal describe it as the largest species

86° SIR C. ELIOT ON NUDIBRANCHS | May 17,

of Trevelyana yet found in East Africa, 3 inches long, and stout

in proportion. The colour was bright vermilion, plentifully be-

sprinkled with slightly projecting spots of a deeper shade. The

rhinophores and gills were small and deep vermilion in colour.

The preserved specimen has greatly shrunk, and is 25 millimetres

long, 14 high, and 11 broad. The colour is dirty white, and no

spots or tubercles are visible. There is no trace of tentacles or of

a mantle-edge, but the frontal veil is a distinct hard ridge. The

foot is grooved in front. The tail is very short. There are

12 small gills set in a circle, bipimnate and in parts tripinnate.

The vent is subcentral and not raised.

Though there is nothing that can be called a labial armature,

the labial cuticle is strengthened with scattered rods of various

shapes. The radula is larger than usual in the genus. It con-

sists of 36 rows, some of which contain as many as 51 teeth,

so that the formula is 36 x 50+1.0.1-+50, but the rows towards

the front are much smaller. The first lateral is large and hamate

(fig. 2a), sometimes with irregular notches or denticles on the out-

side of the hook (figs. 2c & 2d). In several cases the top seemed

to be broken off, and the remaining part was bifid or trifid

(fig. 26). The other teeth are slender and hamate (figs. 2e & 2/).

In all the teeth the hook is directed forwards, not backwards.

The liver is greyish and not very large. In front of the liver,

but quite separate from it and from one another, lie two large

spherical hermaphrodite glands with a diameter of about 5 and

7 millimetres respectively. They are yellowish in colour, and

the surface is covered with knob-like follicles. The verge is armed

with transparent spines. The large pericardium lies in front of

the branchiz, and in the alcoholic specimen is much inflated.

This form is possibly the Stenodoris rubra of Pease (Am.

Journal of Conch. ii. 1866, p. 206), though, if so, “light red

papille ” is a strange description of the raised spots; but the

account given of the animal is not sufficient to admit of

identification.

TREVELYANA CEYLONICA Kel. (Plate III. figs. 3 a—3 c.)

[Kelaart, Ann. & Mag. of Nat. Hist. 3rd ser. vol. i. p. 257,

1858. |

One specimen from the East Coast of Zanzibar.

The notes on the living animal describe it as about an inch long,

creamy white, with bright red dots. The gills were yellow, with

bright red lines down their backs; larger and more feathery than

is usual in the genus. There was a line of bright red round the

edge of the foot.

The preserved specimen is colourless, 15 millimetres long and

6°5 broad. The back is quite smooth, and there is no sign of a

mantle-rim. The pericardium forms a large, much swollen

prominence. The rhinophores are completely retracted. There

are 12 branchiz set in a circle open behind; one is large and

bifid, one is rudimentary. The foot is deeply grooved in front.

No tentacles could be discerned.

1904. | FROM EAST AFRICA AND ZANZIBAR. 87

The buccal mass is rather large, the radula fragile, with a wide

rhachis. There are 21 rows in all, some of the longest of which

contain 24 teeth on each side. The first tooth (fig. 3a) is larger

than the rest, and projects into the rhachis; it is slightly bent, but

hardly hamate. All the first teeth are similar and regular in shape.

The base is somewhat wavy and as if hollowed out. The other

teeth are awl-shaped, with an irregular and somewhat bifid base

(fig. 3c). The liver is yellowish grey and not very large. In front

of it are two hermaphrodite glands, much as in 7’. coccinea, but

smaller, The verge is armed with numerous short thorns of very

varying shape.

I think this animal is probably Kelaart’s 7. ceylonica, for

which the genus was founded, and which appears not to have

been described since; but it is difficult to be certain of the identi-

fication, as he gives no information respecting the radula. The

form and colour agree well, including the red lines on the branchize

and round the foot. ‘The chief difference is that whereas his

specimen has 15-16 pure white branchie “set round a large

disk,” mine had 12 yellow branchie set in a circle open behind.

But his specimen was nearly twice the size of mine, and probably

the larger individuals develop extra plumes which close up the

posterior gap. On the other hand, both specimens agreed in

having rather large feathery branchiz, an unusual character in

the genus. Kelaart says “they resemble a small tuft of marabout

feathers.”

TREVELYANA CROCEA B. (Plate III. fig 4.)

[ Bergh, in Semper’s Reisen, xvi. 2, p. 850, figs. |

More than 100 specimens from the Hast and West Coasts of

Zanzibar, where it is one of the commonest littoral molluses at

certain seasons.

Mr. Crossland, who collected them, informs me that this form

provided a most striking case of the migration of molluscs in

flocks to shallow water for the deposition of spawn.

But few specimens were collected before a certain period of a

few days’ duration, when the sand of Chuaka Baw just below low-

tide mark was occupied by astonishing numbers of these delicate

little nudibranchs. These were not washed up by accident, but

were all actively crawling on the sand among the weeds kc.

Many were i coitw, and when placed in basins of sea-water

most of the specimens were soon engaged in copulation or the

deposition of yellow egg-ribbons. By-and-by the swarm dis-

appeared to some unknown permanent habitat. If this were in

the deeper channels of the bay (1 to 2 fathoms deep at low tide)

they must have been found there by dredging. As this was not

the case, it seems most probable that the migrations of these tiny

animals extend to and from the deep sea three or more miles

away. An almost equally conspicuous swarm was formed by

individuals of Melibe fimbriata, and other species (e. g. Ceratosoma

cornigerum, Chromodoris spp., and Pleurobranchus delicatus)

appeared occasionally for a few days in considerable though

88 “SIR C, ELIOT ON NUDIBRANCHS [May 17,

smaller numbers, being rare or completely absent from the shore

of the Bay at other times.

Most of the animals were of a bright dark yellow with the black

liver showing more or less conspicuously through the transparent

integuments, but the colour ranges in exceptional cases from deep

orange to almost colourless transparency. Many specimens were

infested with small light yellow copepoda found adhering to the

body, especially on and near the gills.

The alcoholic specimens are of a more or less yellowish white.

The largest is 29 millimetres long, 12 high, and 8 broad, but, as a

rule, the back is proportionately broader. The whole body is smooth

and very soft. In most specimens the dorsal area is bounded by a

distinct lateral ridge. It is not visible behind the branchiz, but

extends from them to the front of the head, where, however, it is

not continuous but divided by a deep notch in the middle. In

several specimens this ridge is only clear in places and in a few

it is absent altogether. The rhinophores bear about ten perfolia-

tions and are set in such shallow pits that they can hardly be

called retractile. They are exposed in the alcoholic specimens.

The edges of the pits are smooth. The gill consists of from 20

to 34 leaflets *, set in a horseshoe or circle open behind, and placed

rather far back. The leaflets are flat and compressed and

decrease in size posteriorly. The largest bear on each side about

ten lamelle, the smallest two or three. The whole appearance of

the branchial apparatus is quite unlike what is usual in the

circum-anal plumes of nudibranchs and recalls the prosobranch

gill. The foot is a narrow groove, but has a thin expanded

margin, including which the breadth is 6 mm. in large specimens.

The anterior margin of the foot is grooved and united with the

corners of the mouth, where it is joined by a second ridge, which

runs above it and apparently represents the tentacles. The tail

is bifid.

The radula has a wide bare rhachis, and the formula varies from

about 11 x 104 2.0.24+10 to 15x 14+2.0.24+14. The innermost

tooth is irregular in shape, but consists of a basal portion from

one end of which rises a more or less bent spine, while another

spine is more or less completely developed at the other end. The

second tooth is larger and is more distinctly bicuspid. The other

teeth are unicuspid, awl-like, and hardly bent; and those nearer

the rhachis are rather stout, but they become slender towards

the end of the row. All the different forms of teeth are well

represented in Bergh’s plates. In the nervous system the ganglia

are very distinct. - The liver is large, black, and very soft. On its

anterior portion, and less detached from it than usual in the genus

(e. g., than in 7’, coccinea described above), are two yellowish

hermaphrodite glands of a somewhat irregular shape. Indeed,

though separable from the liver, they cannot be said to be separate

from it. This may be possibly due to the fact that the specimens

are in good condition, so that the membranes connecting the

* The gills as represented in the Plate are not sufficiently numerous.

1904.] FROM EAST AFRICA AND ZANZIBAR. 89°

various organs are fresh and strong, whereas in other cases they

may have dried up or decayed. The verge is armed with numerous

small spines of very variable shape, simple, bifid and trifid. From

the genital mass to the tail extends on each side a long, ramified,

almost arborescent gland, distinctly visible through the trans-

parent body-wall with which it is united.

TREVELYANA BICOLOR (?). (Plate IV. figs. 1 a—-1c.)

[A. & H., Notes on a Coll. of Nud. Moll. made in India, p. 132,

pl. xxix. figs. 11, 12.]

The single specimen, which was captured at Prison Island,

Zanzibar, was 20 millimetres long, with a very long narrow foot,

tapering to a point posteriorly. The whole animal was white,

with projecting spots of bright yellow. The tips of the rhino-

phores and edges of the gills were also bright yellow. The liver

showed through the dorsal integuments as a black mass before

and behind the branchig, and in front of it were seen the yellow

reproductive organs. The branchie were simple and leaf-like and

shrunk together when touched.

The preserved specimen is contracted into a spherical shape,

showing no trace of the raised spots or of a mantle-margin.

The head-parts are much retracted and distorted, but the anterior

margin of the foot seems to have been deeply grooved. The

colour is white, but the black liver is still conspicuous. The

twelve branchiz are set in a complete circle.

The radula consists of 26 rows, the widest of which contain

24 closely packed teeth. The first lateral is large and hamate and

the next much like it. The other teeth are rather stout, of the

bradawl shape or slightly curved. In the pharynx were found the

remains of a small tectibranch, which, to judge from its radula

and stomach-plates, was probably Atys.

I think this form is probably A. & H.’s 7. bicolor. Their

description was made from the drawing which they reproduce

and they saw no specimen. The bicoloration there depicted was

probably due to the liver being seen through the integuments, for

though the picture certainly suggests a black patch on the skin,

it will be seen that this patch occupies exactly the position of the

liver, and that it bears yellow spots like the white part. It is

also possible that Riippell and Leuckart’s 7’. impudica is identical

with this form. They describe it (Neue wirbellose Thiere des

rothen Meers, p. 33) “corpore dilute lacteo; tentaculis superi-

oribus, maculis ocellisque dorsalibus, branchiis pedisque limbo

aurantiacis ; dorso tuberculato; branchiis 12 medium dorsi versus

sitis ; pallio indistincto.”

NemsBrotHa B.

[Bergh, 8. R. xi. p. 450, figs., xvii. p. 980, figs.; id. Beitr.

zu einer Monographie der Polyceraden, ii. p. 658, figs., and iii.

pp. 164-5. ]

This genus is allied to Zrevelyana, but both internally and

externally can be readily distinguished from it. The coloration is

90> SIR C, ELIOT ON NUDIBRANCHS _ [May 17,

generally rather sombre but gorgeous, a prevalent tint being very

dark green or blue with brilliant lighter markings. The gills are

few (3-5), but large and strong. The hermaphrodite gland is as

usual, and not collected into globules. A very narrow labial

armature is present In some species, but usually there is none.

The radula is never very wide and sometimes is very narrow,

consisting of a median plate with from three to twelve laterals, of

which the first is large and hamate and the rest plate-like. The

species are not all equally well known, all our information as to

NV. morosa and cristata coming hitherto from drawings by Semper.

It would appear, however, that some of my specimens should be

referred to the latter species. WV. nigerrima, kubaryana, and

cristata have a fairly broad radula, with about twelve laterals,

and are distinguished by their dark coloration. They are

evidently closely related, and may prove to be merely varieties,

including WV. morosa. JN. gracilis, diaphana, gratiosa, and affinis

are lighter in colour, and have a narrow radula with only three or

four laterals.

Nembrotha is recorded from the Indo-Pacific and West Coast of

Mexico. It is fairly common onthe Hast Coast of Africa.

NemBrorua cristata B. [?=V. nigerrima, var.] (Plate IV.

fig. 2.)

[ Bergh, 8. R. xi. p. 458, pl. xxxiii. fig. 6.]

Three specimens from the East Coast of Zanzibar. The living

animals are described as having a sloping back, long tail, and

narrow foot, somewhat like Ceratosoma. ‘The texture was soft,

and the colour a very dark but brilliant green with black spots,

and also narrow stripes of brighter and lighter green. The gills

were counted as five, and the rhinophore-pockets were raised.

The measurements of the largest alcoholic specimen are : length

54 millimetres, breadth 15, height of body 13, height of branchiz

above body 8. The texture has become hard and wrinkled, the

animals having evidently been strongly contracted. The main

stem of the gills is very thick, strong, and muscular, so that it

almost forms a valve to protect the pinne as in otodoris. The

anterior plume is distinct and separate, but the lateral pairs are

almost confluent, and it is consequently hard to say where one

begins and the other ends, or whether the total. number of

branchiz should be reckoned as three, four, or five. The rhino-

phores are not very large and are completely retracted within

smooth projecting sheaths about 2°5 mm. high. The foot is

narrow. ‘The relations of the external mouth-parts are much

obscured and distorted by the strong contraction which has

affected the whole anterior portion of the body, but it appears

probable that the foot is grooved and notched with the upper

lamina attached to the corners of the mouth, and that the

tentacles are horizontal ridges. There is a very narrow labial

armature, about half a millimetre wide and hardly visible to the

naked eye. Itappears to form a complete ring, and is composed

1904. ] FROM EASY AFRICA AND ZANZIBAR. 91

of a loose mass of long, yellow, transparent rods, irregular in shape

and often bent.

Tn the two specimens dissected, the radula consists of 30 and

31 rows respectively, and the formula of each row is, as a rule,

10+1.1.14+10 or more rarely 114+1.1.1+11. The median tooth

is squarish, not very broad, and bears, as a rule, five denticles on

the anterior edge, but sometimes only four, while in one specimen

there were only three denticles in the hinder rows. The first

lateral tooth is large and sickle-shaped. The corner of the basal

part projects over the rhachidian tooth and creates a false

impression that it is an accessory denticle. The remaining teeth

are generally ten, but sometimes an additional rudimentary one at

the end of the row raises the number to eleven. They are little

more than flat squarish plates, decreasing in size outwards. Only

the first of them shows some traces of a hamate shape.

Allthe internal organs are of a deep black colour, which rendered

their examination difficult. The blood-gland is large. I was not

able to make any satisfactory preparations of the reproductive

organs, but the glans seemed to be armed with a dense mass of

curved rods.

IT think this form must be identified with JV. cristata B., of

which, however, no specimen has been described, all that is known

of it being Semper’s drawing and apparently a few notes. But

it is also not improbable that it is a variety of WV. nigerrima B.,

from which it differs externally in little except the absence of

any red coloration. The number of branchie is, as explained,

uncertain, but the arrangement shown in Bergh’s plate of

NV. nigerrima is certainly not that of these specimens. On the

other hand, the presence of the narrow labial armature is an

argument for identity.

NEMBROTHA CHRULEA, sp. 0.

Four specimens from Sii Island, near Vanga. No notes on the

living animal, except that it was blue and had apparently no red

or green mottlings.

The colour of the freshly-preserved specimens was a fine bright

indigo, varying in intensity in different parts. One of the

specimens was much lighter than the others and also smaller.

The whole of its body and the lighter parts of the other indi-

viduals were marked with deep indigo spots.

The largest preserved specimen is 43°) millimetres long,

18 high, and 12°5 broad. The space from the head to the

branchie is 12 mm. and from the branchie to the end of the tail

22 mm., but the tail is longer in this specimen than in the others.

The shape is somewhat like that of Ceratosoma without lobes, as

the back rises considerably from the head to the branchie. The

integuments are leathery and not at all transparent. The surface

is quite smooth, and there is no indication of a mantle-edge. The

rim of the rhinophore-pockets is only slightly raised. The rhino-

phores themselves are large, entirely retracted, with 25-30 deep

92 SIR C. ELIOT ON NUDIBRANCHS [May 17,

perfoliations. As in the last species, the gills are very thick,

strong, and muscular, apparently five, but in this case, too, the

lateral pairs sometimes coalesce, so that the whole number may be

counted as three or four. They are bipinnate. The oral tentacles

appear as large, distinct tubercles on each side of the mouth, and

were doubtless fairly long in life. The foot is rather broad, with

a Shallow groove in front; the upper lamina is connected with the

sides of the mouth under the tentacles.

The internal organs are mostly of a greyish yellow, not deep

black as in the last species. Though the labial cuticle contained

a few scattered yellowish rods, no connected armature is visible.

The radula much resembles that of JV. cristata, and has for

formula about 27 x 10+1.1.1+10 or occasionally 11+1.1.1+411,

but the median plate is broader, with five distinct denticulations

which do not vary in number. The first lateral has a groove near

the end of the hook, and the next two or three teeth have a rudi-

mentary hamate shape. The liver is large. The upper wall of

the pericardium is very thick and strong. The verge resembles

the figures in Bergh’s plates of Membrotha nigerrima, the glans

being armed with a profuse mass of hamate teeth. Those on the

top seemed rather larger and more curved than in his figures.

This species is closely allied to WV. nigerrima, but appears to

be sufficiently distinguished by (a) its coloration, (6) the only

slightly projecting edges of the rhinophore-pockets, (c) the absence

of a labial armature, (d) slight differences in the radula, (e) another

form of tentacles.

NEMBROTHA AFFINIS, sp. n. (Plate LV. figs. 3 a—-3 d.)

[Cf. W. gratiosa Bergh, Nudibr. of ‘Blake’ Expedition, pp. 172-

175.] .

One specimen caught in a trawl in Chuaka Bay on the Hast

Coast of Zanzibar. Very long and narrow, being 5 centimetres

in length and 1 in height.

The living animal was extremely soft, dull violet-black in

colour, with dull yellow stripes on the sides and somewhat brighter

ones of the same colour on the back. The stems and bases of the

gills were light green, and the same colour occurred between the

rhinophores and round the edges of their pockets. The pinne

of the gills looked black, but when seen by transmitted light were

of a fine purple. The foot was very narrow, and the animal could

not adhere strongly to anything.

The alcoholic specimen is flabby, 28 millimetres long, 5 broad,

and 10 high. As the result of this reduction in size, the yellow

parts look wider and the black parts narrower, so that the animal

appears to be yellow with black stripes, rather than black with

yellow stripes as in Mr. Crossland’s figure. No doubt, however,

the latter is correct ; it represents four lateral yellow stripes and

one medio-dorsal. The stripes are interrupted in places, particu-

larly on the tail, and there are some long yellow spots between

them. The branchie are distinctly only three in number, smaller

1904. } FROM EAST AFRICA AND ZANZIBAR. 93

than usual in the other species, but with a very thick rhachis and

bipinnate. The rhinophores are large and exserted, each bearing

about 35 perfoliations. The rims of the pockets are very slightly

raised. The oral tentacles are two hard black ridges, curved

downwards and sideways. The foot is narrow and grooved in

front.

The buccal mass was extracted, but the animal was not further

dissected in order to preserve the specimen. There is no labial

armature. The formula of the radula is 13x3+1.1.1+3. The

teeth closely resemble those of WV. gratiosa, the chief difference

being that the anterior margin of the wide median tooth (fig. 3 ¢)

is indistinctly bilobed, the right half being always a little higher

thar. the left. The first lateral (fig. 3d) is large, rather irregular

in shape, and with a double hook at the apex.

This form is closely allied to JV. gratiosa, and were the latter

found in the Indo-Pacific region, I should be inclined to regard

them as varieties of one species. But JV. gratiosa is recorded from

the West Coast of Mexico*, which lies outside of the Indo-

Pacific area; and it is therefore probable that the differences pre-

sented by the present animal are real and greater in living indi-

viduals.. (a) It is not mentioned that JV. gratiosa is remarkably

soft. (b) The present specimen shows no traces of ridges near

the rhinophores or on the tail. (c) The coloration of WV. gratiosa

is not dissimilar, but the pattern is spotted whereas here it is

striped. (d) The tentacles do not look as if they had ever been

ear-shaped. (e) The anterior margin of the median tooth is

indistinctly bilobed.

MARIONIA.

[See especially Bergh, in Semper’s Reisen, xv. p. 737, & xvii.

p- 890.]

All the Tritoniade which I have collected in East Africa belong

to this genus, unless the form described as Warionia sp. is regarded

as sufficiently certain to constitute a new generic type. Marionia

is distinguished from its near allies Z’ritonia and Candiella by the

presence of a circle of horny plates or leaves in the stomach. The

velum bears distinct processes, which are often ramified. The edge

of the jaw has one or more rows of denticles, and the radula is

moderately wide. The central tooth is broad and more or less

distinctly tricuspid. The laterals are hamate, but the first one is

larger and clumsier than the others. Provisionally I think it

best to divide the forms here described among six species, but am

by no means certain that they will all prove valid. When more

material can be examined it will probably be found that the

species of MJarionia exhibit many varieties in formand colour and

run one into another. It is also not impossible that the denticu-

* In Bergh’s ‘System der Nudibr. Gasteropoden,’ p. 1145, the locality is given as

“ mare indicum, Amboina,” but this appears to be a slip. ‘The animal is described by

Bergh among the molluscs of Amboina, but is expressly said to come from Mexico.

94 SIR C. ELIOT ON NUDIBRANCHS [May 17,

lation of the jaw varies with age. Of the six species, IZ. pellucida

seems distinct from the others, which are all nearly related to

M. arborescens. WM. levis is distinguished by being quite smooth

and not at all tuberculate. J. ramosa is closely allied to I, arbo-

rescens, and differs chiefly in having unusually large branchie and

appendages. WW. viridescens and albo-tuberculata differ from these

last two forms in having branched processes on the velum, and

are closely allied one to another in structure, though by no means

similar in external appearance.

It is noticeable that in none of these forms is the interior of the

buccal cavity black, and that most of them have only one fully-

developed row of denticles on the jaw.

MARIONIA PELLUCIDA, Sp. 0.

One specimen dredged in 10 fathoms near Wasin, East Africa.

The living animal showed very httle colour but for the pink

liver which shone through the transparent integuments. The back

was sparsely reticulated with vermilion, turning to deep crimson

near the bases of the branchiz, and also bore some opaque white

raised spots. ‘The sides of the body were white and the edge of

the velum sandy-coloured. The velum was not bifid, and bore 12

processes, of which 8 were 3-branched. The branchiz were 13, of

moderate size, directed backwards, The finer branches very

delicate and transparent.

The alcoholic specimen is yellow, with small tubercles of a

lighter colour on the back and sides. It is 15 millimetres long,

5 broad at most, and 4 high. The 13 branchie are rather far

apart from each other; none are large, and the first pair as well

as the last three are minute. The dorsal margin is not very

prominent. The rhinophores are large. The velum as described

above, but though the outermost processes probably represent the

tentacles, they do not seem to be grooved as usual. The long

narrow jaws bear three or four rows of denticles on the edge.

The radula is at most 22+1.1.14+22 x 25, but many of the rows

are much shorter. The central tooth is not very wide and tri-

cuspid, the side cusps being as high as that in the middle. The

stomach has a circular band of about 70 small yellowish plates, all

of much the same size and usual triangular shape.

MARIONIA LEVIS, sp. n. (Plate IV. fig. 4.)

Six specimens from Chuaka, East Coast of Zanzibar, and Wasin,

East Africa. Two were dissected.

The living animals were high and narrow in shape, with a flat

back. The sides were described as white, mottled with translucent

patches. The ground-colour of the back was a light purplish

brown, with stripes of the same colour but darker and others of

white. The branchize and rhinophores were pink with dark red

blotches.

An uninjured alcoholic specimen is 26 millimetres long, 10 high,

1904. ] FROM EAST AFRICA AND ZANZIBAR, 95

and 7 broad in the widest part, but one which was dissected was

about twice as large. The colour has become pale green, with a

white reticulation on the sides and white stripes on the back.

The skin is quite smooth, and there are no tubercles whatever.

There are nine or ten pairs of branchie, of which the last three

are quite small. The rhinophores have long raised sheaths with

simple edges; the club is surrounded by six bipinnate plumes.

The velum bears at each end a small grooved tentacle of the usual

shape and six processes. The two in the middle are simple and

smaller; the other four are larger and branched.

The jaws are white and membranous in the smaller and probably

immature specimen, yellow and corneous in the larger one. In

both there are from 20-30 very large blunt denticles, and also

undulations near the edge of the jaw, which in the larger specimen

sometimes develop into denticles, so that in about half the length

there are two rows of denticles and here and there three. The

radula consists in one specimen of 47 and in the other of 45 rows,

with a formula of about 804 1.1.14 80, which rises to as much as

85 marginals in one and 100 in the other for a few rows. The

central tooth is broad and tricuspid; the median cusp is taller

than the others, but not very pointed; all the cusps are rather

irregular in shape, and have indentations here and there on the

edges. The first lateral tooth is large, blunt, and very different

from the rest in appearance. The othersare hamate. The stomach

has a girdle of about 150 horny, yellow, triangular plates of

different sizes.

J do not think that this species can be identified with any of

the forms the descriptions of which I have seen*. The color-

ation somewhat resembles Tritonia rubra Leuckart and 7’.

hawaiensis Pease, but the other details do not coincide. The

species differs from the others hitherto found in East Africa in

being quite smooth and having no tubercles.

MARIONIA ARBORESCENS B.

[Bergh, in Semper’s Reisen, xvii. pp. 890-894.]

One specimen from near Wasin.

The notes on the living animal suggest that it is the same species

as M. ramosa, and say that it differs chiefly in that the branchie,

rhinophores, and processes of the velum are much smaller. The

colour appears to have been the same as in that species (7. e. cocoa

and green), and it is noted that there was a greenish tinge in the

branchiz. The back was warty.

The alcoholic specimen does not look much like IZ. ramosa. It

is rather bent, but the length appears to be about 21-5 millimetres,

the breadth 11-5, and the height 9. The back and sides are covered

with flat low tubercles and the epidermis comes off in flakes. The

dorsal margin is unusually prominent and projects 3-2 mm. It

* Tn this group as in others I have not access to the descriptions of a few ferms

by the older writers, e. g. Tr. palmeri.

96 SIR C., ELIOT ON NUDIBRANCHS [May 17,

bears eleven pairs of branchie, the main axis of which is bifid and

the secondary axis bifid again. The first pair of branchie are set

at the side of the rhinophores, which appear not to be on the

dorsal margin, but this arrangement may be due to the contraction

of the anterior part of the animal. The velum bears eleven simple

processes of irregular length; the outermost are tentacular and

grooved as usual.

The jaws bear a single row of very large, bent, almost hamate

denticles with slight indications of a second row. The radula con-

sists of 36 rows, with a maximum formula of 27+1.1.14 27, but

in most rows it is only about 154+1.1.1+15. The central tooth

is broad, and, as in J. ramosa, seems to bear five cusps. The

stomach is provided with the usual girdle of about 100 triangular

plates, all of much the same size.

This form appears referable with tolerable certainty to JM.

arborescens B.

MARIONIA RAMOSA, Sp. 0.

One specimen dredged in 5 fathoms, north of Kokotoni,

Zanzibar.

The notes on the living animal are as follows :—“ Colour cocoa-

like. Two rows of big branched processes which are greenish in

their finer divisions. The rhinophores and processes of the velum

very long. The neck part is long and the whole creature has the

shape of Limax. Length about 23 inches.”

The preserved specimen is of a uniform light yellowish green,

much bent, but about 27 millimetres long if stretched out. The

back is only 8 mm. across, but the whole animal looks much

broader on account of the large branchie. These are thirteen in

number, set on the somewhat projecting dorsal margin. ‘The first

are a little behind the rhinophores and the last at the end of the

tail. None are rudimentary, and the longest are 11 mm. long and

almost ribbon-like. The largest tufts consist of three main stems,

each of which is trifid again. The velum is ample and bears, in

addition to two tentacles of the usual grooved shape, twelve simple

digitate processes. The largest are 2 mm. long; the four in the

centre are much smaller than the others. ‘The sides and back are

tuberculate. The rhinophore sheaths are 5°5 mm. high, with

simple but ample and spreading margins. ‘The club is surrounded

by five plumes, once or twice pinnate.

The jaws are not very strong, and, except that the cutting-edge

is yellow, colourless. Both bear about thirty large pyramidal

denticles, at the base of each of which is a small accessory denticle.

In parts there are traces of a second line, which might be

regarded as mere ridges on the first line of denticles, but which

in seven or eight cases seem to be independent formations. The

transparent radula consists of 45 rows, containing at most 29

laterals, so that the formula is 45 x 29+ 1.1.1+29 as a maximum.

The central tooth is much as in Bergh’s plates of Marionia

arborescens (S. R. Heft xvi. pl. Ixxxvili. fig. 34), and, as

1904. ] FROM EAST AFRICA AND ZANZIBAR. 97

there, looks as if there were five cusps, but the median cusp is in

this specimen thinner and more pointed than in the plates. The

liver is large and yellowish. Embedded in the front part of it is

the stomach, consisting internally of a large, soft, laminated

portion, and a ring of about 120 yellowish, fairly stiff, horny

plates. They are not all of the same size, the largest being

2-millimetres long and 1 high, and the smallest about half as

large.

This species will perhaps prove to be only a variety of

M. arborescens, from which it is distinguished chiefly by its long

ribbon-like branchiz, which give it a remarkable appearance. The

jaws also present some differences.

MARIONIA ALBO-TUBERCULATA, Sp. 0.

One specimen from the neighbourhood of Wasin, East Africa.

Dredged.

According to the notes on the living animal the sides were

opaque white, with a reticulate pattern of red-brown. At the

centre of each mesh was a small white projection. The back,

which was dark brown at. the sides and greyish in the centre, bore

a similar arrangement of reticulations and projecting spots. The

sheaths of the rhinophores were tall, and the wavy edges were

turned over outwards. The branchiz were much subdivided, and

very large when fully extended. The main stems were of a light

greenish grey, and the finer branches of a dark yellowish brown.

The velum was plate-like, with five processes on each side, three

of which were branched.

The alcoholic specimen is 45 millimetres long, 15 high, and

13 broad. It does not taper toa point behind. The colour is

dirty yellow with profuse white markings. The stems of the

branchie are spotted and striped with white. There are nine

pairs of branchize of which the fourth is the largest, but the left-

hand plume of this pair is much larger than the other. The

middle and left-hand side of the velum are injured. There

remain on the right-hand side, starting from the inside, (a) a bifid

process, with three branches on each bifurcation, (6) a simply

quadrifid process, (¢) a simply bifid process, (¢d) a quite simple

process, (¢) a tentacle grooved below. Taking into consideration

the notes on the living animal, it appears that there was a similar

arrangement on the left side and that the middle of the velum was

smooth. There is a small oval papilla below the fourth branchia,

close to the dorsal margin. The genital papilla is lower down

between the second and third branchie.

The jaws are yellow, horny, and large, being 9 millimetres long

and 4 wide. They bear a single row of strong denticles, 10 of

which are very much larger than the rest. Under five of the

largest are indications of a second row. The radula is yellow, and

consists of forfy rows with a maximum formula of 95+1.1.1+95.

The central tooth is fairly broad and bears three cusps, of which

that in the middle is pointed and those at the sides blunt. The

Proc. Zoou. Soc.—1904, Vou. IT. No. VII. 7

98 SIR C, ELIOT ON NUDIBRANCHS [ May 17,

first lateral is large and clumsy: the rest are of the ordinary hamate

shape. The internal organs are whitish. From the large buccal

mass issues a tube 14 mm. long and nearly 5 mm. wide. The

interior is lined with folds, and there is a pouch-like diverticulum

in the floor immediately after the buccal mass. The tube is of

much the same size until it dilates into the small stomach

(7-5 mm. x 5°5 mm.), which is under and partly within the liver.

The stomach has a girdle of more than 100 plates, very thin and

membranous, and all of about the same size, namely, 3 mm. along

the base and 1:5 high. They lie in groups so as to produce a

superficial impression of about 12 thick plates. The intestine is

large.

MARIONIA VIRIDESCENS, sp. n.

[?= Tritonia hawaiensis Pease. |

One specimen from near Wasin.

The notes on the living animal are as follows :—“ Sides of foot

light greenish brown, netted with light bright green, which

becomes white near the edge of the back. There is a broad line

of opaque white and green (a mixture resembling verdigris), which

sends out prolongations to the bases of the branchie. Apart

from this line the colour is reddish brown with a greenish network

and white spots. This coloration extends into the main stems of

the branchie, but the finer ramifications are white and the finest

of all bright pinkish brown. The whole coloration is strikingly

beautiful. The velum bears seven processes on each side; only

the largest are branched. Therhinophores project but little from

their pockets, which are as in MZ. albo-tuberculata. The branchize

are kept moving continually, expanding and contracting. The

animal is about 4 inches long.”

The preserved specimen is 42 millimetres long, 21 broad, and 14

high. The shape is not tapering. The back and the sides bear

small flat tubercles. The velum is large; besides the two small

grooved tentacles it bears on each side seven processes, the largest

of which have 2-4 short branches. The central space is wide and

bears four rather indistinct tubercles not amounting to processes.

The rhinophores are entirely retracted, and the club is surrounded

by six bipinnate plumes. There are ten pairs of branchie, of

which the fourth is the largest; they still bear traces of green

colour. Thestout and strong main stem divides into four branches,

each of which bifurcates, and each bifurcation is then 3—4-pinnate.

The arrangement of the smaller tufts is simpler, but none are

rudimentary. The foot is very narrow, being, as preserved, only

2:5 mm. wide. The mouth is large and open, showing the jaws.

Tt is surrounded by a circular disk with thin free margins. All

this portion of the specimen seems to have been somewhat

distorted by the preserving fluid.

The jaws are 9 mm. long and bear a single row of coarse

denticles, of which ten are very large, the rest gradually decreasing

1904. | FROM EAST AFRICA AND ZANZIBAR. 99

in size. There are only very faint traces of a second row. The

radula consists of 37 rows, with a maximum formula of about

90+1.1.1+90. The teeth are of the shape usual in the genus.

The central tooth is finely striated, moderately wide and tricuspid,

the central cusp being pointed, those at the sides blunt. From

the buccal mass issues a long broad tube (4°5 mm. wide), which

passes above and to the left of the genital organs, and then enters

the liver, where it dilates into astomach bearing a girdle of plates.

These are about 120 in number, horny, fairly strong, brown,

triangular, and of various sizes, the largest being 4 mm. long and

1°5 high, the smallest only a quarter of the size and white. The

liver is large, yellowish externally, blackish internally.

In coloration this animal resembles J/. chloanthes B., but can

hardly be identified with that species on account of the differences

in the velum, jaws, central tooth of the radula, and stomach-plates.

I think it is very probably identical with Pease’s Tritonia hawai-

ensis from the Sandwich Islands, but his description is not suffi-

ciently detailed to make identification certain, and the expressions

“ Veil strongly digitate,” “ Tentacles [7. e. rhinophores] retractile

into .. . laciniated sheaths,” hardly apply tothe present specimen.

M. virescens and M, albo-tuberculata are closely allied and possibly

only varieties of one species; but the present specimens exhibit

some differences in the velum, median tooth, and digestive organs,

as well as in coloration.

MARIONTA, species.

One small specimen, dredged in 10 fathoms off Wasin.

Tt was dead when found and of a uniform opaque white. The

velum was hardly digitate, but presented six undulations. The

foot was broad.

The alcoholic specimen has become deep brown and is somewhat

decomposed. It is 5 millimetres long, 2 broad, and 1°5 high.

The back is tuberculate, with a slightly projecting margin, which

bears on each side 6 small branchiz set at a considerable distance

from one another. The rhinophore-pockets are raised and simple.

The velum appears simply circular.

No jaws could be discerned. The radula was extremely small,

and on a superficial examination appeared to be uniseriate, but

on careful investigation was found to have the formula 5.1.5.

The laterals are all alike, thin and hamate. They are folded over

the central tooth, a narrow plate with slight indications of being

tricuspid. The stomach contajned about 80 yellow plates, all of

much the same size.

This is perhaps an immature form in which the jaws are mem-

branous ; but, if so, it is remarkable that the stomach-plates are

fully developed. The extreme narrowness of the radula is also

remarkable. The characters as described above are sufficient to

constitute a new genus, but I hesitate to do this on the evidence

of one minute specimen,

100 SIR C. ELIOT ON NUDIBRANCHS [May 17,

BoRNELLA.

_ The members of this genus are slender, elegant animals, having

on either side of the back a row of cerata mostly divided into 2—4

branches and bearing gills. On either side of the mouth is a

compound tentacular process consisting of a number of conical

tubercles set in a sort of rosette. Over the head are a pair of

large organs called in the following descriptions for brevity’s sake

rhinophore-sheaths, but apparently formed by a fusion of the true

rhinophore-sheaths with a pair of cerata. The pair of cerata after

these organs are called the first pair. The vent is latero-dorsal

between the first and second pair of cerata. The buccal mass 1s

not large, but very muscular ; besides the jaws and radula, there

is alsoa labial armature of scales. The radula consists of a median

tooth, roughly triangular, either smooth or denticulate, and a few

(9-19) smooth hamate laterals, bent somewhat forward. The

innermost are generally very small, and the size increases towards

the outside of the row. There are two stomachs, of which the

second is armed with spines, and two accessory livers, besides the

main mass. Asa rule ramifications of the liver enter the cerata,

but there is some irregularity in this respect. The hermaphrodite

gland lies on the liver; the preputium is smooth or armed with

spines.

There is considerable difficulty in dividing the genus into species.

The colour presents little variety, being in all the known forms

whitish yellow, with a red or yellow reticulation on the back.

On the other hand, there is some variety in both the external

and internal organs. The number of the cerata and their sub-

divisions appears not to be specifically characteristic, but to increase

with age, and is not always the same on the two sides of the

body. The ramification of the liver may be present or absent in

the same species (B. eacepta; see Bergh’s two descriptions), and,

when present, may not extend to all the cerata. The armature of

hooks on the preputium may also be present or absent in the

same species (B. digitata and B. arborescens; see Bergh). Nine

species are recorded, but B. hermanni Angas, caledonica Crosse,

adamsii Gr., sempert Crosse, and hancockana Kel., will hardly

prove valid, for even if they represent specifically distinct forms

they are insufficiently characterised. Of the remaining species

B. calcarata Mirch, from the West Indies, is distinguished by

having appendiculate rhinophore-sheaths and smooth median

teeth. The Indo-Pacific forms fall into two groups—the one

represented by B. digitata, with a single process behind the rhino-

phores, cerata divided into rather long erect fingers, and median

teeth with faint denticulations; the other by J. excepta, with

several processes behind the rhinophores, small fingers on the

cerata protecting the external branchie, and much more distinctly

denticulate median teeth. Whether &. digitata and B. arborescens

are really distinct is discussed below. . stmplew, n. sp., 1s

certainly a separate species, unless it is a monstrosity.

1904. ] FROM EAST AFRICA AND ZANZIBAR. 101

My specimens seem to be on the whole smaller than those of

Bergh and to have fewer cerata.

BorRNELLA DiciTaATA Adams. (Plate IV. fig. 5a.)

[A. & H., Notes on a Coll. of Nud. made in India, p. 140,

pl. xxxili. figs. 8,9; Bergh, S. R. vii. p. 301; id. Danish Exp.

to Siam, Opisthobranchiata, p. 199.]

Several specimens from Zanzibar Harbour (Bawi and Prison

Island).

The living animals were white, with a granulated surface. On

the back was a reticulate pattern of deep orange. The cerata

were tipped with opaque white, below which was a band of bright

orange. The transparency of the body-walls varied in different

specimens. In some the liver and its ramifications were clearly

visible.

The following description, when not otherwise stated, applies to

the largest alcoholic specimen; the others are much like it, but

the smaller ones are only half the size. Length 30 millimetres,

breadth 4, height 8°5; much compressed laterally. On each side

of the mouth is a large branched process with about fifteen sub-

divisions; of these the four or five uppermost are larger and

digitate; the remainder are round and tubercular. The back

bears a pair of rhinophores with appendages, and, as a rule, four

pairs of cerata behind them. The largest specimens have a fifth

pair of small cerata, which in one case are fused together into a

single process. The rhinophore-sheaths are raised; they bear in

front three small digitate processes, and behind one long tapering

process which rises 5°5 mm. above the rhinophores. The first

pair of cerata are divided into two large and two small fingers ;

the right-hand member cf the second pair into two approximately

equal fingers, and the left into two largeand onesmall; the third

into one large and one small finger; the fourth are simple; the

fifth are merely small warts. In the smaller specimens the first

pair of cerata are trifid only, and in the smallest bifid, with indica-

tions of an incipient third digit. It appears probable that the

number of digits increases with size and age. The first pair of

cerata bear three branchie, the second, third and fourth two, the

fifth none. The branchie are all on the inner side of the cerata

and set close together.

The labial armature consists of small overlapping scales, arranged

in fairly regular rows. The edge of the jaws is quite smooth.

The radula consists of 34 rows. The median tooth has a long

central cusp, with from 8 to 10 denticulations or ridges at the base.

In most rows there are 9 laterals, increasing im size from the

innermost outwards, but in some the number rises to 13 and 15.

The walls of the second stomach are raised into folds on which

are set large brown thorns, with rather blunt tips. The ramifica-

tion of the liver appears to be very irregular and to vary in

different specimens. In the largest the arrangement is as follows :—

A single branch runs up into the tall tapering process behind each.

102 _ SIR C. ELIOT ON NUDIBRANCHS [May 17,

rhinophore; the first pair of cerata receive no branches at all ;

the second and third receive on the right hand a branch which

bifurcates, and on the left a simple branch which, in the third,

stops at the base of the cera and does not enter it. The re-

maining cerata receive no branches.

I think these specimens are the B. digitata described by A. & H.

and by Bergh. The best external character seems to be the

tapering, finger-like shape of the cerata and of the process behind

the rhinophores, to which no doubt the specific name is due.

BorNELLA ARBORESCENS Pease.

| Bergh, ‘Neue Nacktschnecken, No. ii.,” Journ. Mus. Godeffroy,

Heft vi. 1874, p. 96; id. 8S. R. xvii. p. 886.]

Several specimens from Mombasa Harbour. Note on living

animals: ‘ Yellowish white, with red reticulations on back and red

tips to cerata.”

The alcoholic specimens are all much of the same size. All are

whiter and more compressed than those of B. digitata, and the

cerata are much smaller. The average dimensions are :—Length

20 millimetres, height 6, breadth 3; rhinophores and cerata

about 2mm. high. The tentacular processes at the side of the

mouth consist of only about six digitations. The rhinophores are

as in B. digitata, but the posterior process is not solong. In most

specimens there are five pairs of cerata, of which the first three

are bifid and the remaining two simple. Each, from the first to

the fourth, bears two gills, the fifth none. The jaws and labial

armature are as in B, digitata; the formula of the radula is

about 40x 9.1.9, rising sometimes to 12.1.12. The teeth are

much as in Bergh’s plates (Journ. Mus. Godef. l. c. plate iv. 12),

but the central cusp of the median tooth is rather longer. The

median tooth is more erect than in #. digitata, and the 8-10

denticles which it bears less distinct and very hard to see. The

other characters are as in B. digitata.

It is not easy to say whether this form is specifically distinct

from B. digitata or, if so, whether it should be called B. arbo-

rescens. It represents, however, at least a well-marked variety or

stage of growth in which the tentacular processes, rhinophores,

and cerata are less amply developed. It could hardly be identified

with B. arborescens on the strength of the original description by

Pease (Amer. Journ. of Conchol. vi. 1871), but in the revised

description by Bergh (Mus. Godef. 7. c.) the chief specific character

seems to be “papillis anterioribus ut plurimum bipartitis.” In

these specimens they are invariably bifid. With regard to the

hooks on the preputium, I was unable to see the difference men-

tioned by Bergh, and found only simple or bifid hooks, not trifid,

in both species.

BorvEwLa Excerpta B. (Plate IV. fig. 56.)

[ Bergh, Challenger Reports, Nudibranchiata, p. 36; id. Danish

Exp. to Siam, Opisthobranchiata, p. 202.]

1904. | FROM EAST AFRICA AND ZANZIBAR. 103

One specimen from the East Coast of Zanzibar. Notes on

living animal: ‘“ Rhinophore bearers very large indeed; colour

whitish, netted with orange.”

The alcoholic specimen is more stoutly built than those already

described. It is somewhat bent, and would be at least 30 milli-

metres long if stretched out. It is 5 mm. high and 4°5 broad.

The rhinophore-sheaths are 8 mm. high, the cerata 5:5.

The tentacular process consists of 11 fairly long digits, all

distinct and none of them merely tubercles. The large rhino-

phores bear 7 digits, three in front quite separate, and five behind

united at the base. Posteriorly there are traces of what may be

a crest. Behind the rhinophore-sheaths are three pairs of cerata,

somewhat resembling those of Doto in general appearance. They

all bear three digits, above which rises the top, covered with knobs.

They also all bear three stout branchiz, two of which are visible

from the outer side and are protected by the digits.

The jaws and labial armature are much as usual; the former

have blunt indentations on the edge. The radula consists of 27

rows with a maximum formula of 16.1.16. In the median tooth

the central cusp is rather longer than depicted by Bergh, and

there are 10-12 denticles and ridges on each side. The second

stomach and the preputium are armed with black spines as

described by Bergh (Chall. Rep. 7. c.). The liver sends branches

into all the cerata except the right-hand member of the first pair,

but not into the rhinophore-sheaths.

Tam somewhat doubtful if this is really Bergh’s B. excepta:

there are differences in the arrangement of the cerata and branchiz

and the rhinophore-sheaths are relatively much larger, On the

other hand, the two specimens examined by Bergh did not agree

in details, and the present animal possesses more or less the

characters common to them.

BORNELLA SIMPLEX, sp. n. (Plate IV. fig. 5c.)

One specimen from Chuaka, East Coast of Zanzibar. The

following are the notes on the living animal :—“ Very like B. digi-

tata, but a distinct species. Anterior tentacles short and simple.

Whole coloration transparent, so that the walls of the heart

are distinctly visible. No opaque white or orange rings on

tips of cerata, but an orange network on the back and a row of

opaque white dots on the sides. Eyes not visible. Length 12

millimetres.”

Superficially the alcoholic specimen looks much like B. eacepta

as described above and has the same Dofo-like cerata, but it is at

once distinguished by having on each side of the mouth not the

usual tentacular rosette, but a single simple tubercle. The left

tubercle is larger than the right. The rhinophore-sheaths bear

six short digitations and a larger rounded knob behind. There

are four pairs of cerata, of which the hindmost are simple warts.

The others are similarly constructed, though the third pair are

smaller than the first two. Hach is divided into four knob-like

104 SIR C, ELIOT ON NUDIBRANCHS [May 17,

divisions, and bears a pair of trifid feathery branchie, one anterior

and one posterior.

The mouth-parts were taken out soon after the specimen was

captured, and as preserved consist of a labial armature and radula,

but no jaws. It is very likely, however, that the jaws had been

lost and were not really absent. The labial armature is much as

in B. digitata. Many of the scalesare heart-shaped. The formula

of the radula is 21x9+4+1+9, the number of laterals being con-

stant in all the rows. The median tooth has 7-8 very strong

denticles on each side of the central cusp, which does not project

much. The laterals are rather short and straight. The second

stomach is armed with spines as in JB. excepta. 'The liver sends

off diverticula into the process behind the rhinophores and all

four pairs of cerata. Those which pass into the rhinophore-

sheath and the fourth pair of cerata are simple, while those that

pass into the other cerata are divided into four branches corres-

ponding to the divisions of the cerata.

The simple tentacles of this animal are a sufficient specific, if

not generic character, provided they are normal. It is possible

that they area monstrosity, for itis not uncommon in nudibranchs

for external processes to remain undeveloped, for example, I have

a specimen of Ceratosoma cornigerum in which the characteristic

lobes are wanting. But apart from the tentacles, this specimen

does not exactly correspond with . excepia, for imstance as

regards the rhinophore-sheaths and radula. The median tooth

has fewer and stronger denticulations; the laterals are fewer,

shorter, and straighter.

PLEUROLEURA ALBA, Sp. 0.

[Cf. Pl. striata van Hass., Eliot in Nudibr. of Maldive and

Laccadive Archipelagoes, p. 566-7. |

Two specimens from Zanzibar. The following are the notes

on the living animal :—‘“ Back white with distinct low ridges,

longitudinal but not parallel to median line, each with a yellow

line along its summit. The rhinophores stand vertically or point

forwards and bear longitudinal perfoliations. The base is white,

the main part black, the apex truncated and yellow. ‘They are

not retractile into pockets, but can be withdrawn under the mantle-

edge. They are not very sensitive. The large velum and the

mantle are edged with bright yellow. Foot not half the width of

mantle. In crawling, the underside of the mantle is applied to

the substratum over which the animal moves. Length 13 milli-

metres, breadth 4 mm.”

The dimensions and colour of the preserved specimen have not

much altered, though the yellow has become faint. The shape is

elongate and tapering. The maximum breadth just behind the

rhinophores is 4 millimetres, rapidly decreasing to 3 mm. and

2mm. One striation runs down the middle of the back ; on each

side of it are six to eight others, not parallel to it and starting

1904. | FROM EAST AFRICA AND ZANZIBAR, 105

from various points. The external characters are those of the

genus.

The mouth-parts on the whole resemble those of Pl. striata

as described by me (J. ¢.). The formula of the radula is

236+1.1.14+6. The rhachidian tooth has a long central cusp

and about six denticles on each side. The first lateralis practically

half the rhachidian tooth, having one tall cusp and about six

denticulations parallel to it and rising from the base on the outside.

The remaining teeth are simply hamate. The jaws are more

membranous than in Pl. striata, and bear six distinct rows of

denticles.

This form is closely allied to Pl. striata, but differs strikingly

in colour, that animal being black with yellow lines. Such

variation in colour is not impossible within the limits of a species,

but in this case it is accompanied by other differences :—(1) The

shape is more elongate; (2) the radula is narrower; (3) the first

lateral is differently shaped. These points seem sufficient to con-

stitute provisional specific rank, though it is quite possible that

the form may ultimately prove a mere variety of Pl. striata.

EXPLANATION OF THE PLATES.

Prater III.

Figs. 1 a-lg. Notodoris minor, p. 84.

la. Lateral view of living animal. 10. Dorsal view of living animal.

1c. Hinder part of body with the valves raised and spread. 1d. Gills

with the valves removed. le. Ventral view of anterior part of body.

1f. Glans penis. 1g. Three teeth.

2a-2f. Trevelyana coccinea, p. 85.

2a-2d. First laterals of various shapes. Teeth from (2 ¢) middle

and (2,f) end of row.

8a-3c. Trevelyana ceylonica, p. 86.

3a. First lateral tooth. 36. Teeth from the middle of arow. 3c. Three

teeth, seen from below and behind.

4, Trevelyana crocea, p. 87.

Puate LV.

Figs. la-le. Prevelyana bicolor, p. 89.

la. Lateral view of living animal, with the liver (the blackish tint)

showing through the translucent body-wall. 16. Ventral view of

crawling animal, showing the proportions of the foot and some of the

internal organs through translucent body-wall. 1c. Gills as seen fully

expanded. Figs. 1 q@and 16 are X 5.

2. Nembrotha cristata, p. 90.

Middle of radula.

3 a-3d. Nembrotha affinis, p. 92.

3a, dorsal and (3 6) lateral views of living animal. 3c, median and

(3 d) first lateral teeth.

4, Marionia levis, p. 94.

5. Median teeth of (5a) Bornetla digitata (p. 101), (66) B. execepta (p. 102),

and (6c) B. simplea (p. 108).

106 ON A NEW SPECIES OF TREE-FROG. [May 17,

2. Description of a new Tree-Frog of the Genus Hyla, from

British Guiana, carrying eggs on the back. By

G. A. BouLencEr, F.R.S., V.P.Z.8.

[Received April 30, 1904. ]

(Plate V.)

HYLA EYANSI.

Tongue subcircular, slightly nicked behind, slightly free behind

and on the sides. Vomerine teeth in a strong transverse series,

narrowly interrupted in the middle, between the very large

choane. Head as long as broad; snout obtusely pointed, a little

longer than the diameter of the orbit; canthus rostralis strong,

straight; loreal region very oblique and concave; nostril much

nearer the tip of the snout than the eye; interorbital region

slightly concave, nearly as broad as the upper eyelid ; tympanum

very distinct, three-fifths the diameter of the eye. Fingers free;

no distinct rudiment of pollex; toes three-fourths webbed, the web

reaching the disks of the third and fifth, two last phalanges of

fourth free; disks a little smaller than the tympanum ; no tarsal

fold. Skin finely granulate above, more coarsely on the belly and

under the thighs, smooth on the throat. Brown above, speckled

and spotted with darker, limbs with regular dark cross-bars ;

white beneath, throat, breast, and limbs speckled with brown.

From snout to vent 75 millim.

The unique specimen, a female, carries its eggs on the back,

fitting into shallow hexagonal impressions in the skin; these eggs,

22 in number, measure 8 or 9 millim. in diameter, and contain

tailed larve with rudimentary limbs and with allantois-like

membranous respiratory organs. In this it agrees with Hyla

goeldi Blgr.*, to which it is nearly related, and with Ceratohyla

bubalus Esp.

This remarkable Frog was obtained at the end of November,

1902, at Groete Creek, Essequibo, by Dr. R. Evans, who has

kindly presented it to the British Museum, together with a sketch

made from the fresh specimen, and showing the embryos in

position ; these have unfortunately come off in transit.

EXPLANATION OF PLATE V.

Hyla evansi, sp.nov. Natural size.

* Cf. Boulenger, P. Z. S. 1893, p. 209, pl. x. figs. 1-8.

+ Cf. Boulenger, P. Z. S. 1908, ii. p. 115, fig. 8.

Ea SOO AG ol MPV

J.Green del.et lth . Mintern Bros .imp.

INA TE VAIN Syl

1904. ] ON THE ANATOMY OF CERTAIN SNAKES. 107

3. Notes upon the Anatomy of certain Snakes of the Family

Boide. By Frank E. Bepparp, M.A., F.R.S., Pro-

sector to the Society.

[Received May 2, 1904. ]

(Text-figures 19-23.)

The Boide are held by most zoologists to occupy a place near

the base of the Ophidian series. This view is based chiefly upon

the paired lungs, the considerable rudiments of the hind limbs,

and upon some other osteological points which are duly summed

up by Boulenger*. The viscera also confirm this opinion; and I

propose in the following pages to call attention to some new

or little-known facts relating to the circulatory system which

collectively support it.

§ Gubernaculum cordis, and Right and Left Aorte.

No member of the genera Python, Boa, Hunectes, and Hryx

which I have dissected possesses any trace whatsoever of a

gubernaculum cordis tying down the apex of the heart to the

walls of the pericardium. It is not altogether unnecessary to

record the absence of a gubernaculum, though it has been stated

that the Lacertilia are to be contrasted with the Ophidia by the

presence in the former and the absence in the latter of this

gubernaculum. A more correct statement would be arrived

at if the word “ generally” were interpolated in both

cases. I find, in fact, considerable vestiges of this tag in several

Ophidia. It occurs for example in Coronella getula. In Colopeltis

monspessulana a thin sheet of membrane runs from the ventricle

some little way above the apex to the vena cava and passes down

the latter to the posterior wall of the pericardium.

In the Hamadryad (Ophiophagus bungarus) the covering mem-

brane of the heart (=visceral layer of peritoneum) is obvious and

can be stripped off. Posteriorly, this membrane forms a tubular

prolongation of which one side is attached to the vena cava at its

entrance to the heart and to the pericardium beyond, while the

other is attached to the pericardium wall behind. The ventricle,

therefore, near to the apex, but on one side, is attached not merely

to the vena cava, but also to the wall of the pericardium.

It isimpossible to speak of these structures but as a gubernaculum

cordis. On the other hand, it is clear that they differ somewhat

from the gubernaculum in the Lacertilia, which has no relation

to the vena cava but attaches the actual apex of the ventricle to

the pericardial wall, and is of more ligamentous consistency. _

The structure is therefore possibly a new one in the Ophidia,

on which view its total absence in the Boide (so far as the

material which I have examined enables me to say) may well be

* Catalogue of Snakes in the British Museum (Natural History), London, 18938.

108 MR. F. E. BEDDARD ON THE [May 17,

a primitive character. There is no @ priori objection to deriving

the Ophidia from some Lacertilian form in which the characteristic

lacertilian “tag” to the heart was absent. In the genus Varanus,

for example, the gubernaculum is absent, as others as well as

I myself have observed; and it may be pointed out that the

Ophidia might well have been derived from some form in which,

as in Varanus, the neck was long, the lungs firmly bound down

to the dorsal parietes, the trachea or bronchi continued for a

considerable distance through the lung, and the urinary bladder

absent.

The fact that in Hryx the right and left aortic arches are equal

in size at their junction to form the unpaired dorsal aorta seems

to me to be undoubtedly a primitive feature. J may furthermore

observe that this feature is figured * by Dr. Gadow as charac-

teristic of Pelophilus (Boa) madagascariensis. Apparently, how-

ever, Python bivittatus has unequal right and left aorte 7. In

other serpents it is common for the right aortic arch to be

smaller than the left, and this is carried so far in Zamenis flagellr-

formis as to give the impression that the right aorta is a mere

inconspicuous forwardly running branch of the left £.

§$ Iniercostal Arteries.

The intercostal arteries in Hryx jaculus show some interesting

features, which are partly indicated in the accompanying drawing

(text-fig. 19, p. 109). In the anterior region of the body the

arteries in question arise from the left aorta immediately after it

has parted company with the anterior vertebral. Anteriorly to this

point some arise from the left aorta, but that region of the body

is supplied from the vertebral artery. The intercostal arteries

which arise from the right aorta and from the first part of the

conjoined aortz are single trunks given off at irregular intervals

not corresponding to the individual vertebra. They join above,

however, to form a continuous and slender dorsal artery which

may be termed the posterior vertebral artery (P.v.); from this

arise at regular intervals the paired intercostals,

After the end of this vertebral artery the aorta continues to

give off the dorsal intercostals which, when they reach the median

dorsal line, run along it for a short distance anteriorly and

posteriorly, giving off as before paired branches to the intervertebral

regions. But there is no formation here of a continuous

longitudinal trunk running over more than three or four vertebre.

So far the arrangement is precisely such as I have lately described

in another Boid, viz. Python spilotes§. But whereas in the last

* Bronn’s Klassen und Ordnungen des Thier-Reichs, Bd. vi. Abth. i. pl. cxxxv.

fig. 1.

+ Loe. cit. pl. xxxiv. fig. 2. The figure is copied from Fritsch.

+ Beddard, “Contributions to our Knowledge of the Vascular System in the

Ophidia,” P. Z.S. 1904, vol. 1. p. 338.

§ “Contributions to our Knowledge of the Circulatory System in the Ophidia,”

P. Z. 8. 1904, vol. i. p. 362. _ ; :

1904. | ANATOMY OF CERTAIN SNAKES. 109

named snake this arrangement of the intercostals persists to the

end of the body, in Hryx jaculus another arrangement comes into

force further down the body which is displayed in the drawing

reproduced as text-fig. 20, p. 110. First of all, ¢. e. anteriorly,

Text-fig. 19.

SENSES SS

(UFR

Ho.

SSS

Part of intercostal arterial system of Hryw jaculus.

Ao., right, Ao., left aortic arch ; es., esophageal twigs; P.v., posterior

vertebral artery; V., vertebral artery.

the intercostal arteries begin to arise regularly from the aorta,

and, when they reach the parietes, do not form short longitudinal

vessels which give off paired branches to several successive

110 MR. F. E, BEDDARD ON THE [May 17,

vertebre ; but each intercostal bifurcates close to the parietes'and

supplies but a single intervertebral area. The arrangement in

this region of the body in fact is precisely like that figured by

Jacquart * for Python sebe, and which I can confirm from my own

dissections of the same serpent. In Python sebe, however, this

arrangement appears to persist throughout the whole body. In

Eryx, on the other hand, a little way back, a third mode of

arrangement of the intercostal arteries occurs. The point of

bifurcation of the single intercostal advances higher up its stem

(@ in text-fig. 20) until (6) a right and left intercostal is established

arising separately from the aorta. A further differentiation is

shown in the case of the intercostals lettered ¢ and d in the

figure referred to. It will be observed that in c the left intercostal

is much thicker than its fellow and than most of the neighbouring

intercostals, while in d the vessel has become single owing to the

complete disappearance of its fellow. These facts indicate the way

in which the irregular intercostal arteries (sometimes right and

Text-fig. 20.

Part of intercostal arterial system of Eryx jaculus.

Ao., aorta; a, 6, c, d, intercostal arteries; C.v.7., C.v.l., right and left posterior

cardinals; G., gastric artery; Z., end of liver; S.mes., superior mesenteric

artery.

sometimes left and at unequal distances) of more modified snakes

have been formed. Obliterate in the accompanying drawing

(text-fig. 20) the finer intercostals and leave only those of

magnified calibre, and the result would be a reproduction of the

intercostal system in many Colubrine Snakes. This double series

of intercostal vessels in Hryx has its counterpart in Hunectes

murinus. In the Anaconda I find both intercostals arising singly

which run for varying distances along the median dorsal line

giving off paired branches to the parietes, and regularly paired

intercostals arising separately from the dorsal vessel It is

important to notice the agreement in these particulars between

Eryx and Hunectes and the difference from Python, since the two

former belong to the subfamily Boine, and the latter, naturally,

to the subfamily Pythonine. It 1s difficult to say which of the

* Ann, Sci, Nat. (4) iv. p. 321.

1904. ] ANATOMY OF CERTAIN SNAKES. 111

two arrangements of the intercostals, the Boine or the Pythonine,

is the more primitive ; perhaps it is the former. In any case it

is from the Boine rather than the Pythonine type that the inter-

costal system of many Colubrine Serpents is derivable.

Lryz conicus differs in detail from Eryx jaculus in the arrange-

ment of the intercostal arteries. Anteriorly, the arteries are

given off singly and regularly, bifurcating just before their entry

into the thickness of the parietes. There is with great regularity

one to each vertebra. The only exception that I noticed was in

the case of one intercostal arising from the right aorta in front

of the junction of the two aorte, and of another some way behind ;

these supplied two intervertebral spaces. This apparently is the

only trace left anteriorly of the arrangement characterising

Eryx jaculus. Far back, much further than in £. jaculus, the

paired arrangement of the intercostals is seen. Still it is evident

that the two species do not differ in the type of the arrangement

of the intercostals.

Eryx johni again shows a fundamental likeness to, but detailed

differences from, the other two species of the genus. The general

agreement is that the anterior series of intercostals arise singly

from the aorta and bifurcate only just before entering the parietes.

They begin to be double in origin shortly behind the liver. In

the region where they are double they are frequently asymmetrical

in both size and in point of origin from the aorta. Thirteen

intercostals arise from the right aortic arch before its union

with the left. There is no trace of any formation of an azygos

median vertebral artery such as occurs in Eryx jaculus. The

intercostal system of this species in fact is somewhat intermediate

between those of the two other species of the genus.

§ Some Visceral Arteries.

Esophageal arteries —I have examined these arteries carefully

in an injected example of Eryx jaculus. It agrees with other

snakes and with the Lacertilia in the fact that the intercostals

(already described on p. 109) arise from the right aortic arch

only. On the other hand, branches to the esophagus, which are

represented in the drawing (text-fig. 19, p. 109), arise either

directly or indirectly from both arches. I have not observed

this double origin of the cesophageal branches in any other

Ophidian ; but, as I am not quite in a position to deny its

occurrence, I cannot emphasise the fact as a characteristic of the

Boide. It is, however, 1 am inclined to think, an anatomical

feature not found in the Lacertilia. The cesophageal vessels, or

rather vessel (for I only noticed one), arises from the left aorta ;

it passes back along the cesophagus, giving off branches to that

gut, and becomes continuous with the first of the csophageal

arteries arising from the conjoined aorte. The right aorta does

not directly give off cesophageal arteries. But from two of the

intercostals of the right aortic arch such arteries arise,

112 MR. F, E. BEDDARD ON THE | May 17,

In Hryxjohni the same series of cesophageal branches arising from

intercostals are present. They arise from intercostals partly

belonging to the right aortic arch, and partly to those arising from

the common trunk. The third intercostal, after the union of the

two aortee, gives off a slender vessel which runs forward and joins a

vessel arising from the last but four of the intercostals belonging

to the right aorta. This longitudinal trunk gives off several lateral

vessels. The third, fourth, and fifth of the intercostals of the

right aortic arch also give off single cesophageal vessels.

I have pointed out in another paper* that the Lizard Pygopus

is unusual by reason of the fact that some of the visceral arteries

arise from intercostals instead of directly and independently from

the aorta. In Hry« johni precisely the same mode of origin

occurs not only for cesophageal arteries, but for a fat-body artery.

This springs from the right-hand intercostal of the seventh pair

after the posterior renal artery.

I observed the exact converse of this state of affairs in

Tropidonotus fasciatus. The right aorta gives off intercostal

branches, but no twigs to the cesophagus that I could find. On

the other hand, a single parietal vessel, accompanied closely by a

vein, enters the parietes a good way to the left of the middle

dorsal line and arises unmistakably from the left aortic arch,

which also, of course, gives off several branches to the cesophagus.

Gastric arteries.—The fact thatin Hrya there are only twot and

in Python spilotes only three gastric arteries, appears to me to be

an archaic point of structure in these Boid snakes. Among the

Ophidia generally there is frequently a large number of gastric

arteries. For example, inthe genus Coluber I have found as many

as ten or eleven. The reduplication of these and other arteries,

so characteristically Ophidian, seems therefore obviously to mark

the more specialised members of the group. The absence of, or less,

reduplication is not therefore inconsistent with the less modified,

more archaic, structure.

Ovarian arteries.—It is at least rare among snakes ¢ for the

arteries supplying the gonads to arise from the aorta opposite to

each other. Asa rule one spermatic or ovarian artery follows the

other in relation to the asymmetrically placed gonads. Never-

theless in a female Hryx conicus the two arteries arose side

by side. They immediately follow, as is usual, the superior

mesenteric. The paired condition of these arteries seems to me

to be a primitive feature in the organisation of this snake.

Renal arteries.—It is the general rule among the Ophidia for

each kidney to be supplied with a considerable number of arteries.

There are, for example, as many as eight m Coluber catenifer.

Among such Boide as I have examined, the number is invariably

one or two arteries only to each kidney. In Python sebe, Hunectes

* Above, p. 12.

+ T could find only one in Hrya johni.

+ I have not myself observed a single instance in the Ophidia except in the case

mentioned above.

1904. } ANATOMY OF CERTAIN SNAKES. i053

murinus, Hryx johni, and Hryx conicus each kidney had only one

renal artery. In Hryx jaculus there was some variation in the

two examples studied. In one there were two renal arteries for

each kidney ; in the other the left kidney had two arteries, the

right only one. Here, it may be remarked in passing, is an apparent

difference between the two species of Hry«x investigated by me,

which f shall refer to again later.

Now, as the kidneys of Hrya are very short * and those of

Coluber catenifer long, as they are in the majority of snakes, it

might be held that the arterial blood-supply had merely a relation

to length. That the character is one peculiar to the Boidze seems

to be shown by the case of Hunectes murinus, for in this serpent

there is only one renal artery to each kidney; and yet those

organs, in the individual which [ dissected, measured respectively

15? and 123 inches in length.

§ Veins of the Posterior Abdominal Region in Kunectes.

The caudal vein emerges from the thickness of the parietes some

way behind the cloaca. When it reaches the level of the cloaca,

two veins, asymmetrically placed with regard to one another,

join it. I suppose that these are the equivalents of the

ischiadic veins of Lizards. Further forward, between the cloaca

and the very anteriorly situated kidneys 7, the caudal vein divides

at once into three branches. The middle one of these is the

right and larger anterior abdominal vein. To the left of this

arises the afferent renal vein of the left kidney, and to the right

of the anterior abdominal a vein which runs over the viscera to

the dorsal surface of the body, where it enters the parietes to the

left of the middle line, after running both forwards and backwards

for a short distance. I am inclined to regard this vein as the equiva-

lent of the lateral abdominal vein of its side in the Lacertilia. Its

place of origin agrees with such an homology, and the shortness

of its course within the body-wall is no reason against the ecom-

parison, since it is of varying length among the Lacertilia. It

might be held that this vein is in reality only the proximal end

of the left afferent renal which has lost its connection with the

remainder of that vein, were it not for the conditions observable

in Hryx to which I recur later (see p. 114). There is, in fact, no

afferent renal on this side of the body springing from the caudal

vein and corresponding in origin to the fully developed afferent

renal, whose origin on the other side of the anterior abdominal

has been already referred to. It rises, I presume, from the eaudal

vein further back.

* In Hryx johni, which measured 26 inches from snout to vent, the kidneys were

respectively = inch (right kidney) and 1 inch (left kidney) in length. In H. jaculus

of 15% inch length to vent, the kidneys were proportionately larger, i. e. inch and

1 inch, and in #. conieus of 27 inches this was also the case ; they measured 14

and 1+ inch.

+ The kidneys are well known to occupy a very anterior position in the Boidz.

Tt has not, I think, been noted that Hryx as well as Humectes agrees in this with

Python and Boa.

Proc. Zoou. Soc.—1904, Vou. IT. No. VIF. 8

114 MR. F, B. BEDDARD ON THE [May 17,

The anterior abdominal vein is double, but the right vein is

very considerably larger than the left, especially posteriorly. The

two join far forward in the immediate neighbourhood of the

pancreas. They are here more nearly equisized. Posteriorly

there is another and the only other junction between these paired

veins not far in front of the trifurcation of the caudal vein. After

this point the left anterior abdominal vein receives a branch from

the body-wall and ceases. It is not directly connected, as already

mentioned, with the caudal vein. Just after its origin from the

caudal vein the anterior abdominal gives rise to the single median

epigastric vein, which runs forward at least as far as the liver, to

which it gives off several branches.

§ Veins of the Posterior Abdonunal Region mm Hryx.

It is remarkable that differences occur in these veins between

the two species Lryx jaculus and EH. conicus, if, that is to say,

Text-fig. 21.

Certain abdominal veins in Erya conicus.

a, b, ¢, parietal veins; Ant.Abd., anterior abdominal ; Ep., epigastric;

L.aff-r., lett afferent renal; R.affz., right afferent renal.

the single example of #., jacwluws which I have examined represents

the normal state of affairs. The veins in question in Zryx conicus

1904. ] ANATOMY OF CERTAIN SNAKES, 115

(text-fig. 21, p. 114) are disposed as follows:—The caudal vein

gives off first the right afferent renal; after the cloaca (¢. e. in front

of that aperture) it gives off two branches to the dorsal parietes

and then divides into three veins as in the Anaconda. Shortly

after its origin, the anterior abdominal gives rise to the single

epigastric vein. The arrangement of veins in this snake is in

Text-fig. 22.

Raf

Loffy.

Certain abdominal veins in Eryx jaculus.

R, branches to rectum; cl., branches to cloaca. Other letters as in text-fig. 21.

fact precisely as in Humectes. In Hryx jaculus, on the other

hand, there are differences which are illustrated in the accom-

panying figure (text-fig. 22). The caudal vein apparently divides

into two—but Iam a little uncertain—and each of these vessels

116 MR. FE. E, BEDDARD ON THE [May 17,

gives off at least one branch to the rectum. A little further

forward each vein certainly divides into two branches. These

branches are, on each side, a renal afferent and a component of

the anterior abdominal vein. Both afferent renals receive, as

usual, branches from the dorsal parietes. The two roots of the

anterior abdominal vein are unequal in size, that of the right

hand being considerably the less. It is important to note this

fact, since in Hryx conicus the left root of the anterior abdominal

vein is the only one which persists. The epigastric vein arises

from the anterior abdominal shortly after the junction of its two

roots. Lryx johni agrees with H. jaculus.

It is a peculiarity of Snakes as contrasted with Lizards, that the

anterior abdominal is occasionally partly double, whereas in Lizards

it is single after the fusion of its two roots. In the Colubrine

and Viperine snakes, so far as my experience goes, the anterior

abdominal is usually single except at its extreme posterior end.

In Zamenis gemonensis the vein bifurcates posteriorly, and after

a very short course ends in minute branches in the fat-body.

In Causus rhombeatus the extent of the bifurcate region of the

anterior abdominal is not much greater; for 8 inches intervene

between the opening of the anterior abdominal into the portal

and its bifurcation posteriorly, which is 2% inches from the vent.

In Boa constrictor, on the other hand, there are 25 inches between

the vent and the fusion of the two anterior abdominals anteriorly,

which point is 10 inches behind the liver, and therefore less from

the point of union of anterior abdominal and portal *.

Tn the Anaconda the double character of the anterior abdominal

vein has been already referred to. In onespecimen of Hryx conicus

it was single throughout. In another it was partly double, as

was also the case with two individuals of Hryx jaculus. In

Python sebe (where it is figured as partly double by Jacquart 7)

the fluctuation of this vein between the single and double condition

was more plainly seen. Just in front of the gall-bladder the

vessel communicates with the gastric portal vein ; from this point

to two inches behind the gall-bladder it is single. Then fora

distance of 44 inches it is formed of two tubes lying side by side ;

these reunite, and finally again separate to form two tubes.

In Eryx johni the vein appears to be single after the union of

its two posterior roots.

Jacquart figures a somewhat different state of affairs in Python.

The anterior abdominal bifureates posteriorly and communicates

with only one afferent renal directly as in the Anaconda. The

other branch only communicates indirectly (by means of small

veins) with the left renal afferent. Hochstetter { observes that in

Tropidonotus natrix and Coluber csculapii there is no direct

connection between the abdominal and the renal afferent veins.

TI can quite confirm this by the conditions observable in Zamenis

* T did not ascertain this measurement.

+ Ann. Sci. Nat. (4) vi. p. 321. { Morph. Jahrb, xix. p. 489.

1904.] ANATOMY OF CERTAIN SNAKES. Thy

gemonensis. I take it that the posterior bifurcation of the

anterior abdominal vein in the last-mentioned Colubrine snake

(and possibly in Cawsus and other forms) is a reminiscence of its

former origin by two roots from the renal afferent veins as in

the less modified Lacertilia, and, as I have already shown, in the

especially “ Saurian” Hryx jaculus and Eryx johni.

It is clear from the foregoing, that the somewhat divergent

opinions of previous authors are partly due to actual differences

in the abdominal and afferent renal veins of different Ophidia,

to our knowledge of which I have been able to add something.

It is furthermore clear that the Boidee contrast with other Ophidia,

so far as observation has gone, in thei greater approximation to

the Saurian type of organisation ; they are, in fact, more primitive

than other Ophidia. In these Ophidia alone is the anterior

abdominal vein connected with one or both of the afferent renals,

and in them there is generally doubling of the anterior abdominal

vein in front of the junction of its constituent veins. Of all the

Boidee whose anatomy is known, Lry«x jaculus (not EH. conicus)

comes nearest to the Saurian type in that its anterior abdominal

vein arises from two distinct roots, one from each of the renal

afferent veins.

§ Remains of Cardinal Veins.

In comparing the venous system of the adult Tropidonotus

with that of the adult Lacerta, Dr. Hochstetter* arrived at the

conclusion “dass das Venensystem der Lacerta. auf einer etwas

niedereren Entwicklungsstufe strehen geblieben ist als das von

Tropidonotus. Bei Lacerta ist das System der Vertebralvenen

noch erhalten, wahrend es bei Tropidonotus nahezu vollig gesch-

wunden ist.” Hochstetter’s statements are also accurate when

applied to other Colubrine snakes. Ihave examined Lioheterodon

madagascariensis with some care from this point of view. In

that snake there is hardly any development of vertebral veins,

such as I shall describe immediately in the Boide. About half-

way down the liver, however, is a longitudinal vessel running for

a short distance, from which arises a tributary to the hepatic

portal system. Another Colubrine snake, viz. Zamenis gemonensis,

showed an interesting persistence of a portion of the posterior

cardinal vein precisely comparable to what is to be found in the

Chameleon t and in Pygopust. The afferent renal artery in

front of the left kidney, instead of ending, as is usual, towards

the anterior end of that gland, passes beyond it and imbeds itself

in the body-wall to the left of the median dorsal line. I could

find nothing to correspond on the right side. If really absent—

and Jam convinced that there is at least nothing really con-

spicuous—this is another example of the asymmetry of Ophidian

* Morph. Jahrb. xix. p. 493.

+ Hochstetter, loc. cit. p. 462, and Beddard, loc. cit. infra.

ft Beddard, “Contributions to the Anatomy of the Lacertilia, No. 8,’ P. Z.S.

1904, vol. 11. p. 12.

118 MR. F, E, BEDDARD ON THE [May 17,

structure. The discovery of this prerenal portion of the posterior

cardinal in an Ophidian removes another point of dissimilarity

between the Ophidia and the Lacertilia.

Text-fig. 23.

Liver and certain adjacent veins in Eryx conicus.

Ao., aorta; cs., cesophageal branches of longitudinally running left posterior

cardinal; Int., intercostal branches of the same; DL, liver; P.v., portal vein ;

St., branches to stomach.

The Azygos vein is admittedly a vestige of the posterior cardinal

1904. ] ANATOMY OF CERTAIN SNAKES. 119

of its side. This vessel appears to be well developed in some

Boid serpents, and that fact appears to me to indicate the retention

of a primitive feature*. In Hryx jaculus, for example, the vein,

which is on the right side, extends back over no less than twelve

intercostal spaces and nearly reaches the junction of the two

aorte. This contrasts with the same vein in Coronella getula,

which only extends over four of these spaces. In a specimen of

Hryx conicus the azygos vein extended only over ten vertebra.

In both specimens of Hryx conicus the azygos, after a break,

reappeared in the region of the liver, where its course is shown in

the accompanying drawing (text-fig. 23, p. 118). It will there be

seen that the vertebral vein with one gap near the anterior end

of the liver runs continuously to a point some little distance

behind the liver. It gives off branches on the one hand

to. the dorsal parietes, and on the other to the portal vein.

Anteriorly to the liver the branches go to the ceesophagus. In Hrya

jaculus (text-fig. 20, p. 110, C.v.1., C.v.r.) there were conspicuous

traces of both posterior cardinals behind the liver. As will be

seen in the drawing referred to, the left posterior cardinal is

continued anteriorly beyond its junction by a conspicuous branch

with the portal trunk in the immediate neighbourhood of the

liver ; posteriorly it ends near to the commencement of the right-

hand vein, the two being therefore supplementary to each other.

The left is considerably the longer. The right extends back a

little way beyond the origin of the superior mesenteric artery

from the aorta. In Python sebe a corresponding vein occurs in

the region of the liver, but it extends both further forward and

backward than I have observed in Hryx. Anteriorly it extends

beyond the junction of the two aorte, and posteriorly it reaches

very nearly to the gall-bladder. I do not for the present suggest

that these longitudinal vessels are more developed in the Boide.

I simply call attention to their arrangement.

§ On the Specific Differences between Eryx jaculus, E. johni,

and Ki. conicus.

These species can be readily separated by external characters,

as Boulenger plainly sets forth in the British Museum ‘ Catalogue

of Snakes.’ They also, however, differ in other points besides

the obtuse or pointed tail, the absent or present mental groove,

and the form of the rostral scale, &c.

There are differences, in the first place, in the form of the

liver. In “ryx jaculus and H. johni the lobes are unequal, and

the right lobe extends further down the body than the left, to

the extent of about half an inch. In Eryx conicus they are as

near as possible exactly equal. As these results depend upon the

examination of two examples of each of the species /. jaculus and

* As I have pointed out in a preliminary account of some of the facts detailed

in the present communication (Ann. & Mag. Nat. Hist. (7) xii. p. 233).

120 ON THE ANATOMY OF CERTAIN SNAKES. [May 17,

Lf. conicus, the character may, I take it, be regarded as a genuine,

if small, difference.

Secondly, the right lobe of the lung extends further back in

Hryx jaculus and H. johni. In those species it reaches the gall-

bladder ; in Hryx conicus it falls short of the gall-bladder by at

least an mch.

The superior mesenteric artery arises in Hryx jaculus and

LH. johni distinctly in front of the gall-bladder. In Eryx eonicus

its point of origin is as distinctly behind the gall-bladder. This

character is perhaps less likely than some others to prove of value

as an absolute mark of distinction.

More important are other differences in the vascular system.

In Fryx jaculus, though it is the smaller species, there are appa-

rently, as a rule, two arteries supplying each kidney; in Lryax

conicus there is but one to each kidney: and here apparently

Lf. johni agrees with H. conicus. The intercostal arteries have a

different arrangement m the two species. In #. jaculus there is a

well-developed posterior vertebral artery formed by the junction of

irregularly arising intercostals; in /. conicus this does not exist

(except as the merest rudiment). The paired intercostals com-

mence further forward in /. jaculus than in Z. eonicus. H. johni

m these particulars is somewhat intermediate. In 4. jaculus

there is a double connection of the anterior abdominal vein with

the afferent renals posteriorly; in #. conicus this occurs only

on one side. HH. johni agrees with H.jaculus. In relation to

these anatomical differences, which appear to me to be fully as

great as those which distinguish either species from Hunectes, I

would draw attention to the restricted range of #. conicus and to

the wider distribution of H. jaculus and EL. johnt.

Résumé of principal facts.

It may be convenient to deduce from the foregoing pages the

main facts in the vascular system of the Boide examined ; such

may be divided into two heads, 7. ¢., those which appear to argue

a basal position among the Ophidia, and those which are of sys-

tematic importance in the group. As to the former it may be

noted that

(1) The heart is always without the least trace of a guber-

naculum cordis.

(2) The two aorte are usually equisized at their point of union

to form the dorsal aorta.

(8) The renal and gastric arteries are much fewer in number

than in other Ophidia, the former consisting generally of

only one artery to each kidney. This distribution has no

relation to the size of the kidney.

(4) The intercostal arteries are always symmetrical and for

the most part regular in their arrangement, frequently in

regular pairs.

2) Te

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1904.] ON ENTOMOSTRACA FROM NATAL. © 121

(5) The anterior abdominal vein always arises from one afferent

renal, rarely from both as in the Saurians. Frequently,

also, it is double.

(6) The existence of veins continuing the azygos posteriorly

is usual,

Facts which are of importance for the systematic arrangement

of genera and species within the family Boidee :—

(1) The Boinz (Hunectes and Hryzx) differ from the Pythoninz

(Python) in that the intercostals are posteriorly paired

arteries, while in the Pythoninz a single median artery

divides into two close to the median dorsal line throughout

the series.

(2) The three species Hry« conicus, E. johni, and EF. jaculus

differ from each other ina large number of anatomical

features.

Besides these points several other anatomical features are of

interest as new or rare among Ophidia. Such are

(1) The continuation of the afferent renal of the left kidney

in Zamenis gemonensis beyond the kidney into the parietes,

as in Chameleon and Pygopus.

(2) The origin of cesophageal arteries not only from the left

aortic arch but from some of the intercostals of the right

half arch.

(3) The fusion of some of the anterior intercostals in Hry«

jaculus (and Python spilotes) to form a continuous longi-

tudinal trunk lying dorsally of the aorta.

(4) Representatives of the lateral abdominal vein of Lizards

appear to exist in certain snakes (e. g. Hunectes and Hryz).

(5) Origin of a fat-body artery in Hryx johni from an

intercostal,

4, On Entomostraca collected in Natal by Mr. James Gibson.

By G. Stewarpson Brapy, M.D., LL.D., D.Se., F.R.S.,

C.M.Z.S8.

[Received March 25, 1904.}

(Plates VI.-VIII.*)

For the opportunity of examining and describing these species

I am indebted to the kindness of Mr. James Gibson, Resident

Magistrate at Greytown, by whom they were collected in the

summer of 1902. All are freshwater species, and were found in

pools in the neighbourhood of Greytown, Natal. The identity of

some of them with European forms is a point of considerable

interest, and indeed the general aspect of the gatherings is quite

* For explanation of the Plates, see p. 127.

122 * DR. G. STEWARDSON BRADY ON [May 17,

similar to what one would expect in ordinary Northern collections.

With the exception of one species, for which I have thought it

best to propose a new generic name—ALctocyclops—all ave refer-

able to familiar European genera.

CYCLOPS LEUCKARTI Claus.

This appears to be less plentiful in the Natal gatherings than

any of the following species, though more conspicuous owing to its

greater size.

The form described by me years ago under the specific name

scourfieldi* has been identified by other authors (Lilljeborg,

Herrick, Schmeil) with C. leuckarti Claus. 1am doubtful as to

the correctness of this identification. Both Schmeily and

Herrick+ figure, with differences, peculiar pellucid marginal

lamine on the last two joints of the larger antenne. I have

been unable to detect any such structure in my British specimens

of C. scourfieldi, neither does it exist in the Natal specimens nor

in others from Ceylon which I refer to the same species. And it

may be noted that Lilljeborg, while accepting Schmeil’s description

and figures as belonging to C. leuckarti, does not himself refer in

his definition to these antennal lamin. Herrick, on the other

hand, figures and describes them, and expresses a doubt as to the

identity of the species with C. scourfieldi. I do not myself

possess authentic specimens of C’. lewckarti, and must be content

for the present to leave the question im suspense.

CYCLOPS PUSILLUS, n. sp. (Plate VI. figs. 11-18.)

Female. Body slender, tapering gradually from before back-

wards (fig. 11); thoracic segments not produced at the angles,

except the last which is extremely short and angulated: abdomen

very slender, the first segment nearly as long as the united lengths

of the following three segments ; furcal segments slender, scarcely

longer than the preceding abdominal segment; principal tail-seta

nearly as long as the whole abdomen. Ovisacs closely adpressed

to the abdomen and containing only a few comparatively large

eggs. Antennules twelve-jointed, slender, somewhat longer than

the entire cephalothorax, bearing numerous long setz (fig. 12).

All the branches of the first four pairs of feet three-jointed ;

spines of the first pair (fig. 17) very slender and setiform ; fifth

pair (fig. 18) rudimentary, almost obsolete, consisting of a minute

papilla from which arise two unequal sete. Length, exclusive of

tail-setee, 0°46 mm.

This is the smallest species of Cyclops with which I am

acquainted. But its general development and the fact of many

specimens bearing ova, indicate that it is not an immature form.

* Brady, G.S. “A Revision of the British Species of Freshwater Cyclopide and

Calanide ” (Nat. Hist. Trans. Northumberland & Durham, vol. xi., 1891).

+ Schmeil, Deutschlands freilebende Stisswasser-Copepoden. 1 Teil, Cyclopide.

1892.

{ Herrick, Synopsis of the Entomostraca of Minnesota, 1895.

1904.] ENTOMOSTRACA FROM NATAL. 123

CyCLOPs GIBSONI, n. sp. (Plate VI. figs. 1-10.)

Female. Seen dorsally the outline is slender, gradually taper-

ing from before backwards, greatest width equal to about one-

third of the length (fig. 1); the second segment as wide as the

cephalic segment, slightly produced and angulated posteriorly,

third segment narrower and scarcely at all produced at the

angles, fourth segment again narrower and without produced

angles; last thoracic segment about as wide as the fourth from

side to side, but much narrowed from before backward, its lateral

angles distinctly produced and bearing a brush of six or eight

rather long sete ; abdominal segments gradually tapering back-

wards, the combined lengths of the second and third equal to that

of the fourth, first segment rather larger than the fourth ; caudal

stylets slender (fig. 10), nearly equal in length to the abdomen,

about five times as long as broad; the innermost of the two’

apical setee is the longer and is about equal in length to the entire

abdomen, outer setee somewhat shorter; the subsidiary sete are

three in number, short, subequal, and arise from the apices of the

stylets; on the side of each stylet not far from the distal end is a

single very minute seta; the larger sete are very finely plumose.

The antennules are twelve-jointed (fig. 2), slender, bearing

numerous sete, and reach backwards as far as to the front of

the second segment: the proportionate lengths of the joints are

indicated in the following formula :—

I 24 Sol 4a Os OME ESes (Olas Rtn

PEL Om eLpe Say ey aEe ae

The mandible is of the usual form, with a rudimentary palp of

three setze, two long and one short (fig. 3); the chewing portion

of the maxilla (fig. 4) is of normal shape, with a laminar palp bearing

several plumose sete. The rami of the first four pairs of feet are

all three-jointed ; terminal joint of the outer branch of the first

pair bearing two slender spines on the outer margin, three long

sete on the inner margin and two shorter apical sete, the outer-

most being finely bipectinate (fig. 7); the terminal joint of the

inner branch has one long apical spine and on its outer margin one

very short spine and one slender seta, its internal margin has

three setz and one at the apex. The second, third, and fourth

pairs (fig. 8) are nearly similar, but with more elongated joints and

more robust spinous armature. The fifth pair (fig. 9) is uniarti-

culate, with three subequal setz, the innermost of which is densely

plumose. Length 1 mm.

Among European species those most nearly allied to C. gibsoni

seem to be C. varicans, C. bicolor G. O. Sars and C. gracilis

Lilljeborg; but all of these have the rami of the swimming-feet

uniformly biarticulate, whereas in C. gibson they are triarticulate.

CYCLOPS FIMBRIATUS Fischer.

This appears to be one of the commonest species in Mr. Gibson’s

124 DR. G. STEWARDSON BRADY ON [May 17,

collection, agreeing down to the minutest details with British

specimens.

EcrocycLops, nov. gen.

Like Cyclops except that the mandibles, maxille, and maxillipeds

are less fully developed and are very indistinctly jointed, the

posterior antenna bears a very stout ciliated seta on the posterior

margin, and the terminal joints of the rami of the swimming-feet

are scarcely at all elongated.

ECTOCYCLOPS RUBESCENS, n. sp. (Plate VI. figs. 19-20;

Plate VII. figs. 21-27.)

Female. Body depressed, rather robust (fig. 19) ; cephalothoracie

segments not produced laterally ; abdominal segments short and

stout, posterior margin of the last segment pectinated, furcal

segments not much longer than broad; principal tail-sete

marginally aculeated, the innermost nearly equalling three-fourths

of the length of the body; ovisacs large and adpressed. Anten-

nules (fig. 20) ten-jointed, much shorter than the first cephalo-

thoracic segment, and rather densely setiferous ; antenne (fig. 21)

robust, anterior margin fringed towards the base with numerous

short sete, penultimate joint with about six larger sete, terminal

joint with several longer sete; on the posterior margin rising

from the base of the second joint is a very long and stout

ciliated appendage. Mandibles bearing a rudimentary palp con-

sisting of three very small sete; maxille (fig. 22) not much

different from those of Cyclops ; anterior and posterior maxillipeds

(figs. 23, 24) small, stout, and indistinctly jointed, the marginal

curved teeth of the anterior pair very robust; rami of the

swimming-feet three-jointed (figs. 25, 26), fifth pair obsolete,

represented only by three short plumose setee (fig. 27). Colour of

the posterior part of the body faint reddish-brown, the anterior

part nearly colourless (in spirit specimens). Length 0°85 mm.

ATTHEYELLA NATALIS, n. sp. (Plate VII. figs. 28-33.)

Female. In general outline like Canthocamptus (fig. 28).

First thoracic segment coalescent with the head; _ posterior

abdominal segments spinulose on the hinder margin ; furca very

short; principal tail-setee equal to at least half the length of the

body. Antennules slender, nine-jointed (fig. 29), shorter than

the first cephalothoracic segment, and very sparingly setiferous ;

antenne (fig. 30) bearing a minute one-jointed secondary branch.

First pair of swimming-feet (fig. 31) short, both branches three-

jointed, outer about equal in length to the first two joints of the

inner branch ; second, third, and fourth pairs having the outer

branch composed of three, the inner of one joint (fig. 32); fifth

pair (fig. 33) foliaceous, two-jointed, basal joint wide and bearing

four long set on its inner and one on the outer lobe, terminal

joint ovate and bearing three unequal apical sete. Length

0-75 mm.

1904. | ENTOMOSTRACA FROM NATAL. 125

CYPRIA CASTANEA, n. sp. (Plate VII. figs. 40-42; Plate VIII.

fig. 43.)

2 Cypria lacustris Lilljeborg.

Shell, seen from the side, subovate (almost semicircular), highest

in the middle, height equal to nearly two-thirds of the length

(fig. 40); extremities broadly rounded, the anterior bordered by a

narrow translucent fillet, dorsal margin boldly and evenly arched,

ventral nearly straight: seen from above (fig. 41) the outline is

narrowly ovate, widest behind the middle, width equal to more

than one-third of the length, tapered to an acute point in front,

rounded behind. Surface perfectly smooth and polished; colour

reddish-brown, without any trace of hairs or sculpture. Length

0-65 mm. Limbs closely resembling those of C. ophthalmica, but

much more slender, both as regards bulk and armature (figs. 42,

43.)

This seems to me to approach most nearly to C. lacustris

Lilljeborg, if indeed that be truly a distinct species, and there

is not much to separate it from the common Northern species

C. ophthalmica, except the absence of shell-markings and the

much more slender build of the limbs, of the post-abdominal

claws, and of the antennal sete. The specific name now proposed

must, however, be looked upon as merely provisional.

CyYPRIS INERMIS, n. sp. (Plate VIII. figs. 44-49.)

Shell, seen from the side, oblong-ovate (fig. 44), rather higher in

front than behind, height equal to less than one-half the length,

dorsal margin gently arched, ventral shightly sinuated ; anterior

extremity evenly rounded, posterior very much narrower and

rounded ; extremities and ventral margin fringed with fine hairs :

seen from above (fig. 45) the outline is ovate, greatest width about

equal to the height and situated near the middle, lateral margins

evenly arcuate, extremities evenly rounded, the anterior rather

the broader of the two. In young specimens the postero-inferior

angle of the right valve bears two sharp teeth (fig. 46). Colour

of the shell greyish white. Length 0°95 mm.

The antenna (fig. 47) bears a brush of four sete which reach as

far as the extremities of the very slender terminal claws, Apex

of the second foot bearing a single slender curved claw (fig. 48)

and a long seta. Post-abdominal rami (fig. 49) long and slender,

finely ciliated on the hinder margin; all the setz slender and

crowded together apically, of these two are stouter than the rest

and marginally pectinated.

This species occurred abundantly in one of Mr. Gibson’s

gatherings, less plentifully in another.

CypRIS ARATRA, n. sp. (Plate VII. figs. 34-38.)

Shell, seen from the side (fig. 34), subovate, highest in the middle,

height equal to more than half the length; anterior extremity

broadly and rather obliquely rounded, posterior much narrower

126 DR. G. STEWARDSON BRADY ON [May 17,

and less distinctly rounded off; dorsal margin boldly arched,

sloping more steeply behind than in front, ventral feebly convex,

with no trace of sinuation: seen from above (fig. 35) the outline

is very broadly ovate, widest in the middle, width equal to nearly

three-fourths of the length, extremities obtusely pointed, the left

valve larger and overlapping the right both in front and behind.

Surface of the shell marked throughout with delicate and closely-

set longitudinal furrows and bearing scattered hairs of variable

length. Colour dusky greenish grey. Length 1:0 mm.

The antenne bear fascicles of setze reaching to the extremity of .

the claws, exactly as in the preceding species; second pair of feet

with a slender terminal claw (figs. 36, 37), one long seta and

several small claw-like processes *; post-abdominal rami (fig. 38)

extremely slender, bearing two apical setze and one much smaller

lateral seta situated not very far from the apex.

The armature of the second pair of feet is here very different

from that of a typical Cypris, but this character seems insufficient

to give it separate generic rank.

STENOCYPRIS PERARMATA, n. sp. (Plate VIII. figs. 50-57.)

Shell, seen from the side (fig. 50), elongated, siliquose, greatest

height equal to rather more than one-third of the length and

situated in the middle; anterior extremity evenly rounded, posterior

suddenly tapered, narrow and rounded off below ; dorsal margin

arcuate, sloping evenly toward the front, more abruptly and with

a slight sinuosity behind, ventral slightly sinuated near the front

but having a longer, shallow sinuation in the middle: seen from

above (fig. 51) the outline is extremely compressed and elongated,

quite four times as long as broad, obtusely pointed in front,

acuminate behind. Surface smooth, slightly setiferous at the

extremities; structure thin and membranaceous t. Length

1-6 mm.

Antepenultimate joint of the antenne (fig. 52) bearing a small

fascicle of setze, two of which are longer than the rest, reaching

beyond the bases of the terminal claws; mandibles (fig. 53) of the

usual form, the biting portion armed with numerous strong teeth ;

maxille (fig. 54) having four slender, elongated lobes, two of the

claws of the second lobe strongly denticulate (fig. 55); terminal

claw of the first foot very long and slender, second foot (fig. 56)

bearing a falcate terminal claw and a very long seta; post-

abdominal rami (fig. 57) slender, bearing two long, strongly

pectinated terminal claws and one very slender seta, and on the

dorsal margin towards the apex armed with a series of about nine

or ten strong spines with smaller sete in the intervals; genital

lobes very similar to those of Herpetocypris and other partheno-

genetic Cypridide.

* The second foot of Cypris ornata, a rare British species, is very similar to this,

and is figured in Plate VII. fig. 39.

+ The specimens, as I received them, had been for a length of time ina formalin

preservative, which may perhaps have led to the disappearance of calcareous

structures.

1904.} ENTOMOSTRACA FROM NATAL, 127

This interesting species seems to occupy an intermediate position

between Herpetocypris and Stenocypris so far as the antennal sete,

at least, are concerned, but from the shell-characters alone one

would undoubtedly assign it to Stenocypris. Professor G. O. Sars in

his description of S'. chevreusxiti mentions the “ partly denticulated ”

spines of the lobes of the first pair of maxille, but does not figure

them, and the antennal sete of that species are certainly much

more fully developed both as to length and number than in

S. perarmata. These points I have been able to confirm from an

examination of British specimens of S. chevreuxit, which I have

been fortunate enough to find in several localities :—very

sparingly in a pond at Pike’s Hill, and in the Hatchet Pond, both

near Lyndhurst, and more recently in considerable quantity in

ditches near the River Arun at Arundel, Sussex. Figures of the

shell, antenna, and post-abdomen of S. chevreuxit are given in

Plate VIII. figs. 58-62.

MACROTHRIX AFFINIS, n. sp. (Plate VIII. figs. 63-65.)

_ Body short, length about one-third greater than the breadth

(fig. 63). Head short and obtuse, with a rounded angle to which

the antennules are attached; dorsal margin gently arched,

obscurely angulated at its junction with the posterior margin

which is finely denticulated, ventral margin convex and fringed

with rather long hairs: antennules (fig. 64) club-shaped, slightly

dilated and truncated distally, crenulated on the outer margin

which bears a few cilia, as also does the truncated extremity.

Post-abdomen (fig. 65) spinulose along the whole of its posterior

border and having a short terminal hooked claw. The antenne

have the normal structure, not presenting any special characters,

nor does the shell exhibit any definite ornament or sculpture.

Length 0°32 mm.

This is a very small species, and is not very unlike the Northern

M. taticornis, though only about half its size.

EXPLANATION OF THE PLATES.

Prats VI.

Cyclops gibsoni, 2, p. 123.

Fig. 1. Seen from above. Fig. 6. Posterior footjaw.

2. Antennule. 7. Foot of first pair.

3. Mandible. 8. Foot of third pair.

4, Maxilla. 9. Foot of fifth pair.

5. Anterior footjaw. 10. Furca.

Cyclops pusillus, 2, p. 122.

Fig. 11. Seen from above. Fig. 15. Anterior footjaw.

12. Antennule. 16. Posterior footjaw.

13. Antenna. 17. Foot of first pair.

14, Mavxilla. 18. Foot of fifth pair.

Ectocyclops rubescens, 2, p. 124.

Fig. 19. Seen from above. || Fig. 20. Antennule,

Fig. 28. Seen from left side.

Fig. 34. Shell seen from left side. i X50

35. Shell seen from above. ,

ON ENTOMOSTRACA FROM NATAL. [May 17,

Prate VII.

Ectocyclops rubescens, 9, p. 124.

Fig. 21. Antenna. Fig. 25. Foot of first pair.

22. Maxilla. 26. Foot of third pair.

23. Anterior footjaw. 27. Foot of fifth pair.

24. Posterior footjaw.

Attheyella natalis, 9, p. 124.

Fig. 31. Foot of first pair.

32. Foot of third pair.

33. Foot of fifth pair.

29. Antennule.

30. Antenna.

Cypris aratra, p. 125.

Fig. 37. The same more highly mag-

nified.

38. Post-abdominal ramus.

36. Extremity of second foot.

Cypris ornata, p. 126.

Fig. 39. Extremity of second foot.

Cypria castanea, p. 125.

Fig. 40. Outline of shell seen fromright | Fig. 41. Outline of shell seen from above.

side. 42. Extremity of second foot.

Prats VIII.

Cypria castanea, p. 125.

Fig. 43. Post-abdominal ramus.

Cypris inermis, p. 126.

Fig. 44. Shell seen from right side. Fig. 47. Antenna.

45, Shell seen from above. 48. Extremity of second foot.

46. Posterior extremity of young 49. Post-abdominal ramus.

shell.

Stenocypris perarmata, p. 126.

Fig. 50. Shell seen from right side. Fig. 55. Maxilla, spiniferous tooth more

51. Shell seen from above. highly magnified.

52. Antenna. 56. Extremity of second foot.

53. Mandible (chewing-end). 57. Apex of post-abdominal ramus.

$4. Maxilla of first pair.

. 58. Shell seen from left side.

59. Shell seen from above.

60. Antenna.

Stenocypris chevreuxii, p. 127.

Fig. 61. Post-abdominal ramus.

62. Group of marginal setz of the

same.

Macrothrix affinis, p. 127.

Fig. 63. Seen from left side. | Fig. 65. Post-abdomen.

64. Antennule.

1904. | DR. A. GUNTHER ON HYBRID PHEASANTS. 129

June 7, 1904.

F. Du Canr Gopmay, Esq., D.C.L., F.R.S., Vice-President,

in the Chair.

The Secretary read the following report on the additions that

had been made to the Menagerie in May 1904 :—

The number of registered additions to the Society’s Menagerie

during the month of May was 276. Of these 102 were acquired

by presentation and 97 by purchase, 8 were born in the Gardens,

61 were received on deposit and 8 in exchange. The number of

departures during the same period, by death and removals,

was 132.

Amongst the additions attention may be called to the following

specimens, all new to the Collection :

1. Three Andaman Banded Crakes (Rallina canningi), pre-

sented by the Government of India on May 2nd.

2. A Yellow-handed Howler (Mycetes beelzebul), received on

deposit on May 13th.

3. An Antilopine Kangaroo (Macropus antilopinus), received

on deposit on May 14th.

4, A Grey Solitaire (/yiadectes wnicolor), received on deposit

on May 30th.

The Secretary reported that he had recently paid a visit to Paris

to inspect two specimens of the Orang-utan (Samia satyrus) which

had been offered for sale to the Society. He stated that they

were very fine animals and fully adult, one of them standing over

five feet in height and the other not much less, the former being

the largest he had ever seen either living or stuffed. Although

they were savage he was able to feed them from the hand with

boiled rice, which constituted their chief food. Owing to his

opinion that they were in indifferent health he did not purchase

them ™*,

Dr. Giinther, F.R.S., Vice-Pres.Z.8., on behalf of the President,

exhibited a series of hybrid Pheasants killed at various times in

the coverts at Woburn, where specimens of many distinct species

had been turned out into the open. He stated that nothing of

a definite nature was known as to their parentage, but proposed

the following determinations tentatively :—

1. Reeves’s Pheasant (Phastanus reevesi) x Common Pheasant.

2. Reeves’s Pheasant x Elliot’s Pheasant (PA. ellioti), An old

female assuming male plumage.

* [I have been informed that both the Orangs died within a few days of my

seeing them.—P. C. M.]|

Proc. Zoou. Soc.—1904, Vor. IT. No. LX. 9

130 SHORT-HORNED BUFFALOES IN THE ANTWERP GARDENS. [June 7,

3. Common Pheasant x Ambherst’s Pheasant (Thaumalea

amherstie). Male.

4, Common Pheasant xX Amherst’s Pheasant. Old female.

5. Cheer Pheasant (Ph. wallichii) x Common Pheasant (? Elliot’s

Pheasant). Male.

6. Cheer Pheasant x Common Pheasant. Male.

7. Silver Pheasant (Gennceus nycthemerus) x Elliot’s Pheasant.

Male.

8. Lineated Kaleege (@. lineatus) x Common Pheasant. Male.

9. Lineated Kaleege x Japanese Pheasant (Ph. versicolor).

Male.

10. Cheer Pheasant x ? Himalayan Monaul (Lophophorus

ampeyanus). Male.

Dr. F. D. Drewitt, F.Z.S., exhibited two fine antlers of the

North-African Red Deer (Cervus elaphus barbarus) and made the

following remarks :—

“These antlers were obtained in the high forest-land of cork,

oak, and cedar, extending far inland on the borders of Tunis and

Algeria.

“One of the most interesting links between the fauna of

North Africa and Europe is this Red Deer living among lions

and panthers.

“‘ Barbary-Deer antlers differ from typical Red-Deer antlers in

having no second tine. This seems a constant characteristic. In

other heads seen in Algeria it was absent. It is absent in a

specimen from Algeria in the Cambridge Museum; also in two

Menagerie specimens in the Natural History Museum (one from

the Gardens and one given by the Duke of Bedford).

“Few Englishmen have seen a wild Barbary Stag. Sir Harry

Johnston is one of the few. He reports that twenty-four years

ago it was fairly common throughout the forest. Now, though

protected by the French Government, it is rare ; forest fires and

poaching Arabs have almost exterminated it—but a few remain.

Fortunately a wild stag among trees, facing its enemy, is some-

times almost invisible at a few feet, the antlers exactly copying

not only the form of a branch but also the bark on it.”

Dr. Drewitt also exhibited a pair of horns (154 inches

in length) of Loder’s Gazelle (@azella leptoceros) from South

Algeria.

Some photographs, sent by Dr. Graham Renshaw, F.Z.S., of a

pair of Short-horned Buffaloes in the Antwerp Zoological Gardens,

were exhibited, and the following note upon them, contributed

by Dr. Renshaw, was read :—

“The difficulty experienced by naturalists in separating the

1904. | ON THE GROWTH OF THE ANTLERS IN THE WAPITT. 131

geographical species or races of the Short-horned African Buffaloes

may render interesting the accompanying photographs, taken by

myself, of a pair of Short-horned Buffaloes which have lived for

many years in the Antwerp Zoological Gardens. Seen sideways,

the horns of both specimens sloped back almost in the plane of the

forehead. Seen from the front, the horns of the bull were broad

and flattened, without transverse markings, and with but a small

development of the boss, which forms a ring-like projection where

the horn springs from ‘the skull. The horns bore considerable

resemblance to those of the Senegambian Buffalo (Los caffer

planiceros), and perhaps the animal is referable to this subspecies.

The horns of the cow, however, were more curved at the tips, and

recalled those of the Lake Tchad Buffalo (Los caffer brachyceros).

Both animals were in the prime of life, of a deep dark blackish-

brown colour; the edges of the ears were heavily fringed with

hair, but there was no hair inside the pinna, A very sparse line

of hair (the individual hairs being about 4 inches long) extended

from the occiput to the root of the tail. The tail was very dark

brown; tail-tuft black. he dorsal line of hair was hardly

noticeable in the cow. The animals were labelled ‘ Bubalus

pumilus, Afr. Australe et Centrale (Buffle nain du Sénégal)’

However, they do not agree at all with Los caffer nanus (Bush-

Cow of the West Coast sportsmen).”

Mr. F. E. Beddard, F.R.S., exhibited and made remarks upon

the skull of the Cape Crowned Crane, Dalearica chrysopelargus,

showing paired lateral and single median bony bosses suggestive

of those of horn-bearing vertebrates.

Mr. R. E. Holding exhibited and made remarks upon a series

of 12 platinotype photographs, three of which are here repro-

duced (text-fig. 24, p. 132), taken by Mr, Henry Irving, of Horley,

showing the growth and development of the horns in the Wapiti

Stag (Cervus canadensis) bred by the Society and now living in

the Gardens. The first of the series was taken on March 20th,

1903, the date on which the horns were shed, and photographs

were taken at fortnightly intervals until the following September,

when the horns were fully adult and almost free from the velvet

covering.

The photographs showed in an interesting manner the rapid

growth of the antlers in the Cervide, the Pe occupied being

six months. On March 20th, 1904, these horns were in the usual

course shed, the weight of the pair being 21 lbs. and length of

beam 453 inches.

Simultaneously with the development of the horns, the photo-

graphs showed the rapid seasonal change in the coat of the

animal, the winter coat beginning to fall off shortly after the

132 ON THE GROWTH OF THE ANTLERS IN THE WAPITI. [June7,

Text-fig. 24.

< Sin?

Antlers of Wapiti, from photographs.

(1904. ] ON THE YOUNG OF THE EGYPTIAN FAT-TAILED GERBILLE. 133

horns were shed. In five weeks the process was complete and

the animal was in its summer coat.

Explanation of Text-fig. 24.

A. Head of Stag, taken April 17th, 1903, three weeks and four days after the horns

were shed. Length of horns about 5 inches. Extremely vascular and sensitive,

the skin of the forehead merging into the “velvety ’’ covering; the burr (e) being

indistinct, the bifurcation between frontal tine (a) and bez tine (6) being just

apparent.

B. Head of same Stag, taken May 14th, viz. three weeks and four days afterwards.

Here a rapid development is seen to have taken place; the frontal tine (a),

bez tine (6), and beam (ce) being well divided, a small supernumerary point (x )

making its appearance at the base of the left horn. Horns still vascular

and warm, and covered with thick epidermis or “ velvet ”’—the burr (e) being

more definite.

C. The same head, taken June 25th, five weeks and four days afterwards. Within

this period the most marked development takes place; the frontal (a), bez (6),

and trez (c) tines being fully developed, and the apex of the beam (d) splitting

up into the characteristic three sur-royals. ‘The velvety covermg is now

shrinking, and the nutrient blood-vessels can be discerned; the burr (e) is

now quite prominent.

On July 24th the horns, though still covered with velvet, are quite adult.

During August the velvet begins to peel off, and by the first week in September

the horns are usually clear.

Mr. Holding also exhibited a fine pair of antlers of the Irish

Red Deer (Cervus elaphus) having 14 points, which had been sent

to him by Sir Douglas Brooke, Bart., F.Z.8., Co. Fermanagh.

They weighed 8 lbs. and were 37 inches in length.

Mr. R. I. Pocock, the Superintendent of the Gardens, exhibited

living specimens of hairless varieties of the Common House-Mouse

(Mus musculus) and Brown Rat (JZ. decwumanus), and remarked

that the available evidence seemed to prove that the abnormality

was always correlated with a wrinkled skin. The living specimens

exhibited further suggested that it was accompanied by weakness

of the eyes, and in the case of the Mouse by the absence, or at all

events diminution, of the smell so characteristic of that animal.

Mr. Pocock also exhibited four young examples, between five

and six weeks old, of the Egyptian Fat-tailed Gerbille (Pachy-

uromys duprest), which were born in the Menagerie and reared by

hand, owing to the death of the mother when the young were only

six days old. One of the young had never opened its eyes and

would probably be permanently blind. There were five young

in the brood, but one had died. Mr. Pocock also showed a

154 LT.-COL. J. MALCOLM FAWCETT ON [June 7,

newly-born example of this species, which was naked and blind

and scarcely distinguishable from the young of the Common Rat.

In this respect the young differed markedly from those of the

Egyptian Spiny Mouse (Acomys), which were born with the eyes

open and the skin covered with hair, the brood consisting of but

two individuals only. This fact further supported the view that

the condition of the young at birth could not be taken as a criterion

of attinity in the case of the Rodentia.

The following papers were read :—

1. On some New and Little-known Butterflies, mainly from

high elevations in the N.E. Himalayas. By Lt.-Col.

J. Matcomm Fawcnrtt*.

[Received April 9, 1904. }

(Plate [X.7)

[The complete account of the new species described in this communication

appears here, but as the names and preliminary diagnoses were published in the

‘ Abstract,’ such species are distinguished by the name being underlined.—E 170k. |

The main object of this paper is to describe some species which

were taken by the native collectors of Mr. J. C. White, British

Commissioner in Sikkim, at high elevations.in the N.E. Himalayas,

on the borders of Thibet. I have been unable to identify them

with any species in the Collection of the British Museum, or in

the Rothschild Collection at Tring, and they were subsequently

sent to Mr. H. J. Elwes, who is unacquainted with them.

Under these circumstances it appears to me necessary that

they should receive names, although future investigation may

perhaps demonstrate some of them to be geographical forms

of species already described. It also appears essential that the

three species of Colias which have been taken at high elevations

on the Sikkim-Thibet Frontier should be figured together; and

for this reason a figure and description of C. eogene var. leecha

(Gr.-Gr.), of wich a figure does not appear to have been hitherto

published, have been added to the figures and descriptions of the

two new species.

At the same time that the above-mentioned species were taken,

there were also four species of the genus Parnassius captured on

the Sikkim-Thibet Frontier: one is clearly P. imperator (described

by Herr Friihstorfer as P. imperator augustus); a second is

P. epaphus var. sikkimensis Elwes ; and the other two have been

* Communicated by the SecRETARY.

+ For explanation of the Plate see p. 141.

ees 1904 oleae

£.C Knight del.et Lth. Mintern Bros imp

BUTTE Re Lib S Ove Dae Nhe eiIMALAVAS.

1904. ] BUTTERFLIES FROM THE N.E. HIMALAYAS. 135

identified by Mr. Elwes as varieties of P. acco and P. delphius

respectively ; the two latter were described as P. acconus $ and

Q by Herr Friihstorfer from the writer’s figures.

A species of Ragadia, from the Battak Mts. in Sumatra, has

been added to the paper, and is here figured for the first time.

Family SAryrina.

1. RaGADIA sIMPLEX Fawcett. (Plate IX. fig. 1.)

Ragadia simplex Fawcett, Ann. & Mag. Nat. Hist. ser. 6,

vol. xx. p. 111 (1897).

Eixpanse 1 in. 6 lin.

Habitat. Battak Mountains, Sumatra, June 1896.

Description.— Vale. Upper side pale fuliginous brown, crossed

by two darker fuscous bands running parallel to one another from

the costa of the fore wing to the inner margin of the hind wing,

the outer being much broadened on the hind wing. There is also

a shorter band proximal to the body, which, starting parallel to

the others on the fore-wing costa, ends at the origin of the sub-

median on the fore wing. From the apex of the fore wing to

the inner angle of the hind wing runs a submarginal row of

small, indistinct, fuscous spots, one in each interspace. The wing-

margins are narrowly fuscous, the colour being bounded by a

narrow submarginal band looped on the fore wing and on the

hind wing parallel to the margin. Under side similar in markings

to the upper, but the ground-colour pale buff and the row of spots

silver instead of fuscous.

The antenne, head, thorax, abdomen, and legs like those of

f. crisia Hiibner, from which this species differs in its generally

pale ground-colour, and in the ocellate band of erisia being

reduced to mere pupils of silver below and fuscous above.

Family NyMPHALINA.

2. MELITHA TIBETANA. (Plate IX. fig. 2.)

Melitea tibetana Fawcett, Abstr. P. Z.S. 1904, No. 9, p. 8,

June 14.

Expanse 1 in. 2 lin.

Habitat. Khamba Jong, Thibet, 15,000 ft. elevation.

Description.— Upper side: both wings pale dull ferruginous,

apex of fore wing pale ochreous; costa and base of wings and

anal margin of hind wing dark fuscous. Fore wing with two

black marks in cell and one at its apex; base of cell inside the

first mark, and between the second and third marks, pale

ochreous ; two transverse series of black discal spots, the inner

series highly angled outwardly above the third median nervule,

enclosing between them a pale ochreous fascia; a marginal series

of pale ochreous lunules inwardly defined with fuscous and

136 LT.-COL. J. MALCOLM FAWCETT ON _ [June 7,

outwardly by a fuscous marginal line, and a pale ochreous lunule

basally between the submedian nervure and first median nervule;

cilia whitish. Hind wing with the marginal fuscous line and

submarginal lunules as in fore wing; the transverse series of

black discal spots and ochreous fascia as in fore wing, and two black

marks in discoidal cell. Under side: fore wing very pale ferru-

ginous, fading into sordid white at the apex and along the costal

margin ; discal markings of upper side obsolescent except near

the apex. Hind wing buff, with a subbasal band, a spot in the

cell, an angulated discal band, and a lunulated submarginal band

all white, defined outwardly by black atoms and fine lines, the

cell and subbasal band surrounded by ferruginous spots.

This species is probably a local race of the variety of W/. sindura

Moore described by Mr. H. J. Elwes in Trans. Ent. Soc. Lond.

1888, pt. i. p. 336, pl. x. figs. 5 & 6, which = MW. sikkimensis

Moore, but differs from it in its much paler coloration, and the

presente on the upper side of the pale ochreous bands and spots

above mentioned, and the more prominent fuscous markings.

Five specimens from Khamba Jong in my possession present

no variation from the above description. J/. sindura var. Elwes,

isan uniformly bright ferruginous insect with no pale fascize or

spots; and I possess six specimens of it from 17,000 ft. in Native

Sikkim, as also six specimens of a larger and darker form from

10,000 ft. in Sikkim interior, in which the submarginal lunules

are obsolescent.

3. ARGYNNIS CLAUDIA. (Plate IX. fig. 3.)

Argynnis claudia Fawcett, Abstr. P. Z. 8. 1904, No. 9, p. 8,

June 14,

Expanse 1 in. 6 lin.

Habitat. Khamba Jong, Thibet, 15,000 ft. elevation.

Drscription.— Upper side rich fulvous (pale fulvous in some

specimens), the base of fore wing narrowly and that of hind wing

broadly irrorated with black atoms. Fore wing with three curved

streaks in the cell, and one at its apex, thicker than the remainder ;

two transverse discal series of black spots, the imner one very

irregular and highly angled outwardly above the third median

nervule; a series of submarginal fulvous cuneiform lunules,

defined inwardiy by a thick black line and outwardly by a thick

black band; cilia white. Hind wing with the submarginal

lunules and transverse series of spots as in the fore wing, except

that the outer series is reduced to four spots (2 subapical and

2 subanal) in some specimens, and to three in others, the remaining

spots being obsolete. Under side: fore wing similar to upper

side but paler, the black marks and spots smaller, and the outer

transverse series of spots obsolete ; apex green; marginal series of

lunules silvery and two silver subapical spots below the costa; no

marginal lines. Hind wing dark green, with a slight bronzy

suffusion ; two basal silver spots below the costa, a linear silver

1904. ] BUTTERFLIES FROM THE N.E. HIMALAYAS. 137

spot defining the apex of the cell and a linear one near the inner

margin ; the space between the two transverse discal rows of black

spots (of the upper side) occupied by a series of linear silver streaks

between the nervules; a submarginal series of triangular silver

spots, much produced ‘inwardly, and between these two series of

silver spots and streaks a faint indication of a submarginal line of

yellow spots ; no marginal lines.

This species is nearly allied to A. clara Blanchard, but differs

from it in its rounder wings, smaller size, and paler pallone On

upper side, outer mar: gins of both wings broadly fuscous, contain-

ing a submarginal series of pale spots, without the submarginal

and mar ginal fine black lines beyond them as in A. clara; the

inner transverse series of discal spots angled outwardly above

3rd median nervule on both wings, and the outer discal series of

black spots is reduced, on the hind wing, to 4 and sometimes

3 spots, the spot on the lower discoidal interspace being always

absent. Under side: the fulvous colour of fore wing is paler and

redder, and the green colour of apex of fore wing and hind wing

is duller and darker ; bronzy suffusion less, and extends to the

margin without the mar ginal fulvous border of 4. clara. The

only variation from the above deser iption in five specimens from

Khamba Jong is that one specimen is more melanic, while another

hasa paler gr Gandleoloue | ‘The aeny, a, var. clarina of Staudinger

(Cat. Lep. Pal. p. 38) cannot be referred to this species, as he says

nothing about his species being smaller than A. clara, and describes

it as having the black spots smaller, whereas this species is more

melanic than A. clara.

Family Lycanipm.

4. Lycmna ArtanA Moore, var. ARENE. (Plate IX. fig. 4.)

Lycena ariana Moore, var, arene Fawcett, Abstr. P. Z.8. 1904,

No. 9, p. 8, June 14.

Expanse | in. 3 lin.

Habitat. Khamba Jong, Thibet, 15,000 ft. elevation, July.

Description.— Male. Upper side: both wings bright cyaneous

blue, margins narrowly black, extending slightly up the nervules ;

cilia broadly white. Under side: both wings purple-grey ; hind

wing suffused with metallic greenish grey at base and on inner

margin. Fore wing: apex of cell defined by a narrow black

streak outwardly surrounded by white; a discal series of five

minute black spots ringed with white, the subcostal spot

evanescent; a marginal series of white spots with minute black

centres, defined inwardly by obsolescent traces of orange lunules.

Hind wing with the marginal white spots larger, and the orange

lunules defining them inwardly more distinct, that at anal angle

geminate ; a subcostal black spot, and a discal series of minute

black spots ringed with white, varymg in number from two to

three, and entirely obsolete in some specimens (this is the form

138 LT.-COL, J. MALCOLM FAWCETT ON [June 7,

figured) ; a thin black streak at apex of cell, in a large cuneiform

white spot, and opposite this spot another large cuneiform white

spot, with its base on the centre of the line of orange submarginal

lunules,

This species differs from Z. ariana Moore from the N.W.

Himalayas in being smaller, and in the discal row of black spots

on the under side being minute, fewer in number, and obsolete, in

some specimens, in the hind wing; and in the discal white spots

at the apex of cell of hind wing being larger.

. Lycmna puEeretes Hiibner, var. praris. (Plate IX. figs.5 g,

5a@.)

Lycena pheretes Hiibner, var. pharis Fawcett, Abstr. P.Z.S8.

1904, No. 9, p. 8, June 14.

Expanse 1 in.

Habitat. Khamba Jong, Thibet, 15,000 ft. elevation, July.

Descriprion.— Vale. Upper side dark purple-blue; fore wing

with a black marginal line; margin of hind wing more broadly

black, especially at apex and costa; cilia broadly white. Under

side: fore wing purple-grey, paling towards apex; a black spot

ringed with white at apex of cell; a discal series of small black

spots ringed with white, varying in number ; in some specimens

only three, usually four. Hind wing pale brownish on the disk,

fading into pale greenish-ochreous at the margin, with the follow-

ing pale ochreous spots: one in and extending beyond discoidal

cell, cuneiform; a discal series of six beyond it, and an indistinct

one basally below the costa.

Female. Upper side dark fuscous, basally irrorated with blue ;

under side as in male.

This form differs from the var. asiatica of Elwes in having

fewer discal black spots on fore wing, and in the size and length

of the pale cellular spot on hind wing.

6. Lycmna (Zrzpra) zERA. (Plate IX. fig. 6.)

Lycena (Zizera) zera Fawcett, Abstr. P. Z.8. 1904, No.9, p.9,

June 14.

Expanse | in. 11 ln.

Habitat. Tounghoo, Burma, June.

Descrietion.—Male. Upper side shining violet ; fore wing with

a pale spot at the apex of the discoidal cell; a broad fuseous

marginal band extending slightly up the nervules, but the inner

border regular and bending inwards on the costa. Hind wing

with the marginal fuscous band broad at the apex, narrowing to

a black marginal line, inside which are placed five fuscous lunules ;

cilia broadly white. Under side pale grey: fore wing with a

fuscous streak, ringed with white, defining apex of cell; a discal

row of six fuscous spots ringed with white; a row of marginal

spots faintly defined imwardly by fuscous Junules. Hind wing:

discal row and a marginal series of fuscous spots as in fore wing ;

1904. | BUTTERFLIES FROM THE N.E. HIMALAYAS. 139

three subbasal black spots ringed with white, the centre spot

inside the cell,

This species is nearly allied to 7. marginata from China, but

differs in the discal row of fuscous spots of the fore wing being

fainter and more regulary on the under side, and the black border

of the upper side being more even in its inner edge and bending

inwards on the costa,

7. Lycana (Nipuanpa) MAncta, (Plate IX. fig. 7.)

Lycona (Niphanda) marcia Bawceth, Abstr. P.Z.8. 1904, No. 9,

p. 9, June 14.

Expanse 1 in, 2 lin.

Habitat. Vounghoo, Burma, June.

Descriprion.— Male. Upper side shining violet, with the spots

of the underside showing up indistinctly. Fore wing with a

narrow fuscous band on the costa and outer margin; hind wing :

costa, apex, and abdominal margin broadly fuscous, and a row of

fuscous lunules on the outer margin ; cilia white between the ends

of the nervules. Under side pale violaceous grey, with the follow-

ing brown marks ringed with white:—fore wing with a short basal

streak below the costal nevvure; two spots, one above the other,

and the lower one the larger, in centre of cell, and below the

lower spot a large triangular space between the first median

nervule and the sabmiedian nervure; a spot defining the end of

the cell; a discal fascia of seven spots, interrupted in the middle,

and with the outer edge of the upper four spots defined outwardly

by fuscous; a pale and indistinct submarginal and marginal

lunular fascia, the lunules of which are largest and most defined

between the median nervules, Hind wing with a blackish-brown

spot at base of cell, two black spots below costal margin, and a

similar spot below the outer one; an irregular row of pale discal

spots, and a marginal row of white lunules, the first three and

the sixth from the anal angle with black centres.

This species is probably a local race of L. (Niphanda) tessellata

Moore, from Penang, but differs from it in its much less heavy

markings on the under side. There is also a specimen (at present

unnamed) in the British Museum which agrees with the specimen

here described,

Family Prerina.

8. CoLiAs BeRYLLA. (Plate IX. fig. 8.)

Colias berylla Fawcett, Abstr. P. Z.8. 1904, No. 9, p. 9,

June 14,

Expanse 2 in. 9 lin,

Habitat. Khamba Jong, Thibet, 15,000 ft. elevation.

Descriprion.—Lemale, Upper side : fore wing pale lemon-

yellow, base and costal margin broadly irrorated with black

atoms; a diamond-shaped black spot at apex of cell; hind or

140 LT.-COL. J. MALCOLM FAWCETT ON [June 7,

outer margin broadly black, irregularly dentate on nervules on its

inner edge, with the following sulphur-yellow spots, viz., five

forming a curved row near the apex from costa to 3rd median

nervule, the lower two being larger than the upper three; also

one spot between the Ist and 2nd median nervules, and one

between the Ist median nervule and the submedian nervure; cilia

of both wings whitish with a pink suffusion. Hind wing black,

basally thickly irrorated with sulphur atoms; an orange spot at

end of cell; anal and outer margin sulphur-yellow, which forms

into a line of lunular spots on outer margin. Body black, head

and antenne pink. Under side: fore wing —disk pale yellow ;

apex and outer margin grass-green, with a submarginal line of

pale yellow spots; a black spot at apex of cell, and three black

discal spots in a row between the median nervules and submedian

nervure. Hind wing grass-green, paler on outer margin, and

irrorated with black atoms; a silver spot surrounded by a pink

area at the apex of the cell.

This species belongs to the Ayale section of the genus Colias,

and is nearest to C. erschoffi Alphéraky, from the Tian Shan Mts.

9. Contas NINA. (Plate IX. fig. 9.)

Coltas nina Fawcett, Abstr. P. Z. 8. 1904, no. 9, p.9, June 14.

Expanse 2 in. 9 lin.

Habitat. Khamba Jong, Thibet, 15,000 ft. elevation.

Descriprion.—lemale. Upper side: fore wing pale sulphur-

yellow, the disc below the median nervure and 3rd median nervule

orange; a black diamond- shaped spot at apex of cell; a broad

black outer marginal band, inwardly irregularly dentate on the

nervules, irrorated with sulphur- yellow scales outwar dly, and a

submar. einal row of ill-defined sulphur spots between the nervules,

that between the 2nd and 3rd median nervules being obsolete.

Hind wing black, irrorated with pale sulphur atoms, most thickly

towards the base and outer mar gin; discoidal cell pale yellow,

with an orange spot at its apex; a submarginal row of pale

sulphur hastate spots; antenne and cilia pinkish. Under side:

fore wing—disk as on upper side; apex and outer margin pale

green, irrorated with black atoms; a black spot at apex of cell,

and three or four ill-defined black spots discally between the

nervules. Hind wing pale green, with spots as on upper side.

This species appears to be a transition between the hyale and

edusa sections of Colias.

10. Conzas EocENE Felder, var. LEEcHI Gr.-Gr. (Plate IX.

figs, 10 g¢, 10a 2.)

C. eogene Felder, Reise Novara, p. 196, t. 27. £. 7, ¢ (1865).

Var. leechi Grum- Grshimailo, Hore Soc. Ent. Ross. xxvi.

p. 382 (1893).

Expanse | in. 4 lin.

Habitat. Native Sikkim, 19,000 ft., July.

1904. | BUTTERFLIES FROM THE N.E, HIMALAYAS. 14]

Descrrprion.— Male. Upper side orange, base narrowly irrorated

with black atoms; a black diamond-shaped spot at apex of cell;

a fuscous marginal band, with a row of pale yellow hastate spots

between the nervules. ‘Hind wing orange ; inner margin green,

irrorated with black atoms ; a paler spot in cell, anda submarginal

row of pale yellow hastate spots; apex and outer margin fuscous.

Under side : fore wing—disk paler than upper side; apex and costa

grass-green, lrrorated with black atoms; a black spot at apex of

cell, and a submarginal row of black spots, ill-defined towards the

apex, and beyond them a row of pale yellow spots. Hind wing

grass-green ; a pale sulphur submarginal fascia, irrorated with

dark green atoms; a silver spot in a reddish area at end of cell.

Head, antennze, and cilia pink.

female. Upper side: fore wing orange, paling on the costa to

yellow; base and median nervules linaeadl with black atoms ;

marginal band and spots as in male. Hind wing black; cell

yellow, with an orange spot at its apex; a marginal band of

sordid-sulphur lunulate spots. Head, antenne, and cilia as in

male.

The specimens in my possession have been identified by Mr. H.

J. Elwes as var. leechi. Herr Frithstorfer erroneously described

two of my figures as representing a new species, under the name

of C. eogene, subsp. miranda (Insekten Borse, xx. 19. 148, May

1903). These figures are introduced into this paper because

this variety does not appear to have been figured previously, and

with a view to complete the series of forms of Colias from the

borders of Sikkim and Thibet.

Since these notes were written I have seen a paper published

by Mr. H. J. Elwes in ‘Iris,’ 1904 (pp. 388 e¢ segq.), on the

subject of the descriptions and figures above referred to. In this

paper Mr. Elwes refers to this species of Colias as “‘stoliczkana,”

although he had previously written “ leechi” below the figures I

submitted to him for identification. The male of this species,

however, differs from the figure of the male of C. stoliczkana

in the British Museum. The var. stoliczkana has hitherto been

recorded only from Ladak, Fergana, and Amdo; while the var.

leecht has been recorded only from China.

EXPLANATION OF PLATE IX.

Fig.1. Ragadia simplex Fawcett, p. 135.

2. Melitea tibetana Fawcett, p. 135.

3. Argynnis claudia Fawcett, p. 136.

4. Lycena ariana Moore, var. arene Fawcett, p. 137.

5. Lycena pheretes Hiibner, var. pharis Paw cett, 6, p. 138.

5a. Ditto, é

6. Lycena (Zizera) zera Fawcett, p. 138.

7. Lycena (Niphanda) marcia Fawcett, p. 139.

8. Colias berylla Fawcett, p. 139.

9. Colias nina Fawcett, p. 140.

10. Colias eogene Felder, var. leechi Gr.-Gr., 3, p. 140.

10a. Ditto, 2.

142 DR. A. G. BUTLER ON SEASONAL [June 7,

2. On Seasonal Phases in Butterflies.

By Ay G.Burunr, Pho Dy WS: eZ Sues

[Received May 10, 1904.]

The fact that Butterflies emerged from the pupa in markedly

different forms at different times of the year was made evident

many years since by the labours of W. H. Edwards in the United

States, by Mansel Weale in South Africa, and by Doherty and

de Nicéville in India; but this fact was misunderstood, and there-

fore not fully accepted by many workers for years afterwards,

whilst not a few are sceptical as to its truth at the present day.

One of the chief reasons for this scepticism is based upon the

unquestionable truth that the dry-season type of a species not

unfrequently emerges in the wet season and vice versa. That I

was myself rendered sceptical for years on this ground will be

seen by referring to some of my earlier papers in which the

question of seasonal forms had to be considered.

In a paper published in 1884 (P. Z. 8. pp. 478-501) I recorded

the remarkable fact that, at Aden, Limnas chrysippus, Hypolimnas

misippus, and Catopsilia florella produced all their varietal phases

simultaneously, and that Zeracolus “nouna” = saxeus (which

actually is the 7’. evagore of Klug) occurred in March, April, and

May, whilst 7. yerburit was also obtained commonly in April

and May; but I did not then fully appreciate the fact that all

these were instances of the simultaneous emergence of phases

characteristic of seasons and climates, and that they represented

the probable condition of all very variable types before seasonal

or climatic changes had begun to act upon them.

As with protective mimicry, the more enthusiastic exponents

of which have frequently erred in supposing that because this

was of use against one enemy, it must necessarily be against all;

so has it been with those who desired to believe in, but failed to

comprehend, seasonal variation. That I misunderstood it myself

in 1886 is clear from the remarks which I made in a paper upon

Lepidoptera from Western India (P. Z. 8. p. 399) respecting the

broods of Belenois mesentina: in 1888 I was no wiser, as my

remarks emphasising the importance of dates of capture in the

case of certain species of Zeracolus clearly show (Ann. & Mag.

Nat. Hist. ser. 6, vol. i. p. 201).

In 1895 (P. Z. 8. p. 727) I hinted at the possibility of Zypan-

artia scheneia and H. hippomene being seasonal forms of the

same species, and in 1896 (P. Z.8. p. 112) I considered this

probable; yet later in that year (P. Z.8. p. 285) I concluded

that this was an error, because both were captured on the same

mountain upon two successive days. At this date, therefore, it

is quite evident that I considered it impossible for wet and dry

phases of any species to occur simultaneously. Indeed, it was

only after reflecting upon the probable identity of 7. yerburw

with the supposed 7’. nowna (P. Z. 8. 1896, p. 247) that I began

1904. ] PHASES IN BUTTERFLIES. 143

to have some idea that because a type of pattern and coloration

was characteristic of a particular season or climate, it did not

necessarily exclude other types: therefore that it was not im-

possible for phases characteristic both of dry and wet seasons or

climates to be sometimes found flying together; that in a very

dry country like Aden it was the rule rather than the exception

for wet, intermediate, and dry phases of a species to occur

commonly together in each brood.

That this polymorphic character was probably of earlier date

than the more or less defined seasonal phases, of such countries

as exhibit great variations of weather at different seasons, seemed

evident to me from the fact that in very moist countries the

extreme dry phase of species is exceedingly rare, and probably

near to extinction. In Precis sesamus, the dry phase of P. natal-

ensis (=calescens) from Southern and Hastern Africa, the seasonal

phases are very distinct, but about equally abundant. In the

wet season, as pointed out by me (P. Z.8. 1898, p. 904), both forms

may be taken flying together in Mashunaland; and on that

ground I proposed that the term “seasonal form” should be

rejected, and the term ‘seasonal phase” substituted *. On the

West Coast P. calescens or natalensis is represented by P. octavia

of Cramer and a number of intermediate phases, but no extreme

dry phase was recorded until 1901, when, im my “ Revision of the

Butterflies of the genus Precis” (Ann. & Mag. Nat. Hist. ser. 7,

vol. vill. p. 205), I mentioned an imperfect example indis-

tinguishable from typical P. sesamus as having been received

from Onitsha on the Niger. The extreme rarity of this phase on

the West Coast, and the probability that it has become absolutely

extinct at Sierra Leone, seem to indicate that it is unsuited to

the conditions of a moist climate; whilst the numerous intergrades

from the dry to the wet phase on the same coast certainly indicate

the transition from fixed varieties, such as obtain where seasons

are well defined, towards a more or less wet type. In Southern

and Hastern Africa intergrades between P. sesamus and P. natal-

ensis are extremely rare, the most striking of such intergrades

being figured by me in 1900 (P. Z. 8. pl. lviii. fig. 1).

As it is by no means rare for individuals of the wet phase of a

species to emerge from the pupa in the dry season, there is no

reason why Lepidopterists should be startled when this occurs.

They should bear in mind the probability of the fact that, as all

the phases of some species occur as simple varieties in extremely

dry countries, they also formerly existed as varieties in other

species; that the latter, as they gradually extended their range,

were subjected to widely different conditions; that then the

summer phase (as we now understand it) was so conspicuous

in the winter, and the winter phase so conspicuous in the summer,

that their chance of survival at the unsuitable season was

lessened ; and thus it came about in course of time that one

variety of the species became the prevalent wet phase, and

* See also P. Z. S. 1900, p. 916.

144 CAPT. R, CRAWSHAY ON THE PREY or THE Lion. [June7,

another the prevalent dry phase. But it is absurd to assume

that the evolution of these seasonal phases is already perfect and

complete over the whole globe, and to speak of the occurrence of

wet and dry phases at any season as “ very remarkable”; as a

matter of fact, it would be more remarkable if they never did

occur simultaneously.

3. The Prey of the Lion.

By Capt. Ricoarp Crawsuay, F.Z.S.

[Received June 3, 1904. ]

Prevailing opinion is so often at variance with my own ex-

perience on the question of the prey of the Lion, that I venture

to offer a few remarks on those experiences covering a period

of many years’ travel and residence in Central Africa. It is

generally believed that the Lion is a fastidious feeder, eating only

what he has himself killed in the shape of the larger mammals.

I have not found the Lion by any means particular in confining

himself to his own kills, or in disposing of carrion when in

tolerably good condition.

As to his prey being only the larger mammals, this is not in-

variably the case, for Iam able to establish one instance at least

of his preying on the Porcupine. Mr. Selous has it on record

that a Lion has eaten the skin of a Sable Antelope preserved with

arsenical soap as a natural history specimen. Nothing as re-

markable as this has occurred within my own knowledge, but I

have known a great many instances of Lions feeding on carrion

of my own killing, such as the carcases of Elephants on several

occasions.

The main purpose of my note is to exhibit two Porcupine-

quills taken from the left fore-paw of a Lion—the skull, skin, and

claws of which I also exhibit. This Lion was shot by myself

two days’ journey N.W. of Kibwezi, Hast Africa Protectorate, in

March 1898. Three quills were found in the paw, one of which

I regret has since been lost. The two exhibited measure each

approximately 1 inch in length. The cartilage in which the quills

were embedded showed no inflammation, so it can be inferred

that these at first most painful inflictions must have been there

for a considerable time.

There is no reason for supposing this Lion killed the Poreupine

under the impulse of hunger, because the country teemed with

game of all kinds—from rhinoceros, zebra, and ostriches, to vast

numbers of antelope, large and small.

My Ahenga, when I questioned them as to whether they knew

other instances of Lions preying on such small game as Poreupines,

seemed amused, and replied :—‘ Oh, indeed! even field rats:

they tear them out of the ground with their claws”!

The larger mammals, no doubt, form the Lion’s chief prey—

Buffalo more especially, when these abounded; but, as can be

seen, there are times when even small rodents are in requisition.

1904. | ON THE ‘* BREPHOS” IN A SKINK,. 145

4. Note on an apparently Abnormal Position of the

“ Brephos”’ within the Body of a Skink (Chaledes

lineatus). By Frank H. Bepparp, M.A., F.R.S.,

Prosector 40 the Society.

[Received May 11, 1904. |

(Text-figure 25.)

Among a number of examples of the Lizard Chalcides lineatus

which arrived in the Society's Menagerie on May 3rd, one was

found to be dead on its arrival, On opening the Lizard, which

proved to be a gravid female, a possible cause of death was at

once obvious. The reptile presented a case of what may be

called extra-uterine pregnancy, and may have been unable to get

rid of the two young ones. That is, at any rate, one way—and

perhaps the most probable way—of interpreting the following

facts. *

The Lizard itself measured 12 inches in length, including the

tail, and the young ones were of the same size, and measured

within a very few millimetres of 35 inches*. I did not attempt

a more exact measurement, as they were somewhat twisted. The

yolk-sac was not fully absorbed, and formed a string-like appendix

of about 3 inch in length, In other respects these foetuses were

exactly like the parent and obviously, at the very least, ready to

be born. Neither brephos was within the oviduct. Both lay, in

fact, apparently in the body-cavity. The position of one of them

is shown in the accompanying drawing (text-fig. 25, p. 146), which

was made before the relations of surrounding organs were much

disturbed. Its head les not far from the head of the parent ;

the end of the tail is nearly on a level with the posterior extremity

of the liver. The second brephos lay much further back, but I

cannot give exact details, as 1t was liberated on cutting open the

body-wall of the parent. Both young ones lay in a slight spiral

curve like the popular representations of asnake. There are some

other facts of interest in connection with the abnormal situation of

the two young Chalcides. The ovaries contained numerous round

white eggs, the largest of which did not measure more than 2 min.

in diameter. I presume, therefore, that they are not nearly ready

for fertilisation. The two oviducts were in a condition corre-

sponding to that of the two ovaries. They showed no signs what-

ever of having recently contained eges or embryos. Nor, on the

other hand, was there anything in the least abnormal about them

or their drawn-out peritoneal orifices. They were quite un

injured by my dissections. It seems to me, therefore, to be

indisputable that at least a great part of the development of the

two embryos must have taken place outside of the oviduct or

oviducts. And it is remarkable that this took so long in

producing an injury grave enough to be fatal to the mother.

Indeed, there were no signs whatever of pathological conditions

* The Scincidie are, at least for the most part, ovoviviparous.

Proc. Zoot. Soc.—1904, Vou. II. No. X. 10

146 ON THE ‘ BREPHOS” IN A SKINK. [June 7,

in the organs neighbouring upon the two young ones ; and one 1s

thereby tempted to speculate as to how far this—as would be

supposed by most—abnormal state of affairs is really abnormal.

Such cases of extra oviducal feetation may be in part responsible

Dissection of Chalcides lineatus.

A, brephos; Z., lung; Zi., liver; St, stomach; Od., oviduct.

for some of the legends concerning the swallowing of their young

by various reptiles for protection’s sake. In the present instance,

for example, as may be seen from the drawing exhibited, an

unskilled observer opening the body of the parent and seeing the

1904. | ON THE VISCERAL ANATOMY OF PELAGIC SERPENTS. 147

head of the brephos lying apparently within the gullet, and at

any rate anteriorly in the body, might arrive at a conclusion

opposed to the real facts which more accurate observation reveals.

One rather important piece of evidence is commonly omitted in

those cases of alleged swallowing of the young, which I refer to

here for other reasons, and only incidentally as concerned with

popular beliefs. Each brephos has not only the small white

vesicle adherent to the body already referred to, but considerable

vestiges of the other embryonic sacs invest and are attached to it.

These are distinguishable by their grey colour, and. are com-

paratively bulky. In the case of one brephos, they are attached

to it; in the case of the other, the young one came away from the

membranes, which were found to be attached to the mesenteries,

and possibly in process of being absorbed. Any suggestion of

protection by the mother within her body of actually born young

would of course be negatived by these facts.

5. Contributions to the Knowledge of the Visceral Anatomy

of the Pelagie Serpents Hydrus platyurus and Platyurus

colubrinus. By Frank E. Bepparpb, M.A., F.R.S.,

Prosector to the Society.

(Received May 20, 1904. |

(Text-figures 26-28.)

Tn the comprehensive works of Milne-Edwards* and Meckel 7

and others, there are numerous references to various points in the

anatomy of the pelagic Ophidia, while a more particular account

of the viscera of one species has been given by Cantor. The

lungs have been particularly dealt with, though very briefly, by

Cope §; while Mi. G. W. Butler || has incorporated remarks upon

some of these snakes into his general papers upon the asymmetry

of the Ophidian lung. Both the last-mentioned papers contain

references to previous literature.

I have had the opportunity of dissecting one example each of

the marine snakes Hydrus platyurus and Platyurus colubrinus

which have been in my possession for some time, the latter

specimen belonging to me, the former to the Prosector’s Stores.

This dissection enables me to add something to our knowledge of

the anatomy of the Hydrophiine, and to compare two quite dis-

tinct generic types. I had not expected to find them so different

as dissection showed them to be.

(1) Platyurus colubrinus.

The specimen of this snake which I dissected measured in all

nearly 17 inches,.of which a little over two belong to the tail.

* “Tecons sur la Physiologie et Anatomie comparée.’

+ ‘Anatomie comparée.’ French Transl.

< “© Observations upon Pelagic Serpents,” Trans. Zool. Soc. 11. p. 303.

§ “On the Lungs of the Ophidia,” Proc. Am. Phi]. Soc. xxxiii. 1894, p. 217.

|| In P. Z. S. 1892, p. 477, and P. Z. 8. 1896, p. 691.

10*

148 MR. F. E. BEDDARD ON THE VISCERAL [June 7,

The location of the different organs of the body and the lengths:

of some of them in relation to the total length of the body differ

in this serpent from the corresponding situation and measurements,

of Hydrus platyurus, which are placed side by side for the pur

poses of easier comparison.

P. colubrinus. — H, platyurus.

Length of body to vent ...... 14 inches. 23 inches.

From tip of snout to base ot

heart. Pelkey ee oereee 53 inches. 5 inches.

From tip of snout to anterior

Gil GE IMEI Lodcoscssescsco006 5% inches. 6 inches.

From apex of heart to anterior

GING GH WAFER soaccochaosooceso0 4 inch, 3 mech.

Iheneth of liver ~...-77-72-.---- 32 inches. 3 inches.

Posterior end of liver to gall-

Alera Gleiskee ee er sec eee une. 12 inches.

Posterior end of liver to

TOL Sl¢ a 7 91 71 7

anterior end of kidneys ... 1g (R), 22 (Lb). 72 (1), 7 (®).

Length of kidneys ........... » > Finch, J inch. 22, 22

Posterior end of kidneys to

5 il 2) (fj D il J1

WOTI tn Os. jase nee ose eeees Ilse (Es)), ge (LI) aire ((les), es (IE)

It is plain from these measurements that the situs viscerwm

shows important differences in the representatives of the two

Hydrophiine genera. In Hydrus the heart is placed much further

forward, i.e. the neck is shorter than in Platyurws, and the

kidneys of the latter are relatively further from the posterior end

of the body. In both, the anterior end of the liver hes close

behind the heart, as is the case in Viperine serpents, a fact also_

noted by Cantor 1m the species dissected by him.

Alimentary viscera.—The liver is long and thin, and thus

characteristically Ophidian. It is very long compared with that of

Hydrus, measuring as it does about one-fifth of the total body-_

length (including the tail), or nearly one-fourth if the tail be

omitted. Taking the length of the body (to the vent) as 100 in

all cases, the following are the proportions of the liver im a

number of serpents.

Zamenis flagelliformis, 193. | Hydrus platyurus, 145.

i gemonensis, 205. Python spilotes, 153.

Coronella getula, 23%. Eryx gaculus, 223.

Coluber melanoleucus, 183. » conicus, 215.

Tarbophis obtusus, 213. Heiterodon platyrhinus, 223.

Naia tripudians, 214. — - Bow constrictor, 174.

Ophiophagus bungarus, 242. | Causus rhombeatus, 232.

Platyurus corallinus, 213. | Lroheterodon madaguseari-

ensis, 201,

The liver shows a peculiarity not without interest, which has

been also figured and referred to by Dr. Cantor * in the Sea-Snake

dissected by him. In Platywrus, however, this character—the

* Toc. cit.

1904. ] ANATOMY OF PELAGIC SERPENTS. 149

division of the liver by furrows running at right angles to the

longitudinal axis of the body—is more marked. 1 counted five of

these transverse furrows, which vary in depth and divide the

liver into a series of segments of hepatic substance. As I shall

point out later (see p. 151), Hydrus platyurus shows the same

“ segmentation ” of the liver. Milne-Edwards observes* of the

liver of Zyphlops that it is “divisé en lobes plats,” and this

lobation is figured also by Cope 7; but it does not appear to be by

any means so regular asin Platywrus. One cannot but put down

this marked lobation to the regular bending of the body im

swimming, and it forms an example of “ segmentation” probably

traceable to a definite mechanical cause.

The gall-bladder gives off a single duct which soon forms a very

complicated network in connection with the hepatic duct. This

network is very much more complex than in Hydrus, and the rete

of ducts is so long before it enters the duodenum that the gall-

bladder can be dissected out and pulled much further away from

the alimentary canal than is possible in Hydrus. The pancreas

seems to me to be smaller proportionately (it certamly is so

actually) than in Hydrus. The coiled region of the mtestine is

very long. When the coils are left undisturbed within their

coelomic space, they measure 7 of an inch, but when unwrapped

no less than 5 inches. The kidneys are approximately equal in

size, each measuring about 2 inch in jiength. They are broad in

proportion to their length, and almost suggest those of the Boid

Eryx. The right, anterior, kidney hardly at all overlaps the left,

which commences where it ends.

S Lung.

It has been pointed out by several zoologists, including Cope <,

that Platywrus and some other genera of Sea-Serpents possess the

tracheal lung found also in a few genera of terrestrial Colubrines.

Cope’s statement on the matter is as follows:—* Finally the

tracheal lung, as I shall call it, is distinct from the true lung in

Platyurus and in Chersydrus. In the former of these genera, the

trachea is not separate fromthe lumen.” Ido not think, however,

that any detailed description of the lung exists. J shall endeavour

therefore to supply this omission by the following description.

There is no trace that I could discover of a second lung. The

single lung extends to within one inch of the cloacal aperture and

ends abruptly without any special diminution of calibre. It les,

posteriorly at any rate, on the right-hand side and is firmly bound

to the dorsal parietes.

The tracheal lung is, as Cope says, distinct from the bronchial

lung; the two are separated at the end of the one and the

beginning of the other by one of the pulmonary vessels which

passes between them, The tracheal lung begins very high up

in the body, close behind the head; it ends posteriorly just in

front of the origin from the heart of the right aortic arch.

* “Lecons sur la Physiologie et Anatomie comparée.’

+ “On the Lungs of the Ophidia,” Proc. Amer. Phil. Sec. xxxini. 1894.

46 ILM. Cit Ws Alf

150 MR. F, E. BEDDARD ON THE VISCERAL [June ig

Until very near its posterior end the trachea forms merely a

gutter along its ventral side, that is it opens freely into it through-

out. Posteriorly, however, there is a distinct tendency for the

lung to be connected only at intervals with the trachea, a hint—

as it appears to me—of the commencing independence of the two

parts of the pulmonary apparatus. The thoracic lung is very

plainly a respiratory organ down to about the middle of the liver

and to a less extent posteriorly. Its texture is like that of many

other serpents, but not like that of Hydrus (see p. 153). It

presents, as is shown in the accompanying drawing (text-fig. 26),

A portion of the internal surface of the lung of Platywrus colubrinus.

a honeycombed and perfectly regular structure. The depressions

are naturally hexagonal through mutual pressure. I could find

no difference in the structure of tracheal and bronchial lungs.

The bronchus is traceable for a long way down the liver. Ata

little distance behind the posterior end of the liver (about ? of an

inch), the lung, which has gradually become very narrow, widens

out into a thin-walled dilatation of quite twice its former width.

This dilatation is 3 of an inch long and fusiform in shape. It is

thin-walled, and thus contrasts with adjacent regions of the lung ;

but on its inner surface a honeycomb pattern is visible, the

depressions, however, being very slightly marked. Behind this

dilatation the honeycomb structure was also plainly visible. The

greater part of the lung in fact seems to be utilised as a respiratory

organ.

It is particularly noteworthy that the posterior region of the

1904. | ANATOMY OF PELAGIC SERPENTS. 151

lung in this serpent has not the excessively thin-walled character

that it has in most land-snakes. In the latter, when dissected, the

anangious region of the lung seems to be little more than a space

between the viscera of the posterior region of the body. In

Platyurus the lung has thick walls throughout. Examined in

transverse sections, the posterior region of the lung, some way

after the dilatation, is seen to be undoubtedly a functional lung,

inasmuch as the blood-capillaries are numerous and approach very

near to the inner surface, being in fact only separated from it by

the pulmonary epithelium. The muscular walls are very thick,

especially the inner layer of circular fibres. Evidently, therefore,

the lung is capable of considerable alterations In size.

Dr. Cantor gives an account of the lungs of Hydrophis schistosa*

Schlegel (= Enkg ydrina valakadien of My. Boulenger’s Catalogue),

somewhat different from the facts as observed by myself in

Platyurus colubrinus ; these differences may be doubtless put down

to the circumstance that the two serpents are of different genera.

In the first place, the tracheal lung would seem to be continuous

with the bronchial lung, though this is not absolutely clear from

Cantor’s figure. The tracheal lung is, moreover, of much less

extent in Hnhydrina. The pattern of the meshwork is quite

different. Finally the dilatation along the course of the pulmonic

portion of the bronchial lung, such as I have found in Platyurus,

is less than in Hnhydrina, and the extreme tip of the lung in

the serpent dissected by me is not tied down to the parietes by

any tag. The dilatation which he does figure is apparently part

of the functional lung.

(2) Hydrus platyurus.

Of this snake the example at my disposal was a female with

the ova immature. The position of the viscera has been already

dealt with in considering Platyurus and a comparison of the two

snakes there instituted.

Alimentary viscera.—The gall-bladder, pancreas, and spleen are

not unlike those of the Sea-Snake figured by Dr. Cantor =. After

receiving the hepatic duct, the bile-duct plunges into the substance

of the pancreas on its way to reach the duodenum. Whether it

forms a reée therein or not, t have not ascertained ; but it is clear

that there is not room for a very extensive one. The pancreas

is much lobulated, and extends in front of, and behind, the gall-

bladder. The liver, as will be gathered from the measurements

on p. 148, is actually, as well as relatively, shorter than that of

Platyurus colubrinus. It is, however, of a more massive structure

and is less divided by transverse furrows into “segments.” I

detected only four of these. The liver also appears to me to be a

little closer to the heart than itis in Platyurus. The coiled region

of the intestine is relatively rather shorter than in Platyurus.

Kidneys.—The kidneys of Hydrus platyuwrus differ markedly

from those of Platyurus colubrinus. They are in the first place

* “ Observations upon Pelagic Serpents,” Trans. Zool. Soe. ii. p. 305.

+ Loe. cit. pl. 57. fig. 1 m. i Loe. cit. pl. a7. figs. 1, 2.

152 MR. F. E. BEDDARD ON THE VISCERAL | June ie

very long and thin, measuring, as has been mentioned, 2? inches

each. While, therefore, the body of Platyurus is rather more

than half the length of that of Hydrws, the kidneys of the latter

are five times the length of those of the former genus. They do

not, however, seem to me to be much, if any, greater in bulk,

since they are extremely thin and narrow, while those of

Platyurus are wider in proportion to their length. Another

difference shown by the kidneys of the two genera is the extent

to which they overlap. In Platywrus the two kidneys hardly

overlap at all, whereas in Mydrus the right anterior kidney

extends alongside of the first two inches of the left kidney. There

is thus a considerable approximation here towards symmetrical

kidneys. :

$ Lung.

The lung-tissue is recognisable from two and a quarter inches

behind the tip of the snout. It begins, therefore, very early in

the neck. The lung-tissue begins gradually between the dorsal

Text-fig. 27.

A portion of the internal surface of the lung of Hydrus platyurus.

non-tused ends of the tracheal rings. This snake has therefore,

as has been stated, a tracheal lung. The trachea, moreover, is

open throughout into the lung, and forms in fact only a gutter

along its ventral surface. The neck of this serpent is short, and

therefore the tracheal lung is not of great extent; it appears to

cease about half an inch in front of the heart, and therefore has

not a course of more than two inches. A sudden widening at that

pomt I take to be the commencement of the bronchial lung.

There is no other differentiation that I can detect. The inner

1904. ] ANATOMY OF PELAGIC SERPENTS. 15s)

surface of the lung is marked by conspicuous thickenings ; but

these do not form a choneyeonip network as in Platyurus.

The folds run across the lung—that is at right angles to its longi-

tudinal axis, and are wavy in outline, which of course allows of

expansion during inspiration (text-fig. 27, p. 152). The trachea (or

bronchus) is continued as a gutter down the lung as far as the com-

mencement of the liver, that is for a very short distance behind

Text-fig. 28.

A portion of the internal surface of the non-vascular part of the lung of

Hydrus platyurus.

the heart. = The lung itself is extraordinarily long; it is indeed

co-extensive with the body-ecavity, reaching as far back as the vent.

Its calibre too is very consider able, and its walls have the tough,

almost shiny, appearance of a fish’s swim-bladder. The dorsal w. all

is firmly fixed to the parietes. The latter part of the lung, 7. e. after

the end of that region which is respiratory, has undergone some

modification in relation to its undoubted function as a swim-

bladder. The folds already spoken of in the vascular region of

the lung persist; but their object is no longer that of merely

increasing the respiratory surface, and thus the efficiency of the

lung as a ane eathing organ, They exist only as bands of muscular

fibre, which, since their direction is mainly circular and parallel

to ond other, must act as constrictors and expel air from

the swim-bladder part of the lung. On a naked-eye inspection of

this portion of the lung, it looks almost as if the bands in question

were the bronchial rings which had in this region taken on a

new development. They are, however, serially continuous not

154 MR. F. E. BEDDARD ON A PARASTERNUM [June He,

with the dwindling bronchial rings, but with the vascular folds

already spoken of. Microscopical examination shows them to he

thick bands of muscular fibres, which are not striated. The

direction of these bands is circular, but they occasionally send off

anastomosing branches as is shown in the accompanying sketch

(text-fig. 28, p.155). Occasionally, too, a small band detaches itselt

from one of the main hoops and ends upon the wall of the lung.

Between the various bands the wall of the lung does not appear

to be muscular. Towards the posterior end of the lung this

arrangement is lost ; there are no longer separate hoops of muscle

but the walls are covered with a single sheet of musculature.

It is clear from the above description, which may be compared

with that of Platywrus on p. 150, that the structure of the lung

differs very considerably in the two species. In Platyurus the

lung has retained to a much greater extent its pulmonary function,

the ‘lung- -substance extending much further back than in Hydrus.

The lung has, however, a less calibre, even proportionately, than

in Hydrus, and there is no trace of so marked a conversion into a

‘“ swim-bladder ” with special muscles effecting its contraction and

expansion as occurs in Hydrus. In the latter genus, moreover,

the whole lung extends further back in the body than it does in

Platyurus.

On the Presence of a Parasternum in the Lacertilian

Genus Tiliqua, and on the Poststernal Ribs in that

Genus. By Frank H. Brpparp, M.A., F.R.S., Pro-

sector to the Society.

| Received June 3, 1904. |

(Text-figures 29 & 30.)

The use of the term “abdominal ribs” for the ventral and

superficially placed cartilages, fibrous bands, or ossifications so

distinctive of certain groups of Reptiles, is open to the objection

that the term “ribs” has already a definite meaning attached

to it. It implies cartilaginous, fibrous, or ossified rods which

have or have had a connection with the vertebral column,

whereas the so-called abdominal ribs have no relation whatever to

the vertebral column, but are purely ventral structures formed

between the plates of the ventral abdominal musculature.

Furthermore, the term ‘abdominal ribs” is actually in use as

descriptive of structures which are real ribs, and which have

nothing to do with what other authors have called abdominal

ribs. For example, Mr. Boulenger, in his ‘ Catalogue of Lizards,’

has referred, under the name of abdominal ribs, to the ventral

region of ribs in the Geckos, Chameleons, and some other forms,

lying behind the sternum, which meet, pair by pair, in the

ventral median line, thus completing a series of hoops encircling

the abdomen. Inasmuch, as Dr. Gadow* justly points out, that

* Cambridge Natural History : Amphibia and Reptiles, p. 504.

1904. | IN LIZARDS OF THE GENUS TILIQUA. 155

these ribs bear “an extraordinary resemblance to the so-called

‘abdominal ribs’ of other reptiles,” it is particularly to be

regretted that the term has been used in so authoritative a

work as the Catalogue to which reference has been made.

Since this confusion has been quite unnecessarily introduced,

it will be as well to adopt the word “ parasternum,” already

used by Fiirbringer, Gadow, and others. The Lacertilia are at

least generally supposed to be without a parasternum, which

is one of the points of difference used to distinguish them

from the genus Sphenodon. The above quotation from Dr. Gadow

implies this general view, which is more explicitly stated in the

‘Royal Natural History’*. I can find no statement in such

works as that section of Bronn’s ‘ Thier-Reich’ which is devoted

to the Lacertilia, as well as in other textbooks, to the effect that

a parasternum is to be found in the Lacertilia; and Iam therefore

free to conclude that the knowledge of its actual occurrence is

at least not widely spread.

In a brief preliminary note in ‘ Nature’ + I pointed out the

existence of a series of abdominal ribs in the Scincoid Lacertilan

Tiliqua scincoides, and I herewith submit to the Society a more

detailed account of these structures (text-fig. 29, p. 156), which

I have not up to the present succeeded in observing in any other

lizard.

The chevron bones of the abdominal-vib system are thin and

not always easy to see; their slender bulk, as it appears to me,

fully accounts for the fact that they have been previously over-

looked. They are not nearly so stout, so numerous, or so closely

adpressed as these bones are in an example of Hatteria of about

the same size as the two specimens of Tiliqua scincoides which

I have examined. For these reasons the bones would very

readily be lost in preparing skeletons of 7iliqua. The distance

separating the chevrons in 7%iliqua is 8 mm., when the abdominal

muscles are gently stretched but not overstretched; the same

distance in the case of the dried abdominal skeleton of Hatteria

was not more than 4 or 5 millimetres. In my specimen of Hatteria

there were quite twenty of the chevrons; I could not detect

more than seven in one of the two specimens of Ziliqua. In the

other specimen these abdominal ribs were hardly at all obvious.

The fact that these chevrons are quite independent of the ribs—

and their purely superficial position, lying as they do in the

ventralmost sheet of muscle of the abdominal wall—is in favour

of regarding them as the homologues of the abdominal ribs of

Hatteria. It remains, however, to be shown that they overlap

the true ribs as the abdominal ribs do in Hatteria. The ribs in

Tiliqua atter the sternum do not apparently reach so far ventrally

as they do in Hatteria. It might therefore conceivably be held

that we had here to do merely with the ventral moieties of ribs

which were defective laterally, and that the condition occurring

preceding ones is the absence of any system of true abdominal ribs or of their

equivalent a plastron ” (vol. v. p. 107).

+ May 5th, 1904, p. 6.

156 MR. F. E. BEDDARD ON A PARASTERNUM [J une 7,

in Tiliqua was an impertect representation of that occurring in the

Chameleon and other lizards, where a number of true ribs join

ventrally behind the sternum. Thus in dAcontias and Typhlo-

saurus, of which the former is a near ally of Zliqua, there

are present these post-sternal ribs meeting in the middle line,

and thus simulating abdominal ribs *. In the Chameleon the

most ventral layer of the abdominal musculature is so thin that it

requires a careful dissection to show that the ventrally meeting

ribs do not lie in this layer but in the deeper layer. In Tiliqua,

on the other hand, the ventral musculature is thick, and the two

Text-fig. 29.

Abdominal ribs of Tiliqua scincoides.

A points to one of these ribs. A meshwork of tendons lies between and over them.

layers are easily distinguishable. When the ventralmost layer is

raised, the abdominal ribs are carried with it, and are seen to end

off in it with shghtly forwardly-curved lateral ends. But—more

important than this—on the deeper layer of the musculature are

two or three cartilaginous bars, lying obliquely (see text-fig. 30,

p. 157), which ave clearly the ventral portions of true ribs such as in

Acontias make complete hoops. The discovery of these entirely

sets at rest any doubt as to the fact that the bars of cartilage which

ule superficially are really quite comparable to a parasternum. The

true ventral moieties of the ribs are not in connection with their

vertebral portions, so far as I have been able to ascertain. They

would thus be very readily lost in macerated skeletons, and might

even be overlooked m a dissection. In his ‘ Catalogue of Lizards

* Peters, ‘ Reise nach Mossambique, Amphibia, pls. xii. & xiii.

1904. | IN LIZARDS OF THE GENUS TILIQUA. 157

in the British Museum, Mr. Boulenger uses as part of his

definition of the Scincide * the sentence ‘ Ossified abdominal ribs

are absent”; and in defining the Anelytropide + he includes in

that definition the phrase ‘“ Abdominal ribs present,” It is not

absolutely clear from the wording used whether these two nearly

related families are to be distinguished (inter alia) by the absence

or presence of “abdominal ribs” (7. ¢., as already pointed out, the

ventral moieties of post-sternal ribs) or by the mere absence or

presence of ossification in such ribs. In any case, not only has

Peters figured complete hoops in Acontias, but I have been able

Text-fig. 30.

Tiliqua scincoides.

Ventral Hap of musculature (B) in which lie the abdominal ribs, turned down to

show the deeper-lying muscles (C), in which are imbedded the true ribs (D).

to show in the present communication that considerable vestiges

of the ventral moieties of post-sternal ribs occur in Tiliqua.

It is to my mind possibly a matter for further enquiry as to

how far the median ventral region of the post-sternal ribs which

actually meet each other behind the sternum may not be actually

a parasternum fused with true ribs. For the median region of

the chevrons, whether of undoubted abdominal ribs or of true

vertebral ribs, appears at the surface of the abdominal musculature.

This median region, therefore, at least of the post-sternal true

ribs may conceivably be a vestige of a parasternum in those

Lacertilia where such ribs occur.

* Vol. iii. p. 130. + Vol. iii. p. 430.

158 DR. E, A. GOELDI ON THE [June 7,

+. On the rare Rodent Jinomys branichii Peters. By

Dr. Emit A. Gortpi, C.M.Z.S8., Director of the Goeldi

Museum, Para.

{Received May 16, 1904. ]

(Plate X.*)

The zoological world was surprised in 1873 by the novelty of

the discovery of a strange, large Rodent, introduced scientifically

by Prof. Peters, then Director of the Berlin Museum, under the

name Dinomys branickii. It was stated that the animal had been

found in the neighbourhood of a town in Peru, wandering about

in an orchard. It was further stated that the natives themselves

were entirely unacquainted with the creature. Prof. Peters

published a somewhat extensive memo on the subject, based

principally upon anatomical features, especially of the skin and the

more or less complete skeleton. Up to this date I have not had

the opportunity of consulting this memoir, which I know only

from a few lines of quotation in recent manuals on mammalogy,

such as Flower and Lydekker, ‘ An Introduction to the Study of

Mammals,’ London, 1891, p. 489; and Beddard, *‘ Mammals,’

London, 1902, p. 495 seg. These citations are barely sufficient

to permit a certain identification of this peculiar form of Rodent

and to exclude the possibility of confusion.

So far as the literature at my disposal goes, there is no indi-

cation of any living specimen having been examined by a

zoologist, nor does there exist any notice of any further specimen

as having been met with since 1873, so that the type specimen at

the Berlin Museum remains till now the only one known to be

in existence.

I have now the extreme good fortune to be able to make

further additions to our knowledge of the subject. I consider

the rediscovery of Dinomys branickii in the Amazonian region,

from a general point of view, as the second most important

* For explanation of the Plate, see p. 162.

+ Since beginning to write the present article I have received both the extensive

memoir of Prof. Peters as well as the preliminary communication on the subject

in the ‘ Monatsberichte der kénigl.-preuss. Akademie der Wissenschaften zu Berlin,

July-August 1873, pp. 551-552. Their contents correspond so nearly to my suppo-

sition that I find no necessity for changing anything m the wording of my note.

The coloured lithographic figure given by Prot. Peters on plate 1. identifies the animal

satisfactorily, but, of course, can make no claim to rival the photographs taken from

the living specimens, which give an essentially different conception of the animal’s

physiognomy. The most noteworthy difference consists in the fierce expression of

countenance, resembling that of an angry rat ready to bite; while my photographs

show a face which might be called the personification of perfect good humour. As

I foresayy, there still exists no information as to the anatomy of the scft parts

of Dinomys, which in all probability is still in reserve for me to bring to hght.—

April 7, 1904. '

+ On reading the memoir of Prot. Peters, I find—what was absolutely impossible

to presume from the brief references in the above-cited manuals—that this type

specimen does not belong to the Berlin Museum, but was only lent to its Director to

be described by him, and that it forms part of the collections of the Warsaw

Museum, for which it had been obtained by the Polish naturalist and explorer

Constantin Jelski.

2 1S, WO4s, woll i Wl 2&,

Bale, Sons and Danielsson, Ltd,

IDL INVOIMEYAS: IB IRUEUIN INC CIN

1904. | RODENT DINOMYS BRANICKII. 159

scientific event that has occurred during the first decennium of

the existence of the Para Museum, after the refinding of the

long-lost Dipnoan Lepidosiren paradoxa, as fully described in

former volumes of the Zoological Society’s publications.

My surprise was unbounded when, a few weeks ago, I received

from a friend of the Museum, personally unknown to me, a cage

containing a pal of live large Rodents entirely new to me,

notwithstanding a twenty years’ acquaintance with the Brazilian

fauna, both from the South and the North. These, after a

quarter of an hour of eager consultation of the more modern

manuals of mammalogy, I recognised as being identical with the

almost legendary Dinomys braniehia, of which science had lost

sight for thirty years.

“It seems to me to be a most opportune occasion to publish a

short description of the external features and the habits of Dinomys,

as it is probable that the memoir of Prof. Peters, the only source

of information on the subject up to the present, is deficient

in these particulars. A ‘series of photographs of the animal,

taken in different attitudes spontaneously assumed, supplement

my verbal explanations.

Doubtless the first impression from the general appearance,

especially as regards size and external features, would ally the

animal closely with the South-American “ Paca” (Agouti paca) ;

and the verdict of the scientist as to its systematic location

has no reason to differ from the popular conception which has

given to our Rodent the local name of “ Paca-rana,” a Tupy-term

meaning “‘ pseudo-paca.” In size the older and larger specimen,

the mother, is almost exactly of the dimensions of an average

Paca ; the young one is approximately half or two-thirds of that

size. The similarity is increased by its capacious head, and the

brownish colour marked with distinctly visible longitudinal rows

of white spots. This similarity, however, begins to diminish as

soon as one undertakes a more mimute examination. It presents

first the striking difference of having a hairy tail of about 2 deci-

metres in length. Still further, as regards the hair, while that of

the Paca is soft to the touch and of even length, resembling

somewhat the fur of the Otter, Dinomys has a rougher coat of

stiffer hairs, uneven in length (these stiffer hairs being white-

tipped and longer, and contributing to interrupt the uniformity of

the general colour), and forming a first transition-step towards

the quills of the coats of the Spiny Rats proper (Loncheride,

Kehinomyide). The white spots are located only on the rear part

of the trunk, beginning from the shoulders. They are separated

in two groups: (1) four longitudinal rows, nearly continuous at

the fore part, the median line of the back being dark; (2) two

lateral areas of more or less circular white spots occupying the

flanks and thighs, scattered irregularly, as it would seem at first

sight, but betraying a tendency to longitudinal arrangement on

closer observation.

The form of the head of Dinomys may be described as sub-

160 DR. E. A. GOELDI ON THE [June 7,

pyramidal, while that of the Paca is subconical, due to its promi-

nent cheek-bones and bulging eyes and the more slender snout.

The former thus resembles more the head of the Capybara.

What gives to the head a peculiar martial aspect are the immense

white whiskers, equal in length to the animal’s head, forming

a thick tuft on each side. The nostrils are §-shaped, and

have the same valvular arrangement for closing the aperture

as the Manatee, for example, showing that the animal is adapted

to a semi-aquatic life. While the prominent eyes of the Paca,

like large black beads, have a glaring glossy aspect, the eyes

of the Dinomys are not at all prominent, are of asoft light brown

colour, the pupil being a vertical slit during the hours of day-

light. They are thus of similar appearance to the eyes of

certain smaller Didelphyidee (Didelphys quica, &c.) and of the large

Two-clawed Sloth (Cholapus didactylus); and this similarity

naturally leads to the conclusion that the animal is rather of

nocturnal habits—a conclusion which is corroborated by actual

observation. Both front and hind feet are fow~toed. The

general build of Dinomys is thick-set and inclined to corpulency.

Due to the fact of setting the whole plantigrade sole on the

ground, the hind feet especially, the Dmomys has a waddling

gait, Al reminds one of an immense rat well advanced in

development towards a bear.

Both my specimens of Dinomys are of a peaceful, phlegmatic

disposition ; they are most excellent boarders in our Zoological

xarden, and cause very little trouble on condition of having

enough to eat. They devour prodigious heaps of provender,

eating being one of thei principal occupations both day and

night. As far as observed up to the present, they show a

preference for bread, squashes, and fruit of different indigenous

palm-trees, such as the orange-coloured * Tucuma ” (Astrocaryum

tucuma). While eating, their favourite position is standing erect

on the hind feet, and in this attitude there is a good opportunity

of admiring the dexterous manner in which they can hold a fruit in

their paws, notwithstanding the lack of the counter-pressure of a

thumb. Such a fruit revolves in their grasp as though it were in

the chuck of a slowly-turning lathe, and the large chips which fly

fill us with respect for the immense gouging faculty of the large

but comparatively shghtly prominent incisors. The erect position

while eating—which is the only attitude thus far observed while

taking fool—reminds one more of the “ Cutia” (Dasyprocta)

among the larger South-American Rodents, and forms a note-

worthy difference from the *Paca,’ which I have never seen

eating in any other position than on all fours.

The predominant feature of the character of Dinomys is a con-

bination of leisurely movements and supreme good nature. It

knows absolutely nothing of haste. Spending the greater part of

the day sleeping in a corner—the mother often lyimg upon the

young one, or standing over it, as if to protect and to keep it

warm—opening its half-closed eyes only when it hears the

‘

1904. | RODENT DINOMYS BRANICKIL. 161

approaching steps of the keeper, it forms the resolution to

move with slow gait, expecting some food, evidently governing

its movements as much by hearing and smell as by sight. It is

not easily irritated, and permits one to stroke and to scratch its

head and back, and only occasionally manifests its displeasure

by a low euttural erowl. I have never yet observed a manifest

intention to bite. When let out of the cage it makes no attempt

to escape, and limits its excursions to an exploration of the

immediate neighbourhood in search of something to eat. It

occasionally scratches itself rapidly with its long claws, which is

the only occasion on which it manifests a capacity for rapid

movements when required. One thing not yet definitely verified

by us is its proclivity for digging, the development of the claws

at least leading to the supposition that the animal is well fitted

for that purpose. The amiable relations always existing between

mother and son prepossess one most favourably as to the natural

disposition of the animals *

This phlegmatic disposition seems to me to be a very precarious

endowment for the struggle for life; and considering the evident

advantages which result to the smaller domestic rodents, such

as rats and mice, from their nervously active constitution, it

would not be strange if the species should tend to disappear.

The apparent rarity of Dinom ys may possibly find its explanation

in the consequences of such a psychological endowment in a more

nervous environment; but it is also possible that this rarity is

because of the circumstance that the real habitat of the species

has not yet been clearly ascertained. As matters now stand, it

would be justifiable to suppose that the true home of Dinomys i 18

not properly in the Peruvian Andes, and that the first specimen

found there was merely a stray individual, and that its actual

habitat may rather be located in the almost unexplored regions

of the eastern slopes and tablelands of the Bolivian and Peruvian

foot-hills bordering on Brazil, including geographically the head-

waters of the rivers Acre, Purts, and Jurua. I shall soon have

occasion to show that a scientific exploration of the region

above described will result in a multitude of great surprises

both from a zoological and a paleontological point of view, of

which the interesting rediscovery of the lost Dinomys branickit is

only a first instalment.

Para, 7th April, 1904.

P.S.—Unfortunately, just before I send this note, the older

Dinomys, the mother, has died owing to a difficult parturition.

One fetus was born under normal conditions, while the other,

apparently on account of its abnormal position, could not be

* After this portrayal of the animal’s peaceful character, it will not seem strange

that the account of the capture of the first individual in 1873 in the Peruvian

mountains, as furnished by Prof. Peters, according to which it was deemed necessary

to deal two powerful sabre-strokes to lay the terrible monster low, always amuses

me.

Proc. Zoot. Soc.—1904, Vou. IT. No. XI. 11

162 ON THE WILD CAT OF TRANSCAUCASIA, [June 7,

delivered. This unforeseen event will now give, much sooner

than I expected, an opportunity to fulfil my promise to undertake

a thorough study of the anatomy of the soft parts, as well as of

the two fully-developed foetuses.

ith April, 1904.

EXPLANATION OF PLATE X.

Dinomys branickii, from photographs of two living examples (a mother and young)

(about <4; nat. size), and the skin of the former (about + nat. size) in the Goeldi

Museum, Para.

8. The Black Wild Cat of Transcaucasia.

By C. Satunin, of Tiflis, C.M.Z.8.

[ Received May 16, 1904. ]

Although the existence ih Transcaucasia of a Black Wild Cat

was known long ago, the animal has never been described nor

scientifically named.

Hochenacker speaks of this cat, so far back as 1837 *, as a

Felis cato affinis; but as all his text is in Latin, this cannot be

regarded as a scientific name forthe animal. I myself have men-

tioned this cat as Felis sp. in my paper on the Fauna of the

Caucasus, as well asin the Catalogue of the Caucasian Museum $f;

but I have not had the opportunity of giving a description of it

until now.

That it really is a Wild Black Cat I knew well, as all the spe-

cimens I have had the opportunity of examining were alike, and

as it is by no means rare in its native haunts.

There remained one important point to decide about this

animal, namely, Was it not a melanie form of the common Wild

Cat, Felis catus L. %

Thanks to the material in the Zoological Museum of the

St. Petersburg Academy of Sciences, where I have found two

mounted specimens, three skins, and three skulls of this cat, I am

now certain of its specific validity, and I name it Melis demon §,

of which the following is the description :—

Size of a big male domestic cat. Colour ranging from black

with a slight reddish tinge to reddish dark brown. ‘This colouring

is somewhat lighter on the under side of the body, on the mner

surface of the extremities, and on the distal under surface of tail.

Very long white hairs are scantily dispersed all over the body.

In a certain light, dark-black transversal stripes are visible on

the sides of the fore part of the body, these stripes being more

conspicuous on somewhat faded skins.

The whiskers, as well as the eyebrow-bristles, are brown.

The tail is considerably longer than in /elis catus.

Claws white, transparent, and with a mother-of-pearl lustre.

* Bull. Soc. Nat. Moscou, 1837, p. 186.

+ Zool. Jahrb., Syst. ix. p. 289 (1897).

‘Museum Caucasicum,’ i. p. 24( 1899). i ;

“Demon” being the hero of everal Caucasian legends, and also of the poem by

Lermontoff with the same title

1904. | ON A NEW FORM OF BUFFALO, 163

The measurements, in millims., of the two mounted specimens

and of the three skins are :—

Mounted specimens. Skins.

I —

(= rae tag aaa

? No. 671. No. 2685. No.2997. No.3001. No.4945.

From tip of nose to base of tail 610 560 6) Oe 3) a odo 0)

Tail, hair included ............ 309 ~=8 40 340 380 370

Ear, from external base ... 51 5] — 54 46

Skull: as compared with the skull of Felis catus that I have

had for comparison, the three skulls of this new eat differ only in

that the frontal part is somewhat narrower. Moreover, in Felis

catus the nasals reach further back than the ascending branches

of the upper mandible, whereas in Melis demon the upper man-

dibular bones go further back than the nasals.

The measurements of two complete skulls, in millims., are :—

No. 2997. No. 3001.

dis :

Gireaibestillemte tas. yek bya 5 20) Lee Ge, Peete eee 98 101

iBasilarWemert byes... 22. tis ea. eee Si 81

Greatest breadth across the zygomatic bones 69:5 71

Smallest breadth behind the postorbital

POIROXGISSIS) AG hn Apt ne ence pan En RRARREMPL tA cfc huo,cio 31°2 34

Smallest breadth between the interorbitals. 19 20

Greatest breadth of brain-case .............0. 46 46°5

Ibrerarerialn Ont lovoragy JRVENIE) A seancoonsboncnotcencosas nr 36 33

Width between upper premolars ............ 38°5 39°D

Condylar lengchyot maxilla) 22-2 .a..ceeeneeeen 61 64

Felis demon is not scarce in the woods of the southern slopes

of the chief range of the Caucasus (Nookhinsky district of Gouv.

Elizabethpol) and the Zakatalsky Province. It used to be

found, according to Hochenacker, also in the mountains of the

Small Caucasus; but nothing is heard of it there at the present

time. According to Radde, this cat inhabits the Kopet-Dagh

Range in Transcaspia; and M. 8, Alphéraky tells me that it is

by no means rare in the woods of Borshom, where it is often

trapped, together with /elis catus and other vermin. Habits

unknown.

9. On a Buffalo-Skull from East Central Africa.

By R. LypexKer.

[Received May 17, 1904. ]

(Text-figure 31.)

[The complete account of the new form described in this communication appears

here; but since the name and preliminary diagnosis were published in the ‘ Abstract,’

the species is distinguished by the name being underlined.—Ep1r7or, |

Through the kind offices of Rowland Ward, Limited, the

British Museum has recently been presented with the skull and

tail-tip of a bull Buffalo, killed by the donor, Mr. Arnold Mathews,

Lie

164 MR. R. LYDEKKER ON A [June 7,

of Ballynahinch Castle, Co. Galway, in the Mfumbiro district, on

a volcanic mountain west of Kiva, on the borderland between the

Congo Free State and German Hast Africa, in forest at an elevation

of between 7000 and 8000 feet. These specimens I cannot identify

with any described form, and therefore propose to regard them as

representing a new race of African Buffalo.

Although I cannot find Mfumbiro in any atlas, I take the

district in question to be the area lying between the Albert

Nyanza and Tanganyika, that is to say, approximately, the Mapi

country.

The following note on the Buffalo to which the specimens

belonged has been communicated by Mr. Mathews :—

“The hide of this animal is not bare like that of the South-

African Buffalo, but covered with a dense crop of black hair all

over. The height of the buffalo is considerably less than that of

the big South and Hast African races. The animal seems to me

to be a race midway between the Congo and the Hast-African

Buffalo. It shows, however, no tendency to red. In addition to

its small size, peculiar shape of horns, and density of pelt, the

only peculiarity I noticed was that the tail had a white tip. I

shot another bull out of the same herd, exactly like the one

of which I sent you the skull, only younger and smaller.”

The skull and horns are evidently those of a fully-adult bull,

and the latter present the following measurements :—

Along outer curve ............ 21°5 inches.

Baisalioirtla e.Witee acl eeecnac cee Ser peta

Gureatestispaniiwas- en. seeaanne 2599)",

Tip to tip interval ............ ISH)

Compared with the various local forms included in my work on

‘Wild Oxen, Sheep, and Goats,’ under the specific title of Bos

(Bubalus) caffer, the skull and horns of the present form come

nearest to those of 5. caffer nanus of the Congo region. They

are, however, considerably larger than the latter ; and the horns

(text-fig. 31, p. 165) are thicker, less sharply incurved, and with a

much more sinuous front surface, bending sharply backwards

immediately behind the basal frontal expansion, and then curving

somewhat forwards with the commencement of the inward

inclination. An interval of about an inch and a half separates

the two horns in the middle of theforehead. Speaking generally,

these appendages are intermediate between those of the Cape and

those of the Congo Buffalo, although on the whole nearer to the

latter. Here it may be mentioned that there is an almost complete

gradation, as regards the extent of the frontal sinuosity, from the

horns of the Cape Buffalo through those of the present and the

Congo forms, to the Senegambian race (B. caffer planiceros), in

which this curvature is practically obsolete.

From the typical red Congo Buffalo (B. c. nanus) the present

form differs by its deep-black coat, whereby it agrees with the

Cape animal, from which, however, it is broadly distinguished by

the dirty-white tail-tuft. In reference to Mr. Mathews’s note,

1904. ] NEW FORM OF BUFFALO. 165

it may be mentioned that two Cape Buffaloes in the British

Museum have a well-developed coat of black hair.

As this Buffalo has obviously nothing to do with the one from

East Africa recently described by Mr. Thomas*, I propose to

regard it as a new race, under the title of Bos caffer mathewsi

(cf. Abstr. P. Z.S. 1904, no. 9, p. 10, June 14), the Museum spe-

cimens being of course the types. It may be shortly defined as a

race of the approximate size of B.c. nanus, but with the abundant

hair black instead of red, and the tail-tuft white ; the horns being

at the same time larger, more curved outwardly, and then inclined

backwards near the lateral bend.

Text-fig. 31.

Front view of skull of male Buffalo, Bos (Bubalus) caffer mathewsi, from

the Mfumbiro district of East Central Africa. + nat. size.

I always feel I owe an apology to naturalists when adding to

the list of subspecies; but the present form has certainly claim

to recognition, on account of its tending to connect the Cape with

the Congo Buffalo—two forms which at one time I had some

compunction in regarding as specifically the same.

* P.Z.S. 1904, vol. i. p. 371.

166 MR. R, LYDEKKER ON A [June 7,

10. The Ichang Tufted Deer. By R. Lyprxxmr.

[Received June 8, 1904. |

(Text-figures 32 & 33.)

[The complete account of the new species described in this communication

appears here, but since the name and preliminary diagnosis were published in the

‘ Abstract,’ the species is distinguished by the name being underlined.—Eprror. |

The genus Hlaphodus has been hitherto known by two species—

the typical ZL. cephalophus and the perfectly distinct H. michianus,

from the Ningpo district, province of Chekiang, on the east

coast of China. J am now able to adda third. A few days ago

Mr. A. E. Leatham called at the Natural History Museum,

bringing with him for determination the skull and skin of a

young male Tufted Deer (Hlaphodus), shot by himself last January

in the mountains near Ichang, province of Hupei, Central China.

Ichang, it may be mentioned, is fully a thousand miles from

Ningpo; and the deer killed by Mr. Leatham was shot high up in

the mountains far away from water, whereas 1. michianus 1s

reported to inhabit the reed-brakes on the Ningpo rivers. On

looking through the specimens in the British (Natural History)

Museum, I found an adult male skin and skull of an Hlaphodus

from Ichang, collected by Mr. F. W. Styan in 1901 (B.M.

INO eS 2c):

Externally, Mr. Leatham’s specimen of the Ichang Tufted

Deer differs from “1. michianus by its decidedly darker and more

uniform colour, which is blackish brown, passing almost into

black on the limbs, while there is more white on the tail, of

which only the basal two-thirds of the upper surface is dark, so

that the whole of the tip is white.

The skin of the adult male sent by Mr. Styan is browner,

except down the middle of the back, but exhibits the same

uniformity in general colour. How different these skins really

are in colour from that of H. michianus, it is not easy to deter-

mine, seeing that the specimens of the latter in the Museum are

more or less faded by exposure.

On comparing the skull of Mr. Leatham’s specimen, which

is immature and retains the milk-molars, with a skull of a male

EH. michianus of nearly the same age, I find that the nasal bones

are absolutely and proportionately shorter in the former, their

length being 17 inch against 24 inches. Moreover, their length

is contained 31 times in the basicranial length, instead of less

than 3 times. Another distinction is to be found in the form of

the anterior upper milk-molar, which is much shorter (antero-

posteriorly) in the Ichang than in the Ningpo skull; and there

also appear to be slight differences in the form of the upper

molars.

Both skulls, it may be observed, show no trace of the pedicles

of the antlers, which must accordingly be very late in development.

Equally noteworthy differences are presented when the adult

1904. | NEW SPECIES OF TUFTED DEDR. 167

skull (text-figs. 32 & 33, A) obtained by Mr. Styan at Ichang is

compared with an adult male skull of 4. michianus from Ningpo

(No. 78.11.14.3) (text-figs. 32 & 33, B), the latter bemg somewhat

older than the former. The Ichang skull shows the shortness of

the nasals characteristic of the immature specimen, the length of

these bones being 23 inches, against 33 inches in #. michianus.

Text-fig. 32.

AN), NOY Np Oe Eo

AM bts ris: ; ye A

Left lateral view of adult male skulls of Hlaphodus michianus (A),

and H. ichangensis (B).

pr.mex., premaxilla; pr.v., preorbital cavity.

The antler-pedicles are much more developed in the Ningpo skull,

and have a much more outwardly-directed ridge connecting them

with the orbit (as shown in text-figure 32, A); but this feature

may be due, to some extent, to difference in age. The flat surface

of the jugal below the orbit is quite narrow in the Ningpo, but

168 ‘MR. R. LYDEKKER ON A [June 7,

broad in the Ichang skull. Very noticeable is the difference in

the form of the preorbital fossa, which is rounder, deeper, and less

oblique in the Ichang skull. More striking still is the difference

in the canines of the two forms, that of the Ichang skull being

nearly twice the length of that of the older Ningpo specimen,

as well as differing somewhat in shape. Such differences cannot

possibly, I think, be accounted for by wear, as I cannot conceive

Text-fig. 33.

Front view of adult male skulls of Hlaphodus michianus (A)

and H. ichangensis (B).

fr-, frontal ; na., nasal.

how such tusks could become worn, while those of the Ningpo

skull show no signs of wear. I may add that the figured skulls

differ in the form and relations of the premaxille, but 1 find this

to be a very variable feature.

Compared with a female skull of the typical Tibetan 2. cepha-

lophus in the British Museum (No. 92.7.13.1), which is the only

specimen of the latter species available, the adult Ichang skull

1904. | NEW SPECIES OF TUFTED DEER. 169

makes a much nearer approach than it does to that of H. michi-

anus, showing a similar conformation of the bones below the

orbit, and apparently also a very similarly shaped preorbital

cavity. The nasal bones are, moreover, of much the same

relative length, their long diameter being one third of the basi-

cranial length. On the other hand, #. cephalophus is at once

distinguished by its greatly superior size, as is shown by the

following table of the basicranial lengths of the four skulls

available for comparison :—

II, TOC OKTIDUIS, WHE Aon aneeoenodsjo6eor 63 inches

: Female. issu doen ces heen Gy

Nchanmesadulitymale yess. at i- Oe iss

Li. cephalophus, female ............... (ace

From this it is obvious that the Tibetan skull cannot possibly

be the female of the form indicated by the adult Ichang specimen.

If a male skull of 1. cephalophus were available for comparison,

I have little doubt that important differences between it and the

Ichang specimen might be detected. As it is, the former species

appears to have a much longer type of skull than the Ichang

animal, from which it is also distinguished (so far as can be

determined) by the much more solid structure of the walls of

the preorbital cavity.

In any case, the Ichang Deer is broadly distinguished from the

Tibetan Hlaphodus cephalophus by its greatly inferior size.

The foregoing differences seem to leave no doubt as to the

specific distinctness of the Ichang Tufted Deer, which may be

named Hlaphodus ichangensis (cf. Abstr. P.Z.S. 1904, No. 9,

p. 10, June 14). Itis characterised, as compared with L. michianus,

by its darker and more uniform colour, whiter tail, smaller antlers,

larger tusks, shorter nasals, and smaller, deeper, and more evenly

circular preorbital fossa; while it is smaller than /. cephalophus.

The type will be the aforesaid adult male skull, collected by

Mr. Styan.

In conclusion, I may take the opportunity of referring to the

skull of a female Tufted Deer (B.M. No. 98.3.7.18) obtained,

together with the skin, by Mr. C. B. Rickett at Fing-ling, Fokien,

lying considerably to the south of Ningpo. This skull is of the

same general type as that of the female of Z. michianus, but

differs by its superior size, the basicranial length in the two

specimens being respectively 63 and 6Linches. The Fokien skull

is further distinguished by the less marked elevation of the

hinder half of the frontal region, which does not develop a

median crest, and by the expansion and flattening of the platform

formed by the base of the preorbital cavity and the anterior

zygomatic root. ‘There are also differences in the form of the

paroccipital process in the two skulls, which, with other details,

and their marked difference in size, appear to justify the reference

of the Fokien Tufted Deer to a separate race, under the name of

Llaphodus michianus fociensis.

170 DR. A. SMITH WOODWARD ON [June 7,

11. On Two New Labyrinthodont Skulls of the Genera

Cupitosaurus and Aphaneramma. By A. Surra Woop-

WARD, DED FuktSs) KeZs:

[Received June 3, 1904. |

(Plates XI. & XII.*, and Text-figure 34.)

Among the remains of Labyrinthodonts acquired by the British

Museum during recent years, there are two skulls of unusual

interest. One was obtained from the Triassic sandstone of

Staffordshire, while the other was found in a formation, appa-

rently of the same geological age, in Spitzbergen. The first

specimen is of importance as displaying the occipital region

of the skull more clearly than any Labyrinthodont hitherto

described; and the second specimen adds facts concerning the

shape and relations of the quadrate bone. Each fossil represents

a new species, while the second is the only skull of a Labyrintho-

dont as yet described from the Arctic Regions.

I. CAPITOSAURUS STANTONENSIS, Sp. nov. (Plate XI.)

The discovery of the Staffordshire specimen in the Lower

Keuper sandstone of Stanton, near Uttoxeter, was briefly noticed

by Mr. John Ward four years agot. The block of sandstone

containing the skull was split along the plane of the cranial roof,

leaving most of the roof-bones adherent to one slab, while the

impression of these, with the rest of the skull, remained in the

counterpart-slab. When Mr. Ward examined the fossil it was

still in the condition. in which it had been exposed by the acci-

dental fracture; so that his description could only be of a general

and preliminary nature. Since its acquisition by the British

Museum, both parts of the specimen have been skilfully prepared

by Mr. C. Barlow; and the principal characters of the skull, as

now observable, are illustrated in Plate XJ.

The skull does not appear to have been much distorted by

crushing, and its shape closely resembles that of the skull of a

broad-nosed crocodile. All the external bones are similarly

ornamented with a coarse network of sharp ridges. The occipital

border is slightly excavated, and the deep notches for the auditory

meatus (aw.) are nearly, if not completely, surrounded by bone.

The specimen is a little fractured in this region. The orbits

(orb.) are set far back, only of moderate size and regularly oval

in shape, with the long axis directly antero-posterior, not oblique.

The pineal foramen (pin.) is a rather large circular vacuity.

* For explanation of the Plates, see p. 176.

+ J. Ward, “On the Occurrence of Labyrinthodont remains in the Keuper Sand-

stone of Stanton,” Trans. N. Statis. Field Club, vol. xxxiv. (1900), pp. 108-112,

pls. iv., v.

PZ. 5 1904 voll Pod.

M L.

J.Green del.et Lith. CAPITOSAURUS STANTONENSIS. Mintern Bros.imp.

Bate aun WOALVELSOU VANVEENVHdV Rarer ee uectney

TXTd 114 vO61 SZ d

1904. ] LABYRINTHODONT SKULLS. 171

The sutures between the roof-bones in the hinder half of the

skull are clear on both sides of the fossil; but the elements in

the rostral region cannot be distinguished. There are the usual

two pairs of small bones forming the occipital margin between

the auditory notches, the middle pair being larger than the lateral

pur. The parietal bones (pa.) are a symmetrical pair, twice as

wide in their posterior half as in their anterior half, and trun-

cated in front where they meet the frontals (/r.). Their maximum

width is less than their total length, and the pineal foramen is

situated at their middle point. The frontals just enter the rim

of the orbit at its antero-internal portion; but the parietals are

completely excluded from this rim by the antero-posteriorly

elongated postfrontals (pif). The squamosal element (sq.) is

longer than broad, articulating in front chiefly with the post-

orbital, but also meeting the postfrontal. The postorbital (péio.)

forms the posterior and half the externo-lateral rim of the orbit ;

the jugal (j.) enters the outer portion of this rim; while the

prefrontal (prf.) completes it in front. The large prosquamosal

(psq.) is about two-thirds as deep as long, tapering to a point

forwards. The quadrato-jugal (qj.) is about two-and-a-half times

as long as deep, also tapering to a point forwards, and extending

shightly downwards below the level of the tooth-bearing edge of

the upper jaw. The jugal (j.), as usual, is irregularly triangular

in shape, with the radiating ridges of its ornamentation pre-

dominant.

At the rostral end of the fossil represented in Plate XI., the

eranial roof and underlying matrix are removed to expose the

palate from above. The antero-posteriorly elongated posterior

nares (pin.) are thus well seen; and the hinder edge of the large

premaxillary vacuity (pv.) is also preserved. The posterior limit

of the premaxilla (pmz.) is distinct on the left side. The exten-

sive vomerine plates (v.), divided by a median suture, are also

well preserved. The premaxillary teeth, shown in transverse

section, are small, uniform in size, and arranged in close series.

The maxillary teeth, exposed in side view, are similar. Hach

tooth appears to have been a hollow cone with plicated walls.

An excavation in the sandstone reveals hollows left by the decay

of the usual large teeth, which occur on the palate in front of and

behind the posterior narial openings.

As already mentioned, the most interesting part of the skull is

the occipital region, which is especially well preserved (fig. 2).

It does not project sufficiently far backwards to admit of the

occipital condyles being seen when the cranium is viewed directly

from above. ‘The position of the foramen magnum (/m.) is clear;

while a vacant space (socc.) between this and the roof-bones of

the occipital border suggests that a supraoccipital element was

present but unossified. Below the middle of the foramen magnum

there is a narrow tongue of bone (bocc.) separated by a little

matrix from the prominent occipital condyles. This is probably

172 DR. A, SMITH WOODWARD ON [June 7,

to be interpreted as a partially ossified basioccipital element

forming only an insignificant part of the occiput. The condyles

themselves (¢.) are convex and seem to have been covered with

unossified cartilage, while they constitute the hindermost end of

a pair of bones which enter extensively into the base of the

cranium and also rise upwards to bound the foramen magnum

on each side. These elements (ex.) therefore exhibit the same

disposition as the exoccipitals of a frog, and must be homologous

with the latter. ‘They are pierced by the foramen for the vagus

nerve, which is well seen on the left side. The posterior face of

each exoccipital immediately above the condyle is impressed with

a triangular fossa, so that the upper end of the bone, partly

obscured by matrix, has the appearance of bifurcating to unite in

jagged sutures with the two bones which descend from the

occipital row of roofing plates. The latter plates are evidently

superficial in origin—either dermal or splint ossifications—but

there is no clear line of demarcation between them and the

iunmediately underlying bones just mentioned. The inner half of

Text-fig. 34.

wt. op PP. soce. SE.

Capitosaurus stantonensis; hinder view of occiput, restored, two-thirds nat. size.

boce., supposed basioccipital; ¢., occipital condyle on exoccipital; ep., epiotic;

ex., exoccipital; fir., foramen magnum; op., opisthotic; psq., prosquamosal ;

pt., pterygoid; ptv., postero-temporal vacuity ; ¢7., quadrato-jugal; gw., quadrate ;

socc., space for supraoccipital ; s¢., supratemporals.

each plate of the median pair would directly overlap the carti-

laginous supraoccipital ; while its outer or lateral half alone is

underlaid by the bone (ep.) which occupies the place of a piscine

or amphibian epiotic. The inner two-thirds of each plate of the

outer pair is underlaid by the second bone (op.) of triangular

shape, which exhibits the usual relationships of an opisthotic.

As shown on the right side of the fossil, the jagged suture

between these two otic bones coincides exactly with that dividing

the two overlying plates. The pterygoids unite in a deeply jagged

suture with the broad basioccipital region, and then expand behind

on each side into a vertical lamina (pi.) which articulates with

1904. ] LABYRINTHODONT SKULLS. 173

the inner edge of the quadrate (qu.). The quadratojugal is in

contact with the quadrate only at its lower end, thus leaving a

rather large postero-temporal vacuity (ptv.) which is especially

well seen on the left side.

The complete occipital region, as here interpreted, is shown

in the accompanying restored sketch (text-fig. 34, p. 172). It

obviously conforms much more closely to the Amphibian type

than to the arrangement in any known reptile. It therefore

agrees with the evidence derived from the palate and the

supratemporal plates, showing that the Labyrinthodonts are

Amphibia.

The characters of the cranial roof, the anterior end of the

palate, and the teeth, prove that the skull now described belongs

to the genus Capitosaurus. It cannot, however, be referred to

any of the known species of this genus. It differs from the skull

of the type-species, C. arenaceus, from the Keuper of Franconia”*,

in the narrower and more tapering form of its rostral region, the

more elongated shape of the orbits, and its more nearly circular

pineal foramen. It is distinguished from the skulls of C. nasutus T

and C. robustus =, among other characters by the slighter excava-

tion of its occipital border and by the shape of the outer pair of

its supratemporal plates. It is also distinguished from the im-

perfectly known skull of C. fronto$ by the different shape of

its auditory notch and the coarseness of its external orna-

mentation. The new specimen therefore represents a hitherto

unrecognised species, which may be named C. stantonensis.

II. APHANERAMMA ROSTRATUM, gen. et sp. nov. (Plate XII.)

The imperfect Labyrinthodont skull from Spitzbergen was

found by Profs. J. W. Gregory and HE. J. Garwood in the Trias

of Sticky Keep, associated with a few short and deep, biconcave

vertebral centra, which probably belonged to the same animal.

It is broken into five pieces, which show little beyond impressions

of the bones. Three fragments are internal moulds, while the

other two pieces bear marks of the exterior. The right half of

the postorbital region and the middle portion of the rostrum are

missing. Enough, however, is preserved to indicate that the

skull must have been of a much elongated shape, with a slender

snout. It is very little depressed behind, and the occipital plane

slopes backwards. Its external surface was ornamented with

large reticulating rounded ridges, which were sometimes inter-

rupted, as shown by the impression of part of the left cheek

* Graf zu Minster, Neues Jahrb. fiir Min. 1836, p. 580; H. von Meyer, ‘ Die

Saurier des Muschelkalkes ’ (1847-55), p. 152, pl. lix. figs. 5-7.

+ H. von Meyer, Palwontogr. vol. vi. (1858), p. 222, pls. xxiv.-xxvi.

t+ H. von Meyer, ‘Die Saurier des Muschelkalkes’ (1847-55), p. 146, pl. lix.

figs. 1-4, pl. Ixi. fig. 10. Cyclotosawrus robustus, Lh. Fraas, Paleoutogr. vol. xxxyi.

(1889), p. 121, pls. ix., x., pl. x1. figs. 1-4.

§ H. von Meyer, Paleontogr. vol. vi. (1858), p. 229, pl. xxviii. fig. 2.

174 DR. A. SMITH WOODWARD ON [June 7,

(fig. 4). The grooves for the slime-canals must also have been

deep and conspicuous, as shown by the same fragment.

The most interesting feature in the skull, its quadrate bone

(figs. 1-3, qu.), is well preserved on the left side and exposed from

behind. This bone is long and narrow, antero-posteriorly com-

pressed, and inclined somewhat backwards. Its inner face is

sheathed by a great vertical plate of the pterygoid, while its

outer or lateral border is in complete contact with the bones of

the cheek. It is quite clear that there was no postero-temporal

cleft or vacuity; and the quadrate bears no impression of the

external auditory meatus.

There are no vacuities in the bony covering of the postorbital

region of the cheek, and the limits of some of its constituent

elements are well shown by the natural internal mould. The

long and narrow ovoid squamosal (figs. 1, 2, sq.) is distinct ; while

the outline of the equally elongated and still narrower postorbital

(pto.) can be traced without difficulty. There can also be little

doubt that the postfrontal (pif) and prefrontal (prf.) exclude the

parietals and frontals from the margin of the orbit. The parieto-

frontal region, however, cannot be satisfactorily interpreted.

It seems probable that the parietals (pa.), which must have been

relatively large, taper rapidly in front, where they extend nearly

to the anterior border of the orkit. They must also have been

either depressed or thickened in the median portion, where their

anterior end articulates with the very narrow pair of frontals

(/r.)— suggesting an arrangement nearly like that represented by

Cope in the skull of Cricotus*. The prosquamosal (psq.) is an

iregularly triangular plate, somewhat longer than deep, with

the apex forwards. The quadrato-jugal (qj.) is nearly twice as

long as its maximum depth. The jugal (j.) is especially elon-

gated and forms the greater part of the infero-external border of

the orbit (orb.), which is oval in shape and directed both laterally

and upwards. There are also some traces of the edge of the

postfrontal (pif), prefrontal (prf.), and lachrymal (/a.) bones,

but the middle part of the rostrum is lost. An internal mould

of nearly the terminal portion of the snout (figs. 6, 7) demon-

strates the slenderness of this region and the elongated shape of

the sasal bones (7.).

The palate is of the typically Labyrinthodont pattern, with a

complete parasphenoid (fig. 6, pas.), which is laterally compressed

in its middle portion. The large posterior lamina of the ptery-

goid, in a vertical plane, abutting on the quadrate, has already

been mentioned. In front of this expansion the pterygoid of

each side curves outwards to meet the jugal (and probably also

the hinder end of the maxilla) in a long suture. Its extent

anteriorly is uncertain owing to the absence of the middle of the

rostrum. Further forwards the palatines (fig. 6, pl.) are rather

* HW. D. Cope, Proc. Amer. Phil. Soc. vol. xvii. (1878), p. 529.

1904. | LABYRINTHODONT SKULLS. 175

broad plates; while the vomers (vo.) bound the antero-posteriorly

elongated posterior nares (pén.) both behind and within. The

only teeth observable are indicated by an impression of the middle

part of the pterygoid region (fig. 5), where they occur chiefly in

two close parallel rows, one on the outer edge of the pterygoid

(pt.), the other on the maxilla (mx.). The teeth of each series

are nearly uniform in size, but those of the pterygoid are some-

what larger than those of the maxilla, They are all cones with

the usual thickened and folded walls. There are also traces of an

irregular patch of minute teeth or tubercles still further back on

the pterygoid.

‘Three vertebral centra found in the same formation and locality

as the skull are deeply-biconcave discs, not pierced by any foramen

for the passage of a remnant of the notochord. One obliquely

crushed specimen is shown of the natural size in Pl. XII. fig. 8,

and another imperfect specimen is similarly represented in fig. 9.

The concavity at the end of the centrum is very slight near the

outer rim and suddenly deepens towards the centre.

It is difficult to determine the precise affinities of so frag-

mentary a Labyrinthodont skull, but if the vertebral centra are

rightly ascribed to the same animal, it evidently represents one

of the higher members of the Order. Though suggestive in some

respects of Cricotus, as already mentioned, the new skull differs

from that of Cope’s genus in the thickening or depression of the

middle part of the parietals, and in the very strong external

sculpture. The associated vertebral centra also differ from those

of Cricotus in being completed discs. Among other Labyrintho-

donts, the specimen from Spitzbergen seems to approach most

closely the skull of 7’rematosaurus*, with which it agrees in its

general shape, external sculpture, dentition (so far as seen), and

the relations of the quadrate bone. It is distinguished, however,

by the peculiar disposition of the parietal bones. The new fossil

thus represents a hitherto unknown genus, which may be named

Aphaneramma and defined as follows :—Skull elongate-triangular,

with the orbits widely separated and situated in its hinder half ;

external bones strongly sculptured, and grooves for slime-canal

deep. Parietal bones extending forwards between the orbits;

frontals very long and narrow: both these elements excluded

by the postfrontals and prefrontals from the orbital border.

A single regular row of small teeth on the pterygoid parallel

with the equally uniform row of teeth on the maxilla; clustered

small teeth or tubercles further back on the pterygoid. Vertebral

centra complete biconcave discs, not perforated. The type-

specimen of the type-species, A. rostratum, is the imperfect skull

now described.

* H. Burmeister, ‘Die Labyrinthodonten aus dem bunten Sandstein von Bern-

burg,’ pt. i. (1849). ‘The minute posterior pterygoid teeth are seen in a specimen of

Trematosaurus in the British Museum (no. R. 1733) and in one in Mr. W. HE.

Balston’s collection.

176 ON LABYRINTHODONT SKULLS. [June 7,

It may be added that vertebra, ribs, and other fragmentary

fossils from the Trias of Spitzbergen have already been referred

to Labyrinthodonts * ; but the specimens described are scarcely

sufficient for exact determination,

EXPLANATION OF THE PLATES.

Prats XI.

Capitosaurus stantonensis, sp. nov. (p. 170); from Lower Keuper, quarries of

Messrs. Peter Ford and Sons, Stanton, near Uttoxeter. [Brit. Mus. no. R. 3174.]

Fig. 1. Upper view of skull, two-thirds nat. size.

2. Occiput of the same, two-thirds nat. size.

Prater XII.

Aphanerammea rostratum, gen. et sp. nov. (p. 173) ; from Trias, Sticky Keep,

Spitzbergen. [ Brit. Mus. nos. R. 8180-82. |

Fig. 1. Upper view of imperfect hinder half of skull, internal mould, one-half

nat. size.

2. Left side view of the same, one-half nat. size.

3. Hinder view of quadrate region of the same, one-half nat. size.

4, Ornament of part of cheek of the same, showing slime-canal, one-half

nat. size.

5. Bases of teeth on maxilla and pterygoid of the same, nat. size.

6. Upper view of end of snout, internal mould, one-half nat. size.

7. Palatal view of the same, one-half nat. size.

8, 9. Imperfect vertebral centra, one-half nat. size.

LETTERING :—aw., opening for auditory meatus; boce., supposed basioccipital ;

¢., occipital condyle; ep., epiotic; ea., exoccipital; fin., foramen magnum ; fr.

frontal; j., jugal; da., lachrymal; ma., maxilla; ma., nasal; op., opisthotic;

orb., orbit ; pa., parietal; pas., parasphenoid; pin., pineal foramen; pl., palatine;

pme., premaxilla; prf, prefrontal; psq., prosquamosal; pé., pterygoid; ptf, post-

frontal; pén., posterior nares; pto., postorbital; ptv., postero-temporal vacuity ;

pe., premaxillary vacuity; g7., quadrato-jugal; qz., quadrate; soce., space for

supraoccipital; sqg., squamosal; sé¢., supratemporal ; v., vomer.

* N. Yakowlew, “Neue Funde von Trias-Sauriern auf Spitzbergen,” Verhandl.

russ.-k. min. Ges. vol, xl. (1902), p. 180, pl. iii. ; “ Nachtrag,” doc. cié. vol. xli. (1904),

pp. 165-169.

1904.| THE SECRETARY ON ADDITIONS 10 THE MENAGERIE. 177

November 15, 1904.

Dr. W. T. Buanrorp, C.LE., F.B.S., Vice-President,

in the Chair.

The Secretary read the following reports on the additions made

to the Society's Menagerie during the months of June, July,

August, September, and October, 1904 :—

The number of registered additions to the Society’s Menagerie

during the month of June was 149, of which 44 were acquired by

presentation, 20 by birth, 15 by purchase, 69 were received on

deposit and 1 in exchange. The number of departures during

the same period, by death and removals, was 121.

Among the additions special attention may be called to :—

1. A male Buffon’s Kob (Kobus kob), new to the Collection,

received on deposit on June 2nd.

2. A male Goral Antelope (Nemorhedus goral), presented by

Major Rodon, F.Z.S., on June 3rd.

3. A male Chimpanzee (Anthropopitheeus troglodytes), from the

Congo, purchased on June 5th.

4. A male Speke’s Antelope (Vragelaphus spekii), new to the

Collection, received on deposit on June 27th.

5. Two Ural Owls (Syrnivm wralense), veceived on deposit on

June 29th.

The number of registered additions to the Society’s Menagerie

during the month of July was 181, of which 74 were acquired by

presentation and 22 by purchase, 63 were received on deposit,

and 22 were bred in the Menagerie. The number of departures

during the same period, by death and removals, was 162.

Among the additions special attention may be called to :—

1. A Riippell’s Colobus (Colobus abyssinicus), deposited on

July 2nd.

2. Two Japanese Bears (Ursus japonicus), one presented by

Miss Violet M. Lakin on July 11th, and the other deposited on

July 28th.

‘3. A male Sing Sing Waterbuck (Kobus wnctuosus) ; 4. A male

Gambian Ourebi (Ourebia wigricaudata); 5. A male Ostrich

(Struthio camelus) ; and 6. Three Levaillant’s Parrots (Pocephalus

robustus): presented by Capt. Sir George Denton, K.C.M.G.,

F.Z.8., on July 14th.

7. Two young male Greater Koodoos (Strepsiceros ludw),

deposited on Suly 15th.

8. A Yellow-crowned Penguin (Ludyptes antipodum), and 9.

Two Rock-hopper Penguins (Ludyptes chrysocome), purchased on

July 23rd.

The number of registered additions to the Society’s Menagerie

during the month of August was 270. Of these 70 were acquired

Proc: Zoou. Soc.—1904, Vou. II. No. XII. 12

178 ON PERE DAVID’S DEER IN HAINAN. [ Nov. 15,

by presentation and 28 by purchase, 87 were born in the Gardens,

and 85 were received on deposit. The number of departures during

the same period, by death and removals, was 182.

Among the additions special attention may be called to :—

1. A female Lar Gibbon (Hylobates lar), presented by Mr. C.

R. Stokoe on August 9th.

2. A Capybara (Hydrocherus capybara), presented by Messrs.

the Liebig Extract of Meat Co. on August 18th.

3. Two young female Gorillas (Anthropopithecus gorilla),

obtained by purchase on August 19th.

The number of registered additions to the Society’s Menagerie

during the month of September was 156, of which 69 were acquired

by presentation, 10 by purchase, and 77 were received on deposit.

The number of departures during the same period, by death and

removals, was 142.

Among the additions special attention may be called to :—

1. A fine specimen of Wolf's Monkey (Cercopithecus wolfi),

deposited on Sept. 5th. This beautiful species is figured in

P.Z.S,. 1894, pl. vii.

2. An Aru-Islands’ Kangaroo (Jacropus brunii), deposited

Sept. 9th.

3. A pair of Kelp Geese (Chloéphaga antarctica), from the

Falkland Islands, presented by Mr. Vere Packe, Sept. 14th.

The registered additions to the Society’s Menagerie during the

month of October were 128 in number. Of these 76 were

acquired by presentation and 12 by purchase, 3 were born in the

Gardens, 36 were received on deposit and 1 in exchange. The

total number of departures during the same period, by death and

removals, was 163.

Among the additions special attention may be called to :—

1. Three Beatrix Antelopes (Oryx leucoryx), one from Aden, a

female, presented by Mr. G. W. Bury on Oct. 5th, and a pair from

the Persian Gulf, presented by Major P. Z. Cox, I.8.C., F.Z.8.,

on Oct. 22nd.

2. Two King Birds of Paradise (Cicinnurus regius), from New

Guinea, purchased on Oct. 15th. New to the Collection.

3. A new species of Moustache Monkey (Cercopithecus, sp. n.),

from the Gaboon, deposited on Oct. 15th.

4, A DuChaillu’s Monkey (Cercopitheeus nigripes), from the

Gaboon, deposited on Oct. 15th.

5. Two Schmidt’s Monkeys (Cercopithecus schnuidti), from the

Congo, deposited on Oct. 17th.

6. Two Blue-tailed Fruit-Pigeons (Carpophaga concinna), from

the Aru Islands, presented by Mrs, Johnstone on Oct. 17th,

Mr. R. Lydekker communicated the result of recent corre-

spondence connected with the sketch by a Chinese artist of a

Deer from Hainan purporting to be Pére David’s Deer (Zlaphurus

1904. | ON THE ANTLERS OF THE ALTAL STAG. 179

davidianus), recently exhibited to the Society (Proc. Zool. Soc. 1904,

ii. p. 83). When this sketch was exhibited it was understood to

have been drawn from a specimen in the possession of Mr. E. T.

C. Werner, then British Consul at Hainan, and was thus taken (in

spite of the addition by the artist of a pair of Peking-Deer antlers)

as evidence of the existence of Hlaphurus davidianus in Hainan.

In a letter dated June 26, Mx. Werner states, however, that the

artist drew the sketch from memory after the death of the animal,

and also that there was considerable doubt as to whether the latter

really was Pére David's Deer (Tst-pu-hsiang) at all.. Nevertheless

the writer expressed his belief that the species did exist in Hainan.

On the other hand, in answer to further enquiries, Mr. Hughes,

now Consul at Hainan, wrote that, so far as he could ascertain,

Pére David’s Deer was unknown in Hainan, and that the drawing

in question must be regarded as a fancy sketch. Under these

circumstances, despite the fact that the sketch undoubtedly

portrayed that species, no credence could at present be given to the

alleged occurrence of Pére David’s Deer in Hainan.

Mr. F. E. Beddard, F.R.S., called the attention of the meeting

to a fact in the life-history of Kangaroos concerning which but

little appeared to be known. An example of J/acropus dorsalis,

which died on the 4th Nov. last, was found to contain a young

one in the pouch which had survived the death of the parent.

The young kangaroo was 6 inches in length (to the root of the

tail) and still perfectly naked. On being removed from the

pouch, it moved its limbs vigorously and emitted a sound which

was rather more voice-like than a hiss.

It was difficult to describe the nature of the sound accurately.

It was uttered at continuous intervals. The production of any

sound in so imperfectly formed an animal was remarkable.

Mr. Frederick Gillett, F.Z.S., exhibited some antlers of the

Altai Stag (Cervus eustephanus), and made the following remarks

on their growth based on his own observations in the Society’s

Gardens :—

“An Altai Stag, purchased by the Society on Aug. 10th, 1897,

although in poor condition and not expected to live, in the

following year produced three sets of antlers and served two

hinds, becoming the parent of a stag and a hind. The young stag

shed his first pairs of antlers in a twelvemonth, and I am able to

show these antlers to-night. This young stag grew very large,

and when two years old bore antlers with twelve points. This

stag was then sold to Hagenbeck.

“The stag now in the Gardens shed a set of antlers on May 28th,

1902, having produced two sets in the year before ; and the three

sets which I exhibit to-night were shed by it on the following

dates :—Jan. 6th, 1903; June 10th, 1903; April 23rd, 1904.”

12 3k

“

180 DR. P. L. SCLATER ON THE OKAPI. [ Nov. 15,

Dr. P. L. Sclater, F.R.S., stated that in July last he had

visited Brussels in order to examine the specimens of the Okapi

(Okapia johnstoni) in the Museum of the Congo Free State at

Tervueren near that city. This he had been enabled to do by the

kind permission of M. Emile Coart, Conservateur du Musée du

Congo. The mounted series of the Okapi in that Museum

consisted of a fine adult pair, of which the male carried short

giraffe-like horns, as shown in a lithographic plate which was

exhibited, while the female had none, and of a pair of skeletons

in which the male had likewise horns but the female was horn-

less. There were also two other mounted specimens of immature

animals. Besides these specimens, Dr. Sclater was informed that

others had been sent from Tervueren by order of King Leopold to

the Museums of Tring, Paris, Stockholm, Madrid, Antwerp, and

Rome. All the specimens, as Dr. Sclater understood, had been

received from the Station of the Congo Free State on the Ituri,

which was practically in the same forest-district as Fort Mbeni,

where Sir Harry Johnston’s specimens had been obtained, although

the Ituri belonged to the water-basin of the Congo, and not to

that of the Nile.

Dr. Sclater also called attention to an article “ dus dem dunkel-

sten Africa,” published in the ‘ Basler Nachrichten’ for May 22nd

last, and subsequently abstracted in ‘Globus’ of July the 21st

last (vol. Ixxxvi. p.. 61), whereby it appeared that the writer,

Dr. T. T. David, a Swiss naturalist resident at Beni on the

Semliki, claimed to be the first European who has observed and

obtained an Okapi in its native wilds. Dr. David had sent one of

his specimens to Prof. R. Burckhardt, C.M.Z.8. (whose former

pupil he had been), for the Zoological Museum at Basel, but

Prof. Burckhardt had informed Dr. Sclater that it was unfortu-

nately received in a bad condition.

The following was an abstract of Dr. David’s principal remarks

in the ‘ Basler Nachrichten ’ :— ;

“The extremely elongated skull of the Okapi presents small rudiments of horns

on the frontal bones. The animal in life has the general bearing of a Tapir; it is

certainly a Ruminant, but its whole appearance, its actions in the swamps in which

it lives, its compressed body and the way in which it carries its head, remind one of

a Tapir and not at all of an Antelope, so that the stuffed examples of this animal in

London and Brussels are quite erroneously set up. The striping of the limbs is

much brighter than that of the Zebras. The back is red, especially so in the male ;

the ears are enormously large, and are furnished with great tufts of hairs standing

up. Small horns are present in some specimens, and, moreover, in both sexes, but

are absent in others, which induces me to believe in the possibility of the existence

of two species of Okapi. The underskin is as thick as in the Pachyderms, which

makes it a very difficult animal to prepare.”

Dr. Sclater concluded by saying that, notwithstanding what

Dr. David had stated and the views of Prof. Lankester and

Dr. Forsyth Major, he was quite unable to believe in the existence

of more than one species of Okapi in the same limited district,

though it seemed that the individual specimens presented some

unusual modifications.

1904. | PROF. J. C. EWART ON EAST-AFRICAN ZEBRAS. 181

Mr. W. B. Tegetmeier, F.Z.S., exhibited a specimen of an

Asiatic King-Crab (Carcinoscorpius rotundicauda) which had

been picked up alive off the Isle of Wight.

Prof. J. C. Ewart, F.R.S., exhibited some skins and a series of

lantern-slides of the Zebras of East Africa, and read the following

note on a form generally resembling in conformation and markings

the Mountain Zebra of South Africa :—

Some years ago Mr. Rowland Ward presented me with a stuffed

Zebra which, though originally “traded out of Somaliland,” has

a general resemblance to the Mountain Zebra of South Africa.

This Zebra (now in the Royal Scottish Museum, Edinburgh) is

in several respects so unlike the other forms hitherto described

that, without waiting for its exact habitat*, it may be worth

while pointing out how it agrees with, and differs from, the true

Mountain Zebra,

Text-fig. 35

Photo by G. A. Ewart.

Ward’s Zebra, to show long ears and face-stripes.

This Zebra (which may be known as Ward’s Zebra) very

closely resembles the Mountain Zebra in height, in the form

and size of the head, ears, and muzzle (text-fig. 35), in the mane,

* It probably inhabits part of the area between the upper reaches of the Tana

River and Lake Rudolf. :

182 PROF, J. C. EWART ON EAST-AFRICAN ZEBRAS. [ Nov. 15

tail, and hoofs, and in having the stripes over the rump arranged

to form the so-called gridiron pattern (text-fig. 36).

It differs from the Mountain Zebra in having a broad dorsal

band (31 inches wide as it crosses the croup), all the hairs of

which are directed backwards—in the Mountain Zebra the dorsal

band midway between the withers and the croup is represented

by a mere line, while from the croup to the mane the hair is.

directed forwards, i.e. the whorl usually at the end of the mane

in the Equide is on a level with the croup. The Mountain Zebra

differs also from Ward’s Zebra (1) in having a dew-lap, (2) in

having decidedly larger front chestnuts, (3) in having a larger

number of stripes running at right angles to the dorsal band,

(4) in having the legs more intensely striped, and (5) in the

ground-colour being nearly white: in Ward’s Zebra, with the

exception of the inner surface of the limbs and under surface of

the body, where white prevails, the ground-colour is of a rich

cream tint.

Text-fig. 56.

Photo by G. A. Ewart.

Ward’s Zebra, to show “gridiron” and broad dorsal band.

In Ward’s Zebra the stripes, except in the vicinity of the

muzzle, are of a dark brown colour, the muzzle and the nostril-

patches are darker than in the Mountain Zebra, and the stripes

above the nostril-patches are of a pale brown hue. The face is

PZ. S VIOG wollte lee ate

H.Gronvold del.et lth. Mintern Bros .imp.

SCOTONYVCREIRIS IWEADIFOIRD I,

1904. ] ON MAMMALS FROM FERNANDO PO. 183

decorated with four pairs of nearly symmetrically arranged stripes,

widest apart on a level with the eyes, and with four pairs of

stripes which meet in the centre of the forehead at or near the

point where the mane terminates four inches below the occipital

crest.

In the above-mentioned stripes, as in those on the sides of the

head and on the neck, there is close agreement between the two

Zebras under consideration, but, as already stated, there are fewer

stripes in Ward’s Zebra in connection with the dorsal band. If

the “gridiron” in the two forms is compared it will be noticed

that in Ward’s Zebra the bars running across the rump are

coarser than in the Mountain Zebra, apparently owing to the

obliteration of several of the intervening light spaces.

In text-figure 35 the colour and great length of the ears in

Ward’s Zebra are well brought out—the ears are longer than in

any of the Mountain Zebras I have had the opportunity of

measuring, and instead of presenting a white tip and a narrow

white band midway between base and apex as in the Mountain

Zebra, the apex is dark, while the proximal part is only faintly

and irregularly pigmented.

If one may judge by the ears, hoofs, and coloration, Ward’s

Zebra is adapted for a habitat similar to that of the Mountain

Zebra; moreover, like the Mountain Zebra, it has the reputation

of being stubborn and intractable.

The following papers were read :—

1. On Mammals from the Island of Fernando Po, collected

by Mr. H. Seimund. By Oxpriztp Tuomas, F.R.S.,

Ags

| Received July 13, 1904. |

(Plate XIIT.*)

(‘The complete account of the new genera and subspecies described in this communi-

cation appears here ; but since the names and preliminary diagnoses were published

in the ‘ Abstract,’ the genera and subspecies are distinguished by the names being

underlined.—EDrror. |

One of the chief desiderata of the British Museum collection

of Mammals has long been a proper series representing the fauna

of the Island of Fernando Po. For from this island there came

in the early days of the study of zoology by British workers

quite a number of specimens, and these were described in the

‘ Proceedings’ of this Society by Mr. G. R. Waterhouse and

others. But owing to age and exposure to light at a time when

the exhibition of types was not thought criminal, the original

specimens, on which all our comparisons depended, have become

so faded that but little use can now be made of them.

* For explanation of the Plate, see p. 187.

184 MR. OLDFIELD THOMAS ON | Nov. 15,

Attention having been again called to this island by the

remarkable ornithological discoveries made there by Capt. Boyd

Alexander, a special collecting-trip in the interests of the National

Museum was rendered possible by the generosity of our President

(the Duke of Bedford), of Mrs. Percy Sladen, and the Hon. Walter

Rothschild. A free passage to the island and back was also given

to the collector by Messrs. Elder, Dempster & Co., through the

kind offices of Sir Alfred Jones.

The collector, Mr. E. Seimund, started in November 1903,

arrived in the island on December 4, and left again in April 1904,

so that he had in all just over four months in which to collect.

The series he obtained is exceedingly valuable for the reasons

above mentioned, as he got good sets of nearly all the species

described so long ago by our predecessors in mammalogical

research ; and these cannot fail to be of constant service to all

workers on the subject.

Of novelties I have only had occasion to describe two—WScoto-

nycteris bedfordi, a Fruit-Bat, and Galago demidoffi poensis, a local

race of the little West-African ‘Galago; but Mr. Seimund has

found on the island several other mammals which had not

previously been recorded from there.

Our knowledge of the Mammals of Fernando Po rests chiefly

on the following literature :—

WarerHouse, G. R.—Descriptions of new Mammals from the

Island of Fernando Po, based on specimens presented by

George Knapp, Esq. P. Z. 8. 1838, p. 57.

Colobus, Cercopithecus, Genetia, Lutra, and Cephalophus.

WarerHouse, G. R.—Descriptions of new Mammals from

Fernando Po, obtained by Mr. L. Fraser during the Niger

Expedition. P.Z.8. 1842, p. 124.

Anomalurus and Squirrels.

ALLEN, W., and Tomson, T. H. R.—Narrative of the Expe-

dition to the River Niger. Appendix, vol. ii. pp. 472 et seqq.

1848.

Most of the Mammals obtained on the Expedition were

collected at Fernando Po by L. Fraser.

Bocace, J. V. Barsoza pu.—Subsidios para a Fauna da Ilha de

Fernao do Pé: Mammiteros. Jorn. Sci. Lisboa, (2) iv. p. 1,

1895.

16 species (none new) collected by Mr. P. Newton.

Bocace, J. V. BArBoza pu.—Faune des Quatre Isles du Golfe

de Guinée: Mammiféres. Jorn. Sci. Lisboa, (2) vii. p. 25,

1903.

Full list of species.

Descriptions of isolated species have also been published by

Gray, Bennett, Ogilby, A. Smith, and others.

The second of Prof. Bocage’s two papers gives a full list of the

1904. ] MAMMALS FROM FERNANDO PO. 185

Mammals of the island, and I have now, by intercalating the

additional species obtained by Mr. Seimund and modifying one or

two doubtful determinations, drawn up a list of the indigenous

species complete to date, with the names of the collectors on whose

specimens the species have been determined.

This contains 36 species, as follows :—

1. Colobus pennanti Waterh.

2. A satanas Waterh.

Bs polycomus Schr.* |

3. Cercopithecus erythrotis Waterh.

4.. o preussi Matsch.

5. Ss martini Waterh.

6. campbelli Waterh.

(6urnetti Gray).

a pogonias Benn.

8. Galago elegantulus Leconte.

9. 3 alleni Waterh.

HOS aS demidoffi poensis Thos.

11. Hypsignathus monstrosus Allen.

12. Rousettus stramineus Geoff.

13. Scotonycteris bedfordi Thos.

14. Rhinolophus landeri Mart.

15. Hipposiderus fuliginosus 'Temm.

16. Nycteris hispida Schr.

17. Mimetillus moloneyi Thos.

[| Glauconycteris poensis Gray F. |

18. Nyctinomus brachypterus Peters f.

19. Crocidura poensis Fraser.

20. Sylvisorex johnstoni Dobs.

21. Genetta poensis Waterh.

22. Poiana richardsoni Thos.

23. Lutra capensis poensis Waterh.

24, Anomalurus fraseri Waterh.

25. Sciurus stangeri Waterh.

265) 55 rufobrachiatus Waterh.

27. 9 punctatus Temm.

28. Funisciurus erythrogenys Waterh.

29. 55 poensis Smith.

30. Mus tullbergi Thos.

31. .,, alleni Waterh.

32. Cricetomys gambianus Waterh.

33. Procavia dorsalis Fraser.

34. Cephalophus ogilbyi Waterh.

35), $5 melanorheus Gray.

36. Manis tricuspis Raf.

1. Cotospus sATANAS Waterh.

Knapp, Thomson. ;

Knapp, Thomson, Newton, Seimund.

Knapp, Thomson, Fraser, Burton,

Seimund.

Knapp.

Thomson.

Knapp, Thomson, Fraser.

Burton, Newton.

Allen, Thomson, Burton.

Seimund.

Newton.

Newton, Seimund.

Seimund.

Thomson.

Fraser, Newton, Seimund.

Fraser, Capt. E. Downes, Seimund.

Seimund.

Downes.

Fraser, Seimund.

Seimund.

Knapp.

Thomson, Seimund.

Knapp.

Fraser, Thomson, Alexander, Newton,

Seimund.

Fraser, Thomson, Newton, Seimund.

Fraser, Thomson, Seimund.

Newton.

Fraser, Thomson, Seimund.

Smith, Thomson, Fraser, Alexander,

Seimund.

Allen, Seimund.

Smith, Newton, Seimund.

Fraser, Alexander, Newton, Seimund.

Fraser, Thomas, Alexander, Seimund.

Thomson, Fraser, Thrupp, Alexander,

Seimund.

Fraser, Newton.

Native skin. Bubi Town, Bantabiri, 500 m.

* There would appear to be some error in the inclusion of Colobus polycomus in

the Fernando Po fauna. Prof. Bocage puts it in on the authority of Gray, who

mentions two of Knapp’s skins as belonging to it. But the list of Knapp’s speci-

mens given by Waterhouse in 1838 does not include it, and until some confirmation

is obtained of its occurrence in the island I think it should be deleted from the list.

+ It is stated by Allen and Thomson (J. c. p. 480) that the specimen described

by Gray as Kerivoula poensis was not obtained in Fernando Po, but at Abo on

the Niger. It has therefore to be deleted from the island list.

{ N. pumilus Dobs. Of. de Winton, Ann. Mag. N. H. (7) vii. p. 38 (1901).

186 MR. OLDFIELD THOMAS ON [ Nov. 15,

2. CERCOPITHECUS ERYTHROTIS Waterh.

Cercopithecus erythrotis Waterh. P. Z.8. 1838, p. 59.

dg. 78,81. 9. 74. Bubi Town, Bantabiri, 500 m.

go. 147. @. 160. Bantabiri, 1800 m.

Well-known as it is by menagerie specimens, few Museums.

possess any wild-killed examples of this handsome monkey, and

the present specimens are therefore most acceptable. The original

types of this monkey, of C. martini, and of Colobus satanas were

native-made skins, presented to the Zoological Society’s Museum

by Mr. George Knapp in 1838, being the earliest zoological

specimens known to have come from Fernando Po.

3. CERCOPITHECUS PREUSSI Matschie.

2 (young). 165. N. Bantabiri, 1800 m.

Two native skins.

Not previously recorded from the island or represented in the

British Museum.

Dr. Matschie, during a visit to London, has examined these

specimens, and considers them to be the same as the species

described by him under the above name. His types came from

Victoria, Cameroons, and he tells me that this southern part of

the Cameroons has a fauna very like that of Fernando Po, such

species as Colobus satanas, Cercopithecus erythrotis, and others.

occurring there without modification. The presence of C. preussi

in Fernando Po is therefore not surprising.

4. GALAGO DEMIDOFFI POENSIS Thos.

Galago denudoffi poensis Thos. Abstr. P. Z.8. 1904, No. 10, p. 12,

Nov. 22.

3. 152, 153, 162. 9. 167. Bantabiri, 1800 m.

“Shot in tree by night.”—E. 8.

Similar in all essential respects to the true @. demidoffi of

continental West Africa, but the under surface is paler, whitish

instead of buffy. General colour above, of specimens in full pelage,

pale russet or cinnamon-brown. Central light line of face white,

contrasting more markedly with the general colour than the more.

or less buffy one of true demidofi. Hairs of under surface slaty

for two-thirds their length, then either white or pale buffy

yellowish. Outer side of limbs like body, a line along the inner

sides pure white; the hairs white to their bases, and forming

prominently white patches below the elbows and thighs. The

corresponding regions in true demidoffi are buffy or yellowish,

never pure white. Upper surface of hands and feet dull whitish.

Tail dark brown, darkening slightly terminally. Ears apparently

rather larger, such measurements as are available running from

27 to 30 mm., as against 24 to 27 mm. in demidoff.

Skull much as in demidofi, though inconspicuously larger.

Dimensions of the type, measured in the flesh :—

Head and body 130 mm.; tail 195; hind foot 46; ear 28.

1904. | MAMMALS FROM FERNANDO PO, 187

Skull—greatest length 38:2; greatest breadth 25-5; interorbital

breadth 5:4; breadth of brain-case 19°6 ; front of canine to back

of m® 12:5.

Type. Adult male, no. 152. B.M. No. 4.7.1.8. Killed 6 March,

1904, at an altitude of 1800 metres.

Compared with 15 well-preserved specimens of the Continental

form from localities ranging from the Gold Coast to Uganda, the

four skins obtained by Mr. Seimund differ so uniformly by the

lightness of their under surfaces, and the pure white of the inner

aspect of their limbs, that I think they should have a special sub-

specific name.

Galago demidoffi has not been previously recorded as occurring

in Fernando Po, though a skeleton obtained there was received

from Sir Richard Burton in 1862, just after the publication of

Gerrard’s ‘ Catalogue of Bones of Mammalia.’

5. RovusETrus STRAMINEUS Geoff.

SLOSS TNOn MASS: 117, VTS s Mosel S enor sll yi

113, 116, 120, 121, 122,123. Bantabiri, 10 m.

3g. 178,181. ©. 87, 105, 106, 179, 184. Bantabiri, 10 m.

“Very common.’—KE.8.

Prof. Bocage also records Hypsignathus monstrosus as having

been discovered in the island by Mr. Newton.

This fine series of specimens shows a peculiar and very unusual

sexual difference in colour which does not seem to have been

previously noticed. The males, without exception, are more or

less brown, the bright yellowish shoulder-patches contrasting

strongly with the general dark colour. The females on the other

hand are, both above and below, of the rich yellowish straw-colour

so often described as occurring in this species.

For the female of any animal to be more richly coloured than

the male is an unusual phenomenon.

6. ScoronycrEeRIs BEDFORDI Thos. (Plate XIII.)

Scotonycteris bedfordi Thos. P. Z.S. 1904, vol. i. p. 372.

@. 31. Fish Town, 10 m.

“Shot during the daytime, hanging on a tree.” —E. 8.

This most interesting Bat is the only new species obtained on

Mr. Seimund’s expedition, and I have thought it worthy of a

figure. It is the first member of the genus to be received by the

Museum, and is therefore a most welcome accession.

The previously known species, S. zenkeri Matsch., was described

from the Cameroons.

The external characters of S. bedfordi have been already

described, but the following measurements of its skull may be of

service :—Greatest length 25-4 mm.; basal length 22:6; zygomatic

breadth 16°5; interorbital breadth 4°8; breadth of brain-case 11-1;

palate length 14; front of canine to back of molar 87 ; front of

lower canine to back of m, 10.

The specific distinction of S. bedfordi rests mainly on the

188 MR. OLDFIELD THOMAS ON [Nov. 15,

conspicuously smaller size of the ears, these being only 11 mm. in

the island species and 17 in S. zenkeri. The skulls, judging only

by Dr. Matschie’s description, seem closely similar.

The cheek-tooth formule of Scotonycteris and Epomophorus are

considered by Dr. Matschie to be P. 3, M. 2, and that of Cynopterus

to be P. 3, M. 2; but the study of a young specimen of the last-

named genus shows that Dobson was perfectly right in giving its

formula as P. 3, M.3*. This young specimen has milk-premolars

present above the second and third cheek-teeth in each jaw, thus

showing them both to be premolars; the minute anterior tooth

has, as usual, no predecessor.

It would follow from this that the formula in Scotonycteris is

P. 3, M. 3, the same as that rightly determined for Hpomophorus

by Dobson.

But further, while correctly determining the teeth of Cynopterus,

Dobson does not seem to have realized that the same formula,

P. 3, M. 3, would certainly be applicable to Vyctymene t (“ Har-

pyra”), to which he assigns P. 2, M. 2.

In Dobsonia, on the other hand, with the same total number of

four cheek-teeth in the upper jaw, the tooth lost has obviously

been the anterior premolar instead of the last molar, so that the

formula should be, as Dobson puts it, P. 2, M. 2. Matschie

erroneously gives it as P. 2, M. 2.

7. HirpposipERrvs FULIGINOSUS Temm.

Go Oly OW, De Ti, Gs, MOO, leaiounalonires, 10) som.

OPV SS shishY Towne ltOim,:

3. 92. Sepopo, 10 m.

6. 04. Taka, 10 m.

“Very common.” —E. 8.

One of these specimens, a female, is bright orange, the others

are of the usual dark sooty brown.

8. NycreRIs HISPIDA Schr. (2).

3. 72. Bubi Town, Bantabiri, 500 m.

Not satisfactorily determinable in the dried condition.

9. MIMETILLUS MoLONEYI Thos.

Bo US Oe Os, OO, WAR rateAloria, IO wi,

“Shot on the wing. Flight swift and with many rapid turns

and twists.”

Mimetinyuus Thos.

Mimetillus Thos. Abstr. P. Z.S. 1904, No. 10, p. 12, Nov. 22.

Type. Vesperugo (Vesperus) moloneyi Thos. Ann. Mag. N. H.

(6) vil. p. 528 (1891).

* In the account of the genus Cat. Chir. B. M. p. 80; but in the synopsis of the

genera on p. 3 the formula is given by an oversight as P.2, M.2. ?

+ Cf. P. Biol. Soc. Wash. xv. p. 198 (1902), where, however, the name is accidentally

misprinted Nyctimene.

1904. | MAMMALS FROM FERNANDO PO, 189

Distinguished from Vespertilio (=Vesperus) by the abnormal

reduction in the size of the wings, which look insufficient to

support so large and heavy a body, and by the remarkable breadth

and flatness of the skull, which resembles in these respects that of

Tylonycteris.

Further study convinces me that this curious Bat, which I

described from a specimen sent home from Lagos by Sir A.

Moloney, should be separated generically from Vespertilio. Its

proportions are quite different from those of any other Bat. as is

shown by the fact that its forearm is barely half the length of

the head and body, its fifth finger barely longer than even this

short forearm, and its third finger is only as much longer than

the forearm as the fifth usually is. The result is that the develop-

ment of the wings recalls that in fetal specimens. The hind

limbs are also abnormally short.

In the original description the wing-membranes were said

to be uniformly brown, but this is a mistake due to the bad

condition of the type. Inward of the fifth finger they are

brown, but those between the third and fourth and fourth and

fifth digits are a transparent whitish, with a few brown spots

terminally.

The penis is remarkable in that it has no reversible prepuce,

the uncovered glans being long, conical, and covered with minute

reversed sete.

The skull, although larger, recalls that of Tylonycteris pach YPUs

by its broad and peculiarly flattened shape. It is not quite so

flat, but its anterior portion is even broader in proportion, the

anteorbital projections being unusually developed. Sagittal crest

practically absent, lambdoid crests strong. No distinct occipital

“helmet.” Median palatal spine longer.

Owing to its short velvety-brown fur and peculiar proportions,

this Bat has a strong superficial resemblance to a Vyctinomus or

Molossus rather than to a member of the Vespertilionide. Hence

the generic name suggested for it.

10. Crocrpura (Croc.) PoENsIS Fraser.

Crocidura poensis Fraser, P. Z.S. 1842, p. 200; Allen &

Thomson, Expedition to River Niger, il. p. 481 (1848).

¢. 36. 2.92. Bantabiri, 10 m.

3. 26. Fish Town, 10 m.

Q. 140. Bilelipi, 500 m.

This Shrew seems to be the same as that afterwards described

from Old Calabar by A. Murray, under the name of Rhinomus

soricoides, apparently in the belief that it was a rodent. His

type specimen, much discoloured, is still in the British Museum.

11. SytyIsoREX JOHNSTONI Dobs.

g. 94. 9. 68,103. Bantabiri, 10 m.

A separate skull (67).

190 MR. OLDFIELD THOMAS ON [ Nov. 15,

SyivrsorEx Thos.

Sylvisorex Thos. Abstr. P.Z.S. 1904, No. 10, p. 12, Nov, 22

Type. Crocidura morio Gray.

African Shrews with white teeth, four upper unicuspids, normal

mandibular dentition, and a short-haired tail without the long

bristle-hairs characteristic of Crocidura.

In 1887* Dr.G. E. Dobson, when describing from the Cameroons

the pigmy Shrew now found by Mr. Seimund in Fernando Po,

included it together with Gray’s Crocidura morio in the genus

Myosorex, » genus founded for the South-African Sorex varius

Smuts. The latter animal, however, is remarkable for the

possession of a minute extra tooth in the lower jaw, as discovered

and described by Dobson; and this character I think of such

importance as to necessitate the species which do not possess it,

but are in other respects allied to J/yosorex, having a special

generic name. Thisnew genus would include the species S. morio

Gray (type), S. gohnstont Dobs., S. sorella Thos., and S. muricauda

Mill

The four Fernando Po skulls of S. gohnstoni differ considerably

in the relative proportions of the upper unicuspids, the second

being much smaller than the third in some cases, as it is in the

type, while in others it is nearly as large. I am inclined to

believe that in the Soricidz generally the systematic importance

of the relative sizes of these teeth has been considerably over-

estimated.

12. PoIANA RICHARDSONI Gray.

Native skin. Bantabiri, 500 m.

13. ANOMALURUS FRASERI Waterh.

3. 58. @. 59,185. Bantabiri, 10 m.

6. 159. N. Bantabiri, 1800 m.

Although several Fernando Po specimens of 4, fraseri, including

the type, are in the British Museum, all are very much faded by

exposure to light, and these fresh topotypes are therefore of much

value.

Among the specimens assigned to this species from the mainland

of Africa are two from the Lower Niger, which a comparison with

Mr. Seimund’s examples shows to be subspecifically separable, as

follows :—

ANOMALURUS FRASERI NIGRENSIS.

Anomalurus fraseri nigrensis Thos. Abstr. P. Z.S. 1904, No. 10,

p. 12, Nov. 22.

Closely similar to the true fraser? in all respects, but the general

colour paler and greyer—body broccoli-brown, membranes smoke-

grey,—and the size, as shown by the skull and teeth, decidedly

smaller, The tail also less bushy.

*S JP a Se USS, Ds BB.

1904. | MAMMALS FROM FERNANDO PO, 191

Coloration of head, under surface, and limbs as in true fraseri.

Skull smaller and with rather a shorter narrower muzzle than

in frasert, the length of the tooth-row decidedly less.

Approximate dimensions of the type, measured in skin :—

Head and body 330 mm. ; tail 235; hind foot (s. u.) 57.

Skull—tip of nasals to back of parietals 53 mm.; zygomatic

breadth 38; nasals, length 14:7, greatest breadth anteriorly 7-4 ;

interorbital breadth 16; breadth of brain-case 26; palate length

from henselion 22°5; diastema 12°5; palatal foramina 6; length

of upper cheek-tooth series 11°9; lower jaw, incisor-tip to

condyle 37; length of lower tooth-row 13°8.

Hab. Abutschi, Lower Niger.

Type. B.M. No. 2.11.10.5. Collected February 1902 by Mr. A.

Braham. Two specimens, adult and immature.

Du Chailluw’s Anomalurus beldeni, from the Gaboon *, con-

sidered by Gray and Alston to be a synonym of 4. fraseri, appears

to me to be referable rather to the red-backed species commonly

known as A. erythronotus M.-Edw. Considering how widely

different in colour the two species are, it seems curious that there

should have been any doubt on the subject; but Du Chaillu’s

description is extremely vague, and it is only from his statement

that ‘on the back the hair is tipped with bright rufous, which

gives a rufous tinge from behind the ears to the lower third of

the body on the median portion to the commencement of the

membranes,” that [ am able to express an opimion on the matter.

This sentence, however, exactly expresses the dorsal coloration

of A. erythronotus, and the locality is approximately the same,

while no examples of A. f/raseri have been since recorded from

the district.

Tf I am right in this identification, the name A. beldeni

will have to stand for the red-backed species, as it antedates

A, erythronotus by many years.

14. Scrurus srancErt Waterh.

Gn Wd, Qe Bs, Lrsw Mopars NO) sei:

3. 73, 80. Bubi Town, Bantabiri, 500 m.

9. 95. Bantabiri, 10 m.

Se LOGs IGS Beli SeNliol aiiOM epee aN ebamtalourr,

1800 m.

Ge lZGaesilelipi 10m:

15. Scrurus RUFOPRACHIAtTUS Waterh.

Ow kon 2e 2h. 34, 37,47. Seeley 22223, 2, +095 » Wish

Town, 10 m.

3. 3,4, 6,10, 39. 2. 7,11, 40, 41,42. Santa Isabel, 10 m.

CO Ma S2AOiG NAD, VAL NGS me GF ih. 4 2. C9789) 269)

Bantabiri, 500 m.

3. 54,139. @. 134, 137. Clarence Mountain, 1800 m,

* P. Bost. Soc. N. H. vii. p. 303 (1861),

192 ON MAMMALS FROM FERNANDO PO. { Nov. 15,

16. FUNISCIURUS ERYTHROGENYS Waterh.

36. 29. ©. 14, 16, 20, 38, 43,50. Fish Town, 10 m.

dg. 91. @. 52. Lepopo Beach, 10m.

Q. 142. Bilelipi, 500 m.

6. 146. N. Bantabiri, 1800 m.

The last three species of Squirrel were all described, together

with <Anomalurus frasert, by Mr. G. R. Waterhouse in the

‘Proceedings’ of the Society for 1842, but during the long

interval since no further Fernando Po specimens of these have

been received. SS. stangeri and S. rufobrachiatus have proved to

be represented on the opposite mainland by forms not specifically

distinguishable from those of the island, while, on the other hand,

nothing to match /’. erythrogenys has been found elsewhere.

17. FuNIsciIuRUS POENSIS Smith.

36.31. @. 27,44. Fish Town, 10 m.

¢. 1,2, 8. Santa Isabel, 10 m.

18. Mus ratrrus L.

¢. Bantabiri, 10 m.

19. Mus rutuBerci Thos.

©. 64, 84,91. Bantabiri, 10 m.

Ge os Ode taka. VO Mme

@. 136. Clarence Mountain, 1800 m.

20. Mus ALLENI Waterh.

@. 101. Bantabiri, 10 m.

21. CRICETOMYS GAMBIANUS Waterh.

. 56, 77, 83, 85, 90, 102, 110. ©. 70, 98. Bantabiri, 10m.

. 129. Clarence Mountain, 1800 m.

Oy Ay

22. PROCAVIA DORSALIS Fraser.

3d. 156, 158, 158 bis. @. 71. N. Bantabiri, 1800 m.

g. 127, 128. Bubi Town, Bilelipi, 500 m.

©. 133. Clarence Mountain, 1800 m.

23. CEPHALOPHUS OGILBYI Waterh.

3d. 75. Bubi Town, Bantabiri, 500 m.

©. 151. N. Bantabiri, 1800 m.

Besides the type and other specimens obtained in the days of

Fraser, the Museum possesses a fine female example of this species,

presented by Capt. Boyd Alexander, who shotit at Moka in 1902.

It was by the examination of this specimen that. I was enabled to

distinguish the Fanti C. brooket from the present species *.

*% Ann. Mag. N. H, (7) xi, p. 283 (1903).

P.Z.S.1904, vol IL. Pl. XIV.

Bale x Danielsson, Ltd Sc.

Engel Terzi,del.

HYLOCHOSRUS MEINERTZHAGENI.

PA L044, voll Pl xe

3 | 5

Engel Terzi.del. Bale « Danielsson.L Sc.

HYLOCH@RUS MEINERTZHAGENTI.

1904.] ON THE FOREST-PIG OF CENTRAL AFRICA. 193

24, CEPHALOPHUS MELANORHEUS Gray.

Ons

® juv. 109. Bilelipi, 10 m.

3. 149, 155, 157, 164. 9. 150,154. N. Bantabiri, 1800 m.

3. 76. 9. 145. Bubi Town, Bantabiri, 500 m.

©. 60, 66. Bantabiri, 10 m.

EXPLANATION OF PLATE XIII.

Scotonycteris bedfordi, p. 187.

2. On Hylochaerus, the Forest-Pig of Central Africa.

By Onprinip THomas, F.R.S., F.Z.S.

[Received October 13, 1904. ]

(Plates XIV. & XV.*)

For some years, dating from the discovery of the Okapi, it has

been known to zoologists that the natives of the Semliki and

other Central African forests had stories to tell about a large pig-

like animal, of whose size and ferocity they gave rather highly-

coloured accounts. Such stories were first brought to Sir H.

Stanley 7 during his Emin relief expedition of 1888-90, and later

on to Sir Harry Johnston £ (who thought they might possibly refer

to a Pigmy Hippopotamus), to Mr. F. J, Jackson, My. W. D.

Doggett, and others.

More recently Lieut. R. Meinertzhagen, of the East-African

Rifles, hearing tales of this Forest-Pig, determined to secure

specimens of it for our National Museum, and it is to his per-

severance and generosity that we are indebted for the specimens

which form the subject of the present paper.

The following extracts from Lieut. Meinertzhagen’s letters to

Prof. Ray Lankester will show under what circumstances he

obtained the specimens here described :—

‘““T was on an expedition near Mount Kenya last February and

one of my men, who had been tracking cattle in the bamboos

about 8000 ft.), reported having killed in the forest a large animal

which he greatly exaggerated as to size. I sent him back next

dlay to see if he could bring in any of the beast. He found that

the Wanderobo had got one and had cut the animal up. He,

however, brought back two pieces of skin$, which was un-

doubtedly pig-skin, but of no pig with which I was acquainted.

The Masai know the animal well and call it ‘ Elguia.’ On moving

* Hor explanation of the Plates, see p. 199.

+ Of. Johnston, in Cornish’s ‘ Living Animals of the World,’ i. p. 267 (1902).

{ P. Z. 8. 1904, i. p. 228.

§ “It was a sow, as the natives had left 2 foetuses.”

Proc. Zoox. Soc,—1904, Vor. II, No. XITI. 13

194 MR. OLDFIELD THOMAS ON THE [| Nov. 15,

round the south-eastern slopes of Mount Kenya I found the

nearly complete body-skin (quite fresh), but cut into four pieces,

in a village, and also a large piece of old skin. These formed my

first consignment to England.

“Tn the following May I was in the Nandi country (E.N.E. of

the Victoria Nyanza), and in the forest I then again heard of

this pig from the natives. Some American missionaries had also

heard of it and had even seen examples. One of their men had

killed one, and I was able to get a skull and another piece of skin

which is identical with the pieces I got from Kenya. I eventually

got a portion of another skull from the same locality (c. 7000 ft.).

The natives all assert that the Pig is essentially a forest animal

and seldom comes into the open, which probably accounts for it.

not having been brought to the notice of sportsmen before.”

Tam also able to add the following extract from a letter written

by Mr. C. W. Hobley, C.M.G., Sub-Commissioner of the Uganda

Protectorate, to Dr. Chalmers Mitchell :—

“T heard of the existence of the Forest-Pig ‘ Hylocherus mein-

ertzhagent’ (as being a separate animal from the Wart-Hog or

Bush-Pig) from the Wanderobo hunting-tribe about a year ago,

and since then have as often as opportunity offered made efforts

to obtain the skin and skull of one. I promised a present of a

cow to anyone who brought me a complete skin and skull. All I

succeeded, however, in obtaining was a shield made of the skin of

this animal. Quite a number of shields made of the hide of this

pig are to be found in EK. Kakumega and Tiriki, and some years

ago I noticed these shields and inquired from what animal they

were derived; my informant, however, misled me by telling me it

was the hide of the ‘ Aard-vark’ or Ant-Bear, and it was only

about six months ago that I discovered my error.

“The Forest-Pig is, I find, well known by the Nandi, Masai,

and Wanderobo. The Masai equivalent is ‘ El Guya’; the Nandi

equivalent is ‘Tumtu’ (which, I believe, means /orest-dweller) ;

the Kakumega and Tiriki equivalent is ‘Mbirri’; some Wande-

robo call it ‘Tum’; there is, however, another word which I have.

for the moment mislaid.

“ All the various tribes acquainted with the beast are united

about its size; the Wanderobo assured me it was as large as a

zebra, and the Kakumega people, who do not know the zebra, said

it was as large as a small ox. These two comparisons agree very

well.

“The only skulls I have seen.are those which Lieut. Meinertz-

hagen obtained from the forest between Tiriki and Kabwaren, and

one which Mr. R. J. Church, of the Uganda Railway (now resident

at Nairobi), had obtained on Mauabout two yearsago. I have not

had an opportunity of comparing the latter with Lieut. Meinertz-

hagen’s specimens, but believe it to be the same; Mr, Church had

a head-skin of his specimen, and I believe it had a lot of white

1904. } FOREST-PIG OF CENTRAL AFRICA. 195

hair on the face. Mr. F. J. Jackson also saw Mr. Church’s

specimen, and may be able to verify this.

“The natives of Western Nandi, in the vicinity of the forest

from which Lieut. Meinertzhagen obtained his specimens, tell me

that before the great rinderpest plague of 1891 these animals

were numerous in the forests of W. Nandi, but the plague killed

nearly all; they spoke of its fierceness, and said it occasionally

attacked women who went into the forest to gather firewood.

“With regard to the distribution of the animal, I have heard

of it being found in the Leikipia forest, the Subugo Leldian on

the EH. side of the Rift valley, in the Kakumega forest, in the

Mau forest near Elgeyo, and I believe it will be found in the

Elgon forests.”

The specimens obtained by Lieut. Meinertzhagen consist of

(1) the imperfect skin, without skull, of a female from Mount

Kenya; (2) a perfect skull, with a piece of body-skin, of a

young adult male from Nandi, near the Victoria Nyanza, altitude

7000 ft.; and (3) an imperfect skull, without lower jaw, of an

old specimen from the last-named place. Specimen 2*, as being

a perfect skull, with a piece of skin, would naturally be selected

as the type of the species.

The first question that presents itself to every naturalist in

connection with such a form is as to whether the animal is most

nearly related to the ordinary Pigs, Sws and its African represen-

tative Potamocherus, or to the aberrant and highly specialised

Wart-Hog, Phacocherus, hitherto separated by a wide gap tT from

every other member of the family.

On a first superficial glance at the skull the answer to this

question would be that the new form was allied to Sus or

Potamocherus, and had nothing to do with Phacocherus, but

further study of the cranial and dental characters gradually

entirely removes such an impression, and indicates that the

animal is a link connecting the two groups, with an undeniable

and perhaps ancestral relationship to Phacocherus. It would, in

fact, appear to be a survivor of an intermediate stage in the spe-

cialisation of the Wart-Hog, its reduced incisors, enlarged upper

canines, complicated molars, and basisphenoid pits all showing

a relationship to that animal, although neither the canines nor

molars are so far advanced in their specialisation. On the other

hand, the general proportions of the skull and teeth are more as

* B.M. no. 4.11.5.14,

+ Gray and, following him, Flower recognised a special family, the Phacochceride,

for the Wart-Hogs, but no other writers have done so, and the present discovery

confirms the judgment of those who included all the Old-World pigs in the Suide.

¢ Attention may be drawn here to the figures and description of a fossil Algerian

pig said to show some relationship to the Wart-Hog, Sus phacocheroides P. Thomas,

Mém. Soe. Géol. France, (8) iii. art. ii. p, 10, pl. iv. figs. 1 & 2 (1884). It is, how-

ever, clearly different from the animal now described. For this reference, and much

other assistance in connection with the present paper, I am indebted to Dr. Forsyth

Major, whose intimate knowledge of the group has been freely placed at my service.

PEs

196 _ MR. OLDFIELD THOMAS ON THE [| Nov. 15,

in Sus and Potamocherus, while the formation of the parietal

region and the characters of the last upper premolar are peculiar

to itself. Of ordinary pigs it shows no special affinity whatever

with its geographical ally Potamochwrus, for it has neither the

characteristic rugosity of the muzzle, the specialised lower canine

section, nor the simple basisphenoid of that genus. In these

latter respects it agrees more with the Sus verrucosus group and

with the Babirussa, but even with them the agreement seems to

be rather in the common retention of primitive characters than

any real near relationship.

With regard to geographical distribution, I have little doubt

that this animal will be found to occur not only in the Kast-

African forests already mentioned, but throughout the great

Congo forest, just as Boocercus does; and Mr, G. L. Bates, the

well-known West-African collector, even tells me that as far west

as the upper waters of the Ja River, French Congo, the natives

speak of a large black forest-pig, which can hardly be anything

else but the present form.

Its habitat being therefore so typically a forest one, I have

proposed to call the genus Hylochwrus, while the species 1s

termed meinertzhagent (cf. ‘ Nature, vol. Ixx. p. 577, 1904), in

honour of the sportsman to whose efforts and generosity the

National Museum owes this interesting and important accession.

The following is a more detailed description of the specimens

before me :—

Body covered thickly and uniformly with black bristles about

3-8 inches in length, oval in section, about 0-4 mm. in the

greater and 0-3 mm. in the lesser diameter. Chest and groins

- with a certain number of whitish hairs. No evidence as to the

existence of a mane, nor are the ears or tail preserved in either

of the specimens.

The skull, as may be seen by the accompanying figures

(Pls. XIV. and XV.), has the general proportions of that of Sus or

Potamocherus, not the very peculiar ones found in Phacocherus.

The crown is very broad, concave above, parallel-sided, the breadth

between the intertemporal fosse approximately equalling that

between the orbits, a state of things very different from that found

in any other recent pig*. Occipital surface broader and lower than

in other genera, concave, its median line with a sharply defined

raised ridge running from the top of the foramen magnum to the

centre of the occipital crest; no such ridge is present in other

genera. Sides of nasal region sloping smoothly outwards, as in

Phacocherus, without any tendency towards the sharpened and, in

old animals, rugose edge characteristic of Potamocherus, though it

should be remembered that the only specimen showing this part

is rather immature. Socket of canines with a prominent longi-

tudinal crest above, probably much more heavily developed in old

age. Zygomata broad and heavy, thickened and strongly convex

* In the Pikermi Sus erymanthius, however, the proportions of the crown are

singularly like those of Hylocherus, widely as it differs in every other respect.

1904. ] FORESI-PIG OF CENTRAL AFRICA. 197

in front of and external to the orbits, this being one of the

few characters in which Hylocherus shows more resemblance to

Potamocherus than to Sus; but even here it is to be noticed that

the spring of the zygomata has not the remarkable abruptness

found in Potamocherus, and that Phacocherus has a still more

prominent zygomatic boss outside and below the eye. Base of

skull with two large sharply defined sphenoid pits, separated from

each other by the high knife-like vomer, the whole structure being

very like the arrangement generally found in Phacochwrus, though

there is considerable variation in this respect between different

specimens of the latter animal. The two skulls of Hylocherus

also differ from each other, as the pits are more deeply hollowed

out and sharply defined behind in the type than in the older

skull. Such pits, present to their full extent in Phacocherus,

are also found, though smaller, in Babirussa, one of the most

primitive of pigs. There is no trace of them in Potamocherus

nor in typical Sus as represented by the Sus scrofa group;

but in the peculiar Sus verrucosus group there is an indication

of a hollowing out in the same region, and in one specimen of

S. verrucosus amboinensis Major there is a deep pit in the

pterygoid on each side of the vomer—evidently a remnant of the

same structure. But this pit is absent in another specimen of

the same form. Bulle reaching up rather more than halfway

towards the hamular processes, thus intermediate between the

low bulle of Phacocherus and those of Potamocherus, which

attain the level of the hamular processes.

Lower jaw posteriorly about as in Potamocherus, not elongated

upwards as in Phacochwrus, but anteriorly it is broad and spatu-

late as in the latter genus, in order to accommodate the widely

splayed canines.

Dentition.—The permanent incisors, as in Phacocherus, are one

in number above on each side and two below, 1° and i’ above and

i, below (all present in Sus and Potamocherus) having been lost.

But they are larger than in the Wart-Hog, and are probably

never shed, as so commonly happens in old individuals of Phaco-

cherus. Moreover, on one side of the single mandible available,

the milk-predecessor of i, is still in place, the retention of the

milk-tooth when the permanent one has been aborted being an

intermediate stage between the complete set of Sus and the

reduced one of Phacocherus. Young examples of this latter do

not show any trace of a milk i,.

The canines present several characters of interest, and are in

many ways of an annectant nature. The upper ones are conical.

very thick and heavy basally, indeed as much so as in Phaco-

cherus, but they taper more rapidly, and in the example before

me do not much exceed in length those of ordinary swine. The

reason of the comparative shortness of the upper canines of Sus

is that they are so placed that the friction with the lower canines

cuts them obliquely across, the lines of the hinder edge of the

upper and front edge of the lower pair, when viewed from above,

198 MR. OLDFIELD THOMAS ON THE [ Nov. 15,

meeting each other at a sharp angle, so that no part of the upper

tooth clears the face of the lower one, On the other hand, the

line of the teeth in Phacochwrus is such as to carry the ends of

the upper pair quite clear of the lower ones. Now in Hylocherus

we find an intermediate condition of set and curvature, and

consequently the upper teeth project a little beyond the ends of

the lower ones, without being so conspicuously developed as in

Phacocherus.

The lower canines are as widely splayed as in Phacocherus, or

even slightly more so, the angle that the line of one makes with

that of the other, when viewed from the front, considerably

exceeding a right angle, this angle slightly exceeding a right

angle in Phacocherus, and falling considerably short of one in all

other pigs. In section the lower canines are of the more primi-

tive shape found in the Sus verrucosus group, the outer face being

nearly equal to the inner, the hinder face being nearly transverse

and passing into the wearing surface without noticeable angle.

[In Potamocherus and the Sus scrofa group the outer face is

much narrower than the inner one, and the posterior face is

directed obliquely outwards and is at a conspicuously different

angle to the surface worn against the upper canine.| Both outer

and inner faces have a median longitudinal ridge, really slight but

appearing conspicuous owing to its being worn white as compared

with the general black colour of the tooth; speaking strictly,

therefore, owing to these median ridges, the section of each canine

is not a triangle but a pentagon.

The premolars appear at first sight to be three in number

above, as in ordinary swine, but closer examination shows the

remarkable fact that in both specimens the true last premolar (p’)

has been entirely suppressed, the tooth standing in its place

being—in the old as well as in the immature specimen—its milk-

predecessor (mp’). This latter tooth is very similar in shape to

the mp* of Phacocherus, in which animal, however, the normal

tooth-change takes place. But in the lower jaw, curiously enough,

mp, has fallen as is usual, and is replaced by p,.

This suppression of p* is a most peculiar character, but occur-

ring in two specimens it does not seem permissible to suppose it

is only an abnormality, a point that in any case future material

will decide. Should it prove a normal character of the animal,

it would indicate that in this one respect Hylocherus has gone

further in the suppression of its anterior cheek-teeth than even

Phacocherus.

Below (one specimen only) there is but one premolar present,

P,» an unusually simple tooth, without secondary cusps of any sort.

In front of it there are the roots of some additional premolars,

but it is not possible to say how many of them have ever been

developed, and for this we must await the advent of younger

specimens.

The molars, above and below, are neither of the complicated

bunodont structure of those of Sus and Potamocherus on the one

1904. | FOREST-PIG OF CENTRAL AFRICA. 199

hand, nor of the specialised hypsodont nature of those of Phaco-

cherus on the other, but at least it may be said that they present

a basis out of which the latter might have been formed. Their

structure will be better seen by the figures than by a detailed

description, though attention may be drawn to the resemblance of

m’ to that of Phacocherus, and to the development of the median

secondary cusps of the lower teeth, these having each its obvious

homologue in the complicated hypsodont molars of the derivative

form,

The measurements of the typical skull are as follows (the

older imperfect skull is only infinitesimally larger, so that the

type has evidently reached its full size) :-—

Greatest median length above 415 mm.; basal length 355 ;

zygomatic breadth 213; nasals, length 235, breadth 52; inter-

orbital breadth 95; tip to tip of postorbital processes 124; inter-

temporal breadth 92; breadth across lateral occipital protuberances

116; height from basion to top of occipital crest 123; least breadth

of maxillary zygomatic processes 57; breadth across sockets of

canines 130; breadth between tips of canines 217; palate length

252; least palatal breadth (between m’*) 44; basal diameter of

canine (c.) 34; horizontal length of p’ 9, p? 12°5, mp* 14, m* 18°5,

m? 26, m® 38.

Lower jaw—length (bone only) 320; breadth across symphysis

at base of canines 111; least breadth across diastema 82; height

at diastema 45; tip to tip of canines 200; lower canines—hasal

diameter of outer face 19, inner face 20, posterior face 12°5;

horizontal length of p, 15, m, 19, m, 26°5, m, 41.

In conelusion, I may congratulate Lieut. Meinertzhagen on the

interest and importance of his capture and ourselves as zoologists

on the discovery of so fine a ‘ missing-link” as Hylocherus proves

to be. And I would express the hope that before long specimens

of this animal may be received in a condition fit for mounting,

while further skulls of different ages will elucidate the develop-

ment of its dentition.

EXPLANATION OF THE PLATES.

Prate XIV.

Hylocherus meinertzhageni.

Skull of type ; upper, lower, and lateral views.

Prate XV.

Hylocherus meinertzhageni.

Figs. 1, 2. Skull of type, anterior and posterior views.

Fig. 3. Upper cheek-teeth, right side, of type.

4. Lower ditto.

5. Upper cheek-teeth, right side, of older specimen.

200 DR. P. CHALMERS MITCHELL ON [ Nov. 15

3. On the Species of Crowned Cranes. By P. Caanmers

Mircuett, M.A., D.Sc., Secretary to the Society.

[Received November 15, 1904. |

(Text-figures 37-40.)

[The complete account of the new species described in this communication

appears here; but since the name and preliminary diagnosis were published in

the “ Abstract,’ the new species is distinguished by the name being underlined.—

Epitor. |

I have had the good fortune to see no less than fifteen living

examples of the Crowned Crane in the course of this year, and

the obvious differences between two types Neestallats included

under the name Balearica pavonina led me to examine the

literature on the subject and the skins in the National Museum,

as well as some shown me by private friends, with the result that

I believe I am able to make a slight addition to our knowledge of

these beautiful birds.

The two major species of this group were figured by George

Edwards, the ‘ Library-Keeper” to the Royal College of Physi-

cians, in his ‘ Natural History of Birds’ (vol. iv. p. 192), published

in 1751. They were not definitely named but described as the

‘“‘ Crowned African Cranes”: the figure in the foreground, which

he supposed to be that of the male of a pair, isa good representation.

of the Cape Crowned Crane; while the other figure, designated

the female by Edwards, is an excellent figure of a West-African

Crowned Crane. The latter figure shows the darker coloration

of the neck and back, and the division of the bare cheek-area into

nearly equal white upper and pink lower half characteristic of the

West-African form; while the other figure shows the grey

coloration of the upper part of the body and the very large

pendent neck-wattle equally characteristic of the Cape form. It

will be more convenient to group my subsequent remarks under

the names of the species.

Balearica regulorum (Benn.). The Cape Crowned Crane.—I

follow Reichenow (Die Végel Afrikas, vol. i. p. 265) in using this

name instead of B. chrysopelargus of the B.M. Catalogue. The

latter name depends on Lichtenstein’s ‘Catalogus Rerum Natu-

ralium Rarissimarum,’ but the specimens on which Lichtenstein

founded his descriptions are not known, and the description of

Ardea chrysopelargus is far too vague to be applied with eertainty

to this or any other Crowned Crane. I have seen four living

examples (of these, three are at present in the Gardens at Regent’s

Park) and a number of skins. The feathers of the neck and back

are Silvery grey. The most striking distinctive characters of the

four species, however, are to be found in the heads, of which I

give outline figures. The large, bare cheek-patch (text-fig. 37,

p- 202) is divided into a small, upper, roughly triangular area which

is bright red in colour; the lower part of the patch, which usually

1904. | THE CROWNED CRANES. 201

follows the curve of the orbit, is characteristically white in the

adult, but may be slightly suffused with pink, as in one of the

specimens at the Gardens. Even in that case, however, and in

dried specimens from which the colour has faded, the two areas

are clearly marked off from one another. The pendent neck-

wattle is very large and bright red. The beak is black, and, as

in all the others, the part of the head not occupied by the crown

is covered with a dense velvety black patch of feathers. The

characteristic ‘‘ crown” in all the species is composed of erect,

bristle-like feathers, each of which has a spiral twist. The crown

is golden or straw-coloured, but in this species each bristle is

tipped with black, and the black often extends a considerable

way down the bristles, darkening the crown. On the surface of

the skull, in the temporal region, is a pair of very strong bony

knobs, described by Mr. Beddard (Proc. Zool. Soc. 1904, vol. ii.

» BI),

: The species is widely distributed in South Africa and extends

a considerable way northwards in East Africa, Reichenow (Joc. cit.

p. 266) gives its northern range as ceasing with the Pangani

River, near Zanzibar. Mr. C. W. Hobley, however, tells me that

a Crowned Crane is abundant in Uganda, and has kindly given

me the head of a specimen from Kavirondo. This undoubtedly

is that of the Cape Crane. Mr. Ogilvie-Grant was kind enough

to show me a fine skin from a similar locality which was also that

of B. regulorum, so that a much more northern range must be

associated with this species.

Balearica gibbericeps Reich.—This species is certainly closely

allied to B. regulorwm, and Reichenow in his most recent work does

not regard it as more than a variety. I have seen neither living

examples nor skins, but reproduce here (text-fig. 38, p. 202), by

Dr. Reichenow’s kind permission, the figure of the head published

in his work ‘ Deutsch-Ost-Afrika,’ Vogel, p. 47. The neck-

wattle is large as in the species just described, and the general

coloration of the head, neck, and back is similar. The striking

difference is the extension forwards and upwards of the bare

cheek-patch on either side, so as to invade the black velvety

patch in a rounded knob-like process. Various localities are

given for this species in the region extending northwards from

the Pangani River towards Uganda, but some of these seem

ascribed to it on the supposition that the Cape Crowned Crane

does not extend northwards of the Pangani. It appears, however,

that the two species overlap, and further specimens of B. gib-

bericeps and information about its exact distribution and relation

to B. regulorum are much to be desired. It is with the hope of

obtaining these that I have copied Reichenow’s figure and borrowed

from his description.

Balearica pavonina (.). The West-African Crowned Crane.—

I have seen seven living specimens and several skins of this

species. Five of the living specimens are at present in the

Gardens—three brought from Nigeria by Lt.-Col. Jackson, one

(=)

Text-fig. 37.

Head of Balearica regulorum (ved area of cheek-patch dotted).

Text-fig. 38.

ANNE Une

edi,

Head of Balearica gibbericeps.

DR. P. CHALMERS MITCHELL ON | Nov. 15,

1904. ] THE CROWNED CRANES, 203

Text-fig, 39,

SS ee

Head of Balearica pavonina (red area of cheek-patch dotted),

Text-fig. 40.

Head of Balearica cecilie (ved area of cheek-patch dotted).

204 ON THE CROWNED CRANES. [ Nov. 15,

obtained from the same locality by Miss Jardine, and another by

Dr. Macfarlane. George Edwards’s figure, already referred to, is

a representation of this bird. It is as large as B. regulorwm,

but much darker in colour, the neck and back, although really a

dark grey, looking almost black in comparison with those of the

Cape Crane. The crown isalmost identical with that of the latter.

The beak is similar, but is horn-coloured towards the tip. The

skull has similar, although smaller bony knobs, but is much

broader and has a larger cranial capacity. The neck-wattles are

red, but, although varying in size, are much smaller than in

L. regulorum, and are visible only when the bird faces the

observer. The bare cheek-patch (text-fig. 39, p. 203) is divided by

a nearly horizontal line extending backwards from the middle of

the orbit into a smaller upper portion which is white, and a

larger lower portion which is red, so that the disposition of the

colours is inverted compared with that in B. regulorum and

B. gibbericeps. The range of this species is given by both

Reichenow and the B.M. Catalogue as extending from Senegal

across to the Upper Nile and Abyssinia. It happens, however,

that all the specimens I have seen, alive or in museums, of this

type (excluding the type about to be described) belong to the

Western portion of this range.

BALEARICA CECILL# Chalmers Mitchell. (TheWhite Nile Crane.)

Balearica cecilie Chalmers Mitchell, Abstr. P. Z. 8. 1904,

No. 10, p. 13, Nov. 22.

The White Nile Crane appears to be the smallest of the Crowned

Cranes as yet known. It is rather darker than the West-African

form, and the crown, especially in the living specimens, is paler

in colour. The beak is shorter and entirely black; the skull and

head are relatively smaller, but the velvety helmet is broader, and

temporal bony knobs appear to be absent. The cheek-patches (text-

fig. 40, p. 203) are rather more rounded than those of L. pavonina

and the disposition of colour is similar, but the upper white

portion is very much smaller, so that at first sight the whole

patch appears to be much redder. The neck-wattles are red, and

small as in B. pavonina.

Early this year Lady William Cecil deposited in the Gardens in

Regent’s Park four Crowned Cranes which she had obtained from

the White Nile, near Khartoum. At first sight they corre-

sponded with Balearica pavonina, and they were registered as

examples of that species. On comparing them with our other

specimens, however, | saw that they differed, and on pointing out

their interest to Lady William Cecil, that lady was kind enough

to present two to the Society. On further investigation, I thought

it necessary to make these birds the type of a new species, which

I propose to associate with the name of the donor. In the some-

what poor collection of skins of Crowned Cranes in the National

Museum, I found only two belonging to this region of Africa—one

1904. ] ON THE MOUSE-HARES OF THE GENUS OCHOTONA. 205

from Fashoda (Hawker Collection), and one from Aboo Zeit,

White Nile, sent by Captain Stanley Flower; and these two were

identical specifically with Lady William Cecil’s specimens *.

Type in the Gardens of the Zoological Society of London,

presented by Lady William Cecil.

Examples of three species have been living throughout the

summer and autumn in the same paddock on the canal-bank in

the North Garden. The three examples of 5. regulorum and one

example of B. pavonina keep together, and the similarity in size is

obvious, while the different coloration of the neck and body and

of the cheek-patches and the large wattles in the Cape form

amply distinguish the species. The two examples of B. cecilie

keep together and away from the others. They are smaller,

darker in the body, lighter as to the crowns and conspicuously

redder as to the cheeks. It would be unwise to attach too much

importance as to the natural grouping of birds in a menagerie,

but it is striking that B. pavonina consorts with B. regulorum,

although the specific distinctness of the two has long been

admitted, and not with B. cecilie, with which it has hitherto been

confounded.

4, On the Mouse-Hares of the Genus Ochotona.

By J. Lewis Bonuore, M.A., F.L.S., F.Z.8.

{Received July 18, 1904.)

[The complete account of the new species described in this communication appears

here; but since the name and preliminary diagnosis were published in the ‘ Abstract,’

the species is distinguished by the name being underlined.—Enrror. |

A large and valuable series of Ochotona from Kashmir, recently

sent home by Col. A. E. Ward, has induced me to take up and

study the whole genus, so far as the Palearctic Region is con-

cerned, with the results given below.

Owing to the fact that these animals live in countries dificult

to reach and, for the most part, inhospitable, the series of skins is

somewhat meagre. Nevertheless I have been enabled to come to

certain conclusions which may serve as a basis for the future

study of the group.

Exclusive of the American forms, the genus is found in

Southern Russia, extending northwards through Persia, Afghan-

istan, Kashmir, Thibet to N.E. Siberia.

Many of the species are closely allied and some, if not all, have

both a summer and a winter pelage. I propose in the first place

to divide the genus into three groups, which may be recognised

by the shape of the incisive and palatal foramina.

* [Since the reading of this paper, four more Crowned Cranes from the White Nile

have been deposited at the Gardens. These are examples of B. cecilie.|

206 MR. J. LEWIS BONHOTE ON THE | Nov. 15,

In the first group, which may be called the

Ladacensis Group, the incisive and palatal foramina are sepa-

rate and distinct, the lower and posterior portions of the

premaxille, although very thin, meeting in the middle line.

In the second group, which may be known as the

Rufescens Group, there is no bony division between the incisive

and palatal foramina, but the incisive foramen is narrow

and slightly constricted at its posterior end, whence it

suddenly broadens out into what represents a large palatal

foramen.

Lastly, there is the

Curzonie Group, in which the large single foramen is practically

triangular in shape, with little or no constriction to mark

the division between the incisive and palatal foramina.

The following is a list of all the names belonging to this genus

under the special groups to which they belong, as well as the

type-locality from which they came. In the body of the paper,

however, I have considered some of them as synonyms or

subspecies.

I. Ladacensis Group.

Type-Locality.

Ochotona alpina ......... Altai.

O. hyperborea.........-.---- N.E. Siberia.

O. normalas “s... 6.0. cases. )

Os FAR UTR AI ossonsico Mouth of R. Maia, N.K. Siberia.

O. cinereoflava .......-.++- :

QB DISGER | Sasa edaso sosonaee

OP IO ATEIRTELOS WERE Riss eee S. of C. Tschukosky.

O. badacensis |.2+.s0c neces Ladak.

OT OU AD OG eos erie es ou os ose 830 Beyond L. Baikal and Desert of

O. erythrotis: ........5.0+-+- Gannsu, N. Thibet. [| Gobi.

Oertlae Bo eS Vernoe Mts., Turkestan.

II. Rufescens Group.

Os TUpeSCONS, i --205- 4-2 Cabul, Afghanistan.

OO SLO RRR se cin oS sotk ete eee N. Thibet.

O. goustlla, | 2.02.2 0008 24020 S.E. Russia and 8.W. Siberia.

OP LAN, Facad. <6) bee ee Kashmir.

III. Curzonie Group.

ON CUPZONME (cecce cnn necen Sikkim.

O. melanostoma .........++- Kuku-noor, 8. Thibet.

Qi QOQUTIGD o.oo spo nsgeniae Dauria.

QEUNGCTIOOIS. a. oe access Doba, Turkestan.

Dis TUPI. SRR eee Pangong Lake, Ladak.

Cugi@S2D eRSeseRBeaesesseS: Kuenluen R., 8. of Sanga Pass.

DsGOUU2S | aocSSeseeeaeenedss Choor Mts.

O. nepalensis ........++-+++- Nepal.

OI ETETSCTIOON. ERR Oe Ladak.

ON mbeianan seen Moupin, N.W. Szechuen.

1904. | MOUSE-HARES OF THE GENUS OCHOTONA. 207

Key to the Species.

A. Palatal and incisive foramina distinct.

a. Bars small, 24 mm. or less.

a2. Colour of upper parts uniform dull rufous.

a3, Large. Hind foot 35 mm. ........2-ccccceceeeeeeeeereeeeee O. alpina.

63. Smaller. Hind foot 26mm. ... sper en aides (OL ANA OTE

67. Colour of upper Pe white or yellowish.

a, Ears rufous ......... Peas eee On agcens7s.

Bo Wars) wititengess sees esos sees mane eee ee een alae O. ogotonda.

b1, Ears large, 281mm. Colour greyish; head and shoulders

rufous.

a2, Post-auricular patch white .............00cc0cceessceesseseene O. rutile.

62. Post-auricular patch red ...... O. erythrotis.

B. Palatal and incisive foramina not distinct.

a. Combined foramen narrowing in centre.

a”, Colour of upper parts uniform.

a, Size large: colour white or ello ish .. .... O. koslowi.

b3, Smaller: colour brown ..........-..20:c000--2-ceeesreeeeeeene O. pusilla.

62. Colour of upper parts not uniform.

a3. Colour brownish, with white collar behind ears ...... O. rufescens.

23, Colour greyish, with red head and shoulders in

summer ........ PB pe OTT

B. Combined foramen not narrowing in n centre.

a2. Ears small, 23 mm. or less.

a3, Size small. Hind foot 25 mm. .... propre OLIN FTOLE.

53. Larger. Hind foot not less than 28 mm. j

as, Colour light.

a. Mouth “black oss doses sages eee easton 0 AO) HeLa OStOMmas

Be= Mouth White: <cscc.<co0.0sec eneestanices-sedeasssassepaaa p ORAMIIICU,

64, Colour darker.

a. Uniform pale brown .... O

Beier ads . CUrzonie.

6°. Dark brown, rufous on head and shoulders. ...... O. rvoylei.

jizy Whee) leneya, PY (aittiny, \ Bapesseeceeano yeas e aaoese csocnacce, | OL MACOS

OcHOTONA ALPINA (Pall.).

Lepus alpinus Pallas, Glires, p. 52, pl. 2 (1778); id. Reise, i1.

p. 701, tab. A (1773); Schreber, Saugthiere, iv. p. 911, pl. 238

1792

ON ys alpinus (Pall.) Cuvier, Rég. Anim. p. 219 (1829);

Waterh. Mamm. ii. p. 15 (1848); Radde, Reisen Siid. v. Ost-

Sibirien, i. p. 232 (1862).

General colour of a uniform reddish brown, sometimes inclined

to greyish. Fur long, soft, slate-grey at the base for about three-

Bye ths of its length, the terminal portion being greyish or whitish

with darker tip. Ears of moderate size, rounded, and thinly clad

with hairs similar in colour to the body-hairs. Under parts

uniform yellowish white, being sometimes rather more rufous

across the breast.

The skull is stoutly built, but long and narrow. The incisive

foramina are small and rounded, rather than narrow and elongate

as is the case with other species. The muzzle is broad and stout,

and the teeth large and strong.

Dimensions (approx. from skin), Head and body 175 mm.;

hind foot 35; ear 20.

Skull. Greatest length 56 mm.; basal length 46; zygomatic

breadth 25; length of nasals 18; length of molar series 10;

interorbital breadth 6.

Habitat. Originally described from specimens from the Altai.

208 MR. J. LEWIS BONHOTE ON THE [ Nov. 15,

I have been unable to find any further particulars as to its

range.

Our knowledge of this species is so scanty that it is impossible

to give any information concerning any changes of pelage it may

undergo, but it does not appear to ever become very light. There

is aspecimen in the British Museum of a uniform deep dark ruddy

brown, marked ‘ Siberia, melanistic variety,” but further material

may prove this to be a normal pelage of the species.

OCHOTONA HYPERBOREA (Pall.). __

Lepus hyperboreus Pallas, Zoogr. i. p. 152 (1831).

Lagomys hyperboreus (Pall.) Wagner, Schreber, Siugeth. Suppl.

iv. p. 121 (1844); Waterh. Mamm. i. p. 30 (1848); Schrenck,

Amurlande, i. p. 147 (1859); Radde, Reisen Siid. v. Ost-Sibirien,

1. p. 232 (1862).

Lagomys hyperboreus, varr. normalis, ferruginea, cinereoflava,

fusca, Schrenck, Amurlande, i. p. 148 (1859).

Lagomys littoralis Peters, SB. Ges. naturf. Fr. Berlin, p. 95

(1882).

This species closely resembles the foregoing, except in its much

smaller size. The five specimens from various parts of Eastern

Siberia which are in the British Museum show a uniformity

quite unusual among members of this genus. The general

colour above is a light brownish rufous (mummy brown, Ridgw.),

which is practically uniform throughout the upper parts, becoming

rather purer rufous on the flanks owing to the absence of black

tips to the hairs. The under parts are of a uniform rufous-white.

The ears are small and scantily covered with whitish bairs.

Skull. Except in its size the skull bears a close resemblance to

that of O. alpina.

Dimensions (approx. from skin). Head and body 160 mm.;

hind foot 26; ear 14.

Skull. Zygomatic breadth 20 mm.; length of nasals 11; length

of molar series 7; interorbital breadth 5.

Habitat. Described by Pallas as inhabiting N.E. Siberia.

Schrenck has recorded it from the mouth of the river Maia on

the mainland opposite Sakhalin, and I have examined specimens

from Ussuri, Kentei Mountains in N. Mongolia, and Yakutsk.

Schrenck describes, under the names given in the synonymy,

several varieties of this species. There can be little doubt as to

the specific identity of these varieties, and they probably repre-

sent the different pelages assumed by this species, but from the

material at my command [I am unable to give any further

information.

I also provisionally place O. littoralis, from a similar locality,

under the same name, as a large amount of material will be

required before these various forms can be elucidated.

OcHOTONA LADACENSIS (Giinth.).

Lagomys ladacensis Giinth. Ann. Mag. Nat. Hist. ser. 4, xvi.

p- 231 (1875); Blanford, J. A. 8. B. xliv. p.110(1875); id. Yarkand

1904. } MOUSE-HARES OF THE GENUS OCHOTONA. 209

Mamm. p. 71, pl. vi. fig. 1, pl. vii. fig. 2, pl. vila. fig. 1 (1879);

id. Faun. Brit. India, Mamm. p. 458 (1891); W. L. Scl. Cat. Cale.

Mus. p. 110 (1891); Biichner, Mamm. Przew. i. p. 185 (1890).

Lagomys curzonie Stoliczka (nee Hodgs.), J. A. 8. B. xxxiv. 2,

p- 108 (1865); Anders. P.Z.S. 1871, p. 562.

General colour light brownish grey, many of the hairs, more

especially along the dorsal region, tipped with dark brown. Head

vather lighter, with dull rufous patch extending from nose to

the crown. Ears moderately large and rounded, clothed externally

with rufous hairs, longer on the inner margin. Under parts

yellowish white, with a trace of a rufous collar, which is also

visinle on the shoulders. Feet clothed with white haus.

The coat just described is that of an individual shot on the

22nd July, and represents a very typical skin ; there are, however,

other forms of pelage which it would be as well to notice, although

I am unable to say exactly in what sequence they may be found.

Specimens in June have, as a rule, very short woolly rufous

couts, sometimes shading to pale yellow and interspersed with

long black hairs.

In September a moult takes place and the new coat is long and

thick and of a warm brownish grey, becoming lighter and paler

along the sides. The rufous markings mentioned in the first

pelage described are present and visible but not so intense.

There is another specimen from the same place and date as the

first mentioned, but the pelage is rather shorter, more woolly,

and not so grey.

In a specimen in fresh pelage at the beginning of October the

hairs are a deep slate-grey at the base, and shade through vinous

to pale white, ending in a buff subterminal ring and a black tip.

IT am of opinion that there is only one moult in the year,

namely in September, and that during the summer the pelage

gradually wears away, giving the animals the different appear-

ances which I have noted above.

Of the skull not much need be said as, except in the foramina

already alluded to, the specific differences between the skulls in

this group are very slight. As, however, the figure given by

Biichner (pl. xxiv. fig. 8) is rather misleading, it may be men-

tioned that the palatal foramen, as shown in fig. 2, much more

resembles that of the type of ladacensis. In other respects

fig. 8 accurately represents ladacensis, and it is unfortunate that

the specimen figured should have been slightly abnormal in an

important feature.

The dimensions of an adult male are :—Head and body 180-5

mm. 3; hind foot 35; ear 24.

Skull. Greatest length 49°5 mm.; basal length 39; zygomatic

breadth 25; length of nasals 15; length of molar series 10;

interorbital breadth 5.

Habitat. Originally described from Ladak. This species has

also been found to the N.E. in Northern Thibet. It is seldom

found at a lower elevation than 14,000 feet.

Type. B.M. 75. 3.30.2. Collected by Col. J. Biddulph.

Proc. Zoou, Soc.—1904, Vor. IT. No. XIV. 14

210 MR. J. LEWIS BONHOTE ON THE [ Noy. 15,

OcHorona oGorona (Pall.).

Lepus ogotona Pallas, Glires, p. 59, pl. 3 (1778); id. Zoogr. i.

p. 157 (1811); Schreber, Siugthiere, iv. p. 915, pl. 239 (1792),

Lagomys ogotona (Pall.) Cuvier, Reg. Anim. p. 219 (1829);

Waterh. Mamm. ii. p. 17 (1848); Radde, Reisen Siid. v. Ost-

Sibirien, i. p. 226 (1862).

Lagomys pallasi Gray, Ann. Mag, Nat. Hist. ser. 3, xx. p. 220

(1867).

Closely allied to O. dadacensis, from which it differs in its much

greyer and lighter colour; the general colour above being of a pale

whitish grey, below white. There is a total absence of the rufous

hairs on the ears so conspicuous a feature in the foregoing species.

The skull differs in no marked respect from that of O. ladacensis.

Dimensions (approx. from skin). Head and body 165 mm.

(6" 7'” as given by Pallas); ear 21; hind foot 35.

Skull. Zygomatic breadth 25 mm.; length of nasals 16; length

of molar series 10; interorbital breadth 3:5,

Habitat. Mountains beyond Lake Baikal and Desert of Gobi.

I have examined only one individual of this species, which,

except for its colour, very much resembles O. ladacensis, of which.

it may eventually prove to be merely an Eastern race.

OcHOTONA ERYTHROTIS (Biichn.).

Lagomys erythrotis Biichner, Mamm. Przewalski, 1. p. 165

(1890); id. loe. cit. pls. xxi. & xxiv. figs. 1-6 (1894).

The following description is taken from Biichner, as I have been

unable to procure a specimen of this species. In summer the

general colour above is of a dull rusty red; the head is similar in

colour to the rest of the body, but the frontal region is somewhat

lighter and the lips yellower. The ear is thickly clothed on both

sides with ferruginous hairs. The chin is yellowish, the chest

reddish, and the remainder of the under parts and feet white.

In its winter pelage, the lips and tip of the nose are white,

the sides of the face and frontal region are whitish brown. The

ears and a small patch immediately behind them are of a rusty red.

The remainder of the body above and below is greyish white

interspersed with blackish or brownish hairs.

The skull, compared with that of O. ladacensis, is rather smaller ;

the muzzle is stouter and shorter, but the skull itself rather nar-

rower. According to Biichner, except for its narrower and more

slender build it closely resembles that of O. vuwtila, the next species.

Dimensions (after Biichner). Head and body 245 mm.; ear 28;

hind foot 42.

Skull. Greatest length 46°5 mm.; basal length 37; zygomatic

breadth 23°7; length of nasals 15:8; length of molar series 8.

Habitat. Gannsu and Burchan Budda Mts., N.E. Thibet.

This species, although clearly belonging to the ladacensis group,

shows, in its external colouring, affinities towards Col. Ward’s new

species, to be hereafter described.

1904. | MOUSE-HARES OF THE GENUS OCHOTONA, 211

OcHOTONA RUTILA (Severtz.).

Lagomys rutilus Severtzoff, Voy. 1873; id. Ann. Mag. Nat. Hist.

ser. 4, xvill. p. 168 (1876); Blanf. Mamm. Yark. Miss. p. 79

(1879); Schaff, Zool. Jahrb. (Syst.) ii. p. 65 (1887); Biichn.

Mamm. Przew. i. p. 160, pl. xx. (1890); id. loc. cit. p. 191 (1894).

This species apparently differs so slightly from the foregoing

that it is doubtful whether they can be considered as more than

geographical races.

A male shot on the 21st June in Turkestan has the whole of

the back behind the shoulders of a whitish grey. The whole

of the head and shoulders, with the exception of the ears, is of

a lighter rusty (lighter, according to Biichner, than in 0. ery-

throtis). The ears are large, being about the same size as those

of O. macrotis, and covered on both sides with short grey hairs,

while a small patch behind the ears, which in 0. erythrotis is

always red, is in this species always white, and forms the most

conspicuous feature distinguishing these two species. The feet

are grey; the under parts whitish, with a more rufous collar

round the throat.

The skull shows hardly any features to distinguish it from the

foregoing.

Dimensions (approx. from skin). Head and body 196 mm.;

hind foot 39; ear 28.

Skull. Greatest length 50 mm.; basal length 42; zygomatic

breadth 24; length of nasals 16; length of molar series 9°5;

interorbital breadth 6.

Habitat. Vernoe Mountains, Turkestan.

It seems to me very doubtful whether these last two species

can really be regarded as distinct, but our knowledge of them

is at present so slight that it seems best meanwhile to consider

them so. In the large ears this species resembles O. macrotis

Giinth., while it is perhaps instructive to note that superficially

the difference, ¢. g. presence or absence of white postoral patches,

between erythrotis and rutila is very similar to that between

roylei and wardi.

OcHOTONA RUFESCENS (Gray).

Lagomys rufescens Gray, Ann. Mag. Nat. Hist. x. p. 266 (1842);

Hutton & Blyth, J. A.S. B. xv. p. 140 (1846); Waterh. Mamm,

ii. p. 20 (1848); Horsf. Cat. E.-I. Mus. p. 149 (1851); Blyth,

Cat. p. 133 (1863); Blanf. E. Persia, p. 83, pl. 6. fig. 2 (1876);

Wood-Mason, P.A.S8.B. p. 173 (1880); Scully, J. A.S. B. lvi.

p- 76 (1886); Murray, Ann. Mag. Nat. Hist. ser. 5, xiv. p. 100

(1884); Radde, Zool. JB. iv. p. 1053 (1889); W. L. Scl. Cat.

Mamm. Cale. Mus. ii. p. 111 (1891); Blanf. Faun. Br. Ind.,

Mamm. p. 458 (1891).

In its winter pelage (Oct.) this species is of a uniform whitish

brown, somewhat paler on the sides and of a pale yellowish buff

beneath. Each hair is slate-grey for its basal two-thirds and

14*

212 MR. J. LEWIS BONHOTE ON THE [ Nov. 15,

then dirty white, with a subterminal buff ring and dark brown

tip. The dark tips are absent on the hairs of the feet and under

parts, and absent or inconspicuous on a patch behind either ear,

which patches tend to meet across the nape. On the frontal

region the buff subterminal ring extends down to the grey base

and is more rufescent. In summer the pelage is similar, but the

colour on the back and head tends to be more rufescent, and that

of the flanks and under parts a purer white. The white patches

behind the ears are larger, and have coalesced on the nape forming

a broad white collar, succeeded posteriorly by a rufescent collar

of about half its width, which gradually merges into the general

reddish colour of the remainder of the body. This rufescent

collar starts from two maroon patches situated on the under side

of the neck, and may thence be traced upwards and backwards,

becoming paler and more rufous in colour, to meet over the

shoulders in the middle line. The maroon patches form a dis-

tinctive feature of this species, but they are not always to be seen

in the winter pelage.

The skuli may best be described by comparing it with that of

O. ladacensis, from which it differs in being broader across the

muzzle. The bulle are more rounded and swollen, thereby nar-

rowing the basioccipital. The main difference, however, lies m

the incisive and palatal foramina, which are not, as in O. ladac-

ensis, separated, but form one large foramen slightly constricted

about one-third of the way from its anterior end. The portion

anterior to this constriction is of uniform width and narrow, the

posterior portion gradually widening out throughout its length.

This distinction forms the difference between the two groups.

Dimensions (approx. from skin). Head and body 175 mm. ;

hind foot 32; ear 22.

Skull. Greatest length 52 mm.; basal length 42; zygomatic

breadth 24; length of nasals 16; length of molar series 10;

interorbital breadth 3. .

Habitat. The type came from Barber’s Tomb, Rocky Hills, near

Cabul; but its range extends throughout Afghanistan, extending

into Persia and Transcaspia.

The white collar and maroon patches on the throat form

characters by which this species may readily be recognised, but

even apart from these it does not bear a very close resemblance to

any of the species which have hitherto been described.

OcHoToNA KosLow! (Biichn.).

Lagomys koslowi Biichner, Mamm, Przewalski, i. p. 187 (1894).

Size rather large. General colour of the upper parts, which is

uniform throughout, pale whitish buff tinged with vinaceous.

Under parts and feet, which are thickly furred, white. Hach

hair is grey for about half its length at the base, the terminal

half being vinaceous buff fading to whitish or ending in a dark

brown tip, which last is never sufficiently conspicuous to modify

1904. | MOUSE-HARES OF THE GENUS OCHOTONA. 213

the general colour. Intermixed with the fur proper are some

long black bristles. I fancy that in its changes of pelage this

species closely follows O. ladacensis, for at the end of summer it

is of a bright golden buff due to the wearing off of the terminal

portions of the winter pelage described above. The long black

bristles, however, do not wear down and thus become more

conspicuous.

The skull, which is well figured by Biichner, is rather short in

the muzzle, giving it a broad and thick-set appearance. The

muzzle is both short and narrow, and possibly in correlation with

this we find the anterior terminal portion of the nasals tending

to turn upwards to a marked extent and having a broad vertical

portion. The foramina are as in O. rufescens.

Dimensions of skin (after Biichner). Head and body 240 mm. ;

hind foot 42; ear 19-5.

Skull. Greatest length 44 mm.; basal length 33; zygomatic

breadth 27-2; length of nasals 14°3; length of molar series 10.

Habitat. Northern Thibet.

This species may easily, apart from skull-characters, be dis-

tinguished from O. ladacensis, with which alone it could be

confused, by the ears being yellow and not rufous, the tips white

not black, and the under parts snowy white and noé yellowish.

OCHOTONA PUSILLA (Pall.).

Lepus pusillus Pallas, Glives, p. 37, pl. 1. (1778); Schreb.

Siugth. iv. p. 906, pl. 237 (1792).

Lagomys pusillus (Pall.), Desm. Mamm. p. eo (1820); Cuv .

Régne Anim. p. 219 (1829); Waterh. Mamm. ii. p. 19, pl. 1

fig. "9 (1848).

This is the smallest species of the genus as yet known. ‘The

general colour above, which is uniform, is dark brown, grizzled

with white where the whitish median portion of each hair shows

through. Under parts white; ears and feet grizzled, the former

having a conspicuous ring of white hairs growing from their

inner margin.

The skull.—From the material at hand Iam unable to say much

about the skull, which is small but well proportioned, the muzzle

being rather stout and short. The foramen is very typical of the

group in which I have placed it.

Dimensions (approx. from stuffed specimen). Head and body

145 mm.; hind foot 27; ear 14.

Skull. Zygomatic breadth 20 mm. ; length of nasals 12; length

of molar series 7 ; length of palate from henselion 8:5.

Habitat. 8.K. Russia and thence eastward to Siberia.

Material is so scarce that it is impossible to say much concern-

ing this species, which may be recognised by its small size and

the white rims to the ears, in which characters it approaches

O. hodgsoni from Kashmir.

214 MR. J. LEWIS BONHOTE ON 'THE [ Nov. 15,

OcHOTONA WARDI Bonhote.

Ochotona ward Bonhote, Abstr. P.Z.S. 1904, No. 10, p. 13,

Noy. 22.

In the summer pelage, worn from June to September, the whole

of the head, shoulders, and fore part of the body (excepting a

small patch behind the ears, which is white) is bright chestnut

(cinnamon-rufous, Ridgw.), becoming more vinaceous on the throat.

The remainder of the upper parts is dark greyish rufous, each

hair being black at its base with a light subterminal annulation,

the tips being either dark or rufous. This latter colour encroaches

greatly on the light portion often to its total exclusion, especially

on the sides of the body. The under parts are white lightly

washed with pale buff. The feet are of the same colour. Ears

moderate in size, very scantily clothed with hair.

In the winter pelage this animal is of a uniform dark iron-grey

all over, with the exception of the light patches behind the ear,

which are white as in summer. Under parts dull white. Slight

traces of rufous are generally to be found at the base of the

shoulders, on the crown of the head, and along the flanks. The

young resemble the adults in winter, but are slightly browner in

general colour and have the rufous on the head and shoulders

more marked.

The skull is very similar to that of the type of O. roylei, as

figured in the original description, and does not show any great

features of note. The combined foramen, while having the narrow

anterior third and the slight constriction typical of the rufescens

group, shows a tendency for the constriction to become less

marked, but it can nevertheless be clearly made out in every

example.

Dimensions of type in flesh. Head and body 187 mm.; hind

foot 25; ear 22°5,

Skull. Greatest length 44 mm.; basal length 37; palatal length

17; length of foramen 12; zygomatic breadth 21; interorbital

breadth 5; breadth of brain-case 17 ; length of molar series 9.

Habitat. Talien, Kashmir, 11,000 feet.

Type (in Coll. Brit. Mus.). A. E.W. No. 56. Ad. d. Collected

on the 8th August, 1903.

In external appearance this species most nearly resembles

O. roylei; the latter, however, is much darker and lacks the

conspicuous white patches behind the ears.

OCHOTONA CURZONIA Hodgs.

Lagonys curzonicee Hodgs. (nec Stoliczka) J. A.S. B. xxvi. p. 207

(1858); Giinth. Ann. Mag. Nat. Hist. ser. 4, xvi. p. 230 (1875);

Blanford, Fauna Br. Ind., Mamm. p. 457 (1891).

This is the first species of the group which I have called after

it and which is very closely related to the rufescens group. The

difference between the palatal foramina of the two is well shown

by a comparison of the figures of O. koslowi (Biichn. Mamm,

1904. ] MOUSE-HARES OF THE GENUS OCHOTONA. 215

Przewalski, pl. xxiv. fig. 14) and of O. dawrica (ibid. pl. xxv.

fig. 2); in which it may be noticed that the sides of the foramen

in O. daurica diverge at once from their anterior point instead of

continuing parallel for the first third of their length, and then

tending to approach again before finally diverging, as in the

rufescens group.

O. curzonie is a pale buff-coloured animal above and rather

lighter below. Along the median area of the back the hairs

are tipped with black and have a subterminal ring of rufous

brown, the extent of the rufous varying in individual specimens.

Behind the ears is a clear patch of a rather deeper buff than

the rest of the body. The ears are of moderate size, clothed on

both sides with fairly long whitish hairs. Feet pale buff.

The skull of this species is small and narrow. Apart from the

palatal foramen, of which mention has already been made, the

chief point of note is the postorbital process of the zygoma, which

is very long and narrow; the posterior nares are similarly modified.

Dimensions (approx. from skin). Head and body 170 mm. ; hind

foot 28; ear 19.

Skull. Palatal length 15 mm.; zygomatic breadth 20; length of

nasals 12; length of molar series 8; interorbital breadth 4.

Habitat. The type-locality of this species is the Chumbi Valley

in the north of Sikhim, whence it apparently extends westward as

far as Kashmir.

The series of this species at my disnosal is so small, that I am

unable to give any particulars of its seasonal changes should any

occur, but from the specimens before me it appears to be a very

uniform species.

OcHOTONA MELANOSTOMA (Biichn.).

Lagomys melanostomus Biichn. Mamm, Przewalski, 1. p. 177,

pl. xxii. (1890).

Except in its slightly larger size I can find, after careful com-

parison of a co-type of melanostoma with the types of curzonia,

no other distinguishing characteristics between these two species,

and possibly a larger series of the latter would prove them to be

identical. Biichner, in his original description, had apparently

overlooked O. cuwrzonie, as he only distinguishes it from O. daurica,

from which it differs in its yellowish under parts and black muzzle,

The general colour of the winter pelage above is sandy brown

grizzled with darker brown or blackish. Each hair at its base

is grey, shading to light brown and ending in a dark tip.

Interspersed over the upper parts are long uniformly black hairs,

The under parts are dirty yellowish white.

In summer the hairs wear down so that the light-brown

subterminal rings become more conspicuous, the black tips being

nearly or quite worn away; the long and uniformly black hairs,

however, remain, so that the grizzled appearance is not altogether

lost, but the animal becomes brighter and browner and the under

parts tend to become of a purer white.

216 MR. J. LEWIS BONHOTE ON THE [ Nov. ies.

The skull, except in size, does not differ from that of O. ew

Zone.

Dimensions (after Biichner). ?Head and body 235 mm.; hind

foot 34; ear 21.

Skull. Greatest length 41 mm.; basal length 36; palatal length

16; zygomatic breadth 21:5; length of nasals 13; length of molai

series 9; interorbital breadth 4.

Habitat. Kuku-noor and Gannsu, N. Thibet.

OcHOTONA DAURICA (Pall.).

Lepus dauricus Pall. Reise, 11. 1776, p. 692.

Lagomys dauricus (Pall.), Biichn. Mamm. Przewalski, i. p. 172,

pl. xxii. fig. 1 & pl. xxv. figs. 1-5 (1890).

The general colour of this species is a very pale buff, lighter

on the flanks and rather yellower along the centre of the back

and over the forehead. Hach hair is dark slate-grey at the base,

succeeded by a whitish portion, increasing in colour towards the

tip, which is sometimes dark brown. Lars well covered with

whitish hairs; feet white. Under parts pure white, having

occasionally a yellowish collar round the neck.

The skulls at my disposal are too fragmentary for a detailed

description. Biichner, however, gives a good figure, from which

it appears to differ but little from the nearly allied species. Its

most marked features are the bulle, which are large, prominent,

and rounded.

Dimensions (after Biichner). Head and body 220 mm.; hind

foot 14:5; ear 19.

Skull. Palatal length 18 mm.; length of nasals 14:5; length

of molar series 8°5.

Habitat. Originally described from Dauria.

Much confusion seems to exist over this species, which closely

resembles at least three others, viz. O. curzonice, melanostoma, and

ogotona.

The black muzzle of O. melanostoma serves at once as a distin-

guishing character, while from O. curzonie the paler colour of the

present species as well as the longer and softer coat form characters

by which it may always be distinguished.

_ From 0. ogotona, to which it bears a greater external resem-

blance, and with which it has been confounded by Biichner, it

may be distinguished by its much smaller size, while the skull-

characters are very distinct.

OcHoTonA MACROTIS (Giinth.).

Lagomys macrotis Giinth. Ann. Mag. Nat. Hist. ser. 4, xvi. p. 231

(Sept. 1875); Blanford, Yark. Mamm. p. 75 (1879); Scully, Ann.

-Mag. Nat. Hist. ser. 5, vol. viii. p. 100 (1891); id. P.Z.S. 1881,

p. 207; W.L.Scl. Cat. Mamm. Ind. Mus. p. 110 (1891); Blan-

ford, Faun. Br. India, Mamm. p. 457 (1891).

Lagomys auritus Blanford, J. A.S8. B. vol. xliv. p. 111 (Oct

1904. ] MOUSE-HARES OF THE GENUS OCHOYONA, 217

1875); id. J. A.S. B. xlvi. p. 326 (1877); id. Yark. Mamm. p. 74.

‘pl. vi. fig. 2, pl. vii a. fig. 2 (1879).

Lagomys griseus Blanford, J. A.S. B. vol. xliv. p. 111 (Oct.

1875); id. Yark.-Mamm.; p. 77, pl. vii. fig. 1, pl. vii. fig. 3

(1879). :

Superticially this species is not unlike a pale form of O. rutila,

but a glance at the skull shows it to belong to the ecurzone

group, and its large ears will prevent confusion with any other

members of that group.

This species is of moderate size, and the general colour above

is pale brownish grey, each hair being dark-coloured at its base,

white in the centre, and pale buff subterminally with a black tip.

Apparently it moults only once a year, in August, but in summer

the hair is much abraded and the animal is then much whiter.

The under parts and feet are white. Along the sides of the face,

across the shoulders, and from the nose over the occiput, the

general greyish colow: is tinged with rufous, this rufous being

more marked in summer; the eye is surrounded by an ill-defined

greyish ring. The inner sides of the ears and a patch behind are

white, the outer sides having dark brown hairs with white tips.

The skull belongs typically to the curzonie group, the sides of

the combined palatal and incisive foramina sloping outwards in

a regular slant from their anterior poimt. Another peculiarity

about the skull is the presence of two small oval foramina above

and in front of the orbit at the anterior end of the frontal bones.

These foramina, which measure about 3 mm. by 1°5 mm., are con-

stant and uniform in all specimens of this species that I have

examined, though they may also be found sporadically throughout

the genus. In other respects there is nothing of note to be

observed in the skulls.

Dimensions (from skin). Head and body 200 mm.; ear 27 ;

hind foot 32.

Skull. Palatal length 17 mm.; zygomatic breadth 23; length

of nasals 14; length of molar series 9.

Habitat. Doba, Kuenluen Mts.; Pamirs; Ladak.

The large eas, correlated with skull-characters, enable this

species to be easily recognised.

There are two species, O. aurita and O. grisea, described by

Blanford, which may probably be assigned to this species. I

have not had the opportunity of comparing any specimens, but

from the description and figures there can be no doubt that, if

not identical, they are very closely allied to O. macrotis.

OcHOTONA ROYLEI (Ogilby).

Lagomys royle: Ogilby, Royle’s Him. Bot. p. lxix, pl. iv. (1839)

id. Geoftr. Voy. Jacquemont, Mamm. p. 62 (1841); Waterh-

Mamm. ii. p. 26 (1848); Adams, P. Z.8. 1858, p. 520; Blyth

Cat. p. 183 (1863); Jerd. Mamm. p. 226 (1867); Blanf. J. A.S. B

xli. pt. 2, p. 35 (1872); Lydekker, J. A. 8. B. xlvi. p. 286 (1877) ;

Scully, Ann. Mag. Nat. Hist. ser. 5, vol. vill. p. 100 (1881)

218 MR. J. LEWIS BONHOTE ON THE [ Nov. 15,

Biichn. Mamm. Przew. i. p. 156, pl. xxiii. figs. 1, 2 (1890); W. L.

Sclater, Cat. Mamm. Calc. Mus. p. 112 (1891); Blanf. Faun. Br.

Ind., Mamm. p. 456 (1891).

Lagomys nepalensis Hodgson, J. A. 8. B. x. p, 854 (1841), fig.

p. 816; id. J. A.S.B. xi. p. 289 (1842); Waterh. Mamm. iu.

p. 24 (1848); Gray, Cat. Hodgs. Coll. p. 21 (1846); Horsfield,

Cat. H.-I. Mus. p. 148 (1851); Giinth. Ann. Mag. Nat. Hist. (4)

vol. xvi. p. 230 (1875).

General colour in winter dark brown grizzled with buff, each

hair being dark brown with a buff subterminal annulation. The

head shows traces of rufous. Under parts white. Feet rufous-

buff. The summer pelage is similar, but the head, shoulders, and

flanks are bright rufous (hazel, Ridgw.), the head being slightly

evizzled with black. Under parts sometimes showing a pale

median rufous streak.

The skull shows no very distinctive features. It is long and

narrow, and the nasal bones, especially at their anterior end,

broad.

Dimensions (from skin). Head and body 175 mm. ; hind foot

Jo eirear 2a:

Skull. Palatal length 15-5 mm.; zygomatic breadth 20; length

of nasals 12; length of molar series 1; interorbital breadth 6;

breadth of nasals 7.

Habitat. Kashmir; Nepal.

The external differences between this species and O. wardi

have been pointed out in the description of the latter species ;

and although somewhat alike, the skull-characters enable them to

be easily separated. The series of this species being very small,

T have been unable to separate roylec from nepalensis. I fancy,

however, that a large series would prove them to be subspecifically

distinct. The type of roylet came from the Choor Mts., a little

to the west of Kumaon, and the type of nepalensis from EK. Nepal,

north of Katmandu.

OcHoToNA HoDGsONI (Blyth).

Lagomys hodgsoni Blyth, J. A. 8. B. x. p. 817, pl. p. 844 (1841) ;

Waterh. Mamm. ii. p. 23 (1848); Gray, Ann. Mag. Nat. Hist.

ser. 3, xx. p. 220 (1867).

Lagomys tibetanus Milne-Edw. Nouv. Arch. Mus. vii. p. 93

(1871); id. Rech. Mamm. p. 314 (1872).

General colour above dull dark reddish brown, shading to pale

buffy brown on the flanks. The under parts are of a dirty white,

having a reddish median band. ‘The upper sides of the feet are

lighter than the rest of the body and slightly tinged with rufous.

The skulls are so fragmentary that Iam unable to give a detailed

description. The incisive foramina are much more triangular in

shape than in roylez, and their margins tend to slope out gradually

from the apex.

Dimensions (as given by Blyth). Head and body 6 ins. (150

mm.); hind foot 1:25 in. (832 mm.).

1904. | MOUSE-HARES OF THE GENUS OCHOTONA. 219

Skull. Palatal length 12mm.; zygomatic breadth 15 ; length of

molar series 6°5; length of nasals 10; interorbital breadth 4.

Habitat. Kashmir; Tibet; Szechuen.

This species is allied to the preceding, the dull pelage of which

it greatly resembles; its much smaller size will, however, enable

it to be easily recognised.

I have seen no specimens from Kashmir, the type-locality of

this species, but there is a series of specimens in the Museum

from K, Sikkim, which agree so closely with Blyth’s description,

that I have no hesitation in referring them to this species. I have

also examined specimens from Szechuen, which are indistinguish-

able from those of E. Sikkim, as well as a single individual from

Gannsu in N. Thibet. M. Milne-Edwards’ species from Moupin

is also, in my opinion, identical with O. hodgsoni, of which it

therefore becomes a synonym.

Since the foregoing has been in the possession of the Society,

Mr. Marcus W. Lyon, jun., has published * an exhaustive paper

on “The Classification of the Hares and their Allies,” founded

almost entirely, as would naturally be the case, on their osteo-

logical characters. He divides the genus Ochotona (l.c. p. 438)

into three subgenera founded on cranial characters, and it is

gratifying to find that his subgenera correspond with the three

groups into which I have found it necessary to subdivide the

genus, as set out in this paper. For the names of his subgenera

Mr. Lyon makes use of two previously existing, viz. Ochotona and

Pika, and coins a new one, Conothoa. The subgenera Ochotona,

Conothoa, and Pika represent respectively the rufescens, curzome,

and ladacensis groups of this paper.

With regard to the actual classification of some of the species,

there are several apparent discrepancies between Mr. lLyon’s

results and my own. In considering these, however, it must be

borne in mind that Mr. Lyons was treating the subject from a

larger point of view and also almost entirely from the osteo-

logical side; whereas in my work geographical distribution and

external characters were more especially studied, and I had the

additional advantage of superior series of skins, owing to many

of the actual types being in the Museum, as well as a nearly

complete set from Biichner of the various species he had described.

The first discrepancy is the grouping together of O. ladacensis

and 0. koslowi. 'The figure given by Biichner of the skull of the

former does not agree with the type skull, as I have already

pointed out (antea p. 209), which undoubtedly belongs to the

same group as O. alpina, and there can be but little doubt that

Biichner’s figure misled Mr. Lyon.

The next discrepancy is with regard to 0. erythrotis, which

Mr. Lyon, on the strength of Biichner’s figure, places in the

curzonie group. I have not been able to examine specimens of

O. erythrotis, but there are in the Museum a series of 0, rutila

* Smithsonian Miscell. Coll. vol. xlv. p. 321 (1904).

220 DR. W. B. BENHAM ON [ Nov. 15,

which, according to Biichner, very closely resemble O. erythrotis.

Biichner’s actual words are :—‘ Die Incisivoffnung ist durch einen

paarigen Vorsprung des Zwischenkiefers in zwei Abschnitte, eine

vordere und eine hintere Incisivoffnung getrennt; in Form und

Grosse gleichen diese Oeffnungen vollsténdig denjenigen bei

L. rutilus” ; and on the strength of this I have no hesitation in

placing O. erythrotis in the ladacensis (subgen. Pika) group, 2

finding with which Mr. Lyon, had he seen specimens of O. ruiila,.

would, I feel, sure agree.

The only other discrepancies relate to O. curzonie, O. daurica,

O. melanostoma, and O. pusilla, but as Mr. Lyon has never seen

specimens or figures of any of these, his conclusions with regard

to them must of necessity be of a rather speculative character.

To sum up shortly, we may fairly consider Mr. Lyon’s con-

clusions as regards the main divisions of the genus to he clearly

borne out by the foregoing pages, and, so far as the Palearctic

species are concerned, this paper may claim to have clearly shown

to which subgenus any particular species should be assigned.

5. On some Edible and other New Species of Harthworms

from the North Island of New Zealand. By W. B.

Benuam, D.Se., M.A., F.Z.S., Professor of Biology in

the University of Otago, New Zealand.

[Received May 31, 1904.]

(Text-figures 41—82.)

The Earthworms that have hitherto been described from New

Zealand by Mr. Beddard and by myself have been collected, with

one exception, from the South Island, and mdeed from the

southern half of that island. The majority of these belong to the

genera Maoridrilus, Notiodrilus, and Plagiocheta, belonging to

the subfamily Acanthodriline, and to the genus Octochetus, of

Michaelsen’s subfamily Octochetine ; and the general facies of our

South Island fauna is very characteristic and quite distinct from

the Australian Earthworms. But I have recently been able to

examine specimens of a number of species from various parts of the

North Island,with the astonishing result that they present astriking

contrast to those of the South Island, and as striking a resem-

blance to the Australian Cryptodrilids. Even in the South Island

we have in two species of Diporocheta, and the lacustrine species.

of Plutellus, representatives of the Australian fauna ; but whereas.

the species just referred to are by no means common, and might

probably be regarded as comparatively recent arrivals, possibly

even accidentally introduced from the adjoining continent, such an

explanation appears to be quite inadmissible for the northern

species; for the new genus Yokea, which I find it necessary to

make, is represented by seven species in quite distant parts of the

North Island; and the Acanthodriline genera are just as scarce

1904. ] EARTHWORMS FROM NEW ZEALAND. at

in the North Island as the Cryptodriline forms are in the South ;

moreover, most of these worms are found in inland places, or in

spots more or less remote from European cultivation ; and, indeed,

some of the species were formerly used as food by the Maoris,

who recognise several different species of Earthworms and give

distinctive names thereto.

Before attempting to explain this Australian faunal resem-

blance of the North Island, we must wait till we can obtain more

material, both from the southern portion of that island and the

northern districts of the South Island.

Another interesting group of Worms is also represented in

the collection, viz., two new species of Rhododrilus—a genus

peculiar to New Zealand, but allied to Wicroscolex, which is an

American form.

Finally, I find it necessary to create a new genus, Dinodri-

loides, for a worm which bears the same relation to Dinodrilus

that Neodrilus bears to Maoridrilus, in that the hinder pair of

prostates has disappeared ; though this is not the only point of

difference from Beddard’s genus.

For the majority of the species described below A am indebted to

Myr. Elsdon Best, of Ruatahuna, who, at my request, took the

trouble to collect, preserve, and despatch to me, in February 1904,

several kinds of worm that were formerly used as food by the

Maories, of which he has written an account in the ‘ Transactions

of the New Zealand Institute’ for 1901, on p. 64, in his article

on “The Food-products of Tuhoe-land.” Tuhoe-land or the

Uvewera Country lies on the eastern part of the North Island,

south of the Bay of Plenty, and not far from Rotorua, so famous

for its baths. The country is much rougher and less affected by

civilisation than elsewhere, and is still peopled by Maories in a

less Kuropeanised condition than in other parts.

In this article Mr. Best enumerates and givesa brief description

of eight different kinds of earthworm as being eaten by the

natives: these are ‘ Kuharu, Noru, Wharu, Tarao, Pokotea, Tai,

Kurekure, and Whiti.” The “two last are famed for their

sweetness and flavour,” and ‘ were reserved as food for the chiefs.

The sweet flavour is said to remain in the mouth for two days,”

though Mr. Best states that he “ cannot speak from experience ”

as to this fact.

In preparing these worms for food, “ those which contain earth

are stripped with the fingers before being prepared for eating,

this forces the earth out of them.” I suppose this means

“* stripping” in the way fish are “stripped” of their milt and ova,

in hatcheries.

“To cook these worms some water is placed in a bowl and

rendered warm (not hot) by means of hot stones. The worms

are then cast into the water and allowed to remain there for some

hours. Before long the worms will have become dissolved or

partially so, but were the water too hot they would not melt.

Some cooked greens are added to the mess, and a prized dish is

yp DR. W. B. BENHAM ON | Nov. 15,

ready; the gods who live for ever would smile at the sight

of it.” ;

‘Worms were preserved in gourds for some time. The best

kinds were favourite 0 matengo of former days; the last food

taken by a dying person is so termed.”

Of the earthworms referred to in this extract, Mr. Best sent

me specimens of “Tarao,” ‘“ Pokotea,” and “‘ Kurekure,” as well

as another earthworm, called by the natives “ Tokerangi,” but not

included in his list of foods.

“Tarao” and ‘“ Tokerangi” belong to the genus Rhododrilus ;

“ Kurekure” and “ Pokotea” to the new genus Zokea. But

‘“ Kurekure ” includes two species, so that we have four (or five if

“'Tokerangi” is eaten) species of edible earthworms, belonging to

two distinct genera. I believe this is the first time that the use

of Oligocheta as an article of diet has been recorded ; for no

mention is made of earthworms in the series of Animals used as

Food enumerated by Professor Lankester in his introduction to

the recently published volume of “ Reports on Economic Zoology,”

issued from the British Museum. mM

The following is a list of the twelve new species of Karthworms

described in the present paper: T desire to express my thanks

to my various friends and correspondents who have so kindly

collected these. and other, worms in out of the way districts. ‘

Fam. MEGASCOLECIDS.

Subfam, ACANTHODRILINA,

Maoridrilus mawiensis.

Octochetus michaelsent.

Dinodriloides beddardt.

Rhododrilus edulis.

Rhododrilus besti.

Subfam. MEGASCOLECIN &.

Tokea, gen. 0.

T. esculenta.

7’. sapida.

T. urewere.

T. hutioni.

T. suteri.

T. kirki.

T. maorica.

MAORIDRILUS MAUIENSIS, sp. n. (Text-figs. 41-44.)

A single incomplete specimen, collected in 1899 by Mr. Suter

and now preserved in alcohol. It is soft and ill-preserved.

Locality. Auckland.

Colour. Pale yellowish, in marked contrast to the usual dark

tint of members of the genus; possibly the pigment has been

dissolved.

1904. | EARTHWORMS FROM NEW ZEALAND.

Dimensions. 80 x 4 mm., for 117 segments,

Prostonnum tauglohic.

Cheta. 8, closely coupled; aa< be=dd.

Chitellum undeveloped.

Henital pores.—The porophores* in 17 and 19 are in line of

chzetee ab, which are absent. - The spermatic groove passes straight

backwards between a and 6, which are present on 18 in the

manner characteristic of the genus; spermathecal pores normal

in number and position, in line ab.

Internal Anatomy.

‘The septa behind segments 8 to 14 slightly thickened.

Text-fig. 42.

Text-fig. 41.

Text-fig. 41. Maoridrilus mauwiensis—A spermatheca (X 12: camera outline): no

structural difference could be detected between the two lobes of the sac, to

enable a distinction of diverticulum and ampulla to be made.

Text-fic. 42. Maoridrilus mauiensis.—A penial cheta (X 30. Laty oc. 1, obj. 2,

camera).

* I have suggested, in an article on MW. uliginosus (Tr. N.Z. I. 1900, p. 125), this

name for papillze carrying the male or prostate pores.

224 DR. W. B. BENHAM ON [ Nov. 15,

Dorsal vessel double ; last heart in 15th segment.

Gizzard large; cesophagus with 3 pairs of glands, quite distinct

and typically developed, situated in segments 13, 14, 15, the last

more dorsally placed and smaller than the preceding.

Sperm-sacs of fair size, in segments 11, 12; botryoidal.

Prostates* normal. Penial chetz delicate, much curved in

an exaggerated S-shape, with the tip spoon-shaped, though‘pointed

and curved in side view.

The spermathece ave of peculiar form (text-fig. 41, p. 225).

Each consists of a bilobed sac; the two lobes are of about the same

size and irregularly ovoid, and joined by a short, narrow isthmus,

whence the muscular duct originates. One might imagine that

one lobe is a “ diverticulum” in the usual sense, but examination

of stained specimens shows no structural difference between them ;

Text-fig. 43.

arr Sis ee

Maoridrilus mauiensis—Tip of penial cheta, side view (X 350. Oc. 1, obj. 7,

camera).

Maoridrilus mauiensis—Tip of penial chieta; view of plane at right angles to the

above (? perhaps the extreme tip is mjured).

the epithelium is folded, irregular, and apparently glandular, as a

quantity of stained material is present in the lumen; I could see

no spermatozoa. As a rule, there is a marked difference in

structure between sac and diverticulum. Unfortunately the

specimen is not sufficiently well preserved to enable me to decide

this question.

At any rate the form of the spermatheca and the arrangement

of the csophageal glands mark the species from any of those

hitherto described.

This is the first species of Maortdrilus described from the

North Island, and is the only specimen amongst the material I

have received from various correspondents collected in several

widely scattered districts.

* No doubt Beddard’s term “spermiducal gland” is in some respects better, but

it is a clumsy term, and when “ duct” is added it is by no means euphonious.

1904. ] EARTHWORMS FROM NEW ZEALAND. 225

OCTOCHETUS MICHAELSEN1, sp.n. (Text-fig. 45.)

A single individual of this worm was collected by Capt. F. W.

Hutton at Wellington. It was broken into four or five pieces

when I received it; in general appearance it agrees with other

species of the genus,

Dimensions. About 210 x 8 mm.; the number of segments was

not counted, as the worm was too greatly contracted and broken to

make the attempt profitable.

The clitellum, though not fully developed, appears to cover

segments 15 to 19.

The porophores are in line of }, as also are the spermathecal

pores, and their position on the body is lateral rather than

ventral, though of course on the under side.

The chat have the usual spaced arrangement, and, in spite of

examination of the skin, I was unable to detect them in front of

the tenth segment.

The arrangement is as follows: d is a little above the lateral

line, so that d-d is about 3 of the circumference, aa=cd=14 ab;

ab=be. The gap aa is wider in the clitellar and preclitellar

segments than posteriorly,

Internal Anatomy,

There are seven very stout septa, the last being behind the

12th segment.

The dorsal vessel is double as far forward as segment 8, and

the last heart is in the 15th segment,

Text-fig. 45,

Octochetus michaelseni.—Spermatheca (X 12): the diverticulum is represented by

several small saccules (D) embedded in the thiekness of the muscular duct.

The long gizzard is in the 6th segment, with the thin septum

five-sixths attached near its anterior margin.

The cesophagus bears a single pair of glands in the 15th seg-

ment and is well marked and hemispherical, though a good deal

compressed antero-posteriorly owing to the contractions of the

body.

The intestine commences in the 16th segment.

Proc. Zoot. Soc.—1904, Vou. II. No. XV. 15

226 DR. W. B. BENHAM ON [ Nov. 15,

With regard to the reproductive system, the testes and ovaries

are on the posterior wall of their segments, as in O. thomasi and

other species. The penial chetze are small and do not appear

internally, but on each of the prostate pores, when examined

under a lens, the broken bases of two dark chzetz are visible.

The spermathece are unsymmetrically developed ; on the right

side the normal two pairs are present in segments 8 and 9, but on

the left side the anterior sac is absent. Each spermatheca is a

simple ovoid sac (text-fig. 45, p. 225), with a short, thick duct

which forms an acute angle with the sac (probably owing to the

contraction of the body), There is apparently no diverticulum,

but in the stained and clarified organ there are seen to be a number

of small saccules, irregularly arranged and embedded in the wall

of the duct ; they do not form any projection, but are situated m

the widest part of the duct, where it is bending to reach the body-

wall ; and though I have not yet sectionised the organ, it appears

as if their presence here caused the duct to be of greater diameter

at this spot.

Remarks.—The form of the spermatheca and the arrangement

of the esophageal glands suffice to distinguish this species from

any hitherto described.

DINODRILOIDES, gen. nov.

Chetze 12 per segment. Clitellum girdle-like, on segments

14-16 (= 3 segments). Prostate pores, one pair, on the 17th,

and male pore on 18th segment. A single spermatheca, opening

at 8/9. Meganephric: pores in line not alternating. Gizzard in

segment 6. 2 pairs of testes, segments 10,11. 2 pairs sperm-

sacs, in segments 11,12. Prostates: a single pair much coiled,

cylindrical.

DINODRILOIDES BEDDARDI, sp. n. (‘Text-figs. 46, 47.)

A single specimen of this interesting worm was collected by

My. H. Suter at Auckland.

The colour (in the specimen preserved in alcohol) is a very

distinctive bluish grey, equally dark in tint along the entire

length, and the pigment extends down the sides to the ventral

surface, which, however, is paler and becomes yellowish in the

hinder region. ‘The clitellum is yellow-brown.

The worm consists of 95 segments, and measures 78 mm. X

4 mm. =f iin

The chete are arranged as in Dinodrilus *—that is to say, there

are 12 in each segment; in 3 couples on each side, the individuals

of which are widely separated, so that the 6 are practically equi-

distant, while the dorsal and ventral gaps are only slightly greater

than the other gaps. The middle couple (c-d) are lateral in

position.

% Prof. Spencer has recently described an Australian Cryptodriline genus, Tricheta,

with six couples of chet (vide Proc. Roy. Soc. 1900, p. 30).

1904. ] EARTHWORMS FROM NEW ZEALAND. 227

Each cheta is carried on a slight papilla (in the preserved

specimen), which is very pale and, indeed, nearly white.

The prostomiwm is only slightly, if at all, embedded, but as the

buccal region is everted, producing wrinkles on the peristomium,

it is impossible to detect the actual limits of the prostomium.

The elitellam is fully developed and very well-defined, both in

front and behind ; it completely surrounds segments 14, 15, 16.

Geméal pores, &e. (text-fig, 46)—The most remarkable thing

about the worm, in which it contrasts with Dinodrilus, is the

presence of only one pair of “ porophores” bearing the pores of

the spermidueal glands, which are situated on the 17th segment

Text-fig. 46.

Dinodriloides beddardi—View ot part of the fore-body, slit open along the dorsal

mid-line, flattened out, and seen from below. (xX about 3.)

The six chet are seen, lettered A-F [a—fin the text] on the right side, at

their true relative distances. ‘The genital pores, nephritic pores, and tubercula

pubertatis (‘T. P.) are also shown.

in line of 6. The papilla is oval, and extends outwards nearly to

the level of ¢; it is traversed by a spermatic groove, which passes

backwards on the next segment, on the anterior margin of which

is a depressed, semicircular prominence, in the same line as the

porophore. ‘The spermatic groove ceases some distance in front

of the cheta 6, so that the male pore is close to the anterior

margin of segment 18.

The ventral surface of segments 16, 17, 18 is pale yellow, and

thus contrasts with the grey tint of the neighbouring segments.

On the 18th segment there seems to be an oval glandular area

15*

228 DR. W. B. BENHAM ON [ Nov. 15,

extending from 6-), and in the prechetal portion of the 16th

segment a pair of similar glands, which perhaps represent tubercula

pubertatis, but they are not well-defined.

The two oviducal pores are visible in the usual segment, in front

of the cheetal space a/b.

The species possesses a single pair of spermathece which open

between segments 8/9; each pore has a very prominent pale

yellowish lip in front and behind, in line dc: further, on each of

the segments 9 and 10 is a round, pitted tubercle in front of

cheeta 6 on each side.

The nephridiopores ave in line d.

Dorsal pores are very evident, and commence behind segment 6,

continuing to the last segments of the body.

Internal Anatomy.

There are no noticeably stout septa.

The dorsal vessel is double right up to the pharynx (as it is in

Dinodrilus) ; the last of the four pairs of hearts is in segment 13,

and of considerable size.

The gizzard is very feebly developed in segment 6; the

cesophagus is dilated in segments 14, 15, 16, but no definite

gland is formed,

The intestine commences in segment 18,

The worm contrasts with Dinodrilus in being meganephric ; and

in the mid-body, at any rate, the large muscular duct is easily

traced, under a dissecting-lens, to the body-wall, which it pene-

trates in line of cheta d to open to the exterior.

The Reproductive system.—There are two pairs of testes and

Text-fig. 47.

Dinodriloides beddardi.—Spermatheca, enlarged.

funnels in the usual segments. The two pairs of sperm-sacs are

in segments 11, 12; of small size, and botryoidal (or racemose) in

form. There is but a single pair of spermiducal glands, which

are thoroughly Acanthodriline in form, each being cylindrical and

compactly convoluted to form a mass of considerable size, provided

with a short narrow muscular duct.

1904. ] EARTHWORMS FROM NEW ZEALAND. 229

There are no penial cheete.

The ovaries are large and occupy the usual segment.

There is a single pair of spermathece in segment 9; each isa

subglobular sac (text-fig. 47) with a muscular duct, narrower than

the sac, which does not graduate into it but suddenly diminishes

indiameter. Two long tubular diverticula open, one on each side,

into the duct close to the body-wall. Each diverticulum, when

extended, is about twice the length of the sac.

Remarks.—This genus, as I have remarked above, bears the

same sort of relation in regard to the prostates to Dinodrilus as

Neodrilus does to Maoridrilus; but in Neodrilus, which Michael-

sen has termed a “ microscolecine form,” the number of testes has

also been reduced. Here, however, the reduction does not occur.

Moreover, Dinodrilus is micronephric instead of meganephric.

We have in New Zealand a series of genera that illustrate the

evolution of Harthworms in a very remarkable manner.

Starting with Wotiodrilus, which on general grounds is con-

sidered by Michaelsen as an ancient genus (and herein J agree

with him), we have a meganephric worm with 8 coupled cheetz,

2 pairs of prostates opening on segments 17 and 19, while the

sperm-ducts open independently on the intervening segment.

From this genus several lines of evolution start :—

(a) Maoridrilus—which differs only in having alternately

arranged nephridia, opening, that is to say, alternately in

relation to the dorsal and ventral couples of cheetze.

(b) Neodrilus—in which the second pair of prostates, second

pair of tubes, and spermathece have disappeared, but male

pore still in the middle of segment 18.

to 12 and widely separated, and in which the male pore 1s

close to the anterior margin of the 18th segment.

(d) Rhododrilus—which still retains 8 clusters, but in which the

male pore has moved forwards to the 17th segment and

opens close to the prostate pore.

Starting again from Votiodrilus, we have Plagiocheta, which

differs from it chiefly in having a considerable number of cheetee ;

and we readily see how this condition may have come about

through a Dinodrilus-like form with 12 chete. But in this

genus Plagiocheta, some, like P. sylvestris, have meganephridia,

others, like P. rossi, have micronephridia *.

Once more reverting to our archaic genus, and imagining the

development of micronephridia, we reach Octochetus, in which the

* See Benham (’02, a). The statement on pp. 287, 289, however, that P. ricardi

and P. montana are similariy micronephric is erroneous. The nephridium is very

small in proportion to the size of the worms, and the tubuli of the meganephridinm

are in tufts, which, under an ordinary dissecting-lens, suggests a series of isolated

micronephridia ; but in P. rossé (laps. cal. rossiz) the meganephridium has broken

up into micronephridia. JI am preparing an article on the nephridia of this and other

genera of New Zealand Earthworms, and for the present refrain from further detail.

230 DR. W. B. BENHAM ON [ Nov. 15,

number of chete remain normal, though they are differently

arranged.

Then there is Dinodrilus, with 12 chetz and micronephridia.

But both these genera have the prostate pores and male pores

arranged as in the archaic form.

It seems to me that Michaelsen is in error in separating these

two genera from other Acanthodriline forms and associating

them in a separate subfamily, the Octochetine, with Hutypheus

and Hoplochetella; for, apart from the micronephric condition,

there is really little to distinguish Octochetus from Notiodrilus ;

moreover their presence in New Zealand indicates their close

association therewith. The step from the Acanthodriline series

to the Megascolecine is a small one, and appears. easily conceivable

from analogy with the origin of Rhododrilus: that is, one prostate

pore has shifted so as to open close to and in common with the

male duct in the 18th segment, for several species of Cryptodriline

worms have prostates similar to those of Rhododrilus.

RHODODRILUS EDULIS, sp. n. (Text-figs. 48-54.)

Two individuals of “'Tarao,” a worm eaten by the Maoris, were

received from Mr. Best preserved in formol.

Dimensions. Length 275 and 285 mm. respectively, with a

diameter of 13 mm. just behind the clitellum. The worm is

cylindrical, tapering only very slightly posteriorly, and then

suddenly decreasing as the anus is approached, so that at segment

m—-8 the diameter is 10 mm., and at m—2 it is still8 mm. The

total number of segments in the larger individual is about 300,

but as the hindmost segments are very closely contracted and

small, and triannulate, it is not easy to count them with absolute

correctness—nor does this matter. There is no doubt that the

living worm was, when extended, at least 15 inches and possibly

more. Mr. Best in a letter says: ‘‘ These Tarao are small, I have

seen them 18 inches in length.”

The segments 2-5 are biannulate,.6—-12 are quadriannulate,

with the chetz in the third annulus, while those of the clitellar

and postclitellar regions are triannulate.

The colour of the “Tarao” is (in formol, which does not cause

the colour to change much) red; the anterior end appears pale

greyish, much paler than the rest of the body, which is pale

reddish, with a tinge of purple on the upper half of the body;

the clitellum is buff.

The prostomium, like the first segment, is a good deal furrowed ;

it is epilobic, being dovetailed into the peristomium for about

half the length of the latter, and it ends in a transverse groove.

The chete are eight in number, rather widely spaced; the

lateral spaces (4c) on each side are nearly equal in the mid-body

and posterior region, while the dorsal space (dd) is about twice

the ventral (aa) and three times the lateral.

As is frequently the case in allied genera, the ventral couple

(a, 6) approach one another in the region of the genital pores;

1904. } EARTHWORMS FROM NEW ZEALAND. 231

so that, on segment 21, ab is only 4 at in the mid-body segments,

and in the subclitellar segments (e.g. 16th) it is even less. At

the same time the ventral gap (aa) enlarges, so that b keeps in its

line, while @ moves outwards. In the preclitellar region this

increased gap remains, and the lateral and dorsal gaps are also

larger, but the gaps ab and cd remain the same size as in the mid-

body ; in other words, the rather increased diameter of the pre-

clitellar region affects the interchetal zone, and not the chetal

spaces themselves.

I measured the spaces on the body by means of dividers, with

the following results :—

Segment Seg. Seg. See.

Mid-body. ‘Tail. Xx1. XVill. XVl. 1X.

AG 5 A 5°25 6 5°25 5°D

ab. 3 2°5 1855 0:75 1 2°5

bb . 4 a0) 6°5 5

cd. 2°5 -2°5 2°5 2°5

Chik ll 9 12

Put in the usual formula for the mid-body, starting with

the smallest gap,

cd <ab<be< aa: aa=2cd : dd=2 aa=3 be.

On segment 17 the ventral cheete are replaced by two long,

delicate penial cheetze.

The clitellwm is saddle-shaped, and though the annuli and inter-

segmental furrows are quite evident ventrally, there is no distinct

latero-ventral margin such as exists in Lumbricus &e. The

glandular tissue extends down to the line of cheta 6. The cli-

tellum extends over the six segments 13-18.

Genital pores, dc.—The male pore is on segment 17, situated

on a low, rounded, and slightly prominent porophore in line with

the chetal gap ab. The pore itself is nearly in line with 6, and

the penial cheetze project therefrom.

Tubercula pubertatis are well-developed, in the form of a series

of transversely disposed, paired oval pads (text-fig. 48, p. 232)

situated on the hinder regions of the segments 12,13, 14, 15, and 19,

20. These pads are developed from the last annulus of the segment

in each case, but the last two pairs appear to be intersegmental,

owing partly to the contraction of the worm, and partly, perhaps,

to their size. These tubercles have not all the same position

relative to the cheetz : those of the 12th, 19th, and 20th seements

extend from the level of 6inwards nearly to the middle line, while

the other three pairs, on segments 13, 14, 15, extend from cheta a

and almost touch in the middle line.

There isa single pair of spermathecal pores on the intersegmental

groove 7/8, in line with 6. As a matter of fact, each pore is

double: 7. ¢., the main sac and its diverticulum open independently

one above the other, but close together, in the furrow.

I was unable to detect either dorsal pores or nephridiopores by

232 DR. W. B. BENHAM ON | Nov. 15,

examination of the surface, owing to the strongly contracted state

of the worm: the nephridiopores, however, are in line ec.

Text-fig. 48.

S—oJ———

——

MW RWS

lw wef

AIAN

18\;

19 TP.

Rhododrilus edulis——Ventral view (X about 8) of clitellar region, showing male

pores (g) and arrangement of tubercula pubertatis (T.P.); AB [aé in the

text], the chetal rows. Segment 16 is unfortunately drawn a little too large.

Internal Anatonvy.

The body-wall is of great thickness and the septa behind

segments 7 to 12 are very stout.

The dorsal vessel is single; the last heart in segment 13, and

rather smaller ones in 10, 11, and 12.

The gizzard, in segment 5, is large, with thick walls, and

contains stones. There is no esophageal gland, nor any dilata-

tion. The intestine commences in the 17th segment, where the

gut suddenly enlarges to twice its previous diameter.

The worm is meganephric, and the nephridia commence as far

forwards as segment 3. Hach nephridium (text-fig. 49) consists

of a bunch of 3-5 loops ventrally (situated in line a—b), whence a

long dorsal loop passes upwards to a point about midway between

ed, and a straight duct leaves the same bunch, and passes to the

body-wall just below c, at which point no doubt it opens

externally.

There is, in addition, a blind cecum or bladder with muscular

wall, which extends almost to the mid-dorsal line: the exact

connections of it I have not determined.

The nephridiostome is small.

1904. | EARTHWORMS FROM NEW ZEALAND. 233

Text-fig. 50.

Text-fig. 49.

we / wr

Text-fig. 52.

Mie

Text-fig. 49. Rhododrilus edulis. —A nephridium, shown in its natural relation to the

wall of the body, as seen when spread out in the normal way. A, B,C, D [a, 4,

c, d in the text |, the position of the four cheete of this side; F, nephrostome ;

V.L., ventral loops or coils of tubules ; D.L., long dorsal loop extending nearly

to cheta D; C.M.,muscular cecum; N.C., nerve-cord ; M.L., dorsal mid-line ;

N.D., nephridial duct.

_ Text-fig. 50. Rhododrilus edulis——The prostate of the left side (enlarged). It

occupies five segments and is provided with a muscular duct, which thickens as

it penetrates the body-wall just below and behind the arcuate muscles (AR.).

The penial cheetal sac has been removed.

Text-fig. 51. Rhododrilus edulis.——A penial chata (80. Zeiss oc. 3, obj. 3, camera.

Text-fig. 52. Rhododrilus edulis.—The tip of a penial cheta (480).

234 DR. W. B. BENHAM ON [Nov. 15,

Reproductive system.—There are two pairs of large, botryoidal

sperm-sacs in segments 11 and 12, attached, of course, to the

anterior wall of these segments.

The prostate (text-fig. 50, p. 233) is tongue-shaped, long, more

or less convoluted, with apex recurved; it extends through

segments 17 to 21; its muscular duct is confined to segment 17 ;

it is rather long, but very narrow where it leaves the gland,

dilating to form a thicker, pear-shaped bulb as it penetrates the

body-wall.

Arcuate muscles are developed in segment 17. :

Associated with each prostate are two sacs containing each two

penial cheete, a long functional and a shorter reserve bristle.

Each penial cheeta (text-figs. 51, 52, p. 233) is stout, nearly

straight, and, compared with the size of the worm, not very long :

it terminates in a rounded knob, which carries at its end a short,

stout, curved, blunt hook. There are no markings at this end;

but about midway along its length there is a series of rather

closely-set, irregularly arranged, short, oblique, and finely serrated

ridges.

The relations of the various parts of the male ducts, as studied

Text-fig. 53.

eas)

Text-fig. 58. Rhododrilus edulis—Diagrammatic sketch of a section through the

porophore, showing the openings of the sperm-duct (S.P.), prostate duct (P.R.),

and the penial cheetze (P.CH.) into a small antrum or chamber, which communi-

cates with the exterior at the apex of the papilla (¢). The sketch is compiled

from a series of sections, and the above pores are not, in reality, in one plane—

the prostate pore being most anterior, and the sperm-pore most posterior of the

three. V. is the ventral side. IL. is laterally placed.

Text-fig. 54. Rhododrilus edulis —Spermatheca (x4). The main sac, or ampulla,

opens externally, independently of the diverticulum (D. W.).

in sections, is as follows (text-fig. 53) :—at the apex of the poro-

phore is a small pore, which leads into a shallow, but laterally

extended chamber, the floor of which rises up as a papilla. At

1904. ] HARTHWORMS FROM NEW ZEALAND. 235

the apex of this “ penial papilla,” as it may be termed, are two

independent apertures, belonging to the two sacs of penial chetee.

At the base of this penial papilla, and on the outer side, there

opens the muscular duct of the prostate, while the vasa deferentia

(which have united) open into a posteriorly placed recess of the

above-named chamber. This chamber is very shallow, and we

practically have fowr pores on the porophore; and probably in a

less contracted condition these four pores would open into a

shallow pit, the margin of which has, in our specimen, closed

over the pit so as to leave only a single small aperture.

The spermatheca (text-fig. 54) lies in segment 8, on each side ;

it is a large and long sac, broadest in the middle, without a

definite duct, but narrowing as it approaches the body-wall. The

diverticulum is narrow and tubular, more than half the length of

the sac and about one-fourth of its diameter; it opens indepen-

dently of the sac itself.

As a matter of detail, the length of the main sac is 8 mm., its

breadth is 2°5 mm.

Loc. Ruatahuua, Urewera Country, North Island of New

Zealand.

RHODODRILUS BESTI, Sp. n. (Text-figs. 55-61.)

A very pale, and probably white, worm, with yellow clitellum ;

its general appearance is that of an Octochetus. A much smaller

worm than the preceding, and known to the Maoris as

“ Tokerangi.”

Dimensions. Length 125 mm., diameter 5 mm. just behind

the cltellum ; with 206 segments, which, with the exception of the

first four, are triannulate.

The chet are spaced, and as measured on the body, ab=cd;

be=aa=1?, ab; dd=25 be=about 4 ab. That is, the interchetal

spaces on each side are equal and less than the lateral space ;

the ventral space is less than half the dorsal space. Relatively

aa is greater than in the preceding species.

Clitellum saddle-shaped ; on segments 4 13—18 (=53 segments).

The glandular tissue extends downwards as far as line of

cheeta a.

Genital pores, &c.—A pair of porophores in segment 17, in line

of @: on each, two pores are readily visible under a lens, and the

penial chetz project from the outer one. There are four pairs of

postgenital tubercula pubertatis, on the posterior annulus of

segments 19, 20, 21, 22, in line with the gap ad, the ventral

margin extending below a; so that they are separated by a

much greater space than in #. edulis. Each tuberculum puber-

tatis is oval, and slightly depressed in its central part, so as

to appear sucker-like (text-fig. 56, p. 236).

In addition to these four pairs, there is on the hinder margin

of segment 11] a pair of rounder swellings—not pitted apparently —

in line with the porophores, and not extending so far outwards

236 DR. W. B. BENHAM ON [ Nov. 15,

as the postgenital tubercle; further, the pair are connected

across the middle line by a transverse ridge.

A single pair of spermathecal pores at the groove 8/9,

Text-fig. 56.

Text-fig. 55. Rhododrilus besti.—Ventral view of the clitellar segments, &c.

(X about 5), showing porophores and arrangement of the tubercula pubertatis

(T.P.).

Text-fig. 56. Rhododrilus besti.—An enlarged view of one of the tubercula puber-

tatis, showing the glandular depression centrally, and also the fact that it is on

the hinder annulus of its segment, though apparently intersegmental.

Text-fig. 57. Rhododrilus besti—Portion of prostate, showing shortness of duct, as

compared with that of R. edulis.

Internal Anatomy.

There are stout septa behind segments 7 to 12. The gizzard is

large and occupies segments 5 and 6. The spermiducal glands

are long, undulating, tongue-shaped, extending into the 24th

segment. The prostrate duct is short, narrow, and only slightly

curved (text-fig. 57).

The sacs of penial cheetee extend from segments 17 to 21. Each

penial cheta (text-figs. 58-60) is much slenderer than in the

preceding species, and larger and of a different form ; it is gently

curved and the blunt point is bent in the opposite direction ;

here it is flattened and slightly excavated, so as to be spoon-

1904. ] EARTHWORMS FROM NEW ZEALAND. 237

shaped when seen from above; in side view, however, the point

is narrower,

Text-fig. 58.

Text-fig. 59. Text-fig. 60.

Text-fig. 58. Rhododrilus besti.—A penial cheta (80).

Text-fig. 59. Rhododrilus besti.—Tip of penial cheta (X480). View of one plane,

showing spoon-shaped excavation.

Text-fig. 60. Rhododrilus besti.—Tip of penial cheta, side view ( X 480), also showing

spoon-shaped excavation.

Text-fig. 61.

Rhododrilus besti.-A spermatheca (X12) (camera outline), showing remarkable

coiled diverticulum (D).

The single pair of spermathece (text-fig, 61) jie in segment 9;

238 DR. W, B. BENHAM ON [ Nov. 15,

each sac is more or less ovoid, somewhat bent at the commence-

ment of the duct (? perhaps artificially by contraction of the

body).

ae duct is short, but wide, and receives a very long, tubular

diverticulum, which is spirally coiled at its lower end, where it

enters the common duct. close to the body-wall. The lower end

of the diverticulum is muscular, though of the same diameter as

the rest,

Loc. Ruatahuua, Urewera Country, North Island, N. Z. <A

single individual,

Remarks.—It will be seen that this species differs from 2. edulis

in details with regard to the arrangement of chet and copulatory

tubercles, in form of penial cheetze and spermatheca, as well as in

size, form, and colour of the worm itself.

The Genus Rhododrilus,

It is clear that in many features FR, edulis and R. besti agree with

Microscolex; nevertheless there are certain points of difference,

which, in view of geographical distribution, may be of more

importance than the resemblances.

For example, most noticeable, but perhaps of no great sys-

tematic importance, is the great size of these two species, and

especially of &. edulis (285 mm.), as compared with the small

worms included in the genus J/icroscolex, ranging as they do

frgm a length of 30 mm, to that of 58 mm.

ut the point upon which I would lay stress is as to the form

and extent of the prostate.

In all the species of MWieroscolex this gland is of comparatively

small size, and limited’ to segment 17, or extends, in

M. hempeli and M. nove-zealandic, into the next segment;

whereas in the two species just described these glands are of

very considerable extent, passing through 5 or even 8 segments.

It is also worth noting, in view of the fact that Beddard is

inclined to lay some importance on the point in some genera,

that in all the species of Jicroscolex the last heart is in segment

12, whereas in my two species it is In segment 13.

Michaelsen points out, too, that in the genus MWicroscolex the

single pair of spermathecze open in the furrow 8/9: this is the

case with one of my species, whereas the pore in the other is

at 7/8.

The gizzard, too, in Microscolex, is “absent or rudimentary,”

which is not true of our edible worms ; and, finally, the clitellum

of the latter is not ‘ complete” or “ girdle-like,” but saddle-

shaped.

These small points of difference necessitate one of three courses :

either we must amend the diagnosis of MWicroscolex, or we must

create a new genus, or we may place our species in the closely

allied genus Rhododrilus. 'The last plan, it will be seen, is the

one I have followed, though even this procedure necessitates the

1904. | EARTHWORMS FROM NEW ZEALAND. 239

alteration in the definition of this genus. But this, it seems to

me, 1s a better plan than to modify the diagnosis of JMicroscolea,

for the latter is an American genus, though one species * at least

has been recorded from New Zealand; whereas the only species

of Khododrilus hitherto described are from this region.

The genus, originally founded by Beddard (89) for R. minutus,

has suffered from its resemblance to J/ecroscolex. Originally

distinguished from it by its author, as having the prostate pore

separate from the male pore, it was, later, included in that genus,

on the discovery that the same arrangement is true of certain

species of the older genus. Michaelsen (00), however, retains it

as distinct, owing to the fact that Rh. minutus has four pairs of

spermathecee.

In 1900 I deseribed a worm from the Chatham Islands as

“* Wicroscolex huttoni,” with two pairs of spermathecze, following

Beddard in disallowing Phododrilus. But since the publication

of that article I have studied a species from Campbell Island

and the Lord Auckland Group, in which there are three pairs of

spermathecee.

Now all these worms agree with the two species described in

the present article in having an elongated prostate, extending

through several segments, and in most of them the gizzard is well

developed; the clitellum saddle-shaped. But the position of the

last heart 1s not constant.

The size of the worms, too, exceeds that of the Species of

Microscolex.

I therefore agree with Michaelsen to resuscitate Beddard’s

genus Lhododrilus to include our New Zealand worms, but

regard the form and extent of the prostate as the chief character,

and one that readily distinguishes it from JWteroscolex.

The characters of the genus as thus modified may be defined as

follows :—

Cheetze 8 per segment, more or less widely spaced. N ephridio-

pores not alternating ; male pores on 17 ; prostate pores one pair,

in 17, opening close to the male pore. Clitellum saddle-shaped,

occupying 4 to 6 segments, 13 (14)—(17) 18. A gizzard in 5.

Testes two pairs. Prostates tongue-shaped, elongated, more or less

undulatory, extending through 4 to 8 segments. Penial cheetz

present.

Distribution. New Zealand and neighbouring islands,

1. Rh. minutus, Beddard, 1889. South Island.

2. Rh. huttoni, Benham, 1900. Chatham Island.

3. Kh. edulis, Benham, sp. n. North island.

4. Rh. besti, Benham, sp. n. North Island.

I have specimens of other species in my possession of which a

description has not yet been published.

* Of a second species, WM. menticola, our information is not altogether sutiicient

to permit us to make use of its characters in this discussion. —

240 DR. W, B. BENHAM ON [N ov. 15

TokEA *, gen. nov.

Characters of the genus.—Cheete 8, spaced, and more or less

equidistant ; clitellum girdle-like (13) 14—-17 (18), 7. e. covers

A or 5 segments.

Male pore common with that of prostate,in 18th segment.

Two pairs of testes in usual segments; two pairs of sperm-sacs in

segments 9, 12; the prostates are long, tongue-shaped, lie below

the gut, close to one another, and extend through several

segments. No penial chete.

The gizzard in 5; no calcareous glands; last heart in 12 or 13.

Micronephric, with meganephore in last few segments, Sperma-

thecze two or three pairs, the last in segment 9.

Distribution, North Island, New Zealand.

1. ToKEA ESCULENTA, sp. n, (Text-figs. 62-67.)

This is one of the edible earthworms alluded to in Mr. Best's

article as “* Kurekure ”—as being a specially tasty article of food.

Asa matter of fact, under this name I find two species recog-

nisable, viz., this and the following.

Mr, Best states that it is ‘“a short red or brown worm about

6 inches in length; found in stony places.”

Of this species I received four individuals.

The colour (in formol) isa rather dark purplish-red, paler below ;

but the pigment extends further round the body than usual; the

anterior end is not perceptibly darker; the clitellum is brown.

Dimensions. The largest is,115 mm. in length; the one studied

is 100 mm. x 6 mm., with 110 segments.

The body is cylindrical.

Prostomium is epilobic, 1/2; the posterior groove is but feebly

developed, but visible when the buccal region is everted.

Chete: the 4 on each side are nearly equidistant, and when

viewed from above both ¢ and d are visible; i. e. d is dorsally

placed and ¢ is on the lateral line. ;

In the mid-body aa=be=cd >ab : dd=nearly 2 aa.

Tn the preclitellar region the gap aw becomes greater and ab

rather smaller.

Clitellum: this is complete (i. e. girdle-shaped) and well

developed over the four segments 14-17, where the interseg-

mental furrows are obliterated. The dorsal surfaces of 13 and 18

are also glandular, but the clitellar colouring is not so definite,

and, moreover, the grooves 13/14 and 17/18 are deep.

Genital pores, kc. (text-fig. 62).—There is a pair of male pores

in the 18th segment; each is a small pit,;in a small, oval, pale

spot, and from this small pit a little papilla projects up to the

level of the body-wall (text-fig, 63).

This oval poriferous area is in line with a.

Tubercula pubertatis, in the form of small paired glands, are

* “Toke” is the Maori for Earthworm.

1904. | EARTHWORMS FROM NEW ZEALAND. 241

present on the hinder margin of segments 16 and 17, lmediad

of the line a.

The two oviducal pores are close to the median line, near the

anterior margin of the 14th segment.

Text-fig. 62.

ae A

——————— saa

2 a

fa)

Text-fig. 62. Tokea esculenta.—Ventral view of clitellar region, &c. (4), showing

male pores and arrangement of tubercula pubertatis and disposition of chate.

This and the views in text-figs. 68, 70, 73, 76, 78, and 80 are somewhat diagram-

matic in that they are represented as flat projections, but the relative spacing of

the cheete, &c., is correct.

Text-fig. 63. Tokea esculenta.—Enlarged view of the male pore, the actual aperture

is situated on a small papilla (C) which projects from the bottom of a pit (B),

the skin round which is paler (A) than the surroundings. The cheta 6 is

shown on the left side.

There are three pairs of spermathecal pores on the hinder

region of segments 6, 7, and 8; these are not intersegmental, but

are situated about midway between cheta @ and the margin, in

each case.

I could not detect dorsal pores.

Internal Anatomy.

There are 8 stout septa, behind segments 7 to 14; but, in com-

parison with the thickness of the body-wall, their thickness is not

so great as one would expect.

The dorsal vessel is single; the last of the four pairs of hearts

is in segment 13. In the heart-segments there is a supra-enteric

vessel, with which the hearts are in part connected.

The worm is micronephric*, and these organs commence in

segment 3. In the last 20 segments of the body there is, in

addition to micronephridia, a pair of compact groups of tubules

* | think this term, which was first employed by Vejdovsky, is preferable to my

term “ plectonephric,” for it is by no means certain that a “network ” of tubules

always (if ever) exists.

Proc. Zoot. Soc.—1904, Vou. I]. No. XVI. 16

242 DR. W. B, BENHAM ON [ Nov. 15,

constituting a meganephridium. In these the nepridial funnel

is present and can be traced into communication with the

nephridium, which opens to the exterior, probably in line with a,

as it is in another species which was more carefully studied in

this respect. I did not trace it in the present species. Not only

is there this meganephric nephridiostome in these hinder segments,

but a similar funnel is present throughout the worm. In the

micronephric segments it is unconnected with the nephridium

and has no external opening.

Text-fig. 64.

Tokea esculenta.—A somewhat diagrammatic drawing of a dissection of the worm,

showing the tongue-shaped form and sub-enteric position of the prostates

characteristic of the genus; the intestine is cut away exposing the glands; the

ventral vessel and nerve-cord remaining éz sitw. INT., mtestine; D.V., dorsal

blood-vessel; V.V., ventral blood-vessel; N.C., nerve-cord.

The presence of a funnel, independent of the micronephridia,

throughout the worm is of very considerable interest. Such an

arrangement has not hitherto been recorded. The funnel is of

the same size and structure as that of the meganephridia, and

both in its large size and peculiar form differs from any funnel

hitherto figured. In each of the species of Yokea the same

general arrangement occurs, though the details as to character

and arrangement and extent of the micronephridial tufts differ in

each case. I am preparing a detailed account of the excretory

1904. | EARTHWORMS FROM NEW ZEALAND. 243

apparatus of these worms, so that I will not further describe it

here.

Alimentary system.—The gizzard is small, rounded, and hidden

by the extrinsic muscles of the pharynx; it lies in segment 5.

There are no definite cesophageal glands, but in segment 15 the

tube is dilated, and its dark vascular wall contrasts with the

paler wall of the neighbouring region, and here its lining is

thrown into a series of horizontal lamelle.

The intestine commences in segment 16; there is no typhlosole.

Reproductive system.—The testes and funnels are free and, like

the ovaries, lie in the usual segments. The two pairs of sperm-sacs

lie in segments 9 and 12: their wall is smooth. Further, on the

anterior wall of segment 13 is a minute curved sac, close to the

gut, whose curvature it follows; it is of about the same size as a

similar sac in the 14th segment, which J take to be the ovisac.

Text-fig. 65.

Tokea esculenta.—An asymmetrical arrangement of the prostates, such as occa-

sionally occurs in some species of the genus; V.V., ventral blood-vessel.

Spencer has noted in some of the Cryptodrilids studied by him

such an extra sac in the 13th segment, which he regards asa sperm-

sac. I could not discover any developing sperms by teasing this

sac on the slide, and I have not yet studied it in sections. I have

met with similar structures in some other species.

16*

244 DR. W. B. BENHAM ON [ Nov. 15,

The prostates have a form and disposition on the body which

appear to be peculiar and characteristic of the genus. Hach

prostate is a long “‘tongue-shaped” gland (text-fig. 64, p. 242),

i. é., though it is tubular in that its axis is hollow, it is not cylin-

drical but somewhat flattened, extending through several segments,

and its apex is usually recurved. The pair of glands, typically,

he side by side, pressed against the body-wall below the gut,

and indeed below the ventral blood-vessel; but in individual

cases the right and left glands are asymmetrically disposed. For

instance, in one case (text-fig. 65, p. 243), the right gland passes

backwards from its duct to the 24th segment, and then bends for-

wards on the left side, and its apex lies in the 21st segment; whereas

the left gland passes forwards into the 16th segment, then curves

round to the right side, passes at first backwards, and then

obliquely to the left side, so that its apex lies in segment 20.

Text-fig. 66.

Text-fig. 67.

Text-fig. 66. Tokea esculenta.—A diagram, compiled from serial sections, showing

the course of the sperm-ducts, their union, and junction with the canal of the

prostate, well within the region of the glandular tissue. PR., prostate; P.R.L.,

its lumen; PR.D., its duct ; S, septum; V.D., vas deferens.

Text-fig. 67. Tokea esculenta.—Spermatheca (X12. Camera outline of mounted

specimen).

The surface of the gland is smooth; it is constricted by the

successive septa, and at its anterior end gives rise to a muscular

duct which passes to the exterior in segment 18. Sections

1904. | EARTHWORMS FROM NEW ZEALAND 245

through this region of the body show that the two sperm-ducts

run back separately along the body-wall as far as the 18th

segment, when, meeting the prostate duct, they pass on to its

dorsal surface into the 19th segment, and after uniting with one

another open into the canal of the prostate gland some distance

from the commencement of the muscular duct (text-fig. 66).

The prostate is traversed by a canal lined with columnar cells ;

at intervals this canal receives small canalicules, around which

the gland-cells are grouped and into which they open. The

arrangement is similar to that described by Miss Sweet for

Plutellus intermedius (loc. cit. fig. 17).

There are no penial cheetze.

There are three pairs of spermathece in segments 7, 8, and 9;

each (text-fig. 67) is an ovoid sac passing gradually into a short

wide duct, which receives the diverticulum close to the body-wall.

The diverticulum is a short ovoid body, with duct; it is about

3 the length of the main sac.

The diverticulum of this and the following species is placed

mediad of the sac.

2. TOKEA SAPIDA, Sp. n. (Text-figs. 68 & 69.)

This species is founded on a single specimen of about twice the

size of the former species, but included with it as “‘ Kurekure ”

by Mr. Best.

Text-fig. 69.

Text-fig. 68.

Text-fig. 68. Tokea sapida, similar view to that in text-fig. 62. (x 6.)

Text-fig. 69. Tokea sapida.—Spermatheca (x 12. Camera outline of mounted

specimen).

In colour it is purple-red, with an imperfectly developed

clitellum of a deeper red than the rest of the body.

Dimensions. 200 x 8 mm., with 190 segments. The pre-

clitellar segments are biannulate.

246 DR. W. B. BENHAM ON [ Nov. 15,

Cheice. ‘These are smaller than in 7”. esculenta, and have a

different arrangement; for, when viewed from above, d is more

laterally placed and ¢ is not visible. In other words, the dorsal

gap (dd) is much greater. The formula is:—ab=ed; be>ab;

aa=14 ab; dd=5 ab. I cannot detect any cheete in front of the

10th segment on examination by a dissecting-lens.

Clitellum: not yet fully developed, as intersegmental grooves

are still unobliterated, though the difference in colour is well

marked. It is “complete,” and covers the five segments 14-18.

Genital pores, dc. (text-fig. 68, p. 245).—On segment 18 is a pale,

tumid, transversely disposed ridge, somewhat enlarged at each end :

it extends from 6-6, and cheta «a appears to be absent. The actual

male pore is uncinate, and on the mesial side of 6, but close to it.

No tubercula pubertatis ave present. The three pairs of sperma-

thecal pores are at 6/7, 7/8, 8/9, in line with the gap ab.

Internal Anatony.

There are eight stout septa, behind segments 6-13, but the

first and last are less stout than the others.

The last heart is in segment 13.

The gizzard, in segment 5, is of large size; the cesophagus

presents no dilatation.

The nephridia have the same general arrangement as in

T. esculenta.

The genital organs agiee on the whole with the latter species ;

but i the single individual in my possession the prostates are

asymmetrically arranged, viz., the left gland extends back to

segment 23, and then bends forwards and ends in the 23rd

segment. But the right gland is bent in an $-shaped manner,

and lies wholly im segments 18, 19, and 20; but, as in the

preceding species, both are pressed against the body-wall.

The spermathece, in segments 7, 8, 9, differ in shape from

those of 7’. esculenta; the diverticulum (text- fig. 69, p. 245) being

globular, with a short duct, opening into the spermathecal duct

close to the body-wall.

Loc. Ruatahuua, North Island, New Zealand.

3. TOKEA UREWER&, sp.n. (Text-figs. 70-72.)

This worm is known to the natives as “ Pokotea,” and is

described by Mr. Best as “a short white worm.” TI received

three specimens, which in formol are quite a pale pinkish brown,

paler still behind the clitellum, which is orange or orange-brown,

with a dusky anterior margin.

Dimensions. 65 to 80 mm. in length, with a diameter of 7 or

8 mm.; there are 78 segments in the larger individual. The

segments are not TRTEIEUO

The prostomium is 4 epilobic ; without a transverse furrow.

The chete have the same general arrangement as in the

1904. | EARTHWORMS FROM NEW ZBALAND, 247

preceding species; when viewed from above d is dorsally and ¢

laterally placed, but in the tail the line ¢ approaches d.

aa=ab=cd; be>ab; dd>be. But one differences, measured

in millimetres, are very slight: thus aa=2°5; be=3; dd=4.

The cliéellum is well marked, complete, and covers the four

segments 14-17; but the ventral surface of the last segment 1s

less glandular than the rest.

Banta pores, de. (text-fig. 70).—On segment 18 is a pair of

subcircular depressions, with a distinct, slightly raised margin ;

this is in line a.

Text-fig. 70.

Tokea wrewere, sunilar view to that im text-fig. 62. (x 6.)

There is asingle, median tuberculum pubertatis, in the form of

an oval, glandular, depressed area, extending from a—a, between

segments 19/20.

The three pairs of spermathecal pores are in line a.

Internal Anatomy.

The seven septa behind segments 6 to 12 are only slightly

thicker than the following.

The last heart is in segment 12.

The micronephridia are more delicate than in the preceding

species, and do not form so dense a covering to the body-wall.

The meganephridia are confined to the ee 12 to 15 segments.

The gizzard (in segment 5) is longer than in 7. agen a,

though not so wide.

The cesophagus is more or less dilated in segments 11-15; and

in the first two of these the thick vascular wall is provided with

Jamelle internally.

The prostates extend back to the 26th segment; the muscular

duct is long, narrow, and more or less undulating, becoming

thicker where it dips into the body-wall.

248 DR. W. B. BENHAM ON [ Nov. 15,

The three pairs of spermathece ave in segments 7, 8, 9; each

sac is ovoid (text-fig. 72), and the duct is much narrower than the

sac; the diverticulum is small, and also has a narrow duct.

Loc. Ruatahuua, North Island, New Zealand.

Text-fig. 71.

N.C.

Text-fig. 71. Tokea urewere.—View of the anterior end of the prostate, showing the

long, narrow and undulating duct. N.C., nerve-cord.

Text-fig. 72. Tokea urewere.—Spermatheca (Xx 12. Camera outline of mounted

specimen).

4, ToKEA HUTTONI, sp. n. (Text-figs. 73-75.)

A single individual collected by Capt. Hutton; it is much

bleached, but was apparently purplish.

Text-fig. 73.

Age Bs wile

Tokea huttoni, similar view to that in text-fig. 62. (x 9.)

Dimensions. 80X5 mm., with 63 segments; but possibly

imperfect *.

Prostomium 3 epilobic; segments of body not annulated.

* On opening the tail, I find no meganephridia ; perhaps the true hinder end had

been severed and the wound healed. ‘The lining of the intestine was continuous

with body-wall round anus, but this region was thin.

“

1904. ] EARTHWORMS FROM NEW ZEALAND. 249

Chete : when viewed from above, both ¢ and d are on the

dorsal surface, and ¢ is well above the lateral margin.

aa=be=cd; ab<te; dd=2b>be.

The clitellum is fully developed, complete, and includes seg-

ments 14 to 17; but both the 13th and 18th segments are

glandular on the dorsal surface, the furrows 13/14 and 17/18

are, however, quite deep, whereas the intervening ones are

obliterated.

Genital pores, tc. (text-fig. 73)—In segment 18 is a slight

circular dépression with raised margin, in line with the gap ab;

the cheeta @ appears to be absent. There is a single median, oval

tuberculum pubertatis on the hinder margin of the 18th segment,

extending from a—a.

The three pairs of spermathecal pores are on the hinder

margins of segments 6, 7, 8, in the gap ab, though nearer to @

than to b.

Internal Anatomy.

There are seven thickish septa behind segments 7-13.

The last heart is in segment 12.

Text-fig. 74.

Text-fig. 75.

Text-fig. 74. Tokea huttoni—View of the prostates, which, though symmetrical,

have assumed a reversed position (? abnormal), with the anterior end directed

forwards. N.C., nerve-cord. ?

Text-fig. 75. Tokea huttoni—Spermatheca (X12. Camera outline of mounted

specimen).

The gizzard is long, narrow, and the wall rather thin.

The micronephridia form a pretty close felt over the body-wall,

and commence (as in other species) in segment 3.

The prostates (text-fig. 74) are shorter than in either of the

preceding species, and both are directed forwards, but lie side by

250 DR. W. B. BENVIAM ON [Nov. 15,

side below the gut. They only extend through the three segments

Toran 16:

The duct is very short and thick. s

The spermatheca (text-fig. 75, p. 249) likewise has a short duct,

and the diverticulum is similar to that of the foregoing species.

Loc. Whangarei, North Island, New Zealand.

Remarks.—Vhis species is evidently nearly allied to 7’. urewere,

but differs from it in the relative position of the male and sperma-

thecal pores, in the position of the tubercula pubertatis, m the

cheetal formula, and especially in the relation of ab to be, as well

as in the less extent of the prostate and its duct.

5. TOKBA SUTERI, sp. n. (‘Text-figs. 76, 77.)

A single individual collected by Mr. H. Suter in 1899. It

differs in general appearance from any of the preceding species, not

only in size, but in colour. It is dark purplish brown throughout

the entire dorsal surface ; the clitellum is reddish purple.

Dimensions. 50X5 mm.; with 86 segments.

Prostomium + epilobic; without transverse groove.

The chete are practically equidistant, 1.e. aa=ab=be=dd

= 2 mm., while cd is only 1°75 mm.

Text-fig. 76.

Text-fig. 77.

A. B.

Text-fig. 76. Tokea suteri, similar view to that im text-fig. 62. (x 9.)

Text-fig. 77. Tokea sutert.—Spermatheca (X 12. Camera outline of mounted

specimen).

On the clitellum the gap aa is a little less than posteriorly, and

in the preclitellar region still less; the gap ab is less in the _

eclitellar region, but not anteriorly. But these differences are

not perceptible till measured with compasses.

Clitellum in segments 13-17 (=5 segments), girdle-like.

Genital pores, ke. (text-fig. 76).—A pair of small papille on

segment 18, in the line of a.

There is a pair of small twbhereula pubertatis—subarcular and

pitted—between 18/29 in line a, and extending further mediad

thereof than do the porophores.

1904. | EARTHWORMS FROM NEW ZEALAND. 2

Ril

The three pairs of spermathecal pores are in line a, at 6/7,

7/8, 8/9.

Internal Anatomy.

There are five slightly thickened septa, behind segments 8

to 12.

The last heart is in segment 12.

The gizzard is feebly developed; its wall is not thicker, though

tougher, than that of the cesophagus.

The prostates are quite typical, and extend back to segment

25, with tips recurved ; the duct is short and narrow.

The spermatheca (text-fig. 77) has a long duct, with a long

narrow diverticulum opening into it at the body-wall,

Locality. Auckland, New Zealand.

Remorks.—Anatomically there is a rather close resemblance

between this and the two preceding species; but in the coloration

and dimensions it is remarkably distinct, while the practical

equidistance of all the 8 chetie, the paired tubercula pubertatis,

and the details as to the prostate and septa, mark it off as

distinct.

6. ToKEA KIRKI, sp. n. (Text-figs. 78, 79.)

Five specimens collected by Professor H. B. Kirk, in 1902, are

about the same size and colour as 7’. esculenta, 7. e. reddish purple,

but rather paler than the latter; the pigment extends further

round to the ventral surface than is usually the case, and ceases

about the level of 4.

The clitellum is brownish red.

Text-fig. 78.

Text-fig. 79.

pie war iG

Text-fig. 78. Tokea kirki, similar view to that in text-fig. 62. (x 7.)

Text-fig. 79. Tokea kirki.—Spermatheca (X 12. Camera outline of mounted

specimen). This has no muscular duct distinct from the sac.

Dimensions. The length varies from 80 to 100 mm., with a

diameter of 6 mm. for 110 segments in the larger individuals.

The chete are practically equidistant ; but on measurement it

is found that ab=be=cd (=1°75 mm.), while aa=dd=2 mm.

252 DR. W. B. BENHAM ON [ Noy. 15

When viewed from above, d is dorsally placed and ¢ rather

above the lateral margin.

The elitellum is not fully developed on any of the specimens,

as the intersegmental furrows are distinct ; but the characteristic

coloration extends over the five segments 13 to 17.

Genital pores, dc. (text-fig. 78, p. 251)—The male pores are

situated in what appears to be a pair of tubercula pubertatis on

segment 18; there is a very large, broad, subcircular papilla on

each side, extending from nearly @ to c; this carries a somewhat

quadrangular depression surrounded by a distinct rim or margin ;

this depression is glandular and has all the characters of a tuber-

culum pubertatis (as seen in section) ; and the male pore is quite

small and situated in the outer edge of this gland, just within the

margin, 2. e. nearly in line 6. The two oviducal pores are situated

close together, in a pale area, near the anterior margin of

segment 14.

There are only two pairs of spermathecal pores, situated at

7/8, 8/9.

Internal Anatomy.

There are no specially thick septa.

The last heart is in segment 12.

The gizzard is quite small; the esophagus is very much dilated

in segment 14, but there is no constriction separating it from the

coma < ; its lining, however, is thrown into a series of horizontal,

lamelliform folds.

The micronephridia, instead of being spread over a great part of

the body-wall, in each segment, form a very distinct and narrow

row close to the septa.

The prostates extend from the 18th to 23rd segments, and each

is provided with a narrow duct.

There are two pairs of spermathece in segments 8, 9; each

(text-fig. 79) is a somewhat pyriform sac, without a distinctly

marked duct; the diverticulum is of the same shape, and also

without definite duct.

Locality. Ohaeawai, North Island, New Zealand.

Remarks.—A very distinct species, characterised both by the

remarkable coincidence of male pore and copulatory tubercle, and

by the possession of only two pairs of spermathece.

7. ToKEA MAoRICA, sp. n. (Text-figs. 80-82.)

I have eight specimens of this species, collected at different.

times by Mr. H. Suter, at different places in the neighbourhood

of Auckland. The colour is a dark purplish brown, more brown

than purple in these alcohol specimens, and resembles that of

T. suteri. The clitellum is paler brown ; the chetz are inserted

in white spots, as in Plagiocheta sylvestris, to which also they

bear a resemblance both in colour and their short, stout form ;

indeed, from a hasty first glance, I had placed them with the

latter genus, awaiting examination.

1904. | EARTHWORMS FROM NEW ZEALAND, 253

Dimensions. The largest specimen measures 70x 4 mm., and

consists of 75 segments; the shortest mature individual measures

25x 2mm., and likewise consists of 75 segments; and the same

number occurs in a worm of intermediate size.

We have here an instance—unique, so far as I am aware—of

an earthworm increasing in length without adding new segments

posteriorly. It is true that we do not know much on this subject ;

but the accepted view is that the number of segments is continu-

ally added to during growth.

With regard to this, Beddard (95, p. 2) states :—‘ There are

at present no exact data as to the constancy of the number of

seoements among Earthworms. In all probability the number is

not absolutely fixed, but there appears to be a mean for each

species round which there is a certain amount of variation.”

Text-fig. 80.

ives B.

Tokea maorica, similay view to that in text-fig. 62. (X 11.)

Prostomiwm i epilobic; without a transverse groove.

Chetal formula :—ab<ecd<be; be=2ab=aa; dd=2 aa.

Clitellum girdle-like, over the five segments 13 to 17.

Genital pores, dc. (text-fig. 80).—On segment 18 is a transverse,

elliptical area, paler than its surroundings; glandular centrally,

with a distinct margin surrounding it. It extends between points

a little outside a, on each side. This is a tuberculum pubertatis

(text-fig. 81, p. 254), and the male pore is situated on the thickened

margin, at each pole of the ellipse in line with a (which is absent

in this segment). There is a second tuberculum pubertatis of the

same shape, but of much smaller size, on the hinder border of

254 DR. W. B. BENHAM ON [Nov. 15,

segment 14, and not quite reaching the line of @ on either side.

This tubercle is absent in some individuals.

The oviducal pores are wide apart, nearly in line of a, but a

little mediad.

The two pairs of spermathecal pores are distinct and slit-like,

about midway between the cheeta @ and the posterior margin of

each of the segments 8, 9.

Dorsal pores are present, at least in the postclitellar region.

Text-fig. 81.

Text-fig. 81. Tokea maorica, the prostates and the gland of the tuberculum

pubertatis (T.GL.).

Text-fig. 82. Tokea maorica—Spermatheca (x 12. Camera outline of mounted

specimen).

Internal Anatomy.

The last heart is in segment 12.

The gizzard, though small, is distinct.

The micronephric tubercles are more delicate than in other

species, and all concentrated to form a latero-ventral mass on each

side.

The prostates extend back to segment 23; the duct is narrow

and rather long, and between the pair of ducts les a great

glandular mass, which thrusts the anterior ends of the prostates

outwards, so that the duct is transverse.

Each of the spermathece (text-fig. 82), in segments 8 and 9,

consists of a pyriform sac with a very thick duct, thicker than

the neck of the sac. A small tubular diverticulum, dilated

terminally, opens into the duct near the body-wall.

1904. ] EARTHWORMS FROM NEW ZEALAND. 255

Localities. Auckland: Waitakerei Bush and Nikau Palm Bush

(near Auckland); North Island, New Zealand.

Remarks on the Genus Token.

The distribution of the seven species attributed to this genus

extends over a considerable portion of the North Island of New -

Zealand, for the localities from which specimens have been

collected are widely separated and cover more than three degrees

of latitude, the most northerly spot being Ohaeawai, practically

in latitude 35° south, and the most southerly spot, Ruatahuua,

is south of 38°.

The really striking fact illustrated by these seven new species

is the presence in the North Island of New Zealand of a Crypto-

driline genus as an important, and apparently predominating,

element in the Earthworm fauna.

Hitherto, as I have remarked, our knowledge of this fauna in

New Zealand has been derived from a study of the South Island

representatives, and here the predominating element is the

Acanthodriline series.

This was recognised some years ago by Beddard, who in his

‘Monograph’ (p. 154) says :—‘‘ New Zealand is essentially

different from Australia. The prevalent forms in Australia are

Perichetide and Cryptodrilide ; the most abundant worms in

New Zealand are Acanthodrilide.”

I have, in my Presidential Address to Section D of the

Australian Association for the Advancement of Science (02, d),

dealt with this difference very fully, and little thought that the

North Island would present so different a set of Harthworms.

Beddard, indeed (1890, p. 285), suggests that “the North

Island may prove to be move ‘ Australian’ in its character when

it comes to be known”; but the grounds for this statement at

that time seem to have been very slender, for only one worm

had been recorded from that island, viz. Schmarda’s “ Hypogeon

orthostichon,” of which nothing was known beyond the few facts

of its external appearance recorded by that zoologist, and sufficient

of its internal anatomy noted by Beddard to indicate its Crypto-

driline affinity. So characteristic, indeed, is the Acanthodriline

group of worms, that doubt has been thrown upon the occurrence

of this species in New Zealand. Captain Hutton (’79) doubted

if on geographical grounds, for Schmarda gives the locality

as “Mount Wellington, New Zealand”; and Hutton, knowing

that there was no mountain of any size called by this name in

New Zealand, stated that probably some confusion with “ Mt. Wel-

lington,” Hobart, Tasmania, had crept into Schmarda’s notes.

In my Presidential Address, I omitted it entirely from the list

of earthworms occurring in this Colony, and even so far as to

read a note (02, c) attributing it to Tasmania. But, in view

of the undoubted occurrence of Cryptodriline worms in New

Zealand, this doubt is itself rendered somewhat doubtful and

256 DR. W. B. BENHAM ON | Nov. 15,

possibly premature. No such mountain occurs on ordinary maps

of New Zealand as “‘ Mt. Wellington,” and I therefore consulted

Prof. H. B. Kirk, who has an extensive first-hand knowledge

of the geography of the North Island: in response to my query,

he informs me that ‘‘ Mt. Wellington is one of the small volcanic

cones just out of Auckland. It is, now, under grass, and is in

a fully cultivated district.”

The geographical doubt is thus set at rest; for Schmarda

visited Auckland, and at that period (1860, about) it is probable

that little or no cultivation had then been carried on on this

small mountain ; and, in light of these new observations of mine,

it seems that we must now admit into our fauna Schmarda’s

species.

Our knowledge of its internal anatomy is due to Beddard (’92),.

who recognised that it belonged to the genus MJegascolides, as he

understood it, but by Michaelsen it is placed in Fletcher's genus

Notoscolex ; but as we did not know the condition of the nephridia

in the poster ior end of the body, it is impossible to be sure of the

correctness of the genus. (See later.)

N. orthostichon appears—from the necessarily brief account

given by Beddard—to differ from the species of Vokea cay in the

size of the prostate, which that zoologist states is “short and

tubular.” He says nothing as to qihes her it is confined to one

segment or extends bey ond - it; but presumably it does not exhibit

the characteristic disposition found in TZokea, though, as we see,

in 7’. hutioni this gland is “short” as compared with its length

in the other species, as it only traverses three segments.

It is vemarkable, however, that it differs fe ‘om all the other

species of Votoscolex, in which the clitellum covers fewer than ten

segments, and in which the arrangement of the cheete is unchanged

at the posterior end of the body (Michaelsen, ’00), in that the

cheete are equidistant (¢.¢. the formula is a)=bc=cd), as is the

case with several of my species; and if we suppose that it does

belong to the same genus as do the seven new species herein

described, it might be suggested that one or other of the latter is

identical with JV. orthostichon, especially as I have obtained two

species from Auckland.

Let us, then, examine such characters which appear to be

specifie as Beddard mentions. Firstly, it has two pairs of

spermathecze, in which it agrees with T. kirki and 7. maorica—

the latter of which came from the Auckland district. The other

Auckland species, 7’. suteri, is out of court, as it possesses three

pairs of spermathece. The two species to which it presents this

resemblance have very different forms of spermathece. Un-

fortunately, Beddard’s description is too meagre to allow us

to judge whether in J. orthostichon the organ resembles either

of them ; for, he says, “ each [spermatheca] has a small diverti-

culum, pyriform i in shape like the main pouch.” Such a general

description applies to several of the above species.

Now, in 7’. maorica the duct of the spermatheca is very wide ;

1904. | EARTHWORMS FROM NEW ZEALAND. 257

and if such a duct exists in JV. orthostichon, it seems unlikely that

it would have escaped Beddard’s notice. Moreover, in 7’. maorica

the chetal formula is different, whereas in 7’. kirki it is similar

to that of WV. orthostichon, and the spermatheca is without a

distinct muscular duct. The only other characters are the extent

of the clitellum and its size.

Beddard gives the former as occupying segments 14 to 17

inclusive, whereas in my species it begins at segment 13; and as

to size, Schmarda gives it as 80 cm. x 4 em., with 60 segments—

distinetly smaller than 7’. kirki, though approximately that of

T. maorica.

It appears, then, to be distinct from either of the two species

possessing only two pairs of spermathece ; for, apart from any

similarities, the prostate in those two species is of considerable

length. But it agrees more closely with 7’. huttoni, in size, in

cheetal formula, extent of clitellum, and small size of prostate ;

but in that species there are three pairs of spermathece.

So far, then, as our information allows us to judge, Schmarda’s

species is distinct from any of those described in the present

paper; and we have not sufficient information to allow us to say

definitely that it even belongs to the genus Tokea, although it

appears not improbable that it does so; but Beddard states that

the sperm-sacs number three pairs, and occupy segments 10,

11, 12—an unusual arrangement, even in WVotoscolex, where 11,

12 is the more general position.

Remarks on the Genera Megascolides and Notoscolex.

It is very evident that this new genus Tokea is nearly allied to

the Cryptodriline genera d/egascolides and Notoscolex; but it

appears to me to differ in details from either of them. It is

notoriously difficult to define these Australian genera; and it is

unfortunate that the opinions of our two leading systematists,

Beddard and Michaelsen, are absolutely at variance as to the

characters and limitations of the genus Megascolides.

This name was given by Prof. M‘Coy in 1878 toa worm, W/. aus-

tralis, which, at a later period, received full anatomical treatment

at the hands of Baldwin Spencer (1888); and this it is which

Michaelsen (1900) takes as the type of the genus, of which he

gives this diagnosis (p. 182, translation): “ Hight cheete ; clitellum

beginning at or before segment 14 and extending over 6 to

93 segments. One pair of female pores; spermathecal pores 2

to 5 pairs, the last being on 8/9. Gizzard 5 or 6, or 5 and 6.

Excretory organs consist of diffuse nephridia, to which is added a

pair of meganephridia in each of the posterior segments. Prostate

tubular (sometimes lobate).” (Italics are Michaelsen’s.)

It will be seen, then, that in some of the above characters the

new genus agrees with MJegascolides, and notably in the co-

existence, in the hinder segments, of meganephridia with micro-

Proc. Zoon. Soc.—1904, Vou. IT. No. XVII. 177

258 DR, W. B. BENHAM ON | Nov. 15,

nephridia, But it differs in that the clitellum in my new species

is less than that characteristic for the above.

The only difference between Megascolides and Votoscolex upon

which Michaelsen lays special stress is the presence of mega-

nephridia posteriorly. The diagnosis of the latter genus he gives

(p. 187) as: “ Cheetze 8, Clitellum, beginning at or before segment

14, extends over 32 to 103 segments. The female pores are

usually 1 pair, seldom unpaired [median]. Spermathecal pores

2 pairs in the intersegmental furrows 7/8.8/9; varely shifted

backwards on to the 8th and 9th segments. One gizzard in 5

or 6. Plectonephric; nephridia diffuse. Prostate usually lobate,

sometimes tubular.” (Italics are Michaelsen’s.)

The two distinguishing characters, then, are the limited number

of spermathece and the diffuse nephridia, without the posterior

meganephridia. It is unfortunate that in a considerable number,

perhaps the majority, of the species included in Wotoscolex the

condition of the posterior nephridia is unknown.

If, then, we depend on these formal diagnoses, we should

no doubt refer my new species to the genus Megascolides,

especially if they had been found in Australia.

On examining the anatomical details of the only species of

Megascolides the anatomy of which is thoroughly known, viz.

WM. australis, we note several differences from Vokea; amongst

others :

(a) The cheetz are in couples, all ventral.

(b) The prostates are compactly coiled, cylindrical tubes, similar

in general shape to those of true Acanthodriline worms ;

they are, too, situated laterally, and confined to segment 18.

substance of the body-wall; which is also the case in the

only other species of the genus that has been investigated

by means of sections, viz. I. ilawarre (see Sweet, p. 113).

(d) The spermatheca has quite peculiar rosette-shaped diver-

ticula.

But the above statements do not apply to each of the other

three species included in the genus ; for the prostates are flattened

and more or less lobate ; the spermatheca in J/. cameron is similar

to that of many other Cryptodriline species. Nevertheless, in all

of them the chet are distinctly coupled; 7.e., the spaces aa

and be are greater, much greater in some cases, than ab, whereas

in Tokea these spaces are more or less equal and the “ coupling”

is quite unnoticeable. It is not probable that any of these

characters by themselves are of generic value. And, as a matter

of fact, when we come to look into the anatomy of these four

species we find numerous differences—which, in the case of other

genera, Michaelsen has considered as of sufficient importance to

deserve generic rank. I refer, for instance, to the fact that

M. insignis and M. cameroni have only one pair of testes, and

this in segment 11. But it is clear that the only characters

1904. ] EARTHWORMS FROM NEW ZEALAND, 259

which these four species have in common, and in which they

agree with Z'okea, is the coexistence in the posterior segments of

meganephridia and micronephridia.

Let us turn to Beddard’s conception of the limits of the genus

Jegascolides (= Notoscolex Fl.) and Cryptodrilus Fletcher (the

history of which is well given in his Monograph, pp. 445 eé seq.).

He draws attention to the fact that in nearly all the worms

described by Spencer, and in a considerable number of those

described by Fletcher (at any rate in such cases in which sufficient

information is given to enable a comparison to be made), there is

a correlation between certain of the internal organs (p. 447), viz.

that “ they have either (1) paired nephridia, tubular spermidueal

glands, and last pair of hearts in the 12th segment; or (2) diffuse

nephridia, lobate glands, and last heart in the 13th segment.”

He decides to regard those species with the first set of characters

as belonging to the genus ‘“‘ J/egascolides,” and those with the

second group as “ Cryptodrilus.” He admits that there are

exceptions to these correlations.

It thus comes about that Michaelsen uses the generic name

Megascolides for “‘ micronephric,” whereas Beddard employs the

same name for ‘ meganephric” worms. But the meganephric

species are placed by Michaelsen in the genus Plutellus, which

is thus extended to include species previously attributed to

Argilophilus of Hisen, Jegascolides of various authors, Cryptodrilus

of Fletcher, as well as Plutellus of Perrier and Benham.

Thus we have this unfortunate confusion in the employment of

generic names :—

Megascolides M‘Coy, in Michaelsen’s sense,

= Votoscolex (part) Fletcher,

= Cryptodrilus (part) Beddard ;

Votoscolex Fletcher, as used by Michaelsen,

= Notoscolex Fl.+ Cryptodrilus F\.

= Megascolides (part) Spencer ;

Cryptodrilus, i Beddard’s sense,

= Cryptodrilus Fletcher,

= Megascolides (part) Spencer,

= Notoscolex (part) Fletcher ;

JWegascolides, 1 Beddard’s sense,

= Cryptodrilus (part) Fletcher, Spencer,

= Megascolides (part) Spencer,

= Argilophilus Kisen,

= Plutellus Perrier.

Tt is natural that ina group like Earthworms a mixture and

confusion of this sort is likely to arise as knowledge advances :

and it remains for Prof. Baldwin Spencer with the large stock of

material in his possession, to endeavour to find some more satis-

factory method of discriminating between genera than those

usually employed.

Of course the correct name that should be applied tothe genera

ele?

260 . DR. W. B. BENHAM ON [ Nov. 15,

depends on the law of priority ; and there can be little doubt but

that Megascolides australis is the “type” of Megascolides M‘Coy ;

but it does not seem quite so clear as to whether the word WVoto-

scolex or Oryptodrilus should be retained to apply to the other

genus. But this thorny matter of nomenclature I regret that I

have not time to discuss thoroughly.

Having thus cleared the ground, satisfactorily as it seems, of

the meganephric species, there remain the large series of micro-

nephric species to be dealt with ; and it becomes a question whether

Michaelsen’s characters are good, viz. :

(a) Absence or presence of meganephridia posteriorly ; and

(6) A limited number (two pairs) or a greater number (up to

five pairs) of spermathece.

In regard to the first, it is admitted that in many species we

are ignorant as to whether the large nephridia are or are not

present.

In the second case, in other genera—e. g., Pheretuma and Mega-

scolex—the number of spermathece has not been used asa generic

character *.

Since, therefore, the seven species of New-Zealand worms here

described agree very closely with one another, and except in one

respect differ from those species referred to Jegascolides by

Michaelsen, and since, too, 1 am not in a position to rearrange

the generic characters of these allied genera, it seems to me better

to erect a new genus. It may be that it is only of subgeneric

rank ; but for the present less confusion will arise, I think, if we

regard it as a distinct genus.

The genus Vokea differs from Megascolides, as defined by

Michaelsen, in the following points :—

(a) The limited extent of the clitellum, in which only 4 or 5.

segments are involved, whereas his minimum is six.

(6) The widely separated cheetee, of which the coupling is not

recognisable.

(d) The position of the point of entrance of the sperm-duct

into the prostate-duct.

(e) The existence throughout the body of nephridial funnels,

although in the greater part of the worm these have no

connection with the nephridia.

These are truly small points of difference on which ‘to form a

new genus; but not smaller than that on which JMegascolides is

distinguished from Wotoscolea, viz., the presence in the former of

meganephridia in the hinder segments of the bodyy. In my

opinion, this is by no means a good line of distinction; and even

* The same remark applies to the position of the last heart, utilised in diagnosing

the genera by Beddard; for in Plagiocheta and Octochetus, as in Tokea, we find

species in which it is in the 12th, and others in which it is in the 13th segment.

+ Or, Maocridrilus trom Notiodrilus, m having nephridiopores alternate, instead.

of in line

a 904. | EARTHWORMS FROM NEW ZEALAND. 261

the separation of Plutellus from these two genera, merely on

account of the presence of meganephridia only depends on a point

quite as small.

T have already described a species of Plagiocheta, viz. Pl. rosst,

in which micronephridia replace the meganephridia of the other

half-dozen species ; and embryology has taught us that the one is

derivable from the other condition—that, at any rate in the genera

Megascolides (Vejdovsky) and Megascolex (Bourne), the earlier

meganephridium breaks up into numerous micronephridia.

Is not the separation of the genera according to the condition

of the excretory system a remnant of my own unfortunate attempt

to classify the families of Harthworms into “ Plectronephrica”

and ‘“ Meganephrica ” 2

But without entering upon the laborious task of essaying to

rearrange the ‘“‘ Cryptodriline ” genera, I will express the opinion

that a careful study of the form and structure of the prostate

(together with other characters) may be more likely to lead us in

the right direction. And, firstly, it seems to me profitable to

distinguish the “tubular ”-cylindrical prostate, such as occurs in

Megascolides ambialis, from the flatter, elongated “tongue-shaped ”

form of gland that occurs in Zokea, some species of Plutellus*, and

others, and the “flattened, lobed, and compact” organ characteristic

of such genera as Pheretima and Megascolez. That these may

form a developmental series, Miss Sweet’s work (1900) has rendered

probable; but they also appear to have structural differences

that may turn out to be of diagnostic character. At any rate,

they are easily recognised macroscopic characters; whereas the

study of a worm from which the posterior end has been acci-

dentally destroyed will not enable me to decide, in all cases,

whether it belongs to the genus Jegascolides or to Notoscolex as

defined by Michaelsen.

It is not difficult to imagine the way in which the genus Zokea

has developed from an Acanthodriline stock, in which the cylin-

drical prostate, instead of being coiled compactly and confined to

its proper segment, has burst away from this limited position, and

elongating backwards has not only become flatter but has lost

somewhat of its smooth external surface. Such a form of

“‘tongue-shaped” gland occurs in Nofiodrilus aucklandicus, in

which each prostate extends through five or six segments. From

such a form Rhododrilus may have developed—the posterior

gland has disappeared, and the sperm-duct has shifted forwards

so as to open close to the anterior gland; whereas in Vokew the

anterior gland appears to have gone, and the posterior gland to

have moved forwards to meet the sperm-duct and to open

externally in the 18th segment.

* In looking through the descriptions of new species of Australian worms

published by Spencer (Proc. Roy. Soc. Victoria in 1892, 1895, and 1900), I find that

the only species that possess elongated “ tongue-shaped”’ prostates (he calls them

“tubular,” but the figures show them to be like those of my species) extending

through several segments belong to the meganephric genus Plutellus in Michaelsen’s

sense.

262 ON EARTHWORMS FROM NEW ZEALAND. [ Nov. 15,

The occurrence of a prostate in MWegascolides australis, similar

to that of a typical Acanthodrilid, seems to indicate that the

confinement to a single segment is related to the cylindrical

form ; and it is admitted by Michaelsen that these “‘ Cryptodriline ”

worms, belonging to the subfamily Megascolecine, are derived

from the subfamily Acanthodriline. And, on the other hand,

the peculiar form of the gland in Wotodrilus aucklandicus

illustrates the relation between a looser structure and the

extension of the gland through several segments. But, although

this transition of form between the “ tongue-shaped” and “ eylin-

drical” tubular prostates seems to occur, yet the flattened form

appears a still later development; it occurs, for example, in the

more modified genera, such as Pheretina.

Miss Sweet has pointed out that in the ‘“ tubular” prostate

there is a lumen running the whole length of the gland; whereas

in the lobate form of this organ there is, typically, no central

lumen, and when it exists it is not only very small, but it has no

epithelium. Unfortunately, the species examined by her do not

belong to the genera under dispute, with the exception of Mega-

scolides awarre, in which the prostate is a “somewhat long and

flattened” organ, and has a structure intermediate in some

respects between a truly “lobate” gland, such as exists in

Megascolex, and a “tubular” gland, such as occurs in Plutellus

and others; for the species referred by her to “ MJegascolides”

belong to the genus Plutellws in Michaelsen’s sense.

Tf I have dwelt so much on the form of the prostate, it is

because it seems to me that Michaelsen has not laid sufficient

stress upon this organ in shuffling the members of the “ Crypto-

driline series.” It alone, I admit, will not serve for generic

distinction ; but it may possibly be useful in the formation of

subgenera.

Dunedin, April 10, 1904.

Bibliography.

89. BeppARD, F. E. “ Oligochetous Fauna of New Zealand,”

in Proc. Zool. Soc. p. 377.

*90. “ Glassification and Distribution of Harthworms,” in

Proc. R. Phys. Soc. Edinb. x. p. 235.

92. “On the Harthworms in the Vienna Museum,” in

Ann. Mag. Nat. Hist. (ser. 6) 1x. p. 113.

OR. “Some new and little-known Oligocheta,” in Proc.

R. Phys. Soc. Edinb. xii. p. 33.

"95. ‘Monograph of the Oligocheta.’ Oxford.

90. Bennam, W. B. “An Attempt to Classify Earthworms,’

in Quart. Journ. Mier. Sci. xxxi. p. 319.

“On Acanthodrilus uliginosus,” in Trans. N.Z.

Inst. xxxi. p. 125.

‘On some Earthworms from the Islands around

New Zealand,” in loc. cit. p. 140.

700 (a).

00 (2).

1904. | ON THE HAND OF THE CHIMPANZEN. 263

’02 (a). Benuam, W. B. ‘On the Old and some New Species of

Plagiocheta,” in Trans. N.Z. Inst. xxxv. p. 284.

02 (6). —— “The Geographical Distribution of Earthworms,

etc.,” in Report A. A. A. 8. Hobart, ix. p. 319.

02 (¢). “Note on a neglected Tasmanian Earthworm,”

in loc. cit. p. 383.

Bourns, A. G. “On certain Points in the Anatomy and Deve-

lopment of some Harthworms,” in Quart. Journ. Mier.

Sci. XXXvi. p. 25.

FirercHer,W. ‘ Noteson Australian Earthworms,” in Proc. Linn.

Soc. N.S.W. (ser. 2) 1886, vol. i.

Hutton, F. W. “Catalogue of the Worms of New Zealand,”

in Trans. N.Z. Inst. xi. p. 317, note.

M‘Coy, F. ‘ Prodromus Zool. Victorie. Decade I. 1878.

Micuartsen, W. “ Oligocheta,” in Das Tierreich. 1900.

Spencer, W. B. “On the Anatomy of Megascolides australis,”

in Trans. Roy. Soc. Victoria, i. 1888, p. 3.

—— “Description of Australian and Tasmanian Karthworms,”

in Proc. Roy. Soc. Victoria, 1892, 1895, 1900.

Sweer, G. “On the Structure of the Spermiducal Glands,

etc.,” in Journ. Linn. Soc. (London), Zool, xxviii. 1900,

joe OY),

Vespoyvsky, Hr. “Zur Entwickl. des nephrid. Apparates v.

Megascolides australis,” in Arch. f. mikr. Anat. xl. p. 552.

November 29, 1904.

G. A. BouLENGER, Esq., F.R.S., Vice-President, in the Chair.

Dr. Walter Kidd, F.Z.S., exhibited a drawing of the extensor

surface of the hand of a Chimpanzee and made the following

remarks :—

In the course of an examination of the papillary ridges in

some specimens of Anthropoid Apes and Monkeys certain groups

of ridges were found on the extensor surface of the terminal

phalanges of the hand, apparently identical with those of the

palmar and plantar surfaces. Three specimens of Chimpanzee

living in the Society’s Menagerie were examined, of the ages

1 year 8 months, 23 years, and 6 years. In the oldest of these,

“‘ Mickie,” the ridges were definite and well-developed on the 2nd,

3rd, and 4th digits on both hands; in the youngest specimen,

“ jack,” they were absent; and in “Jimmie,” 24 years old, they

were small and ill-defined, as if in the process of development.

Direction of Ridges.

Mickie. Ridges longitudinal and reaching to the matrix of the

nail on the 2nd, 3rd, and 4th digits.

264 CAPT. R. CRAWSHAY ON THE [ Nov. 29,

Jimmie showed ridges as follows :—

1st digit none.

{ ana » Oblique.

Right hand ...... 3rd_,, transverse at base of digit.

[as 3? 9 4 bb) . 2?

5th ,, nearly longitudinal.

lst digit none.

2nd ,, oblique.

Leftrhand ecccss 38d! x55 sf

[am a -

5th ,, none.

In these three specimens ridges were absent from the corre-

sponding surfaces on the feet.

The well-defined longitudinal direction of the ridges in Mickie

is worth notice. It must be remembered in this connection that

a Chimpanzee walks with the extensor surfaces of the phalanges

touching the ground and the digits turned inwards, so that their

long axes are at right angles to the line of progression of the

animal, and accordingly the ridges of this part also occupy the

same relative position. There is no correlation in this instance

between the act of prehension and the direction of the ridges,

though it agrees closely with the general rule which obtains in so

many regions, that the ridges lie at right angles to the line of

incidence of the predominating pressure on the part.

The following papers were read :-—

1. Some Observations on the Field Natural History of

the Lion. By Capt. Ricoarp Crawsuay, F.Z.8.

[Received June 10, 1904.]

In offermg these observations I wish it understood at the

outset that I do not pose as a great lion-hunter, nor as having

made lion-hunting a special pursuit, but speak merely as one who,

during a period covering at intervals some seventeen years of

travel and residence in Central Africa, has had many experiences

with Lions.

From what I have read, and still more from the opinions I hear

expressed from time to time, prevailing impressions seem so often

at variance with my own observations that I have thought it

worth while to record these latter.

It has always seemed to me that, though much has been written

respecting the habits of the Lion in his natural state, a great portion

of it is more fiction than fact: this, at least, is my experience.

For one thing, the common opinion of the Lion being an animal of

almost exclusively nocturnal habit—rarely or never seen in the

1904. ] FIELD NATURAL HISTORY OF THE LION. 265

daytime unless roused from his lair in the gloom of some dense

clump of bush or reed-bed—is quite erroneous. It is the fact that

Lions are heard very much more at night than in daytime ; never-

theless they are largely diurnal as well, though usually silent.

I have heard Lions roaring at noon on one or two occasions,

and I have many times heard them as late as an hour or more

after sunrise, and as early in the evening asan hour or more before

sunset. The usual time is at sunset and a little after, and more

especially about dawn in the early morning and thence onwards

until broad daylight. I have seen Lions at all times of day,

under all sorts of conditions, in all sorts of country, in all sorts of

weather from misty rain to the hottest noonday sun of the

tropics.

It is certainly not my experience that the Lion’s habit is to

hide himself in dark cover in the daytime—far from it. Other

circumstances being equal, he likes open country and sunlight,

and goes about and lies out. in it freely, nothing being more to

his liking than some coign of vantage commanding a view of the

neighbourhood, where he can stretch himself out and survey the

prospect. I have seen a Lioness and three cubs lying out on a

river sand-spit in Henga in the full glare of the early afternoon

tropical sun, stretched out on her belly, with her cubs crawling

about her back and neck and tumbling over on to the sand. . On

another occasion about 10 A.m., also in Henga, I saw a Lioness

sitting up on her haunches on a flat-topped ant-hill, watching and

listening to my men talking and laughing as they were skinning

an Impala shot half an hour previously.

The same instant as | made her out and levelled my glasses on

her at a distance of some 300 yards, she slunk down behind the

ant-hill—melting away, as it were, in the endeavour to make her

movement as unnoticeable as possible.

T have seen a Lioness crossing the bare scorching lava-covered

plain intervening between the East-Africa and Uganda Pro-

tectorates in the fierce heat of noon at midsummer, with the heat

radiating in lambent tongues from the ground, giving her the

appearance of being enveloped in fire.

Nothing had disturbed her, as this country was uninhabited by

man; she was making her way to the water.

On another occasion I saw three half-grown cubs near the same

spot in the early afternoon, in the hottest sunlight, playing about

in the open on the banks of the stream.

Once, in Henga, I came across a troup of five Lions on the move

at noonday. It had been raining about an hour before, but the

sun was then out in all its power of midsummer, as they crossed

from open country to go into the scrub on the banks of the

Lunyina River.

In Henga, in 1893, about an hour after sunrise in the hottest

time in all the year, a full-grown dark-maned Lion passed down

the valley below me following a game-track at a long striding

walk, throwing up his head and roaring at intervals as he went,

266 CAPT, R. CRAWSHAY ON THE [ Nov. 29,

making daytime hideous and stampeding the game in ridiculous

fashion. Hartebeestes were sneezing, Reedbuck whistling, and

herds of Zebra thundering about all over the place. J was

actually stalking him at an angle to cut him off, when the late

Surgeon McKay fired at another Lion a few hundred yards lower

down the river, killing his first Lion and spoiling my chance.

In 1893 I had a curious experience with a Lion—also in

Henga,—which, for aught I know, may have occurred to me

oftener without my having been made aware of it. In stalking

two old bull Hartebeestes up a slope in country timbered sparsely

with sapling trees and bushes, I passed within some 20 paces of a

Lion lying on the bare burned ground in the shade of a sapling

without knowing it at the time. On my return from shooting

both Hartebeestes, one of my men followed me and told me,

pointing out the Lion lymg under the tree.

On setting out to stalk the Hartebeestes, I left my two gun-

bearers sitting on the steep slope on the other side of the stream,

and myself descended to cross the stream and stalk my way up the

opposite slope within easy view of themen. What first drew their

attention to the Lion was his moving his head as I passed him all

intent on my stalk, looking neither to my right nor left

According to my men, I passed within 20 paces of him, to wind-

ward, nothing whatever intervening between us but the bare

fire-swept ground. He did not catch my eye as he lay in a black

patch of shadow—so black that when the man afterwards pointed

him out to me fromabove and not in relief, I could not make him

out. As I passed, he lay placidly where he was, merely raising

and lowering his head—like a dog winding game—as he winded

me. Had my eye happened to catch his, he would have behaved

otherwise, no doubt ; he would either have made a demonstration

to put me to flight, or have retreated under protest, grunting, as

Lions usually do.

Another popular idea of the Lion is that he becomes a man-eater

only in extreme old age from force of circumstances. I do not

agree with this at all. I believe rather that he learns this in the

first instance more often under the impulse of hunger or passing

caprice than of failing strength, and having thus overcome the

natural repugnance and instinctive fear which all the lower

animals have for man, finds in him an easy victim and henceforth

constitutes him his special prey.

Tn evolving this habit Lions also evolve extraordinary cunning

not primarily their own. In man they recognise a creature of

higher intelligence than theirs, and pit themselves to meet this.

What they seem to become aware of is that, if they are to over-

come man, they must take him at a complete disadvantage—that

unless they do so he is their master against whom mere force as

applied to other creatures will not prevail.

From contact with man they become extraordinarily conversant

with his habits, using this knowledge against him. Nevertheless,

hough they become extraordinarily cunning in plans of attack,

1904. | FIELD NATURAL HISTORY OF THE LION. 267

they also become abnormally cowardly should such plans fail. I

have known remarkable instances of this.

Another point on which I cannot endorse the general theory is

that of the Lion being a fastidious feeder, eating almost exclusively

his own kills, his prey being the larger mammals—buftalo, zebra,

and antelope. I have not found this to be so as regards his being

a fastidious feeder. In my experience Lions feed freely on carrion

—often far gone in putrefaction. Sometimes also they prey on

such very small game as the smaller rodents.

Mr. Selous has it on record how on one occasion a Lion ate the

skin of a Sable Antelope treated with arsenical soap for pre-

servation as a natural-history specimen. Nothing so remarkable

as this has occurred within my personal knowledge. It is probably

an almost isolated case. As an instance of the Lion’s primary

fear of man and also of his eating carrion, | remember a case to

illustrate this which occurred to me on the western shores of Lake

Nyasa in 1885. I had shot two Elephants in the afternoon, and

after following up the herd with no further results than to have to

shoot a calf practically in self-defence, I returned some time after

dusk to where I had killed my first Hlephant, and there slept under

the belly and between the legs of the animal to be in readiness to

cut out the tusks in the morning. In those days, in that country,

Lions gave little anxiety on the score of being man-eaters; there-

fore no precautions were taken to guard against them, either in

keeping watch or burning large fires. Had I had some of my

experiences of later years I could never have slept as I did then—

lying down anywhere and never keeping a watch or burning a fire

except to cook. Nothing occurred during this particular night to

disturb my rest. It was a surprise, however, when morning light

came, to find on the loose soft ashes of the grass burned the day

before that a troop of Lions had circled round, desirous of feeding

on the carcase, but deterred by the presence of man.

The following night, after the tusks had been cut out and I had

moved camp to a point about a mile away, they returned and fed

on what remained of the flesh, then putrid from exposure to

the sun.

On the Lower Shiri Plains, British Central Africa, in 1885, I

shot a very fine Lion whose stomach was full of Hlephant’s trunk

an Hlephant killed by natives and cut up by them, the knife-

cuts in the flesh and hide being at once noticeable. He had made

a huge and rapid meal—chunks of solid flesh, with pieces of the

hide attached, weighing several lbs. each, had been bolted whole.

He was a very fine Lion, in splendid condition and in the prime

of life, as could be seen from the skull.

As regards Lions preying on other game than large mammals, I

have already mentioned to this Society (P. Z. 8. 1904, vol. 11.

p. 144) an interesting case of a Lion which I believe to have

preyed on porcupines.

In ‘Mammals, Living and Extinct,’ Sir William Flower re-

rarked that probably Lions paired for life: this is so, I think.

268 SIR C, ELIOT ON NUDIBRANCHS [ Nov. 29,

What induces me to this belief is the inconsolable behaviour of the

remaining one of a pair should the other be killed, no matter

whether the survivor be the male orfemale. It is really touching.

I shall never forget the moaning sobs of the mate of the Lion

killed N.W. of Kibwezi during the entire succeeding night, nor

the continuous melancholy roaring of the mate of the “Lioness

killed in Henga in December 1893.

2. On some Nudibranchs from Hast Africa and Zanzibar.

Part VI. By Sir C. Extot, K.C.M.G., late H.M. Com-

missioner for the Hast African Protectorate, F.Z.8.

[Received October 6, 1904. ]

(Plates XVI. & XVII.*)

This paper contains an account of the following Nudibranchs

collected in Zanzibar or East Africa :—

1. Orodoris striata, sp. nD.

. Hexabranchus lacer Cuv., varieties faustus, margmatus,

and moebw.

. Doridopsis tuberculosa (Q. & G.).

. spiculata Bgh.

. pudibunda Bgh.

. nigra (Stimpson).

. denisoni (Angas).

. clavulata A. & Hi.

. rubra (Kelaart).

10. Phyllidia varicosa Lamarck.

11. Ph. nobilis Bgh.

As » var. rotunda, nov.

13. Ph. pustulosa (Cuv.).

14. Phyllidiopsis cardinalis Bgh.

15. Doto africana, sp. n.

16. Fiona? pinnata (Eschsch.).

17. Hervia lineata, sp. n.

18. Phidiana tenuis, sp. n.

19. Facelina lineata, sp. n.

20. Phyllodesmium hyalinwm Khr.

21. Stiliger varians, sp. n.

22. St. wregularis, sp. n.

23. Phyllobranchus prasinus Bgh.

24. Cyerce elegans Bgh.

25. Placobranchus ocellatus Van Hass.

26. Hlysia faustula Beh.

27. HL. marginata Pse.

28. #. dubia, sp. n.

It is very likely that some of the smaller forms are immature,

OOM TE

SISISISISIS)

* For explanation of the Plates, see p. 297.

PZ.S.1904.vol Il. PLXVI.

——

eet

C.Crossland, del

Huth, sc.et imp.

2.6. HERVIA LINEATA.

IIE DIVA INF OU WIL WS ,

It,

AO) ) EVAN CBE INVAY Te INSP eASINAL

SSOP WIN Gi i IRE WON IRULVAINI 1S),

EBELYSIA MARGINATA.

P.Z8.1904.vol. I. PLXVIL.

WA.

C.Grossland, del. Huth, sc. et amp

9. MADRELLA FERRUGINOSA. 10.H. FACELINA LINEATA

te StiniGhR IRREGULARES, 16.168: PLACOBRANCHUS

ie Bal iyAU WIS 47. Pl Yeo Awe Us TAT US ie. MARGINATA.

1904. ] FROM EAST AFRICA AND ZANZIBAR. 269°

but if they are noticed at all it seems simpler to give them a

specific name.

MADRELLA FERRUGINOSA.

In the plates of Madrella ferruginosa in the first of these papers

(P. Z.8. 1902, vol. ii. pl. vi. fig. 6) the central cusp of the median

tooth is represented as much too blunt, and I now give a new figure

of it (of. Pl. XVII. fig. 9). It is really long and rather pointed.

This character is quite clear in the hinder part of the vadula, but

in the front part it would appear that the end of the central cusp.

becomes broken or worn off and the point looks blunt as in the

first figure above referred to.

ORODORIS.

| Bergh, Jour. d. Mus. Godeffroy, Heft viii. 1875, pp. 67-71. ]

ORODORIS STRIATA, Sp. n.

One specimen from Pemba found on the shore crawling among

Ulva. ‘The description of the living animal is as follows :—‘“ Six

inches long. The ground-colour of the back is greenish centrally

and deep green and chocolate laterally. The mantle-edge has a

wide border half an inch wide. The most characteristic external

feature is the presence of numerous prominent, narrow ridges

running over the back. The back also bears fairly large tubercles

over which these ridges continue. There are three circular areas

on each side of a deep green colour where the ridges are absent.

The ridges are usually white, but in places are tinged with

greenish grey. The rhinophores are vertical, the pockets a little

raised at the edges. The gills are 8, fairly large, having a fluffy

appearance, but not very sensitive. The anal papilla and main

rhachis of the gills are pink. The secondary branches are light

brown and the smaller branches white. The gill-pocket is

irregularly lobed. The mantle-edge is soft but stiff in texture,

and may assume a very wavy outline. The under side of the

animal is white, with a narrow, irregular, brownish line near the

junction of the mantle and the foot.”

The preserved specimen has suffered severely from contraction,

and most of the internal organs except the buccal mass have been

lost through a rent in the side. The actual length and breadth

are 71 and 59 mm., but could be increased by at least a centi-

metre each if the animal were straightened out. The unusually

strong and fleshy mantle-edge is 9 mm. thick. The colour is a

uniform yellowish white. The characteristic curved ridges are

still plainly visible, but the tubercles are somewhat obscured ;

there seem to be four between the rhinophores and branchiz and

one behind the branchiz : two can be distinguished on each side.

The pockets of the rhinophores are 4 mm. high. The gill-pocket

is raised and bears 8 irregular lobes. The 8 branchie are strong,

with broad stems, and mostly quadripinnate. The anal papilla is

large, crenulate, and connected with one of the anterior gills by a

270 SIR C. ELIOT ON NUDIBRANCHS [ Nov. 29,

lamina. The foot is strongly grooved in front, but not notched.

On each side of the mouth is a large but not very distinctly

shaped lump, which seems to represent a retractile and furrowed

tentacle.

The labial armature consists of a dense mass of light yellow

bent rods, not bifid, exactly like those in Bergh’s plates of

O. miamirana. ‘The radula is large, broad and tough, yellow

behind, dark brown in front; it consists of 120 rows, the widest

of which have at least 130 teeth on each side of the rhachis. No

rhachidian thickenings are visible, but otherwise the radula closely

resembles that of O. miamirana. The first teeth bear two strong

denticles on each side of the central cusp; the next 10-15 are

denticulate on the outer side only, bearing as many as 10 denticles.

The remainder are simply hamate. The outermost are smaller

and rather irregular, but not denticulate.

The specimen is clearly an Orodoris and closely allied to

O. miamirana, particularly the specimen described by Bergh

(J. c. p. 71) as coming from Zamboanga. It is, however, super-

ficially extremely unlike the preserved specimen of O. mianvirana

given me by Dr. Willey, and I hesitate to regard it as a mere

variety. The main difference is that whereas in O. miamirana

there are ridges on the back formed by compound tubercles, there

are in this form a large number of narrow longitudinal as well as

transverse ridges which pass over the tubercles as well as the

general surface of the back.

HeExasrancuus Khr.

[See (1) Bergh, 8. R. Hefte xiii., xvi., and Supplem.-Heft i.

(2) Eliot in Gardiner’s Fauna and Geog. of Maldive and Laccadive

Archipelagoes, vol. i1. part 1(1903). (3) Bergh im Schauinsland’s

‘Reise nach dem Pacific: Die Opistobranchier. |

The Hexabranchide, which are very common in the Indo-

Pacific, but not recorded from other seas, are large doridiform

animals of brilliant coloration and active movements. ‘They differ

from the eryptobranchiate Dorids chiefly in having no branchial

pocket but a circle of separate branchial tufts, each of which con-

tracts when touched into a temporary hollow which forms at its

base. The texture is soft and smooth, the shape flattish, the

mantle-margin ample, and the tentacles foliaceous. There is a

strong labial armature, and the radula is similar to that of many

Dorids, consisting of simply hamate teeth with the formula 00.0.0 .

The verge is extremely long and the nervous system much

concentrated.

Bergh observes (/. ¢. (3) p. 225): “‘ Hine Reihe von etwa 20 Arten

ist angegeben welche zum allergréssten Theil doch wohl nur

Varietiiten oder Localformen einer sehr verbreiteten laingst

evwihnten Art sind, des Hex. lacer Cuv.”* My own observa-

tions support this, as far as the three forms here mentioned are

* It seems to me quite clear that Cuvier’s Doris lacera is a Hexabranchus, not a

Doridopsis.

1904. | FROM HAST AFRICA AND ZANZIBAR. 271

coneerned. The shape and colour of the living animals are both

very variable. The former can be altered at will from an almost

circular to a long slug-like shape, in assuming which latter the

mantle-edges are folded over the back. The colour varies even

in the same animal, and individuals kept in captivity become

conspicuously paler and duller in a few hours. As is usually the

case with soft Nudibranchs, the preserved specimens are much

subject to distortion in alcohol, and such characters as the

expanded or contracted state of the branchiz and the flat or arched

shape of the back appear to have no specific importance. 1 think

that A. digitatus described by me (J. c.) is a distinct form, but its

state of preservation is such that it is difficult to say whether it

should be referred to Hexabranchus or to a new allied genus.

None of the other specimens which [ have examined show any

material differences in structure or anatomy. The active

habits of the animals perhaps explain how it is that one varying

form is spread over so large an area.

The numerous specimens which I have collected in Zanzibar

and on the Hast Coast of Africa represent three varieties :—

(1) @. faustus B—This form, which is not very abundant,

seems characterised by its prevailing red coloration and the

absence of any white bands. One specimen was of a dark blood-red

all over, with only a few yellowish markings at the sides of the

visceral mass, but in most cases this dark blood-red is confined to

a fairly broad irregular border. The centre of the back is of a

lighter red, with mottlings of various tints of red and orange.

Between this region and the border the colour is of a dull reddish

grey. The specimens obtained of this variety are of moderate

size, not exceeding 8 centimetres in length.

(2) H. marginatus (Quoy & Gaimard).—This variety, which

is also not very common or very large, is characterised by having

a broad red band round the mantle, divided in the middle by

a white line. In addition to this there is sometimes, but not

always, a white edge to the mantle. The middle of the back is

mottled red and orange.

(3) The third variety, which is also the commonest, is the

animal captured by Nees at Mauritius, and described by Bergh

(S. R. xvi. p. 828 ff.) as H. marginatus (Q. & G.). Though it is

probably not specifically separate from H. mar ginatus, Bergh

seems to have overlooked the fact that it constitutes a variety

quite as distinct as the other so-called species. Its chief charac-

teristic is clearly given by Moebius :—‘ Die obere Seite des

Mantelgebriimes nach innen porzellan weiss, nach aussen mit

breitem rothem Saume von welchem abwechselnd kleinere und

erdssere Bogen nach innen laufen.” Though the difference

between this form and Z. Joustus is really only one of degree,

the brilliant white band between the equally brilliant deep red

border and the variegated central region is most conspicuous, and

the animal looks superficially quite distinct from those described

above. I propose to call it var. moebii.

This variety is very common and grows to a very large size,

272 SIR C. ELIOT ON NUDIBRANCHS [ Nov. 29,,

one specimen being as much as 25 centimetres long and 16:5

broad. It is very active, swims rapidly, and has been found on

the surface of the sea a quarter or half a mile from the shore.

The colour of the dorsal surface within the borders is very

varying and may be bright light-red, orange, yellow, sandy,

almost white, or still more frequently mottled with all these

colours. Sometimes the external red border is divided into two

parts by a lighter line as in the last variety. The branchie

exhibit somewhat similar variations of colour, but are generally

of a pale reddish yellow with darker lines on the axes and

white tips to the pinne. Sometimes the main axes are bright

light green.

T have been unable to find any differences of structure between

these three varieties: faustus, marginatus, and moebii. In all

the radula consists of from 30-45 rows of simply hamate teeth,

rather slender, 65-80 on each side of the naked rhachis. The

branchiz are from six to eight, seven being perhaps the commonest

number. Hach so-called branchia consists, as a rule, of four

plumes (but sometimes of three or five) inserted very close

together but not springing from a common stem. Not unfre-

quently one plume is separated a little from the others, and the

individual then appears to have an abnormally large number of

branchie.

DortpDopsis.

This genus is distinguished from the other cryptobranchiate

Dorids by having a suctorial buccal apparatus, with no jaws or

radula. The mouth is a fine pore situated in the anterior part of

the foot ; and the internal organs consist of a buccal cone from

which issues a long tube which is generally twisted and expands

into a dilatation before entering the liver. Beneath the anterior

part of the tube is a large folliculate mouth-gland, generally

double. The true salivary glands appear to be represented by

two nodules at the commencement of the dilatation. The nervous

system is very concentrated. The liver is bifid behind. There is

an armature of minute hooks on the spermatic duct and glans.

On the upper wall of the pericardium are a number of lamelle,

sometimes called the pericardial gill. The animals are generally

soft, more rarely spiculous, either smooth or tuberculate. The

branchiz are rather large and few (rarely more than 8), and

though they are completely retractile, are very commonly ex-

serted in preserved specimens so as to appear at first sight non-

retractile. This feature seems to depend on some peculiarity of

texture, and not on any difference of structure.

The genus is very abundant in the Indo-Pacific, and about

60 species have been described, many of doubtful validity. Even

in dealing with the forms which are adequately described, it is

not easy to draw the line between species and varieties, as nearly

all are very variable both in shape and colour, and the internal

organs present few features which can be safely used for classi-

1904. | FROM EAST AFRICA AND ZANZIBAR. (273.

fication. It is noticeable that none of the species here mentioned

are new, although in other genera novelties have been abundant

on these coasts.

DoridoPsis TUBERCULOSA.

{Bergh, Jour. Mus. Godeffroy, Heft xiv. 1878, pp. 38-40, and

nS. R. xvi. 2, pp. 845-848. ]

Several specimens from the East and West Coasts of Zanzibar-

The following are the notes on one of the living animals :—‘* Gill-

pocket shaped like that of Asteronotus, 5-lipped. Gills five, quadri-

pinnate, large, and brown. Rhinophores bent backwards, pockets.

raised at edges. Body soft but firm; grey with black patches; back

covered with large compound tubercles. Under side of mantle

and sides of foot a dirty brown with white blotches; foot lighter

and without blotches. Free mantle-margin about 20mm. A most

handsome creature.”

The dimensions of the largest alcoholic specimens are, length

97°5, breadth 60°5, height 36 mm. The free mantle-edge is

ample and the foot of moderate size (58 mm. long by 25 broad).

The colour is much as described in the living animals, but in two

of the specimens the tubercles are plentifully besprinkled with

black dots. The whole back is covered with compound tubercles,

which are smaller towards the mantle-edge. The rims of the

rhinophore-pockets are prominent but not conspicuous; the club

of the rhinophores is set at almost right-angles to the thick stalk

and bears 50-60 deep perfoliations. The branchial aperture is

raised and stellate; it can be almost closed by five triangular

lobes which bear tubercles on the outside. The five branchie are

quadripinnate with remarkably stout stems. In one specimen

there are six lobes and six branchie. The anterior part of the

foot is much retracted and distorted in all the specimens, but it

appears to be thickened and notched but not grooved.

The buccal cone is 5°5 mm. long in the largest specimen. From

it issues a long narrow tube (36 mm. by 2 mm.), bent almost into

the shape S on the left side, with a smooth interior, which then

dilates more or less clearly in different specimens into a sausage-

like shape, 25 mm. long by 10 broad. The walls of this dilatation

-are strong and laminated internally. After it the digestive tract

narrows again, and a tube 9 mm. long and 5 broad leads into the

hollow of the liver. This organ is very large (51 mm. long by 38

broad), yellow, and distinctly bilobed behind. The mouth-gland

is large, yellow, and with several lobes. The salivary glands are

small, oval, and set at the top of the sausage-shaped dilatation.

The single blood-gland is elongate and purple. The central

nervous system is as usual in the genus. The pericardial lamine

are distinct and yellowish. The albumen and mucous glands are

very large: the vas deferens is thin, extremely long, and much

convoluted. The verge is armed with numerous colourless spines,

not much bent, rather irregular in ee and rising out us circular

discs: ag

Proc. Zoou, Soc. —1904, Vou. II. No. XVIII. 18

‘274 SIR C. ELIOL ON NUDIBRANCHS [ Nov. 29,

DorIDopsis SPICULATA, B.

[See Bergh, Jour. Mus. Godeff. Heft xiv. p. 37.]

One specimen from Wasin. ‘The notes describe the living

animal as three-quarters of an inch long, high and narrow in

shape. The back was covered with numerous small, fiat-topped

warts and was visibly full of spicules, particularly at the bases of

these warts. The colour was white, except for a row of small

dark grey spots of irregular shape on each side of the visceral

mass and a few other scattered spots. The foot was broad.

The preserved specimen is of a uniform greyish white; the

skin is hard and full of spicules. The length is 9 mm., the

breadth 4, and the height 2°5. The mantle-edge is narrow. The

back is covered with flat warts, bearing smaller tubercles, and

there are also a few simple papille. The branchial opening is not

much raised and slightly crenulate. The branchiz are six, tri-

pinnate, and set in an incomplete circle which is open behind.

The foot is pointed at both ends, with uncertain traces of a notch

and groove in front. The tentacles are small.

The internal organs are mostly yellowish and present nothing

remarkable. The pericardial lamelle and penial hooks are present

as usual in the genus.

This specimen has the main characters of Bergh’s D. spiculata

from the Philippines. The difference in form is not remarkable,

as the animals of this genus frequently alter their shape from

long and high to flat and broad, but the discrepancy in the number

and arrangement of the branchie throws some doubt on the

identification. Bergh’s specimen had only four.

DoRrIDOPIS PUDIBUNDA, B. 2

[Bergh in 8. R. xvi. 2, pp. 844-5. Cf. id. Jour. des Mus.

Godeffroy, Heft xiv. 1878, pp. 33-4.]

Four specimens from Chuaka. ‘The ground-colour of the livin

animals was whitish, but almost hidden by two sets of blotches,

one of which varied from light reddish brown to deep dark brown,

and the other from pale blue to an inky colour. The intensity of

coloration varied in different specimens, and was also greater in

the middle of the back than on the mantle-edges. The mantle-

edge changed considerably in shape and size with the movements

of the animal, The branchize were lined with dark brown and

not very sensitive; the rhinophores dark brown and tipped with

white,

The preserved specimens are stout and high in shape, and have

yetained their colour fairly well. The largest is 30 mm. long,

17 high, and 22 broad. The mantle-edge is crinkled and allows

the sides of the foot to be seen, In one specimen it is much wider

than in the others, and this specimen is flatter than the others. The

back is smooth, arched, and appears to be swollen. The pockets

of the rhinophores and branchiz are slightly raised and smooth.

The rhinophores are somewhat bent backwards and bear about

1904.] . FROM FAST AFRICA AND ZANZIBAR. 275

25 perfoliations. The branchie are six in number, quadripimnate,

low but strong and bushy. The circuit is open behind, and the

subcentral anal papilla very large. The foot is notched in front,

but not grooved; the tentacles are small and adherent. The

integuments of the body are thick and tough.

From the small reddish buccal cone issues a long thin tube, which

is twisted into a shape somewhat like § on the right-hand side.

The mouth-gland is bilobed. The pericardial lamellze are yellowish

and not very distinct. Bergh says, ‘‘ Im Penis wurde eine Haken-

Bewaffnung nicht nachgewiesen ”; and I could find none in two

specimens. In the third there were clearly visible about 12 rather

irregular rows of thick-set, transparent, hooked spines.

These specimens seem referable to the forms described by Bergh

in the passages referred to, though the descriptions are not alto-

gether clear. In the specimens from the Philippine Islands the

back was “aufgedunsen knotig”; in that from Mauritius, “die

Form-Verhaltnissen die gewohnlchen.”

DorIDOPSIS NIGRA (Stimpson) A. & H.

[A. & H., Coll. of Nudibr. Molluscs made in India, p. 128;

Bergh, 8S. R. xvi. p. 842, xvii. p. 963; id. Danish Exped. to Siam,

p. 191; id. Beitr. zur Kennt. Japan. Nudibr. p. 181.]

Forms which seem referable to this species are among the

commonest Nudibranchs on the coasts of both Zanzibar and the

mainland. The species, as well known, is extremely variable

not only in its colour, but also in its shape, in the number and

form of the branchie, the presence or absence of tubercles, and

the configuration of the anterior foot and tentacles, though with

regard to the last point it must be remembered that this is the

part of the body which is most liable to be obscured and distorted

by alcohol. The internal structure of all the varieties seems much

the same, and does not present any very remarkable features.

The organs are mostly yellowish. The tube which issues from the

buccal cone is at first narrow (generally about 1 mm.) and bent

considerably, usually on the left-hand side; it then dilates into a

sausage-shaped expansion of about double the width, after which

it enters the liver. The mouth-gland is distinct and bilobed.

The pericardial lamelle ave well developed. The verge is armed

with spines of irregular shape.

The chief varieties are as follows (the texture is in all cases

very soft) :—

(1) Jet-black, without any spots or markings, but white tips to

the rhinophores. This form does not appear to be common; I

have only two specimens from Mombasa and two from Zanzibar.

The animals are not very large (about 20 mm. long and 10 broad)

and flattish. The branchie are six. In three specimens the

anterior margin of the foot appears to be entire and the tentacles

are very indistinct. In the fourth both the tentacles and the

groove are clear.

(2) Jet-black, with a brick-red band running round the body,

276 SIR C. ELIOT ON NUDIBRANCHS [Nov. 29,

‘ but wider anteriorly than posteriorly and visible from the under

side of the mantle. Rhinophores black with white tips. Foot

broad, grey. Branchie six. Only ones specimen.

(3) Brownish, with a red line much as in the last variety, but

occasionally broken and accompanied by red spots at the sides.

In the centre of the back are some clusters of whitish spots, and

there is a red band round the foot. The genital papilla is yellow

and conspicuous. Branchie six. The foot is narrow, pointed at

both ends, and grooved anteriorly. No tentacles are visible.

Only one specimen. Length 24°5 mm., breadth 8°5, height 5.

(4) The commonest variety of all is of a greenish black, of

varying intensity, with white spots. A. & H.’s figure No. 13

gives a good idea of an average specimen. The animals are about

30 mm. long and 10 broad, and somewhat globular, though, like

most Doridopsids, they can alter their shape. The branchie vary

from 6 to 12 in number and are usually ample and fluffy. Asa

rule, the foot is plainly grooved, and the small flattish tentacles

are distinct. In coloration there are many subvarieties. Generally

there are both clusters of small white spots and scattered larger

white spots. In the specimens where the pattern is most deve-

loped the clusters are arranged in two symmetrical lines down

the back, and the scattered spots are more numerous round the

margin, where they tend to form a border. Irregular black

blotches may be present or absent. But in some specimens the

markings are much reduced, and in one there is only a single

white spot.

(5) A variety with raised tubercular spots is fairly common.

The specimen which shows this feature best is flat and elongate.

Down the centre of the back is an elaborate pattern consisting of

clusters of white spots in two rows and scattered white spots

between them, and round the mantle-margin is a ring of elongate

white spots forming an interrupted border. The spots, especially

those near the edge of the mantle, are distinctly raised and

tubercular. The branchie are fully exposed, 10 in number, and

somewhat stiff and meagre. The anterior margin of the foot is

grooved and the tentacles are distinct. If this specimen were

isolated, I should certainly regard it as specifically distinct, but a

number of intermediate forms seem to show that it passes imper-

ceptibly into variety 4, as regards both the tubercular spots and

the branchiz.

(6) I have once found a specimen of a uniform greyish white,

with no markings and six branchie.

Tam indebted to Mr. F. W. Townsend, of the Indo-European

Telegraph Service, for two specimens from Karachi, which may be

referable to this species, According to his notes and a rough

pencil-sketch, the animal was much broader behind: than before,

of a very deep purple, with a bright crimson line round the

undulated mantle-edge: The branchie were the same colour 2 as

the body and fern- like.

The two -preserved specimens: retain the Goin fairly yell

1904.) FROM EAST AFRICA AND ZANZIBAR. ete

and are hard and stiff. The largest is 26 mm. long and 12 high, |

7 broad. as measured across the rhinophores, and 15 across the '

branchie. The integuments are thick and tough, the intestines

bright yellow, and as usual in the genus. The branchie 8; the |

anal papilla not in the centre but somewhat to the right. ;

These specimens do not look like D. nigra as preserved, and

apparently did not look like it when alive, since they were kept

separate from normal examples of the species, but it is difficult to

formulate any distinguishing characteristics except the stiffness

and thickness of the integuments.

Doripopsis DENISONI (Angas).

D. gemmacea A. & H.

[Angas, Descr. d’espéces nouvelles de Moll. Nudibr., Journal

de Conchy]l. sér. 3, iv. 1. 1864, p. 45; A.& H., Coll. of Nud. Moll.

made in India, p. 126; Bergh, 8. R. xv. p. 694 ff.]

One small specimen from near Wasin, East Africa.

The living animal was long, narrow, and high; the mantle-

edge, though not wide, descended straight down to the ground on

each side, so that the sides of the foot were hidden by it. The

central part of the back was flat and smooth, but bounded on

each side by three large tubercles. Two more stand in the median

dorsal line immediately before and behind the rhinophores. A

few more tubercles are scattered here and there near the sides.

In the centre of the back were two large eye-like spots (similar to

those found in some species of Votarchus), bright blue with rims

of dull dark yellow, and on each side were five double spots (each

like the figure 8) of similar colour. The tubercles were tipped

with brown, underneath which was a band of yellowish white. The

ground-colour of the back and of the bases of the tubercles was

reddish brown with clear lines of yellowish white. On the mantle

was a border formed by alternate blotches of pale yellow and

light aimson. The under surface was of a beautiful pink, deeper

towards the mantle-edge, but without markings. Theedge of the -

gill-pocket was slightly toothed. The gills 5, tripimnate, but not

ample, white, with dark brown lines on the rhachis. The animal

was flexible and lively in its movements; the tops of the tubercles

were often drawn in and then thrust out again. The beating of

the heart was distinctly visible under the skin.

The preserved specimen is 7 mm. long, 3 broad, and 2°5 high.

The mantle-edge is narrow. The lumps on the back remain fairly

distinct. The rhinophores are large, the tentacles small. The

five gills are erect and strong, but not very ample. The internal

organs are yellowish, and seemed to be as usual in the genus.

Pericardial lamellz were not discovered, but this was probably

due to the small size of the specimen.

Angas’s D. denisoni is identified with A. & Hs D. gemmacea

by Bergh, in spite of considerable differences of colour, and

D. gemmacea is said to vary a little in colour and markings. The

present specimen is clearly allied to these forms, and as, in view

278 ‘gIR C. ELIO! ON NUDIBRANCHS [ Nov. 29,

of its small size, it is probably immature, the difference in pattern

is perhaps not specific. The chief distinction is that here we

have yellow ocelli with a blue centre, whereas in A. & H.’s

specimens there were “lozenge-shaped areas of a rich brown

colour, with a few brilliant blue spots in each.”

DORIDOPSIS CLAVULATA.

[A. & H., Notice of a Coll. of Nudibr. Molluscs made in India,

p- 127.]

One specimen dredged between Wasin and the mainland in

10 fathoms.

The following are the notes on the living animal :—“ Very like

D. denisoni. Probably a distinct species, but nearly related.

Shape as D. denisoni, but even longer. The tubercles are not so

large or so definite, and the back is rounded. There are two

small tubercles between the gills and rhinophores, and the rest of

the back is fairly thickly covered with smooth warts. The mantle

has a border formed by blotches of greenish black, white, and

yellowish brown. The whitish patches extend inwards, and there

are three others in the middle of the back. On each side are

three smooth irregular patches of yellowish brown, with scattered

slit-like marks of bright blue. Apart from these various markings,

the ground-colour is a dull red. The tops of the tubercles are

greyish, but this does not altogether hide the underlying red or

white. The edges of the rhinophore and gill-pockets are slightly

raised, and the latter is wavy. Gills 5, large, feathery, tripinnate,

white, with black lines on the rhachis. Rhinophores dark brown,

tipped with white, not bent back. Anal papilla large, and keeps

up a motion like the beating of a heart. Foot pinkish, with a

yellow border, only slightly projecting behind mantle. The animal

is infested with numerous white copepoda, especially about the

ills.”

: The alcoholic specimen is very soft and much bent. It would

probably be at least 25 mm. long if straightened out; width

13 mm., height 12. The mantle not ample but reaching to the

ground. The rhinophore-pockets bear one or two tubercles. The

gill-pocket is indistinctly five-lipped and also irregularly denti-

culate. There are indications of three or four large tubercles on

each side of the back twice as large as the rest, although the

description of the living animal does not notice them. The foot

is deeply notched in front, but not distinctly grooved. The ten-

tacles are thin but distinct. The internal organs are yellow and

as usual in the genus. The mouth-gland falls into two halves,

each with several lobes.

I think that this may be identified with A. & H.’s D. clavulata,

which may perhaps be shown by the discovery of intermediate

forms to be the same as D. denisont. D. nicobarica B. appears to

be akin:

1904. ] FROM EAST AFRICA AND ZANZIBAR. 279

Dorivopsis ruBRA (Kelaart) A. & H.

[A. & H., Notice of a Coll. of Nudib. Moll. made in India, p. 126;

Bergh, Danish Exped. to Siam, Opisthobranchs, pp. 190-1. ]

This form is common both in Zanzibar and on the mainland.

The notes on the living animal describe it as a “‘ Large bright red

Doridopsis, between crimsou-lake and vermilion : the back above

the viscera blotched with small irregular spots of chocolate-brown.

Length two and a half inches, breadth at most two inches; texture

of skin smooth and shiny; body very contractile. Gills 6,

feathery, tripinnate. Gills and rhinophores rapidly and completely

retractile.”

Six specimens are preserved, all much contracted and blistered.

Two are uniformly white; in the rest the dark mottlings remain.

The skin is soft and smooth; the mantle-edge fairly ample; the

foot fairly wide and slightly pointed both before and behind. The

rhinophores are set very far in front and the branchie very far

behind. The pockets of both have slightly raised smooth margins.

The branchie are tripinnate, generally exposed, fairly luxuriant,

and apparently six in all specimens. No head or tentacles are

visible in any of the specimens, but while in the two white ones.

the anterior margin of the foot appears to be simple, it is distinctly

grooved in the others.

The internal organsare mostly of a reddish yellow and arranged

as usual in the genus. The mouth-gland is bilobed. From the

buccal cone issues a long thin tube about 1 mm. broad, which

is generally curved into the shape of §, and then dilates into a

wider portion, about 2 mm. broad, which is constricted before it:

enters the liver. The mouth-gland is bilobed, the eyes large and

distinct. The verge and part of the seminal duct are armed with

small hooks of a rather irregular shape.

The difference of colour in the preserved specimens, coinciding

as it does with a somewhat different shape of the anterior pedal

margin, suggests that the specimens may really belong to two

species, although the notes do not indicate any difference in the

appearance of the living animals. Possibly D. rubra and D. brockia,

as Bergh suggests, are merely varieties of a very variable form.

PHYLLIDIADA.

[See Bergh, ‘ Bidr. til en Monogr. af Phyllidierne,” Naturh,

Tidssk. 3 R. v. 1869; id. Neue Beitr. zur Kenntniss d. Phylli-

diaden, 1876; id. in 8. R. xvi., xvii, Eliot, Nudibranchiata in

Gardiner’s Fauna of Maldives and Laccadives, p. 560 ff.]

The structure of these well-known and unmistakable animals

has been so thoroughly examined by Bergh that it need not be

described here. Five genera have been proposed: Phyllidia

(Cuv.) B., Phyllidiella B., Fryeria Gray, Phyllidiopsis B., and

Ceratophyllidia Eliot. The last three genera have all decided

characters. In /ryeria the vent is terminal and not dorsal; in

280 SIR C, ELIOT ON NUDIBRANCHS * > { Nov. 29,

Phyllidiopsis and Ceratophyllidi« the mouth-parts are much as

in Doridopsis, the glands not being fused with the buccal tube,

and Ceratophyllidia has the additional peculiarity of bearing

stalked globes on the back. The distinction between Phyllidia

and Phyllidiella seems to me less certain. According to Bergh,

(a) the oral tube is symmetrical in Phyllidia, asymmetrical in

Phyllidiella; but I have not found the difference to be clear or

persistent, and even if it is so, I doubt if it is of generic worth.

(0) In Phyllidia, “ Dorsum tuberculis elongatis, plus minusve con-

fluentibus obsitum, medio varicositates longitudinales formant-

ibus.” In Phyllidiella, “* Dorsum proprium tuberculis discretis

vel pro parte confluentibus quincunces formantibus obsitum.”

Even in typical forms it does not appear that this distinction is

clear. Phyllidia elegans (see Bergh’s Monograph, pl. xix. fig. 1)

seems to me to have not “ varicositates longitudinales,” but

groups of confluent tubercles; and, on the other hand, Phyllidiella

pustulosa strikes meas having not so much tubercles arranged in

‘“quincunces,” as compound tubercles arranged in lines. But

in abnormal forms, which are frequent, it is still harder to draw

the distinction.. I have a fine specimen with all the characters of

Phyllidia varicosa, but the median ridges, though very distinct

posteriorly, are broken up in front and unite to form a quincunx

as in Phyllidiella nobilis. Again, in this latter, the quincunces

are often placed so regularly above one another that the tubercles

seem to be arranged in longitudinal lines and not in figures.

1 therefore think it better to abandon Phyllidiella as a separate

genus.

The arrangement of the dorsal tubercles in these forms is so

variable, that it is hard to draw the line between species and

varieties, but at least three certainly specific forms occur in East

Africa, :—

' (1) Ph. varicosa.—Colour black, blue, and orange ; rhinophores

yellow. A black stripe on the foot. Tubercles more or less

fused into ridges, but not compound. Typically, there are three

long ridges down the centre of the back and a number of short

ridges, more or less at right angles to them, running inwards from

the mantle-edge. Buccal mass yellow and very complicated.

(2) Ph. pustulosa—Black and green; rhinophores black, no

black line on foot. Tubercles simple or composed of only two or

three lumps. Buccal bulb partly black.

» (3) Ph. nobdilis—Colour as in Ph. pustulosa, but tubercles

highly compound, sometimes consisting of ten small lumps fused

together. Typically, they are arranged in square or oblong figure

Buccal bulb large, yellow.

That the patterns formed by the tubercles should vary is not

surprising, if we consider that the tubercles have always a

tendency to unite, and may do so more or less decidedly in a

given direction. Thus the typical form of Ph. varicosa occurs

when the lateral tubercles unite in a predominantly transverse

direction. "When however, union in a longitudinal direction

1904. ] FROM EAST AFRICA AND ZANZIBAR. 281

predominates, a variety is formed which seems to have five or

seven longitudinal rows.

As I have mentioned elsewhere, my observations do not confirm

the statements of the older naturalists as to the ‘torpidity and

immobility of the Phyllidiade, which seem in this respect much

like the average Dorid. Ph. varicosa is the most active and

crawls quite rapidly ; Ph. nobilis when in captivity crawled con-

tinually but slowly ; Phyllidiopsis cardinalis was sluggish but not

motionless.

The Phyllidiade are common in the Indo-Pacific, but appear

not to be littoral. They are generally found in a few fathoms or

in spring-tides at extreme low water.

PHYLLIDIA VARICOSA Lam.

[Bergh, Bidr. til en Monogr. p. 500 f.]

This large, handsome animal is common on the coasts of

Zanzibar and the mainland of East Africa. My largest specimen

is 73 mm. long and 32 broad, and specimens measuring 50 or

60 mm. are not infrequent. The colour in life is glossy black,

with slate-blue ridges on which are bright orange-coloured pro-

jections. The rhinophores are bright light yellow. The tubercles

are more or less confluent. Down the middle of the back run

three ridges bearing 12-16 tubercles. From the sides of the

mantle there run inwards about 30 ridges, less uniformly con-

tinuous than those in the centre and bearing each two or three

tubercles. The openings for the rhinophores and anal papilla are

small and placed in tubercles, not on the smooth surface of the

back. The tentacles are digitiform and yellowish. The buccal

mass is large, yellow, and, as Bergh says, ‘‘ magnopere compositus.”

The foot is broad, and bears in the middle a distinct black line

1-5 mm. wide in large specimens. There are sometimes black

mottlings at the side of the line. Variations from the typical

form are frequent. In one specimen the three longitudinal

ridges are all fused together to form a central dorsal prominence,

whose tripartite nature is only obscurely visible. In another the

three unite in the posterior third of the body, though before it

they are separate. In one very fine specimen the three ridges are

very distinct behind, but in the anterior third of the body forma

group of tubercles like those found in Ph. nobilis.

The most distinct variety, however, is one which perhaps

corresponds to Bergh’s Ph. fasciolata*, which also comes from

East Africa (Comoro Islands), and which he appears to regard as

not specifically distinct. It is characterised by having from five

to seven ridges on the back, in which the tubercles are more

distinct and the connecting-lines less developed than in the

typical form. The lateral ridgesare almost entirely absent. The

rhinophores vary from yellow to grey. The bulbus pharyngeus

* Bidr. tilen Mon. af. Phyllidierne, p.507. “ Ph. varicose et eleganti forma et

charactere dorsi affinis, sed rhinophoriis nigerrimis, varicositatibus dorsalibus (7) sat

tuberculosis.”

282 .SIR C, ELIOT ON NUDIBRANCHS [ Nov. 29,

is as in the typical PA. varicosa. This variety when preserved

sometimes superficially resembles Ph. pustulosa, but I think it

should be referred to Ph. varicosa because (1) there are inter-

mediate forms; (2) the bulbus pharyngeus is not black; (3) the

foot is marked with a broad black line.

Pay.irpia (PHYLLIDIELLA) NoBILIs B. (Plate XVI. fig. 1.)

[Bergh, Bidr. til en Monogr. p. 512 ff.; & id. 8. R. xvi. 2,

p. 860 ff. |

This species is common, but does not attain a large size, the

finest specimen being 36°5 mm. long by 18 broad. The back is

black, with sea-green tubercles. The under surface and branchiz

are greenish grey, and there is a yellowish border round the foot.

The rhinophores are black. The patterns on the back vary con-

siderably, but the ground-plan appears to be in all cases two or

three borders running round the mantle and a series of oblong

figures in the centre. The tubercles are mostly compound, and

sometimes consist of as many as ten small lumps fused together.

In one of the most regular specimens there is first a green border

showing hardly any traces of tubercles (this feature appears

invariable), then a circular band of narrow tubercles, then a

similar band of much broader and more composite tubercles.

Down the centre of the back are arranged four oblong figures ;

the sides of each are composed of two tubercles and the top and

bottom of one; within the area are two tubercles. All these

prominences are compound, but.a number of little simple tubercles

are scattered here and there. This arrangement varies consider-

ably owing to the borders and figures running into one another.

Tn other cases the central figures are placed accurately one above

the other, and the tubercles then appear to be arranged in straight

lines. Bergh’s uncoloured plates (S. R., Heft xvi. 2, pl. boxxiv.

figs. 11 & 18) give a good idea of the animal, but in my specimens

the borders are more distinct and circular. The drawing in

Plate XVI. fig. 1, by Mr. Crossland, is a fair representation of

the average Hast-African specimens, though a little less regular

than the one described above. It fails, however, to indicate that

all the larger tubercles are compound. In many specimens the

green parts have a tendency to coalesce and form blotches, but T

have never seen this so highly developed as in the animal figured

in Bergh’s ‘ Opisthobranchs of the Danish Expedition to Siam ’

(plate ii. fig. 15).

The openings for the rhinophores and anal papilla are very

small and situated in or at the edge of tubercles. The buccal

mass is very large and, like the rest of the intestines, yellow, but

in some specimens has a little black pigment in front.

PHYLLIDIA NOBILIS, var. ROTUNDA.

Two specimens constitute a very distinct variety, possibly

meriting specific rank. From the absence of notes it may,

1904. ] FROM EAST AFRICA AND ZANZIBAR. 283:

perhaps, be concluded that the living animals were black and

green like ordinary individuals. As preserved, the back is black,

with brilliant white tubercles; the under side greyish yellow, the

rhinophores black, and the branchie greenish ; several black bands

run from the mantle- edge to the branchiz on the under side. The

tubercles are compound as in the ordinary form but more pro-

jecting, and show a few black depressions between the confluent

lumps. There are no borders or rings of tubercles round the

mantle, but both in the centre and round the margin the tubercles

are arranged in square or oblong figures. The buccal mass is

large and deep orange. The shape of the whole animal is much

broader than usual, being about 23 mm. long by 18 wide, but the

end of the foot and mantle are pointed.

If this animal proves to be sharply distinguished from

Ph. nobilis, it is no doubt a good species, but it will probably be

found to be connected with the ordinary form bya series of links.

It bears a certain resemblance to Van Hasselt’s figure of his

Ph. verrucosa (v. Bergh, “* Die Van Hasselt’schen Nudibranchien,”

Notes from the Leyden Museum, vol. ix. 1887, p. 313, and

plate 6. fig. 7), but does not coineide in details.

Much the same may be said of another specimen in which the

tubercles were pink rather than green in life. The three borders

and central pattern are very regularly developed as in the typical

form described above, but the raised parts are fused into flat,

smooth ridges, with hardly any indication of knobs.

PHYLLIDIA (PHYLLIDIELLA) PUSTULOSA (Cuv.).

[| Bergh, Bidr. til en Monogr. p. 510 ff. }

I have only two specimens of this form, which would hence

appear not to be very common on the East Coast of Africa. It is

more elongated than Ph. varicosa and nobilis, the larger specimen

measuring 33 mm. in length and 11 in breadth. The colour of

the back is a very deep bright black with green tubercles. The

rhinophores and branchial lamelle are also deep black, but the

sides of the body, the foot, and the oral tentacles are slate-

coloured. Many of the tubercles, especially in the centre of the

back, are compound, and are composed of two or three, rarely

four, partly fused together. In the more regular of the two

specimens there are three borders of tubercles round the mantle-

edge, and three groups, composed of four compound tubercles each,

down the centre of the back. These three groups are separated

from one another by straight transverse rows composed of three

tubercles each. In the other specimens, though the number and

character of the tubercles are nearly the same, the pattern is less

regular, and the general impression given is of five moderately

straight lines down the back. The openings for the rhinophores.

and anal papilla are rather large, and are placed not in tubercles

but on the flat surface of the back. The buccal mass is large ;,

parts of it are yellow, but below and in front it is black.

284 SIR C, ELIOT ON NUDIBRANCHS. | Nov. 29,

PHYLLIDIOPSIS CARDINALIS B.

{Bergh, Neue Beitrige zur Kenntniss der Phyllidiaden,

Verhandl. der k.-k, zool.-bot. Gesell. in Wien, 1876.]

‘Four specimens, one from Zanzibar and three from Wasin, were

obtained of this form, which has hitherto been recorded from

Tonga. One specimen (from Zanzibar) is of considerable size,

being 37 mm. long, 15:5 broad,and 13 high. The others are much

smaller, being about 15 mm. long and 7 broad. The notes on the

living animal say that the larger specimen had the under side,

branchiz, and sides of foot light yellow. On the back were dark

reddish-brown blotches bearing black warts alternating with

sandy blotches bearing sandy warts, irregularly arranged. Rhino-

phores dull green, anal papilla bright yellow. The smaller speci-

mens had a lighter coloration. The following are the notes on

one of them :—‘“ Ground-colour light yellow. On the under side

this is only interrupted by dark green dots along the edge of the

foot, and blotches, which are black at their extreme edge, on the

mantle. They appear dorsally as black blotches. On the sides of

the visceral mass are large black blotches, greenish at the edges,

three on one side and two on the other. In the centre of the

back are three large reddish-brown blotches. The tubercles are

the colour of the blotches on which they occur. Between the red

and black blotches is a coarse, clear network of greenish brown.

The rhinophores are dark bright green and the anal papilla bright

yellow.” In the alcoholic specimens all trace of yellow has dis-

appeared, and the general coloration is a dull purplish red with

indications of black spots. It would appear that the alcohol

liberates the red pigment, which then overpowers the other

colour, a phenomenon which I have observed in some preserved

Chromodorids.

The general aspect of the preserved specimens resembles

Doridopsis, the back being covered with pyramidal compound

tubercles not unlike those of D. tuberculosa, but quite different

from those of Ph. nobilis. In the largest specimen there are

three distinct lines of 8-10 tubercles in the central area, two

other lines less distinct, one on each side, and a number of

irregular tubercles round the mantle-edge, arranged in two or

three lines and decreasing in size outwards. There are also

scattered simple tubercles all over the back. The openings of the

rhinophores are smal] and inconspicuous, that of the anal papilla

large and circular. The branchie are dull red; they run up

nearly to the mouth, and are otherwise only interrupted for about

2, millimetres on the right-hand side by the genital papilla. The

smaller, and doubtless younger, specimens are very like the large

ones, but the tubercles are less developed.

. The buccal parts are much as in Doridopsis. From the mouth

issues a thickish tube with laminated walls inside. It is bent

towards the left and back again, and the central nervous system

lies at its posterior end. It is 2°5 mm. broad and about 4:5

long, but would be considerably longer if straightened out. From

—-1904.] FROM EAST AFRICA AND ZANZIBAR. 985.

this issues a much thinner tube, about 5 mm. long and 1°5 broad,

with muscular walls. After a sharp constriction it continues

again for about 5 mm., and enters the liver rather far back. In

the large specimen the posterior part of the tube is much the

same size as the anterior. In the smaller ones it is considerably

more inflated. A purplish gland lies under the first-mentioned

thick portion of the above tract, but is not fused with it as in the

other Phyllidiadee.

DOTONIDA,

DovTo AFRICANA, Sp. n.

One small specimen found on a Sertularian at Chuaka. The

notes on the living animal are as follows:—‘“‘ Ground-colour of

body grey-black with two white stripes; sides of foot also white.

Rhinophore-sheaths large, also grey-black. Six pairs of cerata, of

which the third is the largest and the fifth and the sixth very small.

Cerata yellow-brown, with dark blue tips to the tubercles.”

The preserved specimen is 3 mm. long, and has retained its

coloration fairly well, though the difference between grey-black

and dark blue is not visible. The cerata are relatively large, the

tallest being nearly 2 mm. high. They are of the shape usual in

the genus. The third pair bears sixteen rounded tubercles, that

18 Some rows of four each; the others have fewer tubercles

according to their size, and the sixth pair are simple warts. The

rhinophore-sheaths are large and stout, not much broader at the

top than at the bottom, the edges of the cavity smooth and not

turned outwards. The rhinophores are completely contracted

within the sheaths. In front of each sheath lies a tubercle or

short ridge, pointing towards the edge of the oral veil, which is

large and circular. The anal papilla is yellow, and lies between

the first and second cerata on the right side.

The delicate and transparent jaws, though hardly visible, appear

to be of the generic type, with smooth edges. The radula consists

of a single row of about 70 teeth, of horseshoe-shape, but some-

what more rectangular than usual. The central cusp is well

developed, and there were faint indications of two or three

denticles on each side of it.

This does not appear to be Bergh’s Doto indica or his Doto sp-

(Mal. Unt. 1894, vi. 1, p. 13), and it seems necessary to create a

new species, though in the case of so small an animal there must

always be some doubt whether it has assumed its mature

and specific form. The most distinct character is the presence of

two tubercles in front of the rhinophores. Cf. the ridges in

D. fragilis and pinnatifida.

AMOLIDIADS.

Fiona? pinnava (Hschscholtz).

[Eschscholtz, Zool. Atlas, 1829, p, 14; Bergh, Journ. Mus.

Godefir. Heft 11. 1873, pp- 87-88; “id. Beitr. z. Kennt. der cols,

diaden, i. p. 605.] - .

One specimen was found on a dead nantine: shell ahi was

286 SIR C. ELIOT ON NUDIBRANCHS | Nov. 29,

floating between Zanzibar and Prison Island. On the shell were

also Clytia and a small species of barnacle. The notes on the

living animal are as follows :—“ Foot colourless and transparent.

Upper surface of body has a yellowish-brown tinge, which is

deeper on the tips of the cerata. ‘The liver-canals appear as dark

greenish brown. The foot projects behind the cerata for some

distance. Cerata numerous; they bear on the inside a wrinkled

branchial membrane.”

The preserved specimen is much bent, and would perhaps

measure 20 mm. if straightened out. It is about 5 mm. broad.

The general shape is much that of Fiona nobilis (=marina), as

figured by Alder and Hancock, and the foot does not project

much behind, as it did in the livmg animal. The bare space

m the centre of the back is not large. On each side is a thick-

set longitudinal row of about 60 or 70 cerata. The transverse

arrangement is irregular, but in a given line there are generally

two or three large cerata, and one or two quite small ones out-

side. The cerata ave somewhat thicker and more inflated than

those in specimens of Fiona nobilis received by me from Naples.

The larger ones bear on the inside a wrinkled branchial membrane,

but this is absent on the smaller ones. The pericardial promi-

nence is not conspicuous. The oral tentacles are some distance

above the mouth, and the rhinophores point sideways. No eyes

are visible. Theanal papilla is latero-dorsal, just inside the cerata

about halfway down the right side.

The radula consists of 33 horseshoe-shaped teeth, with a large

central cusp and six denticles on each side of it. At the base of

these main denticles, or between them, are occasional accessory

denticles. The jaws are yellowish, with a single row of rather

-coarse teeth.

I doubtfully identify this form with /. pinnata, recorded from

the Northern and Central Pacific. In favour of the identification

are the facts that the living animal had a projecting tail of some

length, though this character is not clear in the preserved speci-

men, that only the larger cerata have the branchial membrane,

and that the teeth have six denticles on each side of the central

cusp.

Hervia LINEATA, sp.n. (Plate XVI. figs. 2 & 3.)

‘Two specimens from Prison Island, Zanabar. The following

are the notes on the living animal :—‘‘ General body-colour a

translucent white with a slightly red-brown tinge, which is well

marked on the rhinophores, and yather less so on the anterior

tentacles and on the sides of the narrow groove-like foot. The

body is marked with thin clear lines of opaque white. Cerata in

four or five clumps; they are of a chocolate colour, with longi-

tudinal, fine, clear, opaque white lines. Between the rhinophores

and tentacles are two brilliant vermilion blotches. The foot is

narrow and tapering to a tail.”

The preserved specimens are of a uniform dirty yellow, and

1904. | FROM EAST AFRICA AND ZANZIBAR. 287

the largest measures 8 mm. long by 2 broad. The pericardial

prominence is large. ‘The cerata, which are not at all caducous,

are set on low inconspicuous ridges in groups as follows :-—

Left. vight.

BigSbyOLOU Psi 1-c/.l2.~ 92 8 10

DECOM) Meee esis. i) 4

EPelnrasllpaget ein Here). ataic ee 5) 5 5

LENO) NG) oak ec 5 5

itil Tie seca des enicd 3 3

The smaller specimen has only four groups of cerata, as in the

Plate XVI. The first group is composed differently from the

others, and possibly consists of two fused together. The cerata

are longish, but, as preserved, slightly inflated in the middle. The

innermost and outermost are smaller and the median ones largest,

attaining a length of 2mm. ‘The tentacles are longer than the

rhinophores, which are not perfoliate. The foot is produced

anteriorly into tentacular angles.

The jaw bears 20-27 distinct coarse denticles of very irregular

shape. There is a single series of 18 teeth, of the form usual in

Facelina, with a strong central cusp and 10-11 longish, some-

what curved denticles on either side.

I cannot see what is the difference between the genera Hervia

Bergh (1871) and fizzolia Trinchese (1877), and refer this

specimen to the former, since it has priority. But it might equally

well be referred to Rizzolia.

PHIDIANA TENUIS, Sp. 0.

Two specimens from Wasin Island, British East Africa, dredged

in 10 fathoms. The notes on the living animal are as follows :—

“ About half an inch long and very narrow, vermiform. ‘Tail

long, but bearing cerata to the tip. These are nearly all lost,

but were uniformly vermilion in colour. Foot white, body

pinkish, tentacles and rhinophores white. Foot very narrow

behind, broader in front, where it is bordered by a flap on either

side; grooved in front but not produced into tentacular angles.

Tentacles very long and spreading outwards. Rhinophores very

slightly ringed in the distal halves, which are opaque, the proximal

halves being translucent.” Of a second specimen caught in the

same place next day, it is noted that when it was fir st captured

“the cerata looked vermilion and light violet-blue, but subse-

quently became practically colourless with a bluish bloom. The

vermilion liver does not nearly fill the cerata; it is thin and has

numerous short more or less horizontal branches. In this specimen

the tentacles and rhinophores appear of nearly the same length.”

The preserved specimens are of a uniform dirty yellow, and the

largest is 9 mm. long and 2 broad. Both of them have lost

nearly all them cerata, and the disposition of these organs is no

longer plain, but apparently they were set in five groups, without

288 SIR C. ELIOT ON NUDIBRANCHS [ Nov. 29,

counting the small ones on the tail. The cerata are long, cylin-

drical, and transparent, allowing the ramifications of the liver to

be distinctly seen. As noticed in the living animal, these latter

are long and thin, with well-developed knobs or short branches.

The rhinophores and tentacles are also long and thin; the former

bear about 15 rings near the top, but are smooth below. The

foot is very narrow, with thin projecting margins: in front it is

expanded into a semicircular disk ; the anterior margin is grooved,

and the corners are rounded.

The jaws, which were examined in all three specimens, were

transparent and very delicate. No denticles were to be seen on

the edge, and, though it is hard to be certain of their absence

in dealing with such slight and colourless material, it is to be

observed that they were found without difficulty in other similar

forms. The radula consists of a single series of horseshoe-shaped

teeth numbering 23, 20, and 18 respectively, in the three specimens.

Thin, pointed, lateral denticles extend almost up to the tip of the

central cusp: there are as many as 20 on either side, but

sometimes the number sinks to 15.

I think these specimens should be referred to Phidiana, in

spite of the doubt about the jaws, and should form a new species,

chiefly characterised by the large number of lateral denticulations

on the teeth. Also, the branches of the liver, which in other

species are covered with knobs, seem to bear distinct short

branches.

FACELINA LINEATA, Sp. n. (Plate XVI. figs. 4 & 5; and Plate

XVII. figs. 10 & 11.)

Two specimens from Zanzibar.

The notes on the living animal say that it had a general resem-

blance to Hervia lineata, which was caught about the same time,

but the colours were brighter, and there was an orange-red ring

round each of the cerata near the tip. The rhinophores were

jet-black. There were white lines on the body but not on the

cerata, and there were three red blotches between the tentacles

and rhinophores. The tail was long, and there was a very deep

groove along the front of the foot.

The largest of the preserved specimens is 6 mm. long and

2-5 mm. broad, but is evidently much contracted. The colour is

a uniform alcoholic yellow, except that the rhinophores are still

black. The disposition of the cerata is not quite clear, as many

have been lost, but appears to correspond with the drawing. ‘The

genital orifices seem to be below and between the first and second

group, and the vent after the third. The cerata are longish and

cylindrical. The oral tentacles are large and thick, but are clearly

much contracted, as are also the rhinophores, the perfoliations on

which are not so distinct as might be expected from the drawing

of the living animal (Pl. XVI. fig. 4). The narrow foct is

expanded at the sides into thin margins, and anteriorly into

deeply grooved tentacular processes.

1904. | FROM EAST AFRICA AND ZANZIBAR. 289

The radula consists of a single series of teeth of the form usual in

the genus, with a moderately large central cusp and six denticles on

each side, of which the innermost and outermost are the smallest

and the median the largest. The jaws bear a single series of

35 rather irregular denticles, The verge is armed with spines.

This appears to be a new species of Facelina, but I have not had

an opportunity of seeing the description of /. cyanella (Couth.),

which Bergh refers to this genus with a query.

PHYLLODESMIUM HYALINUM Khrenb.

[Hhrenberg, Symbole Physice, series prima, 1831 ; ‘Bergh,

“« Anatomisk Undersodgelse af Ph. hyalinwm,” Naturhist. Foren.

Vidensk. Meddelelser, 1860. |

One specimen dredged in about 10 fathoms near Wasin.

The notes on the living animal are as follows :—‘“ About one inch

long. Body semiopaque, pure white. Rhinophores and tentacles

ditto. Rhinophores slightly annulated, shorter than the tentacles.

Cerata very long and opaque, so that the liver is not visible;

whitish violet in colour; their upper halves and the whole length

of the sides are covered with low rounded projections, between

which dark-brown pigment is found. The first groups of cerata

are almost at the side of the rhinophores, and consist of only two

cerata on each side. The remaining cerata are set in seven pairs

of clumps of four each, and there is a space between the second

and third pairs of clumps. The cerata are somewhat flattened.”

The preserved specimen is 10°5 mm. long and 3 broad, with a

thread-like tail. Relatively to the size of the animal, the cerata

are enormous, the largest beg 8 mm. long. They are flattened,

rather convex on the outer and concave on the inner face. The

edge all round is marked by a line of knobs, which are, however,

more numerous at the top than at the bottom. On the outer side

the whole surface of the upper half is covered with similar knobs.

On the inner side the surface is mainly smooth except at the edges,

but at the very top there are a few knobs. The cerataare largest

on the inside, and gradually decrease outwards. The smallest

appear to have no knobs. The larger are easily detached, and

hence the arrangement of groups was not easy to see in the pre-

served specimen, but it appeared to have been as described in the

notes on the living animal. The rhinophores are short and thick,

set close together, and annulate. ‘The tentacles are longish and

curved. The foot is grooved anteriorly, and produced into mode-

rately long but not conspicuous tentacular angles. The anal

papilla is latero-dorsal, just behind the rather large pericardial

prominence. The genital openings are under the rhinophores.

The jaws bear five or six course deaticles, of which three are

very large indeed, the others smaller. The radula consists ot

sixteen colourless teeth, bearing between 30 and 40 denticles on

each side. The shape is much as in Bergh’s plates (/.c.), but the

denticles are rather longer.

Proc. Zoou. Soc.—1904, Vou. II. No. XIX, 1)

290 SIR C, ELIOT ON NUDIBRANCHS [ Nov. 29,

HERM AIDA,

STILIGER VARIANS, sp. n. (Plate XVI. fig. 6.)

Several specimens from Prison Island in Zanzibar Harbour,

found in green-branched seaweed, in which they are practically

invisible. The colour was very variable, ranging from dark

brown to white, but was as a rule brilliant green. After being

kept in captivity for a night, the animals grew perceptibly

paler. The main colour was largely hidden, except at the sides

of the body and in the centre of the back, by numerous lines

of a deeper colour, generally dark bright green, and in some,

but not all, specimens there were more or less extensive patches

of crimson lake. The form was somewhat elongate, and the

maximum dimensions 10mm. by 2mm. ‘The foot was fairly

broad and green.

The preserved specimens are colourless and semitransparent.

They have contracted into an oval or nearly semicircular form,

and strongly resemble small tufts of seaweed. The centre of the

back is bare, and through its transparent integuments can be

seen a great number of circular folliculate organs which are

apparently the follicles of the hermaphrodite gland. There are

about ten transverse rows of cerata in the largest specimens, con-

taining four (or sometimes five) cerata on each side of the central

space. The two innermost cerata of each row are large (about

3°50 mm. xX 2 mm.) and somewhat inflated. The others are very

much smaller and look like mere tubercles. They all contain

ramifications of the liver, consisting of one large main stem from

which spring three or four quite small and short branches. The

bladder-like pericardial prominence is somewhat elongate; in

front of it and fused with it is the anal tube. The rhinophores

are entire and not grooved. In the largest specimen they are

about 3 mm. long and rather thick, as if contracted. Below

them are two lumps which may be regarded either as a frontal

veil notched in the middle, or as rudimentary tentacles. The

anterior angles of the foot are not much produced. ‘The tail is

pointed but not long.

The radula was examined in several specimens, and was found

in all to consist of four or five teeth in the ascending part and

six in the descending, while the number in the heap did not seem

to exceed six or eight.

The teeth (Pl. XVI. fig. 6) are somewhat like those of Hrcolama

stotti (v. Trinchese, ‘ Aeolidide del Porto di Genova,’ vol. ii.

pl. x. figs. 7 & 8), and have a broad spoon-like hollow into which

the tooth behind fits.

In my account of Mr. Gardiner’s Nudibranchs (‘ Fauna and

Geography. of the Maldive and Laccadive Archipelagoes,’ vol. 1.

pt. i. p. 571, and pl. xxxii. figs. 9 & 10) this animal is erroneously

figured as Hermeea minor. It is, however, not a Hermcea and

not identical with Mr. Gardiner’s specimen. That specimen is a

Hermeea, and may possibly be ZZ. minor, as there suggested.

1904. | FROM EAST AFRICA AND ZANZIBAR. 291

STILIGER IRREGULARIS, sp.n. (Plate XVII. fig. 12.)

Two specimens from Chuaka on the Hast Coast of Zanzibar,

found among Sertularians. The animal had a somewhat peculiar

appearance owing to the hinder cerata being about twice as long

as those in front and spreading out in a fan-like shape. One speci-

men was of a translucent white, but the liver, extending in two

lines down the side of the body and giving off branches to the

cerata, was green, and created an impression that the whole

animal was of that colour. Im the other specimen the branches

of the liver in the cerata were of a dirty yellow, and there was

some reddish-grey pigment in the integuments of the body, so

that the longitudinal liver-tubes were not easily discernible. Near

the head, however, they were distinct and green. The cerata in

this specimen had white spots. The animals were less than

2 mm. long.

Only one specimen has been preserved, and its very small

dimensions rendered examination rather difficult. In the hinder

part of the body there are two longitudinal lines of cerata

arranged in five transverse rows on each side, of which the inner

are two or three times as long as the outer. In the front part

there is a single line of five small cerata, and there are no signs

of others having been detached. The cerata are cylindrical, much

like those of Hermea dendritica, and not inflated or ovate.

Though the surface of the liver-branches is irregular, they do not

appear to have distinct secondary ramifications within the cerata.

No pericardial prominence is visible. The rhinophores are short

and simple; behind them are two very distinct black eyes. The

oral veil is circular and not notched. The foot is truncate in

front; there were no signs of a groove or tentacular prolongations

of the corners.

The radula consists of four teeth in the ascending portion, six

in the descending, and a small heap. The teeth are much like

those of S. varians, but the outline is somewhat simpler and less

wavy (Pl. XVII. fig. 12).

PHYLLOBRANCHIDA.

{ Bergh, in 8. R. i. & xvi.; id. Beitrige zur Kennt. d. Aeoli-

diaden, ix.; A.& H., Coll. of Nudibr. Moll. made in India, p. 145 ;

Trinchese, Aeolididze del Porto di Genova, 1881; Pelseneer, Re-

cherches sur divers Opisthobranches, 1894, pp. 50-52. ]

This remarkable family, which is characterised by its flat leaf-

like dorsal papille, consists of three genera, Phyllobranchus,

Cyerce, and Caliphylla, of which the first two are recorded from

the Indo-Pacific (but Ph. viridis from the West Indies), and the

last from the Mediterranean. They all agree in having flat

leaf-like cerata, an ascoglossan radula and a buccal crop, compli-

cated reproductive organs, and (except Caliphylla) oral tentacles

as well as rhinophores. Cyerce, though very like Phyllobranchus

292 SIR C, ELIOT ON NUDIBRANCHS [ Nov. 29,

superficially, presents many points of difference and is certainly

nob a mere subgenus, as it is considered by Fischer (Manuel de

Conch. p. 343). Externally the chief difference is that the foot

in Phyllobranchus is, as usual, an undivided surface, whereas in

Oyerce there are two distinct parts separated by a transverse

division. In Phyllobranchus the buccal crop is long and twisted,

the vent lateral, and the teeth are preserved in a spiral. In

Cyerce the crop is flat and oval, the vent dorsal, and the teeth

are preserved in an irregular heap. The digestive organs also

present important differences, the chief of which is that whereas

in Phyllobranchus (and in Caliphylla) the liver is ramified within

the dorsal papille, in Cyerce it appears not to enter them at all

and to be wholly contained in the body-cavity.

The function of the large buccal crop is obscure. It would

appear that, asa rule, this organ is only found when the radula

is uniseriate (ascoglossan) or very narrow (Lamellidoris, Gonio-

doris, &e.), and when there are no jaws. On the other hand, its

presence under these conditions does not appear to be necessary

(e. g. Elysiadee and Hermeidz).

PHYLLOBRANCHUS PRASINUS B.

[Bergh, in 8. R. Heft ii. pp. 52-87. ]

Fourteen specimens from Chuaka on the Hast Coast of Zanzibar,

mostly about 3 centimetres long, but two much larger, measuring

over 5 centimetres in life. The animals were found at low tide.

The colour is described as transparent, with small green branching

lines on the cerata, giving on the whole an effect of greyish

green. It was noticed that the cerata break off easily when the

animal is disturbed, and retain the power of independent move-

ment for some time.

The preserved specimens have kept their colour fairly well, but

many of them show a yellowish rim round the cerata, not men-

tioned in the description of the living animal or by Bergh. The

yellow spot on the cerata mentioned by Semper is not visible.

The largest preserved specimen, which has lost nearly all its

papille, is 47 mm. long and 14 broad. The head is separated

from the body by a sort of ridge, which descends and forms

lappets on each side of the mouth. The oral tentacles are un-

divided, about 7 mm. long, and slightly grooved. The rhinophores

are bifid ; the main branch measures 12°5 mm. and the side branch

5-5. Both branches are grooved and also the common stem, but

less distinctly. The rhinophores seem larger than in Bergh’s

specimens; in one specimen only 25 mm. long they measure

12 mm. At the base of the rhinophores the black eyes are

clearly visible. The pericardial prominence lies 14 mm. from

the anterior margin. The opening for the penis is immediately

behind the right tentacle. The large female genital papilla is a

little further back on the right-hand side, under the rhinophores.

Still further back is the large, cup-shaped, anal papilla, under the

anterior end of the pericardial prominence. The foot has thin

1904. | FROM EAST AFRICA AND ZANZIBAR. 293

wide lateral margins and a tail 9 mm. long; the anterior margin

is thick, distinctly, but not very deeply, grooved, and produced

into tentacular expansions at the corners. The centre of the

back is bare, with small! irregular tubercles, whose number varies

greatly in different individuals. The sides are covered with

cerata, which extend right up to the lappets of the mouth. They

are easily detached, and all the specimens have lost many, but

they appear to be arranged in four longitudinal rows on each side,

with a few very small extra ones at the extreme outside. They

consist of an oval plate set on a short stalk; at the junction of

the plate and the stalk is generally a small funnel-like depression.

The edges of the plate are smooth and not denticulate, as in Bergh’s

specimens. On the inner surface are a number of lines bearing

small yellowish tubercles. These lines spring from three main

trunks, but subdivide, so that there are 10-15 at the edge. The

largest papille are those on the inside nearest the centre of the

back. In fine specimens the stalk is about 2 mm. high and the

plate 7 long and 6 broad.

The anatomy of this remarkable animal has been described by

Bergh (/. ¢.) with such elaborate thoroughness that a further

account is hardly necessary, although the function of some of the

internal organs is not clear. Both the digestive and reproductive

systems are extremely complicated. The buccal mass is large

(8°5 by 5 mm.) and striped. The radula is of the ascoglossan

type, the teeth being preserved ina regular spiral, not a heap.

Only two or three teeth are in useata time, but the total number

varies between 40 and 50 in large specimens. The shape of the

teeth is as described by Bergh, not elongate, with an indentation

in the back, and 12-14 squarish denticles on the edges. The

digestive apparatus including the large crop (which measured

35 mm. by 2 mm. when straightened) was as described by Bergh ;

but although I was able to follow the ‘“‘Seitengallengange ” for a

considerable length, I did not succeed in seeing that they form

a complete circuit and unite behind. It seemed clear that the

hepatic system resembles that of the Aeolids, and is ramified

within the cerata.

Though there is no sufficient reason to regard these specimens

as specifically distinct from Ph. prasinus B., points of difference

(such as the shape of the cerata, the length of the rhinophores,

and perhaps the coloration) are not wanting, and may indicate a

distinct variety.

CYERCE ELEGANS B.

[B. in 8. R. Heft ii. pp. 99-113.]

Three specimens from Chuaka. Mr. Crossland says of the living

animal :—‘“ The foot and central part of the body are white, the

sides of the body being dull green (?liver). Cerata very delicate

and colourless. Head, rhinophores, &c. translucent white.”

The preserved specimens are colourless, with semitransparent

integuments which allow the internal organs to be seen, particu-

294 SIR C. ELIOT ON NUDIBRANCHS [ Nov. 29,

larly a large folliculate mass which covers the sides and part of the

centre.

The largest specimen is 20°5 mm. long and 11 broad across the

back. The general construction of the head-parts is as in Phyllo-

branchus, with the dividing line running down and forming

lappets by the mouth, grooved oral tentacles, and grooved bifid

rhinophores about 4 mm. long. At the base of these latter can

be seen the large black eyes. The genital openings are as in

Phyllobranchus, but the intestine terminates dorsally in a cylin-

drical tube set in front of the pericardium and slightly to the

right of the median line. The foot is in two divisions, of which

the anterior is the wider, being 17°5 mm. broad by 7:5 in the

longitudinal direction of the body, whereas the posterior portion is

12 mm. long by 9°5 at its broadest part. Except for this division,

the foot is as in Phyllobranchus prasinus, but the corners of the

anterior margin are not much produced. Nearly all the papille

have fallen off, but it appears that they were arranged at the sides

of the back, leaving the centre bare. The largest are about 5 mm.

high and 3:5 broad. Though they taper towards the base, they have

not a distinct stalk. On their margins are 8-10 yellowish spots.

The anatomy of this species, like that of Phyll. prasinus, has been

elaborately investigated by Bergh (/.c.), but the structure of the

hepatic system is obscure. In my specimens most of the papille

are quite transparent, and it seems clear that they contain no

hepatic branches, and when they are held out from the body

the folliculate mass, which I take to be the liver and which is

distinctly visible, is not seen to send any prolongations into their

transparent bases. On the other hand, the folliculate mass

adheres to the sides of the body-wall in the neighbourhood of the

papillee, which it does not do elsewhere, and must be in immediate

contact with the openings at their bases. Within the papillz are

round bodies which look like minute bubbles, and in some cases 1t

seemed that these bubbles were connected by a system of colourless

canals. I have unfortunately no means here (Hast Africa) of

preparing sections for microscopic examination.

The radula consists of elongated teeth as figured by Bergh (4. ¢.

pl. xv. figs. 5-11), each bearing 12-17 denticles, but the number

of teeth seemed less, and was 14+9 in one specimen and 17410

in another. The second figure in these expressions represents the

teeth which have fallen down into an irregular heap, and this

heap was much smaller than that described and figured by Bergh,

possibly in consequence of the youth of the specimens.

HLYSIADA.

PLACOBRANCHUS OCELLATUS Van Hass.=PL. arcus B. (Plate

XVII. figs. 13 & 13.4.)

[B. in S. R. iii. pp. 147-165, and id. Danish Exp. to Siam,

Opisth. pp. 180-181.]

Two specimens from Prison Island, Zanzibar Harbour. ‘The

larger measured 30 mm. in length, and 15 in breadth when the —

1904. | FROM EAST AFRICA AND ZANZIBAR. 295

sides were folded over the body, but 23 when they were extended.

The ground-colour was mainly whitish, but on the dorsal surface

this was almost entirely hidden by the numerous dark green

branchial ridges. The outer surface of the lateral expansions was

sandy coloured with darker spots, and on the foot were many

irregularly-arranged deep black spots. Along the junction of the

lateral expansions and the foot ran a line of violet rings, and

there were four or five more on the forehead. The inside of the

furrowed rhinophores was violet, as was also the end of the body.

The animal secreted a very abundant mucus.

In the preserved specimen the branchial ridges are very large

and distinct, being as much as 1:5 mm. high. There are about

36 main folds, and smaller ones in between. The buccal mass is

small, only about 1°5 mm. long, and the crop half that length. ‘The

teeth are exactly as figured by Bergh, with about 12 denticles on

the margin, but are less numerous than mm his specimens. In the

radula I found 15 and 14 respectively, and in the heap at the

bottom about 40 and 50.

Two pencil drawings by Mr. Crossland are reproduced because

they show the animal in a somewhat different attitude from that

in which it is ordinarily represented.

HLYSIA FAUSTULA, B.

[B. in 8. R. iv. pp. 186-190. ]

One specimen from Wasin, Kast Africa.

There are unfortunately no notes on the living animal.

The preserved specimen is very flat and crinkled, surprisingly

hike a planarian in appearance, and also somewhat resembling

Tridachia, but the wings show no signs of being joined hehind

the neck. The length is 19 and the breadth 16 mm., but the

form has become somewhat contorted, and these measurements

represent at least 25 and 20 if it were straightened out. The

colour is a uniform pale yellowish grey, with a very distinct deep

black border all round the edge, and a few scattered black dots on

both the upper and lower surface. There are three black dots

on the pericardial prominence, and the anterior margins of the

tentacular groove are black. There is a fine furrow dividing the

foot transversely just below the point where the wings arise,

behind which the foot is not clearly differentiated from the sides

of the body. The anterior margin of the foot is not expanded

into tentacular processes. From the rather large pericardial

prominence issue three vein-like ridges on each side. The two

anterior pairs are simple and have only very slight ramifications

near the edge of the wings. Thethird pair soon divides into three

main branches, which have one or two secondary ramifications.

The tentacles are rather large and broadly opened.

The radula consists of 17 teeth, besides which there is a heap of

about ten disused ones. The shape is exactly that given in Bergh’s

plates (in 8. R. iv. pl. xxii. figs. 15-17), elongate, and with no

trace of denticulations.

I think this specimen may be certainly referred to Z. faustula B.,

296 SIR C. ELIOT ON NUDIBRANCHS [ Nov. 29,

recorded from the Philippines. The specific characteis appeai to

be the coloration, the planarian-like shape, and the elongate

smooth teeth.

ELYSIA MARGINATA Pease. (Plate XVI. figs. 7 & 8; and Plate

XVII. fig. 18.)

[Pease, Amer. Journ. of Conchology, 1871, vol. vi. p. 304; ef.

Bergh, Jour. Mus. Godeffroy, Heft ii. 1873, p. 80, on Llysia

nigrocincta. |

The following are notes on living specimens captured at

Zanzibar :—

1. “Extreme length 1°5 cm. General colour a dull green,

but on the surface are opaque black and white spots. The

tentacles and edges of the wings are bordered with orange-yellow,

white, and on the extreme edge black. The white line is irregular,

Internally the edges of the wing are blotched with white. Hach

black spot has a corresponding orange spot just beneath.”

(Vide figures 7 & 8, Pl. XVI.)

2. ‘In a second specimen the orange line round the wings was

much broken, perhaps a step towards its disappearance.”

3. “Rather more than an inch long, when fully extended.

Tentacles and wings edged with a double border of orange and

black, which is interrupted at the neck. Whole body spotted

with small black flecks. Head and auriform tentacles very large.

Pericardial bulb prominent. Inside of wings not striated, but

the veins can be seen beneath the skin.”

4, “Green, without black spots, but wings bordered with a

double line of orange and black. Length about one inch.”

Unfortunately, only the second and fourth of these specimens

have been preserved.

As preserved, the latter is 1] mm. long and 6 high, the wings

being raised, not spread out, but applied to one another. ‘The

colour is olive-green: the borders have disappeared, but the

outside of the wings is covered with numerous yellowish-white

spots, and there are a few inside. The tentacles are short and

thickish, with traces of black about the groove. The wings are

rather thick, not much indented at the edges, and the posterior

expansion is not ample. The pericardial prominence is distinct,

and the anus lies to the right side of it. The inside of the wings

is smooth and does not bear ridges, but where the animal is

sufficiently thin to be transparent, veins can be seen radiating

from the pericardium.

The foot is not distinctly divided from the body, and the front

part is hardly differentiated from the rest. The radula consists

of 14 teeth, and there are about 12 more of very varying

sizes lying in a heap. The teeth are as in Bergh’s plates of

E. nigropunctata (1. c. pl. xi. fig, 10* and pl. xii. 1), but seem

somewhat more slender. ‘There is no trace of denticulation.

The second specimen is 7°5 mm. long, 7 broad when the wings are

* There seems to be a mistake in the explanation of this plate, figs. 13-26

apparently referring to Cyerce and not to #. nigropunctata as stated.

1904. | FROM HAST AFRICA AND ZANZIBAR. 297

spread out, and 4:5 high when they are closed and folded together.

The colour is yellowish green with black spots within and without,

the borders having disappeared. The shape and other external

characters are exactly as in the specimen just described. The

teeth are also similar but a trifle more bent: there are 16 in the

radula, and about 15 of various sizes in the heap.

I think these specimens must be referred to Hlysia (Pterogastroi)

marginata Pease, although his description is somewhat deficient

in details. In view of the fact that the teeth of the animal

examined resemble those of H. wigrocincta, as described by Bergh,

and that the coloration is clearly very variable, it is probable that

both #. nigrocincta and marginata are varieties of a protean

species ranging from green spotted with black, but without a

coloured border, to green with or without black spots, and a more

or less continuous single, double, or triple border.

Eitysta DUBIA, sp. n. (Plate XVII. figs. 14-17.)

Four specimens from Chuaka, found on Zostera at low spring-

tides. The animals were dark green with a few spots of dull,

light blue. They were about 6 mm. long and 3 broad when at

rest, but when crawling become narrower and more elongated.

They can also swim on the surface of the water foot uppermost.

As preserved, the specimens are of a uniform dark green, and

have somewhat the appearance of a minute Aplysia, as the wings

do not reach to the end of the body but terminate separately,

leaving a distinct tail. The tentacles have become mere knobs,

but were apparently of a fair size in life. The foot is very

distinctly divided from the body by a ridge. The pericardium is

continued into a long median ridge down the centre of the body.

At about the point where the wings end it bifureates. From each

side of this central ridge issue seven or eight vein-like ridges,

much as in Placobranchus. The radula consists of 14 teeth and

about 20 in the heap; they are elongate and slender. The basal

part is nearly as long as the hook and there are no denticulations.

The specimens are perhaps immature.

If Hlysiella is regarded as a separate genus, these specimens

should probably be referred to it, but Bergh (Beitr. zur Kenntniss

der Aeolidiaden, viii. 1886, p. 17) seems doubtful as to the

validity of the genus and its definition. For the present I think

it simpler to refer this form to Hlysia.

EXPLANATION OF THE PLATES.

Prate XVI.

Fig. 1 Phyllidia nobilis, dorsal view (p. 282).

2. Hervia lineata, dorsal view (p. 286).

3. os one of the cerata.

4, Facelina lineata, dorsal view (p. 288).

» one of the cerata.

. Stiliger varians, three teeth (p. 290). The outline of the middle tooth

is coloured red for distinctness.

. Elysia marginata, dorsal view with wings open (p. 296).

99 a dorsal view with wings closed and body elongated

ON DON}

298 MR. ROBERT GURNEY ON FRESHWATER [ Nov. 29,

Puatt XVII.

Fig. 9. Madrella ferruginosa, central tooth (p. 269).

10. Facelina lineata, side view of anterior end (p. 288).

ih, 55 As ventral view of anterior end.

12. Stiliger irregularis, tooth (p. 291).

13, 13a. Placobranchus ocellatus, dorsal views (p. 294).

14, Hlysia dubia, wings open (p. 297).

15. 53 ‘ wines closed and body elongated.

16. 3 » crawling foot uppermost on surface of water.

17 teeth.

18. Blysia marginata, teeth (p. 296).

3. On a small Collection of Freshwater Entomostraca

from South Africa. By Ropert Gurvey, B.A., F'.Z.8.

| Received June 21, 1904. |

(Plate XVIIL*)

The collection which I describe here was kindly entrusted to

me to work out by Prof. Jeffrey Bell on behalf of the Natural

History Museum. The specimens were partly mounted on slides

and partly contained in tubes, and were collected by Major E.

Heckersley, R.A.M.C., from a water-hole on the veld at Kroonstad,

O.R.C. This water-hole was a collection of surface-water, quite

dry in ordinary weather but filled up by thunder-showers.

Unfortunately very few specimens were preserved, but the few

that there are seem of sufficient interest to deserve description,

especially as so little is known at present about the Entomostraca

of South Africa. Of the seven species collected, only three have

been previously described ; of the other four, three are apparently

new and one is represented by only a single mutilated specimen.

The following is a list of the species :—

PHYLLOPODA.

Fam. BRANCHIPODIDA.

STREPTOCEPHALUS DREGEI, G. O. Sars. (Plate X VIII. figs. 1, 2.)

Sars, Arch. f. Math. og Naturv. xxi. no. 4, 1899, p. 19.

Of this species three males and three females were included in

the collection. The male only has as yet been described, so that

I will give a short description of the female.

Female.—Body slender ; thoracic region as long as the caudal

region exclusive of the caudal rami, which are long and densely

fringed with ciliated sete. Superior antennz long and slender;

inferior antenne considerably shorter than the superior, foliaceous,

the tip truncated and with a short conical process. Marsupium

not reaching beyond the second caudal segment: in one specimen

it contains a single row of eggs, each enclosed in a thick capsule

with conspicuous more or less polygonal markings (fig. 2).

* Wor explanation of the Plate, see p. 301.

PZS.1904,vol. I] Pl. XVII.

E.Wilson, Cambridge.

SOUTH AFRICAN FRESHWATER ENTOMOSTRACA.

1904. | ENTOMOSTRACA FROM SOUTH AFRICA. 299

Total length 14 mm.; thorax 7 mm.; caudal rami 3 mm.

The caudal rami of both sexes are stated by the collector to

have been red during life.

One of the male specimens is 25 mm. long—very much larger

than Sars’s two specimens, which measured 16 mm. only.

Fam. LimnADImD&.

ESTHERIA ELIZABETH G. O. Sars.

Sars, Arch. f. Math. og Naturv. xx. 1898.

The collection included several specimens of this species, both

male and female.

While agreeing in all essential particulars with the description

given by Sars, the females differed in having no cilia upon the

base of their caudal claws.

Fam: LIMNETIDZ.

LIMNETIS WAHLBERGI Lovén.

Loven, K. V.-Akad. Handl. 1845, p. 203.

The specimens examined were all females.

CLADOCERA.

Fam. DaPHNnip.

MOINA BELLI, sp.n. (Plate X VIII. figs. 3, 4.)

Dorsal margin of head evenly rounded, without any concavity

above the eye; ventral margin somewhat protuberant; posterior

margin finely ciliated. Fornix well developed and extending over

the eye.

Shell without any trace of striation; ventral margin setose for

about two-thirds of its length. First antenne ciliated all over.

Tail of the usual shape, with eight lateral teeth, the first of which

is bifurcated. Between the bifurcated tooth and the first simple

tooth is a minute elevation covered with cilia, which may represent

a rudimentary tooth. Apical claws armed with a basal row of

secondary denticles and with a ventral chitinous expansion cleft

into teeth. Posterior dorsal surface of tail provided with cilia,

which are more or less arranged in transverse rows. Ephippium

reticulated all over and containing two resting eggs.

Length 1-7 mm.

Several specimens of this species were contained in the collection,

but all were females, one of which was ephippial. The species

very much resembles UW. wierzejskii Richard, and perhaps should

be regarded as only a variety of that species. It is mainly distin-

guishable by the ciliation of the head and first antenne, and by

the structure of the postabdomen.

300 MR, ROBERT GURNEY ON FRESHWATER [ Nov. 29,

DAPHNIA sp.

One of the slides in the collection was unfortunately broken in

transit, and from the debris I separated the dried and crumpled

body of a Daphnia. It is, however, impossible to make out more

than that it belongs to the Daphnia magna group, with a much

pointed head and a very well-developed fornix.

Kam. Lyncrem a.

LEYDIGIA AFRICANA, sp. n. (Plate XVIII. figs. 5, 6.)

In general appearance very like LZ. acanthocercoides. Rostrum

short and acute. Shell showing faint striation. Hye very slightly

larger than the ocellus (fig. 5). First antenna shorter than the

rostrum. Tail closely resembling that of LZ. acanthocercoides, but

differing from it in the presence of a minute tooth at the base of

each of the terminal claws, and in the arrangement of the spines

and cilia. In L. africana only the first seven spines have accessory

spines at their base and the dorsal margin of the tail is not evenly

ciliated, but is provided with a few very small spines (fig. 6).

Length 54 mm.

Four specimens of this small species are included in the collec-

tion, and it is stated by the collector to have been ‘not at all

common.” It differs mainly from ZL. acanthocercoides in its small

size and in the relative proportions of the eye and ocellus, which

are practically the same size.

CoPEPODA.

Fam. CENTROPAGIDA.

LovENULA MEA, sp. n. (Plate XVIII. figs. 7-13.)

Female.—Body slender, with the anterior segment tapering

evenly ; lateral lobes of last segment expanded, symmetrical, and

armed each with two minute spines. ‘Tail consisting of two seg-

ments only, the anterior one produced laterally into lobes which

are asymmetrical. Caudal rami armed with five strong ciliated

setee, and a slender one springing from the dorsal surface be-

tween the two innermost terminal sete; inner edges of the rami

ciliated. Posterior antenne with outer ramus slightly longer

than the inner, but in other respects exactly as in Lovenula

falcifera (Lovén). Posterior maxillipedes very closely resembling

those of L. falcifera, but the terminal part is distinctly composed

of four joints instead of three. The second basal joint bears

terminally a small lobe armed with two sete. In ZL. falcifera

two of the strong terminal spines are borne upon the penultimate

joint, and the third upon a small terminal joint, whereas in the

present species each is borne upon a separate joint. ‘The

swimming-legs agree in all respects with those of L. falcifera.

1904. ] ENTOMOSTRACA FROM SOUTH AFRICA. 301

Last pair of legs with the inner ramus one-jointed and armed

terminally with two short spines; terminal joint of the outer

ramus provided with three stout spines, of which the inner and

stronger is a prolongation of the joint; the other two are jointed,

the middle one being cleft into two nearly equal spines.

Length 3 mm.

Male.—Body rather more slender than that of the female; tail

consisting of five segments, the second and third of which bear a

few very minute spines. Caudal rami asin female; the sete not

asymmetrical as in L. falcifera. Last legs closely resembling

those of that species (fig. 13).

Of this species two females and one male were included in the

collection.

I have been in doubt as to whether to include this new species

in the genus Lovenula Schmeil, or to form a new genus for its

reception. Among the most important characters of the genus

BGroteas, as defined by G. O. Sars (1899)*, are these: that the

caudal rami of the male are asymmetrical, and that the posterior

maxillipeds have only three terminal joints. In both these

characters my species differs from the type. On the other hand,

the agreements are very striking in other respects: in the form

of the swimming-legs, in the two-jointed tail of the female, even

in the form of the posterior maxillipeds (apart from the number

of joints), it agrees with the genus Lovenula or Broteas, and

differs from all other genera. If it is to be included in the

genus Lovenula the definition of that genus will have to be

modified somewhat. Lovenula mea may be said to provide a link

with the genus Diaptomus, being less specialised in the form of

its furca and posterior maxillipeds than are the other two species,

L. faleifera and L. lumellata (Sars).

EXPLANATION OF PLATE XVIII.

Fig. 1. Streptocephalus dregei Sars (p. 298). Dorsal view of head of female. x 18.

2. s oe An egg from the marsupium. X 34,

3. Moina belli, sp. u. (p. 299). Side view of female. 23.

4, 35 A Postabdomen. X 90.

5. Leydigia africana, sp. un. (p. 800). Side view of female. X 60.

6. ie . Postabdomen. 156.

7. Lovenula mea, sp.n. (p. 300). Dorsal view of female. The left furcal ramus

broken off. 18.

iB a Tail of another female, showing spermato-

phore. X 34.

9. 5b Me Posterior maxillipede of female. X 60.

10. 59 s First leg of female. 72.

ale =f i Last leg of female. X 60.

12. Ps 3 Posterior maxillipede of male. » 60.

13. Bs #3 Last pair of legs of male. X 60.

* Arch. f. Math. og Natury. xxi. no. 2, 1899.

302 DR. H. J. HANSEN ON THE [ Nov. 29,

4. On the Morphology and Classification of the Asellota-

Group of Crustaceans, with Descriptions of the Genus

Stenetrium Hasw. and its Species. By H. J. Hanszn,

PhD SMES:

[Received October 17, 1904.]

(Plates XIX.—XXI*.)

I. Introductory Remarks.

The tribe or suborder Asellota is in some respects one of the

most varied and, as to the number of species, probably by far the

richest of all groups of pre-eminently marine Isopoda. A perusal

of the portion in question of Beddard’s account of the ‘Chal-

lenger’ Isopoda (Zoology, vol. xvii.), and of G. O. Sars’s ‘An

Account of the Crustacea of Norway,’ vol. ii. Isopoda, 1896-99,

conveys a fair idea as to the striking differences in general aspect

and in some structural features between the numerous genera,

among which we find such types as Asellus Geoff., Janira Leach,

Munna Kr., Dendrotion G.O.8., Macrostylis G.O.8., Ischnosoma

G. O. S., Desmosoma G. O. 8., Munnopsis M. Sars, Hurycope

G. O. 8. In the work named, Sars describes 42 species referred

to 21 genera; Beddard has established 32 species referred to

15 genera, and 8 of these genera are not found in Norway. In

order to furnish an instance showing how much remains to be

known, | may perhaps state here that from the seas around

Greenland, Iceland, and the Faroe Islands, the Copenhagen Museum

possesses more than 90 species, of which at least 60 are new to

science ; the major part of these new forms were secured by the

‘Ingolf’ in depths between 300 and 1870 fathoms.

Sars refers his 21 genera to five families, but it will be shown

below that four of these are almost artificial, as really good

characters for their separation are wanting. But this distinguished

author has produced a vast number of good figures—with useful

descriptions—of all his forms and of details of their dermo-skeleton ;

moreover almost all really important genera hitherto established

of Asellota have been incorporated in his fine work. For these

reasons I can often, in the following discussion, refer the reader

to his figures as proofs and illustrations for my remarks.

Among the genera not represented in the Norwegian fauna,

Stenetrium Haswell is the most aberrant and important. Of

this genus five species have been established by four authors,

but our knowledge of several essential points of its structure,

especially of the pleopoda, is still imperfect. Most of the figures

of S. antillense, sp. n., were drawn by me more than fifteen years

ago, but publication was, however, postponed, which proved to be

fortunate, as new and interesting species have been received in

* Wor explanation of the Plates, see p. 330.

1S. IOs. Well Jl. IIL, AIK,

x Y AN Z | iy

ue : : :

\ \ i

\ |

H.J.Hansen del. :

E.C. Knight lith. West, Newman imp.

1. STENETRIUM ARMATUM Hasw. 2. S.MEDITERRANEUM 7. sp.

3.§.SERRATUM rsp

PZ. S. IOS. wOllail, IAL 2OG,

H.J. Hansen del. ‘

H.C. Knight lith. West, Newman imp.

1. STENETRIUM SERRATUM nsp 2.8. OCCIDENTALE np.

3. §. ANTILLENSE n.sp

P.Y.S8.1904. voli. Pl. XX1.

\ ZZ SS

\ Z oN

. Z

H.J. Hansen del. ; 5

E.C. Knight lith. West, Newman imp.

1. STENETRIUM ANTILLENSE 7.sp 2.8. SIAMENSE 7p.

3-6. VARIOUS ASHLLOTA.

1904. | ASELLOTA-GROUP OF CRUSTACEANS. 303

later years. In the following pages I describe six species examined

by myself of this type, five of which are considered new to science.

Having now such a rich material of an important and imperfectly

known genus at my disposal, I thought it appropriate to work it

out ; besides, I seize the opportunity of elucidating some points

of the mor phology of the pleopoda in other Asellota, and discussing

the classification of the whole group.

Il. Description of the Genus Stenetrium Hasw.

Body oblong, three or four times as long as broad, rather

depressed, shaped nearly as in Lanira.

Head with the dorsal surface much broader than long; in

advance of the anterior margin is seen a transverse area, the

frontal plate (PI. XIX. fig. 1 a, f), between the insertions

of the antennule. When the head is stretched forward a large,

sloping, anteriorly rounded part is seen in advance of the frontal

plate ; this area is clypeus and labrum, and when the mandibular

palps are in their natural position the distal part of their second

joint and the whole third joint are observed on the surface of the

clypeus. Eyes always distinct, but varying much as to shape,

size, and situation.

Antennule from somewhat shorter to a little longer than the

breadth of the head. Peduncle three-jointed ; basal joint oblong

but rather thick, longer and much thicker than any of the two

others. Flagellum varies much in length and number of joints.

Antenne nearly as long as or a little longer than the body,

very similar to those in Janira. The peduncle consists of four

short and two long joints: the first joint is always well developed

(and shows excellent specific characters) ; third joint on the outer

side with an exopod which is a subtriangular, oblong, setiferous

plate, with the lateral parts bent inwards.

Mouth-parts essentially as in Janira. Mandibles (Pl. XX.

fig. 3a) moderately long; lacinia mobilis of the left mandible

(fig. 3 5) consists of a thick, long, movable process and a few very

broad, long, a little curved sete, pectinate with exceedingly short

teeth along their anterior margin; one of these setz proceeds

from the process itself near its base; the articulating membrane

is broad on the lower side. Lacinia mobilis of the right mandible

(fig. 3c) shows a number (in S. antillense about ten) of very thick

sete, nearly all with saw-teeth. Mandibular palp well developed,

three-jointed; terminal joint rather broad and long, with

a comb of numerous fine sete. Hypopharynx (paragnatha)

(Pl XX. fig. 3d) rather deeply bifid; each half has the

inner margin ee straight and clothed with fine bristles

on its distal part; the anterior angle rather rounded and

the outer margin very convex.—Maxillule (fig. 3) slender; the

inner lobe terminates in three curved, thick, plumose setz ; outer

lobe (fig. 3) with a good number of very thick, curved spines,

coarsely serrate along at least one margin. Maxille (fig. 3 g)

304 DR. H. J. HANSEN ON THE [ Nov. 29,

nearly as long as the maxillule, rather slender; the lobe from

the second joint oblong, as long as the two lobes from the third

joint. Maxillipeds (fig. 34) large; second jomt—the lobe not

taken into consideration—very large, more than twice as long as

broad, its lobe, which is marked off by a transverse suture, is

large, longer than broad, with several small hooks at the inner

margin, while the distal margin is cut off and furnished with

several short sete, some of which are very broad, scale-like; fourth

and fifth joints rather expanded; the two distal joints slender.

Epipod very long, about three times longer than broad. Basal

joint in the adult female without any leaflet directed into the

marsupium.

Thorax shaped nearly as in /anira; anterior lateral angle of

first segment always produced into a triangular, acute, flat

process directed forward*. First pair of legs terminates in a

prehensile hand, the sixth joint being large, compressed, with

the palmar margin armed with processes or remarkable spines or

sete, while the seventh joint together with its short terminal

claw is slender and claw-shaped; this hand shows sometimes

rather little, but often a highly developed, sexual difference, being

frequently not only much larger in the adult male than in the

female, but of quite another shape. The six other pairs of legs

essentially as in Janira; seventh joint terminates in a claw,

beneath which a spine of the same size is seen. Marsupial

lamellee four pairs, proceeding from first to fourth pair of legs.

Abdomen essentially as in Janira; two rudimentary segments

ave observed in front of the large abdominal shield; the latter

has at the end of each lateral margin a small notch, the outer

margin of which is formed by a sharp triangular tooth.

Pleopoda are exceedingly characteristic 7. In both sexes the

third pair is only to a very small extent (Pl. XIX. fig. 1 d) covered

by the first pair (in the female) or the two anterior pairs (in the

male); its sympod is rather small, quadrangular (Pl. XX. figs. 22

and 2); the two-jointed exopod is exceedingly large, scarcely

respiratory, and covers the respiratory endopod, which is unjointed

and several times smaller; the inner margin of each exopod is

straight, and the two exopods touch each other along the mesial

line, constituting together a kind of operculum which covers the

lower surface of abdomen, with the exception of a moderately broad

margin at the sides and behind, and a small portion in front

occupied by the anterior pleopoda. In the female the pleopoda

of the first pair are completely fused (Pl. XX. fig. 2m; Pl. XXI.

fig. 22), constituting a subtriangular more or less oblong operculum,

which is at least three times smaller than an exopod of the third

* According to Haswell this process is wanting in S. inerme Hasw., but in the

sequel it is shown that this species probably does not belong to the genus

Stenetrium.

+ In Section V. of this paper the comparative morphology of the pleopoda in the

Asellota is discussed; in this account of Stenetriwm and in Sections III. and IV.

the structure of the pleopoda is described and the interpretations applied without

explanations.

1904. | ASELLOTA-GROUP OF CRUSTACEANS. 305

pair of pleopoda. Second pair is wanting, as in all Asellota. In

the male the first pair (Pl. XX. fig. 2g; Pl. X XI. fig. 2) is

slightly longer and at the base narrower than the female oper-

culum ; the sympods of the two appendages are completely fused

with each other, forming a short, transverse plate; each pleopod

has one ramus, which is free, oblong, and between two and three

times longer than the sympod ; each ramus can be moved a little by

a tiny muscle (m.) in the sympod. The second pair is a good deal

smaller than the first; each appendage (Pl. XX. fig.2; Pl. X XI.

fig. 2g) consists of an oblong subtriangular plate, the sympod, the

inner margin of which is sinuate, and from the distal end of this

margin arise the two rami. The exopod is very small, slender, a

little curved, scarcely hook-shaped, one-jointed, but in S. stamense

(Pl. XXI. fig. 2h, ex.) a vestige of a division into two joints 1s

observed. The endopod is rather long, very slender, two-jointed,

and strongly geniculate in the articulation (Pl. XXL. fig. 2h, en.) ;

the proximal joint contains a muscle for the movement of the

second, which has no internal cavity, while its end is obtuse and

often furnished with a brush of exceedingly short bristles;

especially in S. antillense, the terminal portion bearing this

brush is distinctly marked off from the joint. The plate-shaped

sympod contains muscles to the rami, two to the endopod, and

at least one to the exopod. The two pleopoda of this pair touch

each other at their base; they are covered by the first pair. In

both sexes the pleopoda of the fourth pair (Pl. XX. fig. 2 £) are

similar as to size and structure; each has a short, broad sympod

and a two-jointed exopod, which is slightly longer and somewhat

broader than the unjomted endopod, and adorned with plumose

setee along the distal part of the outer margin; both rami

are lamellar and both seem to be respiratory. The fifth pair

(Pl. XX. fig. 27) has no discernible sympod and only one ramus,

which is large, unjointed, but otherwise shaped and adorned with

setze like the exopod of the preceding pair, and accordingly it is

in all probability the exopod itself.

Uropoda consist of an unjointed sympod and two unjointed

nearly styliform rami; the exopod is as long as or longer than the

sympod, nearly as long as or somewhat shorter than the endopod.

fl. Comparison between the Genera Stenetrium and Asellus.

Pl. 39 in Sars’s work is filled with figures of Asellws aguaticus L.

{can therefore refer to his good drawings, and give only a new

figure of the second male pleopod.

The essential differences between Asellus Geoff. and Stenetriwm

Hasw. are found in the antenne, the maxillipeds of the adult

female, and some of the pleopoda. The peduncle of the antenne

shows the same number of joints in both genera, but in Asellus

the exopod from the third joint is wanting. In Asellus the basal

joint of each maxilliped possesses in the ovigerous female a rather

large plate, bearimg a number of bristles at the end and directed

Proc. Zoou. Soc.—1904, Vou. Il. No. XX. 20

306 DR. H. J. HANSEN ON THE [ Nov. 29,

backwards; it has been mentioned and well drawn by Sars ; its

function is certainly to produce a current in the water of the

marsupium,

The pleopoda show, however, some more interesting features.

As in Stenetriwm, the three posterior pairs in the male do not

differ from those in the female; the very large two-jointed

exopods cover, as in Stenetriwm, not only the small respiratory

endopods but almost the whole lower surface of the abdomen,

and are freely exposed with the exception of a rather small basal

portion. The fourth and fifth pairs are essentially alike, both con-

sisting of a short sympod and two rami, viz. a two-jointed exopod,

somewhat larger than the unjointed endopod ; consequently we

have here a well-marked difference in the fifth pair between Stene-

trium and Asellus, as in the former genus the exopod—according

to my interpretation above—is unjointed and the endopod want-

ing. In the female the pleopoda of the first pair are not fused

as in Stenetriwm, but independent and originate rather distant

from each other; each pleopod consists of a nearly rudimentary

sympod and a moderately large circular plate distally edged with

plumose setze. In the male each appendage of the first pair consists,

as in Stenetriwm, of a short sympod and a much longer suboval,

movable ramus; but while in the latter genus the two sympods

are completely fused, they are free in Asellus, but yet furnished

with some hooks* along their inner margin, so that they can be

coupled together nearly as the second joint of the maxillipeds. The

second pleopoda in the male are interesting; the distal half of the

left sympod with its rami is shown from below in Pl. XXT. fig. 3.

The sympod is shortly oval, with both rami proceeding from its

end and containing strong muscles for their movement. The

exopod is oblong, nearly lamellar, only a little shorter than the

endopod, two-jointed ; the distal joint is somewhat larger than

the basal one, obliquely oval, with marginal sete and containing a

good-sized muscle. The endopod consists of two movable joints ;

the proximal joint is short, but produced into a long, slender,

curved process, turning inwards and forwards along the inner

margin of the sympod; it contains a small muscle to the second

joint. This is obliquely oval, its distal end rather rounded, but

near the end a few minute teeth and irregular incisions and

depressions are seen; the joint is besides inflated, and the major

portion of its interior is occupied by a large pear-shaped sac, which

opens at a short distance from the end of the joint; the wall of

this sac is well chitinised. Having removed by dissection the

major part of the wall of the joint itself, I was able to examine

the wall of the inner sac. In the female second pleopoda are

wanting.

Before attempting to decide as to the systematic import-

ance of the differences between Stenetriwm and Asellus, it may

* Fach hook is a very thick and rather short spine, the end of which is broadly

rounded, curved very slightly upwards, and the upper surface of its terminal portion

is set with from seven to ten tiny, sharp, oblong teeth.

1904. ] ASELLOTA-GROUP OF CRUSTACEANS. 307

be appropriate to discuss the structure of the other Asellota,

especially their pleopoda. It may be added that, according to

descriptions given by 8. I. Smith and A. 8. Packard, the two

genera Mancasellus Harg. and Cacidothea Pack. are closely allied

to Asellus in the structure of antennz and pleopoda, the essential

difference being that the endopod of the second male pleopod has

no process from the basal joint.

IV. On the Structure of the Asellota, Asellus and

Stenetrium excepted.

Some years ago A. Dollfus described and figured (Bull. Mus.

d’Hist. Natur. Paris, 1898, no. 1, p. 37, figs. 2 & 2a) a very

curious animal, Stenasellus virei Dollf.; unfortunately he had

only one minute and mutilated specimen from fresh water in the

Cévennes. He refers the genus to the Asellota. His description

together with the two figures are certainly sufficient for the

recognition of the species, but not for deciding the question of

the relationship of the genus. The uropods areas in the Asellota,

and the four thoracic legs figured are, so far as can be seen, not

very different from those in Janira; but the animal differs from

all Asellota in two features. The author says: “ Cephalon intime-

ment uni au premier segment pereial,” which is not the case in

any form of the Asellota hitherto known. The other feature is

in the structure of the abdomen. Dollfus writes: “ Pleon a trois

premier segments trés développés”; this agrees well with his

figures, which show the abdomen as consisting of an oblong

“pleotelson” and three segments; these latter are slightly nar-

rower than, and their sum at least half as long as, the posterior

undivided portion. But this abdomen differs much from that met

with in any of the Asellota or any other group of Isopoda. In

Dollfus’s description we find as to the pleopoda only the statement

that they are ‘‘narrow,” and they are nearly invisible in his

figures; besides, he does not mention the mouth-parts. Judging

from all these circumstances I thought that Stenasellus could not

be referred to the Asellota, and in the manuscript despatched to

London in October I added some further critical remarks. But

at the end of November Dr. Armand Viré, the ardent explorer of

French caves, most kindly presented me with three specimens of

Stenasellus virer captured in August 1904. An examination

of these specimens showed that the abdomen has only two free

segments in front of the large “ pleotelson,” and that the animal,

in spite of some differences, is rather allied to Asellus in the

structure of the mouth-parts and the pleopoda. J communicated

my conclusions to Dr. Viré, who allowed me to make the necessary

corrections in the proof, for which I beg him to accept my sincere

thanks. I will therefore state that Stenasellus differs from other

Asellota in having the head fused with the first thoracic segment, in

having the two anterior abdominal segments well developed, while

these are rudimentary in the Asellota (for instance in Stenetrium),

and in a few other points. It must, in my opinion, be established

20*

308 DR. H. J, HANSEN ON THE [ Nov. 29,

as a subfamily of the Asellidee, but I will leave to my friends

Dr. Armand Viré and Mr. Adrien Dollfus the further examination

of the structure of this most interesting type.

Having thus discarded Stenasellus, and omitting Asellus

(Mancasellus, Cacidothea) and Stenetriwm, we shall now consider

the remaining portion of Asellota, which comprises, I think,

about thirty genera, twenty of which are found in Norway.

Sars divides all Asellota into five equivalent families: Asellide,

Taniride, Munnide, Desmosomide, and Munnopside. A perusal

of Sars’s account and of Beddard’s ‘Challenger’ work will show that °

the genera which have been—or must be—referred to the four

latter families present great differences in the shape of the body,

in length and shape of the thoracic legs and their coating of

spines or sete, in the degree of development of the uropoda. and

similar features, but of more essential differences between these

four families scarcely one is to be found. The peduncle of the

antenne is six-jointed; the exopod is sometimes rather large,

sometimes rudimentary or absent. In the shape of the mandibles

the differences between the genera decidedly allied to each other,

and by Sars referred to his Munnopside, are considerably larger

than those which can be pointed out between the families. The

other mouth-parts present no difference worth mention. The

thoracic legs show frequently excellent generic characters, but the

differences are so gradually developed that they are valueless as

distinguishing characters between the families. I will refer the

reader to the good figures given by Sars of the posterior pairs of

legs and their development as natatory organs in Hehinopleura,

Desmosoma, Pseudarachna, and Munnopsis. 'The two former

genera are referred to Desmosomatidx, the two latter to Mun-

nopsidee ; but the development of the legs as natatory organs is

gradual, and the difference between these legs in Pseudar achna

(with their seventh joint long) and Munnopsis (with seventh

joint wanting) of the same family is conspicuously larger than

between Pseudarachna and Desmosoma, which are “agiten vec to

different families. The differences in the uropoda are only of

generic value.—The pleopoda show great uniformity in the

genera and families, but must be treated more in detail.

In the females of the four families recognised by Sars, the

lower side of the abdomen, a more or less broad margin excepted,

and the three posterior pairs of pleopoda are covered by a more

or less vaulted operculum which does not show any suture; it is

the first pair of pleopoda. The second pair is wanting. The third

pair has always both rami; the exopod is sometimes small and

unjointed, sometimes larger and two-jointed, in most cases it is

situated along the margin of the endopod, but sometimes it

overlaps a smaller or larger portion of this plate; furthermore,

the difference in size between endopod and exopod is always at

least considerably smaller than in Asellus or Stenetriwm, and the

two exopods do not constitute together a kind of operculum as in

the two last-named genera. The fourth pair of pleopoda possesses,

1904. ] ASELLOTA-GROUP OF CRUSTACEANS. 309

at least generally, perhaps always, both rami, while the fifth pair

has never more than one ramus, in all probability the exopod.

In the males of the same four ‘“ families” the two anterior pairs

of pleopoda constitute together a kind of large operculum, which

consists of three separate plates coupled together ; generally this

operculum covers completely the three posterior pairs of pleopoda,

but in an undescribed form—rather similar to Janira—tfrom the

Southern Atlantic it reaches beyond the hind margin of these

pairs, but laterally the major portion of the exopod of the third

pair is left uncovered ; it may be added that this exopod is longer

but narrower than the corresponding endopod, and does not cover

half of its area. The central plate of the operculum (Pl. XXI.

figs. 4 & 5) is long, of various breadth, with the lateral margins

more or less concave; it has a conspicuous suture along the

middle, and is more or less cleft at the end; each half consists of

the same parts as in Stenetriwm, viz., an unjointed sympod (s.)

and an unjointed ramus (7.). The twosympods are very long and

coalesced with their inner margins; each of them has the posterior

lateral angle produced so that a triangular more or less deep

incision is seen between their distal parts. The ramus mentioned

is attached to the oblique or sinuate posterior margin of this

produced portion, often, as in Janira, rather well marked off

from it, sometimes, as in Hurycope gigantea G. O. S., fused

with it so completely that a limit between them can be traced

only at their distal end. The rami are not coalesced with each

other, but are at most united by membrane in their proximal part.

Each lateral part of the operculum consists of a large plate with

the outer margin convex, the inner nearly straight or somewhat.

concave: this plate is the distal joint of the sympod, which has

the two movable rami attached to the distal part of the inner

margin, and contains muscles for their movement; in Janira 1

found, besides, a very short part which, I think, must be a

proximal joint of the sympod. The endopod is rather slender,

strongly geniculate, typically two-jointed ; the basal joint is

directed forwards, contains a muscle to the second joint, and at

least sometimes, as in Hurycope gigantea G. O. 8. (Pl. XXT.

fig. 6, 6.) it is divided again into two joints. The distal joint is

directed backwards, curved and always produced into a point ;

often it is long, with the distal part extremely slender; in

Munnopsis typica M. Sars it is even more than half as long as

the whole animal, reaching far beyond the abdomen, and this

uncovered portion is setiform, The joint contains a pear-shaped

or very oblong cavity (Pl. X XI. fig. 6, ¢.), which continues into

a narrow duct (d.) opening at the end of the joint. The

exopod (ex.) is very short, two-jointed ; the distal joint is shaped

as a hook, the function of which is to couple the appendage with

the sympod of the first pair of pleopoda; on the upper (posterior)

side of the sympod an impression and a ridge is formed for the

reception of this hook. The three posterior pairs of pleopoda in

the male are exactly as in the female.

310 DR. H. J. HANSEN ON THE [ Nov. 29,

Before concluding this account an apparent exception may be

mentioned. In Sars’s work pl. 50 is filled with drawings of

Nannoniscus oblongus G.O.8. The author figures two animals

which he believes are female and male of the same species. On

the figure representing the male abdomen from below is seen a

large undivided operculum. In the text he says (p. 120): “It is

a very remarkable fact, that the operculum in neither of the two

specimens examined showed any trace of the usual transforma-

tion, though the male character of the specimens otherwise could

easily be demonstrated, both by the greatly projecting sexual

prominence, and by the presence of well-developed testes shining

distinctly through the integuments in their usual place. In the

Caspian species, on the other hand (of which as yet only a solitary

male specimen is known), the sexual characters were quite

normally displayed.” But such differences between the males of

species belonging to the same genus do not exist; the second

pair of pleopoda with its complex organisation for copulation is

not wanting in the male of one species, and highly developed in

the male of another species of the same genus. What Sars

considers to be the male of JV. oblongus is in reality a female of

another species: I cannot account for the nature of the structure

interpreted by him as testes, but the large spine in front of the

operculum has nothing to do with the ‘sexual prominences”

of the seventh thoracic segment ina male. I may add that Iam

very well acquainted with the genus Vannoniscus ; chiefly from

the ‘ Ingolf’ our Museum possesses examples of about ten species,

all with the globular or ovate organ at the end of the antennule

also found in the two species described by Sars as WV. oblongus

G.0.8:

V. Morphological Interpretation of the Pleopoda in Asellota.

In the three preceding sections the pleopoda and their parts

are mentioned as if the names applied had been generally used

or accepted by carcinologists, but it is far from being so. Some

of the interpretations are new, others not generally accepted ;

for these reasons it may be useful to give a comparative review

of this subject.

I must admit that I have not looked through many of the

descriptions of pleopoda scattered in the literature of the last

fifty years or more in order to be able to point out that an author,

in the description of a genus or a species, might have proposed

one or another of my interpretations; but I am sure that the

major part of them are either new or set forth in some of my

earlier papers. In the account of the Crustacea in ‘ Dijmphna-

Togtets zoologisk-botanisk Udbytte,’ Kjébenhavn, 1887*, I gave

a detailed description (with figures) of Hurycope gigantea G. O.8.;

on p. 202 I stated that the three parts of the male operculum are

respectively the fused first pair and the endopods of the second

* I had received and distributed separate copies of my paper in this work by the

aniddle of July 1886.

1904.] ASELLOTA-GROUP OF CRUSTACEANS. BAe

pair of pleopoda; the latter interpretation is not correct, each

lateral plate being not the endopod but the sympod, bearing its

two rami; furthermore, I described the three-jointed copulatory

organ and the two-jointed hook, but did not perceive that they

are the rami of this appendage. In ‘ Isopoden, Cumaceen und

Stomatopoden der Plankton-Expedition,’ 1895, I wrote (p. 6) :—

“Teh habe Janira Leach, Jolanthe Bedd., Zaera Leach, Munna

Kr., Pleurogonium G. O. 8., Macrostylis G. O. 8., Munnopsis

M. Sars und Hurycope G. O. 8., welche Sars in seinen 3 Familien

vertheilt, sammt A sellus Geoffr. und eine vermuthlich zu Stenetriwm

Hasw. gehdrende westindishe Form, untersucht. Ale die erst-

genannten 8 Gattungen weichen nun griindlich in dem Bau der

Pleopoden von den zwei letztgenannten ab, die sich ziemlich

nahestehen. Bei den ersten 8 Gattungen findet man folgenden

Bau: Bei dem Mannchen bildet das 1. und 2. Paar Pleopoden

zusammen einen grossen, festen, aus drei Theilen bestehenden

Deckel, der vollstindig die drei folgenden, zum Athmen einge-

richteten Paare bedeckt (der Deckel entsteht dadurch, dass das

1. Paar zu einer schmiileren Mittelplatte zusammen gewachsen ist,

wihrend das 2. Paar die breiten, mit Paarungsorganen versehenen

Seitenplatten bildet); bei dem Weibchen bildet das 1. Paar

einen miachtigen, ungetheilten Deckel fiir das 3. bis 5. Paar,

wihrend das 2. Paar giinzlich fehlt. Bei Asellus und Stenetrium

wird der Deckel bei beiden Geschlechtern von ganz andern

Elementen gebildet, namlich von den Aussenisten der 3. Paare

von Plecpoden; diese Ausseniiste sind nimlich zu miachtigen

Platten entwickelt, die in der Mittellinie zusammenstossen und

vollsttindig den kleinen Innenast und die beiden folgenden Pleo-

podenpaare decken; bei dem Minnchen befinden sich vor diesem

Deckel 2 kleine, freie Pleopodenpaare, das 2. Paar mit den

Paarungsorganen; bei dem Weibchen fehlt das 2. Paar, wahrend

sich das 1. Paar in Form von 2 kleinen, freien Platten vorfindet.”

The last sentence in this quotation is erroneous as to Stenetriwm ;

otherwise the whole passage is correct, so far as it goes.—In his

recent work on the Isopoda G. O. Sars says (p. 96) :—‘“ A closer

examination of this compound operculum [in the males of most

Asellota] will, however, soon show, that the suggestion at first

put forward by Dr. Hansen is quite correct.” He then gives an

abstract of my results, already quoted here, and continues: “ By

such an explanation, indeed, more uniformity is obtained, as to

the number of appendages of the metasome, which, in fact, is the

very same in all Asellota, viz., 4 pairs in the female, and 5 pairs

in the male, the additional pair constituting the copulative

appendages.” Sars is thus inclined to follow me as to these

questions, but on the plates (39 and 40) with figures of Asellus

and Janira he names the third pair of pleopoda pip’, the fourth

pair ply’; he goes even so far that on the plates (43 and 44)

with Jaera and Munna boecki he marks the third pair pip’, the

fourth pair plp’, &., but this is inconsistent and rather confusing.

Tt cannot be denied that in all Asellota we have five pairs of

pleopoda in the male, but only four pairs in thefemale. Further-

312 DR. H. J. HANSEN ON THE [ Nov. 29,

more, the third pair in the male is shaped exactly as that pair

which in the female follows the operculum; the penultimate pair

in the male is exactly like the penultimate in the female, but differs

from the preceding and from the last pair. We must therefore

conclude that the three posterior pairs in both sexes are homo-

logous. That the undivided operculum found in all genera, Asellus

excepted, in the female is homologous with the first pair in the

male, must be concluded from the fact that in all these genera the

two appendages constituting this pair in the male have their

sympods coalesced or, as in Stenetriwm, completely fused. The

second pair, which in the male bears the copulatory organs, is

therefore wanting in the female.

Next, the interpretation of the parts constituting the two

anterior pairs in the male must be considered. <Asellus presents

the best starting-point. That the two joints of the first pair in

this animal ave respectively the distal joint of the sympod—its

two proximal joints having disappeared—and one of the rami

must, I think, be admitted, and is easily seen from comparison

with Cirolana, ga, &ec., but it is impossible to decide whether the

distal joint, the ramus preserved, is the endopod or the exopod.

The second pair in Aseliws is easy to interpret: each appendage

consists of the sympod with the two two-jointed rami proceeding

from its distal end; no other interpretation is possible, but the

result is that it is the endopod itself which is transformed as a

kind of copulatory organ, with a cavity in the interior of its

distal joint.

Let us, then, look at the first pair of the male in other Asellota.

In Stenetri wm (PL. XX. fig. 29) the sympods are fused, and the

plate thus formed bears two unjointed rami, but, as in Asellus, it is

impossible to decide whether they are the endopods or the exopods.

Comparing this structure with that in Janira (Pl. XXI. fig. 5),

and especially in the undescribed genus (Pl. X XI. fig. 4), it amust

be admitted that the distal pair of lobes marked off by oblique

lines from the long proximal plate must be the rami found in

Asellus and Stenetrium.

Finally, we must consider the second pair of the male in Stene-

triwm and other Asellota. As in Asellus, we find a sympod with

two rami, the essential difference being that these rami proceed

not from the end but from the inner margin of the sympod. The

most distal ramus, which in all genera, Stenetriwm excepted, is

shaped as and performs the function of a hook, is therefore the

reduced exopod ; asin Asellus it is always two-jointed, Stenetriune

excepted, but even ina species of this genus a vestige of a division

into two joints is discernible. The copulatory organ is the

endopod ; asin dsellus it is two-jointed—in Hurycope I found the

basal joint divided again into two joints (Pl. XXI. fig. 6

the distal joint has an internal cavity, Stenetriwm excepted.

(Beddard, in his ‘Challenger’ Isopoda, has already correctly

interpreted the rami as endopod and exopod in Stenetriwm and

Ischnosoma.) It can be added that we have now found the key

to the interpretation of the endopod of the second pair of pleopoda.

1904. ] ASELLOTA-GROUP OF CRUSTACEANS. 313

in the male of other Isopoda. In Jdothea, Spheroma, Cirolana,

Cymothoa, &c., this endopod is generally described as an undivided

plate with an ‘‘appendix masculina” articulated at its mner

margin: this plate is the first, the “appendix” the second joint of

the endopod. This endopod is therefore two-jointed in all Isopoda,

Epicaridea and Gnathiide excepted; but in most forms only the

second joint is transformed, the first being large and lamellar like

that of the first or the third pair, while in Asellota and Oniscide

both joints are narrow.

VI. The Classification of the Asellota.

The tribe or suborder Asellota is very sharply defined from

all other Isopoda, but its subdivision into families isa matter of

considerable difficulty. As already stated, G. O. Sars in 1897

divided the Asellota into five families, but four of these are

far from distinct from each other; moreover, other objections

can be raised. His family Desmosomatide is in reality a rather

mixed company: such genera as MMJacrostylis and Ischnosoma

differ strongly from each other in most features; MVannoniscus

and especially /schnosoma are far from being closely related to

Desmosoma, &ec. I have, for the rest, alre eady, on p. 308, pointed

out several difficulties as to these four families; it may be added

that from the ‘ Ingolf’ we have several new forms which differ

rather or very considerably from the genera of Sars, so that an

attempt at arranging them within his families will aggravate the

state of things. When nature has not worked out groups well-

defined from each other we can of course subdivide a tribe or

suborder into families, founding them on some points in the general

aspect of the animals, but their number and limitation must

then bea matter of personal opinion, and many other authors will

propose the establishment of other or of new families not better

than those first erected. It is, in my opinion, to be preferred to

keep a very large group of genera in the same family, a large number

of species in the same genus, than to subdivide respectively the family

or genus into fanilies and genera with new names, when sharp

lines of distinction are not to be found in nature.

It is well-known that differences in the structure of the

abdominal appendages are among the most important characters

for dividing the order Isopoda into tribes or main-families. That

considerable importance must be ascribed to the above-named

differences in the structure of the pleopoda in Asellus, Stenetriwm,

and other Asellota, will probably be admitted, these differences

being much sharper than those met with in any other external

organ. In the Plankton paper I wrote in 1895 the long passage

quoted above on differences in the pleopoda between Asellus and

Stenetriwm on one side, and several other genera of Asellota

on the other, and continued :—‘“ Es ist anzunehmen, dass alle

existirenden Gattungen in die eine oder dieandere dieser zwei nach

iusserst scharfen Kennzeichen getrennten Gruppen eingefiigt

werden koénnen, welche also die 2 Familien bilden, in welche die

314 DR. H. J. HANSEN ON THE [ Nov. 29,

Asellota am besten getheilt werden konnen.” But two years later

Sars added very much to our knowledge of numerous genera

of Asellota; furthermore, in 1895 I saw a very large number of

undescribed forms—among which several new genera—unknown

to me, and received and studied more closely many examples of

Stenetrium. The question as to the classification of the Asellota

can therefore now be reconsidered on a broader base.

Putting Stenetriwm aside, it will probably be admitted that the

differences in the pleopoda between <Asellus (with MJancasellus

and Cecidothea) and other Asellota would justify the division of

the Asellota into two families. But the structure in Stenetriwn

gives rise to considerable difficulty. Both in Stenetriwm and

Asellus the two anterior pairs of pleopoda in the male and the

first pair in the female are quite small, and overlap only a small

proximal portion of the following pair, the exopods of which are

very large and constitute a complete covering for the respiratory

lamelle ; furthermore, in the second pair of the male the end of

the endopod is blunt and the exopod not developed as a hook for

coupling together the two anterior pairs.

In all other Asellota the first pair in the female is very large

and covers the following pairs completely; in the male the two

anterior pairs constitute together a large operculum formed by

coupling of three plates, which cover the following pairs in their

whole length and, with a single exception, also in their whole

breadth ; the exopods of the third pair are, therefore, generally

invisible, in the instance alluded to partly visible from below at

the side of the operculum, but in this animal (Plate XXT. fig. 5)

they are yet of moderate size, and their inner margin rather

distant from the mesial line; furthermore, in the second pair of

the male the end of the endopod is acute, the exopod hook-shaped

and adapted for coupling. On the other hand, Stenetriwm differs

from Asellus and agrees rather well with other Asellota in some

particulars, viz.: in the male the sympods of the first pair of

pleopoda are fused with each other, and the rami of the second

pair are attached to the inner margin of the sympod; in the

female the pleopoda of the first pair are fused with each other, in

both sexes the last pair has only one ramus. The genus is dis-

tinguished among all other Asellota by the curious feature that

the endopod of the second pair in the male is without an internal

eavity in its distal joint.

That Stenetriwm differs less than Asellus from the other

Asellota is thus easily seen, and the question arises as to the

systematic importance of the differences and similarities. Ought

Stenetriwm to be placed together with Asellus or established in a

family of its own? Considering all particulars, I am now inclined

to prefer the latter alternative. The Asellota will therefore be

divided into three families—Asellide, Stenetriide, n. fam., and

Parasellide, n. fam.

The first-named family comprises the genera Asellus Geoftr.,

Mancasellus Harg.,and Cecidothea Pack.; the second family only

its single genus; the Parasellide all the other genera of Asellota.

1904. | ASELLOTA-GROUP OF CRUSTACEANS. 315

Diagnoses of these three families may now be given :—

A, ASELLIDZ.

First pair of pleopoda in the male small, the sympods free, very

short, together much broader than long, with coupling-hooks

along their inner margins; rami (only a single pair) movable,

much longer than the sympods.—Appendages of same pair in the

female attached rather far from each other, each consisting of a

minute sympod and a circular ramus of moderate size and edged

with setee.—In both sexes this pair overlaps only a small basal

portion of third pair.

Second pair in the male small, situated above and not coupled

with the first pair. Rami attached to the distal margin of the

sympod; endopod not geniculate, its distal joint inflated, containing

a large cavity and its end obtuse; exopod nearly as long as the

endopod, its distal joint movable, lamellar, with marginal sete.

Third pair in both sexes has the exopods very large, touching

each other along the mesial line, and constituting a complete

covering for the endopod and the following pairs ; this operculum

is freely exposed except at the base.

Fifth pair with endopod and exopod well developed.

B. STENETRIIDA.

First pair of pleopoda in the male small, the sympods com-

pletely fused with each other, very short, together much broader

than long; rami movable, much longer than the sympods.—Ap-

pendages of same pair in the female completely fused, constituting a

small oblong operculum without suture or marginal setze.—In both

sexes this pair overlaps only a small basal portion of third pair.

Second pair in the male small, situated above and not coupled

with the first pair. Rami attached to the distal part of the inner

margin of the sympod; endopod strongly geniculate, its distal

joint rather narrow, without internal cavity, and with the end

obtuse ; exopod very short, several times shorter than the endopod,

unjointed (at most with a vestige of a division mto two joints),

narrow, scarcely hook-shaped, at most with a single seta.

Third pair in both sexes has the exopods very large, touching

each other along the mesial line, and constituting a complete

covering for the endopod and the following pairs ; this operculum

is freely exposed except at the base.

Fifth pair with only one ramus, in all probability the exopod.

C. PARASELLIDA.

First pair of pleopoda in the male large ; the sympods coalesced

with each other, together longer than broad, with the lateral

margins concave ; rami immovable, much shorter than the sym-

pods.—Appendages of same pair in the female completely fused,

constituting a very large operculum without suture or marginal

sete.—In the female this pair covers completely the following

pairs; in the male it reaches beyond the distal margin of the

316 DR. H. J. HANSEN ON THE [ Nov. 29,

following pairs, the lateral portions of which are generally com-

pletely covered by the second pair.

Second pair in the male large; the major portion of the

sympods situated outside and coupled with the first pair. Rami

attached to the distal half of the inner margin of the sympod ;

endopod strongly geniculate, its distal joint slender, containing

an internal cavity and distally produced into a point; exopod very

short, many times shorter than the endopod, two-jointed, narrow,

hook-shaped, without sete.

Third pair in both sexes has the exopods of moderate size, not

touching each other in the mesial line, generally completely

covered by the first pair (in the female) or by the two anterior

pairs (in the male); only in the male of a single form their

exterior portion is uncovered.

Fifth pair with only one ramus, in all probability the exopod.

VIL. The Species of the Genus Stenetrium.

The genus was established in 1881 by A. Haswell on two

species, S. armatum Hasw. and S. inerme Hasw. The first-

named must be regarded as the type for the genus; besides it

will be proved below that according to Haswell’s description and

figures of S. inerme this species, in all probability, cannot be

referred to the same genus.

Though I have exainined only one of the five species referred to

Stenetrium by previous authors, I have deemed it useful to incor-

porate them in the analytical key, and to describe them as well as

possible, applying some of the characters found in the descriptions

of the authors, and adding others drawn from their figures, hoping

that these are ‘tolerably correct as to the details in question. It

has not been my intention to mention features showing differences

of slight or no value for the determination of the species.

Conspecius of the Species*.

A. Basal joint of antennule, seen from above,

anteriorly at the outer side produced into an

oblong acute process, or at least (in S. has-

wellii) with a conspicuous tooth marked off

from the oblique front margin by an inden-

tation.

a. First thoracic legs with the upper distal

corner of fifth joint + not produced into a

process.

a. Hand of first legs in both sexes conspi-

cuously more than 3 3 as long as deep...... 1. S. armatum Hasw.

6. Hand of first legs in the male (2) less than

~ as long as “deep Cae one specimen

known) ........ sissscreeercenceceee Oe Se fractum Chilton.

* In the key S. znerme Haswell is omitted; this species is mentioned below after

the descriptions of the other forms.

+ First joint of these legs is fused with the thoracic segment; the following lone

joint, which apparently is ‘the fir st, is here and in the sequel regarded as the second,

according to the morphological interpretation of these legs; in the six other pairs

of thoracic legs the first short joint is, as in all Asellota, ‘not fused with the thorax

but movable.

1904. ] ASELLOTA-GROUP OF CRUSTACEANS. 317

&. First thoracic legs with the upper distal

corner of fifth joimt produced into a long

process.

@. Basal joint of antennulz, seen from above,

anteriorly at the outer side produced into

a long process reaching considerably

beyond the distal margin of second joint. 3. S. mediterraneum, sp. 0.

3. Basal joint of antennule, seen from above,

without any real process, but at the outer

distal angle with a conspicuous sharp

tooth, well marked off from the oblique

front margin by an indentation, and far

from reaching to the distal end of second

FOOLS. “Goo ann sodnde 96 236429 ane doEHSb5s009607002050 4, S. haswellii Bedd.

B. Basal joint of anteunule, seen from above,

with the exterior half of the distal margin

transverse, outer angle at most rectangular

and acute, but without process or tooth.

a. Lateral corner of the head, seen trom above,

produced into an acute process. yes

rather large, oblong.

a. Abdominal shield on each lateral margin

with about five sharp teeth. First tho-

racic legs with the upper distal angle of

fifth joint produced into a long process... 5. S. servatwim, sp. un.

8. Abdominal shield on each lateral margin

with only the tooth at the notch. First

thoracic legs with the upper distal angle

of fifth joint rectangular, without process.

+ In the male the lower margin of the hand

has its proximal half concave, and at

the distal end a low broad process with

three er four teeth nearly equal in size

placed in a convex line. In the female

the angle between the palmar and the

lower margin of the hand measures

about 110°; the hand is a little more

than 2 as long as deep ............--.-----

++ In the male the lower margin of the

hand has its proximal half convex,

and at the distal end two processes

separated by a rather deep incision ;

each process terminates in two teeth.

In the female the angle between the

palmar and the lower margin of the

hand measures about 125°; the hand

is twice as long as deep*.................

+++ In the male the lower margin of the

hand has its proximal half convex, and

at the distal end a narrow, moderately

long process, with the end bifurcate,

and sometimes besides a feeble tooth

on its proximalmargin. Inthe female

the angle between the palmar and

the lower margin of the hand mea-

sures less than 100°, and the hand is

a little more than 2 as long as deep ... 8. S. antillense, sp. n.

}. Lateral corner of the head, seen from above,

without any process. Eyes small, sub-

CINCUIAL eee eee eee etc corona oeeetnes

6. S. occidentale, sp. n.

7. S. stebbingii, H. Richardson.

9. S. siamense, sp. uv.

nN —__—__<_<<$<$<$__—LK<———

* AJl these characters have been derived .from the figures given by Miss Harriet

Richardson.

318 DR. H. J. HANSEN ON THE [ Nov. 29,

1. STENETRIUM ARMATUM Hasw. Ovigerous female (and adult

male, after Haswell*), (Plate XIX. figs. 1 a1 d.)

1881. Stenetriwm armatum Haswell, Proc. Linn. Soc. New South

Wales, vol. v. p. 479, pl. xix. fig. 1 [teste Haswell].

1882. Stenetriwm armatum Haswell, Catal. Austral. Stalk-

and Sessile-eyed Crust. p. 308.

1884. Stenetrium armatum Haswell, Proc. Linn. Soc. New

South Wales, vol. ix. pp. 1009-1010, pl. li. figs. 1-12

[teste Zool. Rec. and Beddard].

Head has its upper surface—the frontal plate excluded—

more than twice as broad as long; the lateral part is expanded

and flattened, the anterior corner produced into a rather long

acute process with a minute tooth on the outer margin; the front

margin outside the base of each antennula produced into a rather

large, triangular, very acute process. Hyes semilunar, long, ob-

lique, with the posterior outer margin rather close to the lateral

margin of the head.

Antennulee have the second joint of the peduncle slightly shorter

than the third and rather thick; flagellum of the female is much

shorter than the peduncle, with about seven joints.

Antenne have the basal joint, seen from above, anteriorly at

the outer side produced into a rather long acute process reaching

slightly beyond the end of the second joint and with a small saw-

tooth on the outer edge. .

First thoracic legs.—In the female the distal half of the upper

part of third joint is expanded, compressed, and produced into a

long curved process ; nearly the whole upper side of fourth joint

is expanded, compressed, and produced into a long process

directed forwards; fifth joint with upper distal angle subrect-

angular and without process, lower angle rounded. The hand

_ not fully twice as long as deep; upper margin rather convex,

lower margin at least as long as the depth, straight, with many

long set; the angle between the lower margin and the palmar

edge measures nearly 110°, is somewhat rounded, with a long

strong spine; palmar edge nearly straight, with several thick

sete serrated along their upper margin. Seventh joint with the

short claw claw-shaped ; the lower margin of this joint armed

with a close row of short spines with a few saw-teeth along the

lower margin.—In the male third, fourth, and fifth joints in all

probability as in the female, the hand is, according to Haswell’s

figure, oblong as in the female, but it has two deep incisions in

the palmar edge, and the process between them is bidentate ; the

‘claw ” reaches a little beyond the palm.

Abdominal shield is somewhat broader than long. Each lateral

margin has about four obscure saw-teeth, besides the usual

rather long tooth at the notch. The posterior margin is strongly

convex and a little sinuate.

* Mr. R. I. Pocock has kindly traced for me the figures given by Haswell in his

first paper.

1904. ] ASELLOTA-GROUP OF CRUSTACEANS. 319

Uropoda considerably less than half as long as the abdominal

shield ; the endopod somewhat longer than the exopod.

Length of a female with marsupium 5-2 mm.; Haswell gives

the length 4 inch.

Occurrence. According to Haswell this species has been captured

on the south-eastern coast of Australia: Port Jackson, Port

Stephens, Griffiths’ Point (Victoria). Of a specimen in the

British Museum (from Griffiths’ Point) I have figured the abdomen

from below; two specimens (from Port Jackson), belonging to

the Museum in Dundee, have kindly been forwarded me, and my

three other figures of this species were taken from one of these

specimens, an almost full-grown female.

Remarks. This species is easily distinguished from all following

forms, S. fractwm Chilt. excepted, by the shape of the head and

basal jomt of antennee, together with the joints of first thoracic

legs. Its differences from S. fractum are mentioned in the

following description of this species.

2. STENETRIUM FRACTUM Chilton.

1887. Stenetrium fractum Chilton, Transact. and Proc. New

Zealand Institute, 1883, vol. xvi. p. 249, pl. xviii.

figs. 3 a-f.

Chilton described and figured a single specimen, the body of

which had been ‘‘ much crushed.” Unfortunately, he says nothing

as to the shape of the head; but judging from the antenne and

the shape of abdomen, I think that the species must be related

to S. armatum Hasw. Most of the characters given below have

been selected among his statements, other characters have been

derived from his figures*.

Antennule.— First joint of the peduncle large, as broad as

long ; second equal in length to the first, but more slender; third

rather longer than the second, ... flagellum about half as long

again as the third joint of peduncle, consisting of about five

joints ....”

Antenne have the basal joint ‘produced acutely at its extero-

distal angle” ; according to fig. 36 the process does not reach the

end of second joint.

First thoracic legs.—Fourth joint much expanded above and its

upper corner produced into a rather long triangular process, which

seems to be a little shorter and thicker than in §. armatum.

Fifth joint shaped about as in the last-named species. Hand very

deep, about #as long as deep, thus proportionately considerably

deeper than in the female of S. armatwm; upper margin as in

that species ; lower margin seems to be a little convex ; the angle

between this margin and the palmar edge almost 120°; palmar

margin straight, armed with thick sete pectinate along their

upper margin and a spine of very moderate length at the lower

angle. Seventh joint and claw as in the female of S. armatum.

* My friend Dr. W. T. Calman has kindly sent me a copy of Chilton’s description

and tracings of his figures.

320 DR. H. J. HANSEN ON THE [Nov. 29,

Abdominal shield—Lateral margins “irregularly serrate,”

“ending posteriorly in a sharp point followed by a small concave

indentation”; the posterior margin almost as in S. mediterranewm ;

its middle portion is somewhat produced, so that a rather low

rounded lobe or protuberance is formed, and almost each lateral

half of the margin is somewhat concave.

Uropoda, according to fig. 3, much less than half as long as

the abdominal shield; endopod considerably longer than exopod.

Length “about 4 inch.”

Occurrence. Lyttelton Harbour, New Zealand.

Remarks. Chilton supposes that his specimen was a female ;

judging from the extreme depth of the prehensile hand, I think it

was a male. It is distinguished from S. armatwm Hasw. by a

shorter process from the basal joint of the antenne, by the

serration of the lateral margins and the shape of the posterior

margin of abdomen, and by the extreme depth of the prehensile

hand in the sex described.

3. STENETRIUM MEDITERRANEUM, sp.n. Adult male and ovigerous

female. (Plate XIX. figs. 2 a—2h.)

Head has its upper surface—the frontal plate excluded—not

fully twice as broad as long; the lateral part is strongly expanded

and flattened, the lateral corner produced into a rather long acute

process without distinct tooth on the outer margin; the front

margin outside the base of each antennula produced into a mode-

rately small, triangular, acute process. Eyes long, oblique, rather

narrow but broader behind than in front; their outer margin 1s

strongly convex, the inner concave; posterior part of their outer

margin rather close to the lateral margin of the head.

Antennule have the second joint of the peduncle moderately

slender and a little shorter than the third ; flagellum in the male

somewhat shorter than the peduncle, with eleven or twelve joints,

in the female considerably shorter than the peduncle, with about

nine joints.

Antenne have the basal jomt, seen from above, at the outer

side produced into a long, rather narrow, acute process reaching

far beyond the distal end of second joint ; external margin of the

process with a couple of long sete inserted at a minute saw-tooth

in front of the middle.

First thoracic legs rather similar in both sexes, of moderate

length. Third, fourth, and fifth joints with the upper part

strongly expanded, compressed; the expansion begins rather near

their base, and is in front produced into a triangular acute process,

which is rather broad and moderately long on third and fourth

joints, somewhat longer and much narrower on the fifth. In the

male the hand is somewhat less than twice as long as deep; upper

margin somewhat convex, with a few shorter sete; lower margin

as long as the depth, straight, with numerous very long hairs;

the angle between lower margin and palmar edge measures about

120°; palmar edge is a little sinuate, with a moderately large,

1904. ] ASELLOTA-GROUP OF CRUSTACEANS. 321

rounded tubercle below its middle and a couple of minute tubercles.

or denticles above the large tubercle; the edge is besides furnished

with some robust sete, increasing much in length downwards and

pectinate along their upper margin ; lower end of palmar edge

armed with a strong long spine, the structure of which is shown

in fig. 2e; seventh joint with its short claw claw-shaped, reaching

slightly beyond the palmar edge, along its lower margin with fine

setee and a row of small strong spines adorned with a few saw-

teeth on their lower margin.—In the female the hand is a little

smaller and a little shorter in proportion to the depth than in the

male, but it differs especially in the palmar edge, which is feebly

convex and quite without tubercles; seventh joint and claw as in

the male.

Abdominal shield a little broader than long. Each lateral

margin with four or five minute spines placed at rather long

intervals, and terminating in the usual triangular tooth at the

conspicuous notch; behind this notch a minute indentation is.

observed. Posterior margin has its middle portion produced so

that a rather low rounded lobe is formed, and almost each half of

the margin is moderately concave.

Uropoda considerably more than half as long as the abdominal

shield; exopod slightly longer than syne but considerably

shorter than endopod.

Distal joint of the endopod of second ak pleopoda unusually

slender and not widened at or beyond the middle; a short ter-

minal portion only half as broad as the remainder, ‘with the end

cut off transversely and without any brush.

Length of the largest male 5°5 mm., of an ovigerous female

6 min.

Occurrence. Some specimens were taken by the author at

Siracusa, Sicily, in depths from 12 to 25 fathoms; four specimens.

were secured by the Danish botanist, Dr. Borgesen, at Ajaccio,

Corsica.

Remarks. This fine species is easily distinguished by the very

long process on the basal joint of antenne, by first thoracic legs in

both sexes, and by the shape and armature of abdomen.—None of

the forms mentioned in Carus’s ‘ Prodromus Faunz Mediterranese’”

ean be referred to Stenetriwm, and the present species seems to be

new, though it is probably widely distributed in the western half

of the Mediterranean.

4, STENETRIUM HASWELLII Bedd.

1886. Stenetriuvm haswelli Beddard, Proc. Zool. Soc. London,

1886, p. 103.

1886. Semarian haswelli Beddard, Isopoda ii. in ‘ Challenger’

Rep. vol. xvii. p. 9, pl. iv. figs. 1-8.

The only specimen hitherto known is a male described and

figured by Beddard. From his long description most of the

characters given below have been selected; some of my state-

ments have been derived from his figures, and from two sketches

Proc. Zoou. Soc.—1904, Vou. II. No. XXI. 21

322 DR. H. J. HANSEN ON THE [ Nov. 29,

kindly dvawn for me from the type specimen in the British

Museum (Natural History) by Dr. W. 'T. Calman.

Head.—The lateral part is exceedingly expanded and anteriorly

produced into a very large, broad and long, triangular, acute

process, reaching forward nearly as far as the front margin of

second joint of the antenne; the front margin outside the base of

each antennula produced into a rather broad, moderately long,

triangular acute process. Eyes long, very narrow, feebly curved,

oblique but essentially transverse, with their posterior end rather

far from the lateral margin of the head.

_ Antennulz have the second joint of the peduncle rather slender

and somewhat shorter than the third; flagellum very much longer

than the peduncle, consisting of numerous joints.

Antenne have the basal joint, seen from above, rather small,

somewhat oblique; without any real process, but at the outer

distal angle with a conspicuous sharp tooth, well-marked off from

the oblique front margin by an indentation, and far from reaching

to the distal angle of second joint.

First thoracic legs elongate, slender, but widening distally, with

a large hand. Third joint long, with the distal part of the upper

side considerably expanded and produced into a rather long,

oblong-triangular process directed essentially forward ; fourth and

fifth joints each considerably shorter than the third, but the distal

half of the upper side is more expanded and produced imto an

oblong-triangular acute process, which is long on the fourth, very

long on the fifth joint. The hand is (according to Beddard’s fig. 4)

large and very deep, not fully 4 as long as deep; upper margin is

strongly convex and furnished with very long hairs on its distal

half; lower margin is straight, with long hairs; the angle between

lower margin and palmar edge measures about 100°, but is some-

what rounded ; palmar edge a little shorter than the lower margin,

straight, with a “row of serrate spines, below which a few fine

slender hairs,” and at its lower end a stout but moderately short

spine. The “claw” of normal size and shape, with serrate spines

along its lower edge.

Abdominal shield nearly as long as broad ; according to sketches

and notes by Dr. Calman, the lateral margin, though ‘a good

deal chipped and broken,” “‘is very finely serrated, at least in

places,” and terminates behind in a small tooth at the usual

notch ; posterior margin shaped almost as in S. serratum.

Uropoda, according to Beddard’s figure, more than half as long

as the abdominal shield; rami, according to his text, “subequal

in size.”

Distal joint of the endopod of second male pleopoda slender and

not expanded at or beyond the middle, terminating in a small

brush.

Length of the single male specimen 16 mm.

Occurrence. ‘Challenger’ “Station 320, off the Rio de la

Plata, February 14, 1876; lat. 37° 17'S., long. 53° 52’ W.; 600

fathoms.”

1904.) ASELLOTA-GROUP OF CRUSTACEANS. 323

Remarks. This deep-sea species is the largest form of the genus

hitherto known; it is easily distinguished from all other species

by the very large lateral processes of the head, the shape of the

eyes, of the basal joint of antennz, and of the first thoracic legs.

5. STENELRIUM SERRATUM, sp.n. Ovigerous female. (Plate XTX.

figs. 3a-3d; Plate XX. fig. 1 a.)

Head has its upper surface (the frontal plate excluded) nearly

twice as broad as long; the lateral part is somewhat expanded

and flattened in front, and produced into a moderately large acute

process; the front margin outside the base of each antennula

produced into a rather large process, which is broad at the base,

while its distal part is shaped as a narrow acute hook curved

somewhat inward. Hyes large, oblong, very oblique, the outer

margin extremely convex, the inner very concave; their posterior

part overlaps the lateral margin of the head itself.

Antennule have the second joint of the peduncle moderately

robust and somewhat shorter than the third; flagellum 9-jointed,

as long as the sum of the two proximal joints of the peduncle.

Antenne have the basal joint, seen from above, distally cut off

transversely, its outer angle acute, but not produced into any

process.

First thoracic legs rather short. Third joint distally widened

but without process; fourth jomt with the upper part nearly

from the base strongly expanded, compressed and distally pyro-

duced into a process of moderate length and breadth; fifth joint

similarly expanded and distally produced into a long slender

process. Hand a little more than twice as long as deep; upper

margin rather feebly convex, with a few sete; lower margin

straight, only half as long as the upper, with numerous very long

hairs ; distal end as long as the lower margin; palmar edge very

oblique, a little simuate, furnished with six very stiff sete, pro-

portionately long and increasing in length downwards, pectinate

along their upper margin, and at the end of the edge a moderately

robust very long spine pectinate as the sete; the angle between

palmar edge and lower margin measures about 130°. Seventh

joint with its claw claw-shaped, reaching a little beyond the lower

end of the palmar edge; the joint is adorned below with serrated

spines and fine curved sete as in S. mediterraneum, but the spines

are less numerous, only about ten.

Abdominal shield is somewhat broader than long; each lateral

margin is adorned with five small nearly spiniform processes,

increasing in size backwards and placed at regular intervals, the

last of these processes being that at the usual notch. Posterior

margin is rather evenly but moderately curved.

Uropoda wanting,

Length of the single adult female 6 mm.

Occurrence. West Indies: St. Thomas, one specimen (A’rebs).

Remarks. This species is easily distinguished from S. armatum,

S. fractum, and S. mediterraneum by the absence of a process

o]*

324 DR. H. J. HANSEN ON THE [Nov. 29,

from the basal joint of antenne. The processes from the front

margin of the head are more produced and much more curved,

the processes or teeth on the lateral margins of abdomen con-

spicuously longer than in any other species hitherto discovered. »

6. SrENETRIUM OCCIDENTALE, sp.n. Adult male and ovigerous

female. (Plate XX. figs. 2 a-2 7.)

Head shaped as in S. antillense (Pl. X XI. fig. 16); its upper

surface (the frontal plate excluded) is considerably less than

twice as broad as long; the lateral part, seen from above, is

feebly expanded and produced into a small acute process; the

front margin outside the base of each antennula produced into a

broad but rather low process, with the end obtuse. Eyes of

moderate length, oblong, somewhat curved, very oblique and con-

siderably removed from the lateral margin of the head.

Antennule with the second joint slightly longer than the third,

moderately robust; flagellum in the male 9-jointed and as long

as the sum of the two distal joints of peduncle, in the female still

shorter, with four or five joints.

Antenne have the basal joint distally cut off transversely, its

outer angle without process and measuring about 90°.

First thoracic legs very different in adult specimens of the two

sexes, but in immature males nearly as in adult females.—In the

male they are rather long, robust; third joint is distally much

expanded above and produced into a triangular process directed

upwards; fourth joint expands above gradually from the base,

forming a broad but rather low process, a portion of the inner

surface of which is furnished with numerous exceedingly long

hairs. Fifth joint has the upper margin very short, without any

process, but it expands below, its lower margin is several times

longer than the upper, and besides it is produced into a very long

oblong-triangular process, the inner side and both margins of

which are closely set with long or very long hairs; the upper

margin of the process is straight nearly to the insertion of the

hand, and the distance from this insertion to the end of the pro-

cess is longer than the distance from the insertion to the base of

the joint. The hand is very large, a little broader near the end

than at the base, two and a half times longer than deep; upper

margin strongly convex, lower margin rather concave from the

base to the distal process, which occupies the major portion of

the short palmar edge; this process is low, broad, its margin more

or less convex and divided into three or four teeth; the lower

major portion of the inner surface of the hand is closely set with

very long hairs. Seventh joint very long, much curved, especially

at some distance from the base, claw-shaped, with fine simple

hairs spread along both margins and on the mner side, but with-

out spines; the claw itself is very short.—In the female the legs

are much shorter than in the male, robust; process on third joint

proportionately a little longer and broader, that on fourth joint a

little longer than in the male; fifth joint much smaller than in

(1904, ] ASELLOTA-GROUP OF CRUSTACEANS. 325

the male, its lower process small. Hand much smaller than in

the male, subtriangular, a little more than half as long again as

deep; upper margin very convex, two and a half times longer

than the lower, which is straight, with many long hairs; distal

end somewhat longer than the lower margin; palmar edge feebly

convex, with an angular notch at the lower end, so that the usual

spine, which is strong and moderately long, is situated a little

behind the edge; the edge from the “claw” to the notch is

occupied by five or six saw-teeth gradually increasing in size

downwards, and besides adorned with some stiff sete pectinate

along their upper margin ; finally, the angle between lower margin

and palmar edge measures about 110°. Seventh joint with the

claw regularly claw-shaped, when extended reaching slightly beyond

the notch mentioned; the major portion of the lower margin of

the joint is adorned with rather slender spines, serrate along the

lower margin, and some fine hairs.

Abdominal shield slightly broader than long; lateral margin

unarmed, only with the usual tooth and notch at the end;

posterior margin, reckoned from the notch, is strongly and rather

evenly curved.

Uropoda considerably more than half as long as the abdominal

shield ; exopod slightly longer than the sympod and much shorter

than the endopod.

Second joint of the endopod of second male pleopoda with the

distal half considerably broader than near the base, the end very

obliquely rounded, the lower surface at the end set with numerous

very short hairs.

Length.—Both sexes similar in this respect, measuring about

373 mm.

Occurrence. West Indies: St. Thomas. Several specimens,

among which five adult males, were taken, 12.11.1888, by H. Kier,

Captain i in the Danish Navy.

Remarks. In the outline of the head, the position of the eyes, the

short third joint in the antennular peduncle, the first pair of legs,

and, above all, in the shape of the hand in the male, this Sorin

differs abundantly from the preceding species ; it 1s closely allied

to the two following species, and the differences are mentioned

below.

7. STENETRIUM STEBBINGII H. Richardson.

1902. Stenetriwm stebbingi Harriet Richardson, Trans, Conn.

Acad. vol. xi. p. 295, pl. xxxix, figs. 46-49.

Of this species I have seen no specimens. According to the

description and the figures published by Miss Richardson it is

very closely allied to S. occidentale and S. antillense, but the

figures representing the first thoracic leg in male and female

show some differences which I hope really exist, and if so, they

are sufficient for the separation of this form from S\. occidentale

and S. antillense.

The figure showing the head with antennule and four proximal

326 DR. -H, J. HANSEN ON THE [ Nov. 29,

joints of antenne is certainly not correct in several details,

especially in the shape of the eyes and of the basal joint of the

antennee, but, so far as I can see, the head with its upper appendages

does not present differences of importance from those of my two

allied species.

First thoracic legs present real differences. In the male the

process from the lower side of the fifth joint is even longer than

in S. occidentale, and besides not triangular but of about the same

depth for two-thirds of its length. The hand is longer and pro-

portionately more slender than in my two species; it agrees with

S. antillense and differs from S. occidentale in having the proximal

half of the lower margin convex, but it differs from S. antillense

in being nearly three times as long as deep, with the proximal

half not deeper than the distal one; the palmar edge is occupied

by two moderately low, narrow processes with the end bifurcate.

(This description is derived from fig. 48; my statement concerning

the two bifurcate processes does not agree with the author's

description: “.... three large spines, the inner one being bifur-

cate,” but I hope that the figure is correct.) In the female the

lower process on the fifth joint is longer than in S. occidentale :

the hand is, according to fig. 49, nearly twice as long as deep, thus

more slender than in my two species, besides the angle between

the lower margin and the palmar edge is larger, measuring about

125°.

Length has not been stated by Miss Richardson ; according to

the degree of enlargement of her figures, the species must be

larger ‘than S. antillense or S. occidentale.

Occurrence. Bermudas. Many specimens were taken, at least

some of them, “in corallines, at low water.’

8. STENETRIUM ANTILLENSE, sp.n. Adult male and small immature

female. (Plate XX. figs. 3a-37; Plate XXI. figs. 1 ale.)

This species is so closely allied to S. occidentale and S. stebbingti

that it is preferable to point out the differences instead of giving

a complete description.

The head is shaped as in S. occidentale ; the eyes have the same

position.

Antennulz have second and third joints subequal in length;

flagellum in the male about as long as the sum of the two

preceding joints, with from nine to eleven joints.

Antenne have their basal joint as in S. occidentale.

First thoracic legs show some important differences in their distal

half.—In the male the fifth joint is below as much produced as in

S. occidentale, but the process is differently shaped: its proximal

half is expanded above and excavated on the upper half of the

outer side in order to receive the proximal lower part of the hand ;

the oblong-triangular, freely protruding part of the process looks

therefore much shorter than in S. occidentale, in which it is

regularly oblong-triangular and quite free to about the articula-

tion of the hand. The hand is deeper than in SS. occidentale and

1904.| ASELLOTA-GROUP OF CRUSTACEANS. 327,

has a different shape; it is slightly more than twice as long as

deep, conspicuously deeper at a shorter distance from the base

than at the distal end; the upper margin is less convex than in

S. occidentale, while the lower margin is considerably convex in.

its proximal and concave in its distal half; the distal process is

longer but much narrower than in JS. occidentale, bifurcate at

the end and sometimes with a feeble tooth on its posterior margin ;

distribution of hairs as in the species mentioned. The “claw,”

formed by the seventh joint and the claw itself, slightly longer than.

in S. occidentale and more hairy at the lower margin.—In the

young—probably also in the adult—female the hand differs some-

what in shape from those of the two preceding species: as in

S. occidentale it is a little more than half as long again as broad,

but the lower margin is comparatively longer, measured to the

base of the lower saw-tooth of the palmar edge slightly more than

half as long as the upper margin, and the angle between the

lower margin and the palmar edge is less than 100°; the notch at

the distal end of the lower margin is longer than in S. occidentale.

Abdominal shield is slightly longer than broad, otherwise as in

S. occidentale.

Uropoda seem to be only a little more than half as long as the

abdominal shield; the rami—preserved only in the young

female—a little shorter than in S. occidentale.

Length of the largest male 4°5 mm.

Occurrence. West Indies. Two adult males and a young female

were found on corals presented to the Zoological Museum in

Copenhagen by Mr. G. A. Hagemann. The corals were said to

be from “ deep water,” which probably signifies about 100 fathoms.

Remarks. 1t is certain that S. occidentale and S. antillense are

valid species; having seen no specimens of S. stebbingit it has

been necessary to rely on Miss Richardson’s drawings, and I

believe that neither of my species is identical with this form from.

Bermudas. It may be added that the hand of the first legs in the

male is similar to that of the second legs in species of /schyrocerus,

a genus of Amphipoda; furthermore that the hand of the female in

Stenetriwm is stmilar to that of the second legs in species of MJetopa,

another genus of Amphipoda, and in the latter genus specific

differences similar to those pointed out in females of Stenetriwm

are well known.

9, STENETRIUM SIAMENSE, sp.n. Male and not quite full-grown

female. (Plate X XI. figs. 2 a—27.)

Body narrower than in the preceding forms, slightly more than

four times as long as broad.

Head has its upper surface (the frontal plate excepted) half

as broad again as long; the lateral part is not expanded, with the

front portion of the lateral margins converging and the angle

not produced, obtuse; front margin outside the base of each

antennula produced into a broad short triangle with the apex

rounded. Hyes small, rounded, placed near the lateral margin.

328 DR. H. J. HANSEN ON THE [ Nov. 29,

Antennulz have the second joint of the peduncle a little

shorter than the third and rather slender; flagellum in the male

about as long as the sum of the two proximal joints of the peduncle,

six-jointed.

Antennee.—Basal joint is distally cut off transversely, its outer

angle without any process measuring more than 90°.

First thoracic legs very different in the two sexes.—In the adult

male they are robust and elongate, a little longer than thorax.

Third joint is oblong, gradually expanded above and produced

into a rather large, oblong, acute process directed forward; fourth

joint very large, gradually much expanded above and produced

into a large plate-shaped process with the end acute; upper margin

and inner side of the expanded part of third and fourth joints

with numerous exceedingly long hairs. Fifth jomt very large,

without dorsal process but strongly expanded below, forming a

large subtriangular plate which is slightly produced below in

front, with the lower anterior angle rounded and the distal

margin nearly as long as the joint; the upper part of the joint

is distally vaulted on the exterior side and situated in a much

higher plane than the large lower part, which is a thin plate;

in this way a kind of excavation, sharply marked off above,

is formed on the outer side of the joint, and the hand can be

turned downwards and backwards so that. a portion of its inner side

becomes overlapped by the thin plate of the fifth joint ; the lower

margin and inner side of this thin plate are furnished with hairs

and sete of very moderate length. The hand is large, not very

long but very deep, about 4 as long as deep; from the basal articu-

lation it expands suddenly and strongly downwards so that a free

and rather long posterior margin is formed, and the posterior

angle is broadly rounded; the upper margin is feebly convex,

the lower margin somewhat shorter than the distal end; the

palmar edge is very oblique, with two teeth above the middle and

a rather small spine at the lower angle which is produced into a

triangular tooth ; at the palmar edge and on the lower margin a

number of sete of very moderate length. Seventh joint with

the claw claw-shaped, rather small and not reaching the lower end

of the palmar edge; lower margin of the joint with a row of fine

spines.—In the female the first legs are very much smaller than in

the male, not elongate. Third and fourth joints as to shape and

hairs essentially as in the male, yet somewhat shorter in pro-

portion to length; fifth joint similar to that in the other sex,

but the upper angle is produced into a rather short acute process,

and the lower plate-like expansion is narrower as compared with

the upper portion, which is rather feebly vaulted. The hand is

nearly twice as long as deep; the lower margin is rather convex

in its proximal half, but a posterior margin, as found in the male,

is not developed; the palmar edge is very oblique, and the angle

between this edge and the lower margin measures nearly 130°,

the edge shows a rudimentary tooth, several stiff sete pectinate

above, and at the lower end a notch so that the usual strong spine

1904.] ASELLOTA-GROUP OF CRUSTACEANS. 329

is placed behind the main portion of the edge (as in the females of

S. occidentale and S. antillense). The “claw” reaches just to the

end of the edge; its lower margin with a number of fine, scarcely

serrate spines.

Abdominal shield slightly longer than broad. The lateral

margin unarmed, only with the usual tooth at the notch ; posterior

margin strongly and evenly convex.

Uropoda wanting in my specimens.

Second joint of the endopod of second male pleopoda increases

gradually somewhat in breadth from the base for two-thirds of

its length; the distal oval portion is marked off by small lateral

indentations, and the outer portion of its lower surface is adorned -

with a brush of very short fine hairs; the margin turning out-

wards has a small spe directed forwards a little in front of the

indentation named.

‘Length of an adult male 4 mm., of the largest immature female

3°74 mm.

Occurrence. Gulf of Siam: between Koh Mesan and Koh

Chuen, 38 fathoms (one male) and 15 fathoms (two immature

females). The specimens were found in sifted bottom material

taken by Dr. Th. Mortensen in the beginning of February 1900.

Remarks. This small slender species is easily distinguished

from all preceding forms by the small rounded eyes and the com-

plete absence of any process from the lateral margin of the head.

First thoracic legs in the male deviate much from those in all

other forms hitherto known.

10, STENETRIUM INERME Hasw.

1881. Stenetriwm wnerme Haswell, Trans. Linn. Soc. New South

Wales, vol. v. p. 480, pl. xix. fig. 2 [teste Haswell].

1882. Stenetrium inerme Haswell, Catal. Austr. Stalk- and

Sessile-eyed Crust. p. 309.

This species is mentioned only for the sake of completeness,

because I think that it does not belong to Stenetriwm. In his

‘Catalogue’ Haswell describes some characters which raise doubt

as to the correctness of his own reference to this genus, the

type of which is S. armatum Hasw.; tracings of figures* in

his first paper corroborate this doubt, According to his fig. 2,

the first thoracic segment is laterally not produced into an acute

process directed forwards as in all other species; in the text

(Catal. p. 309) he says: “ Lateral borders of anterior thoracic

segments not much produced, bilobed,” which thus agrees with

the figure. Furthermore, the figure of the maxilliped, the de-

scription of the antennule, having their basal joint ‘“ very short

and broad,” and abdomen, having the lateral margin “ entire,”

thus without notch, all these particulars do not agree with features

met with in all other species of Stenetrium ; unfortunately Haswell

omits the pleopoda. On hand and “claw” of first thoracic legs

' * Kindly forwarded to mea long time ago by Mr. R. I. Pocock, now Superintendent

of the Zoological Gardens, London.

330) DR. H. J, HANSEN ON THE [ Nov. 29,

he says: “ propodos subtriangular in outline, the palm transverse,

concave, armed with a few short bristles, and defined by a promi-

nent acute tooth; dactylos much longer than the palm;” this

description agrees well with his fig. 2¢, but hand and especially

the ‘“ dactylos” differ much from the structure in all other species

of Stenetrium, and agree much more with the same parts in males

belonging to a genus allied to Janira, The length is “ about

=3;in.” Haswell’s type is from Port Jackson.

EXPLANATION OF THE PLATES.

Prate XIX.

Fig. 1. Stenetriam armatumn Hasw. (p. 318).

Fig.1a. Head with front end of thorax of a female, from above; X 19. .f,, frontal:

plate; p., process from basal joint of antenna; s., squama of antenna.

16. Distal part of first right thoracic leg of a female, from the exterior side

(from behind) ; Xx 30.

1c. Abdomen of a female, from aboner X19. Sete and uropoda wanting.

1d. Abdomen of a female, from below; X9. a, first pair of pleopoda con-

stituting a small operculum ; 6, sympod of third pleopoda.

Fig. 2. Stenetrium mediterranewm, sp. n. (p. 320).

Fig.2a. Head of an adult male, from above; X 22.

26. Distal part of first left thoracic leg of an adult male, from the exterior

side; X 30.

2c. Palmar edge ss the hand and seventh joint with claw of the leg shown in

fig. 26; xX 5

2d. Subdistal GEES from the lower side of seventh joint of the leg show nin

fig. 26; X 212.

é. Spine from fie end of palmar edge of the lez shown in fig. 2b; 212.

f. Distal part of first left thoracic leg of an adult female, from the exterior

side; X 30.

g. Abdomen of an adult male, from above; X 20.

h. Second left pleopod of an adult male, from in front; xX 45.

Fig. 3. Stenetrium serratum, sp. n.; adult female (p. 323).

Fig.3a. Head, from above; X 22

3. Lateral part of the two anterior thoracic segments with their legs, from

above; X 22.

3c. Distal part of first left thoracic leg, from the outer side (from behind) ; F

x 63.

3d. Distal part of sixth jomt, seventh joint, claw and spine below the claw of

second left thoracic leg shown in fig. 36; X79. s., exceedingly broad

spine inserted at the end of sixth joint on its posterior side, and over-

lapping the proximal portion of seventh joint.

PratE XX.

Fig. 1. Stenetriwm serratum, sp. n. (continued) (p. 328).

Fig.1a. Abdomen of an adult female, from above; X15. Uropoda wanting.

Fig. 2. Stenetriwm occidentale, sp. n. (p. 324).

Fig.2a. Left antennula of an adult male, from above; X 39.

2b. First’ left thoracic leg of an adult male, from the exteri lor side (from:

behind); X 39.

2c. First left thoracic leg of an immature male, from the exterior side; X 39.

2d. First left thoracic leg of an adult female, from the exterior side; x 39.

2 e. Palmar edge and seventh joint with claw of the leg shown in fig. 2d, from:

the interior side; X 97.

2 f. Abdomen of an ovigerous female, from above; X 28.

2g. First pair of pleopoda of an adult male, from in front; 51. m., muscle

to the ramus.

1904.| ASELLOTA-GROUP OF CRUSTACEANS. dol

Fig. 2h. Left pleopod of second pair of the same male, from in front; X 51. The

—

muscles in the sympod are seen. 1908 ;

2 i. Left pleopod of third pair of the same male, from in front; X 51.

2k. Left pleopod of fourth pair of the same male, from in front; x 51.

2 1. Left pleopod of fifth pair of the same male, from in front; X pie

2m. First pair of pleopoda, constituting a small operculum, of an adult female,

from in front; X 451.

2n. Lett pleopod of third pair of the same female, from in front; X 51.

Fig. 3. Stenetrium antillense, sp. n. (p. 326).

.3a. Left mandible of a male, from below; X 49.

3 6. Distal part of the mandible shown in fig. 3a, from below; X 12o.

3c. Distal part of right mandible of the same male, from below; X 120.

3d. Hypopharynx (paragnatha) of the same male, from below; X 49.

3 e. Left maxillula of the same male, from below; X 49.

3 f. Distal part of the lobe from third joint of the maxillula shown in fig. 3.4;

from below; X 185.

3g. Left maxilla of the same male, from below; X 49.

3h. Left maxilliped of the same male, from below; x 49.

3%. Distal portion of first left thoracic leg of an immature female, from the

exterior side; X 53.

Prats XXI.

Fig. La. Stenetrium antillense, sp. nu. (continued) (p. 326).

Fig.1a. Adult male, from above; X 9. Several appendages omitted.

1b. Head of the adult male shown in fig. 1a, trom above; X 24.

1c. First left thoracic leg of the adult male shown in tig. 1 a, from the exterior

side; < 16.

1d. Major part of fifth joint and prehensile hand of first right thoracic leg of

another male, from the exterior side; X 28.

1e. Abdomen of an immature female, from above; X 36.

Fig. 2. Stenetrium siamense, sp. n. (p. 327).

.2.a. Head of an adult male, from above; X 33.

2b. First right thoracic leg of an adult male, from the exterior side (from

behind) ; 33.

2c. First right thoracic leg of a not quite full-grown female, from the exterior

side; X 33.

2d. Palmar edge and “claw” of the leg shown in fig. 2c. from the exterior

side; X 86.

2 e. Abdomen of an adult male, from above; X 28. Uropoda wanting.

2 f. First pair of pleopoda of an adult male, from in front; < 50. m., muscle

to the ramus.

24. Second left pleopod of an adult male, from in front; x 50.

2h. Distal part of the sympod, endopod, en., and exopod, ewr., of the pleopod

shown in fig. 2g, from in front; X 95. In the sympod the muscles for

the movement of both rami are shown.

2 i. First pair of pleopoda, constituting a small operculum, of a not quite full-

srown female, from in front; > 50.

Figs. 3-6. Parts of various Asellota.

Fig. 3. Distal part of sympod with both rami of second left pleopod of the male of

Asellus aquaticus L., from in tront; * 54 (p. 306). Three muscles to the

rami are shown in the sympod; in the first joint of the endopod a muscle to

the second joint is seen, in the distal joint of the exopod a muscle producing

the movement of this joint is plainly visible. The dotted line on the distal

joint of the endopod indicates the outline of the sac within.

4. First pair of pleopoda and second right pleopod of a male of an undescribed

form rather allied to Zaniza, seen from in front; < 43 (p. 309). 7., ramus,

and s., sympod of first left pleopod; ¢., second pleopod, with the short hook-

shaped exopod, while the elongate endopod has been omitted.

5. First pair of pleopoda of a male of Ianira maculosa Leach, from in front ;

X 32 (p. 809). 7., ramus; s., sympod.

6. Distal inner part of left second pleopod of a male of Hurycope gigantea

G. O.Sars, from in front; xX 12. . (p.309), basal joint of endopod divided

into two parts or joints ; c. (p. 309), cavity within the distal joint of the

endopod; d. (p. 312), duct from this cavity; ex., two-jointed exopod.

332 | MR. G. A, BOULENGER ON.THE [Nov. 29,

5. On the Lacerta depressa of Camerano.

By G. A. Boutencsr, F.R.S8., V.P.Z.S.

[ Received October 20, 1904. ]

(Plate XXII.*)

Much work has been done lately on the various forms which

cluster round the common Lacerta muralis of Europe, and the

younger herpetologists have shown an ever increasing inclination

towards multiplying species. Whether this narrower conception of

species will result in a better understanding of the distribution and

phylogeny of this difficult group, is a question I will not at present

discuss. But I may say that these recent systematic attempts

seem to justify the view expressed by Dr. Gunther thirty years

ago, that ‘Such nominal species rarely survive their author ;

but before they are merged again in the synonymy, they are the

cause of much unnecessary trouble, and being founded on slight

individual peculiarities, they are frequently mistaken, rarely

recognised.”

The history of Lacerta depressa illustrates the case in point.

Some timo ago I received from my friend Dr. F. Werner a

lizard from the Bithynian Olympus, near Brussa (altitude 1500—

1800 metres), which he referred to L. depressa of Camerano, and

particularly to the var. rudis of Bedriaga. An account of this

and other specimens of the same form is given by Werner in his

valuable paper on the Reptiles of Asia Minort. This lizard

differs so considerably from the specimen of ZL. depressa (one of

the types, received from the Turin Museum) preserved in the

British Museum, that I could not satisfy myself of the correctness

of Werner’s identification, notwithstanding his express statement

that this was arrived at after comparison of one of the types from

Trebizond, entrusted to him by the Turin Museum.

In order to clear up my doubts, I applied to my friend Prof,

Camerano for the loan of the specimens of ZL, depressa $ preserved

in the Museum under his charge; and my request having been

kindly granted, I wish to lay before the Society the result of my

examination.

This shows that, as could be gathered from Bedriaga’s descrip-

tion ||, the species is made up of several distinct forms (I would

call them varieties). There is no evidence that the specimens all

came from Trebizond, for De Filippi, their collector, himself says

of the species referred to L. taurica ¥, ‘‘trovanno commune da

Trebisonda a Tiflis.” It is remarkable that the descriptions of

both Camerano and Bedriaga, who had access to all the specimens,

* For explanation of the Plate, see p. 339.

+ Introduction to Catalogue of Fishes, viii. p. vi.

t Sitzb. Ak. Wien, cxi. 1902, p. 30, pl. m1.

§ Podarcis depressa, Camerano, Atti Acc. Tori. xiii. 1878, p. 539.

|| Abh. Senck. Ges. xiv. 1886, p. 272.

€ Arch. per la Zool. 11. 1863, p. 386.

oue1dUIeD vssdddad VLYHOV'I

op WEqQoV' 9 | ez ‘ojoyd udaery'¢

TOG OCIS Zict

1904. | LACERTA DEPRESSA OF CAMERANO. aon:

six in number*, though very detailed, do not cover by any means.

the variations which the specimens show. I quite agree with

Boettger + in regarding ZL. depressa and. its varieties as not

specifically separable from the Z. muralis of S.W. Asia, which

embraces the forms named L.chalybdea, L. saxicola, L. portschinskii,.

and L. defilippui. With these forms I propose to deal elsewhere,

and my object on the present occasion is merely to give accurate

descriptions of the individual specimens which are the types of

L. depressa, One of them is in the British Museum, one has

passed into the collection of Dr. de Bedriaga and is not available

to me at present, whilst the four others are preserved in the Turin

pais e if deal with the five specimens in order of size.

1. Male——Snout obtusely pointed; the greatest depth of the

head equals the distance between the eye and the tympanum.

Rostral not entering the nostril, forming a narrow suture with

the frontonasalt; frontal as long as its distance from the end of

the snout ; a complete series of granules between the two principal

supraoculars and the supraciliaries; fourth supraocular divided

into two; parietals once and a half as long as broad, not in con-

tact with the upper supraocular § ; the upper border of the parietal

very slightly concave in front for the accommodation of a rather

large upper temporal shield; occipital half as long, but a little

broader than the interparietal ; temporal scales small and granular;

tympanic and masseteric shields well developed, the latter oval,

oblique, and separated from the upper temporal by two series of

granules; four upper labials anterior to the subocular, the lower

border of which is nearly as long as the upper.

Collar even-edged, composed of 11 plates; 31 scales on a line

between the symphysis of the chin-shields and the median plate

of the collar.

Body much depressed. Dorsal scales roundish-hexagonal, flat,.

faintly keeled on posterior part of back; 60 scales across the

middle of the body, 3 or 4 transverse series corresponding to a

ventral plate, 44 to the length of the head. Ventral plates in 6

longitudinal and 26 transverse series. Anal plate preceded by a

nearly equally large shield, the two plates bordered by a semi-

circle of 9 small plates.

Hind limb reaching the collar. Scales on upper surface of tibia.

rhomboidal, keeled, and a little larger than the dorsals, 8 trans-

verse series of the former corresponding to 10 of the latter.

Femoral pores 22-21. 27 lamellar scales under the fourth toe.

Upper and lateral scales at the base of the tail strongly keeled,

those on the sides raised behind, subtrigonal, squarely truncate ;

the whorls alternately longer and shorter, but not very markedly

* Bedriaga says seven, but this is probably through a Japsus calami.

+ Ber. Senck. Ges. 1892, p. 141.

+ As occurs sometimes 1 in both the f. typiea and the var. bedriage.

& Méhely, Ann. Mus. Hung. ii. 1904, p. 367, has strangely overrated the syatemane

importance of this character, which is inconstant not only in Asiatic examples but

also in the Huropean (numerous exceptions from France, Spain and Portugal, Italy,,

Islands of Mediterranean) and North African.

334 MR. G. A. BOULENGER ON THE | Nov. 29,

<0; the.fouth whorl behind the postanal small scales contains 29

scales.

Back spotted all over; sides darker, with whitish ocellar spots ;

upper surface of head unspotted ; labials black-edged; uniform

white beneath.

9, Female.—Head as in the preceding specimen, but smaller and

less strongly depressed, its greatest depth equalling the distance

between the centre of the eye and the tympanum. Rostral as in

the preceding, but separated from the frontonasal, the nasals

forming a short median suture; frontal, supraciliary granules,

and parietals as in the preceding ; fourth supraocular undivided ;

occipital very short, a little narrower than the interparietal, and

separated from it by an additional shield; masseteric shield sepa-

vated from the upper temporal shield by a single series of granules;

five anterior upper labials; subocular as in the preceding.

Collar even-edged, composed of 9 plates; 25 scales between

the chin-shields and the collar.

Body much depressed. Dorsal scales oval and distinctly keeled ;

51 scales across the middle of the body, 3 corresponding to a

ventral plate, 35 to the length of the head. Ventral plates in

6 longitudinal and 27 transverse series. Anal plate preceded by

a large but narrower plate and bordered on each side by 3 small

plates.

Hind limb reaching the axil. Scales on upper surface of tibia

rhomboidal and strongly keeled, much larger than the dorsals, 6

of the former corresponding to 10 of the latter. Femoral pores

18-18. 25 scales under the fourth toe.

Upper caudal scales very strongly keeled, those on the sides

yaised, subtrigonal, almost spinose, truncate behind ; the whorls

alternately longer and shorter, and the scales of the two median

dorsal series of every other whorl much wider than the others ;

28 scales in the fourth whorl.

'The black spots form two irregular series along the middle of

the back; a dorso-lateral series of white black-edged ocelli; a

large ocellus above the shoulder; black dots on the head; the

labials black-edged ; lower parts uniform white. °

3. Gravid female* —Head very small and even rather more de-

pressed than in specimen No. 1, its greatest depth hardly equalling

the distance between the eye and the tympanum ; snout acutely

pointed. Rostral not entering the nostril and separated from the

frontonasal by the nasals, which form a rather long median suture ;

frontal as long as its distance from the end of the snout; a

complete series of granules between the two principal supraoculars

and the supraciliaries ; parietals once and half as long as broad,

in contact with the upper postocular, the outer border nearly

straight ; three small occipitals in a transverse line; temporal

scales small.and granular ; tympanic and masseteric shields well

* Boettger was therefore not justified in saying (Ber. Senck. Ges, 1889, p. 205) of

L. depress: “Sowohl De Filippi, als auch Camerano, v. Bedriaga und Boulenger

kaunten nur die mehr graugriin oder olivengrau gefarbte Jugendform derselben.”

1904. | LACERTA DEPRESSA OF CAMERANO. 335

developed, the latter oval, oblique, and separated from the upper

temporal shield by one series of granules; four upper labials

anterior to the subocular, the lower border of which is nearly as

jong as the upper.

Collar even-edged, composed of 10 plates; 31 scales between

the chin-shields and the collar.

Body much depressed. Dorsal scales round and flat, perfectly

smooth, 54 across the middle of the body, 3 or 4 corresponding to

a ventral plate, 51 to the length of the head. Ventral plates in

6 longitudinal and 29 transverse series. Anal plate preceded by

a large but narrower plate and bordered on one side by two small

plates, on the other by three.

Hind limb reaching the wrist of the adpressed fore limb;

‘scales on the upper surface of the tibia round, feebly keeled, con-

siderably smaller than the dorsals, 14 of the former corresponding

to 10 of the latter. Femoral pores 17-18. 28 scales under the

fourth toe.

Caudal scales as in specimen No. 2, but the keels somewhat less

developed.

Uniform greyish above, with mere traces of dark spots and

ocelli on the sides (bleached 2); labials not dark-edged ; lower

parts white.

4. Female.—Head as in a typical L. muralis, its depth equal

to the distance between the centre of the eye and the tympanum ;

snout obtusely pointed. Rostral not entering the nostril and

separated from the frontonasal by the nasals, which form a rather

long median suture ; frontal as long as its distance from the end

of the snout; a complete series of granules between the two

pvincipal supraoculars and the supraciliaries; parietals once and

a half as long as broad, not in contact with the upper postocular,

the outer border very slightly concave for the accommodation of the

large upper temporal; occipital not quite half the length of the

interparietal but a little broader ; temporal scales small and

granular ; tympanic shield well developed ; masseteric shield very

small, separated from the upper temporal by two or three series of

granules; four upper labials anterior to the subocular, which is

considerably narrower beneath than above.

Collar even-edged, formed of 10 plates; 25 scales between the

chin-shields and the collar.

Body moderately depressed. Dorsal scales round, flat, per-

fectly smooth, 48 across the middle of the body, 3 corresponding

to a ventral plate, 35 to the length of the head. Ventral plates

in 6 longitudinal and 28 transverse series. Anal plate bordered

by a semicircle of 4 rather large plates.

Hind limb reaching elbow of adpressed fore limb. Scales on

upper surface of tibia round, smooth, smaller than the dorsals,

12 of the former corresponding to 10 of the latter. Femoral

pores 16-16. 26 scales under the fourth toe.

Caudal scales moderately keeled, as m a typical L. muralis; 32

scales in the fourth*whorl,

336 MR. G. A. BOULENGER ON THE [ Nov. 29,

Black dots irregularly disposed on the middle of the back;

a dark lateral streak, formed of confluent vermicular spots,

enclosing light ocelli above the shoulder ; upper surface of head

without spots; labials not dark-edged; lower parts white.

5. Male*.—Shape of head and head-shields as in specimen

No. 1, but nasals forming a short suture behind the rostral, fourth

supraocular undivided, and lower border of subocular distinctly

shorter than upper.

Collar even-edged, composed of 8 plates; 32 scales between the

chin-shields and the collar; gular fold very indistinct f.

Body much depressed. Dorsal scales roundish-hexagonal, flat,

smooth, very faintly keeled on posterior part of back; 61 scales

across the middle of the body, 3 or 4 corresponding to a ventral

plate, 42 to the length of the head. Ventral plates in 6 longi-

tudinal and 25 transverse series. Anal bordered by a semicircle

of 8 small shields, the right median of which is twice as large as

the left.

Hind limb reaching the shoulder. Scales on upper surface of

tibia rhomboidal, keeled, and a little larger than the dorsals, 8

transverse series of the former corresponding to 10 of the latter.

Femoral pores 18-18. 26 scales under the fourth toe.

Caudal scales as in No. 1, but the whorls more distinctly unequal

in length, alternately longer and shorter ; 26 scales in the fourth

whorl.

The specimen is much bleached ; in what remains of the mark-

ings it agrees with No. 1.

The following measurements are taken from the five specimens,

in all of which the tail is either reproduced or partly broken off :—

I 2: 3. 4. 5.

mm, mm, mm. mm. mm.

From snout to vent ....... .. 68 60 53 53 52

<5 jp thovaey Ihvemloy sae5 2s) 23 18 KS) 21

Iuength of head ............:.. 17 15 11 12 133

Wadi brOb rh Cagiva. sane losers 11 9 Ths Wi Z 8

Depth of heady 2. ..2...5e- a0 6 6 4 54 5

One MIMO), cence dececcnerneetiges 23 20 16 16 19

Titan clasannalo uey, brocilo te eiice ane icies 40) 32 24 24 28

POO bie oars acct cists Saroosteamcicds 20 We 14 14 16

The first and largest specimen, the male of which measure-

ments have been given by Camerano%, having smaller dorsal

scales (60 in a transverse series), less spinose caudal scales, and

22-21 femoral pores, clearly represents Bedriaga’s var. modesta §,

and is identical with the smaller male specimen referred to as

* The specimen preserved in the British Museum.

+ Quite distinct in the other specimens.

{ The female specimen of which comparative measurements have been given is

evidently the one which has passed into Dr. de Bedriaga’s private collection.

§ Bedriaga divided the species into two varieties: modesta and rudés. The name

modesta had been previously proposed for a variety of Z. maralis by Himer.

1904.] LACERTA DEPRESSA OF CAMERANO, 337

L. depressa in the British Museum Catalogue of Lizards*. Should

Camerano’s species be broken up into several forms, this specimen

must be regarded as the restricted type of LZ. depressa. It is pro-

bably from Trebizond, since Boettger’s specimens from that locality

agree closely with it’; but it must be noted that specimens from

Shusha, E. Karabagh, received from the Senckenberg Museum

as L. muralis, var. defilippii Boettgert, belong, in my opinion,

to the same variety. Méhely’s var. depressa from Shion in Trans-

caucasia § is also probably the same thing.

The second specimen, with larger dorsal scales (51 across the

body), with larger and more strongly keeled scales on the tibia,

with more strongly raised keels on the caudal scales, and with 18

femoral pores, has been specially selected by Bedriaga as the type

of var. rudis. But it is remarkable that no allusion should have

been made to its having five anterior labials instead of four, the

number unreservedly given in the diagnosis of LZ. depressa. The

specimen from Batoum noticed by Boettger || is probably correctly

referred to this form; it has 5 anterior labials on one side, 4 on

the other, and 46 scales across the body.

The third specimen answers in all important respects to the

figure of L. portschinskii of Kessler, from Tiflis, the Russian

description of which was translated in 1879 by Bedriaga **.

Bedriaga then identified L. portschinskii with L. depressa; he after-

wards in his monograph published in 1885, most emphatically

repudiated this identification and placed L. portschinskii simply

in the synonymy of his LZ. mwuralis fusca, whilst regarding

L. depressa as a distinct species having much less in common with

L. muralis than with L. oxycephalatr.

The fourth specimen agrees entirely with the Persian lizards

described by De Filippi and by Blanford, and may be referred to

the var. defilippi Camerano, of which var. persica Bedriaga, is a

synonym.

The fifth specimen, as stated above, should he referred to the

var. depressa, sensu stricto.

Comparing the L. depressa of Werner with these specimens, I

find it does not agree with any of them, differing in the shorter

limbs, a character emphasised by Werner in his description.

But it agrees with specimens from Lake Gokcha which, in my

opinion, represent the L. chalybdea of Eichwald (LZ. muralis, var.

saxicola Bedriaga). In the following table I give the measure-

%* Vol. ili p. 34 (1887).

+ Ber. Senck. Ges. 1892, p. 141. 58-60 scales across body, 30-32 gular scales,

8 tibial scales corresponding to 9 orl10 dorsals.

tL. c. p. 144. These Shusha specimens are regarded by Boettger as connecting

the var. defilippii Camer. with the var. raddii Boettg.

§ Dritte Asiat. Forschungs. Graf. E. Zichy, ii. Zool. p. 54 (1901).

|| Ber. Senck. Ges. 1889, p. 204.

#& Aych. f. Nat. 1879, p. 308.

++ The British Museum possesses a quite similar specimen from Elizabethpol,

among several received trom the St. Petersburg Museum, the largest of which agrees

with Eversmann’s figure of L. saxicola.

Proc. Zoo. Soc.—1904, Vou. IT. No. XXII. 22

308 ON 'THE LACERTA DEPRESSA OF CAMERANO. [ Nov. 29,

ments of two female specimens of the same size, the first of which is

one of Dr. Werner’s L. depressa from the Bythynian Olympus, near

Brussa, altitude 1500 m.; the second forms part of a small series

from Yelenowka, Lake Gokcha, altitude 2000 m., received from

the St. Petersburg Museum in 1886. Scaling and coloration, in

these two examples, are practically identical :-—

ik. 2.

mm. mm,

ROTI SAG LWG WO) WEG osoonsoscavsoooon0gococnnnener 70 70

ms see nb Hone Mimic ah. thon spine 22 22

enethyotdnead setae crac aens tc. ieeeiath is sentsete 15 14

IWirclthv or Wear acs ache aoassen ocesb ears 10 9

Deyo tla ores Carel ae verre serene ehes ca cette ta ace re rn 6 6

TMOre pinnate eee atch tesotet nicks nis tnewreicei acces 20 al

TEV Tin cle ilo sey sect eto aetna a caren Groeh ise 29 30

OO Unsaic. cher bun chiacabaaives aaatnw nen vai rin smtente 15 16

On the other hand, I shall surprise many by stating that

T have before me a female collected by Dr. Gadow in the Serra

Estrella, Portugal, which, in form, coloration, and lepidosis, is,

with exception of the tibial and caudal scales, a very good match

for the var. rudis of LZ. depressa. The wall-lzard from the High

Pyrenees, mentioned by Bedriaga and suspected by Meéhely to

represent a distinct species, is much nearer to the typical form,

from which it should certainly not be separated. Curiously, its

collector, Lataste, was struck with its general resemblance to

L. vivipara, jast as Werner was when observing his supposed

L. depressa on the Olympus near Brussa.

Since writing these notes, I have received, through the kind-

ness of Dr. A. N. Kaznakoft, Director of the Caucasian Museum,

Tiflis, a female specimen from Tchorok, Caucasus (Coll. Radde &

Kiénig) which entirely agrees with the var. radis. The scales

on the tibia are very large, rhomboidal, strongly keeled; 6

oblique series correspond to 10 transverse series of dorsal scales ;

the latter are very distinctly keeled, and number 45 across the

middle of the body. Four upper labials anterior to the sub-

ocular; rostral shield forming a suture with the frontonasal ;

first and fourth supraoculars broken up into three small shields ;

parietal not touching the upper postocular. Femoral pores 18-17.

26 lamellar scales under the fourth toe. The hind limb reaches

the shoulder. The plate in front of the preanal transversely

enlarged.

[P.S. Jan. 9, 1905.—Thanks to the kindness of my friend

Dr. de Bedriaga, I have been able to examine the female specimen,

alluded to above, still preserved in his private collection. It

belongs to the var. rudis, having 52 scales across the middle of the

body, very large and strongly keeled tibial scales, and almost

spimose seales on the sides of the base of the tail. The only

important characters in which it differs from the female No, 2

are the presence of only four anterior upper labials and the

1904. | ON OLD PICTURES OF GIRAFFES AND ZEBRAS. 339

absence on one side, and the very small size on the other, of the

masseteric shield. Femoral pores 17-19. From snout to vent

70 millim.

Dr. de Bedriaga has also sent me on loan a large male, measur-

ing 83 millim. from snout to vent, from Batoum, received from

the St. Petersburg Museum, which also belongs to the var. rudis.

The cheeks are much swollen, and the depth of the head equals

the distance between the centre of the eye and the anterior

border of the tympanum. The hind limb reaches the shoulder.

45 scales across the middle of the body. 21—22 femoral pores.

25 scales under the fourth toe. Rostral forming a narrow suture

with the frontonasal. Only three labials anterior to the sub-

ocular. This specimen, which I have carefully compared with

Bedriaga’s L. depressa, var. rudis, confirms Boettger’s identifica-

tion of a Batoum specimen in 1889. |

EXPLANATION OF PLATE XXII.

Three of the type specimens of Lacerta depressa, Camerano; enlarged figures

showing upper and side views of head and posterior part of back, with hind limb

and base of tail.

a. Male (no. 1, i, 333) X 2; 6b. Femaie one 2; p. 334) x 23;

. Female ne 2, p. 834), X

6. On Old Pictures of Giraffes and Zebras.

By R. LyDEKKER.

| Received October 7, 1904. |

(Text-figures 85-89.)

The Natural History Branch of the British Museum has re-

cently received from the Lord Chamberlain, through My. Lionel

Cust, Surveyor of the King’s Pictures and Works of Art, four

photographs from paintings of Giraffes and a Zebra preserved in

the Royal Collection, some of which are of considerable interest

from an historical point of view.

The painting from which text-figure 85 is taken represents an

immature Nubian Giraffe presented in 1827 by Mohamed Ah,

Pasha of Egypt, to his Majesty George the Fourth. This animal,

which survived but a short time at Windsor, was the first

received alive in Britain, and one of the four first imported into

Europe in modern times. Of its three fellows, one was sent

by the Pasha to the Sultan of Turkey, the eecond to Vienna,

and the third to Paris, where it attracted an enormous amount of

attention *. Although, owing to the immature condition of the

animal, the frontal horn is not fully developed, the paimting dis-

plays all the characteristics of the typical Nubian race of Giraffa

camelopardalis, such as the net-like style of the markings, the

white ‘ stockings,” and the comparatively large size of the spots

on the upper part of the legs. A portrait of My. Cross, the

* See Renshaw, ‘ Natural History Essays,’ p. 105 (1904).

29%

340 MR. R. LYDEKKER ON OLD PICTURES [ Nov. 29,

aninal-dealer, is introduced into this picture, which is the work

of James Laurent Agasse.

Text-fig. 85.

George the Fourth’s Nubian Giraffe at Windsor.

(From a Painting in the Royal Collection.)

The first picture (text-fig. 85) represents the animal in its

surroundings at Windsor; the painting reproduced in text-fig. 86

depicts, on the other hand, a Giraffe in its native country, or

what is intended therefor. Whether this painting portrays the

same animal as the first it is not easy to determine. Apparently,

however, the portrait is that of a Nubian Giraffe, although it is

by no means so good as the first; the animal apparently had

1904. | OF GIRAFFES AND ZEBRAS. 341

a distinct frontal horn, although it is much concealed by the

halter.

Text-fig. 86.

George the Fourth’s Nubian Giraffe in its native country.

(From a Painting in the Royal Collection.)

The third picture (text-fig. 87) represents a group of Giraffes

which appear to be intended for the Southern or Cape form

(G. c. capensis), as the old bull has no frontal horn, while the

markings are of the blotched instead of the netted type, and the

lower parts of the legs are spotted, although not quite so fully as

they ought to be. The painting, which is by R. B. Davis, and

dated September 1827, is described, however, as representing

“two Giraffes belonging to George [Vth ;” but on the back is

342 MR. R. LYDEKKER ON OLD PICTURES [ Nov. 29,

written “portrait of the Giraffe belonging to his Majesty.” It

is difficult, however, to believe that the artist did not take a por-

trait of the Cape Giraffe for his model, and he may have copied

Paterson’s specimen in the British Museum. If, as I think, it

represents the Cape Giraffe, the painting is of very considerable

Text-fig. 87.

Group of Cape (?) Giraffes.

(From a Painting in the Royal Collection.)

interest, as that race now appears to be extinct. Both Agasse

and Davis were well-known animal painters in the first quarter

of the last century.

The fourth painting (text-fig. 88) represents a specimen of the

Mountain Zebra (Hquus zebra) in two positions; the characteristic

Text-fio, 88.

1904. | OF GIRAFFES AND ZEBRAS. 343

“ ovidivon” on the rump, the broad stripes on the thighs, and the

white under-parts being remarkably well shown. The picture is

stated to have been drawn from life at the Hague by C. Kehrer ;

but there is no record at Windsor of the original model having

(From a Painting in the Royal Collection,)

Mountain Zebr

Text-fie. 89,

344 ON OLD PICTURES OF GIRAFFES AND ZEBRAS. | Nov. 29,

been there. Car] Christian Kehrer, who painted hunting-scenes

and portraits, and is, I take it, the painter in question, was born

in 1758 and died in 1833.

In this connection | may mention that a few years ago my

friend Dr. F. H. H. Guillemard sent me a copy of an old print

bearing the following legend :—

“The portraiture of the Zebra or Wild Ass, drawn from the

life. This beautiful animal was brought from the Cape of Good

(From an Old Print.)

€

Queen Charlotte’s Mountain Zebr

P.Z.S.1904,vol.Il. Pl. XXII.

J.Smut delet ith - Mintern Bros .imp.

NCES OP LOMLSES..

1904. | MR. R. LYDEKKER ON TWO LORISES. 345

Hope by Sir Thomas Adams in the Zerpsicore Man-of-War and

presented to her Majesty [Queen Charlotte}. 1762.”

I think there is considerable probability that this print (text-

fig. 89) and the royal picture were taken from one and the same

animal, although the general drawing and the details of the

stripes are far less true to nature in the former than in the latter ;

but Kehrer’s portrait could not have been done at the Hague

while the animal was en route for England, and there is no record

of its having been taken there later. If both pictures represent

the same animal, Queen Charlotte’s Zebra, as the individual repre-

sented in the old print may be called, appears to have been the

first of its kind ever brought to England.

7. On Two Lorises. By R. LypDEKKER.

{ Received October 31, 1904. |

(Plate XXIIT.*)

The Trustees of the British Museum have recently purchased

from Rowland Ward, Ltd., two mounted specimens of Lorises

belonging to forms hitherto unfigured, and one of which I regard

as new; the first specimen being a Slow Loris (Vycticebus), and

the second a Slender Loris (Loris). Both of these genera, it may

be observed, appear to be represented only by a single species, if

we except the ill-defined .V. menagensis of the Philippines.

Whereas, however, several local forms of the Slow Loris have

been recognised, the Slender Loris has hitherto been undivided.

As regards the Slow Loris, Messrs. Stone and Rehn, in the

‘ Proceedings’ of the Philadelphia Academy for 1902 (pp. 138 &

139), recognised five local forms, namely, Vycticebus tardigradus *

typicus of India, V.t. javanicus, NV. t. malayanus, NV. t. natune, and

iV. t. hilleri (of Sumatra); the two last being described for the

first time. In addition to these there is the Tenasserim form, of

which no examples were at the time available.

The first two of the five races mentioned above are grouped

together in a section characterised by the general colow: being

ashy grey, slightly tinged with rufous, while the crown of the

head is not marked by a large patch of brown. In the three

yemaining races, on the other hand, the general colour is rutfes-

cent grey, and the crown of the head has a large brown patch.

Omitting mention of the Natuna Islands’ form, the Malay race—

as represented in the collection of the British Museum by three

mounted specimens from Penang (Pl. XXIII. fig. 1), the gift of

Capt. Stanley Flower—is characterised by the general rufescent

grey tone of the fur, and the strongly pronounced rufous-brown

crown-patch and dorsal stripet. This crown-patch has a pair of

lines extending transversely outwards to the ears, and another

* For explanation of the Plate, see p. 346.

+ Messrs. Stone & Rehn substitute the name coucang.

{ This does not accord with Messrs. Stone & Rehn’s description.

346 MR. R. LYDEKKER ON TWO LORISES. [ Nov. 29)

and broader pair passing obliquely downwards to the eyes, which

they encircle. Above the eyes the white interocular stripe

expands markedly.

A very different-looking creature is the Sumatran Slow Loris

(LY. ¢. hilleri), now, L believe, for the first time figured (Pl. XXIII.

fig. 2). In this race the general colour is bright rufescent chest-

nut slightly washed with grey; the dorsal stripe and crown-patch

being a deeper rufous, faintly bordered with brown, and on the

whole less distinct than in WV. ¢. malayanus. The crown-patch is

much less distinctly divided into two pairs of lines than in the

latter; the blotches over the eyes being much broader, and the

interocular white streak consequently much reduced in width.

The plate illustrates very clearly the marked distinction between

the grey Malay phase and the rufous Sumatran phase of the species ;

it would be interesting to know the reason for this very strongly

marked local difference in colour.

Passing on to the Slender Loris (Loris gracilis), I find that in

the typical Indian form of this animal, as represented by a couple

of mounted specimens from Madras recently presented to the

British Museum by Mr. E. Thurston (Pl. XXIII. fig. 3), the

general colour is pale mouse-grey passing into pure white between

the eyes, on the sides of the face, under-parts, &e. On the face

the white interocular streak extends some distance on the fore-

head above the line of the eyes, and then divides into a pair of

bands which pass outwards in front of the eyes, and thus cut off

a small patch above each of the latter from the grey of the rest

of the head. ‘These patches are practically restricted to the areas

above the eyes, there being little or no grey fur on the outer side

of and below the latter.

The Ceylon Loris, on the other hand, of which the British

Museum has recently purchased a mounted specimen (Pl. XXIII.

fig. 4), is a rufous instead of pale grey animal ; the general colour

of the upper-parts and eye-patches being pale rufescent brownish

erey silvered with white; the crown and back being darker than

elsewhere. Moreover, the eye-patches are much larger, the brown

extending round the outer side of the orbits to occupy a con-

siderable area below them. Again, with the exception of the

interocular stripe and the sides of the face (and even these are by

no means pure white) the under-parts are cream-colour or pale buff,

instead of white. These differences, I submit, amply demonstrate

the right of the Ceylon Loris to subspecific distinction, and i

accordingly propose to eall it Loris gracilis zeylanicus, taking the

British Museum specimen as the type.

EXPLANATION OF PLATE XXIII.

Fig. 1. Head of Nycticebus tardigradus malayanus (p. 345).

2. Head of Nycticebus tardigradus hilleri (p. 346).

3. Head of Loris gracilis typicus (p. 346).

4, Head of Loris gracilis zeylanicus (p. 346).

1904. | ON A NEW GAZELLE FROM PALESTINE. 347

December 13, 1904.

Hersert Druct, Esq., F.Z.8., Vice-President,

in the Chair.

The Secretary read the following report on the additions to the

Society’s Menagerie during the month of November 1904 :—

The number of registered additions to the Society's Menagerie

during the month of November was 150. Of these 68 were ac-

quired by presentation and 14 by purchase, 26 were received on

deposit, 34 m exchange, and 8 were bred in the Menagerie. The

number of departures during the same period, by death and

removals, was 179.

Amongst these, special attention may be called to the fol-

lowing :—

1. A Ferret Badger (Helictis personata) from Burmah, presented

by Capt. Burnett on Nov. 4th. This species has not been exhibited

previously in the Gardens.

2. Two Chimpanzees (Anthropopithecus schweinfurthi), a male

and female, from the Bagomo Forest, Uganda, presented by

Mr. Stanley C. Tomkins, C.M.G., on Nov. 7th. These animals

are nearly adult, the male being the finest Chimpanzee ever shown

in the Gardens.

3. A male specimen of the Senegal race of the African Buffalo

(Bubalus centralis), deposited on Noy. 21st. No example of this

subspecies has been exhibited previously in the Menagerie.

4, Three Mouse-Hares (Ochotona roylei) from Baluchistan, pre-

sented by Col. Chas. E. Yates on Nov. 28th. These interesting

little Rodents are also new to the Collection.

Mr. Oldfield Thomas, F.R.S., exhibited specimens of a pair of

Gazelles from Palestine which had been sent by Dr. Selah Merrill,

U.S. Consul at Jerusalem, to Dr. Sclater, and had been passed

on by the latter to the British Museum *.

The Gazelle proved to have no relationship to Gazella dorcas,

the only species as yet recorded from Palestine, but to be nearly

related to the Edmi or Atlas Gazelle (Gazella cwviert), of which,

although distinct, it might be considered a local representative.

As no Gazelle of this type was found in the intermediate countries

of Tripoli and Egypt, so that connecting links were unlikely to

oceur, it seemed more in accordance with modern practice to give

the Palestine form a binomial rather than a trinomial name.

The Gazelle was therefore proposed to be called :—

GAZELLA MERRILLI Thos. (Text-fig. 90, p. 348.)

Gazella merrilli Thos. Abstr. P. Z.8. No. 12, p. 19, Dec. 13, 1904.

Colour and general appearance exactly as in G. cuvieri, the

* [The complete account of the new species described in this communication

appears here, but since the name and preliminary diagnosis were published in the

PP Seth ome cane 2 seew

* Abstract,’ the species is distinguished by the name being underlined—Ep1ror. |

348 ON A NEW GAZELLE FROM PALESTINE. [ Dec. 13,

coarse hair, the markings, knee-brushes, and tone of body-colour

all just as in that animal, of which a figure and description had

been published in the ‘ Book of Antelopes’*.

But in size of skull and in the development of the horns the

Palestine Gazelle was markedly inferior to that from the Atlas,

as might be seen from the measurements given below, while the

curvature of the horns was distinctly different. In G. cwvieri the

horns, which might attain to 12 or 13 inches in height and have

up to 24 well-developed rings, were very slightly curved back-

wards below and equally slightly—indeed scarcely at all—recurved

forwards at their tips. In the Palestine form, on the other hand,

the horns in an old male were short, thick basally, and markedly

S-shaped, curved backwards below, and distinctly recurved for-

wards at their tips; the ridges, which were less well defined than

in cwvert, numbered only about 10-12. Viewed from in front,

the horns were evenly divergent, without lyration.

Text-fig. 90.

Skull and horns of Gazella mervilli.

The horns of the female were of fair relative development, nearly

four inches in length, and about half an inch in diameter at the

base, smooth throughout, slightly curved upwards.

The skull was decidedly smaller than in G. cwvieri, the nasals

shorter, and the premaxillary bones did not in either specimen

* Vol. ili. p. 109 (1898).

1904.| ON THE BRITISH SPONGES OF THE GENUS LEUCCSOLENIA. 349

reach more than two-thirds up towards the nasals, while in

cuviert they articulated broadly with the latter bones. The bulle

were more rounded, and the pair of prominences on the basis of

the skull over the basilar suture were much more developed than

in the single old skull of G. cwviert available. From G. arabica,

to which there was a certain resemblance in the set and curvature

of the horns, G. merrilli was distinguished by its conspicuously

greater size.

The typical male skull of G. merrilli measured as follows :—

greatest length 194 mm.; basal length (c.) 170; greatest breadth

88; muzzle to orbit 99; length of upper-tooth row 60.

Horns— ¢. Length over curves anteriorly 241; basal cireum-

ference 108. 2. Length 98; basal circumference 41.

Type. &, B.M. No. 4.12.18.1. Killed 11th December 1903.

Presented by Dr. Selah Merrill through Dr. P. L. Selater.

Hab. Hizmeh, just north of Jerusalem.

The following extract from a letter from Dr. Merrill, dated

31st October 1904, will give an idea of where this new Gazelle

was found :—

“The male Gazelle was brought to me Dee. 11th, 1903, and the

female on February 7th, 1904. Both were shot by a hunter

whom I know, and who has served me at times during several

years past. His home is at Hizmeh, five or six miles north of

Jerusalem; he is a plain simple man, and probably never goes as

far as ten miles from his village. Hizmeh is near Wady Farah,

two hours north-east of Jerusalem. All that region for many

miles north of Wady Farah, where the tableland of Judea breaks

down to the Jordan valley, is very wild and has never been very

carefully explored.”

Mr. Thomas had much pleasure in naming this new Gazelle,

which represented a type quite new to the Palestine Fauna, in

honour of its discoverer Dr. Selah Merrill.

The following papers were read :—

1. The Characters and Synonymy of the British Species of

Sponges of the Genus Leucosolenia. By E. A. MIncuin,

University College, London.

[Received November 15, 1904. |

(Text-figures 91-98.)

I. IyrrRopucrory.

The following memoir is an attempt to fix the nomenclature

and to define the characters of the British species of Ascons

belonging to the section for which, in my opinion, the name

Leucosolenia is the correct taxonomic designation. In order to

350 PROF. E. A. MINCHIN ON THE BRITISH [ Dec. 13,

carry out this intention a large number of specimens, including

the types of earlier authors, have been carefully examined. This

task has been forced upon me as the result of some investigations

upon the development of the various forms of spicules in sponges

of the genus Leucosolenia, which it is hoped to publish shortly.

It was found that in describing the spicule-development it would

be necessary either to use names for the species which were in-

correct or else to employ a nomenclature at variance with that in

the current literature dealing with these sponges. And since the

utmost confusion exists with regard to the designation of these

species both in the labels of museums and collections, no less than

in even the most recent works dealing with them, 1¢ seemed worth

while trying to give a thorough description of their distinctive

characters once and for all. It was my original intention to

have included the Mediterranean species in ‘this memoir, but

lack of material foreed me to confine myself for the pr esent to

the British forms, which, moreover, are the more important from

the taxonomic point of view, as including the earliest described

sponges of the genus.

The generic name Lewcosolenia Bwk. is used by different authors

in different senses, but is employed in this memoir in the same

sense aS in my former publications [15, 16], namely, to include

those Ascons which form a natural group distinguished by the

following characters :-—

1) The sponge-body or colony grows in a more or less erect

form with relatively large, distinct oscular tubes. (2) All three

kinds of spicules are present—monaxon, triradiate, and quadri-

radiate. (3) The triradiate systems have two paired angles, less

than 120°,and an unpaired angle greater than 120°, corresponding

to a straight median ray and two curved lateral rays. (4) The

collar-cells have the nucleus apical in position, situated close

under the origin of the flagellum. (5) The larva is an amphi-

blastula, and the first spicules formed are monaxons. Of these

characters, all except (2) ave family characters, distinguishing the

Leucosoleniide from the Clathrinide ; the latter having a reti-

culate form, equiangular trivadiate systems, basal nucleus in the

collar-cells, and parenchymula larva, the first spicules to be formed

being triradiates. The presence of all three kinds of spicules

distinguishes Zewcosolenia from Haeckel’s genus Ascyssa,in which

monaxons alone are alleged to be present—a genus which, if it

exists, should probably be placed in the family Lewcosolenride.

IJ. HisrortcaAn Review AND CrIvrIciIsM.

The earliest descriptions of species of Ascons were based en-

tively on outward form and appearance, and though the spicules

were noticed and figured no use was made of differences in spicu-

lation in order to distinguish the species. The external characters

of an Ascon are not a very safe guide, as a rule, to its specific

identification. In some cases, however, a species has a typical

1904. | SPONGES OF THE GENUS LEUCOSOLENIA. 301

form which enables one to recognise it almost with certainty, or

at least to distinguish it from others belonging to the same fauna

by simple inspection. That is the case particularly with Lewcoso-

lenia botryoides, the first-named Ascon species described in 1786

from the British coast (locality Emsworth, between Sussex and

Hampshire) by Ellis and Solander [7] under the name Spongia

botryoides. In this case the form is so characteristic that there

can hardly be any doubt as to the species which is represented by

the authors’ figure. The next species of Lewcosolenia to be

described was the Svongia complicata of Montagu [17], also from

the British coast; and in this case it is more difficult to be

positive, but, on the whole, it is highly probable that Montagu’s

figures represent a specimen of the sponge for which his name

is retained, following Haeckel, in this memoir. The arborescent

mode of growth depicted is a feature extremely characteristic, if

not absolutely distinctive, of this species, at least as fav as the

British fauna is concerned. On the other hand, more doubt

attaches to the third species described, the Spongia confervicola of

Templeton, 1836 [22], which is evidently a Lewcolosenia from the

figures, but of which the specific identity remains doubtful.

Templeton’s memoir is freely annotated by ‘G. J.,” apparently

George Johnston, and the footnote to S. confervicola is ‘* Spongia

complicata G. J.” I am more inclined to the opinion, however,

that the figure of S. confervicola represents a specimen of the

later described species ‘‘ Aseandra variabilis” of Haeckel, though

Haeckel himself identifiesit with botryoides. In any case, as the

point cannot be determined, Spongia confervicola must be ve-

garded as a nomen nudum without importance for taxonomic

nomenclature. Johnston, in his work of 1842 [14], ignores both

Spongia complicata, as Fleming [8] had done before him, and

S. confervicola; both these authors recognise only Spongia

botryoides, and regard Montagu’s S. complicataas merely a variety

of the former.

In the meantime different authors had subdivided the compre-

hensive genus Spongiw into various genera, and Templeton was

one of the last to employ the name for any calcareous sponge.

Fleming in 1828 [8] proposed the generic name Grantia for all

calcareous sponges, putting G'. compressa Fabry. as the first, and

G'. botryoides Ell. & Sol. as the second species; compressa must

therefore be regarded as the type species of Grantia. Grant in

1833 [11] proposed the genus Lewconia for calcareous sponges,

putting as his first species mivea, which is therefore the type

species of this genus. In 1834 de Blainville [6]-proposed the

name Caleispongia in exactly the same sense as Grantia, putting

also compressa first and botryoides second, so that this generic

name becomes a synonym of Grantia. A great advance was made

by Bowerbank [1], who, in 1864, further subdivided the calcareous

sponges. Pointing out that Grantia botryoides was quite different

from either the G. compressa (Fabr.) or G. nivea (Grant) of

Fleming, he retained Fleming’s genus Grantia for compressa,

302 PROF. E. A. MINCHIN ON THE BRITISH [ Dee. 13,

Grant’s genus Leuwconia for mivea, and placed botryoides in a new

genus Leucosolenia. In making these changes Bowerbank acted

with perfect correctness, according to accepted modern rules of

nomenclature; and it is clear that for the species botryoides the

generic name Leucosolenia has the priority over all other generic

names for it, or for other species associated with it generically.

Leucolosenia is, in short, the first generic name put forward which

has an undoubted Ascon as the type species.

Bowerbank added various species to his genus Leucosolenia,

amongst them forms which, in my opinion, cannot be associated

generically with botryoides, and therefore do not belong to the

genus Leucosolenia as here understood, but to that section of the

Ascons for which I employ the generic name Clathrina (Gray,

1867). Moreover, Bowerbank did not properly understand the

distinctions between the different species which he dealt with, so

that different species are found confused together in his monograph

in an extraordinary manner, and his descriptions are sometimes

quite incorrect. Thus the specimen described and figured as

L. botryoides in vol.i. of his monograph (p. 164, figs. 347, 348,

pl. xxvi.) does happen to be a genuine specimen of botrs yoides.

This can be seen at once from ints figure 348, which is extremely

characteristic, and I have been able ae examine this specimen and

have figured its spicules (text-fig. 98, figs. 27. a—-g, p. 390). On

the other hand, the specimen figured as L. botryoides, in vol. 111.

pl. 11. fig. 1, 1s a specimen of the species described by Haeckel

under the specific name variabilis, and the description given by

Bowerbank of the triradiate spicules as ‘“‘ equiangular” (vol. 11.

p- 28, vol. 111. p. 7) can be seen, even from his figures, to be in-

correct. Bowerbank further described a new species under the

name “ Lewcosolenia contorta.” I hope to discuss the rather com-

plicated question of the characters and synonymy of this species

in another memoir, the true contorta being a Clathrina. I will

only say here that amongst specimens identified by Bowerbank as

contorta I have found a Clathrina species mixed up with speci-

mens of Leucosolenia complicata and variabilis. Bowerbank

himself considered (vol. 1. pp. 30, 31) that his species contorta

might be synonymous with Montagu’s species complicata, but

was more inclined to regard Montagu’s figure of the latter as

being “‘a very characteristic figure of Spongia botryoides Ellis &

Solander,” and thought it better under the circumstances to reject

the term complicata altogether. Finally, in vol. iii. of his mono-

eraph, Bowerbank described and figured a sponge found in

Brighton Aquarium under the name of Leuconia somesii (pp. 334—

332, pl. xci. figs. 6-17). A glance at his figures makes it obvious

that this sponge is a Leucosolenia, but his description is inade-

quate for determination of the species. Having been able to

examine Bowerbank’s types of this species in the British Museum,

I found them to be merely aberrant specimens of Lewcosolenia

variabilis (Haeckel), as Topsent had already suspected, charac-

terised by the great development in the number and length of

1904. | SPONGES OF THE GENUS LEUCOSOLENIA. 303:

the monaxons, and, as shown by Bowerbank’s figures, a great

tendency to the production of abnormal forms of triradiates (see

text-fig. 95, fige. 18 a—e and 19a-h, p. 379). Leuconia somesii

may, in short, be characterised as an interesting aquarium variety

of Leucosolenia variabilis, showing modifications parallel to those

described by Bidder for Sycon raphanus*, growing in the Naples

Aquarium. Bowerbank himself was struck by the resemblance of

this sponge to a Leuwcosolenia, and particularly to the specimen

figured by him in pl. iii. fig. 1, which, as stated above, was

actually a specimen of ZL. variabilis; he remarks that the only

other known British calcareous sponge with which this species is

likely to be confounded is Leucosolenia botryoides, but “ only in

its young and immature state.” Leuconia somesii must therefore

be put as a synonym of Haeckel’s species variabilis.

Enough has been said to justify the criticism made above that

Bowerbank did not grasp the real distinctions between the species

of his genus Leucosoleniat. It is the great merit of Haeckel,

whose name marks the next epoch ¢ in our knowledge of calcareous

sponges, that he was the first to understand the great importance

of the spicules in specific determinations, and to give descriptions

of the species by which they could be recognised clearly. It

may be said, in short, that previous to Haeckel’s great mono-

graph no species of calcareous sponge was really adequately

characterised, and that Haeckel was the first to show how this

should be done. Hence, where previous descriptions of a cal-

careous sponge leave us in doubt as to its identity, Haeckel’s

determination of its characters fixes the application of the name.

Had Haeckel carried out his own method with accuracy and

conscientiousness it would not have been necessary for the present

paper to be written, but unfortunately this is far from being the

case. Of the specimens which I have been able to examine,

some have been through Haeckel’s hands and have been identified

by him, and these show, in many cases, the most extraordinary

errors of identification, as will be evident from the descriptions

and figures given below. It will be made clear, also, that in two

cases at least he founded unnecessary species simply as the result

of overlooking spicules in certain specimens which he found to

be present in others of the same species. Haeckel’s numerous

species of Ascons require, one and all, a thorough re-examination,

and there can be no doubt that a careful revision would result in

* Quart. Journ. Micr. Sci. v. xxxviii. p. 10.

+ To the well-known monographs of Oscar Schmidt [19, 20], more or less con-

temporary with Bowerbank’s writings, further reference is not necessary here, since

of Leucosolenia, im the sense used in the present memoir, only two species are

described, DT. licberkiihnii, which is not a British form, and L. fabricii, which

appears to be a synonym of complicata.

+ Both of Haeckel’s works, the ‘Prodromus’ (1870) and the ‘ Monographie ’

(1872), were published at dates betw een those of the second and third volumes of

Bowerbank’s ‘ British Spongiade ;” but they were not noticed by Bowerbank, and

belong in all respects to a subsequent epoch. Of Bowerbank’s species, only Leuconia

somesii was described after Haeckel’s monograph, and is therefore not noticed by phe

latter.

Proc. Zoo. Soc.—1904, Vor. If. No. XXITT. 23

354 PROF. E. A. MINCHIN ON THE BRITISH | Dec. 13,

many, perhaps the majority, of his specific names becoming

synonyms.

There is, however, a further blemish on Haeckel’s work, which

has been the cause of all the universal confusion in the nomen-

elature of these sponges. Haeckel took no notice in his final

monograph * of any generic names for caleareous sponges used

before him. Making a clean sweep of all previous names, he set

up twenty-one new genera, seven of which were Ascons. Such

a proceeding could not, of course, be tolerated, being a flagrant

violation of the rules of taxonomic nomenclature which have

long been followed in this country, and now are universally ac-

cepted abroad also. Hence Haeckel’s system has undergone

various modifications at the hands of subsequent writers.

The first work of primary importance dealing with calcareous

sponges after Haeckel was Poléjaeff [18], who reverted to Bower-

bank’s use of the name Zeucosolenia to denote all Ascons. In

works of later date some systematists have followed Poléjaeff in

the use of the name Zeuwcosolenia, as, for example, Topsent ;

others have used modifications of Haeckel’s system, as, for example,

Lendenfeld ; and others, again, have used Leucosolenia in some

special sense, as, for example, Breitfuss. In 1896 [15] I put

forward a scheme of classification for Ascons which has not been

followed by subsequent writers, but to which I still adhere. It

would be foreign to the purpose of the present memoir to discuss

the classification of Ascons generally, but in order to justify my

use of the name Leucosolenta | put forward three propositions :—

(1) That the oldest Ascon genus, Leucosolenia Bowerbank, is a

valid genus, founded in a perfectly correct manner, its type

species being LZ. botryoides, the oldest described species of Ascon.

(2) That therefore the generic name Leuwcosolenia has priority

over all others for this species, and the combination Leucosolenia

botryoides is one that should never be disturbed.

(3) That therefore in any scheme of classification in which

other Ascons are placed in the same genus as botryoides, they also

should be termed Leucosolenia.

If these three propositions are accepted, it becomes of extreme

importance to describe accurately the specific characters of Lewco-

solenia botryoides. I think I may claim to have done so in the

present paper, and from the description below it will be imme-

diately apparent that the restriction of the name Leucosolenia to

Ascons without monaxon spicules, as done by Breitfuss, is an

error, caused by Haeckel’s incorrect description of the species

botryoides.

As regards the specific determination of Ascons, the chief

criticism which I have to make, with regard both to Haeckel and

to post-Haeckelian systematists generally, is that sufficient account

is not taken in their descriptions of the great variability of the

* In Haeckel’s ‘ Prodromus’ [12] he put forward a scheme of classification in

which previous generic names were used, but in his ‘ Monographie’ [13] he com-

pletely altered both his classification and his nomenclature.

1904. | SPONGES OF THE GENUS LEUCOSOLENIA. 355

spicules, not only in different specimens but even in any given

specimen. In order to separate the essential from the accidental

in the description of an Ascon, not only should many specimens

be studied, if possible, but in each specimen all possible forms of

spicules should be drawn. One instance from Haeckel’s mono-

graph will suffice to illustrate this point. Under the genus

Ascandra (t.c. p. 81) the species botrys, which I have shown

below to be a synonym of botryoides, is characterised as having

‘“triradiates obtuse-angled, rays 8 times as long as thick,” while

the species witida is distinguished from it by having “ triradiates

right-angled, rays 4 times as long as thick.” Now if the reader

will turn to my figures of the spicules of botryoides given below

(text-figs. 97 & 98, pp. 388, 390), or, better still, will examine a spe-

eimen for himself, and compare the spicules with Haeckel’s figures

of the triradiates of botrys and nitida*, it will be found that in any

specimen the triradiates vary in slenderness from the types figured

by Haeckel for botrys to those given for nitida, and that their

paired angles vary from obtuse, in the more slender spicules, to

right angles in the thicker forms. There is therefore no argu-

ment to be drawn from Haeckel’s descriptions and figures against

putting these species together and considering botrys and nitida

as synonyms of botryoides. Whether they are really distinct or

not can only be determined by fresh investigation of the speci-

mens. Until that has been done we are justified in striking out

two of the three names.

Another point in which variation occurs commonly is the re-

lative frequency of a given type of spicule in different specimens.

Thus in some specimens a form of spicule may be abundant, which

in others may be so scarce that much searching may be necessary

to find itt. As negative characters require much greater caution

to affirm than positive ones, it is only after very careful investi-

gation that one should declare a type of spicule to be absent in

any specimen which agrees in all other respects with other spect-

mens in which it is present; even then it would be most unsafe

to separate such a specimen as a distinct species on this character

alone. Yet it is in this way that Haeckel separated (in error, as

it has proved) the species botryoides and complicata trom their

synonyms botrys and pinus.

The extreme uncertainty and doubt which attach to all iden-

tifications of Ascon-species in works dealing with them, have, it

may be pointed out, one important consequence: that, namely,

of rendering utterly worthless all statements concerning their

geographical distribution. It is, indeed, my firm conviction that

the study of the distribution of Ascons, perhaps of all Calcarea,

requires to be commenced de novo, and to be preceded by an accurate

study of their specific characters. Until 1t is possible to have

* The extremely fanciful, if artistic, curves which Haeckel introduces into his

drawings of spicules must be discounted in making comparisons.

+ Compare the very important observations of Topsent (Arch. Zool. Exp. (3) vii.

p. 43) on the different types of spiculation in Cliova celate at different ages.

YORK

396 PROF, E. A. MINCHIN ON THE BRITISH [ Dec. 13,

confidence in the correctness of the identification of a species,

statements as to its occurrence and distribution are of no value

whatever.

Il]. Metyops ann MaArTerriAt.

The present investigation has been directed mainly towards

a thorough examination of the spiculation, For making prepa-

vations of calcareous spicules, the reagent used by me is ; Kau de

Javelle. The piece of sponge selected is first of all, in the case of

spirit-specimens, placed in water for a minute or two, and then

put into a test-tube with a small quantity of Hau de Javelle,

barely more than enough to cover the bit of sponge. In a few

minutes the soft parts are dissolved, and with gentle shaking the

sponge disappears, being resolved into a cloud of spicules, No

heating is necessary, but the Hau de Javelle loses its powers after

a few months, and should have been recently made up. The test-

tube is then filled up with distilled water, shaken up well, and

put aside to stand undisturbed until the spicules have fallen to

the bottom, which they do in the course of a few hours. As much

as possible of the liquid is then decanted off, care being taken not

to disturb the spicules settled at the bottom, after which the test-

tube is again filled up with distilled water, shaken up, and left to

settle again. After a third washing with distilled water in this

way, the Hau de Javelle is sufliciently removed, and the tube is

then filled up with strong alcohol (90 per cent.), in which spicules

settle much more quickly than in water*. After two washings

with alcohol, the spicules are ready to be mounted. This is done

simply by drawing them up with a pipette from the bottom of

the alcohol in the test-tube, placing them on a slide, and burning

off the alcohol, leaving the spicules dry on the slide. <A drop of

Canada balsam is then put on them, and on that a cover-slip.

In this way very clean preparations of the spicules can be obtained.

Tt is advisable, however, not to defer the examination of them

too long, as even in Canada balsam they become corroded sooner

or later, and in some samples of the mounting medium the cor-

rosion proceeds rapidly. Hence attention should be paid to the

purity, that is to say the non-acidity, of the Canada balsam

employed.

For drawing the spicules I have used in all cases a camera

lucida with Zeiss’s ocular II. and objective D, giving a magnifica-

tion of 320 (reduced in the illustrations in this memoir to 300).

My method is first to draw the commoner forms of spicules seen,

and then’ to hunt carefully through the slide and draw every

spicule found differing at all markedly from those already drawn..

The process is a ‘eins one, and it is too tiring to attempt the

examination of more than two specimens a day at the utmost.

But only in this way is it possible to frame an idea of the great

* The process of washing the spicule can be greatly hastened by using a centri-

fugal machine, but as there is great danger of their then becoming caked through

interlocking of the spicule-rays, I have preferred the slower and surer method

described abov e.

1904. ] SPONGES OF THE GENUS LEUCOSCLENTA. $57

range of variations shown by the spicules in every specimen. It

is, Moreover, extremely easy to overlook inconspicuous forms of

spicules, It was not until T had studied carefully many specimens

of botryoides and variabilis that IT became aware of the invariable

presence in both of slender straight barbed monaxons (text-fig.

94, fig. 104, 1, p. 377; text-fig. 97, fig. 220, u, &e., p. 388).

For the study of the species dealt with in the present memoir

i have examined specimens from various sources. Besides those

which I have collected myself at Plymouth, Roscoff, and elsewhere,

or which have been sent me by friends, 1 have had access to

specimens of historic importance in the private collection of

Canon A. M. Norman, and in the collections of the British

Museum and the Berlin Museum. Among those who have given

me specimens my thanks are especially due to Mr. Walter

Garstang, who sent me numerous specimens from the neighbour-

hood of Plymouth, and Monsieur HE. Topsent, who sent me

specimens from Fiance. Canon Norman, with great liberality,

placed his collection at my disposal and allowed me to examine

all his specimens*. The information I acquired in this way was

most valuable, since his collection comprised type specimens of

Bowerbank and others, as well as many which had been through

Haeckel’s hands, identified by him, and returned with the labels

written or endorsed by Haeckel with his own hand. Not less

valuable, and more numerous, were the specimens in the British

Museum of Natural History, for access to which I have to thank

Dr. A. C. L. Giinther, F.R.S., who permitted me to examine all

the specimens, and Mr. Kir! kpatrick, who most kindly looked

them out for me and instructed me as to the various handw ritings

on the labels. For examination of the specimens in the Berlin:

Museum I am indebted to Dr. Weltner, who, by the kind per-

mission of Professor Moebius, was so good as to send me small

pieces of the Ascons in the Berlin collection, to enable me to study

their spiculation.

A list of the specimens from various sources which I have thus

been able to examine is given below, following the descriptions of

the species, from which it will be seen that my identification of a

specimen is often very different from that of its previous label.

T have only to add that, in the case of each specimen mentioned,

the spicules have been carefully drawn by me with the aid of the

camera lucidain the manner describedabove. The illustrations to

this memoir are tracings from a selection of the drawings so

made.

LV. GENERAL REMARKS ON THE CHARACTERS OF THE

SPECIES OF LEUCOSOLENLA.

The external form and mode of growth in the genus Lewcosolenia

* Tt is now ten yearssince 1 examined Canon Norman’s specimens, and nearly as

long since I obtained for study the specimens from the British and Berlin Museums.

Much stress of other work has prevented the completion of these investigations.

358 PROF, BH. A. MINCHIN ON THE BRITISH [ Dec. 13,

are subject to considerable variations, due to the particular en-

vironment and conditions under which the sponge grows in each

case. The variability of form is, however, combined always with

constant and characteristic features, and may be compared with

the different forms which a creeping plant will assume under

different natural conditions. Too much has been made, in many

works, of this variability, and not enough of the constancy which

underlies it. No one who has a working acquaintance with

Ascons can ever mistake a Leucosolenia for a Clathrina, except:

perhaps in the very youngest stages of growth; there is no need

for me to repeat here what I have said in previous memoirs as to

the diagnostic importance of the form of the sponge for dis-

tinguishing the two genera. It is, moreover, by no means difficult,

indeed it is usually very easy, to recognise at sight all the species

occurring in any particular locality, when one has once become

intimate with their characteristic features. Yet from looking at

Haeckel’s plates of these sponges, the impression gained is that

Ascons have no characteristic generic or specific features except

in the spiculation. Haeckel’s artistic pencil has misled him, and

others, upon this point, and his plates fail to portray the natural

appearance of the sponges.

Speaking generally, there are three principal types of body-form

occurring in ‘the species of Leucosolenia. If the sponge be growing

on a bare rock, or on the stem of a large alga, it will creep over

it, sending out anastomosing basal stolons from which oscular

tubes arise at intervals. Such a specimen has been figured by me

elsewhere [16, fig.5]|; it was found growing over a granite rock at

Roscoff, and is now in the British Museum. This type of colony

may be designated the spreading form; it is not at all common,

since it may be supposed to be a rare occurrence for the sponge to

find a rock-surface unoccupied by other competitors. More usually

these sponges are found growing crowded up amongst alge and

various organisms, often in muddy situations, or creeping over

the seaweeds, and they then assume a form which may be termed

bushy; an example of this mode of growth has been figured by

me in the case of a specimen of JL. lieber kithnii |. ¢. fig. 3] which

came from the keel of a ship moored permanently in the Porto

Militare at Naples, and which was growing in a luxurious forest

of alg, hydroids, barnacles, worm-tubes, &c. The bushy form is

the commonest type of Lewcosolenia-colony. In a third modifica-

tion the sponge forms a creeping or arborescent growth usually

closely applied to its support, but sometimes branching out under

favourable circumstances into tree-like growths [1. c. fig. 4].

Although all these three modifications of form merge into one

another, it is convenient to classify them into the three principal

types noted above.

A few words upon the characters of the spicules will not be out

of place here. The three kinds of spicules found in Leucosolenta

may be classified into: ) monaxon spicules, simple needle-like

forms; (2) triradiate systems, with or without the addition of a

1904. | SPONGES OF THE GENUS LEUCOSOLENIA. 359

fourth or gastral ray. In the monaxons a proximal end imbedded

in the wall of the sponge is to be distinguished from a distal end

projecting freely into the water. The shaft of the monaxon is

generally thickest towards the proximal end, at which it tapers

rapidly to a blunt or moderately sharp point. The distal end

usually has a barb or “ lance-head,” frequently rudimentary or

absent. The barb is in reality a double bend in the axis of the

spicule, and is comparable to a very thick bayonet, rather than

to a spear-head*. In cases where it is absent, the distal ends of

the monaxons become excessively sharp and fine.

A remarkable point with reference to the monaxons of Leuco-

solenia, which I have found to hold good, not only with regard to

the species described in this memoir, but also for all other species

that I have examined, is that the monaxons can be separated more

or less easily into two varieties, distinguished by the fact that

one kind appears very refringent, the other, by comparison, pale,.

under the microscope. The refringent monaxons are always scarcer

than the pale ones, but their peculiar optical property makes it very

easy to find them, especially under low powers (Zeiss, Oc IT. Obj. B).

In form the two kinds of monaxons may not differ essentially,

but the refringent ones always show certain characteristics which

may be summed up by saying that they tend to be straighter,

more slender, and sharper than the others, and their distal barb is

less distinct or absent. When examined by means of polarised

light, the conditions are reversed, since the pale monaxons light

up brightly between crossed prisms, while the refringent forms.

remain dark or feebly illuminated. This is particularly well seen

in the small monaxons of L. variabilis (see below, p. 380), where the

curved forms light up most brilliantly with crossed prisms, while

the straight refringent forms remain quite dark or only slightly

illuminated in all positions, when the stage of the microscope 1s

rotated. This shows clearly that the difference between the two.

types is due to a difference in the relation of the axis of crystal-

lisation to the form of the spicule.

The triradiate systems which do not acquire gastral rays do not

differ in any other structural feature from those which, by doing

so, become quadriradiates. In some cases the triradiates and

quadriradiates may differ in size, and their relative abundance

varies greatly in different specimens. In each triradiate system

we have to distinguish, as has been said above, an unpaired or

posterior ray, which in the oscular tube points away from the

oscular opening, and two paired lateral rays; the latter make

with the posterior ray paired lateral angles less than 120° and

sometimes almost approaching 90°; the lateral rays at their

junction enclose an unpaired anterior angle, which is greater

than 120° in proportion as the lateral angles are less. The lateral

rays are nearly always distinctly curved; the posterior ray is

normally straight.

* This point is, unfortunately, not very well brought out in the drawings accom-

panying this memoir.

360 PROF, E. A. MINCHIN ON THE BRITISH [ Dec. 13,

Since the triradiate systems lie in the wall of a hollow cylinder,

the three rays are never in the same plane, but are disposed in

such a manner that if the spicule be viewed in a direction cor-

responding to the axis of the unpaired ray, the two lateral rays

appear to meet at an angle less than 180° on the gastral side,

greater than 180° on the dermal side, This is an important fact

to bear in mind when studying the spicules in preparations. If

the spicule be lying on the slide with its dermal face uppermost,

then the points of the three rays touch the slide, but their

junction is raised off it; hence the spicule appears from this

aspect perfectly symmetiical, with two lateral rays of equal length

and similar curvature, but each of the three rays is slightly fore-

shortened. If, on the other hand, the spicule be lying on the

slide with its gastral face uppermost it may lie so as to appear

symmetrical, but more usually it is found lying with one lateral

ray and the posterior ray flat on the slide, the other lateral ray

pointing obliquely upwards. Hence when the two rays which

lie flat are in focus the third ray is out of focus, and when drawn

with the camera it appears foreshortened, giving the spicule an

asymmetrical appearance. To this fact is due the foreshortened

appearance of one of the two lateral spicule-rays in many of

my drawings, especially of Z. botryoides, in which the thickened

T-shaped triradiate systems are very concave on the gastral face.

V. DESCRIPTION OF THE SPECIES.

1. LEUCOSOLENIA COMPLICATA.

Spongia complicata Montagu, 1812, Wernerian Memoirs, i1.

De oe pl nix fies 23:

Spongia botryoides pars Grant, 1826, Hdinb. New Phil. Journ.

i [> HOE

Grantia botryoides pars Fleming, 1828, HEGSt. Brit. Animals,

p. 525.

Grantia botryoides pars Johnston, 1842, Brit. Spong. and Litho-

phytes, p. 178.

Leucosolenia contorta pars Bowerbank, 1866, Mon. Brit. Spong.

ul. p. 9; 1874, ui. pl. iii. figg. 5-10.

Leucosolenia botis youdes Gray, 1867, P.Z.8. p. 555

Leucosolenia fabriciti O. Schmidt, 1869, Mitth. naturwiss. Ver.

Steiermark, 11. p. 91.

Leucosolenia Jabrictt O. Schmidt, 1870, Grundz. Spong.-Faun.

Atl. Geb. p. 73.

Olynthus hispidus Haeckel, 1870, Jena. Zeitschr. v. p. 237.

Olynthus pocillum Haeckel, 1870, 1. c. p. 237.

Leucosolenia ameboides Haeckel, 1870, 1. c. p. 2438.

Leucosolenia (Leucelia) complicata Haeckel, 1870, 1. ec. p. 2438.

Leucosolenia fabricti Haeckel, 1870, 1. c. p. 2438.

Asculmis seu Ascandra armata Haeckel, 1872, Kalkschw.

pp. 77-79, pl. 13.

1904. ] SPONGES OF THE GENUS LEUCOSOLENIA. 361

Ascortis seu Ascandra fabricit Haeckel, 1872, l.c. p. 71, pl. 11.

fig. 3, pl. 12. figg. 3 a—37.

Ascandra complicata Haeckel, 1872, 1. ¢. p. 93, pl. 15. figg. 1 a—

Ascandra pinus Haeckel, 1872, 1.c. p. 105, pl. 16. figg. 34-37%

and pl. 19. ;

@ Ascandra contorta Barvois, 1876, Embryol. d. q. Eponges d. 1. |

Manche, Ann. Sci. Nat. (6) 111. p. 36.

Ascandra complicats Bowerbank and Norman, 1882, Mon. Brit.

Spong. iv. p. 226.

Ascandra complicata Fristedt, 1887, ‘ Vega’ Exped., Vetenskapl.

Jakttagelser, iv. p. 406.

Leucosolenia pinus Topsent, 1891, Arch. Zool. Exp. (2) ix.

p. 925.

Leucosolenia conrplicata Levinsen, 1893, Vid. Ud. Kanonbaaden

‘Hauchs’ Togter, v. p. 424.

Leucosolenia complicata Weltner, 1894, Wissensch. Meeresun-

tersuch. n. F. i. p. 325.

Leucosolenia complicata Minchin, 1896, Ann. Mag. Nat. Hist.

(6) xvill. p. 359.

Leucosolenia complicaia Bidder, 1898, P. R. Soc. Ixiv. p. 69.

Ascandra complicata Breitfuss, 1898, Arch. Naturges. lx. 1.

p. 213.

Ascandra contoria Breitfuss, 1898, lc. p. 214.

Ascandra fabricti Breitfuss, 1898, l.ec. p. 214.

Ascandra fabricti Breitfuss, 1898, Mém. Acad. St. Pétersbourg,

(8) vi. p. 7.

Ascandr« contorta Breitfuss, 1898, |. c. p. 15, pl. i. fig. 1.

Ascandra fabricit Breitfuss, 1898, Arch. Naturges. lxiv. 1. p. 285.

Ascandra contorta Breitfuss, 1898, 1. c. p. 285.

- Ascandra complicata Breitfuss, 1898, |. ec. p. 285.

Ascandra fabricti Breitfuss, 1898, Ann. Mus. Zool. Acad. St.

Pétersbourg, p. 17.

Ascandra complicaia Breitfuss, 1898, |. ec. p. 27.

Ascandra contorta Breitfuss, 1898, 1. c. p. 27.

Leucosolenia complicata Minchin, 1900, in Lankester’s Treatise

on Zoology, ii. Sponges, p. 5, fig. 5.

Ascandra complicata Aynesen, 1901, Bergens Mus. Aarborg,

1900, no. 5, p. 13.

Ascandra armata Ayrnesen, 1901, I. c. p. 13.

Leucosolenia complicata Rousseau, 1903, Mém. Soe. Malae.

Belgique, xxxvii. p. 7, fig. 3.

Leucosolenia fabricit Rousseau, 1903, le. p. 6, fig. 2.

Leucosolenia complicata Allen, 1904, J. Mav. Biol. Assoc. n. s.

vii. p. 185.

I commence with this species as being the most easily identified

of the three British Leucosolenias, although, curiously enough, it

is more often found incorrectly determined than either of the

others. While the systematists previous to Haeckel for the most

part considered it a synonym of botryoides, recent authors have

362 PROF, E. A, MINCHIN ON THE BRITISH [ Dec. 13,

generally confused it with Clathrina contorta, impossible as this

might seem to anyone acquainted with the two species, and con-

sidering not only the sharp differences in spiculation, correctly

described by Haeckel, but also the complete dissimilarity im

external form and appearance between the two sponges; nothing

could be imagined, in fact, more unlike than full-grown colonies

of the two sponge species.

(a) Laternal Characters.

Leucosolenia complicata occurs commonly in either the bushy

or the arborescent form. My specimens from Plymouth are all

of the former type; I have it not only from rock-pools along the

shore, but also dredged from deep water off the Mewstone. This

delicate sponge is rarely found in situations in which it is hable

to be left dry at low tide. Jt forms compact colonies in which

numerous closely-set oscular tubes arise from, and partly conceal,

a basal growth of finer tubes forming a reticulum attaching it to

the substratum. In my specimen from the Mewstone the oscular

tubes show a marked tendency to assume the characteristic tree-

like form, especially towards the centre of the bushy colony.

The largest specimens of JZ. complicata that I have seen were

collected in the Zosiera-beds at Roscoff, close to the Laboratory,

where this beautiful sponge, favoured by the shelter afforded,

grows in profusion and in the most luxuriant manner. Asarule,

it does not grow on the Zostera itself but on the stronger and

tougher alge found associated with it. The sponge itself is so

fragile that it is scarcely possible even to lift a large specimen out

of the water without pieces breaking off, and I have found it

impossible to transport them entire. These specimens, and all

others which I have collected at Roscoff, show the typical arbor-.

escent form by which the sponge can be recognised at a glance, a

mode of growth perhaps correlated with the clean granite rocks

and sand, and the pure water, very free from mud and sediment,

at Roscoff, while in Plymouth Sound the conditions are more

estuarine. In the largest specimens from the Roscoft Zostera-

beds the arborescent growth differs, in a manner which strikes the

eye at once, in different portions of the sponge-colony. In the

deeper parts, close to the stems of the supporting seaweed, the

oscular tubes form a looser, more straggling growth, apparently

due to the fact that they grow more rapidly in length than do the

diverticula which arise from them. Higher up the oscular tubes

are found growing vertically upwards, and at the same time

sending out on all sides a profusion of diverticula which become

oscular tubes and throw out other diverticula in their turn, with

the result that the sponge assumes the pinetree-like form figured

and described by Haeckel from the coast of Normandy under the

name Ascandra pinus (Monographie, vol. ii. p. 105, vol. iii. pl. 19).

Haeckel’s figure represents this form fairly well, except that, as

usual, he puts more curves into the branches than they should

1904. | SPONGES OF THE GENUS LEUCOSOLENTA. 363

have. The straightness of the oscular tubes is rather a marked

feature of the “ pinus” form. Where Haeckel’s figure is most in

error is in representing this tree-like form as an independent .

growth arising by a massive trunk from a solid rock-foundation,

instead of being merely the upper part of a large colony. It is

these pinetree-like portions which generally drop off by their own

weight when the sponge is gathered, or become detached during

transport of the specimen, and it was doubtless to a fragment of

this kind that Haeckel’s sense of artistic completeness supplied

the lacking foundation. I have also found ZL. complicata at

Roscoff growing amongst the stems of algz on rocks and isolated

boulders, in situations where it is left dry at the spring-tides,

though not at ordinary tides. Under these conditions also the

sponge shows the characteristic arborescent growth, but clings

close to the seaweeds and is never independent of them to any

great extent *.

Comparing different specimens of this sponge, I find that the

most constant feature of its habit of growth is, that the erect and

often very long oscular tubes never grow to any length without

throwing out diverticula, which, in their turn, give rise to other

diverticula and soon form oscula at their distal extremity. Hence

the oscular tubes of Z. complicata are always beset with diverticula

to a greater or less extent, thus contrasting with the long, smooth,

usually slender oscular tubes characteristic of LZ. variabilis. The

body-wall is usually thinner and more delicate than in variabilis,

and the natural contour is a pure creamy white, except when

obscured by sediment and the numerous diatoms and other organ-

isms which settle on the exterior of the sponge.

Another distinctive feature of this sponge is the shortness of

the oscular rim—that is to say, of that portion of the oscular tube

immediately surrounding the oscular opening which is not lined

by collar-cells.

(b) Characters of the Spiculation.

(a) Triradiate and Quadriradiate Systems.—(1) The ordinary

triradiates, such as are found in every specimen (text-fig. 91, fig. 1

a-d, &c., p. 364), have the rays slender and tapering gradually to

sharp points. The unpaired ray is straight and distinctly longer than

the paired rays, or at least equal to them in length; the exceptions

to this rule are so rare that they may be termed abnormalities. The

paired rays show a more or less distinct double curvature; proxi-

mally they slope very slightly backwards for about two-thirds of

their length, while at their distal extremity they curve forwards

rather more sharply. The unpaired angle is only slightly greater

than 120°; so that the system often appears nearly equiangular,

but is never quite so. The usual length of the unpaired ray is

* Tam informed that Zostera-beds similar to those at Roscoff occur also at

Jersey, and it is probable that from them come the specimens of this sponge sent

out by Hornell’s Zoological Station, and found in various collections with the label

Leucosolenia contorta.

PROF. E. A. MINCHIN ON THE BRITISH

Text-fig. 91.

| Dec. 13,

1904. } SPONGES OF TH£ GENUS LEUCOSOLENIA, 365

from 100 to 120m, the thickness about 6; the paired rays

range from 75 y to 90 uw in length, with a thickness of about 7 pu.

(2) Besides the ordinary triradiates just described, there occur in

some specimens triradiates with shorter and more thickened rays

(text-fig. 91, fig. 2d—f, p. 364; text-fig. 93, fig. 7c, p. 369). The

proximal curve of the paired rays is scarcely noticeable ; while the

distal curve is rather accentuated. The length of the rays is about

70 « or rather more in some specimens, the thickness 9 or 10 yp.

(3) The ordinary quadriradiates have the basal rays similar to (1)

and the gastral rays of moderate length, curving forwards at the

tip, smooth and tapering evenly to a point (fig. 17). The gastral

vay is implanted on the unpaired ray of the basal system, distinctly

behind the central point of the junction of the three rays compos-

ing it. In some specimens the quadriradiates are, on the average,

of slightly larger dimensions than the triradiates. (4) Occasionally

a gastral ray is found developed on the thickened triradiates (2),

but this is rather a rare type of spicule (text-fig. 93, fig. 7e, p. 369).

(b) Monaxon Spicules—Three kinds occur constantly, and are

the most diagnostic feature of the species. (1) Large spicules with

distinct lance-heads (text-fig. 91, figg. lo—lq, p. 364). The cylin-

drical shaft is always more or less curved, sometimes irregularly, and

tapers rather rapidly to a sharp point at the proximal extremity,

but at the distal end remains of even thickness or diminishes only

very slightly and almost imperceptibly up to the large, broad, sharp-

pointed lance-head. The length is usually from 190 to 280 p, the

thickness 9 or 10. (2) Large spicules without lance-heads, be-

longing to the category of refringent monaxons(text-fig. 91, fige. 1m,

1m, p. 364). These are generally fewer in number than (1), and

each is usually nearly straight or but slightly curved. The shaft

is thickest about one-fourth of its length from the proximal end,

whence it tapers rapidly to a point proximally and very gradually

to a sharp point distally. Sometimes there is a slight indication

of a rudimentary lance-head distally, but usually there is none.

Length usually about 200 nu, greatest thickness 8or9 px. (3) Small

and slender spicules, usually with no trace of a lance-head, and,

with rare exceptions, perfectly straight (text-fig. 91, figg. 17

to 1 @, &e.). The shaft is thickest close to the proximal end,

where it tapers rapidly to a point. From the region of greatest

thickness the shaft tapers extremely gradually to the very sharp

Explanation of Text-fig. 91 (opposite).

Spicules of Leucosolenia complicata.

Figg. la—-1gq. Spicules of a quite normal specimen from Roscoff, one of the same lot

as that figured in Lankester’s ‘ Treatise on Zoology,’ part ii. Porifera, p. 5, fig. 4.

a-d, triradiates;. e-2, quadriradiates; 7-1, slender monaxons; m, », large

monaxons without lance-heads; o-g, large monaxons with lance-heads. —

Figg. 2-21. Specimen dredged on Duke Rock, Plymouth (Garstang), showing

thickened triradiates in addition to ordinary forms. a, ordinary triradiate ;

6 & c, quadriradiates ; df, thick triradiates; g-i, slender monaxons; 7, large

monaxon without barb; & & /, large barbed monaxons.

366 * PROF. E. A. MINCHIN ON THE BRITISH | Dee. 13

distal extremity, which in some specimens shows slight indications

of a lance-head, but more usually does not. These spicules are, as

a rule, short, from 70 to 140 » in length, the greatest thickness

about 3; but they are subject to extraordinary variations in

length, which are described in more detail below.

Variations of the Spiculation.—(1) As regards the relative number

of the spicules. In some specimens no examples of the thickened

triradiates (a, 2) are to be found, or only after much searching; in

others, on the other hand, they are relatively abundant (text-

fig. 91, figg. 2d-f, p. 364), and by their presence give a distinct

facies to the general spiculation, which might lead at first sight to

the impression that the specimen represented a distinct species or

variety. The relative numbers of triradiates and quadriradiates

are also subject to great variation.

The large barbed monaxonsare always abundant, and constitute

a marked feature of the species. Even more characteristic are the

small slender monaxons (6, 3), which are usually very abundant ;

but in some specimens they are relatively scarce, and require to be

searched for carefully. ‘They are also very liable to be broken, by

reason of their slender proportions. As pointed out below, through

Haeckel having overlooked the monaxons in some specimens, he

was led to make two species, pinus and conplicata, characterised

by their presence and absence respectively. I have never, how-

ever, found the small monaxons entirely absent in any specimen

I have examined, not even in the specimen identified by Haeckel

himself as complicata (text-fig. 92, fige. 3 a—3 m, p. 367). The large

unbarbed monaxons (6, 2) are usually much less abundant than

the other two kinds, but can always, in my experience, be found.

(2) As regards the form and size of the spicules. The triradiate

systems vary considerably, both in length and thickness of the

rays. Any specimen shows a considerable range of variations in

this respect ; but in some specimens the spicules show a tendency

to be constantly smaller, in others again constantly larger. The

greatest extremes of variation that have come under my notice, as

regards length of the rays, are shown by the specimens Nos. 3 and 9

of my list given below. In the former, which was a very small

Explanation of 'Text-fig. 92 (opposite).

Spicules of Leucosolenia complicata.

Pigg. 3a-3m. Specimen from Scarborough in Canon Norman’s collection, identified

by Haeckel as Ascandra complicata. a-c, triradiates; d & e, quadriradiates ;

JF, slender monaxons (absent according to Haeckel), showing distinct traces of

barbed heads; j-l, large barbed monaxons; im, large monaxon without barb.—

Figg. 4 a-4.7. Specimen from North Harbour, Peterhead Beach (British Museum,

Bowerbank Collection, No. 987), showing the remarkable length reached by some

of the slender monaxons (d—7) and the small size and relative slenderness of the

large barbed monaxons (a-c).—Figg. 5 a-5 h. Monaxon spicules of a specimen

from the British Museum (Reg. No. 15.1.9.82-34), showing the manner in which

some of the slender monaxons (a—d) are not only of great length but of unusual

thickness and curvature. Other monaxons of the slender type are quite normal

(e-g). hk, one of the large barbed monaxons.

368 PROF. E. A. MINCHIN ON THE BRITISH [ Dee: 13,

colony, the unpaired rays vary from 70-90 p, the paired rays from

65-80 p in length (text-fig. 93, fige. 6a-6h, p. 369). In the latter

(text-fig. 93, figg. 8a-8/, p. 369) the unpaired rays vary from

120-170 p, the paired from 105-125 ». With the increased size

the rays also tend to increase in thickness, reaching 10 p or —

slightly over in their thickest part. This is especially well seen

in a specimen sent me by Topsent from Banyuls-sur-Mer, which

is remarkable for the general thickness of its triradiates (text-

fig. 93, fige. 9a-9d, p. 369). It is interesting to note that it is

in those two specimens with exceptionally large triradiates that I

found barbed monaxons of the largest size, reaching a length of

425 pp in No. 9 and 360 ,. in Topsent’s specimen, while in No. 3

these spicules were below the average in length (150-200 p); the

small monaxons, on the other hand, did not show any noteworthy

variation in those three specimens. Topsent’s specimen possesses

further interest as being the only example of this sponge which I

have seen from the Mediterranean.

The large barbed: monaxons (0, 1) show great variations as

regards curvature and length, but remain remarkably constant in

their general appearance, as well as in thickness. Only in one spe-

cimen, No. 4 of my list, have I remarked a tendency to be below the

normal in this respect (text-fig. 92, figg. 4a—4c, p. 367). In length

they may vary from 80 to nearly half a millimetre, perhaps even

more in some cases. The unbarbed large monaxons vary from

straight to slightly curved, and may also varyin dimensions. But

the greatest variations ave shown by the slender monaxons (b, 3).

In the first place, while, as a rule, they show no trace of a lance-

head, in some cases they exhibit very distinctly a rudimentary barb

at the distal end. This is the case in the specimen in Canon

Norman’s collection identified by Haeckel as complicata, No. 1 of

my list (text-fig. 92, fige. 3-3 7, p. 367). But the greatest variation

is seen in length. In almost every specimen they vary greatly in

Explanation of Text-fig. 93 (opposite).

Spicules of Lewcosolenia complicata.

Figg. 6a-6h. Spicules of one of the type specimens of Bowerbank’s Leucosolenia

contorta (British Museum, Bowerbank’s Collection, No. 988, left-hand middle spe-

cimen), showing triradiate systems of unusually small size. a & 6, triradiates ;

c,a quadriradiate; d & e, slender monaxons showing distinct barbs ; f & g, large

barbed monaxons; /, a large monaxon without barb.—Figg. 7 a-77. Spicules of

a specimen in the British Museum (Reg. No. 95.4.6.1) labelled “ Leacosolenia

hotryoides”’ in Bowerbank’s handwriting and “type sp.” in Carter’s handwriting.

a & 6, ordinary triradiates ; c, a thickened triradiate; d, an ordinary quadri-

radiate; e, a thickened quadriradiate; f & g, slender monaxons; h, a large

monaxon without barb; i & 7, large barbed _monaxons.—Vigg. 8 a—-8f. Spicules

of a specimen in the British Museum (Reg. No. 95.4.6.2) labelled “ Lewcosolenia

botryoides” in Bowerbank’s handwriting and “ type sp.” in Carter’s handwriting,

showing all the spicules above the average in size. a, a triradiate; 5, a quadri-

radiate; c &d, slender monaxons; e & f, large barbed monaxons.—Figg. 9 a—9 d.

Quadriradiate spicules of a specimen from Banyuls-sur-Mer, sent to me by

Topsent, showing the unusually large size and thickness reached by some of the

spicules (6 and d).

1904. | SPONGES OF THE GENUS LEUCOSOLENIA. 369

: |

Text-fig. 93.

fa)

Proc. Zoou. Soc.—1904, Vou. Il. No. XXIV. 24

370 PROF, E. A. MINCHIN ON THE BRITISH [ Dec. 13,

this respect; but in the specimen (No. 4) already referred to as

having a tendency to diminution in the thickness of the large

monaxons, the slender forms are found varying from short ones

of 50 in length to long hair-like forms exceeding half a milli-

metre in length (text-fig. 92, figg. 4d-49, p. 367). The monaxons

showing this extraordinary development also tend to become

abnormal in form; for whereas the ordinary examples of this

type of spicule are characterised by their perfect straightness and

absence of any curvature, the elongated forms become irregularly

curved and wavy. I consider the special development of these

spicules in this specimen as an abnormality, due perhaps to some

special conditions of its habitat, of which unfortunately no record

is preserved. Another specimen in which the slender monaxons

are greatly elongated, but to nothing like the same extent, is seen

in No. 10 (text-fig. 92, figg. 5a-5g, p. 367). This specimen is

unique, in my experience, in another respect. While in all other

specimens I have seen the monaxons are distinguished by their

slenderness, whatever their length, from the other classes of

monaxons present, in this specimen these slender forms show a

tendency to become thickened and at the same time evenly curved,

thus indicating a transition to the thicker types.

This sponge, as has been said, was first described by Montagu

in 1812 under the name Spongia complicata ; but its distinctness

was not generally recognised until Haeckel in 1872 gave it precise

characters, thus determining definitely the species which must

henceforth bear the name complicata, whatever Montagu’s specimen

may have been.

Unfortunately, Haeckel overlooked the small monaxons in some

of his specimens, for which he retained the name complicata, but

saw them in a specimen for which he founded a new species, pinus,

which must therefore rank as a synonym. There are, moreover,

other species in his monograph which must be put as synonyms

of complicata until further evidence be forthcoming as to their

distinctness.

First and foremost among these is Ascortis fabricii, the Lewco-

solenia fabricii of Oscar Schmidt. There is nothing whatever in

Schmidt’s or in Haeckel’s descriptions to differentiate this species

from complicata. The diagnostic feature, absence of quadri-

yadiates, is Insufficient, since Haeckel records the frequent presence

of this type of spicule, and founds on it a “ connexive Varietit”

which he names Ascandra fabricti. This proves that the diagnostic

absence of quadriradiates was in reality merely scarcity, a common

variation of this sponge. The small monaxons are not mentioned

by either writer, but Schmidt’s brief description testifies to his

having made a most cursory examination of the sponge, and

Haeckel overlooked them also in his complicata, so that this point

1904. | SPONGES OF THE GENUS LEUCOSOLENIA. 371

counts for nothing. Haeckel’s figure of fabricii, evidently much

reconstructed and embellished, may be taken to represent an

arborescent specimen of complicata.

Secondly, Haeckel’s species Asculmis arnvata appears to me to be

founded simply on the converse variation of complicata—that is

to say, on specimens (Olynthus-forms) in which triradiates were

scarce. Here also we have a ‘“connexive Varietiit,” Ascandra

armata. There is nothing in Haeckel’s description to separate

this species from complicata. Haeckel’s two varieties of Asculmis

armata, named by him var. norvegica and var. pocillwm, are formed

on variations in the length of the gastral rays, which can be found

im any specimen of the sponge *.

T consider it also highly probable that Haeckel’s Ascyssa acufera

will prove, when re-examined, to be a specimen of this species.

Since in Lewcosolenta the monaxons are always the first spicules

to appear at the metamor phosis, every species of Lewcosolenia is at

first an “ Ascyssa.”

Haeckel further made two varieties of complicata. The first he

named hispida, which was characterised by having the “lateral rays

straight or only slightly curved; monaxons also slightly curved,

with lance-head scarcely distinct.” The second, named amboides,

has “lateral rays strongly curved in the form of an S$; monaxons

also slightly curved, with lance-head sharply distinct.” Since

all the variations of the spicules mentioned can be found in any

specimen, it is not necessary to cumber taxonomy with these

names.

Breitfuss, in his memous on calcareous sponges [2-5 |, seems to

have consistently confused this species with Clathrina contorta ;

he has certainly done so, as pointed out below, in his ‘ Catalogue

of the Calcarea in the Berlin Museum’ [3]. In his work upon the

calcareous sponge-fauna, of the White Sea ty he figures (pl. 1. fig. 1),

under the name Ascandra contorta,a sponge which is certainly not

the species with which it is identified, but is clearly a Lewcosolenia,

and resembles the ordinary arborescent form of complicata. The

description of the spiculation is inadequate even for determining

the genus, but, so far as it goes, agrees with L. complicata.

The diagnostic features of this species are :—(1) the elongation

of the unpaired ray of the triradiate systems relatively to the

lateral rays ; (2) the presence of two distinct forms of monaxons—

the first small, straight, slender, usually without a barb at the

distal extremity; the second large, curved, thick, usually with a

distinct barb: the former are often scarce, but never apparently

entirely lacking, pace Haeckel.

* Haeckel at first considered the Spongia pocillum of O. F. Miller (1776, Zool.

Dan. Prodr. p. 256) and Fabricius (1780, Faun. Greenland, p. 449) to be identical

with Asculmis armata; but as it is quite impossible to identify Spongia pocillain

from their descriptions, it must be considered a nomen nudum, of no systematic

importance.

24*

372 PROF. E. A, MINCHIN ON THE BRITISH [ Dec. 13,

(d) List of Specimens examined *.

(a) From Canon Norman’s Collection,

1, Dried specimens collected at Scarborough by Bean, sent to

Haeckel for examination, and returned by him with the

following label in his handwriting :—

“ Ascandra complicata H.

“ (Spongia complicata Montagu).

“Scarborough (Bean).”

These specimens were of the utmost importance, as they

showed that the small monaxons supposed to be absent in

Ascandra complicata H.. and present in A. pinus H. were

simply overlooked by Haeckel in the specimens referred

by him to the former species (see text-fig. 92, figg. 3 f-2,

p. 367).

2, Dried specimens labelled ‘“Zezcosolenia contorta, Sinner

in Bowerbank’s handwriting. According to information

given me by Canon Norman, the specimens seen by me

were not sent to Haeckel, but are of the same lot as the

type sent to him, and are equally types of Ascandra

contorta H. My preparation shows typical spicules of

Leucosolenia conplicata mingled with spicules of Clathrina

coriacea. These two species often grow in the closest

proximity; and I have a series of sections of Clathrina

coriacea showing tubes of Leucosolenia complicata growing

side by side with those of the Clathrina.

(6) From the British Museum.

3. Bowerbank Coll., No. 988. Seven dried specimens stuck

on a card, and labelled in Bowerbank’s handwriting

“« Leucosolenia contorta, Guernsey.” These specimens are

the types of Bowerbank’s species LZ. contorta, figured by

him in Mon. Brit. Spong. pl. 1. figg.5-10. One specimen

is much larger than the others; it 1s stuck at the top of

the card, over the middle. The other six specimens are

arranged in two vertical rows, three in each row, along the

two sides of the card. Of the seven specimens I have

examined six, that is to say, all except the right lower

specimen, which is very small. The large specimen

(Bowerbank’s fig. 7) is a Clathrina sp. which agrees with

Haeckel’s Ascandra contorta in spiculation, except for the

absence of monaxons, which I have not been able to find.

The five smaller specimens examined by me are one and

all of them unmistakable specimens of Leucosolenia com-

plicata, but being very young colonies the spiculation is

sometimes rather aberrant, especially in the triradiates ;

the three types of monaxons, however, show the specific

characters quite invariably. The spicules are generally

* In the lists of specimens enumerated by me, I count only public specimens, so

to speak, without mentioning the many examples I have studied in my own or other

private collections. ¥

1904. ] SPONGES OF THE GENUS LEUCOSOLENIA, 373

small, as shown in text-fig. 93, fig. 6, p. 369, which is drawn

from the left-hand middle specimen. The large monaxons

are often short and stumpy, especially in the right-hand

middle specimen, which is further unique, in my ex-

perience, in that the unpaired rays of some of the

triradiates are shorter than the paired rays. In general,

these young specimens do not show the characteristic

elongation of the unpaired ray so markedly as the larger

specimens examined by me.

4, Bowerbank Coll., No. 987. Labelled, in Ridley’s hand-

writing, “ Grantia botryoides. North Harbour, Peterhead

Beach. No. 4, 1851. J.S. Bowerbank.” A somewhat

abnormal specimen of Z. complicata (see text-fig. 92,

figg. 4a-49, p. 367).

5. Bowerbank Coll., No. 986. Bowerbank’s label, copied by

Ridley, is as follows—“ Grantia botryoides John. Guern-

sey. Mr. Buckland.”

6. Register No. 72.5.4.1 a. ‘“ Leucosolenia botryotdes,’ Vigo

Bay, Saville Kent. :

7. Bowerbank Coll., No. 992. Dried specimens stuck on a

card, labelled “Grantia botryoides,’ locality Orwell

River.

8. Register No. 95.4.6.1. Labelled “ Leacosolenia botryoides ”

in Bowerbank’s handwriting and “type sp.” in Carter’s

handwriting, meaning apparently Bowerbank’s type (see

text-fig. 93, fige. 7 a—-7j, p. 369).

9. Register No. 95.4.6.2. Labelled exactly as the last (see

text-fig. 93, fige. 8a-8 f, p. 369).

10. Register No. 85.1.9.32-34. Labelled “ Calcarea, about

15 fathoms off Port St. Mary, I. of Man. J. Lomas, Esq.”

Rather an abnormal specimen of complicata (see text-

fig. 92, figg. 5 a—5 h, p. 367).

11. Register No. 85.3.6.6. A beautiful specimen labelled

“2% Leucosolenia botryoides. Jersey, Saville Kent,” in

Carter’s handwriting.

12. No. 1780. Labelled “ Ascandra contorta H. Jersey”™*.

2, LEUCOSOLENIA VARIABILIS.

2 Spongia confervicola Templeton, 1836, Magazine of Nat. Hist.

ix. p. 470, fig. 67.

Grantia botryoides var. himantia Johnston, 1842, Brit. Spong.

and Lith. p. 179, pl. xxi. fig. 3.

Leucosolenia (Leuciria) variabilis Haeckel, 1870, Jena. Zeitschr.

v. p. 243.

* In the Catalogue of the Calcarea of the Berlin Museum, published by Breit-

fuss [3], this specimen, No 1780, and others are put down with the labels copied from

the bottles, without, apparently, any attempt at verification. The author is evidently

unacquainted with <Ascandra (Clathrina) contorta, otherwise a glance at the

specimen would have prevented his making this error: but it is surprising that he

did not examine the spiculation.

374 PROF, E. A. MINCHIN ON THE BRITISH [ Dec. 13,

Sycorrhiza corallorrhiza Haeckel, 1870, |. c. p. 249.

Ascandra variabilis Haeckel, 1872, Kalkschwimme, 11. p. 106,

iu. pl. 16. figg. 4 @-4/ and pl. 18.

Ascortis corallorrhiza Haeckel, 1872, 1. ec. p. 78, pl. 11. fig. 4,

pl. 12. figg. 4 a—47.

Leucosolenia botryoides Bowerbank, 1874, Mon. Brit. Spong.

ii. pl. i. figg. 1-4.

Leuconia somesii Bowerbank, 1874, 1. ce. p. 334, pl. xci.

fige. 6-17.

Ascandra tenuis Schuffner 1877, Jena, Zeitschr. xi. (n. F. iv.),

p. 406, pl. xxv. fig. 8.

Ascandra variabilis Bowerbank and Norman, 1882, Mon. Brit.

Spong. iv. p. 227.

Ascandra botryoides Fristedt, 1885, K. Vetensk.-Akad. Hand-

lingar, xxi. no. 6, p. 9.

Leucosolenia variabilis Topsent, 1891, Arch. Zool. Exp. (2) 1x.

p. 525.

Leucosolenia variabilis Topsent, 1894, Rev. Biol. Nord France,

Vil. p. 2

Leucosolenia variabilis Minchin, 1896, Ann. Mag. Nat. Hist.

(6) xvinl. p. 359.

Ascandra variabilis Breitfuss, 1898, Arch. f. Naturges. [sin. i.

p. 215.

Ascandra variabilis Breitfuss, 1898, Arch, f. Naturges. lxiv. 1.

p- 286. ;

Ascandra corallorrhiza Breitfuss, 1898, 1. e. p. 285.

_ Ascandra variabilis Breitfuss, 1898, Mém. Acad. St. Péetersbourg,

(8) vi. p. 16.

Ascandra corallorrhiza Breitfuss, 1898, 1. c. p. 9.

Ascandra variabilis: Breitfuss, 1898, Ann.; Mus. Zool. St.

Pétersbourg, 1898, p. 28.

Ascandra corallorrhiza Breitfuss, 1898, 1. ec. p. 17.

Leucosolenia variabilis Minchin, 1900, in Lankester’s Treatise

on Zoology, ii. Sponges, p. 5, fig. 5.

Ascandra variabilis Arnesen, 1901, Bergens Mus. Aarbog, 1900,

No. 5, p. 15.

Ascandra corallorrhiza Arnesen, 1901, 1. ec. p. 14.

Leucosolenia variabilis Rousseau, 1903, Mém. Soc. Malae.

Belgique, xxxvii. p. 8, fig. 4.

Leucosolenia variabilis Allen, 1904, J. Mar. Biol. Assoc. n. 8. vil.

p. 185.

(a) Hxternal Characters.

L. variabilis is known to me both in the spreading and the

bushy form. Of the former type I have two typical specimens,

both from Roscoff. A portion of one of them has been figured

by me elsewhere [16, p. 5, fig. 5]. It was growing on a granite

rock, and an attempt was made to detach the piece of stone to

which the sponge was attached by means of hammer and chisel ;

as aresult of this somewhat violent treatment, the slab broke

1904. } SPONGES OF THE GENUS LEUCOSOLENIA. 37D

across under the middle of the sponge-colony, which was thus

detached in two halves and somewhat damaged, but still showing

well the peculiarities of this mode of growth. The spiculation ot

this specimen was found to be normal in all respects. My second

specimen of the spreading form was removed entire from the rock

on which it grew ; it is a small colony remarkable for the close

network of basal tubes, giving the sponge almost the appearance

of a Clathrina, were it not for the characteristically large oscular

tubes. The spiculation of this specimen shows a comparative

scarcity of monaxons and unusually small triradiate systems; in

other respects, however, the characters are typical (text- fig. 95,

figg. 14 a-14f, p. 379). Johnston [14, pl. xxi. fig. 3] has figured

a typical spreading specimen of variabilis under the name of

Grantia botryoides var. himantia, which Haeckel has wrongly

placed as a synonym of Clathrina coriacea.

L. variabilis occurs most commonly in the bushy form as a

compact reticulum of fine anastomosing basal tubes from which

arise the stouter oscular tubes, often closely packed and of con-

siderable length. In this form it may be found in rock-pools

attached between the stems of algze, or creeping over the algex

themselves. In the former situation the basal portion of the

sponge is often half buried in mud and sediment, and, doubtless

in consequence of this, the oscular tubes grow to a oveat size and

length. I have such a Specimen from Plymouth, in which the

oscular tubes average 1*°5 cm. in length, reaching in some cases

2 em.; the spiculation of this specimen (text-fig. 95, figg. 15 a-15f;

p. 379) is remarkable for the large size of the triradiate systems.

It is more usual, however, for the sponge to form a creeping

erowth over the alge themselves—either twining amongst the

filaments of confervee, as Haeckel’s fig. 6 on pl. 18 of the ‘ Mono-

graphie’ [13] shows fairly correctly, or spreading over the stems

of stouter seaweeds. These creeping forms are commonly found

at Roscoff in situations where they are left dry at all tides, growing

amongst the stems of the dense growth of alge covering isolated

boulders s on the seashore. The youngest colonies form a delicate

network spreading over the seaweed stems and sending up oscular

tubes at intervals, thus having a form similar to the spreading

colonies already described ; but with further growth the basal

tubes form a compact tangled mass from which long oscular tubes

arise or hang down as the case may be.

In all the modifications of form exhibited by this sponge the

most constant feature is to be found in the oscular tubes, which

show little or no tendency to throw out diverticula except near

their base. Hence the sponge shows numerous oscular tubes

arising from the basal reticulum, each long, slender, generally

slightly curved and smooth, 7. e. free from diverticula from near

their base up to the oscular opening. We find the sharpest con-

trast, as has already been stated above, with LZ. complicata in this

respect, and the typical arborescent form of the latter is never

found in Z. variabilis.

376 PROF, E, A. MINCHIN ON THE BRITISH [ Dee. 13,

Haeckel has given a plate (Monographie, pl. 18) intended to

represent the form-variations of Z. variabilis, but, in my opinion,

the greater number of his figures are untrue to nature. The

most charitable interpretation that can be put upon this plate is

to suppose that many of the specimens figured are incorrectly

determined and are not ZL. variabilis at all. Thus fig. 12 is

evidently a Clathrina, and fig. 15 is either a contracted Clathrina,

probably coriacea, or possibly a specimen of variabilis im which,

owing to rough usage, all the oscular tubes have been knocked

off. Fig. 2 probably represents some of the Sycons which, as I

have stated below, are commonly found growing in company with

L. variabilis, and the same interpretation possibly applies to

fige. 1 and 3, the latter being contracted ; what figg. 10 and 13

may be I should not lke to assert, but they are certainly not

specimens of variabilis. On the other hand, figg. 8 and 14 are

probably specimens of variabilis, which owe the extraordinary

curves exhibited by their oscular tubes to shrinkage as the result

of desiccation. The imaginative figures given by Haeckel on this

and other plates are responsible for the general opinion, far from

being true, that any Ascon may assume any form in which any

other species of Ascon occurs.

As compared with other species, I find a tendency in variabilis

for the oscular rim to be of greater length. In young specimens

this issometimes very marked indeed. The body-wall of variabilis

is generally thicker and stronger than in Z. complicata, and the

sponge is evidently hardier, as seen from the more exposed

situations in which it grows.

b) Characters of the Spiculation.

(a) Triradiate and Quadriradiate Systems.—(1) The ordinary

triradiates (text-fig. 94, figg. 10a—10c, &c., p. 377) have the rays

slender or of moderate thickness and tapering gradually to sharp

points. The unpaired ray is straight, and distinctly shorter than,

very rarely as long as, the paired rays. The unpaired angle is always

much greater than 120°. The paired rays usually show a distinct

double curvature, each ray curving first backwards, then forwards

Explanation of Text-fig. 94 (opposite).

Spicules of Leucosolenia variabilis.

Figg. 10 a—10 p. Spicules of a specimen with monaxons not reaching a large size,

from Roscotf. a-c, ordinary triradiates; d, e, T-shaped triradiates (in this

specimen not greatly thickened) ; f, g, brackets; h, 2, ordinary quadriradiates ;

Jj, bracket with long gastral ray ; &, U, straight slender monaxons ; m-~p, ordinary

curved monaxons.—Figg. 11 a—1lg. Spicules of another specimen from Roscoft,

in which all the monaxons are small. a, b, triradiates; c, T-shaped quadri-

radiate; d, ordinary quadriradiate; e, straight slender monaxon; f, g, curved

monaxons.—Figg. 12 a—-12,7. Spicules of a specimen from Roscoff in which the

monaxons vary up to a large size. a, triradiate; 6, quadriradiate; e, d, straight

slender monaxons; e-j, curved monaxons.—Fig. 13. Monaxon of large size from

a Roscoff specimen.

SPONGES OF THE GENUS LEUCOSOLENIA. aad

1904.]

378 PROF. E. A. MINCHIN ON THE BRITISH [ Dec. 13,

at the distal extremity, but the curvature may be very slight, all

three rays then being practically straight. The unpaired rays

vary from 65-83 w in length and are about 6 or 7 in thickness ;

the paired rays range from 80-100 u in length, with a breadth of

7 or 8. (2) In most specimens thicker triradiates occur (text-

fig. 96, fig. 21 ¢, p. 381) with the anterior angle approaching 180°, so

that the spicule appears nearly T-shaped. The paired rays, seen

in facial aspect, may appear nearly straight, but more usually the

proximal backward curve of the ray is short, while the distal

forward curvature extends over more than half the length of the

ray. The system as a whole is strongly concave on the gastral

face, so that when seen in facial aspect from the gastral side one

of the two paired rays is usually very much foreshortened, and

therefore appears different in length and curvature from the

other. The rays reach 10 in thickness. (3) In all specimens

peculiarly modified triradiates occur which may be termed brackets

(text-fig. 94, figg. 10 f, 10g, p. 377). ‘The two paired rays are

straight or curved, and more or less normal as regards length and

proportions, though in some cases rather slender; the unpaired

ray, on the other hand, is short and strongly curved out of the

plane in which the paired rays lie, so that in a facial view of the

spicule it is not in focus at the same time as the paired rays.

The unpaired ray is often irregularly curved, and may be reduced

to a mere knob or even to the vanishing point. (4) The ordinary

quadriradiates (text-fig. 94, figg. 104, 102, &e., p. 377) have the

basal rays similar to (1); the gastral ray is of moderate length,

smooth, and evenly curved at the tip. (5) The thickened T- ‘Shaped

triradiates, as in (2), may develop a gastral ray, which is then

rather thick and set very upright on the basal system (text-fig. 94,

fig. lle, p. 377; text-fig. 96, fig. 21g, p. 381). (6) The brackets

may also bear a gastral ray, which is then usually of very great

Explanation of Text-fig. 95 (opposite).

Spicules of Leucosolenia variabilis.

Figg. 14a-14f. Spicules of a specimen from Roscoff showing triradiates of un-

usually small size. a, 6, triradiates; c, quadriradiate; d, e, curved monaxons ;

J, straight monaxon.—Figg. 15 a-15 f. Spicules of a specimen from Plymouth

showing triradiate systems unusually large. a, 6, triradiates ; c, quadriradiate;

d, e, curved monaxons; f, straight monaxon.—Figg. 16a-16g. Spicules of a

specimen from Bantry Bay, Ireland, in Norman’s collection, identified by Haeckel

as Ascandra variabilis. a, trivadiate; b, c, quadriradiates; d, bracket with

greatly elongated gastral ray ; e, f; curved monaxons; g, bayonet-like monaxon.

—Figg. 17 a, 176. Bayonet-like monaxons of a specimen from a tide-pool below

the Hoe, Plymouth.—Figg. 18 a-18 e. Spicules of the specimen from Brighton

Aquarium figured by Bowerbank (‘ British Spongiade,’ 11. pl. xci. fig. 7) under

the name Lewconia somesii (Brit. Mus. Bowerbank Coll. 1019). a, 6, tri-

radiates ; c, quadriradiate; d, e, curved monaxons showing their great variation

in size.—Figg. 19a-19h. Spicules of another of Bowerbank’s specimens of

Leuconia somesii (I. c. pl. xci. fig. 6) (Brit. Mus. Bowerbank Coll. 1017) show-

ing a tendency to develop abnormal forms of the triradiates and monaxons of

great length, irrespective of thickness. a-—e, triradiates ; f—h, monaxons; h, on

account of its great length, has been drawn in two pieces.

380 PROF, E, A, MINCHIN ON THE BRITISH [ Dec. 13,

length, reaching 120 p or more, and tapering gradually to a sharp

point (text-fig. 94, fig. 107, p. 377; text-fig. 95, fig. 16 d, p. 379).

(b) Monaxon Spicules—Three kinds occur, two invariably, the

third only to be found in certain specimens. (1) The ordinary

monaxons (text-fig. 94, figg. 12e-12 7, &e., p. 377) ave curved and

have distinct lance-heads, and vary greatly in size, in the same or

in different specimens, ranging from 80 to 320 » in length and

2 to 9 in thickness; they cannot, however, be divided into two

distinct classes as has been done by Haeckel, since every possible

gradation is found from the smallest to the largest. The shaft is

generally more or less straight for its proximal half or two-thirds

and then has a slight bend, the distal fourth or sixth, however,

with the lance-head, being nearly straight. The shaft is thickest

in the region between a third and a half of its length from the

proximal end. The proximal half or third is cylindrical and

tapers abruptly to a point; the distal half or two-thirds tapers

gradually and almost imperceptibly, dwindling to about half the

greatest thickness when it reaches the small lance-head. (2) Very

slender monaxons occur (text-fig. 94, fige. 10%, 107, 12, 12d,

p. 377), easily found on account of their very refringent appearance

and sharp outline, though they are much less abundant than (1).

They have straight, very slender shafts and distinct lance-heads,

and vary from 70 to 110 in length, never reaching a large size.

(3) In some specimens fairly large bayonet-like spicules occur

(text-fig. 95, figg. 16g, 17a, 176, p. 379) with a sharp bend in the

middle, sometimes represented only by a swelling. They reach

about 200 » in length, varying in thickness from about 7-10 p.

Variations of the Spiculation. (1) As regards the relative numbers

of the Spicules—In some specimens the triradiates and quadri-

radiates are all of the ordinary slender type (a, 1 & 4) and none

of the thickened T-shaped forms (as 2 & 5) are to be found, or

only after much searching. In others they are more easily found.

The brackets, with or without gastral rays (a, 3 & 6), are never

very abundant and often scarce. Of monaxon spicules, the barbed

forms (6, 1 & 2) are always present more or less abundantly,

but in some specimens I have not succeeded in finding the large,

bayonet-like forms (6, 3), in spite of much searching ; in others

they are easily found, and I have also come across one i situ in

the body-wall of the sponge.

Explanation of Text-fig. 96 (opposite).

Spicules of Sycon sp. and Leucosolenia variabilis.

Figg. 20a-20w. Spicules of a Sycon found commonly occurring in company with

L. variabilis, specimen from Roscoff. a-e, dermal triradiates; f-j, tubar tri-

radiates; &, 7, quadriradiates from oscular rim; m-—p, gastral triradiates and

quadriradiates ; g-w, monaxons.—Figg. 21a—p. Spicules of a specimen of

L. variabilis from Bergen, Norway, identified by Haeckel as Ascandra varia-

bilis (Berlin Mus. no. 417), showing admixture of Sycon spicules similar to those

in the preceding figure. a-c, triradiates of L. variabilis; d-h, quadriradiates

of L. variabilis; i, 7, monaxons of ZL. variabilis; k, 1, tubar trivadiates of the

Sycon; m-p, monaxons of the Sycon.

SPONGES OF THE GENUS LEUCOSOLENIA. 381

1904. |

382 PROF, E, A. MINCHIN ON THE BRITISH [ Dec. 13,

(2) As regards the form and size of the Spicules.—The ordinary

trivadiates and quadriradiates vary considerably in size both m

different and in the same specimens, though here, as in other

species, one specimen may be found characterised by spicules

generally above the average in size, another by spicules below

the average. The smallest triradiates I have observed, occurring

in a specimen from Roscoff, have the unpaired rays about 60 p,

the paired rays about 70 in length (text-fig. 95, fig. 14, p. 379).

The largest I have seen have unpaired rays 95-100 p, paired rays

135-140 p, in length (text-fig. 95, fig. 15). In thickness also they

vary considerably, ranging from 4-10 (text-fig. 96, fig. 21a,

p. 381, text-fig. 95, fige. 15 6, 156, p. 379), and, in the latter case,

only differing from the T-shaped forms by the paired angles and

the even curvature of the paired rays. On the other hand, as

regards form and proportions, the triradiates are remarkably

uniform, the unpaired ray being almost invariably shorter than

the paired rays, usually very obviously so, and only very excep-

tionally equalling them in length.

The brackets are very variable both as regards the form and

degree of development of the unpaired ray, and as regards the

length of the gastral ray when developed; the latter always

greatly exceeds in length, however, the gastral rays of the ordinary

quadriradiates.

The curved monaxons (6, 1) are extremely variable in size.

The normal condition is for every intermediate grade to occur

between spicules of about 80 in length and others three times as

long. In some specimens, however, only small monaxons occur,

in others the majority of them ave large, reaching nearly 400 in

length and correspondingly thickened; but in the latter case

small forms are always present in addition to the large, though

they may be scarce by comparison.

The aberrant specimens of ZL. variabilis from Brighton Aqua-

vium, which were described by Bowerbank under the name

Leuconia somesii (text-fig. 95, figg. 18, 19, p. 379), are characterised

by an extraordinary development of the monaxons, which, mm

addition to ordinary small and large forms, reach in other cases a

great length (as much as 1000) combined with unusual slender-

ness for such a length (2-5 p). The triradiates of these specimens

are also characterised by great range of variation in size, and

by the large number of abnormalities in this class of spicules.

In certain specimens of this sponge which I have been able to

examine—specimens important for the reason that they had all

been through Haeckel’s hands—I was puzzled by finding certain

types of spicules in my preparations in addition to those already

described (text-fig. 96, figg.21 4-21 p). The specimens in question

were, first, four in Norman’s collection (Nos. 1-4 of my list

below); secondly, one in the Berlin Museum (No. 13 of my

list) (text-fig. 96, fig. 21, p. 381). All these five specimens

show the typical spiculation of Leuwcosolenia variabilis, the

specimen from Polperro (No. 5) alone showing an admixture of

1904. | SPONGES OF THE GENUS LEUCOSOLENIA. 383

the spicules of Z. complicata, so that the two species were evi-

dently growing in close proximity, as so often occurs; but all

of them show, as has been stated, additional spicules. Since

L. variabilis forms a dense tangle amongst the seaweeds, hydroids,

and other organisms, and might easily be involved in this way

with quite distinct calcareous sponges, the idea occurred to me

that the spicules in question might belong to another sponge ;

and on examining carefully my specimens of ZL. variabilis from

Roscoff, and picking them over under a lens, 1 was able without

difficulty to trace these spicules to their source. Growing on the

seaweeds, in closest proximity, very often, to the tubes of the

Leucosolenia, were numerous small Sycons, ranging in size from

minute Olynthi to specimens a centimetre or more in height, but

all alike showing a characteristic spiculation, which accounted at

once for the intrusive spicules in my preparations. J have not

been able to identify this sponge in Haeckel’s monograph, neither

among the Sycons nor the Leucons. J! content myself, therefore,

for the present, with describing the spiculation, and leave the

identification of it to a future period, or to others. Five classes

of spicules can be distinguished :—(1) Dermal triradiates (text-

tig. 96, figg. 20 a—20 e, p. 381) tending to be irregular in form, often

nearly equiangular; (2) tubar triradiates (2. e. from the walls of

the flagellated chambers), with long straight unpaired ray, and

shorter lateral rays characteristically curved, the unpaired angle

very obtuse (text-fig. 96, figg. 20/20 J, p. 381); (3) gastral trira-

diates and quadrivadiates, with long straight unpaired ray, shorter

paired rays slightly curved, gastial ray present or absent, unpaired

angle nearly a right angle or less, at any rate considerably less than

120° (text-fig. 96, fige. 20 m-20 p, p. 381); (4) quadriradiates

from the oscular rim, the three basal rays nearly equal in length,

the unpaired angle nearly 180°, so that the spicule is nearly

T-shaped (text-fig. 96, fige. 20 £, 201, p . 381); and (5) monaxons of

very characteristic form, the acer half nearly straight, taper-

ing to a point, the distal half curving evenly outwards, without

any diminution in thickness, to the conspicuous, bluntly pointed

lance-head. It is the presence of these monaxons that was so

often noticed in my specimens of Leucosolenia variabilis ; ib is

probable that such monaxons may often find then way even into

pure cultures of the Leuwcosolenia, for if thrown off by the Sycons

and washed about in the water they might easily come to stick to

the Leucosolenia or to the seaweed on which it was growing.

ce) General Remarks on the Species.

Haeckel was the first to recognize the specific distinctions of

this sponge and to describe accurately its distinctive characters,

although it was common enough in collections long previous

to Haeckel’s monograph. Bowerbank, as has been already pointed

out, figured this sponge and its spicules under the name ZL. botry-

oides, and again in another place as a new species under the name

384 PROF. E. A. MINCHIN ON THE BRITISH [ Dec. 13,

Leuconia somesii; why he should have called these specimens

Leuconia is a mystery to me.

Haeckel gave the species the name variabilis on account of “the

unlimited changeableness of its form as a whole, as well as of its

specific skeleton-structure.” I have already expressed my opinion

upon Haeckel’s figures of the external form ; as regards the spicu-

lation, Z. variabilis is variable certainly, but not more so than other

Ascons. The frequent association, mentioned above, of this species

with a heteroccele sponge, and the constant contamination, so to

speak, of spicule-preparations of the Ascon by spicules not properly

belonging to it, may account for Haeckel’s noticing in this instance

the variability of the spiculation.

Haeckel described variabilis as having two kinds of monaxons,

small and large; butfrom his figure it is evident that the smaller

kind seen by him were the small curved monaxons, and he did not

notice that they are connected by every possible gradation of size

with the large ones. He overlooked the small straight monaxons,

as I must also confess to have done till quite recently. Haeckel

made four specific varieties—cervicormis, confervicola, arachnoides,

and hispidissima,—based on variations of the relative numbers of

the different sorts of spicules, and leading up to his so-called

“‘eonnexive varieties,” distinguished, in his usual way, by ringing

the changes on the generic name, such as Ascaltis, Ascortis,

Asculmis, ov Ascyssa variabilis. None of these varieties appears

to me to have any taxonomic value except hispidissima, which

might be retained for forms such as were named by Bowerbank

Leuconia somesii: i. e. for those in which the monaxons are

excessively developed in size and number to form a furry pro-

tective covering. In my specimens from Roscoff, even from the

same rock, I find some which, viewed with a lens, appear smooth,

others which appear hispid ; the difference between them is

merely one of the length attained by the monaxons (compare

fige. 10-13, text-fig. 94, p. 377). Thesame is true of L. complicata,

and there can be no doubt that these sponges respond readily in

this manner to differences in their surroundings.

Of other species of Ascons in Haeckel’s monograph, I feel no

doubt whatever that his Ascortis corallorhiza is founded on

a specimen of this species with rather large and thick spicules

(compare figg. 15 a-15/, text-fig. 95, p. 379). Here also we have

a connexive variety, Ascandra corallorrhiza, mentioned.

Systematists subsequent to Haeckel have for the most part

recognized and identified this sponge correctly. A specimen,

however, in the British Museum from Trieste, labelled Z. varia-

bilis, is certainly not this species. L. variabilis does not, to the

best of my belief, occur in the Mediterranean ; but it is impossible,

I repeat, to make definite statements about the distribution of

Ascons in the present confused state of their nomenclature *,

* Kirkpatrick, in 1901 (Brit. Mus. Rep. “Southern Cross’ Collections, p. 317),

identified an Antarctic sponge as L. variabilis ; but having been able, by the author’s

kindness, to examine the specimen I am in a position to state that it is not D. varia-

Bilis but a species, apparently new, allied to L. complicata.

1904. | _ SPONGES OF THE GENUS LEUCOSOLENIA. 38

The diagnostic features of this sponge are :—(1) The constant

shortness of the unpaired ray of the triradiate systems relative

to the lateral rays, and the wideness of the anterior angle,

frequently nearly 180°; (2) the two kinds of monaxons, both

barbed—straight refringent ones always slender and small, rela-

tively scarce; and curved ones, varying from very small to very

large, always abundant.

(d) List of Specimens examined.

(a) From Canon Norman’s Collection.

1. Specimen from Bantry Bay, Ireland, with Norman’s label

“* Leucosolenia botryoides,” endorsed by Haeckel ‘A scandra

variabilis.” (See text-fig. 95, figg. 16 a—-16 g, p. 379.)

2. Specimens from Shetland, with label in Bowerbank’s

handwriting “ Grantia botryoides Johnston, more largely

developed than usual.” Also a label by Norman “ Zeuco-

solenia botryoides, very large, Shetland,” across which is

written in Haeckel’s handwriting “A scandra variabilis H.”

* These specimens are of the same batch as No. 8 below.

3. Specimens received by Norman from Haeckel, collected at

Bergen, Norway, with printed label “ Ascandra varia-

bilis H.”

4, Specimen received by Bowerbank as a type of Leucosolenia

contoria, from Guernsey; sent to Haeckel for examination,

and returned with label in Haeckel’s handwriting as

follows :—

“ Ascandra contorta H.

“(Leucosolenia contorta Bwhk.)

“* Guernsey. Bowerbank.”

The specimen is a quite typical variabilis, with a slight

admixture of Sycon spicules.

5. Specimens growing on seaweed with Girantia compressa,

from Polperro, Cornwall, and identified by Haeckel as

Ascandra variabilis: my preparations show a mixture of

spicules of variabilis and complicata.

(6) From the British Museum.

6. Bowerbank Coll., No. 1017. The type specimen of Bower-

bank’s “ Leuconia somesw,” figured in Brit. Spong. iii.

pl. xci. fig.6. With label in Bowerbank’s handwriting—

“ Leuconia somesii Bwk., sent to me by Mr. H. Lee,

15.9.73, said to have been found alive in the aquarium ”

(i. e. at Brighton). (See text-fig. 95, figs. 19a-19h,

. 379.

f Dowereale Coll., 1018. Another specimen with Bower-

bank’s label “ Leuconia somesii, Brighton Aquarium, sent

by Mr. Lee, 5.11.73.” Figured in Brit. Spong. iii. pl. xci.

fig. 8.

8. Serevent: Coll., 1019. Another specimen with label

similar to the last. Figured in Brit. Spong. iii. pl. xci.

fig. 7. (See text-fig. 95, figeg. 18 a-18¢, p. 379.)

Proc. Zoou. Soc.—1904, Vou. II. No. XXV, 25

386 PROF. E. A. MINCHIN ON THE BRITISH [ Dee. 13,

Yes)

. Bowerbank Coll., 985. Specimen figured in Brit. Spong.

iii. pl. iii. fig. 1, with label in Bowerbank’s handwriting

“ Leucosolenia botryoides, Shetland.” A quite normal

specimen of variabilis with fairly large spicules.

10. Bowerbank Coll., 979. Labelled in Bowerbank’s hand-

writing “ Leucosolenia botryoides, Shetland.”

11. Bowerbank Coll., 964. Label as in the foregoing.

12. Register No. 47.9.7.112. Label “Grantia botryoides var.”

Holy Island, Johnston Collection.

[Register No. 83.12.4.17, labelled “ Ascandra varia-

bilis H. Adria, Triest,” is a specimen of a species totally

distinct from variabilis, which I have not yet been able

to identify with certainty. |

13. No. 417. Specimens received from Haeckel; locality

Bergen. My preparation shows quite typical spicules

of variabilis mixed with those of a Sycon. (See text-

fig. 96, figg. 21 a—21 p, p. 381.)

3. LEUCOSOLENIA BOTRYOIDES.

Spongia botryoides Ellis & Solander, 1786, Nat. Hist. Zoophytes,

p. 190, pl. lviii. figg. 1-4.

Spongia botryoides Montagu, 1812, Wernerian Mem. ii. p. 89.

Spongia botryoides pars Grant, 1826, Edinb. Phil. Journ. xiv.

p. 839; Edinb. New Phil. Journ. i. p. 169.

Spongia botryoides pars Grant, 1827, Edinb. New Phil. Journ.

Tok, foe ALES » «

Grantia botryoides pars Fleming, 1828, Hist. Brit. Animals,

. 525.

‘ Calcispongia botryoides de Blainville, 1836, Manuel d’Actino-

logie, p. 531.

Grantia botryoides pars Johnston, 1842, Brit. Spong. and

Lithophytes, p. 178.

Leucosolenia botryoides Bowerbank 1864, Mon. Brit. Spong. 1.

p. 164, pl. xxvi. figg. 347, 348; (1866) ii. p. 28.

Grantia lieberkiihnii Schmidt, 1866, Spong. Adriat. Meer.

Suppl. i. pp. 8, 20.

Leucosolenia botryoides Gray, 1867, Proc. Zool. Soc. 1867,

. 550.

2 Leucosolenia botryoides Schmidt, 1868, Spong. Adriat. Meer.

Suppl. 11. p. 31.

Olynthium nitidum Haeckel, 1870, Jena. Zeitschr. v. p. 237.

Olynthium splendidum Haeckel, |. c. p. 237.

Leucosolenia (Leuceria) botryoides Haeckel, 1870, 1. c. p. 243.

Leucosolenia granti Haeckel, 1870, 1. c. p. 243.

Ascaltis botryoides Haeckel, 1872, Kalkschwimme, 1. p. 65,

pl. 9. fig. 10, pl. 10. figg. 7 a—7e.

Ascandra botrys Haeckel, 1872, 1. c. p. 101, pl. 16. figg. la-lf.

Ascundra nitida Haeckel, 1872, 1. c. p. 103, pl. 16. figg. 2 a-2 q,

Tle Tid paiasy Sa 774 WOE Sy

1904. | SPONGES OF THE GENUS LEUCOSOLENIA. 387

Ascandra botryoides Fristedt, 1885, K. Vetensk.-Akad. Hand-

lingar, xxi. no. 6, p. 9.

Ascaltis botryoides Hanitsch, 1890, Trans. Liverpool Biol. Soc.

iv. p. 223.

Leucosolenia botryoides Minchin, 1896, Ann. Mag. Nat. Hist.

(6) xviii. p. 359.

Leucosolenia botryoides Breitfuss, 1898, Arch. f. Naturges. lxiil.

re Oy ZAO

Ascandra botrys Breitfuss, 1898, 1. c. p. 213.

Leucosolenia botryoides Rousseau, 1903, Mém. Soc. Malac.

Belgique, xxxvu. p. 9, fig. 1.

Leucosolenia botryoides Allen, 1904, J. Mar. Biol. Assoc. n. s.

vii. p. 185.

(a) Haternal Characters.

This sponge is only known to occur in one kind of situation and

under one form. It is always found growing over alg, forming

a basal reticulum of finer tubes attaching it to its support, from

which arises a dense cluster of short smooth oscular tubes. This

extremely characteristic form is well seen in the figures of Ellis

and Solander [7], Bowerbank [1, vol. 1. fig. 348], and even

Haeckel [3, pl. ix. fig. 10], and enables this species to be distin-

guished at once. The form of the sponge is usually compared to

that of a bunch of grapes, hence the specific name botryoides ; but

since the oscular tubes are at least five or six times as long as

they are broad, a. bunch of bananas would be a ‘NeMaee comparison

as regards form.

if have not found botry yoides at Roscoff, but at Plymouth it is

not uncommon. In Wembury Bay I found it extremely abun-

dant, competing with Grantia compressa for the occupation of the

algee on the overhanging sides of rocks, in situations where the

sponges are left suspended high and dry at lowtide. Its mode

of growth may be compared with the busny form of variabilis

growing in similar situations. In fact, the only difference between

the two sponges under these circumstances is that in botryoides

the oscular tubes are shorter in proportion to their length and

more thickly clustered together, so that the distinctiveness of the

external form of botryoides is more apparent than real as regards

mode of growth. The oscular rim in this species is short or

of moderate length, and the oscular tube narrows more or less

rapidly towards the opening, but in both these respects also the

difference from variabilis is rather one of degree. The wall of the

oscular tube is greatly thickened by quantities of stout spicules,

and the sponge is much tougher and SHOE EB: than either of the

preceding species.

(b) Characters of the Spiculation.

Since the spiculation of this species is modelled, so to speak,

25%

[ Dec. 13,

F. E. A. MINCHIN ON THE BRITIS

388

Text-fig. 97.

j !

\ Dray

1904.] SPONGES OF THE GENUS LEUCOSOLENIA. 389

almost exactly upon that of Z. variabilis, and every class of

spicule found in the one is found also in the other, it is only

necessary to enumerate them in the same order as has been done

for L. variabilis, and to give at the same time an account of their

differences in the two species, and of their variations in the species

under consideration.

(a) Triradiate and Quadriradiate Systems.—(1) The ordinary

triradiates (text-fig. 97, figg. 22 a—-22.d, p.388) have exactly the same

characters as in variadilis, the unpaired ray short, the unpaired

angle obtuse. They vary from slender spicules, the rays about

3 p in thickness, to thick ones with the rays reaching about 10 pu.

There is often a tendency noticeable for the unpaired ray to be

irregularly curved or bent. This is to be ascribed to the great

number of spicules, closely packed in the body-wall of this sponge,

in consequence of which tbe growing spicule-rays encounter

obstacles to their growth which cause them to be deflected from

their normal straightness. (2) The thickened T-shaped triradiates,

comparatively few and far between in variabilis, are exceedingly

abundant in botryoides, and constitute the most marked feature

of the species (text-fig. 97, figg. 22 f-22 7, p.388). They are thick,

generally irregularly curved, with the paired rays usually much

longer than the unpaired one, and when seen from the gastval

aspect, if the spicule be lying so that one of the two paired rays

and the unpaired ray are in focus at the same time, the other

paired ray appears greatly foreshortened, giving the spicule an

asymmetrical appearance (text-fig. 97, figg. 22h, 227, p. 388).

(3) The brackets are more abundant than in variabilis and show

more tendency to modification and reduction of the unpaired ray

(text-fig. 97, figg. 221, 22 m, p. 388), so that while in variabilis

they might be overlooked, in botryoides they are easily found and

recognised. (4) The ordinary quadriradiates are similar to those

of variabilis (fig. 22e). (5) The T-shaped quadriradiates are

fairly abundant (figg. 227, 22%) and, as in variabilis, the gastral

ray is very upright or even inclined backwards in its basal two-

thirds. (6) The brackets may develop a gastral ray, which, as

described for variabilis, is usually straight and may reach a great

length (figg. 22 m, 220). In some specimens this type of spicule

is very abundant.

Explanation of Text-fig. 97 (opposite).

Spicules of Leucosolenia botryoides.

Figg. 22 a-22 0. Spicules of a specimen from Wembury Bay, Plymouth. a-d, or-

dinary triradiates; e, a quadriradiate; fi, thickened J-shaped triradiates ;

j» &, V-shaped quadriradiates; J, m, brackets; n, 0, brackets with gastral rays;

p-s, ordinary curved monaxons ; ¢, uw, straight slender monaxons.—Figg. 23 a—

23 p. Spicules of a specimen from Berwick Bay in Norm2n’s Collection, identified

by Haeckel as Ascaltis botryoides. a, 6, ordinary triradiates; c, d, ordinary

quadriradiates ; e-h, T-shaped triradiates; i, a T-shaped quadriradiate; j—n,

ordinary monaxons; 0; p, straight slender monaxons.

390 PROF. E. A. MINCHIN ON THE BRITISH [ Dec. 13,

Text-fig. 98.

= ~265 2672 26% 267 2

1904. | SPONGES OF THE GENUS LEUCOSOLENIA. 391

(b) Monaxon Spicules—(1) The ordinary curved, barbed forms

(text-fig. 97, fige. 22 p-22.s, p. 388) have the same characters as in

variabilis ; but they are usually short (60 u.-80 » in length), and

though they may vary in size, they never reach quite the length

which they do in variabilis; I have found none exceeding

300 in length, and such dimensions are quite exceptional

(text-fig. 98, fig. 26 g, p. 390). Further, they have a tendency to

be less curved and slightly thicker, in proportion to their length,

than in variabilis, and not to dwindle so much in thickness

just below the lance-head as do the monaxons of the latter

species; the features described are most marked in specimens

in which the monaxons are for the most part short (fig. 22). The

monaxons are always abundant and never absent, not even in the

specimens identified by Haeckel as Ascaltis botryoides (fig. 23), in

which, therefore, they are supposed to be wanting. (2) The

slender, straight, refringent monaxons are always present and

have the same characters as in L. variabilis (figg. 22¢, 22 x).

(3) The bayonet-shaped monaxons are not to be found in most

specimens, but in some they are fairly common (figg. 25 7, 257),

and show the same features as in LZ. variabilis.

Since this sponge was first named by Hllis and Solander in

1786, it was for many years the species to which all British Leuco-

solenias were referred, both in works dealing with sponges and

in the labels of museums and collections, until Haeckel in 1872

first pointed out the distinctive characters of the species and gave

a diagnosis of it. Haeckel, however, as has been pointed out,

wrongly separated this species into two, one with, the other with-

out monaxons, so that the characters which he ascribed to his

unnecessary species boirys are those which should be given to

botryoides. It is incomprehensible to me how Haeckel came to

Explanation of Text-fig. 98 (opposite).

Spicules of Leucosolenia botryoides.

Figg. 244-247. Spicules of a specimen from Portrush in Norman’s Collection

identified as Ascandra botrys by Haeckel. a, ordinary triradiate ; 6, ordinary

quadriradiate ; c, d, T-shaped triradiates; e-h, curved monaxons; 7, straight

slender monaxon.— Figg. 25 a-257. Spicules of a specimen from Heligoland in

the Berlin Museum (No. 1777), showing the bayonet-like monaxons. 4a, tri-

radiate, rather unusually slender; 6, T-shaped triradiate; c, d, brackets ;

e, straight slender monaxon; f-, curved monaxons, reaching a considerable

length ; 2,7, bayonet-like monaxons.—Figg. 264-261. Spicules of a specimen

from Liverpool in the Berlin Museum (No. 1763), showing the monaxons of

great length. a, 6, ordinary triradiates; c, quadriradiate ; d, T-shaped triradiate ;

e, f, brackets; g-k, ordinary curved monaxons; 7, bracket with gastral ray.—

Figg. 27 a-27 g. Spicules of a specimen in the British Museum (Bowerbank Coll.

992), figured im Bowerbank, Brit. Spong. i. fig. 848 (see p. 164). a, ordinary

triradiate; 6, ordinary quadriradiate; c, T-shaped triradiate; d, e, ordinary

nonaxons ; f, g, vefringent monaxons.

392 PROF. E. A, MINCHIN ON TNE BRITISH [Dec. 13,

make such a blunder*, since not only have I found monaxons well

developed and abundant in his type specimen, No. 1 of my list

below, but it is evident he himself saw them, since he founded

one of his usual “ connexive” varieties, Ascandra botryoides.

Haeckel further distinguished two varieties, ellisti and solanderit,

the former with the “apical ray of the quadriradiates slightly

curved, the lateral rays 7 or 8 times as long as thick, the unpaired

angle 130°-150°” ; the latter with the “apical ray of the quadri-

radiates straight, the lateral rays 5 or 6 times as long as thick,

the unpaired angle 150°-180°.” Since all these variations can be

found in any specimen, Haeckel’s two varieties may be struck out

of the systematic list. As I have also pointed out above, there is

nothing in Haeckel’s description of Ascandra nitida to separate

it from botryoides; A. nitida is, in fact, distinguished from

A. botrys by the same characters practically as Ascaltis botryoides

var. solanderia from var. ellisir.

Of writers subsequent to Haeckel, Fristedt alone [9] seems to

have penetrated Haeckel’s mistake, since he calls the species

Ascandra botryoides, as Haeckel ought to have done. Fristedt

has, moreover, gone a step further, and has put variabilis as a

synonym of botryoides, in which he has been followed by Vasseur

[23]. I have been sorely tempted to follow Fristedt’s lead also,

and to place both forms as well-marked varieties of one species,

for which, of course, the name botryoides would have to be main-

tained ; the form ordinarily known as botryoides could then be

called botryoides var. typica, and the other botryoides var. varia-

bilis. As has been shown above, the difference between the two

forms is purely one of degree in every respect. As regards spicu-

lation, they are in complete agreement, every form of spicule

occurring in the one being represented also in the other, and the

special features of botryoides are merely an exaggeration of those

of variabilis. As regards external form, variabilis occurs in a

variety of situations and consequently varies in form; the fact

that in botryoides the habits both of situation and growth are

constant, is in itself highly suggestive of its being a form-variety

adapted to a particular environment. Hxamination of the sponge

brings to light a further very important fact, bearing directly

upon the question under discussion, namely, that the thickened

triradiate systems so highly characteristic of botryoides are nearly

absent, or comparatively scarce, upon the basal network of tubes

by which the sponge is attached to its support, but are greatly

developed as a protecting and supporting layer upon the erect

* The only source I can suggest for Haeckel’s error with regard to the monaxons

of botryoides is the fact that Bowerbank also failed to see monaxons in this species,

and considered their absence as one of the characters distinguishing it from his

Leucosolenia contorta. If my notion be correct, this is a curious case of successive

incarnations of an error, manifesting itself first in Bowerbank 1866, then in Haeckel

1872, and for the last time, let us hope, in Breitfuss 1898. <A still more remarkable

point in this history is the fact pointed out above, that the specimen from which

Bowerbank described the spiculation of botryoides was really a specimen of

uariabilés (Haeckel), while the speeimen of “contorta” from which he figured

mouaxons was in reality a specimen of complicata !

1904. | SPONGES OF THE GENUS LEUCOSOLENIA. 393

oscular tubes. The basal network thus completely resembles

variabilis in its spiculation. For these reasons I should certainly

place the two forms as varieties of one species were I now

describing them for the first time.

I refrain at present, however, from taking the step of uniting

the two forms, for the reason that they have long been kept

distinct, and that they are generally very easily recognised and

distinguished from each other. ‘To call them one species, definite

proof of relationship should be obtained. If, for instance, larve

of a typical specimen of boiryoides were cultivated and found to

give rise to variabilis as well as to botryoides, according to the

conditions of growth, no one could doubt any longer that the

alleged two species were one. The experimental test of their

relationship should not be difficult to obtain; but until it has

been obtained, the two forms may be provisionally kept distinct

under their old names, in spite of the fact that the diagnostic

features of botryoides are purely relative as compared with

variabilis.

(d) List of Specimens examined.

(a) From Canon Norman’s Collection.

1. Dried specimen received from Johnston and sent by him

to Haeckel for examination; figured by Haeckel, Die

Kalkschwamme, pl. 9. fig. 10. (Haeckel’s figure is,

however, very much reconstructed, and does not bear a

close resemblance to the original.) With a label in

Haeckel’s handwriting :—

“ Ascaltis botryoides Hkl.

“ Grantia botryoides Johnston.

“ Leucosolenia botryoides Bwhk.

“(vera !).

‘“‘ Berwick Bay, Johnston.”

Also with label by Norman :—

** Leucosolenia botryoides.

‘“¢ Berwick Bay, Feb. 22nd, 1851.

“ Dr. Johnston.”

This most precious specimen is therefore one of Haeckel’s types,

and it shows clearly that, in a manner analogous to the case of

L. complicata mentioned above, Haeckel overlooked the small but

very abundant monaxons (see text-fig. 97, figg. 23 a—23 p, p. 388) ;

that therefore the species should have been placed as Ascandra

botryoides in his system; and that consequently the species A scandra

botrys H.. should be struck out of the system, becoming a synonym

of L. botryoides.

2. Dried specimen received from Dr. KE. Perceval Wright, by

whom it was sent to Haeckel for identification. With

label in Wright’s handwriting :—

“ Ascandra botrys, Portrush, BE. P. W.

“Specimens named by Haeckel.”

(See text-fig. 98, fige. 24 4-247, p. 390.)

394 PROF. E. A. MINCHIN ON THE BRITISH [ Dec. 13,

(6) From the British Museum.

3. Bowerbank Coll., 992. Dried specimen figured by Bower-

bank, Brit. Spong. i. fig. 348; described p. 164. (See

text-fig. 98, fige. 27 a-27 g, p. 390.)

4. Bowerbank Coll., 980. Dried specimen, with label in

Bowerbank’s handwriting—‘ Leucosolenia botryoides,

Gulot Caves, Sark, J.S. B.”

Bowerbank Coll., 981. Label in Bowerbank’s handwriting

— Leucosolenia botryoides Bowk. Fowey. C. W. Peach.

Very fine. O.C.”

6. Bowerbank Coll., 981. Label in Bowerbank’s handwriting

—‘‘ Leucosolenia botryoides, Scarborough, Bean.”

7. Register No. 47.9.7.109. Label ‘“ Grantia botryoides.”

Locality Berwick Bay, Johnston Collection.

8. Register No. 47.9.7.107. Label as last: locality Bangor,

County Down, Ireland, Johnston Collection.

9. Register No. 87.6.2%.4. Label “ Caleareous Sponge,

Shetland, E. M. Nelson, Esq.”

[The specimen No. 82.3.6.36, labelled ‘“ Leweosolenta

botryoides B. Australia. ?sp.,” in Carter’s handwriting,

is a totally distinct species. |

10. No. 1763, “ Ascaltis botryoides,” Liverpool. A specimen

ot L. botryoides in which the monaxons attain an un-

usually large size, though the designation Ascaltis implies

their total absence!* (See text-fig. 98, figg. 26 a—26 J,

p- 390.)

ll. No. 1777. “ Ascandra botrys,” Heligoland. (See text-

fig. 98, fige. 25 @—257.)

BIBLIOGRAPHY.

1. Bowrerpank, J.S. A Monograph of the British Spongiade.

Ray Society, 3 vols.: vol. i. 1864; vol. ii, 1866; vol. 11.

1874.

2. Brerrruss, L. L. Kalkschwammfauna des weissen Meeres

und der Hismeerkiisten des europdischen Russlands. Mém.

Acad. Imp. Sci. St. Pétersbourg, (vili.) vi. 41 pp., 4 pls.

3. Breirruss, L. L. Catalog der Calcarea der zoologischen

Sammlung des kéniglichen Museums fiir Naturkunde zu

Berlin. Arch. f. Naturges. lx. 1. (1895) pp. 205-226.

4. Brerrruss, L. L. Die arktische Kalkschwammfauna. Arch.

f. Naturges. lxiv. i. (1898) pp. 297-316.

5. Brerrruss, L. L. Note sur la faune des calcaires de Océan

Arctique. Ann. Mus. Zool. Acad. Imp. Sci. St. Peéters-

bourg, 1898, pp. 12--27.

* Breitfuss in his Catalogue [8] puts this specimen down as “ Leuwcosolenia

botryoides,” and since the name Leucosolenia is used by him to denote Ascons in

which monaxon spicules are lacking, it follows, either that he did not examine the

specimen, or that he overlooked the very conspicuous monaxons.

1904. ] SPONGES OF THE GENUS LEUCOSOLENIA. 395

6. pe Buarnvittz, H.M.D. Manuel d’Actinologie ou de Zoo-

22.

23.

24.

phytologie. Paris, 1834, 2 vols.

. Euuis, J., & Souanper, D. The Natural History of many

Curious and Uncommon Zoophytes. London, 1786.

. Fuemine, J. A History of British Animals. Edinburgh

and London, 1828.

. Frisrept, K. Bidrag till Kannedomen om de vid sweriges

vestra Kust lefvande Spongie. K. Svensk. Vet.-Ak. Handl.

xxl. 1885, no. 6.

. Grant, R. E. Remarks on the Structure of some Calcareous

Sponges. Edinb. New Phil. Journ. i. 1826, pp. 166-170.

. Grant, R. EH. Lectures on Comparative Anatomy, IV.

Lancet, i. 1833, pp. 193-200.

. Harcxen, KE. Prodromus eines Systems der Kalkschwimme.

Jen. Zeitschr. v. 1870, pp. 236-254; (translation) Ann.

Mag. Nat. Hist. (4) v. (1870) pp. 176-191.

. Harcker, H. Die Kalkschwimme, eine Monographie.

Berlin, 1872, 3 vols.

. Jounston, G. A History of British Sponges and Lithophytes.

1842.

. Minourn, E. A. Suggestions for a Natural Classification of

the Asconide. Ann. Mag. Nat. Hist. (6) xviii. pp. 349-362

(1896).

. Mincuin, H. A. Sponges, in: Lankester, ‘A Treatise on

Zoology,’ part u. A. & C. Black, London, 1900.

. Montacu, G. An Essay on Sponges, with Descriptions of all

the Species that have been discovered on the Coast of Great

Britain. Wernerian Memoirs, 11. 1812, pp. 67-122, pls. 11.—

XV1.

. PonésarFF, N. Report on the Calcarea dredged by H.M.S.

‘Challenger’ during the years 1873-1876. Chall. Rep. Zool.

vill. 2 (1883).

. Scumipt,O. DieSpongien des adriatischen Meeres. Leipzig,

1862.

. Scomipt, O. Grundziige einer Spongien-Fauna des atlant-

ischen Gebietes. Leipzig, 1870.

. ScuurFnerR, O. Beschreibungen eimiger neuer Kalk-

schwimme. Jen. Zeitschr. xi. (n. F. iv.) 1877, pp. 403-433,

pls. xxiv.—xxvi.

Trempteton, R. A Catalogue of the Rayed Animals found in

ireland. Magazine of Nat. Hist. ix. 1836, pp. 466-472,

fige. 66, 67.

Vasseur, G. Réproduction asexuelle de la Leucosolenia

botryoides (Ascandra variabilis Haeckel). Arch. Zool. Exp.

vill. 1879, pp. 59-66.

Wettner, W. Beitriige zur Fauna der sudéstlichen und

dstlichen Nordsee. I. Spongien. Wissenschaftliche Meeres-

untersuchungen, n. F. 1. 1894, pp. 325-328.

| Addendum.—Since the foregoing memoir was written and

396 MR. MARTIN JACOBY ON NEW SPECIES OF [ Dec. 13,

read before the Zoological Society, a paper has appeared on the

‘Plymouth Marine Invertebrate Fauna” in the Journal of the

Marine Biological Association (n.s. vii. 2, Dec. 1904), in which

notes furnished by myself and Mr. Bidder are mixed up together

with rather contradictory results (p. 185). Thus Leucosolenia

botryoides is stated to occur ‘on the shore between tide-marks,

not abundant except in certain localities (#. A. W/.); rocks under

the Hoe, in abundance (6. P. B.).” LZ. variabilis is put down as

‘commons everywhere in rock-pools between tide-marks (/.

A.M.).” The fact is that Bidder has not recognised variabilis

and botryoides as distinct, and, taking the characters of botryoides

from Bowerbank (whose figured specimen of this species was

really variabilis), Bidder has not unnaturally identified sponges

as botryoides which should have been named variabilis. Bidder’s

note on the occurrence of botryoides should therefore be trans-

ferred to variabilis, and then it will be perfectly correct. The true

botryoides is certainly very far from abundant on rocks under the

Hoe; I doubt if it ever occurs there. ]

2. Descriptions of Thirty-two new Species of Halticine

(Phytophagous Coleoptera) from South and Central

America. By Marvin Jacosy, F.E.S.

[Received November 25, 1904. |

Inst of the Species*.

Lactica nicotine. Mexico. Acanthonycha costatipennis. Brazil.

» decorata. Peru. ss antennata. Brazil.

» maculicollis. Peru. Sophraenella fulva. Amazons.

» posticata. Peru. Blepharida flavocostata. Mexico.

5 discicollis. Peru. is multimaculata. Mexico.

, rufobrunnea. Peru. Prasona peruviana. Peru.

> baer. Peru. Systena melanocephala. Peru.

» argentinensis. Argentine. » argentinensis. Argentine.

Agasicles vittata. Peru. » antennata. Amazons.

Disonycha amazonica. Amazons. Pseudogona discoidalis. Argentine.

3 peruana. Peru. Ay militaris, Panama.

ws albicincta. Peru. 9 pallida. Costa Rica.

Acanthonycha peruana. Peru. Oxygona amazonica. Amazons.

Af geniculata. Peru. 5 capitata. Peru.

Bs dimidiata. Peru. Crepidodera longicornis. Peru.

us stali. Costa Rica. | Hippuriphila catharine. Brazil.

LAcTICA NICOTINA, sp. Nn.

Flavous or fulvous, antenne (the basal three joints excepted)

black, head and thorax impunctate; elytra dark metallic blue or

violaceous, finely punctured anteriorly, the posterior portion very

obsoletely so.

Length 3-33 millim.

Head impunctate, the frontal tubercles very distinct, carina

short, not widened anteriorly, palpi robust ; antennz extending to

% The types of all the species described here are contained in the author’s collection.

1904. ] AMERICAN PHYTOPHAGOUS COLEOPTERA. 397

about the middle of the elytra, black, the lower three joints and

the base of the fourth flavous, third and fourth joints equal,

slightly longer than the second; thorax twice as broad as long,

the sides nearly straight, with a narrow margin, the anterior

angles obliquely thickened, the basal suleus deep, straight, and

bounded at the sides by equally deep perpendicular grooves, the

surface impunctate, flavous or fulvous; scutellum fulvous; elytra

without basal depression, the base distinctly, closely and somewhat

regularly punctured, the punctuation gradually diminishing in

size towards the apex; underside and legs fulvous, the tarsi

sometimes piceous.

Hab. Mexico (found in tobacco), Jalapa.

This species was not known to me when I described the

Central American Phytophaga for the Biolog. Centr.-Americana.

It must be placed near LZ. dives Har. and ZL. clara Har., and is

also closely allied to several other Mexican species of Lactica; but

it differs from all in one or more details, notably in the distinct

frontal tubercles, colour of the antenne, and of the under side,

and the fine elytral punctuation, &e. There are five specimens

before me.

LACTICA DECORATA, Sp. n.

Fulvous; antennz, the underside and legs more or less black ;

thorax impunctate, with very deep sulcus; elytra minutely punc-

tured, a transverse band at the base and a large spot near the

_ apex metallic blue or greenish.

Var. a. Hlytra metallic blue, the lateral margins narrowly and

the apical one broadly fulvous.

Var. 6. Antenne, under side, and legs entirely fulvous.

Length 7-74 millim.

Head with a single puncture near each eye, the rest of the

surface impunctate, frontal elevations absent, clypeus convex

between the antenne, labrum black; antenne more or less piceous,

sometimes nearly fulvous, robust, extending beyond the middle of

the elytra, the second and third joints short, nearly equal, following

joints elongate; thorax scarcely twice as broad as long, the sides

straight, obliquely narrowed anteriorly, almost concave, the anterior

angles strongly oblique, the basal sulcus slightly sinuate, very deep

at the sides, less so at the middle, the surface entirely impunc-

tate, pale fulvous; scutellum fulvous or piceous; elytra with an

obsolete depression below the base, extremely finely and remotely

punctured, fulvous, a transverse band at the base, not extending

to the lateral margins and downwards to about the third portion

of the elytra, and a transversely shaped shorter band or spot below

the middle, metallic greenish or bluish; underside and legs black,

abdomen more or less fulvous.

Hab. Peru.

A handsome species, somewhat allied in coloration in regard

to the variety to ZL. marginata Clark, but the elytra of metallic

coloration and the underside black. I have looked upon the

398 MR. MARTIN JACOBY ON NEW SPECIES OF [ Dec. 13,

banded form as the typical one, and that in which the bands

occupy the entire disc of the elytra as the variety. I cannot

distinguish a specimen in any way from the others, having the

antenne, underside, and legs fulvous.

LACTICA MACULICOLLIS, sp. n.

Testaceous, the head and a spot at the anterior part of the

thorax rufous; elytra elongate, scarcely perceptibly punctured,

testaceous, a transverse band at the base rufous.

Length 4 millim.

Head entirely impunctate, rufous, frontal elevations absent,

carina elongate, strongly convex, eyes very large; antenne fulvous,

long and slender, the third joint slightly longer and thinner

than the second; thorax about one half broader than long, the

sides straight, the basal sulcus broad and rather deep, the surface

testaceous, impunctate, with a rufous spot at the middle of the

anterior margin, anterior angles obtuse, thickened; scutellum

testaceous; elytra not perceptibly punctured, even under a strong

lens, elongate and parallel, testaceous, with a transverse rufous

band at the base extending to the lateral margins, this band

scarcely occupies a third portion of their length ; underside and

legs pale testaceous.

Hab. Peru.

Of less than half the size of ZL. rufobasalis Jac. and distinguished

by the coloration of the head and thorax.

LACTICA POSTICATA, Sp. n.

Head, the basal joints of the antennz, the thorax, and the

anterior femora flavous; elytra dark violaveous, finely punctured

anteriorly ; the posterior legs blackish.

Length 3 millim.

Head impunctate, the frontal elevations feebly raised ; carina

short, not widened in front ; antenne long and slender, black, the

lower three and the base of the fourth joint flavous, third and

fourth joints equal; thorax about twice as broad as long, the sides

nearly straight, the anterior angles oblique, the basal sulcus

straight and deep, bounded laterally by very deep perpendicular

grooves, the surface impunctate, flavous; scutellum flavous; elytra

without a trace of a basal depression, violaceous, with indistinct

rows of fine punctures at the anterior portion, extremely sparingly

and finely punctured near the apex; the breast and the anterior

femora flavous; the abdomen, the posterior femora, and the tibie

and tarsi piceous.

Hab. Peru.

Closely allied to Z. elegantula Har. (Coleopt. Hefte, xiv. p. 16) ;

but the frontal elevations of the head feeble, the latter without

punctures, the third and fourth joints of the antennz of equal

length, the elytra without any basal depression and some fine

punctures posteriorly, and the legs differently coloured. L. boli-

viana Jac. is of double the size and strongly punctured.

1904. | AMERICAN PHYTOPHAGOUS COLEOPTERA. 399

LACTICA DISCICOLLIS, sp. n.

Flavous, the head and the disc of the thorax piceous, the latter

impunctate ; elytra flavous, extremely minutely punctured.

Length 3 millim.

Head impunctate, nearly black ; eyes very large, the diameter of

each exceeding the intermediate space, frontal elevations obsolete ;

antenne flavous, the second to the fifth joints gradually elongate ;

thorax one half broader than long, the sides straight, the anterior

angles oblique, the basal sulcus and the perpendicular grooves very

deep, the surface impunctate, piceous, all the margins narrowly

flavous; scutellum piceous; elytra narrow and parallel, fulvous,

scarcely perceptibly depressed below the base, very finely punc-

tured and somewhat regularly arranged in rows, more irregularly

and finely so near the apex; underside pale flavous, legs flavous.

Hab. Peru.

A small species, principally distinguished by the colour of the

thorax. There are six specimens before me.

LAcTICA RUFOBRUNNEA, Sp. n.

Elongate and parallel, reddish-fulvous; the antenne, the apex

of the posterior femora, and the tibie and tarsi black ; thorax

impunctate, deeply sulcate ; elytra extremely minutely punctured.

Length 53 millim.

Head with some deep punctures at the vertex, the frontal

tubercles entirely obsolete, carina linear, short; antenne ex-

tending to the middle of the elytra, black, the third and fourth

joints equal, terminal joints slightly shorter ; thorax scarcely one-

half broader than long, distinctly narrowed anteriorly, the sides

straight, the anterior angles oblique, the surface impunctate, with a

broad transverse sulcus and deep perpendicular grooves; scutellum

rather small; elytra elongate, without basal depression, extremely

minutely and closely punctured; legs black; the base of the

posterior femora and the underside fulvous.

Hab. Marcapata, Peru.

Allied to Z. clypeata Baly; but smaller and shorter, of dark

reddish colour, not testaceous, the head not punctured in front,

the encarpe obsolete, the antenne entirely black, the thorax much

less transverse with the sides straight, the anterior angles not

dentiform.

LACTICA BAERI, sp. n.

Rufous ; antenne (the basal joint excepted) and the tibiz and

tarsi black, head and thorax impunctate; elytra very finely and

rather closely punctured, metallic blue.

Length 5 millim.

Head impunctate, rufous, the frontal elevations small and

transverse, carina elongate and narrow; eyes widely separated,

moderately large; antenne extending beyond the middle of the

elytra, black, the basal joint and the base of the following two

joints fulvous, the third joint distinctly shorter than the fourth,

400 MR. MARTIN JACOBY ON NEW SPECIES OF [ Dec. 18,

the following elongate; thorax twice as broad as long, the sides

rounded with comparatively broadly reflexed margins, the basal

suleus rather deep, the surface impunctate, rufous; scutellum

fulvous; elytra nearly parallel, not depressed below the base,

the shoulders prominent, the surface finely and closely punctured,

the punctures distinct to the apex; underside and legs rufous,

the tibie at their outer edge and the tarsi black.

Hab. Prov. Huallaga, Rio Mixiollo, Peru (@. 4. Baer). Col-

lection M, Clavareau and my own.

This species of Lactica is allied to L. dives Har.; but differs from it

in the black tibie, the rufous coloration, and blue not violet elytra,

and from the other species of the genus in the same differences of

coloration and the impunctate head; in one of the specimens the

head has a distinct fovea at the middle of the vertex, which is

wanting in the other or is scarcely indicated.

LACTICA ARGENTINENSIS, Sp. 0.

Flavous, antenne and legs black; head and thorax impunctate ;

elytra dark violaceous, microscopically punctured.

Length 53 millim.

Of broad and nearly parallel shape, the head very broad and

convex, without scarcely an indication of tubercles or carina; the

clypeus broad, convex, scarcely separated from the face; labrum

and palpi black; antenne robust, black, the second and third

joints short, nearly equal, fourth and following joints about one half

longer ; thorax twice as broad as long, narrowed anteriorly, the

anterior angles oblique, the basal suleus deep as well as the per-

pendicular lateral grooves, the surface impunctate, pale flavous ;

scutellum triangular, flavous; elytra convex, without trace of a

basal depression, dark violaceous blue, remotely and extremely

minutely punctured ; underside flavous, legs black.

Hab. Tucuman, Argentine Rep.

This is a well distinguished species on account of the broad and

convex shape of the head, which nearly forms an uninterrupted

smooth surface. I received two specimens of this insect from

Mr. C. Bruch, of the La Plata Museum.

AGASICLES, gen. nov.

Body elongate and glabrous ; antenne filiform, the third and

following joints elongate, terminal ones shorter; thorax as long as

broad, the sides straight, the surface without sulcus ; elytra wider

at the base than the thorax, obsoletely punctured, their epipleure

broad ; legs stout, the posterior femora strongly incrassate, their

tibia: mucronate, the other tibie unarmed, the first joint of the

posterior tarsi as long as the following joints together, claws

appendiculate ; prosternum narrow; mesosternum elongate, rather

broad, deeply triangularly emarginate at the base ; anterior coxal

cavities open; pygidium not covered by the elytra, convex, broadly

rounded.

1904. ] AMERICAN PHYTOPHAGOUS COLEOPTERA. 401

Amongst the genera of Halticine with open coxal cavities and

a non-suleate thorax, I know of none in which the latter part is of

similar shape, that is as long as broad; this character and the

general elongate shape, which resembles somewhat a Donacia,

will help to distinguish the genus.

AGASICLES VITTATA, Sp. n.

Black, the lower three joints of the antenne fulvous, thorax

with an eneous gloss; elytra nearly impunctate, flavous, the sutural

and lateral margins as well as the apex and a longitudinal stripe

at the disc, abbreviated at each end, black.

Length 6 millim.

Head deeply and irregularly punctured, «neous, the frontal

elevations feebly marked, subquadrate; clypeus broad, raised at

the middle, triangular; palpi fulvous, the terminal joint small,

acutely pointed ; antenne extending to the middle of the elytra,

black, the lower three joints fulvous, third joint shorter than the

fourth, both thickened at the apex, the following joints shorter ;

thorax with straight sides, the angles obtuse, the surface flat,

dark neous, extremely minutely granulate and punctured;

scutellum broadly rounded at the apex, coloured like the thorax ;

elytra narrow and parallel, extremely minutely punctured, a

sutural and lateral narrow stripe widened into a spot at the apex

and a longitudinal band at the middle, not extending to the base

or the apex, greenish-black, the vest of the surface flavous; under

side and legs black ; last abdominal segment deeply longitudinally

sulcate, its apex strongly emarginate.

Hab. Prov. Huallaga, Rio Mixiollo, Peru.

Of this species 2 single specimen is contained in my collection

which I received from M. Clavareau at Brussels, another is in that

gentleman’s posession. The specimen before me is probably a male,

judging from the structure of the last abdominal segment and the

prominent pygidium.

DISONYCHA AMAZONICA, Sp. nD.

Head, the breast, and the legs pale piceous; thorax testaceous,

impunctate ; elytra minutely punctured, black, the lateral margins

and a narrow transverse band at the middle flavous.

Length 8 millim.

Head impunctate, the vertex piceous, the lower portion tes-

taceous; eyes large, oblong, with a single deep puncture near

their inner margins, frontal tubercles obsolete; clypeus strongly

convex; labrum black; antenne obscure fulvous, the basal joimt

flavous, third joint shorter than the fourth ; thorax twice as broad

as long, the sides feebly rounded, narrowly margined, the anterior

angles obliquely rounded, the posterior ones acute, oblique, the

surface impunctate, shining, obsoletely transversely sulcate near

the base ; scutellum black ; elytra extremely finely punctured,

black, this colour interrupted at the middle by a narrow transverse

Proc. Zoou. Soc.—1904, Vou. II. No. XX VI. 26

402 MR. MARTIN JACOBY ON NEW SPECIES OF [ Dec. 13,

flavous band, and at the sides by the flavous lateral margins ;

abdomen testaceous, breast and legs pale piceous,

Hab, Amazons.

The coloration of this species entirely resembles that of many

members of the genera Oedionychis and Asphera; but the shape

of the thorax and the non-inflated claws, as well as the elongate

metatarsus of the posterior legs, prove the insect to belong to:

Disonycha.

DISONYCHA PERUANA, Sp. n.

Fulvous, the antenne, tibie, and tarsi black, testaceous above ;.

the head with one, the thorax with five black spots; elytra

impunctate, a narrow sutural and discoidal stripe and the lateral

margins black.

Length 7 millim.

Head impunctate, testaceous with a central black spot, frontal

elevations feebly raised; antenne black, not extending to the middle

of the elytra, the basal two joints fulvous below, third joint slightly

shorter than the fourth ; thorax one-half broader than long, the

sides nearly straight, the anterior angles thickened, the posterior

ones strongly oblique, the surface impunctate, testaceous, with

five obsolete piceous spots, four anteriorly and one elongate at the

middle; scutellum black; elytra impunctate, flavous, narrowly

edged with black, and a similar narrow stripe at the middle which

does not quite extend to the apex, elytral epipleurz testaceous.

within ; under side testaceous, femora darker, tibie and tarsi

black.

Hab. Peru.

A proportionately large-sized species, and differing from most of

its congeners in having the lateral black elytral stripes not placed

close to but at the actual margins; the narrow shape of the stripes

and the entirely impunctate elytra will assist further in the

determination of the species.

DisonycHA ALBICINCTA, Sp. Nn.

Black; thorax pale flavous, impunctate ; elytra bluish black, a

very narrow transverse band before the middle, another near the

apex, connected at the sides by a longitudinal stripe, white.

Length 6 millim.

Head with a few deep punctures near the eyes, black, very

shining, frontal elevations absent, carina very narrow ; antenne

black, the basal joint flavous below, fourth joint much longer

than the third, apical joint extending beyond the middle of the

elytra; thorax scarcely twice as broad as long, the sides straight,

narrowly marginate, the anterior angles thickened, obtusely

rounded ; the disc impunctate, with a very obsolete transverse

groove near the base, pale yellow, scutellum black ; elytra micro-

scopically punctured, elongate and parallel, bluish black, metallic,

with two very narrow transverse bands connected at the lateral

margins by an equally narrow longitudinal stripe, the first band

1904. | AMERICAN PHYTOPHAGOUS COLEOPTERA. 403

immediately before the middle, the second near the apex ; under-

side and legs black.

Hab. Peru and Bolivia.

A very distinct species, well distinguished by the narrow white

elytral bands connected at the sides. Seven specimens are con-

tained in my collection.

ACANTHONYCHA PERUANA, Sp. Nn.

Elongate and parallel, reddish fulvous, antennz and legs black,

posterior femora fulvous at the base; thorax impunctate; elytra

metallic green, finely punctured and wrinkled.

Length 5 millim.

Head impunctate, fulvous, shining, the frontal tubercles

strongly developed, rather broad ; antenne very slender, extending

beyond the middle of the elytra, fulvous, the second joint piceous,

third one-half shorter than the fourth joint; thorax not broader

than long, the sides nearly straight and forming a slight angle

before the middle, anterior angles slightly produced outwards, the

dise very obsoletely transversely sulcate near the base, reddish

fulvous, very shining and impunctate ; scutellum fulvous, broad ;

elytra wider at the base than the thorax, parallel, very narrowly

margined, metallic greenish, the surface very finely and closely

punctured and transversely wrinkled, the extreme lateral margins

and the epipleure fulvous, the latter very broad anteriorly ; below

and the legs pale fulvous, the anterior tibiz rather darker.

Hab. Peru.

Acanthonycha was established by me in the Biolog. Centr.-

Amer. for a species placed by Baly in Pelonia (P. elegantula),

from which it differs very materially in the structure of the thorax

and other details; the species described here differs from that of

Baly by the sculpturing of the elytra, which are finely wrinkled

instead of smooth.

ACANTHONYCHA GENICULATA, Sp. n.

Reddish fulvous, the antenne and legs (the base of the posterior

femora excepted) black, thorax impunctate; elytra dark blue,

finely rugose and punctured.

Length 5 millim.

This species agrees in most structural details with the preceding

one, of which it may possibly be a variety only; but the antenne

are more robust, the joints less elongate and entirely black; this

is also the case with the legs, excepting the basal greater portion

of the posterior femora which is fulvous, the colour of the elytra

also is dark blue instead of green. There are two exactly similar

specimens before me.

Hab. Peru.

ACANTHONYCHA DIMIDIATA, Sp. 1.

Fulvous; the antenne, tibie and tarsi, the posterior legs and

the abdomen black; thorax with a deep transverse sulcus; elytra

26*

404 MR. MARTIN JACOBY ON NEW SPECIES OF [ Dec, 13,

minutely punctured, the posterior portion metallic blue, the

anterior one fulvous.

Length 7 millim.

Head produced, impunctate, fulvous, the frontal elevations

strongly vaised, tuberculiform ; clypeus thickened, flavous ;

antenne extending to two-thirds the length of the elytra, black,

fourth and following joints very elongate, the third shorter ;

thorax subquadrate, the sides distinctly angulate before the middle,

the anterior angles acute, the disc with a deep transverse sulcus

near the base and a depression anteriorly (accidental?) impunctate,

fulvous; elytra extremely minutely punctured, fulvous, with a

slight purplish gloss, the posterior portion, from immediately below

the middle, bright metallic blue; elytral epipleurze very broad,

fulvous anteriorly, blue at the posterior half; underside fulvous,

the abdomen and the posterior legs as well as the tibize and tarsi

black.

Hab. Peru.

Excepting in the deep thoracic sulcus, this species (of which I

possess a single, apparently female, specimen) agrees in all

structural details with the others, but is much larger and well

distinguished by its coloration; thoracic feeble sulci can also be

perceived in the other species of this genus.

ACANTHONYCHA STALI, Sp. 0.

Fulvous; the antennee (the basal joint excepted) and the tibie

and tarsi black ; thorax impunctate, shining; elytra very minutely

punctured, rather flattened.

Length 6 millim.

Head impunctate, shining; antennz extending to the middle

of the elytra, black, the basal joint fulvous, third joint much

shorter than the fourth; thorax subquadrate, of the same shape

and smooth as in the allied species, more transverse however,

and distinctly angulate before the middle in the female insect ;

elytra obsoletely depressed below the base and longitudinally

so within the shoulders, very finely and closely punctured, the

posterior femora rather strongly incrassate.

Hab. Tucurrique, Costa Rica.

Rather larger than the other species of the genus, the antenne

with fulvous basal joint, the fourth joint proportionately longer,

the elytra differently coloured, and the posterior femora more

strongly incrassate. I cannot discover any sexual characters in

the structure of the last abdominal segment, which seems the same

in all the specimens before me, but the shorter antennz and more

transversely shaped thorax, with its strongly angulate sides in one

of the specimens, are probably the female characters.

ACANTHONYCHA COSTATIPENNIS, Sp. 0.

Fulvous; the antennz (the basal joints excepted) and the tibize

and tarsi black; thorax subquadrate, impunctate ; elytra dark blue,

1904. | AMERICAN PHYTOPHAGOUS COLEOPTERA. 40d:

closely punctured, the sides with a longitudinal costa, preceded by

a sulcus (@ ).

Length 53 millim.

Head deeply inserted, the frontal elevations strongly raised,

clypeus with an acute carina between the antenne, the latter

black, the basal three joints fulvous, the third joint one-half

shorter than the fourth, the following joints slightly widened ;

thorax subquadrate, of the same shape as usual, fulvous, the base

with an obscure transverse depression, another one still less

distinct being placed near the anterior margin ; scutellum fulvous ;

elytra closely and finely punctured, with an acutely raised costa

from the shoulders to below the middle and preceded by a

longitudinal sulcus ; posterior femora but slightly thickened, tibize

and tarsi black.

Hab. Santa Catarina, Brazil (/ruhstorfer).

At once distinguished from the other species of the genus by

the cost of the elytra in connection with the close punctuation,

their epipleure are broad but the prosternum is very narrow. ‘The

two specimens before me are probably females, and in the male

the elytral costz may be absent.

ACANTHONYCHA ANTENNATA, Sp. 0.

Fulvous ; antennz robust, black, the basal three joints fulvous ;

thorax impunctate; elytra dark blue, very finely and closely

punctured, their epipleurze fulvous; tarsi piceous.

Length 4 muillim.

Of this species only a single male specimen is before me, which

may possibly be the male of A. costatipennis; but the antenne

differ so much from those of that imsect and of the other species

of the genus that I must look upon the species as distinct :

these organssare proportionately short and stout, the third joint

is one-half shorter than the fourth and the latter and the following

joints are triangularly elongate, but not filiform; the under side

is clothed with fine yellow pubescence, the legs are more robust

than usual and the tarsi rather widened. ‘The species can only be

compared to those in which the elytral punctuation is distinct, but

the short antenne will distinguish it from any of them.

Hab. Santa Catarina, Brazil.

SOPHRAENELLA, gen. nov.

Ovate, convex ; antennz short, gradually transversely widened,

the basal joint club-shaped, the second very short, third joint

triangularly elongate, the following joints transversely sub-

quadrate, terminal joint short, ovate; eyes reniform, deeply and

broadly emarginate; palpi with the penultimate joint strongly

incrassate ; thorax nearly three times broader than long, the sides

narrowly margined, the anterior angles oblique, the surface

without sulcus; scutellum broadly triangular; elytra convex,

broadly ovate, their epipleuree moderately broad at the base,

406 MR. MARTIN JACOBY ON NEW SPECIES OF | Dec. 13,

disappearing below the middle; legs short and robust, the posterior

femora strongly incrassate, their tibize with a short spur, broadly

channelled near the apex, tarsi short and broad, claws appendi-

culate; prosternum narrowly elongate, mesosternum subquadrate,

the anterior cotyloid cavities open.

I propose this genus for a robust-looking species of fulvous

coloration, which, in the short, transverse joints of the antenne,

resembles much the genus Sophraena Baly, but which may be at

once distinguished by the elongate and reniform shape of the eyes ;

there is also a difference in the proportionate length of the joints

of the antennz. In Wephrica Har. the eyes are likewise kidney-

shaped, but not to such an extent as in the present genus, and the

antenne are filiform. As to Sophraena, Baly did not mention

the state of the anterior cotyloid cavities when he described the

genus; Chapuis in his ‘Genera’ has placed the species near

Oxygona and states that the cavities are closed. As the opposite is

the case in this genus, the latter must find its place near

Rhopalotoma Clark, but the genus stands isolated in that group by

the shape of the antenne.

SOPHRAENELLA FULYVA, Sp. 0.

Fulvous; the antenne (the basal joint excepted) black; thorax

finely and closely punctured; elytra strongly convex, punctured

like the thorax.

Length 7 millim.

Head very flattened (like that of a Cryptocephalus), impunctate

with the exception of a single puncture near the eyes, the latter

occupying nearly the entire sides of the head; frontal elevations

absent; antenn only extending to the base of the elytra, black,

the basal joint entirely and the second one partly fulvous; thorax

nearly three times broader than long, the sides nearly straight,

narrowed anteriorly, the anterior angles oblique, the surface

closely and finely punctured, posterior margin rather rounded and

produced at the middle; scutellum broadly ovate; elytra not wider

at the base than the thorax, very closely and scarcely more

strongly punctured than the latter.

Hab. Amazons.

The resemblance of the head to that of a species of Crypto-

cephalus is very striking, if the antenne are not considered.

BLEPHARIDA FLAVOCOSTATA, Sp. n.

Flavous, the antenne, tibise, and tarsi black; thorax with five

black spots, finely punctured ; elytra closely and strongly punctured,

fuscous, each elytron with five narrow, flavous, longitudinal costee ;

sides of the breast and abdomen black.

Length 9 millim.

Head with the frontal elevations broad and strongly raised, the

carina linear ; antenne extending to the middle of the elytra,

black, the third and fourth joints equal, the terminal joints shorter ;

thorax transverse, about twice and a half broader than long, the

1904. | AMERICAN PHYTOPHAGOUS COLEOPTERA. 407

sides straight, the posterior angles oblique, the disc very finely

and irregularly punctured, flavous, with five black spots (2, 3),

those near the posterior angles larger and of transverse shape ;

scutellum broadly triangular, the base fuscous, clothed with fine

yellow hairs, the apex fulvous ; elytra widened towards the middle,

very broad, of fuscous colour, closely, strongly, and somewhat

rugosely punctured, with five flavous longitudinal costee on each

elytron, placed at equal distances, the first subsutural one short and

extending to the suture at its posterior end, the third and fourth

coste joined at the apex ; the femora and the rest of the under

side flavous, the sides of the breast and of the abdomen, the tibiee

and tarsi black, claws bifid.

Hab, Cuernavaca, Mexico (Dr. A. Fenyes). :

I only know two specimens of this species, one of which I

received from M. Clavareau at Brussels.

BLEPHARIDA MULTIMACULATA, Sp. Nn.

Fulvous; thorax very finely and sparingly punctured ; elytra

dark fulvous or piceous, the suture more or less fulvous, closely

longitudinally costate, the interstices strongly punctured, the costz

with numerous small whitish spots.

Length 4 millim.

Head impunctate, fulvous ; antenne extending slightly beyond

the middle of the elytra, fulvous; all the joints, with the excep-

tion of the second, elongate, the third slightly shorter than the

fourth joint; thorax of equal width, twice as broad as long, the

sides feebly rounded ; the surface covered with a few fine punctures,

irregularly distributed ; scutellum piceous; elytra dark chestnut-

coloured or piceous, strongly and closely punctured in rows, the

interstices longitudinally costate, to the number of about ten on

each elytron, with numerous small round yellowish-white spots,

irregularly placed and different on each elytron; legs flavous, the

sides of the breast blackish.

Hab. Atlixco, Tepetlapa, Mexico.

This species, of which I received several specimens some time

ago from M. Clavareau, was not known to me at the time of the

publication of the Biologia Centr.-Amer.; it resembles greatly in

coloration several Eastern species, and must not be confounded

with B. marmorata Jac., likewise from Mexico, as in that msect

the elytra are not costate, the colour is black, and the spots much

fewer in number.

PRASONA PERUVIANA, Sp. n.

Greenish-testaceous, antenne fulvous, head with a black spot ;

thorax impunctate, transversely suleate ; elytra closely punctured,

each elytron with two elongate spots at the base and a round one

at the apex, black.

Length 7 millim.

Head broad, impunctate, greenish, the vertex with a black spot ;

antennze about two-thirds the length of the body, pale fulvous,

- 408 MR. MARTIN JACOBY ON NEW SPECIES OF [ Dec. 13,

the third and fourth joints equal; thorax twice as broad as long,

the sides subangularly rounded before the middle, constricted near

the base, the anterior angles blunt, the posterior ones acute, the

surface with a rather deep, transverse, sinuate sulcus near the

base, extending to the sides, the disc impunctate ; elytra wider at

the base than the thorax, transversely depressed below the latter,

very closely and distinctly punctured, the punctures somewhat

regularly arranged in rows, the ground-colour of a pale greenish-

testaceous, the base with two elongate black spots, one near the

margin, the other near the scutellum and extending nearly to the

middle of the elytra, another round black spot is placed near the

apex ; underside and legs coloured like the upper surface.

Hab. Mar capata, Peru.

I possess five similarly marked specimens of this distinct s species.

SYSTENA MELANOCEPHALA, Sp. 0.

Flavous, the head, the terminal joints of the antenne, and

the breast black ; thorax impunctate, flavous ; elytra extremely

minutely punctured, flavous, the suture at the base, a humeral spot

and a transverse band near the apex black.

Length 8 millim.

Head impunctate, very shining, black, the frontal elevations

and the clypeus flavous; antennz extending to the middle of the

elytra, flavous, the apical four joints black, third joint one-half

shorter than the fourth ; thorax one-half broader than long, the

sides nearly straight, the anterior angles thickened, posterior ones

distinct, the surface very obsoletely sulcate near the base, extremely

minutely punctured, flavous, shining; scutellum black ; elytra

punctured like the thorax, flavous, a narrow sutural stripe at the

base and a subtriangular humeral spot as well asa transverse band

near the apex black, this band extends down the suture to the

apex, its posterior edge is consequently deeply concave ; under-

side and legs flavous, the breast and the tarsi black.

Hab. Marcapata, Peru.

A single specimen is contained in my collection. This species

much resembles in its elytral markings certain forms of Diabrotica.

SYSTENA ARGENTINENSIS, Sp. n.

Testaceous, sides of the thorax with a black stripe; elytra

minutely punctured, the suture, a submarginal, medially widened

stripe, and a spot at the base and near the apex black.

Length 5 millim.

Head broad, pale fulvous, microscopically punctured, without

any tubercles or ridges ; clypeus pale flavous, deflexed ; antennz

not extending to the middle of the elytra, testaceous, the third

and fourth joints elongate and equal, terminal joints distinctly

shorter; thorax subquadrate, about one-half broader than long,

with a very obsolete transverse depression near the base, impunc-

tate, the sides with a narrow longitudinal black stripe; elytra

very finely punctured, testaceous, a sutural band, not extending

1904. | AMERICAN PHYTOPHAGOUS COLEOPTERA, 409

to the apex, a stripe, widened medially, near the lateral margins

and likewise abbreviated posteriorly, black, this stripe connected

at its apex with an elongate spot, forming a hook, another small

black spot is placed near the scutellum ; under side and legs

testaceous.

Hab. Prov. Tucuman, Argentine Rep. (C. Bruch).

Of different elytval marking than any other species of the genus

described. In the collection of the La Plata Museum and my own.

SYSLENA ANTENNATA, sp. n.

Flavous, the apical joints of the antenne black ; thorax impune-

tate ; elytra fulvous, extremely minutely punctured.

Mas. Antenne robust, the basal joint strongly thickened, deeply

concave above. Fen. Antenne of normal shape.

Length 5 millim.

Mas. Head impunctate, with a short central groove, the frontal

elevations flat and broad, carina linear; antenne stout, the lower

six joints flavous, the others black, basal joint short and very thick,

deeply hollowed out at the apex, the latter produced into a point,

second joint short, third one-half longer, the following joints

more elongate ; thorax nearly twice as “broad as long, the’ sides

rounded, the disc very obsoletely transversely sulcate near the base ;

elytra wider at the base than the thorax, reddish-fulvous, extremely

minutely punctured, their epipleuree broad and distinct to the

apex; under side and legs flavous, posterior femora rather strongly

inerassate, their tibie with a short spine; metatarsus as long as

the following joints together, claws appendiculate ; prosternum

very narrow, the cotyloid cavities closed.

Hab. Amazons.

The structure of the antenne in the male differs from that of

any other species known to me; in the female, however, the basal

joint is long and slender and the following two joints are both

short and equal and of flavous colour only) all the others being

black: no other differences of importance are present.

PSEUDOGONA DISCOIDALIS, Sp. 0.

Fulvous; the antenne, the apex of the posterior tibie, and the

tarsi black ; thorax impunctate ; elytra parallel, impunctate, black,

a sutural band, transversely widened before the middle and near

the apex, flavous.

Length 4 millim.

Head im punctate, fulvous, with a deep fovea between the eyes,

base of the clypeus very broad between the antenne ; ; eyes large

and round ; labrum and palpi piceous, the latter slender ; antenne

filiform, black, the basal jomt more or less fulvous, fourth and

following joints slightly triangularly widened, not longer than the

third joint which is more slender, terminal joint extending to the

middle of the elytra; thorax about one-half broader than long,

the lateral margins slightly rounded at the middle and constricted

at the base, the anterior angles obtuse; the disc transversely

410 MR. MARTIN JACOBY ON NEW SPECIES OF [ Dec. 13,

convex, impunctate, fulvous; scutellum black; elytra with a few

minute punctures at the base, rest of the surface impunctate, the

sides broadly black, the sutural portion in shape of a flavous

longitudinal band which is transversely widened below the base

and near the apex; underside and legs pale fulvous, the apex

of the posterior femora and the tarsi black, the posterior metatarsi

elongate, claws appendiculate ; anterior cotyloid cavities closed.

Hab. Cordova, Argentine Rep. (C. Bruch).

I received two exactly similar specimens of this species from

Mr. Carlos Bruch.

PSEUDOGONA MILITARIS, sp. n.

Head, the lower joints of the antenne, and the tibie and tarsi

black ; thorax fulvous, impunctate ; elytra opaque, black; a small

spot near the scutellum, a transverse band at the middle and the

apex flavous or whitish ; femora and abdomen fulvous.

Length 54 millim.

Head entirely black, very shining and impunctate, frontal

elevations transverse, short; ; antennze extending beyond the middle

of the elytra, black, the apical four joints fulvous, these rather

stouter than the rest of the joints; thorax of the same shape

and colour as in the preceding species, shining and impunctate,

scutellum black; elytra with a shallow transverse depression below

the base, opaque, extremely minutely punctured, black, the lateral

margins, a narrow small spot near the scutellum, a transverse

narrow band at the middle and the apex broadly, pale fulvous;

the breast, tibie, and tarsi black.

Hab. Panama.

This species must not be mistaken for P. panamensis Jac.,

which it resembles somewhat in its markings; but the last-named

species has very shining elytra without basal depression, set with

two large fulvous spots placed at the base, whilst the posterior

fulvous band is placed below, not at the middle; the antenne also

have the last three joints flavous instead of four.

PSEUDOGONA PALLIDA, sp. n.

Testaceous, the head, the antennz (more or less) and the tibize

and tarsi black; thorax fulvous, impunctate; elytra extremely

minutely punctured, testaceous, an elongate humeral spot black.

Var. Elytra without spots, the middle with an obsolete trans-

verse flavous band.

Length 5 millim.

Of elongate, subcylindrical shape; the head impunctate, black ;

the clypeus flavous ; eyes very large; palpi filiform, pale; antenne

extending to the middle of the elytra, flavous, the intermediate

jomts more or less black, the apical three joints always pale, the

third and following joints nearly equal in length; thorax sub-

quadrate, scarcely broader than long, the sides rounded, the angles

not prominent ; the disc convex, very shining, fulvous, impunctate ;

scutellum small; elytra with indistinct rows of minute punctures,

1904. | AMERICAN PHYTOPHAGOUS COLEOPTERA. 411

only visible under a strong lens, pale testaceous or flavous, the

shoulders with an elongate black spot; femora robust, flavous

like the under side, tibiz partly or entirely as well as the tarsi

black.

Hab. Tucurrique, Costa Rica.

Of the genus Pseudogona, established by me in the Biologia

Centr.-Amer., four species are now known ; they are neat-looking

insects, and distinguished by the subquadrate thorax without sulcus

in connection with the closed cavities, &c. The present species

seems variable in regard to coloration ; the entirely flavous under-

side and the markings of the elytra (when present) distinguish the

species.

OXYGONA AMAZONICA, Sp. 0.

Flavous; the antenne piceous, thorax impunctate; elytra ex-

tremely minutely punctured, a transverse band at the base and

an elongate spot near the apex obscure piceous.

Length 6 millim.

Head impunctate, with a central fovea, clypeus broad between

the antenne, deflexed anteriorly ; antennz extending to the middle

of the elytra, piceous, the third and following joints nearly equal ;

thorax more than twice as broad as long, of usual shape, the sides

slightly rounded, with a narrow margin, the anterior angles oblique,

the disc impunctate, flavous; elytra with a slight depression below

the base, impunctate, with the exception of a few fine punctures

at the base, the latter with a narrow transverse piceous band,

the apex with a similar elongate spot not extending to the

margins; underside and legs flavous, the apex of the posterior

femora and the tarsi sometimes stained with piceous.

Hab. Amazons.

I possess two exactly similar specimens, which differ from their

congeners in the design of the elytra.

OXYGONA CAPITATA, Sp. n.

Testaceous, the base of the head, the thorax, and the scutellum

obscure piceous, thorax impunctate ; elytra finely and closely punc-

tured, antennze pale piceous.

Length 53 millim.

Head impunctate, the vertex pale piceous, the lower portion of

the face flavous; frontal elevations strongly raised, pear-shaped ;

clypeus flattened anteriorly ; antenne extending to two-thirds the

length of the elytra, pale piceous, the third and fourth joints

elongate, equal; thorax twice as broad as long, the sides very

slightly constricted at the base, the anterior angles slightly pro-

duced outwards, thickened, the surface rather convex, pale piceous,

impunetate; scutellum piceous; elytra very finely and rather

obsoletely punctured, much paler than the thorax; under side and

legs coloured like the elytra.

Hab. Peru.

There are two exactly similarly coloured specimens before me,

a2 ON NEW AMERICAN PHYLOPHAGOUS COLEOPTERA. [ Dec. 13,

which I must separate from O. luridus Cl. and O. simplex Clark,

of which I have compared the types in the British Museum.

The head and thorax differ in coloration from these, the head in

the present species having no fovea at the centre, the anterior

angles of the thorax are dentiform and produced, not obsolete as

in O. simplex, and the general size is much smaller.

CREPIDODERA LONGICORNIS, sp. n.

Oblong-ovate, black, the basal joints of the antennz and the

legs fulvous; thorax subquadrate, minutely punctured, deeply

suleate ; elytra convex, finely punctate-striate ; antenne very long ;

legs robust.

Length 4 millim.

Of broadly ovate shape, the head impunctate, black; frontal eleva-

tions broad, trigonate, strongly raised ; clypeus convex between the

antenne, its anterior margin of the shape of a semicircular ridge ;

antenne nearly extending to the apex of the elytra, black, the

lower three joints fulvous, the third joint longer than the others,

basal joint thickened, elongate-cylindrical, terminal joint about

as long as the third; thorax convex, but slightly broader than

long, the sides rounded at the middle, the angles not produced,

the disc microscopically punctured, with a deep transverse sulcus

near the base, bounded at the sides by a perpendicular groove ;

elytra wider at the base than the thorax, widened towards the

middle, with a distinct basal transverse depression, the shoulders

prominent, the surface finely punctate-striate, black and shining,

the interstices flat; legs robust and short, fulvous, all the femora

thickened, the tibize strongly widened at the apex, the first joint

of the tarsi broadly flattened.

Hab. Peru.

I know only a single specimen of this species, which, by its

robust legs, the long antenne, and general coloration, is well dis-

tinguished. In the last respect, the species resembles evidently

C. peruviana Harold (Deutsche ent. Zeitsch. 1877, p. 130), but

it is much larger, the antenne are differently coloured, and the

elytral punctuation is distinct to the apex.

HIPPURIPHILA CATHARIN A, Sp. nN.

Black below, above greenish-zeneous, the basal joints of the

antenne fulvous ; thorax very strongly punctured, with a feeble

transverse sulcus; elytra strongly punctate-striate, the base with

a transverse depression.

Length 2 millim.

Short, convex, and ovate; the head impunctate, eneous, a single

punctiform impression above each eye, the latter rather large,

from the upper margins of which a deep narrow groove extends

obliquely to the base of the antennz, both grooves meeting at the

centre ; clypeus convex between the antenne, widened anteriorly ;

PZ. Sls04 vol de Plea:

Pree en

H.Grénvold del. et hth, Mintern Bros .imp.

SIMIA VELLEROSUS (Gray)

(very old male)

1904. | ON ANTHROPOID APES. 413

antenne rather short and robust, black, the lower three joints

fulvous, the second and third ;joints small, equal, the terminal six

joints rather strongly widened, but longer than broad; thorax

scarcely twice as broad as long, the sides nearly straight, obliquely

angulate before the middle, very narrowly margined, basal margin

oblique at the sides, rather strongly produced towards the scutellum,

preceded by a shallow transverse suleus which is bounded at the

sides by a deep perpendicular groove, the surface rather convex,

somewhat closely and strongly punctured ; elytra with a shallow

depression below the base, the shoulders rather prominent and

smooth, the surface strongly and regularly punctate striate, the

interstices impunctate and flat; under side and legs black, base of

tibie and the tarsi more or less fulvous, prosternum narrowly

elongate; with a longitudinal shallow sulcus,

Hab. Santa Catarina, Brazil.

The short, ovate shape of this species, short and rather robust

antenne, and the produced basal margin of the thorax at the

middle agree best with the species at present placed in Hippuri-

phila instead of Crepidodera proper ; four specimens are contained

in my collection.

3. Notes on Anthropoid Apes.

By the Hon. Watrer Roruscuinp, Ph.D, F.Z.8.

| Received December 13, 1904. |

(Plate XXTIV.* and Text-figuires 99-117.)

Although, from the earliest times, beginning with Hanno’s

Gorille, we find the writings of observers of nature filled with

accounts of hairy wild men, and, in later days, many descriptions

by zoologists of anthropoid apes, it was only after the appearance

of Du Chaillw’s book that universal attention was turned to

these creatures.

Prior to 1870, several so-called species both of Gorilla, Anthro-

popithecus, and Simia auct. had been established, but until lately

the majority of zoologists maintained that there was only one

rather variable species each of Gorilla, Chimpanzee, and Orang-

Outan. Professor Matschie’s articles on the genus Gorilla

(Sitzungsb. Ges. naturf. Freunde, 1903, pp. 253-259, and 1904,

pp. 45-53) and his articles on the species and races of Chimpanzee

(Sitzungsb. Ges. naturf. Fr. 1900, pp. 77-85, and 1904, pp. 55-69)

have, however, once more aroused the greatest interest in the

question of the true status of our knowledge of the anthropoid

apes. In the first place, Professor Matschie insists, and, I believe,

rightly, that the Gibbons (//ylobates) should form a separate

* For explanation of the Plate, see p. 440.

4l4 [ Dee. 13,

THE HON. WALTER ROTHSCHILD ON

family—Hylobatidee—and not be united with the great man-like

apes. Although I propose to deal mainly with the Chimpanzees,

I must allude to the genera Gorilla and Pongo, as I differ slightly

from Professor Matschie.

In the first place, Professor Matschie himself gives only

TABLE of Comparative Measurements

of the Skulls of Gorilla gorilla and G'. gorilla matschiet.

Hinder surface of Head :

Greatest breadth at the Mastoid

processes.. wee 144, 147, 155, 162 mm. 170 mm.

Breadth above the Mastoid pro-

CeSSeS ...... .... 116, 120, 121, 123 mm. 163 mm.

Length from centre of Crista

lambdoidea to basal edge of

Foramen magnum ......... 90, 100, 101, 103 mm. 109 mm.

Foramen magnum :

THGSTERID, coccroececuasdaeaccccneccsene Uy G5 Sin CHI raha. 40 mm.

Breadth .. ..... 26, 28, 29, 30 mm. 35 mm.

Breadth on outside of Occipital

CONG ylesmeee teers ecese ees: 45, 45, 51, 51 mm. 59 mm.

Basioccipitatl :

Length from front edge of

Foramen TORFATUIN — sccosncoaeen 45, 45, 46, 49 mm. 35 mm,

Occipital condyles 5

Breadth at base ............... 31, 32, 35, 38, 39 mm. 40 mm.

Breadth at anterior edge ...... 23, 24, 29, 30 mm. 20 mm.

Vomer:

TET scosconnccedonnenecaercceceece — UKs IMS), IIS), is) sony. 19 mm.

ISSRSEO ND Me hae aenace SauBooabeses cocognaG 16, 17, 18, 19 mm. 10 mm.

Basisphenoid :

Width at Foramen rotundum... 58, 58, 58, 59 mm. 62 mm.

Width at apex of Petrous

portions of Temporal ......... 32, 33, 35, 85 mm. 30 mm.

Pterygoid processes sh San

Length ........ 64, 65, 68, 69 mm. 51 mm.

Breadth, singly .. 17, 18, 18, 19 mm. 24 mm.

Breadth across ...... 60, 60, 60, 62 mm. 79 mm.

Articular es of Lower « Jaw:

Width.. : 32, 34, 36, 36 mm. 42 mm.

Coronoid pi ocess :

Greatest width .......... 26, 30, 30, 34 mm. 21 mm.

Width between Cor onoid. pro-

cess and Articular condyle... 34, 34, 34, 41 mm. 21 mm.

Width from outside of Coronoid

process to outside of Articular

condyle .........000ceceeeeeeeeeeee 70, 73, 75, 79 mm. 56 mm.

Nasals:

THSneAtIN “Gonecasagssoccdoaarboeeaonmne | Glen 240), Ail ei) vervasr, 45 mm.

Width 28, 29, 33, 86 mm. 29 mm.

Premaxilla:

Length : 31, 31, 34, 36 mm. 39 mm.

Br eadth across s the canines.. 55, 70, 71, 73 mm. 90 mm.

Zygomea :

Breadth at molar portion ...... 30, 32, 32, 38 mm. 43 mm.

Breadth at narrowest part ...... 16, 16, 17, 17 mm. 16 mm.

Length of skull front of Arcus

superciliaris to front of Pre-

maxilla ......0..-.0:--0.--- 145, 146, 150, 153 mm. 160 mm.

1904. | ANTHROPOUD APES. 415.

doubtful characters, from lack of material, for Gorilla castuneiceps

of Slack. The latter author gives as one of the principal characters

(if not the principal) of his species, the red crown; now I have

seen a good many Gaboon and Ogowe Gorillas, and I have found

the red colour so variable that I am forced to regard Gorilla

castaneiceps merely as a casual aberration of Gorilla gorilla. The

cranial characters, as given by Matschie, appear to me also very

uncertain, On the other hand, the Gorilla manyema of Alix and

Bouvier I believe to be a very large ape of the group of Simia

vellerosus Gray, and not a Gorilla at all, although Professor

Matschie places it as a synonym of Gorilla castaneiceps. While

I consider G. castaneiceps to be an aberration only of G. gorilla,

I think Professor Matschie was rather bold to unite all South

Camaroons Gorillas with the typical Gaboon G. gorilla. The

Camaroons specimens I have seen appear to me to have shorter

and stouter limb-bones, much longer hair, and the skulls show

as a rule, though not always, a higher crista sagittalis. The

facial portion is also shorter than in G. gorilla. These characters

are more or less given by Matschie as probable points of distinction

between G. castaneiceps and G'. gorilla, but Slack did not found

his species on these characters. Professor Matschie has separated

the North Camaroons form of Gorilla as G. diehli on the evidence

of eight skulls, all of which have the planwm nuchale much wider

than high. Jam inclined to think that the N. and 8, Camaroons

Gorillas are merely geographical races of the Gaboon and Ogowe

Gorilla gorilla, while, owing to the presence of full beard and the

skull having certain very peculiar differences, the Gorilla from

Kirunga, in German Kast Africa, ought to be upheld as a species,

at least till we can examine fuller material. I propose to call the

S. Camaroons race Gorilla gorilla matschiei, subsp. nov. Hair

longer than in Gorilla gorilla, whole back and fore part of legs

much greyer, limbs much shorter and stouter; crest of skull

generally higher and rising closer to the arcus superciliaris; skull

generally shorter: female much greyer,

From the foregoing particulars it will be seen that Gorilla

gorilla and G, gorilla matschiei differ widely in the proportions of

their skulls. (I have compared five fully adult males of equal

size, all much above the averge size.) The most striking differ-

ences are certainly in the shape of the hinder surface of head

and the basioccipital bone, as well as the very widely different

portion of the lower jaw comprising the coronoid process and the

articular condyle. I have compared numerous other Gorillas’

skulls—in all 27 $ and @, adult and young—in my possession,

both from the Gaboon and the Camaroons, but they are all more

or less imperfect or less adult than the five compared, so that the

measurements could only have been partially given, therefore I

did not think it advisable to quote them in this paper.

The casts of the type skulls ¢ 2 of Gorilla diehli Matschie

agree perfectly with two skulls wanting the lower jaws which I

416 THE 11ON. WALTER ROTHSCHILD ON | Dec. 13,

possess, and which were brought back by Mr. G. L. Bates from

the Camaroons, they being native killed, while the cast of

G. beringeri Matschie shows differences from all skulls known

to me.

According to this classification, the species and subspecies of

Gorilla would stand as follows :-—

GORILLA GORILLA (Savage & Wyman), (Text-figs. 99 & 100.)

Boston Journal of Natural History, vol. vy. p. 419 (1847).

Synonyms: Gorilla gorilla Is. Geoftr., 1852. Troglodytes savaget

Owen, 1848. Pithecus gesilla Blainv., 1859. Satyrus adrotes

Mayr, 1856. Chimpanza gorilla Haime.

Text-fig. 99.

a We

Skull of Gorilla gorilla (Savage & Wyman). (Side view.)

Aberration: castaneiceps Slack Proc. Acad. Nat. Sci. Phila-

delphia, pp. 159-160 (1862).

Habitat. Gaboon and Ogowe Region.

GoRILLA GORILLA MATSCHIET Rothsch. antea, p. 415. (‘Text-

figs. 101 & 102, pp. 418, 419.)

Habitat. Southern Camaroons.

1904. | ANTHROPOID APES, 417

Text-fig. 100.

Ze Hl

¢ <1 pitep i

. os J

ee:

Skull of Gorilla gorilla (Savage & Wyman). (Front view.)

Proc. Zoou, Soc.—1904, Vor. Il. No. XXVII. 27

418 THE HON. WALTER ROTHSCHILD ON | Dec. 13,

Text-fig. 101.

PAs

Mii

SSS

Skull of Gorilla gorilla matschiei Rothsch. (Side view.)

GoRILLA GORILLA DIEHLI Matschie. (Text-fig. 108, p. 420.)

Sitzungsber. Ges. naturf. Freunde Berlin, 1904, p. 52.

Habitat. Northern Camaroons.

GoRILLA BERINGERI Matschie. (Text-fig. 104, p. 421.)

Sitzungsber. Ges. naturf. Freunde Berlin, 1903, pp. 253-259.

Habitat. Kirunga, Ya Sabinyo Volcano, German East Africa.

The genus and species of Chimpanzees now must be considered,

and the first and most vexed question is that of the correct

nomenclature. In common with Mr. Oldfield Thomas and most

of the continental and American zoologists, I adopt, as the starting-

point, Linneus’s tenth edition of the ‘Systema Nature’ (1758).

This being the case, I must now go into the changes it necessitates.

In the first place, Anthropopithecus Blainville, 1838, must sink, as

we find by the help of Palmer & Merriam’s ‘Index Generum

Mammalium,’ p. 109, that there are the following generic names

older than Anthropopithecus, viz.:—Troglodytes Geoffroy, 1812;

Pan Oken, 1816; and Theranthropus Brookes, 1828. Troglodytes

was used as a name for the Wren in 1806 by Vieillot, while

1904. ] ANTHROPOID APES. 419

Text-fig. 102.

Ly W WS

oo5) n ANS SS

dd pi Si

Mr 4 ) ‘

‘ WN

BEE

Skull of Gorilla gorilla matschiei Rothsch. (Front view.)

Mimetes of Leach, 1820, another name for the Chimpanzee, was

also preoccupied. I, therefore, who, in opposition to Professor

Matschie, consider Oken’s names applicable, would have had to

accept Pan as the generic name of the Chimpanzee, as do many

American writers, but for the fact that a still older name exists.

Linneus describes as the first species of his genus Sima in the

27*

420 THE HON. WALTER ROTHSCHILD ON | Dec. 13,

Text-fig. 103.

My MU 7

Shy

Ye)

Yi) {iy

Yee

Yi!

LUNs ne

ty, wot

PUN ue

/ is RNS WW) yr _

COsater ane

Skull of Gorilla gorilla diehli Matschie. (In Tring Museum.)

‘Systema Nature,’ 1. p. 25 (ed. x. 1758) an anthropoid ape as

follows :—

“ Satyrus. 1. 8. ecaudata subtus nuda. Syst. Nat. vi. p. 3.

Satyrus indicus Tulp. obs. IIc. 56.

Habitat in Africa. Asia.

Magnitudine puerl sexennis. Dorsum crinibus nigris_ hir-

sutum ; subtus s. antice undique glaber.”

Tulp described and figured an ape which was brought from

West Africa and presented to Prince Frederick Henry of Orange,

and which lived some years in Kurope.

Linneus copied Tulp’s description almost word for word, and,

as quoted above, expressly states that the S. satyrus was black on

the back. It was only in his twelfth edition (1766) that Linneus

calls the Orang-Outan Simia satyrus, and says it is red-haired,

but he had already, in the ‘Ameenitates Academice,’ vi. p. 69

(1763), named the red-haired animal Simia pygmeus. Not only,

1904. ] ANTHROPOID APES. 42]

Text-fig. 104.

LAL i

Hi 74

ab tin dsa

VEU NDCHEL ” 4

\ ee 4 A cA

————F

navola@.

Skull of Gorilla beringeri Matschie.

however, can we prove that Linneus’s Simia satyrus is really a

Chimpanzee, but we can even distinguish the exact race to which

the name applies, for Tulp’s description and figure show an ape

the hair of which is not parted in the centre of the head, and

with a short thick beard clothing the cheeks and leaving the chin

bare. ‘These characteristics are found in the Chimpanzees from

the coast-lands of the South Camaroons and the Gaboon and

Ogowe districts. It is therefore necessary to adopt the name of

Simia L. for the genus of the Chimpanzees, and the famous

““Tschego” proves to be the veritable Simia satyrus. According

to Palmer, on the other hand, we have the following generic

terms for the Orang-Outan or Maias :—

Satyrus Lesson, 1799; Pongo Lacepede, 1799; Pithecus

G. Cuvier, 1800; Lophotus G. Fischer, 1813; Faunws Oken,

1816; Macrobates Billberg, 1828; and Brachiopithecus Sénéchal,

1839. As Satyrus of Lesson and Pongo of Lacepede are of equal

date, I think we must adopt, as the least confusing name, Pongo

of Lacepéde, and therefore the correct name of the Orang-Outans

as a group is Pongo pygmcus (Linn.).

Professor Matschie, in his article on the Chimpanzees, Sitz. Ges,

422 THE HON. WALTER ROTHSCHILD ON [ Dee. 13,

naturf. Fr. 1904, pp. 55-69, acknowledges seven species of the

genus Simia. According to the view of the value of various

animal forms which I take up, as a large proportion of these

represent one another geographically, they ought to be treated

only as subspecies. This diversity of opinion between Professor

Matschie and myself is more apparent than real, for in many

cases Professor Matschie regards what I call “species” as genera

or subgenera, while he considers what I call “subspecies” to be

species, thus only differing in the terms to apply to certain

categories of individuals. According to our present state of

knowledge of the Chimpanzees, there are two very well-defined

groups, namely, the Simia satyrus group, with black or blackish-

brown faces when adult, and the Simia pygmeus group, with pale

faces both in the adult and young stages. From this it will

be seen that I differ entirely in one point of nomenclature from

Professor Matschie—namely, I hold that a specific name can be

used for a species, even if previously used in a different sense,

so long as the species first denoted by the name has since been

placed in another genus. I therefore consider Simia pygmeus

applicable to one of the races of Chimpanzee, because Linnzeus’s

Simia pygmeus must now stand as Pongo pygmeus. In addition

to the seven forms of Simia recognised by Matschie in his paper

(Sitzungsber. Ges. nat. Fr. Berl. 1904, pp. 54-69), there are several

more, among which is a pale-faced Chimpanzee which comes from

some part of the French Congo, which I propose to name Sima

pygmeus raripilosus, subsp. nov., distinguished from other forms

of Simia pygmaeus by the sparse, almost absent, beard, narrow

protruding face, and very long limbs, largish ears, rounded fore-

head, and only partially divided hair on the head.

Professor Matschie’s paper gives the forms as follows :—

1. Simia satyrus L. Syst. Nat. 1. p. 25 (1758). (Text-figs. 105,

106, 107, & 109, fig. 1, pp. 423, 424, 426.)

Synonyms: Sima troglodytes Gm., 1788; Troglodytes niger

Geoffr., 1812; Vvroglodytes koolo-kamba Du Chaillu, 1861; 7Z'ro-

glodytes aubryi Gratiolet & Alix, 1866; Psewdanthropus fuliginosus

Schaufuss, 1875.

Distribution. Lower Guinea from Sanaga in the Camaroons to

the Ogowe.

Distinctive characters. Eaternal: hair of head not parted in

centre; hair on forehead falls out in adults, but not so far as level

of ears. Hars medium size, 65 x 50 millimetres. A narrow beard

of thick short hairs pointed downwards surrounds the face except

chin, which is sparsely covered with dark grey hairs. Arms very

long, exceeding 700 mm. Colour of face in the young leather-

yellow, in adult animals blackish brown.

Cranial characters: facial portion of skull very narrow, much

constricted behind the canine teeth; the greatest breadth of the

skull is never more than 1 mm. greater at the canines than at

the molars. The brain-case is an elongate egg-shape, measuring

from the glabella to the protuberantia occipitalis in the 5 g

1904. } ANTHROPOID APES. 423

Text-fig. 105.

Head of Simia satyrus Linn. (From life.)

Text-fig. 106.

ING

Pen

nine

: Wl

a i)

Skull of Simia satyrus Linn, (Fully adult.)

424 THE HON, WALTER ROTHSCHILD ON [ Dec. 13,

Text-fig. 107.

& EVR i

Skull of Simia satyrus Linn. (Fere adult.)

134-142 mm. and in the 9 2 122-130 mm. ‘The thinnest place

in the zygomatic arch is 5-9 mm. high (always over 7 mm. in

old gS).

2. SimiA CALVus (Du Chaillu), Proc. Boston Soc. Nat. Hist. vii.

p-. 296 (1861).

Distribution. Interior of Gaboon Region and Southern

Camaroons.

Distinctive characters. Haternal: hair of head not parted in

centre, falls out when adult to behind base of ears; the ears are

enormous, 80x53 mm. at least. Beard laterally thin and longer

than in S. satyrus, and does not join under the chin. Chin

sparsely covered with white hairs. Length of arms in adult

animals 600 mm. Colour of face in adult animals brownish

black ; eyes light brown, in young animals wood-brown, ears

yellowish.

Cranial characters : facial portion of skull apparently very wide,

as it is expanded behind the canines. The greatest width at the

canines 1s 5 mm. narrower than at the molars. The brain-case is

round, the greatest length from the glabella to the protuberantia

occipitalis externa is the same in ¢ ¢ and @ Q, viz. 127-139 mm.

1904. | ANTHROPOID APES. 425

and over the arch of the forehead 141-160 mm. The breadth at

the canines is 50-60 mm. in the ¢ ¢ and 54-68 mm. in the 9 @.

The thinnest place in the zygomatic arch is 7-10 mm. (always at

least 9 mm. in old ¢ CS).

3. SIMIA VELLEROSUS (Gray), P. Z. 8S. p. 181 (1862). (Plate

XXIV. and Text-figs. 108 & 109, fig. 2.)

Distribution. Northern Camaroons and higher mountains

further south.

Distinctive characters. External: ears very small, 50 x 45 mm.

Beard very long and thick, completely surrounding face. Arms

very long, at least 750 mm. in length; face brown (IV. &.).

(Colour of hair in fully adult old ¢ 3 is yellowish grey.—IV. Rh.)

Text-fig. 108.

/ Li

\OeCMen y

frm SA iT

De y Wis

C hg)

a Iii

Skull of Simia vellerosus (Gray).

Oranial characters : facial portion of skull slender, flat in front

of nostrils; breadth behind canines from 1 mm. less to 1 mm.

more than between the molars. Brain-case, measured from the

glabella to the protuberantia occipitalis eaterna much shorter in

the ¢ $6 than in the 9 @, viz. 131-132 mm. and 137 mm.

respectively ; measured over the arch of the forehead 150-152

mm. and 160 mm. respectively. Breadth at canines in 3 o

62-63 mm., in 2 9 55mm. The thinnest place in the zygomatic

arch is 6-9 mm. high.

426 THE HON. WALTER ROTHSCHILD ON [ Dec. 13,

Text-fig. 109.

Lett canine tooth of :—l. Simia satyrus Linn.

is ys 2. Simia vellerosus (Gray).

4, SIMIA SCHWEINFURTHI Giglioli, Ann. Mus. Civ. Genova, 111.

p. 135 (1872). (Text-fig. 110, p. 427.)

Synonyms: Z'roglodytes marungensis Noack, 1887.

Distribution. Niam Niam to South-eastern Soudan, and from

Lake Tanganyika to Uganda Protectorate (and perhaps to Lake

Chad and Wadai.—IV’. 2.).

Distinctive characters. Haternal: jface, when young, pale,

when adult, according to Matschie, dark. ars very large (no

exact measurements known). eard enormously long and thick.

Chin thickly covered with long white hairs. Arms very long

indeed. Hair generally very long and thick.

Cranial characters: facial portion of skull extremely narrow, at

the very outside only 55 mm. broad. behind the canines, but not

so wide even at the widest part of the palate. DBrain-case almost

round, of equal length in both sexes, measured from the glabella

to the protuberantia occipitalis externa 128-133 mm., measured

over the arch of the forehead 150-160 mm. Breadth between

the molars 51-55 mm., between the canines 49-55 mm. The

thinnest place in the zygomatic arch is 4-7 mm. high.

5. SmmiA Fuscus (Mayer), Abh. und Ber. Mus. Dresd. No. 14,

p. 7 (1894-1895).

Distribution. ¢ Probably between Liberia and Togoland.

Distinctive characters. External: hair-whorl on the top of the

head, from which hair falls on all sides. Har blunt, almost flat

at the top. Region of eye darker than nasal region. Beard long

and entire.

Cranial characters : facial portion of skull slender. About the

same width behind the canines as at the molars, viz. 53 mm.

Brain-case in a Q skull measures in length from the glabella to

the protuberantia occipitalis externa 128 mm., and over the arch

of forehead 155 mm. Zygomatic arch at its thinnest place is

still 8-5 mm. high.

1904. ] ANTHROPOID APES. 427

Text-fig. 110.

Simia satyrus schweinfurthi (Gigl.). (From a photograph from life.)

6. SIMIA LEUCOPRYMNUS Lesson. 1831.

Synonyms: Sima pygmeus Schreber, Siiugthiere, Taf. 1B

(1796).

Distribution. Probably Sierra Leone and Western Liberia.

Distinctive characters. KHxternal: hair of head parted in the

centre, very thin on the strongly-arched forehead. Hars very

large and rounded at the top. Legion of the eyes as pale as rest

428 THE HON. WALTER ROTHSCHILD ON [ Dec. 13,

of face. Chin sparsely clothed with brown hairs. Beard short

and thick and surrounding chin.

Cranial characters: brain-case flat; occipital region flat and

much lengthened ; facial portion of skull slender, not expanded

at the molars.

7, SIMIA CHIMPANSE (Mayer) Arch. Naturg. xxii. (i.) p. 282

(1856).

Distribution. Gambia and Senegambia.

Distinctive characters. Haternal: hair of head parted in centre ;

ears large; beard laterally long, standing out from the face ; chin

free and clothed with white hairs.

To these forms I must add the following :—

SIMIA RARIPILOSUS Rothsch. antea, p. 422. (Text-fig. 111.)

Distribution. Probably some part of the interior of French

Congo.

Text-fig. 111.

Head of Simia pygmeus raripilosus Rothsch. (From life.)

Distinctive characters. Haternal: hair of head not or partly

1904. | ANTHROPOID APES. 429

parted in centre; hair very sparse all over body ; arms very long ;

beard almost absent.

Since writing the paper (Sitzungsb. Ges. naturf. Fr. Berl. 1904,

pp. 55-59) Professor Matschie has examined over one hundred

Chimpanzees, including the types of Zrog. aubryi, Tr. niger,

Tr. calwus, Tr. koolookamba, Tr. tschego, Tr. troglodytes, Tr.

fuliginosus, Tr. marungensis, and T'r. leucoprymnus, besides many

skulls, and in consequence has found much in the above-quoted

article which requires altering. With such of those alterations

as he has communicated to me I quite agree. The conclusions

both he and I have come to make it clear that there are at least

four distinct species of Chimpanzee (Simia) all living side by side

throughout the greater part of their range, while I, personally,

maintain there are five such species. The five species, according

to the geographical and physical position of various portions of

their range, again fall into a number of well-developed subspecies

= geographical races; and we find that at present we have

twelve named races belonging to five species; while Professor

Matschie, in a forthcoming paper, proposes to describe, besides

others, two new forms from Liberia, two from Central Congo, and

three from the Uelle region. At present I only propose to deal

with the twelve named forma of which the following are the Key

and Synopsis, which latter, I hope, will be understood by aid of

the former ; but the notes following them will no doubt clear up

much.

KEy.:

Ce i i ary

Face of adult black or blackish brown

Face of adult pale

Co bo

Peewee eee tees tees esse eee esse eee es es eeeeeesses

{ Hair long, harsh and black; arcus superciliaris strongly

| developed.............. chieebtien Simia satyrus marungensis.

Hair long and soft, generally yellowish grey in very

old animals. Last lower molar very small ; facial

| portion of skull very short ; canines very large.

Simia vellerosus.

| Hair long and soft, sooty-brown to black.

Simia vellerosus fuliginosus.

Hair and beard very long, limbs long; head narrow

02 and very high, face olive. brown.

‘ Simia satyrus schweinfurthir.

| Hair black, short, and harsh; head round, ears enormous.

Simia koolookamba.

| Hair short, harsh ; head long; arcus superciliaris

strongly developed. Last lower molar showing four

| very ill-defined tubercles, facial portion of skull

a Camimesismialllliracce tn saseeaes. esas saas Sinia satyrus.

Hair “plack and short, ears very small; face very

prognathous ; last lower molar with five large tuber-

cles as in Gorilla ; eyes very wide apart ... Simia aubryi.

430 THE HON. WALTER ROTHSCHILD ON [ Dec. 13,

3 Region round eyes dark......... Simia pygmeus chimpanse.

: Region round eyes paleshike rest.of face seaesseneeaae: 4

Beard short, sparse, almost absent; hair short, thin,

| and black ; face pale, blotched with dark tan.

Simia pygmeus raripilosus.

| Face pale flesh-colour, beard thick and long; hair

long, thin, and black ; head round, ears set on low.

4d Simia pygmeus.

Hair, even when young, mostly reddish ; hair on chin

| whitish ; hair of head spreading from central whorl,

beard entire and long......... Simia pygmeus fuscus

Hair on head sparse, ears large, chin-hairs brownish.

L Simia pygmeus leucoprymnus.

This Key is as close as I have been able to work, but when

some of the remaining subspecies are described it must be revised.

SYNOPSIS.

1. Simia vellerosus (Gray). Camaroons.

la. Simia vellerosus fuliginosus (Schaufuss). Congo region.

2. Simia satyrus (Linn.), Camaroons and Gaboon.

2a. Simia satyrus marungensis (Noack). Central Congo. (Text-

fig. 112, p. 431.)

9b. Simia satyrus schweinfurthi (Giglioh). Soudan and Uganda.

ae ui Awe §

3. Simia koolookamba (Du Chailla). Camaroons and Gaboon,

4. Simia aubryi (Gratiolet & Alix). Camaroons and Gaboon.

5. Simia pygmeus Schreber. Congo. (Text-fig. 113, p. 432.

5 YI Z co, ‘ to} ?

5a. Simia pygmeus fuscus (Mayer). Gold Coast 2

5b. Simia pygmeus leucoprymnus (Lesson). Sierra Leone and

8. Liberia.

5e. Simia pygmeus chimpanse Matschie. Gambia. (Text-

fig. 114, p. 433.)

5d. Simia nyqmeus raripilosus Rothschild. French Congo.

Y. S

In Professor Matschie’s previously cited paper, first of all, under

the head of Simia satyrus Linn. he has confused four species,

viz. S. satyrus, S. koolookamba, S. aubryi, and S. vellerosus, in fact

all the black-faced species. As now ascertained, Linnzeus’s species

must stand as Sima sArykus Linn. Syst. Nat. 1. 25 (1758).

Synonyms: Simia troglodytes Gm., 1788; Troglodytes niger

Geoftr., 1812; Zroglodytes tschego Duvernoy, 1855; and Z’roglo-

dytes calvus Du Chaillu, 1861.

Troglodytes aubryi Gratiolet & Alix, 1866, must stand as a good

species as StmzA AuBRyI (Grat. & Alix).

Troglodytes koolookamba Du Chaillu must stand as a distinet

species as Smita KOOLOOKAMBA (Du Chaillu). (Text-fig. 115, p. 434.)

Simia calvus Matschie (nec Du Chaillu) is the same as Simia

vellerosus (Gray).

Simia vellerosus Matschie (nec Gray) is a mixture of S. velle-

rosus and S. aubryi.

1904. | ANTHROPOID APES. 431

Pseudanthropus fuliginosus Schaufuss is the Loanga subspecies

of Gray’s Troglodytes vellerosus, and will have to stand as StMIA

VELLEROSUS FULIGINOSUS (Schauf.).

Simia pygmeus Schreber is not a synonym of 7’. lewcoprymnus

Lesson, as Professor Matschie has placed it, but is a distinct sub-

species,

Text-fig. 112.

Head of Simia satyrus marungensis (Noack). (From life.)

In the preceding pages I have endeavoured to compress into

concise limits all that could be ascertained of the natural divisions

and classification of the African anthropoid apes in the light of

modern study and investigations. We now come to the Asiatic

anthropoids; and if the Gorillas and Chimpanzees offer serious

difficulties to the student, the Asiatic Orangs present difficulties

ten times worse. I do not for a moment wish to assert that my

conclusions are even as closely correct as in the Gorillas and

432 THE HON. WALTER ROTHSCHILD ON [ Dec. 13,

Chimpanzees, but seeing that we are faced by two distinct problems,

when trying to classify the large red apes of Borneo and Sumatra,

I think the present arrangement clears up a few difficulties and

is an advance on former classifications. The first of the above

problems connected with the Orangs is, that throughout their

entire range two forms are found living side by side which are

extremely different in appearance. In one of these the adult

males are very large and have huge callosities on each side of the

face, in the other the adult males are smaller and have no sign of

any face- (i.e. cheek-) callosities. There is considerable diversity

Text-fig. 113.

Head of Simia pygmeus Schreber. (From life.)

of opinion as to what is the correct position to assign to these two

forms. Professor Matschie not only considers them to be distinet

species, but even goes so far as to say they are distinct genera. I

cannot at all agree to even considering them distinct species, but

feel sure they are only dimorphic phases of one species.

The second problem presented by the Orang-Outans is whether

there are a number of different species or whether there is only

one variable species consisting of a number of geographical races

or subspecies.

Professor Matschie inclines to the former view, while Dr. Selenka

1904. ] ANTHROPOID APES. 433

takes the latter. I am convinced that this is the right view to

take, and that many who side with Professor Matschie go too far

in splitting up the forms of Orang. Dr. Selenka gives a very

plausible and, I believe, well justified explanation for the exist-

ence of a number of local races in Borneo, viz., that the Orang-

Outans cannot swim and can only climb mountains, when bare of

trees, with difficulty ; and as Borneo is intersected in all directions

by broad rivers and high mountain-vanges, the Orangs in the

various districts are almost as much isolated as if confined to

separate islands.

Text-fig. 114.

Head of Simia pyymeus chimpanse Matschie. (From life.)

Dr. Selenka separates 8 races of Orangs from Borneo and

Sumatra, 4 with cheek-callosities and 4 without, so that, as I

consider these two forms dimorphic phases, he distinguishes

actually 4 distinct subspecies. Professor Matschie distinguishes

14 races, or, as he calls them, species, from Borneo and Sumatra,

Proc, Zoom Soc —lo04e vious lle No, XOXOVNES 523)

A434 THE HON. WALTER ROTHSCHILD ON | Dee. 13,

or according to my view 7 distinct subspecies, each being dimorphic.

His 3 additional subspecies are Bornean, and will he described in

his forthcoming paper. As I cannot find in previous writings on

the Orangs any descriptions which can be employed to denote

other forms than the 4 dimorphic subspecies recognised by

Dr. Selenka, I shall only deal with them, and leave Professor

Matschie to work out any forms, in addition to these 4, which may

exist in collections.

Text-fig. 115.

Head of Simia koolookamba (Du Chaillu). (Very young animal.)

(From life.)

Dr. Selenka, in the Sitzungsber. Akad. Wissensch. Berlin,

xvi. pp. 381-392, gives distinctive characters of lis 8 races as

follows :-—

Borneo.

A. 3 ¢ with Cheek-callosities.

1. Pithecus satyrus lundakkensis.—Haiv deep reddish brown,

rarely brownish yellow. Skull mikrencephalic and micro-

gnathous; cubic capacity, J g 420-450 cb.cm., 2 2 350-

390 cb.cm, 4th molar rarely present.

1904. | ANTHROPOID APES. 435

_ Pithecus satyrus batangtuensis.—Hair deep brown. Skull

mikrencephalic and micrognathous; cubic capacity 400-437

eb. cm. in 6 G, 350=420 chem. in 9 2°

3. P. satyrus dadappensis.—Hair dark reddish brown. Skull

megalencephalic and macrognathous ; cubic capacity in dd

470-534 cb.em., in 9 Q 360-490 cb. cm.

B. ¢ ¢ without Cheek-callosities.

P. satyrus genepaiensis.—Hair deep reddish brown. Skull

megalencephalic and macrognathous ; cubic capacity in ¢ d

390-435 cb.cm., in 2 9 360-410 cb. em.

P. satyrus skalauensis. — Hair dark brownish red. Skull

megalencephalic and brachygnathous ; cubic capacity in

3 g 440-500 ch. cm., in 2 Q 330-440 cb. em.

P. satyrus tuakensis.— Hair stiff and coarse, rusty yellow to

rust-red; skin reddish ; shape clumsy and expression coarse.

Skull mikrencephalic and maecrognathous. Malays call this

form “Maias kesar,” the ‘ Coarse Orang,” and this is the

form almost always seen alive in Europe as it is hardier and

travels better.

(Sy!

SUMATRA.

_ Pithecus sumatranus deliensis—Hair brownish to foxy red.

Face black. Old males with large cheek-callosities, mesen-

cephalic ; cubic capacity In do 445-485 cb. cm., in 9

340 cb. em.

8. P. sumatranus abongensis.—Hair deep brown. Old males

without cheek-callosities.

“i

Except that Dr. Selenka entirely overlooked the fact that of the

8 forms recognised by him 5 had already been named and de-

scribed, and that therefore only 3 of his names can stand, he was

most accurate in his distinctions, which is not to be wondered at

as he has over 300 skulls, about 100 complete skeletons, about 100

skins, and many embryos and young for comparison, an amount

of material obtainable nowhere else in the world. In addition to

these 8 forms Dr. Selenka described, tentatively, a ninth form

without cheek-callosities as P. satyrus rantaiensis, which will

probably prove to be one of Prof. Matschie’s 3 additional species.

The following table will explain the relationship of the various

subspecies of Orang-Outan, so far as the present state of our

knowledge enables me to judge.

The table, however, requires considerable explanation, for

although I have employed similar nomenclature In my entomo-

logical articles, it is something quite new when employed in

connection with Vertebrata. As I have shown previously in this

paper, the first name applied to the Orang-Outan after 1758, our

nomenclatorial starting-point, was Simia pygmeus of Linneus,

436 THE HON, WALTER ROTHSCHILD ON [ Dec. 13,

1763, and therefore, as the first acceptable generic name is Pongo,

we find that the name for the Orangs as a group is Pongo

pygmeus (Linn.). Now this animal of Linneeus’ s is clearly shown

by the description and Edwards’s figure to have been a form

without cheek- callosities, while we “Gin in Schreber an ape,

entitled Simia agrias, which equally clearly is a young specimen

of a form with cheek-callosities. But of neither of these forms

or phases have we any data which can satisfactorily determine

to what subspecies they belong; therefore the names of these

two phases can only, I think, with any propriety, be applied to

the Landak-Sarawak form, and we arrive at the following

combination :—

form. dimorph. py gmoeus (without callosities).

Pongo pygmvcus, | form. dimorph. agrias (with callosities).

But because this three-legged appellation applies only to one

race of Orang-Outans, it is necessary to give names not only to

each of the three other subspecies as a whole, but also to each

dimorphic phase of each subspecies. Now Dr. Selenka called the

phase with cheek-callosities from the Landak region Pithecus

satyrus landakkensis, while he named the phase without cheek-

callosities P. satyrus tuwakensis; to these, however, I apply the

names of Simia pygmeus of Linnzeus and S. agrias of Schreber, so

we get as the first Bornean subspecies of Pongo pygmeus :—

form. dimorph. agrias (with cheek-callosities).

Pongo pygmeus (Text-fig. 116, p. 437.)

PYygMeus, form. dimerph. pygmeus (without cheek-callo-

sities). (Text-fig. 117, p. 438.)

The phase with cheek-callosities from the Batangtu region was

named Pithecus satyrus batangtuensis by Dr. Selenka, while he

named the phase without cheek-callosities P. satyrus skalauensis ;

of these, one had, however, been described previously under the

name of Simia wurmbii K. Geoft.; therefore the second Bornean

subspecies must stand as follows :—

form. dimorph. wurmbii (with cheek-callosities).

form. dimorph. skalawensis (without cheek-

callosities).

Pongo pygmeus

wurmbia,

Dr. Selenka has called the phase with cheek-callosities from the

Dadap-Genepai region Pithecus satyrus dadappensis, while he

gave the name P. satyrus genepaiensis to the phase without cheek-

callosities from the same region; therefore the third Bornean

subspecies must stand thus :—

form. dimorph. dadappensis (with cheek-callo-

Pongo pygmceus sities).

dadappensis, form. dimorph. genepaiensis (without cheek-

callosities).

The Sumatran Orang with cheek-callosities had the name of

1904. ] ANTHROPOID APES. 437

Pithecus sumatranus deliensis bestowed upon it by Dr. Selenka,

while he called the phase without the callosities P. swmatranus

obangensis. Unfortunately, however, both these forms had had

previous names given to them, viz. Simia abelii by Clarke and

Text-fig. 116.

\ \\

ANS

NOS N

NEN AAS

GD SARS

——

yy)

S==

Skull of Pongo pygmeus forma agrias (Schreber). o. seif ee DIR

Simia bicolor by Isidore Geoffroy respectively ; therefore the fourth

and Sumatran subspecies of Pongo pygmaeus stands as follows :—?

Pongo pygmeus | form. dimorph. abelii (with cheek-callosities).

bicolor, | form. dimorph. bicolor (without cheek-callosities).

After these explanations, I think the following synoptical table

of the divisions and subdivisions of the Orang-Outan, Pongo

438 THE HON. WALTER ROTHSCHILD ON [ects

pygmeus (Linn.), will give succinctly the true nomenclature and

relationship of the different forms.

Text-fig. 117.

ay:

Skull of Pongo pygmeus forma pygmeus (Linn.).

Entire Species.

Poneco pyem#us (Linn.).

Bornean Subspecies.

1. Landak Subspecies.

form. dimorph. agrias (Schreber).

Pongo pygmeus

form. dimorph. pygmceus (Linn.).

pygmeus,

2. Batangtu Subspecies.

Pongo pygmeus { form. dimorph. wurmbii (K. Geoff.).

wurmbii, | form, dimorph. skalawensis (Selenka).

3. Dadap-Genepai Subspecies.

Pongo pygmeus { form. dimorph. dadappensis (Selenka).

dadappensis, | form. dimorph. genepaiensis (Selenka).

1904. | ANTHROPOID APES. 439

Sumatran Subspecies.

Pongo pygmeus { form. dimorph. abelii (Clarke).

bicolor, | form. dimorph. bicolor (1. Geoft.).

IT will now briefly summarise the results arrived at in this

paper. I acknowledge, tentatively, 2 species of Gorilla, one with

3 subspecies; but eventually, with more material available, I think

we shall find only one species, Gorilla gorilla, with 4 or more local

subspecies. I have acknowledged 5 species of Chimpanzee, for

which 1 employ the generic name Simia, as the oldest name given

to a Chimpanzee was Linneus’s Simia satyrus for the Tschego.

T characterise 3 local races of Sima satyrus, 2 of Simia vellerosus,

and 5 of Simia pygmeus, while as yet only one race each of Simid

aubryi and Simia koolookhamba are known tome. Of Orang-Outans

Pongo, | can recognise only one very variable species, which can

be divided up into a number of subspecies. I have characterised

4 such, each with a dimorphic phase, but our knowledge is so im-

perfect that Tonly wish to accept these 3 Bornean and | Sumatran

races for the present, until a fresh lot of material arrives.

Professor Matschie, as a result of his last journey, is preparing

« paper describing a much larger number of forms of Orang and

Chimpanzee than I have dealt with im this paper, dividing them

also into several genera; but, while fully awake to the possibility

of a large number of additional forms existing, I have noticed

here only such forms as are known to me at the time of writing.

In conclusion, | only wish to explain the standpomt I have

taken up in writing this paper. My first contention relates purely

to nomenclature. Hitherto, at least in Great Britain, zoologists

have been divided as to the date to take as the starting-point for

zoological nomenclature : ornithologists and entomologists taking

Linneeus’s X11. edition of the ‘Systema Nature’ of 1766, while

mammalogists take the X. edition of 1758. Also it has been

customary for different zoologists to admit or disallow various

changes in nomenclature. This variety of opinions has led to

much confusion, and I therefore consider, as all writers on

mammals of recent years and also the bulk of German and

American zoologists, that the only way to obtain a uniform and

final nomenclature is to adopt the tenth edition of Linneeus, and

adhere absolutely to the strictest law of priority in nomenclature,

however intrinsically absurd or unsuitable a name may be.

IT now come to my other contention. | Much discussion has taken

and is taking place as to the naming or not of local (¢.e. geographical)

races. The zoologists of the old school maintain that such races

should not be named, and any variation of less than specific value

should be ignored as regards the nomenclatorial point. The

younger generation, however, declare that any distinction, how-

ever slight, ought to be signified by a name so long as it has

geographical foundation. J am of the latter opinion. I am in

favour of this method for many reasons; one of which is, that by

AAO ON ANTHROPOID APES. [ Dec. 13,

distinguishing all local races by a name we prevent the creation

of useless synonyms by forcing the inexperienced student to

study all of these before describing what appears to him a new

species.

As to the method of denoting by names geographical races, there

are many views, but I consider that much the most practical

method is to add a third name to the two already possessed by the

species. This method is no novelty, for it has been done since

the time of Linnzeus, the third name being coupled to the first

two by the term ‘ varietas.” So long as “varietas” was only

used to express a ‘ geographical race” it answered very well, but

soon it was also applied to individual variations and confusion

reigned supreme. I consider, therefore, that it 1s important to

abolish “ varietas” from our nomenclature entirely, as it has so

often been wrongly used, and to substitute the term “ sawbspecies ”

for “geographical races” and the term “ abberatio” for “ indt-

vidual variations.” Thus the South Camaroons Gorilla would be

called

Gorilla gorilla subspecies matschier ;

but this interpolation of the word subspecies makes the name very

long and cumbersome, so that I and most Continental and

American zoologists have agreed to leave out the term ‘ sub-

species” and to write the names of geographical races thus:

Gorilla gorilla matschiei. It is seen, therefore, that this so-called

innovation is no innovation at all, but simply the using of the

long-established formula for local races in an abbreviated and

more convenient form—i. ¢., instead of writing Gorilla gorilla

varietas matschiei, we simply leave out the word “ VARIETAS” or

its equivalent.

The chief reason, however, why I hold that geographical races

ought to be named and diagnosed is that it facilitates so much the

study of geographical distribution. Also the habits of local races

ave often widely different, and it prevents errors if differences in

habits can be correlated to outward differences.

Some zoologists maintain that it is a mistake to describe ‘* seb-

species,” as we cannot tell where individual variation ends and

geographical variation begins.

This holds good only in the case of Reptiles, Fish, Mollusca,

and most probably in the majority of the lower invertebrates ; but

in Insects, Birds, and Mammals it is practically always possible

to tell whether a difference is racial or individual, and, | believe,

even in the previously mentioned groups it will eventually be

possible to define geographical races.

EXPLANATION OF PLATE XXIV.

Simia vellerosus (Gray) (very old male): p. 425.

PZ SS NIOA vol din Pl 2X

J. Green del. et lith. Minvern Bros. imp.

NUNCIRLOCISIDVAIINGS CS SILMUOIVAIES IWAINIO)SIRURILILIS ISLONNL JUN DIAS -

1904. ] ON INDIAN AND BURMESE LAND-SHELLS. 44]

4, Descriptions of Indian and Burmese Land-Shells referred

to the Genera Macrochlamys, Bensonia, Taphrospira,

(gen. nov.), Microcystina, Huplecta, and Polita. By

Wo De BrANrorpy Cb. LED HRs Savile ZnS.

[Received October 31, 1904. ]

(Plate XXV.*)

The following pages contain diagnoses of several Indian land-

shells that are believed to be undescribed. The majority belong -

to the genus Macrochlamys, and have been met with in the course

of an attempt at monographing the forms found in the Empire of

British India. Only those kinds have been described of which

the localities are believed to be accurately ascertained, which are

sufficiently distinct to render it probable that they may be recog-

nised from their description, and of which type specimens are

available for deposit in the British Museum.

Some of the specimens are from my own collections made in India

and Burma, others are from the British Museum accumulations,

whilst for others [am indebted to Colonel Godwin- Austen, Colonel

Beddome, and Mr. Hugh Fulton. I have to acknowledge the

valuable assistance and advice of Mr. KH. A. Smith in my examina-

tion of the British Museum specimens.

BENSONIA NEPALENSIS, Nevill MS. (Plate XXYV. fig. 1.)

Testa aperte perforata, conoideo-depressa, sublenticularis, tenwis,

superne oblique et rugose plicata, subtus nitidior atque striatula,

haud decussata ; spira depresso-conoidea, sutura vix impressa ;

anfr.6, conveat, regulariter accrescentes, ultimus non descendens,

ad peripheriam carinatus, subtus twmidus ; carind versus

aperturam minus acuta ; apertura obliqua, rotundato-lunata,

marginibus convergentibus ; peristoma tenue, margine colu-

mellarit curvato, denique vertical, wndique expansulo, juxta

perforationem late refleco, Diam. maj. 23:5, min, 21, alt.

12 mm.

Hab. Khatmandu, Nepal.

Near B. camura Bs., but distinguished by a lower spire, smaller

umbilicus, stronger sculpture, the absence of decussating striation

and by less acute carination.

The types are in Col. Godwin-Austen’s collection now in the

British Museum.

TAPHROSPIRA }, gen. nov.

Testa depressa vel globoso-depressa, tenuis, cornea, ab ila Macro-

chlamydis Bs., fossd spirali extra suturam in omnibus anfract-

ibus tantum diversa.

Typus 7’. convallata, Bs.

** For explanation of the Plate, see p. 447.

+ Tappos, a ditch or trench; o7eipa, spiral .

28*

442,

DR. W. T. BLANFORD ON INDIAN [ Dec. 13,

Animal not known.

So far as is known, the following species should be referred to

this genus :—

T. convallata Bens. Tenasserim.

T. bathycharax * Bens. MS. Andaman Islands.

T. compluvialis Bf. Arakan Hills, W. side.

T. excavata, sp.nov. Hills south of Assam.

TAPHROSPIRA EXCAVATA, Sp. nov. (Plate XXV. fig. 3.)

Helix compluvialis, Hanley & Theob. Conch. Ind, pl. 88. figs. 1,

4 (1874), nec Nanina compluvialis BIf. (1865).

Testa perforata, globoso-depressa, tenuis, Jusco-straminea, cornea,

diaphana, nitida, minute transversim atque subobsolete sub

lente longitudinaliter striata ; spira parum easerta, aprce

obtuso, fossa suturali lata sed parwm profunda ; anfr. 5,

superne extra fossam suturalem acute angulati, ultimus ad

peripheriam rotundatus, subtus inflatus, versus aperturam

vie descendens ; apertura parum obliqua, rotundato-lunaris,

superne emarginata, fere eque lata ac alta ; peristoma tenue,

vie sinuatum, margine columellari superne verticali, breviter

triangulatim refleco. Diam. maj. 15, min. 13, alt. 9 mm.

Hab. ad Asalu in provincia Cachar septentrionali (Godwin-

Austen), necnon in montibus Khasi dictis teste Nevill.

This species resembles 7’. compluvialis, for which it has been

mistaken, but which is a much less globose form and smaller, with

a smaller, more oval and more oblique mouth, and closer and more

distinct longitudinal striation under the microscope. 7’. complu-

vialis is figured for comparison, Pl. XX V. fig. 4.

MacrocHLAMYSs KULUENSIS Nevill MS. (Plate XXYV. fig. 5.)

Testa aperte perforata, subumbilicata, subgloboso-depressa, tenuis,

translucens, nitida, pallide cornea; spira conoidea, sutura

impressa ; anfr. 54, conveat, ultimus valde major, ad peri-

pheriam rotundatus, subtus tumidus ; apertura obliqua, rotun-

dato-lunata, subceque lata ac alta; peristoma tenue, rectum,

margine columellari verticali, triangulatum refleco. Diam,

maj. 12, min. 10°5, alt. 7 mm.

Hab. in pago Himalayano Kulu (coll. 6.M,).

Near Jf. glauca and WM. nuda, but distinguished by much more

open perforation and rounder mouth.

MACROCHLAMYS SUPERFLUA, sp. nov. (Plate XXYV. fig. 7.)

Testu perforata, depressa, tenuis, polita, translucens, via striata,

sub lente lineis impressis subconfertis spiralibus, decussatula,

luteo- vel fusco-cornea ; spira parum elevata, sutura bene

impressa ; anfr. 6, convear, ultimus valde latior, ad peripheriam

rotundatus, subtus convexus ; apertura obliqua, subovate

lunata, latior quam alta; peristoma tenue, margine basali

* Described by Mr. Hugh Fulton, Journ, Mal. x. p. 99 (1908).

1904. | AND BURMESE LAND-SHELLS. 443

recto, columellari superne verticali aique sublate reflexo, tunc

oblique curvato. Diam. maj. 30, min. 17-5, alt. 10 mm.

Hab. in valle fluminis Tista, in provincia Sikhim interiore

Himalayana satis frequens (W. ?. B.).

This is a much larger shell than J/, sequaa with a different

sculpture. The spaces between the longitudinal impressed lines

appear papillose when considerably magnified.

MacrocuLamys (?) atomA Fairbank MS. (Plate X XV. fig. 6.)

Testa aperte perforata, depressa, discoidea, solidula, nitida,

vitrea, lineis impressis parallelis subdistantibus spiralibus vel

concentricis undique sub lente striata, albido-cornea; spira

Sere plana, sutra impressa ; anfr. 33, regulariter accrescentes,

superne convert, ultimus non descendens, ad peripheriam

rotundatus, subtus plano-convexus ; apertura parum obliqua,

lunata ; peristoma tenue, marginibus supero basalique leviter

arcuatis, columellart fere horizontah, haud reflexwo. Diam.

maj. 1:5, min. 1:2, alt. 0°6 mum.

Hab. prope ripas fluminis Godavari (W. 7. B.).

This minute shell, found commonly amongst the debris left

behind by river-floods, was named in MS. by the late Rev. S. B.

Fairbank 40 years ago, and was mentioned in a report of mine

written in 1866 and published in my absence in the Records of the

Geological Survey of India, vol. i. 1870, p. 62. The typical

specimens are from Paitan near Ahmednagar, and I have spe-

cimens from various places as far to the south-east as the first

barrier on the Godavari at Dumagudem, also from the Wardha near

Nagpur, and I believe I had some from the Nerbudda Valley.

This is probably not a Macrochlamys but the animal is unknown,

only dead shells having been found. The only allied Indian form

appears to be WZ. anone G.-A., from Calcutta, a much less depressed

shell.

MACROUHLAMYS PRAVA, Sp. hov. (Plate XXYV., fig. 9.)

Testa perforata, subgloboso-depressa, tenuis, nitida, polita, quasi-

obsolete sub lente spiraliter striata, rufescenti-fusca ; spira via

elevata, conoidea, sutura parum impressa ; anfr. 5, convexi,

ultimus latior, ad peripheriam rotundatus, subtus twmidius-

culus; apertura fere verticalis, subovato-lunata ; peristoma

tenue, rectum, margine columellari superne fere verticali ibidem

reflexiusculo. Diam. maj. 10°5, min. 9, alt. 5°5 mm.

Hab. ad urbem Beypur prope litus Malabaricum (Fairbank) :

etiam ad latus occidentale montium Nilgiri (W. 7. B.), et Anaimalai

(Beddome), preterea in provincia Travancore (Day).

common and rather widely spread form, resembling the

Arakan species JZ. kuwmahensis Theob. & Stol. in form.

MACROCHLAMYS RUTILA, Sp. nov. (Plate XXV. fig. 11.)

Testa perforata, depressa, tenuis, vitrea, polita, dense minuteque

regulariter sub lente et supra et subtus longitudinaliter

444

DR. W. T. BLANFORD ON INDIAN [ Dec. 13,

(spiraliter) striata, castanea ; spira humilis, sutura leviter

impressa ; antr. 6, convexi, ultimus latior, ad peripheriam

rotundatus, subtus convexus ; apertura parum obliqua, lunata ;

peristoma tenue, margine superiore via arcuato, basali subrecto,

columellari obliquo, breviter triangulatim refleco. Dian. maj.

vie 15, min. 13, alt. 6 mm.

Hab. in dumeto ‘ Anagundi shola’ dicto montium Anaimalai

(Leddome).

MACROCHLAMYS CHAOS, sp. nov. (Plate XXV. fig. 8.)

Testa perforata, conoideo-depressa, subglobosa, tenwis, natida,

vitrea, transversim striatula, sub lente lineis minutis crebris

flexuosis spiralibus subobsolete sculpta, pallide fulva ; spira

parum elevata, apice acuto, sutura impressa ; anfr. 54, con-

veaiusculi, ultinvus latior, ad peripheriam rotundatus, subtus

converus ; apertura obliqua, rotundo-lunata, latior quam alta ;

peristoma pertenue, rectum, margine columellari curvato, superne

verticalt et breviter triangulatim refleco. Diam. maj. 16,

min. 14, alt. 8 mun.

Hab. in Burma, ad Thayet Myo atque haud procul ab Ava

(IV.

T. B.).

Very near the Bengal J/. swbhjecta Bs., but distinguished by the

presence of longitudinal sculpture. It is also more vitreous and

less

globose, with a narrower last whorl and smaller mouth.

MACROCHLAMYS NOTHA, sp. nov. (Plate XXV. fig. 19.)

Nanina petasus Bf. Jour. As. Soc. Beng. 1865, pt. 2, p. 86, nec

Benson.

Testa perforata, depressa, parum polita, via striatula, lineis con-

centricis confertis eax tuberculis minutissimis constantibus sub

lente undique ornata, flavescenti-fulva ; spira breviter conordea,

apice subacuto, sutura impressa; anfr. 6-7, convexiusculi,

lente accrescentes, ultimus ad peripheriam rotundatus, subtus

convecus ; apertura via obliqua, fere verticalis, lunata ; peri-

stoma tenue, margine basali arcwato, columellari expanso,

oblique curvato, denique ad perforationem verticali. Diam.

maj. 11:5, min. 10°5, alt. 6 mm.

Hab. in. montibus Aracanensibus inter Prome et Tongoop ;

necnon prope Thayet Myo, in provincia Pegu Burmannica.

Very close to J/. petasus Bs., but distinguished by want of

labiation and presence of longitudinal sculpture.

MACROCHLAMYS NOXIA, sp. nov. (Plate XXV. fig. 14.)

Testa minute et subobtecte perforata, depressa, tens, nitida,

polita, haud usquam striata, pallide castanea, subtus, nist juxta

peripheriam, albescens ; spira via elevata, conoidea, sutwra

impressa ; anfr. 6, convext, haud celeriter crescentes, ultumus

ad peripheriam rotundatus, subtus convexus ; apertura parum

obliqua, late lunata ; peristoma tenue, margine basali arcuato,

1904. ] AND BURMESE LAND-SHELLS. 445

columellart obliquo, superne breviter refexo. Diam. maj. 9,

min. 8, alt. 4 mm.

Hab. ad latus occidentale fluminis Irawadi in pago Bassein

provincie Pegu Burmannice (IW. 7. B.).

Varietas, Fare 5 munita, et spira aliquanto elevatiuscula

prope portum Akyab in provincia Arakan invenitur.

This is near J. hypoleuca Bf. from Upper Pegu, best dis-

tinguished by being narrowly perforate and more depressed, and

by the complete want of sculpture.

MACROCHLAMYS CURVILABRIS, sp: nov. (Plate XXYV. fig. 13.)

Testa perforata, depressa, glabra, polita, via striatula, fusca ;

spira parum elevata, sutura bene impressa ; anfr. 53, convexi,

regulariter crescentes, ultimus versus aperturam anecnuluan

descendens, ad peripheriam rotundatus, subtus plano-convenus ;

apertura obliqua, late lunata ; peristoma obtusum, intus albo-

labiatum, simwosum, margine supero vic arcuato, externo

sinuato, basali valde arcuato, columellari obliquo, vix relleaius-

culo. Diam. maj. 5°5, min. 5, alt. 2 mm.

Hab. in montibus Aracanensibus ad latus occidentale pagi

Burmannici Prome (W. 7’. B.).

This is almost a miniature of the Tenasserim J/. aspides Bs.

MACROCHLAMYS SPRETA, sp. nov. (Plate XXYV. fig. 12.)

Testa minute et subobtecte perforata, depressa, tems, nitida,

polita, lineis impressis spiralibus sub lente undique ornata,

pallide castanea, subtus circa perforationem albescens ; spira

parum elevata, conoidalis, sutura via impressa ; anfr.d, convexi,

ultimus majusculus, ad peripheriam rotundatus, subtus con-

vexus ; apertura obliqua, subovato-lunaris ; peristoma tenue,

rectum, margine columellari obliquo, leviter refleco. Diam.

maj. 8, min. 7, alt. 3° mm.

Hab. ad Thamandewa in pago Bassein et in aliis partibus

provincie Pegu Burmannice (W. 7’. B.).

Near I. subpetasus Nevill and J. noxia, but easily recognised

by the spiral striation.

MAcROCHLAMYS PATENS, sp. nov. (Plate XXV. fig. 15.)

Testa anguste sed perspective wmbilicata, conoideo-depressa, sub-

lenticularis, nitida, polita, lineis inpressis spiralibus haud

crebris spatus incequalibus discretis undique sub lente onli,

fulwo-fusca ; spira conoidea, sutura parum impressa ; anfr. 43

convexiusculi, ultimus paulo latior, ad peripheriam obtuse

angulatus, subtus convexus, circum umbilicum compressus ;

apertura diagonalis, fere trapezoidalis, subsecuriformis ; peri-

stoma tenue, rectum, margine columellari obliquo, triangulatim

reflexo. Diam. maj. 7, min. 6, alt. 3°5 mm.

Hab. in Pegu; circa Thayet Myo, Bassein, &c. (W. 7. B.).

446 DR. W. T. BLANFORD ON INDIAN [ Dec. 13,

MACROCHLAMYS PSEUDOCHOINIX, sp. nov. (Plate XXV. fig. 10.)

Testa subobtecte perforata, depressa, tenuis, glabra vix politula,

subobsolete plicato-striata et lineis minutis confertis sub-

flexuosis spiralibus undique decussatula, fusco-cornea ; spira

fere plana, sutura viz impressa ; anfr. 5, planiusculi, ultumus

valde latior, ad peripheriam rotundatus, subtus twmidus ;

apertura obliqua, subdiagonalis, magia, subovato-lunata ;

peristoma acutum, margine supero arcuato, columellari juxta

perforationem vertical, breviter reflexo, perforationem partum

tegente. Diam. maj. 14, min. 12, alt. 7 min.

Hab. in insula ‘Great Cocos’ dicta in sinu Bengalensi.

Near the Andaman J. choinix, but more tumid beneath, less

flat above, and with much stronger spiral sculpture. For this

shell I am indebted to Mr. Hugh Fulton. The Great Cocos is one

of a group of islands between the Andamans and Cape Negrais in

Arrakan.

MicrocystIna sTUARTI Godwin-Austen MS. (Plate XXV.

fig. 16.)

Testa imperforata, vel subperforata, convexo-depressa, pertenris,

nitida, politissima, minute, haud crebre, sub lente undique

lineis parallelis striata, luteo-fusca; spira parum elevata,

convexo-conoiden, sutura vic impressa; anfr. fere 5, plano-

convext, ultimus non descendens, ad peripheriam rotundatus,

subtus convexus ; apertura obliqua, lunata ; peristoma tenue,

margine columellari obliquo, superne in angulum prominentem,

perforationem claudentem, dilatato. Diam. maj. 4:5, min. 4,

alt. 2°35 mm.

Hab. in insulis Andamanicis (coll. G.-A.).

Near the Nicobar WM. rinki Morch, but smaller, rather thinner,

more closely wound and imperforate,

MICROCYSTINA SHEVAROYANA, sp. nov. (Plate XXYV. fig. 17.)

Testa aperte perforata, subwmbilicata, conoideo-depressa, tennis,

translucens, polita, dense minute et subflexcuose spiraliter

undique sub lente striata, succinea; spira parum elevata,

conoidea, sutura impressa ; anfr. 6, superne conveai, ultimus

ad peripheriam rotundatus, subtus convexiusculus ; apertura

parum obliqua, lunata; peristoma tenue, marginibus supero

basalique leviter arcuatis, columellari obliquo, obtuso, expansi-

usculo atque superne reflexiusculo, yuata perforationem angu-

latim dilatato. Diam. maj. 8°25, min. 7:5, alt. 4 mm.

Hab. in montibus Shevaroy dictis, Indiz meridionalis (W. J.

Daly).

Near the Ceylon I. bintennensis (M. perfucata var. bintennensis

G.-A.), but distinguished by more open perforation, more numerous

whorls, and paler colour. I am indebted to Mr. Hugh Fulton

for specimens of this shell.

1904. ] AND BURMESE LAND-SHELLS. 447

EUPLECTA PULCHELLA, sp. nov. (Plate XXYV. fig. 18.)

Testa subobtecte perforata, conoideo-depressa, fere lenticularis,

solidiuscula, succineo-cornea, decussatim striatula, superne

sub lente liris obliquis flexuosis graniferis et lineis spiralibus

impressis pulchre ornata, subtus glabra, polita; spiraconoidea,

apice obtuso, sutwra impressa ; anfr. 5, conveai, ultimus versus

aperturam aliquanto descendens, ad peripheriam obtuse angu-

latus, infra convexus, circa perforationem compressus ; apertura

magna, diagonalis, rotundo-lunata ; peristoma tenue, superne

juata anfractum penultimum arcuatum, margine columellari

obliquo, expansiusculo, ad perforationem ldtius reflexo et

incrassato. Diam. maj. 11:5, min. 10, alt. 6°5 mm.

Hab. in montibus Anaimalai dictis Indiz australis (Beddome).

This shell, whilst in some respects recalling that of 4. layardi,

is distinguished by its large diagonal mouth and granular sculp-

ture in flexuous transverse lines.

Pourra (?) TURBINATA, sp. nov. (Plate XXYV., fig, 2.)

Testa aperte umbilicata, depressa, tenuis, pallide cornea, nitida,

polita, sub lente minutissime spiraliter striata ; spira convexa ;

anfr. 43, convexi, ultimus undigque rotundatus fere teres ;

apertura obliqua, rotundo-lunata, marginibus convergentibus ;

peristoma tenue, margine superiore arcuato, columellari vix

superne verticali, regulariter curvato, haud refleco. Diam.

maj. 4°5, min. 4, alt. 2 mm.

Hab. in sammis montibus Nilgiri dictis Indie meridionalis ad

alt. 7000 ped. (Leddome).

us is a very near ally of the Ceylonese Polita ? notabilis Sykes.

It is smaller and the aperture is more oblique.

co Ont OO em ob

EXPLANATION OF PLATE XXV.

. Bensonia nepalensis, nat. size, p. 441.

. Polita(?) turbinata, nat. size and X 3, p. 447.

. Taphrospira excavata, nat. size, p. 442.

. T. compluwialis, nat. size, p. 442.

. Macrochlamys kuluensis, nat. size, p. 442.

- M.(?) atoma, nat. size, and two views X 15, also sculpture further enlarged,

p- 443.

. M. superflua, nat. size, p. 442.

. WM. chaos, nat. size, p. 444.

. WM. prava, nat. size and X 2, p. 443.

. WM. pseudochoinix, nat. size, two views, p. 446,

. M. rutila, nat. size, p. 443.

. MW. spreta, nat. size and x 8, p. 445.

. M. curvilabris, nat. size, and two views, X 3, p. 445.

. WM. nowia, nat. size and X 2, p. 444.

. M. patens, nat. size, and two views X 8, p. 445.

. Microcystina stuarti, nat, size and X 3, also basal and columellar margins

of peristome further enlarged, p. 44.6.

. Microcystina shevaroyana, nat. size, p. 446.

. Huplecta pulchella, nat. size, and sculpture much enlarged, p. 447.

. Macrochlamys notha, nat. size, p. 444A,

448 DR. W. G. RIDEWOOD ON THE CRANIAL [Dec. 13,

5. On the Cranial Osteology of the Clupeoid Fishes. By

W.G. Ripewoop, D.Sc., F.Z.S8., Lecturer on Biology

at St. Mary’s Hospital Medical School, London.

[Received November 11, 1904. |

(Text-figures 118-143.)

INTRODUCTION.

In the spring of 1896, Prof. G. B. Howes suggested to me

that an investigation on the structure of the skull in the lower

Teleostean fishes would be a profitable piece of research, since

there was every prospect of the results proving a valuable means

of testing the validity of the existing schemes of classification of

the fishes in question, and because an accurate knowledge of the

structure of the skull in the lower Teleostean fishes was essential

to a successful study of the remains of those extremely interesting

extinct fishes which lie on the boundary-line between the 'Tele-

osteans and the Ganoids.

The investigation has proceeded slowly and intermittently,

owing to repeated interruptions and to pressure of other work ;

but sufficient progress has now been made to allow of the publica-

tion of some of the results. Descriptions of the skulls of Hops,

Megalops, and Albula, together with some general observations on

the Teleostean skull, have already appeared in the ‘ Proceedings

of the Zoological Society,’ 1904, 11. pp. 35-81, and observations

on the cranial osteology of the Mormyride, Notopteridz, and

Hyodontide in the Journal of the Linnean Society, xxix. 1904,

pp. 188-217. A third paper, on the skull of the Osteoglosside,

Pantodontide, and Phractolemidz, has just been completed, and

has been offered to the Linnean Society; the present contribution

deals with the skull of the Clupeoid fishes.

Eleven genera are considered in this paper, namely :—Chiro-

centrus, Clupea, Pellona, Pellonula, Pristigaster, Hyperlophus,

Chatoéssus, Dussumieria, Hngraulis, Coilia, and Chanos. A

“Summary ” of the observations and some ‘“‘ Comments ” thereon

are given on pp. 488-493.

Skulls of Chirocentrus, Clupea, Chatoéssus, Hngraulis, Coilia,

and Chanos were specially prepared for the purposes of the

investigation, and were disarticulated according to the method

explained in the ‘ Proceedings of the Zoological Society,’ 1904, 11.

p. 36; the other skulls examined are the property of the British

Museum, and I take this opportunity of acknowledging my

indebtedness to Mr. G. A. Boulenger for offering to me every

facility in his power for the examination of the skulls under

his charge.

CHIROCENTRUS DORAB.

The only published figure of the skull of Chirocentrus that

I have been able to discover is a not very serviceable sketch of

1904, | OSTEOLOGY OF CLUPEOID FISHES, 449

the hinder part of the cranium, seen from above, given by Klein

(Jahresh. Ver, vaterl. Naturk. Wiirtt. xli, 1885, pl. 3. fig. 82).

Material examined—In addition to a skull (A) specially

prepared for the purposes of this investigation from an alcohol-

preserved specimen kindly furnished by Prof. Howes, two skulls

were examined, one (B) belonging to a complete skeleton, marked

89.2.1.2059, in the Osteological Collection of the British Museum,

and another (C) bearing no distinctive number.

Text-fig, 118.

pro af bo

Cranium of Chirocentrus dorab. A, dorsal view; B, left side; C, back view.

For explanation of lettering see p. 493.

Cranium (text-fig. 118).—The parietals are separated by the

supraoccipital, and there is a small median fontanelle, divided

Proc. Zoou. Soc.—1904, Von. Il. No. XXIX. 29

450 DR. W. G. RIDEWOOD ON THE CRANIAL | Dee. ay

longitudinally by a bar of cartilage, between the supraoccipital

and the frontal bones (wanting, however, in specimen B), and

another fontanelle between the anterior ends of the frontals.

The posterior temporal groove is deep, but is not roofed over.

The groove narrows away to a point anteriorly, and situated near

its termination is a temporal foramen leading into the upper part

of the cranial cavity. Behind the temporal foramen is a deep

pre-epiotic fossa, bounded above and in front by the parietal,

below by the squamosal, and behind by the epiotic, and ter-

minating blindly against the deeper part of the supraoccipital.

The lateral temporal groove above the postfrontal is wide and

shallow.

The opisthotic is very small, and abuts on the squamosal,

exoccipital, and pro-otic, but does not touch the epiotic. The

pro-otie is extensive, and nearly reaches the back of the cranium.

There is a small bulla, containing a vesicle of the swim-bladder,

situated a little above the centre of the latero-ventral face of the

pro-otic, and another bulla occurs in the squamosal and projects

slightly above the floor of the posterior temporal groove. An

auditory fenestra, bounded by the pro-otic, exoccipital, and basi-

occipital, is present in the side of the cranium.

At the front of the eye-muscle canal is situated a small basi-

sphenoid, bounded laterally by processes of the alisphenoids which

descend to meet the pro-otics. There is no descending part of the

basisphenoid to bisect the eye-muscle canal. The parasphenoid

projects a little behind the basioccipital, and has right and left

posterior wings which bound the posterior outlet of the eye-

muscle canal. At its anterior end, where it meets the vomer,

the parasphenoid becomes considerably broader than in its

middle part. The vomer projects in front of the mesethmoid, and

neither it nor the parasphenoid bears teeth. The ethmoid region

is comparatively short and well ossified, and there is no cartila-

ginous tract intervening between the mesethmoid and prefrontal.

The two prefrontals are closely united in the median plane of

the head.

Temporal and Preopercular Series * (text-fig. 119, p. 451).—The

post-temporal is comparatively large and has three limbs. The

upper one rests over the epiotic; the opisthotic limb is long, rod-

like, and very slender ; and the third, which is pointed and shorter

than the other two, passes forward to touch the back of the

supratemporal, The supratemporal is roughly trivadiate, and its

antero-dorsal ray or limb is the largest ; it forms a lateral wall to

the unenclosed posterior temporal groove. The upper and lower

limbs of the preopercular are set at a very wide angle. The

interopercular has at its anterior end an ascending process which

flanks the inner surface of the lower part of the symplectic.

* The reasons for including the preopercular and interopercular bones in this

series, and for excluding them from the opercular and branchiostegal series, are

given in a former paper (Proc. Zool. Soc. 1904, ii. pp. 68 and 75). For reasons given

in the same paper, it is considered expedient to regard the post-temporal as a con-

stituent of the skull.

1904. | OS 2EOLOGY OF CLUPEOID FISHES. ADI

Circumorbital Series (text-fig. 119).—The total number of

circumorbital bones on each side is nine; the largest is that

which lies antero-ventrally to the eye. The nasal is very small.

Manillary Series (text-fig. 119).—Both premaxilla and maxilla

bound the gape above. They both bear long, pointed teeth, and

the anterior tooth of the premaxillary series is much larger than

the others. As already noticed by Valenciennes (Hist. Nat.

Poiss, xix. 1846, pp. 150, 152, and 154), the premaxilla is firmly

attached to the maxilla in Chirocentrus, whereas in Clupea it is

readily movable upon the maxilla. The two maxille meet one

another in front of the mesethmoid, behind and above the

premaxillary symphysis. There are two surmaxille.

Text-fig, 119.

Chirocentrus dorab, vight side of skull. For explanation of lettering see p. 493.

Mandibular Series (text-figs. 119 and 120, p. 452).—The dentary

bears eight or ten large, curved, pointed teeth, four or five of which

are firmly anchylosed to the bone*. The angular is distinct, and

there is a very small sesamoid articular; the endosteal articular

is not distinct from the ectosteal articular.

Hyopalatine Series (text-fig. 120, p. 452).—The hyopalatine arch

is short and deep. The hyomandibular has a single broad head for

articulation with the cranium. ‘The metapterygoid extends high

up the outer face of the hyomandibular, and the symplectic is

small and lies nearly in a line with the axis of the hyomandibulayr.

The symplectic, when viewed from ths buccal aspect, is largely

(not entirely) hidden by a downgrowth of the hyomandibulay,

* Smith Woodward (Brit. Mus. Cat. Foss. Fishes, iv. p. 88) says of Chirocentrus :

“Teeth .... . firmly fixed in shallow sockets.’ He mentions, however, but one

extinct species of Chirocentrus (C. polyodon), and states that it is “doubtfully of

this genus,”

29*

452 DR. W. G. RIDEWOOD ON THE CRANIAL [ Dec. 13,

and in an external view is barely visible between the quadrate

and the preopercular. This concealment of the symplectic is

utilised as a family character by Boulenger (Ann. Mag. Nat.

Hist. (7) xiii. 1904, p. 164). The palatine is short and com-

pletely ossified, and has two distinct articular heads, one for the

mesethmoid and vomer, and one for the prefrontal; its ventro-

external surface enters into extensive synovial articulation with

the maxilla. There are four or five minute teeth on the palatine,

but none on the pterygoid bones.

Text-fig. 120.

Sop ae

EU gaya VB? Up

Chirocentrus dorab, hyopalatine arch, opercular bones, and mandible, left side,

mesial aspect. For explanation of lettering see p. 493.

Opercular Series (text-figs. 119 and 120).—The opercular bone

is of normal proportions; the subopercular is of small vertical

extent, and has at its anterior end a strongly developed process

rising against the anterior edge of the opercular. The branchio-

stegal rays are eight in number. The front four are shaped like

the blade of a scythe, and there is a fairly regular transition in a

backward direction to the larger and lamellate posterior members

of the series. The first five are attached to the outer surface of

the ceratohyal, and the other three to the outer surface of the

epihyal.

Hyobranchial Series (text-fig. 121, p. 453).—The interhyal is

ossified. The upper hypohyal is considerably smaller than the lower.

The glossohyal is small and bears a few minute teeth on its supero-

lateral edges. The first basibranchial is short, the second and

third are exceptionally long. A long dentigerous plate overlaps

the three basibranchial bones, and a similar plate, of large size,

but readily removable, and not shown in the figure, overlies the

common cartilage of the fourth and fifth basibranchials. The

dorsal parts of the branchial skeleton appear short in proportion

to the ventral, but the disparity is probably to be accounted for

1904. | OSTEOLOGY OF CLUPEOID FISHES. 453

by the abnormal length of the latter. The first pharyngo-

branchial is conical, fully ossified, and in a line with the first

epibranchial; there is no spicular bone. The second pharyngo-

branchial is triangular, and about 24 or 3 times as long as broad.

The third pharyngobranchial is long ; its anterior part is slender

and rod-like, and runs along the mesial edge of the second

pharyngobranchial.

Text-fig. 121.

hb?

Chirocentrus dorab, hyobranchial skeleton, dorsal view. The epibranchials and

pharyngobranchials of the right side are not shown. For explanation of

lettering see p. 493.

CLUPEA FINTA, etc.

In 1820 Weber (De Aure et Auditu Hominis et Animalium,

pl. 8. figs. 64-66) gave three views of the cranium of the Herring,

and a remarkably accurate description of the osseous bulle that

enclose the dilatations of the anterior end of the swim-bladder.

Rosenthal’s figures (Ichthyotomische Tafeln, Aufl. 2, Berlin,

1839, pl. 4) are moderately accurate, but of no particular value ;

and Briihl’s figures (Vergl. Anat. aller Thierklassen, Abschn. 1,

Skelettlehre der Fische, 1847, pl. 5. fig. 32 and pl. 10) are copied

from those of Weber and Rosenthal. The figure of the skull of

the Herring in the ‘ Histoire Naturelle des Poissons’ of Cuvier

and Valenciennes (pl. 593) is hardly worth mentioning. The

side view of the complete skeleton of the Alose given by Agassiz

in his ‘Recherches sur les Poissons Fossiles* (Atlas, v. pl. L)

is good, but only the superficial bones are seen, and none of

them are named.

454 DR. W. G. RIDEWOOD ON THE CRANIAL [ Dec. Mae

Hyrtl, in his paper on the Accessory Branchial Organ of

Clupeoid Fishes (Denkschr. Akad. Wiss. Wien, x. 1855, pl. 1.

figs. 2 and 3) has published figures of the branchial skeleton of

Clupea thryssa (Meletta thryssa) and Clupea mattowocca (Alausa

tyrannus), but they are of little service in the present connection.

Vrolik (Niederl. Arch. Zool. i. 3, 18738, pp. 268-270, and figs. 28

and 29) has given a short description and two figures of the

cranium of the Herring; and Matthews has contributed a very

complete account of the whole skull of that fish, with observa-

tions also on the skulls of Clupea finta, Clupea pilchardus, and

Olupea sprattus (Fifth Rep. Fish. Bd. Scot. 1887, pp. 274-292,

and figs. 15, 17, and 18). Three good figures of the cranium of

the Herring appear in Fries’ ‘Scandinavian Fishes” (ed. 2,

by F. Smitt, vol. ii. 1895, p. 949).

Material examined.—The description below applies mainly to.

Clupea finta, but Clupea harengus, Clupea sapidissima, and Clupea

sprattus were also examined. The hyobranchial skeleton de-

scribed is that of Clupea harengus. All the skulls were specially

prepared for the purposes of the investigation.

Cranium (text-fig. 122, p. 455).—The parietals are separated by

the supraoccipital. The posterior temporal groove (mastoid groove

of Fries, /.c.) is not roofed. Near its anterior end is an oval

temporal foramen (Fries, /.c. p. 947), bounded by the parietal

and frontal, and occupied by a fatty mass in the’ fresh fish, but

leading directly into the cavum cranii in the dried skull; behind

it is the pre-epiotic fossa, bounded by the parietal, squamosal, and

epiotic, and ending blindly against the supraoccipital. The pre-

epiotic fossa is relatively larger in Clupea harengus and Clupea

sprattus than in Clupea finta and Clupea sapidissima.

In the middle of the pro-otic bone is a conspicuous bulla con-

caining a spherical vesicle of the swim-bladder. A second bulla

is present in the squamosal, but this is not visible in Clupea finta

axcept by breaking open the bone; in Clupea harengus it is just

visible on the surface. Clupea sprattus has no squamosal bulla

(Matthews, .¢., and Ridewood, Journ. Anat. and Phys. xxvi.

1891, p. 36, and fig. E, p. 32). In Clupea harengus the duct

that leads through the exoccipital bone from the pro-otic and

squamosal vesicles to the swim-bladder is dilated and fusiform

in shape. The auditory fenestra, in the ventro-lateral aspect of

the cranium, has an irregular outline; it is bounded by the

pro-otic, exoccipital, and basioccipital, and leads into the peri-

lymphatic cavity of the ear.

The cpisthotic is moderately small; it touches the pro-otic,

squamosal, and exoccipital bones, but not. the epiotic. The

basisphenoid is small and transversely set, and its descending

process is a mere spicule of bone which fails to reach the para-

sphenoid. The orbitosphenoid sends forward a process which

meets a backwardly directed process of the united prefrontals.

The parasphenoid is straight, or nearly so, in Clupea finta, but

the middle part is slightly depressed in the other species examined.

1904. ] OSTEOLOGY OF CLUPEOID FISHES. 455

The posterior wings, on the right and left sides of the posterior

opening of the eye-muscle canal, are thin and delicate. They

become separate from one another beneath the middle part of the

pro-otic (they separate in front of the pro-otic in Clupea harengus),

Text-fig. 122.

vor

pro

Cranium of Clupea finta. A, dorsal view; B, left side; C, back view.

E For explanation of lettering see p. 493.

(ey

and extend a considerable distance behind the occiput. There

are no teeth on the parasphenoid nor on the vomer in Clupea

finta, but vomerine teeth are present in Clupea harengus and

456 DR. W. G. RIDEWOOD ON THE CRANIAL | Dee. 13,

Clupea spratius. There is a median fontanelle between the

mesethmoid and the frontal bones,

Temporal and Preopercular Series (text-fig. 123),—The post-

temporal is rather large and slender. It has two long limbs

attached to the epiotic and opisthotic respectively ; the third

limb, that which carries the sensory canal, is very short, and

touches the back of the supratemporal. The supratemporal is

elongated horizontally; its sensory canal, entering behind from

the post-temporal, divides, as usual, into one branch going

upward to the parietal and another which passes forward into

the squamosal. The axis of the upright limb of the preopercular

is more oblique in Clupea harengus than in Clupea jfinta, and the

interopercular bone and the lower limb of the preopercular are

relatively longer.

Circumorbital Series (text-fig. 123)—There are eight bones of

this series—a small nasal, and seven bones disposed around the

eye. The orbital ring is incomplete above.

Text-fig. 123.

pop top gq eH

Clupea jinta, right side of skull. For explanation of lettering see p. 493.

| fiaxillary Series (text-fig. 123)—The gape is bounded above

by the maxilla and premaxilla, both of which bear a row of

minute teeth along the lower edge. In Clupea harengus there

are rarely more than three teeth in each premaxilla, but in

Clupea jinta the number is between twenty and thirty. Two

surmaxillary bones are present on each side.

Mandibular Series (text-figs. 123 and 124, p.457).—The mandible

is rather high in proportion to its length, particularly so in Clupea

harengus. In Clupea finta the highest point of the mandible lies

1904. | OSTEOLOGY OF CLUPEOID FISHES. 457

over the hinder half of the ramus ; in Clupea harengus and Clupea

sprattus it lies over the front half. The mandibular symphysis is

more in advance of the premaxille in the two latter species than

in the first. The angular bone is distinct. There is a small

sesamoid articular in Clupea finta, but not in Clupea harengus.

There are about five teeth situated in a row at the anterior end

of the dentary in both species.

Hyopalatine Series (text-fig. 124).—As has been pointed out

by Matthews (/.c. p. 289), the hyomandibular of Clupea finta

articulates with the cranium by two distinct heads, whereas in

Clupea harengus the hyomandibular has a single broad head.

On comparing Clupea finta and Clupea harengus, the quadrate

is seen to be more forwardly rotated in the latter, and the

hyomandibular to slope more forward. The angle in the middle

Text-fig. 124.

ope.._ if Be

SO/27 aan

SOP

Clupea finta, hyopalatine arch, opercular bones, and mandible, left side, mesial

aspect. For explanation of lettering see p. 493.

of the ectopterygoid, also, is smaller, so that the forward displace-

ment of the quadrate-articular joint does not affect the front part

of the hyopalatine arch. It results, however, in the forward

extension of the mandibular symphysis in front of the pre-

maxillaries, and is accompanied by an elongation of the inter-

opercular and the lower limb of the preopercular.

The symplectic is rather small in Clupea finta; it is relatively

longer and more slender in Clupea harengus. The lower end of

the symplectic of Clupea finta is abruptly terminated, and is

not enveloped by the quadrate as is so generally the case. The

palatine is shorter and broader in Clupea finta than in Clupea

harengus, and has a distinct cartilaginous head for articulation

with the prefrontal, whereas in Clupea harengus it is the pre-

458 DR. W. G. RIDEWOOD ON THE CRANIAL | Dec. 13,

frontal which has the boss of cartilage, the palatine merely

offering a flat facet for articulation with it.

I fail to comprehend the meaning of Owen’s remark (Anat.

Vert. i. 1866, p. 117) that “in the Shad the palatine articulates

with the premaxillary as well as the maxillary.” The palatine

does not touch the premaxilla in Clupea finta.

There are no teeth on the palatine or pterygoid bones in Olupea

finta, but two or three minute teeth occur on the palatine of

Olupea harengus, while in Clupéa sprattus there is a row of

minute teeth on the palatine, extending back even on to the

entopterygoid. Matthews states (/.c. p. 291) that in the Sprat

there are a few delicate teeth on the metapterygoid. These

I have failed to discern.

Text-fig. 125.

bb 4£5(ct) eb?

Clupea harengus. A, hyobranchial skeleton, dorsal view. The epibranchials and

pharyngobranchials of the left side are not shown. 5B, fourth and fifth

branchial arches of the right side, mesial aspect, more enlarged than fig. 125 A.

For explanation of lettering see p. 493.

1904, | OSTEOLOGY OF CLUPEOID FISHES. 459

Opercular Series (text-figs. 123 and 124, pp. 456, 457).—The

opercular and subopercular bones are of average proportions; the

latter is relatively larger in Clupea harengus than in Clupea finta.

There are seven branchiostegal rays in Clupea jfinta—five rather

slender ones borne by the ceratohyal, and two broader ones

attached to the outer face of the epihyal. In Clupea harengus

the numbers are five and three respectively.

Hyobranchial Series (text-fig. 125, p. 458).—The lower hypohyal

of Clupea harengus is larger than the upper. The urohyal is large

and extends back behind the posterior extremity of the third

basibranchial. The glossohyal is a conical cartilage, flanked on

its upper surface by a thin membrane-bone. This is covered by

mucous membrane bearing small teeth; but the teeth, although

they may leave scars when removed, are not intimately attached

to the bone. The only elements of the hyobranchial skeleton

which bear teeth anchylosed with the bone are the fifth cerato-

branchials.

The second basibranchial bears fixed on its upper surface a

toothless membrane-bone which extends forward and backward

over, but is readily removable from, the first and third basi-

branchials. The equivalent membrane-bone in Clupea finta does

not extend over the third basibranchial. The fourth and fifth

basibranchials are represented by an elongated cartilage which is

continued back in the form of a slender rod some distance behind

the point at which the fifth ceratobranchials are attached.

The fourth epibranchial is rather large and flat, and serves

to support the wall of the epibranchial organ. Its posterior

extremity is united with the upper end of the fifth cerato-

branchial by a ligament which is the exact equivalent of that

slender bar of cartilage which in Hngraulis represents the fifth

epibranchial (cf. text-figs. 125 B and 135 .B, pp. 458 and 475).

In Clupea alosa Gegenbaur (Morph. Jahrb. iv. Suppl. p. 24 and

pl. u. fig. 18, Clupea vulgaris or Alosa vulgaris) has figured a

fifth epibranchial cartilage. The first pharyngobranchial is small

and cartilaginous, but a well-developed spicular bone rises

vertically from the upper surface of the anterior extremity of

the first epibranchial.

PELLONA MOTIUS.

In Pellona motius (Brit. Mus. 1888.11.6.64, E. C. Madras) the

top of the cranium is much narrower than in OClupea, and the two

principal longitudinal ridges on the upper surface are closer

together and more nearly parallel. The part of the cranium

behind the orbit is greatly reduced in an antero-posterior direc-

tion, but not in a vertical direction. The temporal foramen is

smaller than in Clupea and the pre-epiotic fossa shallower.

The pro-otic, squamosal, and exoccipital appear to be completely

hollowed out for the accommodation of the cecal diverticula of

the swim-bladder; but the exact relations of the bulle can only

be made out by freely incising the hinder part of the cranium,

460 DR. W. G. RIDEWOOD ON THE CRANIAL [ Dec. 18,

which in the circumstances was not permissible. The pro-otic

bulla is elongated in a direction transverse to the axis of the

skull, and in a posterior view of the cranium a portion of the

squamosal bulla is visible on the mesial side of the descending

ridge of the epiotic bone. An auditory fenestra is present.

The opisthotic bone is small and flat; the orbitosphenoid does

not extend to the prefrontals; and there is no distinct fenestra

between the mesethmoid and the frontal bones, although the

anterior ends of the right and left frontal bones are separated

by a narrow space. The parasphenoid has the same relations

as in Clupea; there are no teeth on the parasphenoid and

vomerine bones.

The post-temporal and supratemporal bones resemble those

of Clupea, except that both are relatively shorter. There is a

small nasal on each side of the head, and seven bones around

the eye.

Vestigial teeth occur on the edges of the premaxilla and

maxilla, The maxilla and two surmaxille are of greater vertical

extent and less horizontal extent than in Clupea. The mandible

is rather short and high, and the highest point is over the middle

of the length of the ramus. The angular bone is small and

distinct. There are six or eight minute teeth at the front of

each dentary bone.

The hyomandibular articulates with the cranium by a single

broad head. The ectopterygoid is sharply bent, even more so than

in Clupea harengus.. The buccal surfaces of the entopterygoid,

ectopterygoid, and palatine are covered with a kind of shagreen

of closely-set, minute denticles.

The bones of the opercular series are much the same as in

Clupea harengus, but they are relatively smaller. The interhyal

is ossified. The urohyal has the form of a thin, triangular plate

of considerable vertical extent.

The fourth epibranchial is relatively less expanded than in

Clupea. The three basibranchial bones, the glossohyal, and the

first and second hypobranchials are covered with a shagreen of

denticles similar to that found on the entopterygoid.

PELLONULA VORAX.

Pellonula vorax (Brit. Mus. 89.11.20.11, Stanley Falls) has a

skull which bears a fairly close resemblance to that of the

Herring. The upper surface of the cranium is smoother, and

the part of the cranium behind the orbit relatively shorter.

The bony bullee resemble those of Pellona rather than those of

Clupea, and a portion of the squamosal bulla is visible in a

posterior view of the cranium on the mesial side of the descending

ridge of the epiotic bone. The exoccipitals and basioccipital were

not present in the specimen examined, so that the presence of an

* auditory fenestra could not be determined.

The orbitosphenoid does not extend to the prefrontals. <A

1904. } OSTEOLOGY OF CLUPEOID FISHES. 461

median fontanelle occurs between the mesethmoid and the

frontal bones. There are no teeth on the parasphenoid and

vomer. .

The relations of the post-temporal are the same as in Clupea,

but the bone is shorter; the vertical limb of the supratemporal

is longer than the horizontal limb.

Each premaxilla bears about eight fairly large and pointed

teeth, but the maxilla is edentulous. The highest point of the

mandible is in advance of the middle of the length of the ramus.

At the front of the dentary are five or six rather long, pointed

teeth, smaller, however, than those of the premaxilla. The

angular bone is distinct.

There is a single row, in places a double row, of small teeth

running along the middle of the convex surface of the ento-

pterygoid, and a row of about six larger teeth, as large as those

of the dentary, set transversely across the front of the palatine.

There are six branchiostegal rays on each side; those which

are attached to the ceratohyal resemble in shape the branchi-

ostegal rays of the Anchovy rather than those of the Herring,

z.e. they are not curved rods, but have the form of plates, each

with a conspicuous antero-ventral projection.

PRISTIGASTER TARTOOR,

Pristigaster tartoor (Brit. Mus, 1889.2.1.2026, Malabar, F. Day)

has a high, narrow skull, partaking of the general lateral com-

pression of the body. The parietal bones are separated; the

temporal grooves are shallow, as in Clupea; there is a temporal

foramen and an auditory fenestra. In the squamosal bone is a

spherical vesicle of the swim-bladder, the bony envelope of which

projects into the large shallow pre-epiotic fossa. The vesicle in

the pro-otic is much larger than that in the squamosal bone, and

its bony envelope bulges upon the ventro-lateral face of the

pro-otic, and also projects in two places on the anterior or orbital

face of the pro-otic bone.

There is a small opisthotic bone, a small basisphenoid, and an

orbitosphenoid ; the orbitosphenoid extends forward towards the

prefrontals, but fails to reach them. There are right and left

posterior wings to the parasphenoid, and the eye-muscle canal

opens between them.

The post-temporal has three limbs; the anterior or supra-

temporal limb is unusually long, the deep or opisthotic limb is

long and slender. The supratemporal is triradiate, and the

upright ray, passing to the parietal bone, is longer than the

other two, The upright limb of the preopercular is about twice

as long as the horizontal limb, and makes with it an angle of

130 degrees.

The nasal is a small, tubular bone. Around the orbit are

seven bones: two narrow postorbitals, two suborbitals, the second

larger than the first, two preorbitals, the lower larger than the

462 DR. W. G. RIDEWOOD ON THE CRANIAL [ Dec. 13,

upper, and a long tapering bone lying antero-dorsally to the orbit.

The gape is small, and its upper margin is formed almost entirely

by the premaxille, although these are small bones, while the

maxille are large. A single row of minute teeth occurs along

the edge of the premaxilla, and these are met by a similar row

along the sharp front edge of the dentary. The ventral edge of

the maxilla is provided with a single row of teeth, but, as in

Olupea, these do not bite against any mandibular teeth. There

are two surmaxille above each maxilla.

The angular bone is distinct from the articular. The ventral

surfaces of the palatine and entopterygoid bones are extensively

covered with minute teeth; there are no teeth on the vomer.

The subopercular is reduced in size; the branchiostegal rays are

five or six in number on each side. A spicular bone is present,

and has the form of an upright rod which broadens out at its

lower end.

HYPERLOPHUS COPII.

The genus Diplomystus was established by Cope in 1877 (Bull.

U.S. Geol. Geog. Surv. Territ. i. 1877, p. 808) for certain

extinct species of fish allied to Clupea, but differing in the

possession of “a series of dorsal scuta, which extend from the

supraoccipital region to the base of the dorsal fin.” The genus

has since been found to be represented at the present day by

species living in the rivers of New South Wales and Chili.

The generic name Diplomystus is, 1 understand from Mr. Bou-

lenger, preoccupied by a Siluroid fish, or, to be more exact, the

name of this Siluroid (Diplomyste, Dumeéril; Diplomystes, Bleeker ;

Diplomystax, Giinther) so closely resembles the name Diplomystus

as to render the latter invalid. In such case the wisest plan is

to apply to “ Herrings with occipito-dorsal serrature” the name

Hyperlophus, as suggested by Ogilby in 1892 (Rec. Austral. Mus.

ii. 1892, p. 26).

The specimen examined is one in the British Museum Collec-

tion, prepared from a fish about 33 inches in length, and marked

“ Hyperlophus copii, 97.10.27.38, N. 8. Wales, Ogilby.”

The parietal bones are separated; the temporal grooves are

as in Clupea; a temporal foramen is present and an auditory

fenestra. The pre-epiotic fossa is present, but with the exception

of its upper part it is largely obliterated by the bulging of the

squamosal bulla. The vesicle of the swim-bladder in the pro-otic

bone is large, and its bony envelope projects upon the ventro-

lateral face of the pro-otic. The opisthotic is small; the basi-

sphenoid cannot be recognised in the specimen under consideration ;

the orbitosphenoid extends forward to meet the prefrontals.

There are right and left posterior wings of the parasphenoid,

and the eye-muscle canal opens between them.

The post-temporal has an epiotic and an opisthotic limb, but

no supratemporal limb. The supratemporal is triradiate, and the

ray which passes to the parietal is longer than the other two.

1904. | OSTEOLOGY OF CLUPEOID FISHES. 463

The upright and horizontal limbs of the preopercular are about

equal in length, and enclose an angle of 100 degrees. The nasal

bone is small and tubular; there is a postorbital bone of moderate

size, two suborbitals, two preorbitals, the upper one smaller than

the lower, and a narrow bone lying antero-dorsally to the orbit.

The gape is so small that, although the premaxilla is small and

the maxilla large, the latter forms only a small portion of the

oral border when the mouth is opened to its widest extent. There

is but a single surmaxilla*, which from its shape and position is

clearly to be identified with the posterior of the two present in

Clupea. The angular bone is distinct. The mandibular ramus

stands high, and the highest point is situated far forwards; the

outline of the ramus is intermediate between that of Clupea

harengus and that of Chatoéssus (text-fig. 128, p. 466). There

are no teeth on the dentary, and none on the premaxilla, maxilla,

palatine, entopterygoid, ectopterygoid, and vomer.

The opercular bones are normal; there are five branchiostegal

rays on each side; the interhyal is bony ; there are two hypohyals,

the lower larger than the upper. The glossohyal is long, narrow,

with minute teeth; the urohyal is large and extends backward

considerably behind the posterior limit of the epihyal.

CHATOESSUS EREBI.

Material examined.—In addition to a skull specially prepared

for the purposes of this investigation from an alcohol-preserved

specimen kindly furnished by Prof. G. B. Howes, two skulls were

examined, belonging to complete skeletons in the Osteological

Collection of the British Museum (67.5.6.99 and 67.5.6.5, both

from Cape York, N. Australia).

Cranium (text-fig. 126, p. 464).—The cranium is stout and

rather broad, and is remarkable for the spines that project from

the squamosal, postfrontal, and prefrontal bones. The middle part

of the parasphenoid is greatly depressed, which gives an appearance

of considerable depth to the middle of the cranium.

The parietals are separated by the supraoccipital. There is an

oval temporal foramen, situated near the anterior end of the

posterior temporal groove, bounded by the frontal and parietal,

and leading directly into the cranial cavity. Behind this, and at

a slightly lower level, is a deep pre-epiotic fossa, bounded above

by the parietal and epiotic, and below by the squamosal and epiotic.

It extends inward and upward as far as the supraoccipital.

There is a subspherical cavity in the pro-otic, and another,

rather larger, in the squamosal, for the accommodation of vesicles

of the swim-bladder; but since these do not appear as bullate

projections on the surfaces of the bones, it is necessary to dissect

* Smith Woodward (Brit. Mus. Cat. Foss. Fishes, iv. p. 139) records Diplomystus

as having two surmaxille. There is a possibility that one surmaxilia had been lost

from each side of the skull examined by me before it came into my hands, but from

the appearance of the maxillary series of bones I do not think that this is at all

likely.

464 DR. W. G. RIDEWOOD ON THE CRANIAL [ Dec. 13,

the skull to determine their presence. The opisthotic is moderately

small, and more ventral than posterior in position ; it touches

the pro-otic, squamosal, and exoccipital, but is remote from the

Text-fig. 126.

Cranium of Chatoéssus erebi.—A, dorsal view; B, left side; C, back view.

For explanation of lettering see p. 493.

epiotic. Anauditory fenestra is present, bounded by the pro-otic,

exoccipital, and basioccipital. At the back of the cranium are

1904. | OSTEOLOGY OF CLUPEOID FISHES. 465

two depressions, bounded externally by the epiotics. The epiotic

has on its postero-superior surface a large, oval, smooth facet

for articulation with the post-temporal; and from the apex of the

epiotic there projects backward into the muscles of the trunk a

separable osseous brush. A similar brush projects back from the

apex of the supraoccipital, which has no crest or spine.

The parasphenoid extends very nearly to the posterior end of

the basioccipital, and although the right and left sides of the

posterior outlet of the eye-muscle canal are formed by vertical

lamine of the parasphenoid, there are no projecting posterior

wings of this bone. The middle portion of the parasphenoid

bears a sharp ventral keel. Neither parasphenoid nor vomer

bears teeth. A small basisphenoid is present, but it has no

descending portion bisecting the eye-muscle canal. The orbito-

sphenoid is fairly large, and has a forwardly directed process that

meets a backward growth of the combined prefrontals. A

fontanelle is present in the roof of the skull between the mesethmoid

and the frontals.

Text-fig. 127.

Chatoéssus erebi, right side of skull. For explanation of lettering see p. 493.

Temporal and Preopercular Series (text-fig. 127).—The post-

temporal is large, and the attachment of its upper limb to the

postero-superior surface of the epiotic is quite intimate, and not

by means of a broad loose ligament. The deep limb is rod-like

and is attached to the back of the opisthotic. The third limb is

Proc. Zoou. Soc.—1904, Vou. II, No, XXX, 30

466 DR. W. G. RIDEWOOD ON THE CRANIAL | Dec. 13,

very short and is united with the back of the supratemporal.

The supratemporal has the usual triradiate arrangement of the

sensory canal,

The horizontal limb of the preopercular is a little more than

half as long as the upright limb, and makes with it an angle of

about 95 degrees. The interopercular is large, considerably larger

than the subopercular, and comes into direct contact with the

angular bone, without the intervention of the usual ligament.

Circumorbital Series (text-fig. 127, p. 465).— There are nine bones

of this series. The nasal is roughly rectangular in shape, with a

narrow tube for the sensory canal passing lengthwise over its

surface. The form and relations of the other bones are readily

to be comprehended by a glance at text-fig. 127.

Maxillary Series (text-fig. 127, p. 465).—The gape is very small

and bounded above by the premaxille. The maxilla is only slightly

longer than the premaxilla. Fries (Scand. Fishes, ed. 2, 11. 1895,

p- 952) states that the premaxilla is longer than the maxilla in

the Chatoéssinze, but such is not the case in the specimens now

under consideration. The front part of the maxilla is of con-

siderable vertical extent, and the posterior extremity is also

expanded. The premaxilla is flattened and scale-lke, with a

sharp lower edge. Neither maxilla nor premaxilla bears teeth.

A single surmaxilla of small size is present on each side.

Text-fig. 128

Chatoéssus erebi, hyopalatine arch, opercular bones, and mandible of left side,

mesial aspect. For explanation of lettering see p. 493.

Mandibular Series (text-figs. 127 and 128).—The remarkable

shape of the mandible is doubtless due to the reduction of the

gape not being accompanied by an adequate forward displacement

of the quadrate-articular joint. The coronoid process thus comes

1904. | OSTEOLOGY OF CLUPEOID FISHES. 467

to le over the anterior half of the ramus; it is formed mainly,

but not entirely, by thedentary. The angular is distinct. There

are no teeth.

Hyopalatine Series (text-fig. 128, p. 466).—The hyomandibular

articulates with the cranium by two heads: a small anterior

and a broad posterior one. It is long, and its axis slopes forward

and makes an angle of about 110 degrees with the symplectic,

which is disposed almost horizontally. The extremity of a

triangular process which rises upward and forward from the

external face of the hyomandibular lies over the outer surface of

the postfrontal spine, and forms with it a kind of sliding joint.

The palatine is very short and broad, and has a single head for

articulating with the ethmoid region of the cranium. There are

no teeth.

Opercular Series (text-figs. 127 and 128, pp. 465, 466).—The

opercular and subopercular bones are of average proportions. The

branchiostegal rays are six in number; the first three are rather

slender and closely set, and situated some distance in advance

of the other three, which are greatly expanded and overlap the

former three. The first four are attached to the outer edge of the

ceratohyal, the fifth les over the suture between the ceratohyal

and epihyal, while the last is attached to the outer edge of the

epihyal.

Hyobranchial Series (text-fig. 129, p. 468).—The hyoid is short,

so short, indeed, that the end of the epihyal is only slightly behind

the end of the first hypobranchial. The lower hypohyal is about

twice as large as the upper. The glossohyal consists of a stout

rod of cartilage, with a small membrane-bone on the posterior

part of its upper surface. The urohyal is a strong bone, the

sections of the posterior half of which have the form of an

inverted Y. The third basibranchial is nearly as large as the

second, and the cartilage that represents the fourth and fifth

basibranchials is drawn out to a considerable length.

Concerning Chatoéssus Giinther states (‘Study of Fishes,’ 1880,

p. 657) that the branchial arches form two angles, one pointing

forwards and the other backwards. This statement, which

appears to be based upon the description given by Valenciennes

(Hist. Nat. Poiss. xxi. 1848, p. 96), does not, I believe, refer to

the skeletal arches as one might suppose on reading the passage,

but is an allusion to the remarkable continuation of the series of

gill-filaments from the top of the epibranchials, particularly the

first, backward along the side of the parasphenoid.

The anterior ends of the fourth ceratobranchials are expanded

both forwards and backwards, so that while not relinquishing

their connection with the anterior ends of the fifth ceratobranchials,

they nearly touch the mesial ends of the third ceratobranchials.

The fifth ceratobranchials are considerably expanded, as also are the

fourth epibranchials. The latter have a curved wall of cartilage

extending upward and inward from the outer edge of the bone,

in relation with the epibranchial organ.

30*

468 DR. W. G. RIDEWOOD ON THE CRANIAL [ Dec. 13,

In the first two branchial arches the epibranchial is longer

than the ceratobranchial, but in the third the reverse is the case.

The third hypobranchials are small and concealed in a dorsal

view by the breadth of the posterior end of the third basibranchial.

Text-fig. 129.

Chatoéssus erebi, hyobranchial skeleton, dorsal view.—The epibranchials and

pharyngobranchials of the right side are not shown. For explanation of

lettering see p. 493.

Whereas in most forms the fourth pharyngobranchial is repre-

sented by a cartilage with a removable dentigerous membrane-

bone on its ventral surface, in Chatoéssus the membrane-bone

(devoid of teeth) flanks the cartilage on three sides, ventral,

mesial, and dorsal, and simulates an ossified fourth pharyngo-

branchial. The first pharyngobranchial is cartilaginous, and a

large spicular bone is present. There are no teeth on any part

of the hyobranchial skeleton.

DUSSUMIERIA ACUTA.

Material ewamined.—In addition to a skull specially prepared

for the purposes of this investigation from an alcohol-preserved

specimen from Madras kindly furnished by Mr. G. A. Boulenger,

another skull was examined, belonging to a complete skeleton in

the Osteological Collection of the British Museum (Brit. Mus.

1889.2.1.2038, Bombay).

Cranium (text-fig, 130, p. 469).—The cranium has all the cha-

1904. OSTEOLOGY OF CLUPEOID FISHES. 469

racteristic features of that of Clupea. The posterior teinporal

groove faces more laterally than in Clupea owing to the width of

the cranium in the squamosal region being proportionately less.

The temporal foramen near the anterior end of the groove is a

Text-fig. 130.

Cranium of Dussumieria acuta.—A, dorsal view ; B, left side; C, back view.

s, point of attachment of the spicular bone to the pro-otic. For explanation

of other lettering see p. 493.

narrow horizontal slit, but the pre-epiotic fossa is large. This

fossa is bounded above by the parietal, below by the squamosal, in

front by the parietal and squamosal, and behind by the epiotic.

It is about half as deep as wide, and the inner wall is mainly

470 DR. W. G. RIDEWOOD ON THE CRANIAL [ Dec. 13,

membranous, but partly formed by lamellar extensions of the

supraoccipital, epiotic, and squamosal bones.

There is an auditory fenestra between the pro-otic, exoccipital,

and basioccipital. A large osseous bulla, enclosing a terminal

vesicle of the swim-bladder, occurs in the pro-otic, and another

in the squamosal. The bony envelopes of the horizontal and

posterior vertical semicircular canals are very prominent , the

former is conspicuous on the hinder part of the squamosal bulla,

the latter forms the posterior vertical edge of the epiotic and

exoccipital. The posterior depression lying to the mesial side of

this edge is rather deep. A smal] opisthotic bone is present.

The supraoccipital has no crest and separates the two parietal

bones.

The posterior third of the length of the parasphenoid consists

of a pair of parallel lamin bounding the sides of the eye-muscle

canal. A process of the alisphenoid runs along the basal portion

of the anterior edge of the postfrontal spine, which is not the

case in Clupea. A basisphenoid is present, but it has no vertical

part descending towards the parasphenoid, The orbitosphenoid

is produced forward to meet the back of the combined prefrontals.

The vomer is strap-shaped and bears a long median row of teeth

which Valenciennes (Hist. Nat. Poiss. xx. 1847, p. 468) failed to

notice. There is no dorsal fontanelle between the mesethmoid

and the frontals.

‘SOp /

Dussumieria acuta, right side of skull. For explanation of lettering see p. 493.

Temporal and Preopercular Series (text-fig. 131).—The post-

temporal has a long limb resting upon the epiotic prominence, a

shorter, rod-like limb attached to the back of the opisthotic, and

1904. | OSTEOLOGY OF CLUPEOID FISHES. 471

a still shorter lamellar limb overlapped by the back of the supra-

temporal. The supratemporal forms a kind of outer wall to the

large pre-epiotic vacuity. There is also a pair of ordinary body-

scales, about as large as the supratemporals, which project back

from the transverse ridge of the parietals. These, together with

the supratemporals and the preoperculars, form a kind of girdle

around the top and sides of the hinder part of the head. The

preopercular is triangular in shape, with a concave anterior

border; the horizontal limb is more than half as long as the

upright limb, and makes with it an angle of about 100 degrees.

Cirewmorbital Series (text-fig. 131, p. 470).—This series consists

of seven bones: a small nasal, a long narrow bone overlying the

prefrontal, and five others of moderately large size.

Maxillary Series (text-fig. 131, p. 470).—The premaxilla is un-

usually short, and has the form of a flat, square scale, overlapping

the anterior portion of the maxilla, and with teeth along its lower

edge. The maxilla is fairly long and is toothed along the whole

length of its lower edge. There are two surmaxille.

Mandibular Series (text-figs. 131 and 132).—The mandible

closely resembles that of the Herring, and the highest point lies

over the anterior half of the ramus. Long, curved, pointed

teeth, similar to those of the maxilla and premaxilla, occur along

the antero-superior edge of the dentary. The angular bone is

separate; there is no distinct sesamoid articular.

Text-fig. 132.

pop op

Dussumieria acuta, hyopalatine arch, opercular bones, and mandible of left side,

mesial aspect. For explanation of lettering see p. 493.

Hyopalatine Series (text-fig. 132).—The hyomandibular has two

distinct heads for articulation with the cranium, the posterior

slightly larger than the anterior. ‘The symplectic makes an angle

of about 125 degrees with the axis of the hyomandibular; it is

flat, and is largely overlapped on its inner surface by the inter-

472 DR. W. G. RIDEWOOD ON THE CRANIAL | Dec. 13,

opercular, The palatine, ectopterygoid, and entopterygoid bear

teeth. The ectopterygoid has an outwardly projecting ledge

which serves to support the eyeball, and which comes into close

relation with the upper edge of the middle of the three large

bones of the suborbital series. The ectopterygoid is curved in the

middle of its length, but not sharply bent.

Opercular Series (text-figs. 131 and 132, pp. 470, 471).—The

gill-cover is large, and the opercular aud subopercular bones are of

corresponding proportions. The branchiostegal rays are thirteen

in number; eight of these are set on the outer side of the lower

edge of the ceratohyal, and five on the epihyal. The shape is

approximately the same in all, but the hinder ones are larger than

those in front. Valenciennes (Hist. Nat. Poiss. xx. 1847, p. 469)

and Giinther (Brit. Mus. Cat. Fish. vii. 1868, p. 466) put the

number of branchiostegal rays as fifteen.

Hyobranchial Series.—The lower hypohyal is slightly larger

than the upper. The posterior one-third of the glossohyal

cartilage is ossified; a narrow membrane-bone with a row of

teeth along the middle covers the whole. The first, second, and

third basibranchials are covered by separate membrane-bones,

each bearing a narrow central row of teeth, but the investing

lamina of the second basibranchial extends forward over the

hinder part of the first basibranchial. The urohyal is long and

slender. The fourth epibranchial is about as much expanded as

in the Herring. The first pharyngobranchial is cartilaginous,

and a spicular bone rises from the front end of the first epi-

branchial, and is attached by ligament to the pro-otic at the point

marked s in text-fig. 130 B, p. 469.

ENGRAULIS ENCRASIGHOLUS.

The chondocranium of a young Anchovy of 25 mm. length has

been described and figured by Pouchet (Journ. Anat. et Phys.

1878, p. 75, and figs. 49 and 50), but the account has no important

bearing in the present connection.

Cranium (text-fig. 133, p. 473).—The most remarkable feature

about the general aspect of the cranium is the considerable vertical

extent of the orbital region, a feature directly related to the large

size of the eyes. The parietals are separated, and on the roof of

the cranium is a pair of fontanelles, each bounded by the frontal,

parietal, and supraoccipital. On the upper surface of each frontal

bone are two cross bars or arches of bone, the anterior one strictly

transverse, the second oblique.

A distinct opisthotic is not present; the deep or opisthotic

limb of the post-temporal is attached to a process of the exoccipital

lying immediately over the foramen for the tenth nerve. The

posterior temporal groove is shallow, and the temporal foramen,

near its anterior end, is large and bounded by the frontal and

parietal, with sometimes also a small portion of the squamosal

below. Owing to the large size of the squamosal bulla that

1904. ] OSTEOLOGY OF CLUPEOID FISHES. 473

encloses a vesicle of the swim-bladder, the pre-epiotic fossa above

it is rendered very shallow as compared with that of Clupec.

The auditory fenestra, opening into the perilymphatic cavity,

is bounded by the pro-otic, exoccipital, and basioccipital, as mm

Clupea.

Text-fig. 133.

Cranium of Hngraulis encrasicholus, lett side. For explanation of lettering

see p. 493.

The foramen for the trigeminal nerve is of large size ; it faces

more anteriorly than usual, and is bounded by the alisphenoid

and pro-otic, instead of by the pro-otie alone. The pro-otic bulla,

like that of the squamosal, is large; it is not spherical, but is

drawn out in a direction nearly at right-angles to the median

plane of the head (see Journ. Anat. and Phys. xxvi. 1891, p. 36,

and p. 382, fig. D). A basisphenoid is present, but it has no

descending limb. The articulation for the hyomandibular is

situated rather far forward; the anterior head of the hyo-

mandibular articulates with the postfrontal, and the hinder with

the squamosal.

The lateral temporal groove is broad and shallow. Removal of

its floor exposes a fairly large cavity opening laterally by two

apertures—one oyer the articular process of the squamosal for the

reception of the posterior head of the hyomandibular, and the

other immediately in front of this. The cavity is roofed by

the frontal, and is bounded in front by the postfrontal, behind

by the squamosal, and below mainly by the pro-otic.

The parasphenoid is depressed at about the middle of its length,

doubtless in relation with the large size of the eyes. It does not

project behind the occipital articulation, but it ends posteriorly

in a pair of parallel lamelle on the right and left sides of the

posterior outlet of the eye-muscle canal. Both parasphenoid and

vomer are edentulous. The front of the cranium is formed by

the mesethmoid, and not by the vomer (cf. Clupea). The mes-

ethmoid is large, of considerable vertical extent, but thin; the

prefrontals are relatively small. The orbitosphenoid is small, and

AT4. DR. W. G. RIDEWOOD ON THE CRANIAL [ Dec. 13,

makes no attempt to support the great membranous interorbital

septum, neither does it send a process forward to meet the

prefrontals as it does in Clupea.

Temporal and Preopercular Series (text-fig. 134).—The post-

temporal has the usual three limbs. The longest is the upper

limb, loosely attached by ligament to the upper surface of the

epiotic. The opisthotic limb is delicate and rod-like, and is

attached, as already mentioned, to a process of the exoccipital,

there being no distinct opisthotic bone. The horizontal limb,

bearing the sensory canal, touches the back of the supratemporal.

The supratemporal has the usual triradiate form. The axis of

the preopercular slopes downward and backward, and its lower

limb is very short, and instead of being forwardly directed is

vertical. The interopercular is remarkably small.

Text-fig. 134.

’

SOP top cr sm

Engraulis encrasicholus, right side of skull. For explanation of lettering

see p. 493.

Oircumorbital Series (text-fig. 134).—The bones of this series

are six in number. he largest lies below and behind the eye,

and above this bone are two small postorbitals. Both nasal and

preorbital, particularly the latter, are relatively larger than in

Clupea.

Mavillary Series (text-fig. 134).—The premaxilla is small, and

bears teeth on the posterior half of its lower edge. The two

premaxille extend, without meeting, below the mesethmoid, and

not in front of it as in Clupea; it is, in fact, the projection of the

ethmoid region in advance of the mouth that gives the character-

istic appearance to the head of Hngraulis. The maxilla is long,

and is toothed all along its edge except at the anterior end, where

it is overlapped by the premaxilla. There are two surmaxille, a

broad one behind and a long thin one in front. They lie in an

extensible membrane which is continuous with the lower edge of

1904. | OSTEOLOGY OF CLUPEOID FISHES. 475

the suborbital plate, and they form a triangle with the posterior

half of the maxilla when the mouth is widely opened.

Mandibular Series (text-figs. 134 and 135 A).—The mandible is

long, and the dentary bears teeth along the whole of its upper

edge. The coronoid process is situated far back, and does not

rise high in proportion to the length of the ramus. The angular

is not distinct from the articular, and there is no sesamoid

articular.

Text-fig. 135.

Engraulis encrasicholus—A, hyopalatine arch and mandible of left side, mesial

aspect ; B, fourth and fifth branchial arches of left side, mesial aspect ;

C, hyoid of right side, with branchiostegal rays and subopercular bone, external

view. For explanation of lettering see p. 493.

Hyopalatine Series (text-fig. 135 A).—The most striking features

of the hyopalatine arch are related to the backward thrust of the

quadrate articulation. The axis of the quadrate slopes backward ;

the axis of the hyomandibular also slopes backward, but the

symplectic is not in the same line with it, as might have been

expected. The hyomandibular articulates with the cranium by

two distinct heads, and the opercular head is fairly long. The

edges of the palatine and ectopterygoid bear microscopic teeth,

476 DR. W. G. RIDEWOOD ON THE CRANIAL [ Dee. 18,

and a row of similar teeth occurs on the middle of the palatal

surface of the entopterygoid.

Opercular Series (text-figs. 134 and 135 C, pp. 474, 475).—The

opercular bone is remarkably large; the subopercular is compa-

ratively small, and extends forward beneath the interopercular,

and is bound to the posterior part of the outer face of the epihyal

in such a way that it has a tendency, during the process of dis-

articulation of the skull prior to the maceration of the parts, to

leave the opercular bone and to come away with the hyobranchial

skeleton. In such cases it is liable to be mistaken for the hinder-

most of the branchiostegal rays (text-fig. 135 C, p. 475), a fact

long ago noted by Valenciennes, who wrote (Hist. Nat. Poiss. xxi.

1848, p. 11) concerning the subopercular, ‘il faut faire attention

de ne pas le confondre avec un rayon de la membrane branchio-

stege.” This is an argument in favour of regarding the opercular,

subopercular, and branchiostegal bones as components of one and

the same series, to which series the preopercular and interoper-

cular do not belong.

The branchiostegal rays are usually ten in number on each

side, sometimes nine or eleven; I have not seen as many as

thirteen, the number given by Valenciennes (J. c. p. 11) and

Giinther (Brit. Mus. Cat. Fishes, vii. p. 386). The first nine are

situated on the outer face of the whole length of the ceratohyal,

the last on the epihyal. They form a well-graduated series, and

are flat, not rod-like. When, as above mentioned, the sub-

opercular is left attached to the epihyal, it is seen that the

subopercular and the last branchiostegal ray resemble one another

closely in shape and size, but that there is some discontinuity in

the series owing to the interval between the subopercular and the

last branchiostegal ray being greater than that between the last

two branchiostegal rays (text-fig. 135 C, p. 475).

Hyobranchial Series.—The interhyal is a long and rod-like bone

(text-fig. 135 A, p.475). The hyoid is long, the posterior end of the

epihyal being close to the posterior end of the first ceratobranchial.

The lower hypohyal is much larger than the upper, and excludes

the latter from union with the ceratohyal (text-fig. 135 C, p. 475).

The glossohyal is extremely reduced, and consists of a small cone

of cartilage, with a small cap of edentulous membrane-bone on its

upper surface. The urohyal is long and slender.

A narrow membrane-bone, with crowded small teeth, extends

over the whole length of the second basibranchial, and projects

forwards over the posterior half or more of the first basibranchial

and backwards over the anterior sixth of the third basibranchial.

The second basibranchial is remarkably long, a fact which disturbs

the parallelism of the first and second ceratobranchials. ‘The

first hypobranchials are long, almost as long as the first cerato-

branchials; the second, however, are small and triangular, and

are fused with the sides of the posterior end of the second basi-

branchial. The third hypobranchials are normal, and slope

obliquely down the sides of the third basibranchial.

1904. | OSTEOLOGY OF CLUPEOID FISHES, ATT

The epipharyngeal teeth and the hypopharyngeal teeth on the

fifth ceratobranchials are larger and more pointed than the teeth

of the basibranchial skeleton. A slender cartilaginous fifth

epibranchial is present, confluent at its upper eud with the

cartilaginous posterior part of the fourth epibranchial (text-fig.

135 B, p. 475). Between the cartilaginous and ossified parts of

the fourth epibranchial is a fenestra closed by membrane. The

first pharyngobranchial is cartilagmous and small; the spicular

bone is remarkably long and slender.

CoILIA NASUS.

Material examined.—In addition to a skull specially prepared

for the purposes of this investigation from an alcohol-preserved

specimen from Kiu Kiang kindly furnished by Mr.G. A. Boulenger,

another skull was examined, belonging to a complete skeleton in

the Osteological Collection of the British Museum (Brit. Mus.

91.1.31.30, Shanghai).

Cranium (text-fig. 136, p. 478).—The cranium is broad and

short, and its posterior surface, instead of rising vertically from

the basioccipital, slopes very much forward. The parietal bones

nearly meet in front of the supraoccipital. The posterior temporal

groove is shallow, and has neither a pre-epiotic fossa nor a temporal

foramen, although on the course of the suture between the frontal

and the parietal bones is a minute depression, large enough to

admit the point of a pin, which evidently represents the last

trace of the closed foramen. There is no auditory fenestra

between the pro-otic, exoccipital, and basioccipital bones.

The bony bulle containing the cecal diverticula of the swim-

bladder are large and prominent. The pro-otic one occupies

nearly the whole of the ventro-lateral face of the pro-otic bone,

and its upper extremity projects upwards into a lateral vacuity

of doubtful homology, but evidently corresponding with that

cavity which in Hngraulis lies beneath the lateral temporal groove,

bounded above by the frontal and squamosal, below by the post-

frontal, pro-otic, and squamosal, behind by the squamosal, and in

front by the postfrontal. The squamosal bulla is just visible in

the hinder part of this vacuity, and also above it, in the floor

of the posterior temporal groove. It is freely open below into a

large exoccipital bulla*, the equivalent of the fusiform dilatation

of the air-duct which in Clupea leads to the pro-otic and squamosal

vesicles. Behind this, however, is another smaller exoccipital

bulla which is blind posteriorly, and communicates anteriorly

by a narrow neck with the former exoccipital vesicle.

The squamosal bone takes no part in the articulation for the

head of the hyomandibular; this is formed mainly by the post-

frontal, but partly also by the pro-otic. The appearances are

such as strongly to suggest that the posterior head of the hyo-

* These relations can only be made out by making suitable incision into the

cranial bones, d

478 DR. W. G. RIDEWOOD ON THE CRANIAL [ Dec. 13,

mandibular of Hngraulis and its corresponding articular facet

in the squamosal bone have disappeared in the present genus.

In text-fig. 138 (p. 481) the part marked hm’ is the articular head

Text-fig. 136.

Cranium of Coilia nasus.—A, dorsal view; B, left side; C, back view. s, point

of attachment of the spicular bone to the pro-otic; ¢f’, last trace of

temporal foramen. For explanation of other lettering see p. 493.

of the hyomandibular, the part above and behind the fenestra is

comparatively remote from the side of the cranium.

1904. | OSTEOLOGY OF CLUPEOID FISHES. 479

The parasphenoid has no ascending processes in the region of

the pro-otics, but there is a well-marked process of each pro-otic,

directed forward and downward, which is suturally united with

the supero-lateral edge of the parasphenoid. The parasphenoid is

short, and underlies but a small portion of the basioccipital. It

has no posterior wings, and the eye-muscle canal does not open

behind, although there is a minute depression at the back of the

parasphenoid, in the region where the opening might be expected

to occur, The anterior part of the parasphenoid is broad,

V-shaped in section, and united by elaborate jagged sutures with

the prefrontals and vomer, The vomer has five or six teeth on

each side, the two patches being widely separated. The mesethmoid

projects considerably in advance of the vomer and has a dorsal

crest ; yet the ethmoid region as a whole is short.

Running across the roof of each frontal is an arched bar of

bone. The alisphenoids face forwards, 7. ¢. they lie transversely

to the cranial axis instead of sloping forward and inward. The

orbitosphenoid is fairly large, and is tubular in its hinder two-

thirds. It passes back between and below the alisphenoids, and

comes into extensive relation with the pro-otic bones. The

tubular part does not lie close up under the median suture of the

frontals, but is separated by a vertical fenestrated sheet of the

orbitosphenoid bone. The opisthotic is absent, and there seems

to be no basisphenoid.

Text-fig. 137.

pe By cor

pop

Coilia nasus, right side of skull. For explanation of lettering see p. 493.

Temporal and Preopercular Series (text-fig. 137).—The post-

temporal has but one limb, a long one, the anterior extremity of

which rests on the epiotic prominence. This limb is concealed

in text-fig. 137 by the supratemporal. ‘The rest of the bone

480 DR. W. G. RIDEWOOD ON THE CRANIAL | Dec. 13,

consists merely of a semi-tubular scale touching the back of the

supratemporal, The supratemporal is a tubular scale which does

not exhibit the usual triradiate character, since the parietal

division of the sensory canal branches in the skin just anterior to

the supratemporal bone. The axis of the preopercular slopes

strongly backward; the anterior edge of the bone is concave and

not angulate, so that the proportions of the upper and lower

limbs cannot be determined.

Cireumorbital Series (text-fig. 137, p. 479)—The nasal is very

small. Beneath the nasal sac are two bones, both rather firmly

attached to the prefrontal. There are three postorbitals and three

suborbitals, the former series making an acute angle with the

latter.

Maxillary Series (text-fig. 137, p. 479).—The extraordinary

length of the maxilla is one of the most remarkable features of the

fish under consideration; indeed, it is difficult to understand what

purpose the teeth on the hinder part of the maxilla can possibly

serve. A similar prolongation of the maxilla is met with in some

species of Hngraulis, attaming a maximum in Hngraulis mystav

and Engraulis setirostris (Cuvier and Valenciennes, Hist. Nat.

Poiss. xxi. 1848). The actual length of the projecting part of

the maxilla varies in different specimens of Coilia, doubtless

owing to fracture during life; but 1t may be taken as a rule that

the toothed part of the maxilla is two-and-a-half times as long as

the tooth-bearing part of the dentary. The premaxille extend

below the mesethmoid, not in front of it; they nearly touch, but

do not actually meet in a symphysis. There are two surmaxille.

Mandibular Series (text-figs. 137 and 138, pp. 479, 481).—The

mouth is so large that the ramus of the mandible is longer than

the cranium. The dentary and articular components of the coro-

noid process are separated by a short interval. This separation,

however, although striking, is evidently a feature of no great

importance; it occurs in an exaggerated form in Gonorhynchus,

it occurs in some Percoid and Berycoid fishes, and doubtless in

many other groups. The dentary bears teeth similar to those on

the maxilla and premaxilla. The angular is not distinct from the

articular, and there is no sesamoid articular.

Hyopalatine Series (text-fig. 138, p. 481).—In relation with the

great size of the mouth the quadrate articulation is thrown far

back, and both quadrate and hyomandibular bones are backwardly

rotated. The head by which the hyomandibular articulates with

the cranium is small and single (see p. 478). The symplectic is

not in a direct line with the axis of the hyomandibular, but forms

an angle of about 140 degrees with it. The metapterygoid is

large, and the entopterygoid small. The ectopterygoid is nearly

straight, and the palatine articulates with the prefrontal by two

contiguous heads, which are right and left, not anterior and

posterior. Teeth are borne on the edge of the palatine and the

anterior part of the edge of the ectopterygoid.

Opercular Series (text-fig. 137, p.479).—The opercular bone has a

1904. ] OSTEOLOGY OF CLUPEOID FISHES. 481

strongly marked backward slope ; the opercular and subopercular

bones are of average size, and take no share in that enlargement

which is so marked in the bones of the mouth. There are eleven

branchiostegal rays on each side. The first is attached at the

front of the ceratohyal, the ninth at the junction of the ceratohyal

and epihyal, and the remaining two on the epihyal. They are

all curved rods except the last two, which are somewhat

lamellate.

Text-fig. 158.

Am . Am

Coilia nasus, hyopalatine arch and mandible of left side, mesial aspect : him', head

of hyomandibular articulating with the cranium. For explanation of other

lettering see p. 493.

Text-fig. 139.

Coilia nasus, hyobranchial skeleton, dorsal view. The epibranchials and pharyngo-

branchials of the right side are not shown. For explanation of lettering see

p. 493.

Hyobranchial. Series (text-fig. 139).—The interhyal is a long,

rod-like bone. There are two hypohyals, the upper one small

and situated antero-superiorly to the lower, which alone is in

Proc. Zoou. Soc.—1904, Vou. Il. No. XX XT. 31

482 DR. W. G. RIDEWOOD ON THE CRANIAL [ Dec. 13,

contact with the front of the ceratohyal. The glossohyal is an

ill-defined cartilage of small size. The second basibranchial is

large in comparison with the first and third, but it is a hollow

shell of bone, the interior of which is occupied by a fatty mass.

The dentigerous membrane-bone that covers it overlaps the

posterior three-fourths of the first basibranchial and the anterior

third of the third basibranchial.

The second hypobranchials are fused with the sides of the

second basibranchial, but the line of demarcation is obvious. The

third hypobranchials are rather long, and slope forward and

downward at the sides of the third basibranchial. The urohyal

is long, and extends considerably behind the posterior end of the

third basibranchial. The epibranchials are longer than usual in

proportion to the ceratobranchials; the fourth is moderately

expanded in a vertical direction.

The first pharyngobranchial is cartilaginous; and the spicular

bone projects upward, backward, and outward, and is attached to

the pro-otic bone at the point marked s in text-fig. 136 B, p. 478.

The cartilaginous plate that represents the fourth and fifth

basibranchials is continued back between the closely approximated

anterior ends of the fifth ceratobranchials, and projects some

distance behind as a free rod of cartilage.

CHANOS SALMONEUS.

The accessory branchial organ of Chanos, briefly alluded to by

Johannes Miiller (‘ Bau und Grenzen der Ganoiden,’ Berlin, 1846,

pp. 74 and 75), has been described and figured by Hyrtl, but the

relations of the skeletal parts to this organ are not shown

(Denkschr. Akad. Wiss. Wien, xxi. 1863, pp. 1-10 and pl. 1;

also Sitzungsber. Ak. Wiss. Wien, xlii. 1, 1861, pp. 155 & 156).

Material ecamined.—In addition to two skulls (a large one, A,

and a small one, B) specially prepared for the purposes of this

investigation from alcohol-preserved specimens kindly furnished

by Mr. G. A. Boulenger, a third skull (C) was examined, belonging

to a skeleton in the Osteological Collection of the British Museum

(Brit. Mus. 98.9.13.1, Tongatabu).

Cranim (text-fig. 140, p. 485).—The cranium is broad and flat-

tened. The parietals are separated by the supraoccipital, but above

the supraoccipital there lie two sensory-canal scales of the transverse

commissural system which in old specimens (A and C) fuse with

the right and left parietals, and thus produce the effect of a false

union of the two parietals over the supraoccipital (text-fig. 140 A,

p- 483). The posterior temporal fossa is large and completely roofed

in. Its inner wall is formed by the supraoccipital and epiotic, its

floor and outer wall by the postfrontal and squamosal, and its root

by the frontal, parietal, squamosal, and epiotic. Its anterior end

is blind, and lies over the middle part of the postfrontal bone.

Hyrtl (.c. p. 8) has pointed out that the great posterior temporal

1904. | OSTEOLOGY OF CLUPEOID FISHES. 483

vacuity is occupied by the upper longitudinal trunk-muscle, and

contains nothing else.

Text-fig. 140.

Cranium of Chanos salmoneus.—A, dorsal view ; B, left side; C, back view.

For explanation of lettering see p. 493.

Owing to the considerable extent to which the epiotic bones

project posteriorly, and to the hollowness of the posterior surface

oil.

484 DR. W. G, RIDEWOOD ON THE CRANIAL [ Dec. 13,

of the supraoccipital, there is a large depression at the back of the

cranium between the two epiotics. This is not subdivided by a

median crest of the supraoccipital, but a supraoccipital spine lies

dorsally to it, and splits up posteriorly into seven or eight fine

bony filaments, compared by Hyrtl (l. c. p. 3) with the “ossified

tendons of birds. The two exoccipitals are produced backward

and upward into pointed plates, which form a roof not only for

the medulla oblongata, but for the anterior part of the spinal cord

as well.

The squamosal is produced into a spine which slopes backward,

outward, and downward, and extends to the posterior end of the

post-temporal. The opisthotic is small, and is applied to the

inner side of the basal portion of this spine, but it also touches

the exoccipital. The articular surface for the head of the hyo-

mandibular slopes downward and forward. ‘The lateral temporal

groove above the postfrontal spine is to a large extent roofed over

by projecting eaves from the frontal and squamosal bones, mainly

the former. Incision into the squamosal and pro-otic bones fails

to disclose the presence of bullae for the lodgment of cecal

diverticula of the swim-bladder. There is no subtemporal fossa.

There is no orbitosphenoid nor basisphenoid. The parasphenoid

is rather sharply bent at about the middle of its length, and at

this point there is on the ventral surface a well-marked oval

depression. The ascending wings of the parasphenoid rise

moderately high up the fr ont of the pro-otics. The parasphenoid

does not extend as far posteriorly as the hind end of the basi-

occipital, and the eye-muscle canal does not open behind. Neither

the parasphenoid nor the vomer bears teeth. A considerable

proportion of the ethmoid region remains cartilaginous. The

pr efrontals do not meet one another mesially, and the mesethmoid

is a thin horizontally disposed lamina of bone of ectosteal origin.

Projecting backward and outward from each side of the back

of the cranium are two strong tendon-bones, or intermuscular

bones. ‘The larger of the two arises from the dorso-latero-posterior

part of the basioccipital, and is attached to the upper end of the

clavicle. The other arises from the back of the exoccipital,

immediately posterior to the aperture for the exit of the vagus

nerve, and terminates in muscular tissue. The deep limb of the

post-temporal, attached to the opisthotic, is parallel to these, and

its resemblance to them is very striking.

Temporal and Preopercular Series (text-fig. 141, p. 485),—The

post-temporal tends to fuse with the supraclavicular. Its upper

limb is long, and lies so far over the epiotic as to touch the supra-

occipital. The deep limb is about half as long as the former and

is attached to the opisthotic. A study of the skull of Chanos

leaves little room for doubting that the opisthotic limb of the

post-temporal belonged perils to the same category as the

tendon-bones that project back from the exoccipital, but has now

become united by its posterior end with the post-temporal bone

(see preceding paragraph).

1904. | OSTEOLOGY OF CLUPEOID FISHES. 485

The third limb of the post-temporal is wanting, since the supra-

temporal overlaps the body of the post-temporal. The supra-

temporal is a flat scale of bone, with the usual triradiate sensory

canal near its lower edge. It covers in the space between the

epiotic limb of the post-temporal above and the squamosal spine

below.

Text-fig. 141.

st sht

SOp pop

Chanos salmoneus, right side of skull. For explanation of lettering see p. 493.

Overlapping the antero-superior part of the opercular bone is a

flat bone which may be termed the “subtemporal.” A branch of

the sensory canal passes from the supratemporal down the

anterior edge of the bone on its way to the preopercular, which

fact, taken in conjunction with the position of the bone below the

squamosal and above the preopercular, points to the conclusion

that the bone is the homologue of that which, in the Salmon,

Parker (Phil. Trans. Roy. Soc. clxii. 1873, p. 99 and pl. 6.

fig. 1, st) erroneonsly called the supratemporal.

In relation with the forward displacement of the quadrate

articulation, the imteropercular and the lower limb of the pre-

opercular are much elongated in a horizontal direction.

Circumorbital Series (text- -fig. 141).—There are eight bones of

this series. The orbital ring is complete, there being two elongated

supraorbitals which meet above the eye, Inve anterior of the two

is in the large specimens examined (A and C) swollen after the

manner of the frontal and some other bones of Hphippus. The

nasal 1s remarkably small and hable to be overlooked.

Maxillary Series (text-fig. 141)—The premaxilla and maxilla

are short and broad, the gape being greatly reduced in size.

Except when the mouth is opened to an unnaturally wide extent,

the premaxilla alone bounds the gapeabove. Both the premaxilla

and maxilla are thin, curved scales with sharp lo-ver edges, devoid

of teeth. There is no surmaxilla.

486 DR. W. G. RIDEWOOD ON THE CRANIAL | Dec. 13,

Mandibular Series (text-figs. 141 and 142)—The mandible is

of remarkable shape, since the anterior part 1s quite thin, having

the form of a curved rod, whereas the coronoid process is broad

and stands high. The coronoid process is formed entirely by the

dentary, there being no ascending process of the articular. A

sesamoid articular is present; it lies above the anterior part of

the endosteal articular, and is slightly movable m a wet pre-

pavation, The angular is distinct from the articular. There are

no teeth.

Text-fig. 142.

er Uj

Chanos salmoneus, hyopalatine arch and mandible of left side, mesial aspect.

For explanation of lettering see p. 493.

Hyopalatine Series (text-fig. 142).—The hyomandibular arti-

culates with the cranium by a single broad head, the upper edge

of which, instead of being horizontal, slopes downward and forward.

In relation with the reduction in the size of the mouth, the

quadrate has been dvawn so far forward as to have parted from

the symplectic and metapterygoid entirely, and it is attached

to the ectopterygoid in such a way that a fair proportion of

this bone lies behind it. The hyomandibular appears to be but

little affected, and its axis is nearly vertical. There is an angle

of about 110 degrees between the axes of the symplectic and

hyomandibular. No teeth occur on the palatine and pterygoid

bones.

Opercular Series (text-fig. 141, p. 485).—The opercular and sub-

opercular bones are of average proportions; the ascending process

of the subopercular which rises along the front edge of the oper-

cular is high and slender. There are only four branchiostegal rays

on each side. They are long, and rather broad and lamellate. ‘Two

are attached to the outer face of the posterior part of the cerato-

hyal, and two on the outer face of the epihyal. On examining

the skull of Chanos, one cannot fail to be impressed by the regular

manner in which the branchiostegal rays grade off into the

opercular and subopereular bones, whereas the preopercular and

interopercular fail to fit into the series.

Hyobranchial Series (text-fig. 143, p. 487).—The most striking

features of the hyobranchial skeleton are the smallness of the hyoid,

1904. ] OSTEOLOGY OF CLUPEOID FISHES. 487

the great size of the uvohyal, and the enlargement of the hinder

parts for the support of the wall of the epibranchial organ. The

structure of this organ has been described by Hyrtl (i. ¢. p. 4).

The lower hypohyal is considerably larger than the upper.

The hind end of the glossohyal overlaps the front of the first

basibranchial, and a thin flake of bone projects from the back of

the first basibranchial over the front of the second. A similar

Text-fig. 145.

Chanos salmoneus, hyobranchial skeleton, dorsal view. ‘The epibranchials and

pharyngobranchials of the vight side are not shown. For explanation of

lettering see p. 493.

flake extends from the second basibranchial over the front part

of the third. he anterior half of the glossohyal is cartilaginous,

but is covered by a membrane-bone which is continued back over

the endosteal glossohyal. The third basibranchial is larger than

the first and second put together—as a rule the second basi-

branchial is the largest of the three. The third hypobranchials

are fused with the sides of the third basibranchial, but the

boundary line between them is not obliterated.

There is a symphysis of considerable length between the two

fifth ceratobranchials, and from the front of this symphysis there

passes forward a bar of cartilage which, in front of the anterior

ends of the fourth ceratobranchials, enlarges into a roughly hexa-

gonal plate. The separation of the anterior ends of the fourth

and fifth ceratobranchials is noteworthy. Behind the symphysis

of the fifth ceratobranchials is a horizontal tract of cartilage,

488 DR. W. G. RIDEWOOD ON THE CRANTAL | Dec. 13,

continuous with the pair of great cartilages which curve upward

from the suturally united posterior ends of the fourth and fifth

ceratobranchials. The upper ends of these great cartilages meet

in the middle line above, but do not fuse ; from the anterior edge

of each there stretches forward an extensive tract of thin, but

tough membrane, which fills in the angle between the body of the

fourth epibranchial and the remarkably long process of the same

which slopes upward and backward from near its anterior end.

The first three epibranchials are of approximately the same

length, but the fourth is longer and wider. The first pharyngo-

branchial is a forwardly directed cartilage of conical shape, and

there is an upright spicular bone rising from the junction of this

with the front of the first epibranchial. There are no teeth on

any part of the hyobranchial skeleton.

SUMMARY.

On comparing the foregoing accounts of the cranial osteology

ot Chirocentrus, Clupea, Pellona, Pellonula, Pristigaster, Hyper-

lophus, Chatoéssus, Dussumieria, Engraulis, Coilia, and Chanos,

the most important features of resemblance and difference appear

to be as follows.

The parietal bones are rather small in size*, and are separated

the one from the other by the supraoceipital. In Cozlia, however,

they nearly meet in front of the supraoccipital, and in Chanos,

owing to the fusion of the commissural sensory-canal bones with

the parietal bones, the latter appear to meet over the top of the

supraoccipital 7. There is a fontanelle between the anterior ends

of the two frontal bones in Ohirocentrus, Clupea, Pellonula,

Pellona (a mere cleft in this genus), and Chatoéssus, but not i the

other six genera. In Coilia and Hngraulis the mesethmoid pro-

jects considerably in advance of the vomer, which is not the case

in the other genera.

Of the genera under consideration, the only one in which the

posterior temporal groove is roofed over is Chanos. The temporal

foramen appears to be a special feature of the Clupeoid skull. It

is an aperture, usually oval in shape, with the long axis horizontal,

bounded by the frontal and parietal bones. In no instance have I

‘found the postfrontal bone forming part of the boundary, although

Boulenger regards this as the normal conditiont. The only

departure from the general rule that has come within my know-

ledge is in the case of Hngraulis, in which a small portion of the

squamosal may come between the parietal and the frontal in the

* Small is, of course, but a relative term, and opinions may differ as to the employ-

ment of the word. ‘‘Trés petits,” the expression used by Boulenger (Poissons du

Bassin du Congo, 1901, p. 123), appears to me to convey an exaggerated idea of the

smallness of the parietal bones. I leave the figures that illustrate this paper to speak

for themselves.

+ Cope, it is worth noting, included the Lutodiride (i. e. Chanide) under the

heading “ Parietals united” (Trans. Amer. Phil. Soc. n. s. xiv. 1871, p. 455).

+ Poiss. Bass. Congo, 1901, p. 123. “Un grand trou de chaque coté du crane

bordé par le frontal, le postfrontal, et le pariétal.”

1904. | OSTEOLOGY OF CLUPEOID FISHES. 489

lower edge of the foramen, but does not invariably do so. The

temporal foramen is reduced to a pin-point depression in Cozlia,

and is wanting entirely in Chanos.

The pre-epiotic fossa, another Clupeoid feature, is a depression

at the side of the cranium, bounded by the epiotic, squamosal, and

parietal bones. It is wanting in Chanos; in Coilia it is entirely

obliterated, and in Pristigaster and Hyperlophus largely filled up

by the prominent squamosal bulla. The pre-epiotic fossa is pro-

bably homologous with the “lateral cranial foramen” of the

Mormyroid fishes and otopterus. The supratemporal bone les

over the aperture of the fossa in the Clupeoid fishes, but being a

reduced bone it serves less obviously as a cover for the fossa than

does the supratemporal for the foramen in the Mormyride. In

Notopterus the supratemporal bone is reduced im size quite as

much as in the majority of the fishes now under consideration,

and more so than in Dusswmieria. The pre-epiotic fossa is

bounded by the parietal, epiotic, and squamosal, whereas the

foramen of MNotopterus and the Mormyride is set a little lower

down, and is surrounded by the epiotic, squamosal, and exoccipital,

the parietal in these fishes occupying a position nearer the median

plane of the head, and being shut off from the foramen by the

union of the squamosal with the epiotic. The suggestion 1s

strengthened by the fact that in Dusswmieria the bottom of the

fossa is unossified and membranous. If in the preparation of the

skull the membrane be removed, the fossa appears as a large

foramen leading into the cranial cavity.

The lateral temporal groove, situated above and behind the

postfrontal bone, is not well marked im the Clupeoid fishes, with

the exception of Chanos ; in most cases it 1s broad and shallow,

and is barely recognisable as a groove. There is no subtemporal

fossa. The opisthotic bone is as a rule present, but small; it is

wanting in Hngraulis and Coilia.

The bull in the pro-otic and squamosal bones for the lodgment

of cecal diverticula of the swim-bladder are remarkably constant.

Both, however, are wanting in Chanos, and the squamosal bulla

is not present in Clupea sprattus, although it occurs in the other

species of Clupea examined. The auditory fenestra, bounded by

the pro-otic, exoccipital, and basioccipital bones, is also a dis-

tinctive feature of the Clupeoids, although it occurs also in Hyodon,

in which it is bounded by the same three bones. It is wanting

in Ohanos and Coilia. The orbitosphenoid and basisphenoid bones

are variable in size; both are wanting in Chanos. In Clupea,

Hyperlophus, Chatoéssus, and Dussunueria the orbitosphenoid

meets the prefrontal bones.

The eye-muscle canal opens behind in all but Chanos and

Coilia, and except in these two genera and Chatoéssus the para-

sphenoid is produced into a pair of posteriorly directed vertical

laminz of bone. In Chatoéssus the vertical laminz are present

at the sides of the posterior opening of the eye-muscle canal, but

they are not produced backward into freely projecting wings.

490 DR. W. G. RIDEWOOD ON THE CRANIAL [Decrees

Not one of the eleven genera under consideration has parasphe-

noidal teeth, and only Coilia, Dussumieria, Clupea harengus, and

Clupea sprattus have teeth on the vomer.

The post-temporal bone has an epiotic limb in all, and a deep

ov opisthotic limb in all but Coilia. In Chatoéssus the union

between the post-temporal bone and the epiotic is more of the

nature of a synovial articulation than a ligamentous connection.

The supratemporal bone is of the same character in all, and in

all but Coilia the sensory canal that it carries is trivadiate. A

subtemporal or supraopercular bone is present in Chanos.

The nasal bone is small and loosely embedded in the skin, and

there are from five to eight bones disposed around the eye. In

Chatoéssus and Chanos the premaxilla bounds the upper border of

the mouth; in the other nine genera the gape is bounded above

by both premaxilla and maxilla, although in Pristigaster and

Hyperlophus the maxilla takes buta smallshare. In Chirocentrus,

Clupea, Pellona, Pristigaster, Dussumieria, Engraulis, and Coilia

both maxilla and premaxilla bear teeth, although in Pellona the

teeth are vestigial: in Pellonula the premaxilla bears teeth, but

not the maxilla; in Chanos, Chatoéssus, and Hyperlophus both

premaxilla and maxilla are edentulous. The premaxilla is firmly

united to the maxilla by fibrous tissue in Chirocentrus. Two

surmaxille are present on each side of the head in all but //yper-

lophus, Chatoéssus, and Chanos ; Hyperlophus and Chatoéssus have

one, and Chanos none.

The angular is a distinct bone in all but Hngraulis and Coilia.

The einglosiee il part. of the articular can be recognised, but in no

case is it separate from the ectosteal part. A small sesamoid

articular is found in Chirocentrus, Clupea finta, and Chanos.

The hyomandibular articulates with the cranium by two heads in

Clupea finta, Hyperlophus, Chatoéssus, Dussumierca, and Engraulis,

and by a single head in Chirocentrus, Clupea harengus, Pellona,

Coilia, and Chanos. In Ooilia the single head present 1s probably

equivalent to the anterior of the two heads found in Eng graulis.

In Chanos alone of the genera under consideration is the quadrate

separated from the symplectic and metapterygoid. Teeth are

borne by the palatine bone in Chirocentrus (4 or 5 minute teeth),

Clupea sprattus (a row of teeth), Clupea harengus (2 or 3 minute

teeth), Pellona, Pellonula, Pristigaster, Dussumieria, Engraulis,

and Coilia; the palatine is edentulous in Chanos, Chatoéssus,

LTyper lophis, and Clupea jfinta. The ectopterygoid is toothed in

Pellona, Dussumieria, Hngraulis, and Coilia, but not in Chiro-

centrus, Clupea, Pelloniale, Pristigaster, Hyperlophus, Chatoéssus,

and Chanos; the entopterygoid is toothed in Clupea sprattus (a

few teeth), Pellona, Pellonula, Pristigaster, Dussunieria, and

Engraulis, but not in the other forms examined.

The branchiostegal rays are most numerous in Dussumieria (13),

Engraulis (11), Coilia (11), and are fewest in Chanos (4). Chiro-

centrus has 8, Clupea 7 or 8, Chatoéssus 6, Hyperlophus 5.

The interhyal is bony in all, except, perhaps, in Chanos. In all

1904. ] OSTEOLOGY OF CLUPEOID FISHES. 491

of the forms studied there are two hypohyals on each side, and

the lower of the two is larger than the upper. The first pharyngo-

branchial ts ossified in Chirocentrus, but remains cartilaginous in

the other genera; the ossified ligament known as the spicular

bone is present in all except Chirocentrus. In Hngraulis and

Coilia the second hypobranchials are fused with the sides of the

second basibranchial, and in Chanos the third hypobranchials are

fused with the sides of the third basibranchial, but in none of the

three cases ave the sutures obliterated.

COMMENTS ON THE SKULL OF THE CLUPEOID Fisues.

Of the eleven genera the skulls of which have been described

in the previous pages the greatest interest centres around Chanos

and Chirocentrus. As regards the others, the craniological

characters are such as would justify the placing of Engraulis and

Coilia in one family, the Engraulide, and Clupea, Pellona,

Pellonula, Pristigaster, Hyperlophus, Chatoéssus, and Dussumieria

in another, the Clupeidee.

Both Hngraulis and Coilia have a large gape, a backwardly

thrust quadrate bone, and a large and prominent mesethmoid.

The skull of Coidlia differs from that of Hngraulis in several

respects, but these are all of such a nature as might be explained

by high specialisation; such characters, for instance, are the

reduction of the temporal foramen, the obliteration of the pre-

eplotic fossa, the absence of the auditory fenestra, the absence of

paired posterior wings of the parasphenoid and the closure of the

eye-muscle éanal, the absence of the opisthotic limb of the post-

temporal, and the loss of the posterior of the two heads by which

the hyomandibular articulates with the cranium. The paradoxical

extension of the maxilla behind the mandibular articulation is

foreshadowed in some species of Hugraulis, e. g. LH. mystax and

Hi. setirostris. .

The skull of Chatoéssus conforms with the Clupeoid type, in

spite of certain aberrant features, such as the absence of projecting

wings from the back of the parasphenoid, the intimate articulation

of the epiotic limb of the post-temporal with the cranium, the

small size of the mouth, the bounding of the mouth above by the

premaxilla alone, the absence of teeth, and the loss of one of the

surmaxille. The characters of the skull do not warrant the

separation of Chatoéssus from the Clupeide to constitute a distinct

family, although possibly on other grounds the action of Gill

(Smithsonian Miscell. Coll. No. 247, 1872, p. 17) and Jordan and

Gilbert (Bull. U.S. Nat. Mus. No. 16, 1882, pp. 262-274) may

prove to be justifiable.

Chirocentrus agrees so closely in the structure of its skull with

the Clupeide, that appeal must be made to other organs of the

body for evidence to support the views of those who would make

of it a distinct family, the Chirocentride (e. g,, Valenciennes,

Hist. Nat. Poiss. xix. 1846, pp. 150-168; Kner, Reise der

492 DR. W. G. RIDEWOOD ON THE CRANIAL | Dec. 13,

Fregatte Novara, Zool. 1. 1869, Fische ; Cope, Trans. Amer. Phil.

Soc. n.s. xiv. 1871, p. 455; Gill, 1. c. p. 17; Smith Woodward,

Brit. Mus. Cat. Foss. Fishes, iv. 1901). Their justification appears

to lie mainly in the fact that Chirocentrus possesses in its intestine a

spiral valve which is not present in Clupea and its allies (Valen-

ciennes, l. c. p. 160 and pl. 565), in the absence of ceca (ibid.

p. 162), and in the presence of a pseudobranch in Chirocentrus

and its absence from most Clupeoids (Miiller, Abhandl. Akad.

Wiss. Berlin, 1844 (1846), p. 191). The only eraniological

differences worth mentioning are the considerable depth of the

posterior temporal groove of Chirocentrus, the small size of the

orbitosphenoid, the firm union between the premaxilla and maxilla

(Valenciennes, /. ¢. pp. 150, 152, and 154), the move or less com-

plete concealment of the symplectic (Boulenger, Ann. Mag. Nat.

Hist. (7) xiii. 1904, p. 164), the bony nature of the first pharyngo-

branchial, and the absence of a spicular bone. The value of the

evidence of the last two items is certainly not great, for except

when (as in Zlops) both ossified first pharyngobranchial and

spicular bones are present, it is not possible to deny absolutely that

what appears to be the first pharyngobranchial bone is not the

spicular bone which has become shortened and thickened and more

forwardly directed than usual.

The resemblances existing between the skeleton of Chirocentrus

and that of such extinct forms as Portheus, [chthyodectes, and

Saurodon, which attained their maximum development in Creta-

ceous times, suggest that the former genus is a survival of an

ancient type (see Smith Woodward, /. ¢. p. vii); but the teeth of

the existing Chirocentrus are not lodged in distinct sockets as are

those of the Saurodontide.

As regards Changs, the evidence of the skull favours the view

of separating the genus from the Clupeide, and of according it a

family rank. Chanos has experienced a variety of treatment at

the hands of taxonomists. It was first regarded as a species of

Mugil (Forskaél, Desc. Anim. 1775, p. 74; Gmelin, Syst. Nat.

Linn. i. 3, 1788, p. 1398), and later as a species of Lewciscws (Gray

and Richardson, Dieffenbach’s ‘Travels in New Zealand,’ 1843,

ii. p. 218). Valenciennes (Hist. Nat. Poiss. xix. 1846) placed it

with Gonorhynchus among the ‘‘ Malacoptérygiens intermeédiaires

entre les Brochets et les Clupes.” Giinther (Brit. Mus. Cat. Fishes,

vil. 1868) placed it in a group ‘“* Chanina” of the family Clipeidee,

and Kner (Reise der Fregatte Novara, Zool. 1. 1869, Fische), Cope

(Trans. Amer. Phil. Soc. n. s. xiv. 1871, p. 455), and Gill (Smith-

son. Miscell. Coll. No. 247, 1872, p. 17) separated it from the

Clupeidee and placed it in a family of its own.

The inclusion of Chanos within the family Albulide, a step

which has commended itself to so experienced an ichthyologist as

Smith Woodward (Brit. Mus. Cat. Foss. Fishes, iv. pp. 60 and

64), is justified in so far as the posterior temporal fosse are rooted

over, which is not the case in any other Clupeoid fishes, and in the

presence of a well-marked lateral temporal groove, partially roofed

1904. | OSTEOLOGY OF CLUPEOID FISHES. 493

over ; also in that the bulle in the squamosal and pro-otic bones

present in other Clupeoid fishes for the lodgment of cecal

diverticula of the swim-bladder are wanting, as also are the

auditory fenestra, the pre-epiotic fossa, and the temporal foramen

also in the fact that the parasphenoid is not produced into a pair

of posteriorly directed wings. On the other hand, such features

as the reduction in the size of the mouth are as likely to be due to

convergence as to genetic affinity.

The absence from the skull of Chanos of the orbitosphenoid and -

basisphenoid bones is a mark of degradation, and supports neither

the hypothesis of a natural affinity between Chanos and the

Albulide, nor that which wouldassociate Chanos with the Clupeidee ;

and the same may be said of the absence of teeth, the reduction in

the number of branchiostegal rays, and the absence of surmaxille.

The closure of the posterior outlet of the eye-muscle canal is a

mark of specialisation, and in this respect also Chanos differs from

both the Albulide and the Clupeide (except Coilia). The absence

of a subtemporal fossa, which is present in the Elopide and

Albulidee, but absent from the Clupeoid fishes, is an argument

against Woodward’s view; and again, the parietals meet one

another in Albwla, whereas in Chanos they are in reality separated

by the supraoccipital, although a secondary union may be brought

about by means of the commissural sensory-canal bones which fuse

with the parietals.

Abbreviations employed in the Text-figures.

af, auditory fenestra. | mx, maxilla.

al, alisphenoid. 2, Nasal.

an, angular. op, opisthotic.

ar, articular. ope, opercular.

bb, dentigerous plate covering the or, orbitosphenoid.

basibranchials.

bo, basioccipital.

br, branchiostegal rays.

bs, basisphenoid.

cb, cevatobranchial.

ch, cevatohyal.

cor, circumorbital bones.

ct, cartilage.

d, dentary.

eb, epibranchial.

ecp, ectopterygoid.

eh, epihyal.

enp, entopterygoid.

e0, exoccipital.

ep, epiotic.

f; frontal.

gh, 2lossohyal.

hb, hypobranchial.

hh, hypohyal.

hm, hyomandibular.

ih, mterhyal.

top, interopercular.

me, mesethmoid.

mpt, metapterygoid.

p, parietal.

pb, pharyngobranchial.

pef, pre-epiotic fossa.

pl, palatine.

pm, premaxilla.

pof, posttrontal.

pop, preopercular,

prf, prefrontal.

pro, pro-otic.

ps, parasphenoid.

pt, post-temporal.

ptf, posterior temporal fossa.

g, quadrate.

sar, sesamoid articular.

sm, surmanilla.

soc, supraoccipital.

sop, subopercular.

sp, spicular bone.

sq, Squamosal.

sé, supratemporal.

sy, symplectic.

tf, temporal foramen.

v, vomer.

‘ very Hak i

pe ge Shs

rt caduiy pM

Abranis, 62.

Acanthias

vulgaris, 23.

Acanthodrilus

uliginosus, 262.

Acanthonycha

antennata, 396, 405.

constatipennis, 396,

404,

dimidiata, 396, 403.

geniculata, 396, 403.

peruana, 396, 403.

stali, 396, 404.

Acerina, 28.

Acipenser, 33.

Acontias, 156, 157.

Mega, 312.

Agasicles, gen. nov., 396,

400.

vittata, 396, 401.

Agouti

paca, 159.

Alausa

tyrannus, 454.

Albula, 39, 41, 52, 53, 54,

55, 56, 58, 60, 61, 62,

64, 66, 69, 70, 71, 72,

73, 75,76, 77, 78, 79, |

80, 448, 493.

conorhynchus, 47, 48, |

50, 52, 55.

Alepecephalus, 63, 65, 69,

71, 78, 79, 80.

Alestes, 58, 62.

Alosa

vulgaris, 80, 459.

Amia, 37, 39, 41, 54, 56,

57, 59, 60, 63, 64, |

68, 77.

Amphibolurus

barbatus, 83.

Anabas, 81.

Anomalurus

beldeni, 191.

INDEX.

Anomalurus

erythronotus, 191.

fraseri, 185, 190, 191,

192.

— nigrescens, 190.

Anthropopithecus, 413,

418.

gorilla, 178.

schweinfurthi, 347.

troglodytes, 177.

Aphaneramma, gen. nov.,

NGOs Ne, Wey).

rostratum, 173,

176.

175,

Aplysia, 297.

Arapaima, 56, 57, 59, 60, |

61, 62, 63, 64, 67, 69, |

1; Ts 72, 73, 0S, 7 |

Us Wes

gigas, 5D.

Ardea

chrysopelargus, 200.

Argilophilus, 259.

Argynnis

clara, 137.

— clarina, 137.

claudia, 136, 141.

Ascaltis, 384.

botryoides, 386, 387,

389, 391, 392, 393,

394.

Ascandra

armata, 360, 361, 371.

botryoides, 395, 374,

387, 392.

botrys, 355, 386, 387,

391, 392, 393, 394.

complicata, 305, 3861,

366, 3/0, 3/1, 372

38).

contorta, 361, 371, 372,

373, 385.

corallorrhiza, 374, 384.

ellisit, 392.

olZ, |

Bathythrissa, £

Aseandra

fabricii, 361, 370.

nitida, 355, 386, 392.

pinus, 355, 361, 362,

368, 372.

solandervi, 392.

tenuis, 374.

variabilis, 351, 3874,

378, 380, 385, 386.

(Clathrina) contorta,

373.

Ascortis

corallorrhiza, 374, 384.

Jabricit, 861, 370.

Asculmis

armata, 360, 371.

— norvegica, 371.

— pocillum, 3871.

Ascyssa, 390.

acufera, 371.

variabilis, 384.

Asellota, 310, 313, 314.

Asellus, 302, 305, 306,

307, 308, 311, 312,

313, 314.

aquaticus, 305, 331.

Aspheera, 402.

Attheyella

natalis, 124, 128.

Babirussa, 197.

Balearica

cecilia, 203, 204, 205.

chrysopelargus, 131,

200.

gibbericeps, 201, 202,

204.

pavonina, 200, 201, 203,

204, 205.

regulorum,

204, 205.

201, 202,

2, 53, 54,

55, 76.

496

Bathythrissa

dorsalis, 52, 55.

Belenois

mesentina, 142.

Belone, 72.

Bensonia

camura, +41.

nepalensis, 441, 447.

Blepharida

flavicostata, 396, 406.

marmorata, 407.

multimaculata, 396,

407.

Boa, 107, 118.

constrictor, 116, 148.

Boocercus, 196.

Bornella

adamst, 100.

arborescens, 100, 102.

calcarata, 100.

caledonica, 100.

digitata, 100, 101, 102,

103, 104, 105.

excepta, 100, 102, 108,

104, 105.

hancockana, 100.

hermannt, 100.

sempert, 100.

simplex, 100, 103, 105.

Bos

caffer brachyceros, 131.

— mathewsi, 165.

— nanus, 131,

165.

— planiceros, 131, 164.

(Bubalus) caffer, 164.

(—) caffer mathewsi,

165.

Botia, 62.

Broteas, 301.

Bubalus

centralis, 347.

pumilus, 131.

164,

Calcispongia

botryoides, 386.

Caliphylla, 291, 292.

Canchroma

cochlearia, 82.

Capitosaurus

arenaceus, 173.

fronto, 173.

nasutus, 173.

robustus, 178.

stantonensis, 170, 172,

173, 176.

Oaranx, 56.

Carcinoscorpius

rotundicauda, 181.

Carpophaga

concinna, 178.

INDEX.

Catopsilia

florella, 142.

Catostomus, 58.

Causus, 117.

rhombeatus, 116, 148.

Cephalophus

brooket, 192.

melanorheus, 185, 193.

ogilbyt, 185, 192.

Ceratohyla

bubalus, 106.

Ceratophyllidia, 279,

280.

Ceratosoma

cornigerum, 87, 104.

Cercopithecus

sp., 178.

burnetti, 185.

campbelli, 185.

erythrotis, 185, 186.

martini, 185, 186.

nigripes, 178.

pogonias, 185.

preussi, 185, 186.

schmidti, 178.

wolfi, 178.

Cervus

canadensis, 131.

elaphus, 133.

— harbarus, 130.

eustephanus, 179.

hortulorum, 88.

Cetorhinus, 29, 31, 33.

maximus, 28, 29.

(Selache) maximus, 25.

Cheetodon, 29.

Chalcides

lineatus, 145, 146.

Chameleon, 6, 14, 16, 22,

vulgaris, 7, 8, 12.

Chanos, 58, 59, 60, 61,

62, 65, 66, 67, 69, 70,

71, 78, 74. 75, 76, 78,

79, 80, 448, 482, 484,

488, 489, 490, 491,

492, 498.

salmoneus, 55,482, 488,

485, 486, 487.

Chatoéssus, 58, 59, 60, 61,

62, 65, 66, 70, 71, 73,

74, 78, 79, 80, 448,

463, 467, 468, 488,

489, 490, 491.

erebi, 55, 463, 464, 465,

466, 468.

Chersydrus, 149.

Chilonyeteris, 5.

Chimera, 28.

Chimpanza

gorilla, +16.

Chirocentrus, 58, 60, 61,

62, 65, 70, 71, 75, 78,

79, 80, 448, 451, 488,

490, 491, 492.

dorab, 55,'78, 448, 449,

451, 452, 453.

polyodon, 451.

Chlamydosaurus

kingii, 82.

Chloéphaga

antarctica, 178.

Choleepus

didactylus, 160.

Chromodoris

sp., 87.

Cicinnurus

regius, 178.

Cirolana, 312, 313.

Citharinus, 58.

Clathrina

sp., 372.

contorta, 352, 362.

cortacea, 372, 375, 376.

Cliona

celata, 355.

Clupea, 23, 58, 60, 61, 62,

66, 70, 71, 72, 74, 78,

79, 448, 451, 459,

460, 461, 462, 463,

469, 470, 473, 474,

477, 488, 490, 491,

492.

alosa, 80, 459.

jinta, 55, 65, 70, 74, 80,

453, 454, 455, 456,

457, 458, 459, 490.

harengus, 61, 70,74, 80,

454, 455, 456, 457,

458, 459, 460, 463,

490.

mattowocca, 454.

pilchardus, 454.

sapidissima, 454.

spratius, 62, 454, 456,

457, 458, 489, 490.

thryssa, 454.

vulgaris, 80, 459.

Cobitis, 58, 62.

Coecidothea, 507,

314.

Ceelopeltis

monspessulana, 107.

Coilia, 61, 62, 68, 65, 69,

70, 71, 78, 74, 75, 76,

78, 79, 448, 488, 489,

490, 491, 493.

nasus, 55, 477, 478,

479, 481.

Colias

berylla, 139, 141.

edusa, 140.

308,

Colias

eogene, 140.

— leechi, 134, 140, 141.

—- miranda, 141.

erschoffi, 140.

hyale, 140,

nina, 140, 141.

stoliczkana, 141.

Colobus

abyssinicus, 177.

pennanti, 185.

polycomus, 185.

satanas, 185, 186.

Coluber, 112.

esculapti, 116.

catenifer, 112, 1138.

melanoleucus, 148.

Conothoa, 219.

Coregonus

albula, 28.

Jera, 23.

pollan, 61.

Coronella

getwla, 107, 119, 148.

Crepidodera

longicornis, 396, 412.

peruviana, 412.

Cricetomys

gambianus, 185.

Cricotus, 174, 175.

Crocidura

morio, 190.

poensis, 185, 189.

Cromeria, 79, 80.

Cryptodrilus, 259, 260.

Cyclops

aati 123:

jfimbriatus, 123.

gibsoni, 123, 127.

gracilis, 128.

leuckartt, 122.

pusillus, 122, 127.

scourfieldi, 122.

varicans, 123.

Cyclotosaurus

robustus, 173.

Cyerce, 291, 292.

elegans, 268, 298.

Cymothoa, 313.

Cynelurus

jubatus, 82.

Cynopterus, 188.

'ypria

castanea, 125, 128.

lacustris, 125.

ophthalmica, 125.

Cyprinus, 58, 62.

Cypris

aratra, 125, 128.

inermis, 125, 128.

ornata, 126, 128.

Proc. Zoot. Soc.—1904, Vou. Il. No. XX XIT

INDEX.

Daphnia

sp., 300.

magna, 300.

Dasyprocta, 160.

Dasyurus

viverrinus, 2.

Dentrotion, 302.

Dermochelys

coriacea, 2.

Desmosoma, 302, 308,

313.

Dicotylichthys, 77.

Didelphys

quica, 160.

Dinodriloides, gen. noyv.,

221, 226.

beddardi, 222, 226, :

228.

Dinodrilus, 221, 228, 229, |

230. |

Dinomys

branickii, 158-162.

Diodon, 77.

Diplomystax, 462.

Diplomystus, 462, 465.

Diporocheeta, 220. |

Disonycha

albicincta, 396, 402.

amazonica, 396, 401.

peruana, 396, 402.

Dobsonia, 188. |

Doridopsis, 270, 272, 280. |

brockii, 279.

clavulata, 268, 278.

denisoni, 268, 277, 278. —

gemmacea, 277.

nicobarica, 278.

nigra, 268, 275, 277.

pudibunda, 208, 274.

rubra, 268, 279.

spiculata, 268, 274.

tuberculosa, 268, 278,

284.

Doris

lacera, 270.

Doto

sp., 285.

africana, 268, 285.

fragilis, 235.

indica, 285.

pinnatifida, 285.

Dussumieria, 60, 61, 62, |

67, 70, 71, 76, 78, |

448, 488, 489, 490,

491.

acuta, 55, 468, 469.

470, 471. |

Echinopleura, 308.

Ketoeyclops, gen.

122, 124.

novy., |

AQT

Kctocyclops

rubescens, 124, 127, 128.

Hlaphodus

cephalophus, 166, 168,

169.

ichangensis, 168, 169.

michianus, 166, 167,

168, 169.

— fociensis, 169.

Elaphurus

davidianus, 83,178,179.

Elops, 37, 41, 44, 45, 47,

54, 55, 56, 58, 60. 61,

62, 64, 67, 70, 71, 72,

73, 74, 75, 76, 77, 79,

80, 448, 492.

saurus, 38, 40, 42, 55.

Hlysia

dubia, 268, 297, 298.

Saustula, 268, 295.

marginata, 268, 296,

297, 298.

nigrocincta, 296, 297.

migropunctata, 296.

(Pterogastron) margi-

nata, 297.

Hlysiella, 297.

Engraulis, 60, 61, 62, 63,

65, 69, 70, 71, 78, 74,

75, 76, 78, 79, 80,

448, 459, 474, 477,

478, 480, 488, 489,

490, 491.

encrasicholus, 5d, 472,

473, 474, 475.

nuystax, 480, 491.

setirostris, 480, 491,

Hnhydrina

valakadien, 151.

Ephippus, 485.

Hpomophorus, 188.

Equus

zebra, 342, 343, 344.

Ercolania

siottt, 290.

Eryx, 107, 108, 110, 112,

WS; LIA 19s 121,

149.

conzceus, 111, 112, 113,

WA GS Wats

119, 120, 121, 148.

Jjaculus, 108, 109, 110,

V1, 113; 114 115,

WING. The TLD). ex}.

121, 148.

gold, Milt, We te}.

ING, tale, ILS), 11940),

121

Ksox, 28.

Kstheria

elizabethe, 299.

498

Hudyptes

antipodum, \77.

chrysocome, \77.

Hunectes, 107, 110, 115, |

TiS, WO), Wen.

murinus, 110, 113.

Huplecta

layardi, 447.

pulchella, 447.

Eurycope, 302, 311, 512.

gigantea, 309, 310, 381.

Kutypkeeus, 230.

Facelina

cyanella, 289.

lincata, 268, 288, 297, |

298.

Faunus, 421.

Felis

sp., 162.

catus, 162.

demon, 162, 163.

leo, 144.

Fiona

marina, 286.

nobilis, 286.

pinnata, 268, 285, 286.

Fistularia, 65.

Fryeria, 279.

Funisciurus

erythrogenys, 185, 192.

poensis, 185, 192.

Furipterus, 6.

Galago

alleni, 185.

demidoffi, 186, 187.

— poensis, 184, 185,

186.

elegantulus, 189.

Gasterotokeus, 65.

Gastrosteus, 63, 74.

Gazella

arabica, 849.

cuviert, 347, 348, 349.

dorcas, 347.

leptoceros, 130.

merrilli, 347, 348, 349. |

Genetta

poensis, 185.

Gennzus

lineatus, 130.

nyethemerus, 130.

Girafta

camelopardalis, 339.

— capensis, d41.

Glauconycteris

poensis, 185.

Goniodoris, 292.

INDEX.

Gonorbynchus, 58, 59,

60, 63, 65, 66, 69,

Wy We Wes, Te 15,

78, 80, 480, 492.

Gorilla, 413, 414, 429,

439.

beringeri,

421.

castaneiceps, 414, 415,

416.

diehli, 415.

gorilla, 414, 415, 416,

417, 439.

— dichli, 418, 420.

— matschiei, 414, 415,

416, 418, 419, 440.

manyema, 419.

Grammicolepis, 56, 60.

Grantia

botryoides, 351, 360,

313, 885, 386, 393,

394.

— himantia, 373, 375.

compressa, 389, 387.

licberkiihnii, 386.

nived, dol.

Gymnarehus, 56, 64, 65,

71, 72, 76.

niloticus, 5d.

Harpyia, 188.

Hatteria, 155.

Helictis

personata, 347.

Hermea

dendritica, 290.

minor, 290.

Hervia, 287.

lineata, 268, 286, 288,

297.

Heterodon

platyrhinus, 148.

Heterotis, 60, 62, 68, 64,

GY, WO, al, 728 745

75, 77, 78, 80, 81.

niloticus, 55.

Hexabranchus

digitatus, 271.

lacer, 268, 270.

—— faustus,

212:

marginatus, 268,

271, 272.

— mocbii, 268, 271,

272.

Hipposiderus

Fuliginosus, 185, 188.

Hippuriphila

catharine, 396, 412.

Hoplochetella, 230.

416, 418, |

268, 271, -|

| Ischnosoma,

Hydrocherus

capybara, 178.

Hydrocyon, 62.

Hydrophis

schistosa, 151.

Hydrus

platyurus, 147-154.

Hyla

evansi, 106.

goeldi, 106.

Hylobates, 413.

lar, 178.

Hylocherus, 193, 196,

197, 198, 199.

meinertzhagent, 194,

196, 199.

| Hyodon, 56, 60, 61, 62,

63, 64, 67, 70, 71,

72, 73, 74, 77, 78,

79, 80, 81.

alosoides, 5d.

Hypanartia

hippomenc, 142.

scheencia, 142.

Hyperlophus, 448, 488,

489, 490, 491.

copti, 462.

Hypogzeon

orthostichon, 255.

Hypolimnas

misippus, 142.

Hypsignathus

monstrosus, 185, 187.

Taera, 311.

Janira, 302, 503, 304,

307, 309, 311, 312,

330, 331.

maculosa, 331.

Ichthyodectes, 492.

Idothea, 3138.

| Iguana, 7, 8, 9, 10, ii,

Tolanthe, 311.

302, 312,

313.

Ischyrocerus, 327.

Istieus, 52.

Kerivoula, 4.

poensis, 188.

Kobus

kob, 177.

unctuosus, 177.

| Labeo, 62.

Lacerta, 117.

chalybdea, 338.

defilipptt, 333.

depressa, 332, 333, 334,

307, 308, 339.

Lacerta

depressa modesta, 336.

— rudis, 332, 336, 337,

338, 339.

muralis, 332, 333, 395,

336, 337.

— defilippii, 337.

— fusca, 337.

— persica, 337.

— saxicola, 337.

oxycephala, 337.

portschinskii, 337.

saxicola, 333, 337.

taurica, 332.

typica, 339.

— bedriage, 333.

vivipara, 338.

Lactica

argentinensis, 396, 400.

baeri, 396, 399.

boliviana, 398.

clara, 397.

clypeata, 399.

decorata, 396, 397.

discicollis, 396, 399.

dives, 397, 400.

elegantula, 398.

maculicollis, 396, 398.

marginata, 397.

aicotineg, 396.

posticata, 396, 398.

rufobasalis, 398.

rufobrunnea, 396, 399.

Lagomys

alpinus, 207.

auritus. 216.

cinereoflava, 208.

curzonie, 209, 214.

dauricus, 216.

erythrotis, 210.

ferruginea, 208.

fusca, 208.

griseus, 217.

hodgsont, 218.

hyperboreus, 208.

koslowt, 212.

ladacensis, 208.

littoralis, 208.

macrotis, 216.

melanostoma, 215.

nepalensis, 218.

normalis, 208.

ogotona, 210.

pallasi, 210.

pusillus, 215.

roylei, 217.

rufescens, 211.

rutilus, 211, 220.

tibetanus, 218.

Lamellidoris, 292.

Lates, 60,

INDEX.

Lepidosiren, 23.

paradoxa, 159.

Lepidosteus, 30, 56, 68

osseus, 56.

viridis, 56.

Lepus

alpinus, 207.

dauricus, 210.

hyperboreus, 208.

ogotona, 210.

pusilla, 213.

Leuciscus, 62, 492.

Leuconia

nivea, 851, 352.

somesit, 352, 353, 374

378, 382, 384, 385.

Leucosolenia, 349-396.

ameboides, 360, 371.

arachnoides, 384.

hotryoides, 351, 352

353, 354, 355, 357

360, 368, 375,

383, 385, 386, 387

389, 391, 392, 393

394, 395, 396.

cervicornis, 384.

complicata, 352, 353

355, 360, 361, 362

363, 365, 366, 368

SiO), Bi, BIZ, Blo

375, 376, 3838, 384

392, 393.

confervicola, 384.

contorta, 302,

363, 368, 372, 585

392.

fabricii, 353, 360, 361

370, 371.

granti, 386.

hispida, 371.

hispidissima, 384.

lieberkiihnii, 359,

358.

pinus, 361, 366.

variabilis, 352, 393

357, 359, 3638, 373

374, 375, 376, 378

380, 382, 383, 384

389, 391, 596.

(Leucelia) complicata,

360.

(Leneeria) botryotdes,

Syeyn

386.

(Leuciria)

373.

Leydigia

acanthocercoides, 300.

africana, 300, 301.

Limnas

chrysippus, 142.

variabilis,

> |

;

499

Limnetis

wahibergi, 299.

Lioheterodon

madagascariensis, 117,

148.

Lophius, 58, 63.

Lophophorus

impeyanus, 130.

Loris

gracilis, 346.

— typicus, 346.

—- zeylanicus, 346.

Lota, 25.

Lovenula

Jalcifera, 300, 301.

lamellata, 301.

mea, 300, 801.

Lucioperea, 25.

Lutra

capensis poensis, 185.

Tuvarus, 53.

Lyceeaa

ariana, 138.

— arene, 137, 141.

pherctes asiatica, 138.

— pharis, 158, 141.

(Niphanda) marcia,

139, 141.

(—) tessellata, 139.

(Zizera) zera, 138, 141.

Macrobates, 421.

Macrochlamys

anone, 443.

aspides, 4405.

atoma, 443, 447.

chaos, 444, 447.

choinix, 446.

curvilabris, 445, 447.

glauca, 442.

hypoleuca, 445.

kuluensis, 442, 447.

kumahensis, 445.

notha, 444, 447.

noxia, 444, 445, 447.

nuda, 442.

patens, 445, 447.

petasus, 444.

prava, 448, 447.

pscudochoinix,

447.

rutila, 443, 447.

seguax, 443.

spreta, 445, 447.

subjecta, 444.

subpetasus, 445.

superflua, 442, 447.

Macropus

antilopinus, 129.

brunti, 178.

dorsalis, 179.

446,

500

Macrostylis, 302,311,313. |

Macrothrix

affinis, 127, 128.

laticornis, 127.

Madrella

Jerruginosa, 269, 298.

Maneasellus, 307,

314,

Manis

tricuspis, 185.

Maoridrilus, 220,

229.

manwiensis, 222, 223,

D228.

uliginosus,

Marionia

223,

308,

INDEX.

291

aul,

Microscolex,

239:

hempeli, 238.

huttoni, 239.

monticola, 239.

nove-zealandie, 238.

Mimetillus, 188.

moloneyt, 185, 188.

Moina

belli, 299, 301.

wierzejskii, 299.

Molossus, 189.

Mormyrops, 60, 64, 79.

deliciosus, 55, 67.

Mormyrus, 60, 79.

oxyrhynchus, 67.

Neodrilus, 221, 229.

Nephrica, 406.

Nestor

notabilis, 82.

Notiodrilus, 220, 229,

230, 260.

aucklandicus, 261,

262.

Notodoris

citrina, 84, 85.

gardineri, 84.

minor, 84, 105.

Notopterus, 60, 61, 62,

63, 64, 65, 67, 70,

es, 13, TE 0:

77, 78, 489.

sp., 93, 99.

albo-tuberculata,

97, 98, 99.

arborescens, 94, 95,

Mugil, 59, 492.

Munidopsis

polymorpha, 82.

Munna, 302. 311.

kapirat, 55.

Notoscolex, 256, 257,

258, 259, 260, 261.

orthostichon, 256, 257.

Sie | boechi, 311. Nycteris

chloanthes, 99. Munnopsis, 302, 308, hispida, 185, 188.

levis, 94, 105. 311. Nycticebus

pellucida, 94. | — typica, 309. menagensis, 31d.

ramosa, 94, 95, 96. Mus tardigradus hillert, 345,

virescens, 99. alleni, 185, 192. 346.

viridescens, 94, 98. | decumanus, 133. — javanicus, 345.

Megalops, 37, 44, 45, 54, | musculus, 133. — malayanus, 345,

55, 56, 59, 60, 61, rattus, 192. 346.

62, 63, 64, 65, 67, | tullbergi, 185, 192. — natune, 345.

70, 71, 72, 73, 74, | Mustelus, 29. — typicus, 345.

76, 77, 78, 79, 80, | Mycetes Nyetinomus, 189.

448, | beelzebul, 129. brachypterus, 185.

cyprinoides, 43, 46,55. | Myiadectes pumilus, 185.

priscus, 45.

Megascolex, 261.

Megascolides, 256, 257,

258, 259, 260, 261,

unicolor, 129.

Myosorex, 190.

Myotis, 4.

Mystacops, 6.

| Nyctymene, 188.

Ochotona, 205.

262. Myzopoda alpina, 206, 207, 208,

ambialis, 261. aurita, 2-6. 219.

australis, 257, 256, aurita, 206, 217.

cinercoflava, 206, 214.

curzome, 206, 207,

215, 216, 217, 220.

daurica, 206, 207, 215,

260, 262, 263. |

cameroni, 258.

dlawarre, 258, 262.

insignis, 258.

Naia

tripudians, 148.

Nannoniscus, 313.

Meletta | oblongus, 310. 216, 220.

thryssa, 454. | Natalus, 3, 4, 5. erythrotis, 206, 207,

Melibe Nembrotha 210, 211, 219, 220.

finbriata, 87. affinis, 90, 92, 105. | ferruginea, 206.

Melitza cerulea, 91. fusca, 206.

sikkimensis, 136.

sindura, 136. | 105.

tibetana, 135, 141. | diaphana, 90.

Metopa, 327. | gracilis, 90. |

Microcystina gratiosa, 90, 92, 93. |

bintennensis, 446. kubaryana, 90.

perfucata bintennensis, morosa, 90.

446. | nigerrima, 90, 91, 92.

rinki, 446. | Nemorhedus

shevaroyana, 446,447. | goral, 177.

stuart, 446, 447. Neoceradotus, 23,

cristata, 90, 91, 92, grisea, 206, 217.

hodgsont, 206,

213, 218, 219.

hyperborea, 206, 207,

208.

koslowi, 206, 207, 212,

214, 219.

ladacensis, 206, 207,

208, 209, 210, 212,

213, 219.

littoralis, 206, 208.

207,

Ochotona

macrotis, 206, 207, 211,

216, 217.

melanostoma, 206, 207,

215, 216, 220.

nepalensis, 206, 218.

normalis, 206.

ogotona, 206, 207, 210,

216.

pusilla, 206, 207, 213,

220.

roylei, 206, 207, 214,

217, 218, 347.

riufescens, 206, 207,

211, 213, 214, 215.

rutila, 206, 207, 210,

211, 217, 219, 220.

wardi, 206, 207, 214.

Octochtus, 220,

230, 260.

michaelsent, 222, 225.

thomast, 226.

Oedionychis, 402.

Okapia

Johnstont, 180.

Olynthium

mitidum, 386.

splendidum, 380.

Olynthus

hispidus, 860.

pocillum, 360.

Ophiophagus

bungarus, 107, 148.

Orodoris

miamirana, 270.

striata, 268, 269.

Oryx

leucoryx, 178.

Osmerus, 28, 49, 61.

Osphromenus, 81.

Osteoglossum, 36, 57, 5

60, 61, 62, 63, 6

66, 67, 69, 70, 7

72, 73, 74, 75, 7

78.

bicirrhosum, 63.

formosum, 63.

leichardti, 55, 63.

Ostracion, 65.

Ourebia

nigricaudata, 177.

Oxygona

amazonica, 396, 411.

capitata, 396, 411.

luridus, 412.

simplex, 412.

Pachyuromys

dupresi, 133.

Pan, 418, 419.

Pantodon, 69, 71, 75.

929 |

a-d, |

INDEX.

Parnassius

acco, 136.

acconus, 135.

delphius, 135.

epaphus sikkimensis,

134.

imperator, 134.

— augustus, 134.

Pellona, 448, 488, 490,

491.

motius, 459.

Pellonula, 448, 488, 490,

491.

vorax, 460.

Pelonia

elegantula, 403.

Pelophilus

(Boa) madagascart-

ensis, 108.

Perameles

obesula, 2.

Perea, 23.

Percichthys, 60.

Petrocephalus,

(ise

bane, 55, 67.

Phacocheerus, 195, 196,

197, 198, 199.

Phasianus

ellioti, 129.

reevesi, 129.

versicolor, 130.

wallichi, 130.

Phelsuma, 7, 16, 20, 21,

22.

madagascariensis, 18,

19.

Pheretima, 261, 262.

Phidiana, 288.

tenuis, 268, 287.

Pholidophorus, 69.

Phractolemus, 69.

ansorgit, 5D.

Phyllidia, 279, 280.

elegans, 280. |

fasciolata, 281. |

nobilis, 268, 280, 281,

282, 283, 297.

— rotunda, 268, 282.

pustulosa, 268, 280, |

283. |

varicosa, 268, 280, 281,

282, 283.

Phyllidiella, 279.

nobilis, 280.

pustulosa, 280.

Phyllidiopsis, 279, 280. |

TO, |

cardinalis, 268, 281.

284.

Phyllobranchus,

Qo |

292, 294. |

501

Phyllobranchus

prasinus, 268, 292,

293, 204.

viridis, 291.

Phylodesmium

hyalinum, 268, 289.

Phyllopteryx, 69.

Pika, 219.

Pithecus, 421.

gesilla, 416.

satyrus bataungtuensis,

435, 456.

— dadappensis, 435,

456.

— genepaiensis,

436.

—- landakkensis, 434,

436.

— rantaiensis, 439.

— skalauensis, 435,

436.

— tuakensis, 435, 486.

sumatranus abongensis,

435, 437.

— deliensis, 435, 437.

Placobranchus, 297.

ocellatus, 268, 294.

Plagiocheta, 220,

260.

montana, 229.

ricardi, 229.

rosst, 229, 261.

sylvestris, 229.

Platyurus

colubrinus, 147-154.

corallinus, 148.

Pleurobranchus

delicatus, 87.

Pleurogonium, 311.

Pleuroleura

alba, 104.

striata, 104, 105.

Plutellus, 220, 259, 261,

262.

intermedius, 245.

Podarcis

depressa, 332.

Pceocephalus

robustus, 177.

Poiana

richardsoni, 185, 190.

Polita

notabilis, 447.

turbinata, 447.

Polyodon

spathula, 22-35.

Polypterus, 77.

Pongo, 414, 421, 439.

pygmeus, 421, 436,

438.

— abelit, 437, 439.

435,

999

229)

502

Pongo

pygmeus agrias, 436,

— bicolor, 437, 439.

— dadappensis, 436,

438.

— genepaiensis, 436,

438

— pygmeus, 436, 438.

— skalauensis, 436,

438.

-— wurmbit, 436, 438.

Portheus, 492.

Potamocherus, 19, 196,

197, 198.

Prasona

peruviana, 396, 407.

Precis

calescens, 143.

natalensis, 143.

octavia, 148.

sesamus, 143.

Pristigaster, 448, 488,

489, 490, 491.

tartoor, 461,

Procavia

dorsalis, 185, 192.

Prochilodus, 62.

Protopterus, 23.

Psephurus, 33.

Pseudanthropus

fuliginosus, 422, 451.

Pseudarachna, 308.

Pseudogona

discoidalis, 396, 409.

militaris, 396, 410.

pallida, 396, 410.

panamensis, 410.

Pteraeolidia

sempert, 83.

Pygopus, 7, 8, 9, 14, 16,

22, 112, 117, 121.

lepidopus, 12, 13, 15,

17, 18.

Python, 107, 110, 113,

116, 121.

bivittatus, 108.

sche, 110, 112, 116,

119.

spilotes, 108, 112, 121,

148.

Quiscalus

versicolor, 2.

Ragadia

crisia, 135.

simplex, 135, 141.

Rallina

canningi, 129.

Regalecus, 58.

INDEX.

Rhacolepis, 54, 67.

Rhinoceros

simus, 1.

Rhinodon

typicus, 23.

Rhinolophus

landeri, 185.

Rhinomus

soricoides, 189.

Rhododrilus, 221, 222,

230, 288, 239, 261.

besti, 222, 235, 236,

237, 239.

edulis, 222, 230, 232,

233, 234, 235, 238,

2B),

huttoni, 239.

minutus, 239.

| Rhopalotoma, 406.

| Rizzolia, 287.

Roceus, 78.

Rousettus

stramineus, 185, 187.

Salmo

Sarcodaces, 67.

Satyrus, 420, 421.

adrotes, 416.

indicus, 420.

Saurodon, 492.

Scaphirhynchus, 35.

Sciurus

punctatus, 185.

rufobrachiatus, 185,

191, 192.

stangeri, 185, 191,

192.

Scomber, 59.

Scotonycteris

bedfordi, 184, 185, 187,

193.

zenkeri, 187, 188.

Scotophilus

temminckii, 5.

Sebastolobus, 78,

Simia, 413, 419, 422.

abelti, 437,

agrias, 436.

aubryt, 429, 430, 459.

caluus, 424.

chimpanse, 428, 430,

433.

Juscus, 426.

koolookamba, 429, 430,

434, 459.

leucoprymnus, 42/7.

pygmeus, 420, 422,

427, 480, 431, 482,

435, 436, 439.

— fuscus, 430.

—- leucoprymnus, 430.

Simia

pygmeus raripilosus,

422, 428, 430.

rariptlosus, 428.

satyrus, 129, 420, 421,

422, 423, 424, 426,

429, 430, 439.

— marungensis, 429,

430, 431.

— schweinfurthi, 427,

429, 430.

schweinfurthi, 426.

troglodytes, 422, 430.

vellerosus, 415, 425,

496, 429, 430, 439.

— fuliginosus, 429,

430, 431.

wurmbii, 436.

Sophraena, 406.

Sophraenella, gen. nov.,

396, 405.

fulva, 396, 406.

Sorex

varius, 190.

Spaniodon, 54.

Sphezeroma, 315.

Sphenodon, 155.

Sphyrena, 59.

Spongia

botryoides, 351, 352,

360, 386.

complicata, 351, 352,

360, 370, 372.

confervicola, 351, 373.

pocillum, 371.

Stenasellus, 307, 308.

viret, 307.

Stenetrium, 302, 303,

304, 305, 306, 307,

308, 309, 311, 312.

313, 314, 316, 321,

327, 329, 330.

antillense, 302, 308,

305, 317, 324, 325,

326, 327, 329, 331.

armatum, 316, 318,

319, 320, 323, 329,

330.

fractum, 316, 319,

323.

haswellti, 316, 317,

321.

imerme, 304, 316,

329.

mediterraneum, 317,

320, 323, 330.

occidentalis, 317, 324,

325, 326, 327, 32%,

330.

serratum, 317, 322,

323, 330,

Stenetrium

stamense, 317, 327,

331.

stebhingti, 317, 325.

Stenocypris

chevreuxii, 127, 128.

perarmata, 126, 127,

128.

Stenodoris

rubra, 86.

Stiliger

wreguaris, 268, 291,

298.

» varians, 268, 290, 291,

297.

Strepsiceros

kudu, 177.

Streptocephalus

dregei, 298, 501.

Struthio

camelus, 177.

Sus, 195, 196, 197.

erymanthius, 196.

phacocheroides, 195.

scrofa, 198.

verrucosus, 196, 197,

198.

— aimboinensis, 197.

Sycon

sp., 380.

raphanus, 393.

Sycorrhiza

corallorrhiza, 374.

Sylvisorex, 190.

gohnstoni, 185,

190.

morio, 190.

muricauda, 190.

sorella, 190.

Syngnathus, 65.

Syrnium

uralense, 177.

Systena

antennata, 396, 409.

argentinensis, 396, 408.

melanocephala, 396,

408.

189,

Taphrospira, gen. nov.,

441.

bathycharax, 442.

INDEX.

Taphrospira

compluvialis, 442, 447.

convallata, 441, 442.

excavata, 442, 447.

Tarbophis

obtusus, 148.

Tarentola, 7, 21, 22.

annularis, 20.

Tarpon

atlanticus, 43, 44,

47.

Teracolus

evagore, 142.

nouna, 142.

saxeus, 142.

yerburti, 142.

Thaumalea

amhersti@, 130.

Theranthropus, 418.

Thrissopater, 54.

Thyroptera, 5.

Tiliqua, 10, 19.

seincoides, 154-157.

Tokea, gen. nov., 220,

222, 240, 257, 298,

260, 261.

esculenta, 222, 240,

241, 242, 243, 244.

huttoni, 222, 248, 249,

256, 257.

kirki, 222, 251, 257.

maorica, 222, 252, 253,

254, 256, 257.

sapida, 222, 245.

sutert, 222, 250, 256.

urewere, 222, 246, 247,

248.

Tragelaphus

spekit, 177.

Trematosaurus, 175.

Trevelyana

bicolor, 85, 89, 105.

ceylonica, 85, 86, 87,

105.

coccinea, 89,

105.

crocea, 87, 105.

timpudica, 89.

Tricheeta, 226.

Trichosurus

vulpecula, 2.

87, 88,

THE END.

503

Tritonia

hawaiensis, 95, 98, 99.

palmert, 95.

rubra, 95.

Troglodytes, 418.

aubryt, 422, 429, 430.

caluus, 429, 430.

Ffuliginosus, 429.

kooloohamba, 422, 429,

430.

leucoprymnus,

431.

marungensis, 426, 429.

niger, 422, 429.

savaget, 416.

troglodytes, 429.

tschego, 429, 430.

vellerosus, 431.

Tropidonotus, 117.

Jasciatus, 112.

natrix, 116.

Tupinambis

teguexin, 10.

Tylonycteris, 189.

pachypus, 189.

Typhlops, 149.

Typhlosaurus, 156.

429,

Ursus

japonicus, 177.

Varanus, 10.

Vespertilio, 189.

Vesperugo

(Vesperus) soloneyi,

Xiphactinus, 62, 71.

Zamenis

Slagelliformis,

148.

gemonensis, 116,

121, 148.

Zizera

marginata, 139.

Zostera, 297.

108,

117,

Printed by Taytor and Francis, Red Lion Court, Fleet Street.

- PROCEEDINGS

| GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

ZOOLOGICAL SOCIETY

OF LONDON,

1904, vol. EI.

PART I.

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MAY anv JUNE., ‘ “ansorian ts : i

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Part I.

May 3, 1904.

Dr. Graham Renshaw, F.Z.8. Exhibition of a drawing of a young African Elephant ....

The Secretary. Exhibition of a photograph, taken by Mr. Frank Haes, of a Quagga ....

Mr. F. E. Beddard, F.R.S. Exhibition of the brain of a Troupial (Quiscalus versicolor)

iMteahen Dy; WOTIB) «6\far:. ss leven nies jes Miegee Bites where hse ous iala ts ci xo aut *

Mr. R. H. Burne, F.Z.S. Exhibition, on behalf of Prof. Stewart, of specimens of the

female reproductive organs of certain _ Marsupials and photographs of a Leathery

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eee ee eee ee ee FO ee tec eres se sets esses sere es FOSS epeorer sees tense neetes

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_ 1. On the Osteology and Systematic Position of the rare Malagasy Bat Myzopoda aurita.

By Oxpriaip Tuoms, F.B.S., F.Z.8. (Plate I.)

ope eesreecerGeese eres ete oreo eeseve

Contributions to the Anatony of the Lacertilia.—(8) On some Points in the Vascular

System of Chameleon and other Lizards. By Frank E. Bepparp, M.A., F.RB.S.,

(Prosecbor sto hexSOciabys as ste oss sea lal talons tale re eee panies ae Cif ln nineteen ie

3. Notes on the Gill-rakers of the Spoonbill Sturgeon, Polyodon spathula. By A. 1D. Inms,

B.Sc. (Lond.), Assistant, Demonstrator in Zoology in the University of Birmingham.

Ea AI) rea tie eiote case ieleyecs <0 jatc ited Aten So aeicE Tete pietepaipie sicisinicisiniel< c'e ais tale ae ieta .

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May 17, 1904.

The Secretary. Report on the Additions to the Society’s Menagerie in April 1904 ..

eeee

Dr. W. T, Calman, F.Z.8. Exhibition of, and remarks upon; a blind Crustacean (Mu-

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Mr. F. E. Beddard, F.R.S. Exhibition of, and remarks upon, a specimen of the ere

Chiamydosaurus kingtt with femoral pores

wees ee weet ane seer MS cece ee OF ee teeeee te

Mr. Oldfield Thomas, F.R.S. Exhibition of, on behalf of the President, a sketch by a

Chinese artist of Pére David’s Deer from Hainan

ee ey

1. On some Nudibranchs from Hast Africa and Zanzibar.—Part V. By Sir C. Exror,

K.C.M.G., late H.M. Commissioner for the East African Protectorate, BZS.

(Plates IIT. & IV.)

ee cere ce ee eco sre seston ere ee tsoeseoeecesesesseeer SF ee sere se esas

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Cs}

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Pee eset ener cre Pees eesene

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oe")

106

107

[ot taae

THE ZOOLOGICAL SOCIETY

OF LONDON.

Tuts Society was founded in 1826 by Sir Sramrorp Rarries,

Mr. J. Sasiye, Mr. N. A. Vieors, 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.

COUNCIL.

HIS GRACH THH DUKE OF BEDFORD, K.G., President.

Srr AnexanDER Barren, Br.

Wittim T. Branrorp, Ese.,

C.LE., LLD., F.R.8., Vice- ||

| E.G. B. Muapz-Watpo, Esa.

President.

GrorceE A. Boviencer, Esaq., |

F.R.S., Vice-President.

Tuomas H. Burroveues, Esa.

Freperic G. D. Drewirr, Ese., |)

MEDS ER.C.P.

Hersert Druce, Ese., F.LS.,

Vice-President.

Cuartes Drummonp, Esa,

| WPreasurer.

Freperick Gittert, Esa.

F. Du Cant Gopman, Hsa.,

D.C.L., F.R.S., Vice-President.

| AtBert GintHer, Esa., M.D.,

Pa.D., F.RS., Vice-President.

Str Epmunp Gites Loner, Br.

P. Caatmers Mircuett, Esa.,

M.A., D.Sc., Secretary.

| Tae Hon. L.Watrer Roruscuinp,

D.8c., M.P.

Howarp Saunpers, Esa., Vice-

President,

Davin SHarp, Ese., M.D., F.R.S.

OLpFIeELD Tuomas, Esa., F.R.S.

Surron Tris, Ese.

Cuartes S, Tomes, Esq., M.A.,

F.R.S,

Aveustus F, Wiener, Ese.

The Society consists of Fellows, and Honorary, Foreign, and

Corresponding Members, elected according to the By-Laws.

The Gardens in the Regent’s Park are open from Nine o’clock a.m.

till Sunset.

The Offices (3 Hanover Square, W.), where all communications

should be addressed, are open from Ten till Five, except on Satur-

days, when they are closed at Two o'clock P.m.

The Library (8 Hanover Square), under the superintendence of

Mr. F H. Warernouss, Librarian, is open from 10 a.m. to 5 P.M.,

on Saturdays to 2 p.m. It is closed in the month of September.

The Meetings of the Society for General Business are held at the

Office on the Thursday following the third Wednesday in every

month of the year, except in September and October, at Four p.m.

The Meetings for Scientific Business are held at the Office twice -

a month on Tuesdays, except in July, August, September, and

October, at half-past Eight o’clock p.m.

The Anniversary Meeting is held on the 29th April, at Four p.m.

TERMS FOR THE ADMISSION OF FELLOWS.

FeLtows pay an Admission Fee of £5, and an annual Contri-

bution of £3, due on the 1st of January, and payable in advance,

or a Composition of £30 in lieu thereof; the whole payment,

including the Admission Fee, being £35.

No person can become a Frttow until his Admission Fee and

First Annual Subscription have been paid, or the annual payments

have been compounded for.

Frttows elected after the 30th of September are not liable for

the Subseriptions for the year in which they are elected.

PRIVILEGES OF FELLOWS.

Frttows have Personal Admission to the Gardens with Two

Companions daily, upon signing their names in the book at the

entrance gate.

Frxtows receive a Book of Saturday and a Book of Sunday Orders

every year. These Orders admit two persons to the Gardens on each

Saturday and two on each Sunday in the year. But the Saturday

Orders are not available if the Fettow shall have used his privilege

of personally introducing two companions on the same day.

Ferttows also receive every year Twenty Free Tickets (Green),

each valid for the admission of one adult any day of the week,

including Sunday. Children’s Tickets (Buff) can be had in lieu of

Green Tickets in the proportion of two Children’s Tickets to one

Adult’s. These Tickets, if not made use of in the year of issue, are

available for following years.

In no case can two children be passed through the gates as

one adult.

Frtiows, if they wish it, can exchange the Book of Saturday

Orders for Twenty Green Tickets available for any day. The Book

of Sunday Orders can also be exchanged for a similar packet of

Twenty Tickets. These books must, however, be returned entire,

and the exchange can only be made during the year of their issue.

The annual supply of Tickets will be sent to each Fetrow on the

Ist of January in every year, on his filling up a form of Standing

Order stating in what way they should be made up, and to what

address they should be sent. Forms for this purpose are supplied

on application.

The Wire of a Fettow can exercise all these privileges in his

absence.

Ferttows have the privilege of receiving the Society’s Publications

on payment of the additional Subscription of One Guinea every

year. 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. Frntows are likewise entitled to purchase the

Transactions and other Publications of the Society at 25 per cent.

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 1871, if above the value of Five pounds.

Fettows also have the privilege of subscribing to the Annual

Volume of the Zoological Record for a sum of £1, payable on the

Ist July in each year, but this privilege is forfeited unless the

subscription be paid before the 1st of December following.

Frttows may obtain a Transrerarts Lyory Ticket 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 Frtiow who intends to be absent from the United Kingdom

during the space of one year or more may, upon giving to the

Secretary notice in writing, have his name placed upon the

“ dormant list,” and will be thereupon exempt from the payment of

his annual contribution during such absence.

Any Frttow, having paid all fees due to the Society, is at liberty to

withdraw his 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, M.A., D.Sc.,

Secretary.

3 Hanover Square, London, W.,

October, 1904.

MEETINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON

FOR

SCIENTIFIC BUSINESS.

(AT 3 HANOVER SQUARE, W.)

Session 1904-1905.

1904.

Tuxzspay, Novemper 15 and 29 | Tuxspay, Drceuser 13

1905.

TurEspay, JANUARY 17 Turspay, Aprm ..18

- Frpruary 7 and 21 ar May .... 2iandl6

5 NEN OHA i) 4. teil xe JUNE eee nO

The Chair will be taken at half-past Eight o'clock in the Evening

precisely.

LIST OF THE PUBLICATIONS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

TE 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 attached to each annual volume of

the “ Proceedings,” to illustrate the new or otherwise remark-

able species of animals describedin them. 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,

on the first of the months of June, August, October, and

April, the part published in April completing the volume

for the last half of the preceding year. From January 1901

they have been issued as two half-yearly volumes.

The “Transactions” contain such of the more important

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 25 per

cent. less than the price charged for them to the Public. A

further reduction of 25 per cent. is made upon purchases of

Publications issued prior to 1871, 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 £1

(which includes delivery in the United Kingdom only),

payable on the Ist 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.

[ October, 1904. }

TRANSACTIONS OF THE ZOOLOGICAL SOCIETY OF LONDON.

4to. 16 vols. and Index. Ale Price to the

ellows. Public.

Vol. _I., containing 59 Plates.... (1833-35) .... £3 18 6.... £418 Ot

shane tesl 5 ZL 1h * Vt SERLSSO Ai ee ele OO ae 5 6 6f

uel; 5 633, sens (RS42—49) eee, TOMS mone ALS SOF

pt a OES _ Tl gs te sist GUS G2) aap One nO ae 8) 2) 16

‘o, e 67. %, “ers 4 1(1862-66)) «#5 ed oC TORO

op AINE if CB ey eon (CISBI=72) cog LO) 4 O. 13 12 0

“WILDE. ii ey Boon CEO) so55 O SB 12 a 90

x. - Oe once (ClWEHT®) sono IO O 8, 13° 0

Index, Noles Ie= Nua un leek na Arh GERRI) Sona OOO 010 0

Vol. XI., containing 97 Plates.. (1880-85) .... 912 0. 12ZMGSO

Fe Sale A) ae bo USO 80) Gago Wests = Os 7 4 0

ep GUI Hi Oe) by pee ELSIE OS ee GUASaaann SIL @

jy bye D2 ay ay ak (1898-1901) . sa) HO 714 0

5 eM Tees, GF om oo (Aine 1008) 25) 1 896. 110 0

PeeXOV Uae ree, Ble pel od (Aw, Maas go OAR G.. 018 0

oy os Willen i ety | I Ne (Oe MOS). I Gs TL AORO

PROCEEDINGS OF THE COMMITTEE OF SCIENCE AND

CORRESPONDENCE OF THE ZOOLOGICAL SOCIETY OF

LONDON. 8vo. 2 vols. (Letterpress only). rice t Beige vo tng

Fellows.

lene Iie ISOSES IOI EhYOs Goncooeo al joer AsiGds nha mOSai

Pulomlisa ze PMs as OT 5 Oe ceoe AS, Os Coes OSs

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 I. 1833.1 vol. 8vo. 4s. 6d. .. 6s. | Part JX. 1841.1 vol. 8vo. 4s. 6d. .. 6s.

PIT, 1884 selfs ad PAsaGWA euGs, 5 TIO) eM GEE. Go,

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» WML 130, 25. Gil, 5, G54 KV. 1847) ort) sede aly Gulesee

» VIII. 1840. 49 4s. Gd. .. 6s.F Thadlexz 1880-1847. 95 4s, 6d. .. 6s.

8vo. 13 vols. an Index. (Second Series.)

Letterpress only. With Plates coloured.

Price to Price to the Price to Price to the

Fellows. Public, Fellows. Public.

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ray EW 45,6) Sod Go. ONT, 1°40 219 eee aor

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XXVIII. 1860. 55 Agu OAs. (Be OSE te sects LIU @ YP ie

Index 1848-1860, in ae 6d. Gs.

i Out of print.

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. t Out of print.

PROCEEDINGS or tar GENERAL MEETINGS ror SCIENTIFIC

BUSINESS or tar ZOOLOGICAL SOCIETY OF LONDON.

8vo. 7 vols.

Price to Price to the

Fellows. Public.

WG Os ORG RSR Mona nis soo oho som bOUK aoc bodobe UID DD COC IS onda os 24s,

o BOL, 5 ite a letheueeteterYeveRoNscede aye Aten eRee hee tenet yore teh eat WSs." srecrste 24s.

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5 ny OLS (ERIE Jo) ag Soon oe cogs obo b 00000 sores iat OSH amare 12s,

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. (Highth Edition.) 8vo.

1883. Cloth, 4s. 6d.

List of the Vertebrated Animals now or lately Living in the Gardens

of the Zoological Society of London. (Ninth Edition.) 8vo.

1896. Cloth, 6s.; Paper, ds.

Catalogue of the Library of the Zoological Society of London.

(Fifth Edition.) 8vo. 1902. Cloth, 6s.; Paper, 5s.

These publications may be obtained at the Socrery’s OFFice

(3 Hanover Square, W.), at Messrs. Lonemans’ (Paternoster Row,

E.C.), or through any bookseller.

ME ZOOLOGICAL “RECORD:

—r0se300—

VHE object of the Zootocrcat Record 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 Inlex 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’ is published by the Society at the

price of 30s. per volume. But all Members of the Zoological Society

of London have the privilege of receiving it, including the cost of

delivery (within the United Kingdom), at a subscription price

of 20s. per annum. ‘This Subscription is due on the Ist of July in

every year, and the privilege of Subscription is forfeited unless the

amount be paid before the lst of December following.

The Zoological Society, having purchased the entire stock of

the ‘Zoological Record,’ is able to supply complete sets. The

thirty-seven Volumes to the end of the nineteenth century, and the

Index-Volume (1880-1900) in addition, will be supplied for £15

net (or without the Index-Volume, for £14 10s. net). Volumes of

any single year (exclusive of the last five volumes and Vol. 6) can

likewise be supplied at 10s. per volume net.

The price of the Index Zoologicus (Index-Volume 1880-1900)

is 20s., to Fellows 18s.

Members of the Society wishing to subscribe to the ‘ Record’

are requested to apply at this office for a Form, to be returned

when filled up and signed by the subscriber. In order to facilitate

the payment of the subscription, a Banker’s Order Form is also

supplied te those who prefer that mode of payment. This order,

when filled up and signed, should be sent to the Society’s office for

registration ; it will then be sent to the Agents named therein.

Learned Societies and Institutions and members of the former

Zoological Record Association are permitted tv subscribe to the

‘Record’ on the same conditions as are accorded to Members of

the Zoological Society.

Commencing with Vol. xxx1x., the divisions of the ‘ Zoologival

Record’ may be obtained separately as shown on the next page.

SEPARATE DIVISIONS OF THE ZOOLOGICAL RECORD.

At present each Volume of the Zoorocican Record consists of

20 separately paged Divisions. These may be obtained separately,

in paper covers, stitched and lettered.

The following are the Divisions and their net prices, viz. :—

So.

List of abbreviations of journals, ete. (0)

Special Records, viz. :—

I. General Subjects .. 2 6

II. Mammalia 2 Om

III. Aves Sas erage @ @

IV. Reptilia and Batrachia. . 26

V. Pisces 4G

VI. Tunicata iL @

VII. Moilusca 4 0

VIII. Brachiopoda .. ke

X. Crustacea 280

XI. Arachnida wi)

XII. Myriopoda LO

XIV. Echinoderma 3 6

XV. Vermes .. a

XVI. Coelenterata . Il @

XVII. Spongise ms 0)

XVIII. Protozoa Bosse Watare the Zn)

Index of new names of genera and subgenera BW)

On receipt of the price any Division will be forwarded as soon

as ready.

These separate Divisions can be obtained from the Zoological

Society, 3 Hanover Square, London, and also from the following

Agents: Friedliinder & Sohn, 11 Carlstrasse, Berlin; and Librairie

A. Hermann, 6 rue Sorbonne, Paris. Cheques and Post-Office

Orders should be made payable to ‘The Zoological Society,” and

crossed ‘* Drummond’s.”

P. CHALMERS MITCHELL,

Secretary.

October, 1904.

ZoouoGgicaL Society or Lonpon,

3 Hanover Squares, W.

LIST OF VOLUMES or raz “ZOOLOGICAL RECORD,’

The Record of Zoological Literature, 1864-1868. Vols. 1.-v:

Kdited by Atsrrt C. L. G. Ginrner, M.A., M.D., Ph.D., F.Z.S., &c.

Price 10s. each Volume. Net.

The Record of Zoological Literature, 1869. Vclume Sixth.

Edited by Atsert C. L. G. Ginruer, M.A., M.D., Ph.D., F.R.S.,

F.Z.8., &c. London, 1870. Price 30s.

The Zoological Record for 1870-1872, Vols. vir.-1x. Edited

by Atrrep Newron, M.A., F.R.S., F.L.S., V.P.Z.8., &e. Price 10s.

each Volume. Net.

The Zoological Record for 1873-1883, Vols. xxx. Edited by

Epwarp CaLpwe tt Ry, F.Z.8., M.E.S. Price 10s. each Volume. Net.

The Zoological Record for 1884, 1885, Vols. xx1., xx11. Edited

by F. Jerrrey Bert, M.A. Price 10s. each Volume. Net.

The Zoological Record for 1886-1890, Vols. xxm1.—xxvit.

Edited by Frank E. Bepparp, M.A., F.Z.8. Price 10s. each

Volume. Net.

The Zoological Record for 1891-1897, Vols. xxy11.—xxxIv.

Edited by D. Sarr, M.A., F.R.S., F.Z.8., &. Price 10s. each

Volume. Net.

The Zoological Record, Volume the Thirty-fifth ; being Records

of Zoological Literature relating chiefly to the year 1898. By J.A.

Thomson, R. Lydekker, R. Bowdler Sharpe, G. A. Roulenger, W. A.

Herdman, E. R. Sykes, E. A. Smith, G. C. Crick, A. W. Brown,

D. Sharp, F. A. Bather, Florence Buchanan, R. T. Gunther, and

R. von Lendenfeld. Edited (for the Zoological Society of London) by

Davin Suarp, M.A., F.R.S., F.Z.8., &c. London, 1899. Price 30s.

The Zoological Record, Volume the Thirth-sixth ; being Records

of Zoological Literature relating chiefly to the year 1899. By

J. A. Thomson, R. Lydekker, R. Bowdler Sharpe, G. A. Boulenger,

W. A. Herdman, E. R. Sykes, E. A. Smith, G. C. Crick, A. W,

Brown, D. Sharp, F. A. Bather, A. Willey, and R. von Lendenfeld.

Edited (for the Zoclogical Society of London) by Davin Suarp, M.A.,

F.RS., F.Z.8., &. London, 1900. Price 30s.

The Zoological Record, Volume the Thirty-seventh; being

Records of Zoological Literature relating chiefly to the year 1900.

By J. A. Thomson, R. Lydekker, R. Bowdler Sharpe, G. A. Boulenger,

W. A. Herdman, EH. R. Sykes, E. A. Smith, G. C. Crick, A. W.

Brown, D. Sharp, F. A. Bather, A. Willey, and E. A. Minchin.

Edited (for the Zoological Society of London) by Davin Saarp,

M.A., F.R.S., F.Z.S., &c. London, 1901. Price 80s.

The Zoological Record, Volume the Thirty-eighth; being

Records of Zoological Literature relating chiefly to the year 1901.

By J. A. Thomson, R. Lydekker, R. Bowdler Sharpe, G. A. Boulenger,

Alice L. Embleton, E. R. Sykes, E. A. Smith, 8. Pace, Albert

Brown, D. Sharp, F. A. Bather, and E. A. Minchin. Edited (for

the Zeological Society of London) by Davin SHarp, M.A., F.R.S.,

F.Z.8., &c. London, 1902. Price 30s.

The Zoological Record, Volume the Thirty-ninth ; being Records

of Zoological Literature relating chiefly to the year 1902. By

D. Sharp, R. Lydekker, R. Bowdler Sharpe, G. A. Boulenger, W. T.

Calman, E. R. Sykes, E. A. Smith, Alice L. Embleton, F. A. Bather,

ki. A. Minchin, and H. M. Woodcock. Edited (for the Zoological

Society of London) by Davin Suarp, M.A., F.RS., F.Z.S., &e.

London, 1903. Price 30s.

Index Zoologicus. An alphabetical list of names of genera

and subgenera proposed for use in Zoology, as recorded in the

Zoological Record, 1&80-1900; together with other names not

ineluded in the ‘ Nomenclator zoologicus’ of 8. H. Scudder. Com-

piled (for the Zoological Society of London) by Caries Owen

Warernouse and edited by Davin Suarp, Editor of the Zoological

Record. London, 1902. Price to Fellows, 1&s.; price to the

public, 20s,

Tluse publications may be obtained at the Socrery’s Orrice

(3 Hanover Square, W.).

IWOs A

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

May 3rd, 1904.

G. A. Boutencer, Esq., F.R.S., Vice-President, in the Chair.

The Secrerary exhibited, on behalf of Dr. GrAnaAM RENSHAW.

F.Z.8., an outline sketch of a young African Elephant, mounted

in the Museum of the Jardin des Plantes, Paris, which showed

an unusual development of hairs on its body.

The Secretary also exhibited a photograph of the Quagga,

taken by Mr. F. Haes from the last female specimen that had

lived in the Gardens.

Mr. F. E. Bepparp, F.R.S., exhibited the brain of a Troupial

(Quiscalus versicolor) in the hinder part of both cerebral hemi-

spheres of which was an entangled mass of Nematoid worms

lying below the pia mater. The bird was reported to have

dropped down suddenly from its perch “in a fit.”

Mr. R. H. Burne, F.Z.S., exhibited specimens of the female

reproductive organs of Derameles obesula, Trichosurus vulpecula,

and Dasyurus viverrinus; the specimens had been presented to

the Museum of the Royal College of Surgeons by Professor Hull.

Mr. Burve also exhibited photographs of the rare Leathery

Turtle (Dermochelys coriacea).

Mr. Ouprietp Tuomas, F.R.S., read a paper on the osteology

and systematic position of the rare Malagasy Bat Myzopoda

aurita. This Bat was considered to form a special family, the

Myzopodide, which appeared to be most nearly allied to the

American Vatalide and Mormopide.

* This Abstract is published by the Society at 3 Hanover Square, London,

W., on the Tuesday following the date of Meeting to which it refers. It will

be issued, free of extra charge, to all Fellows who subscribe to the Publications,

along with the ‘ Proceedings’ ; but it may be obtained on the day ef publication

at the price of Sixpence, or, if desired, sent post-free for the sum of Six

Shillings per annum, payable in advance.

Mr. F. EK. Bepparp, F.R.S., read a third of a series of papers

on the Anatomy of the Lacertilia, which dealt with points in the

vascular system of Chameleon and other Lizards.

A communication from Mr. A. D. Tums, B.Se., contained notes

on the gill-ralkers of Polydon, in which he pointed cut that these

processes in this Fish were setiform structures disposed along

both the anterior and posterior edges of the first four branchial

arches and along the anterior edge of the fifth arch also. Hach

gill-raker consisted of a basal portion, which was attached by

means of elastic fibres to the cartilage of the branchial arch, and

a long, free shaft portion which formed its principal part. The

matrix of a gill-raker contained numerous lacunar spaces which

were connected with one another by means of canaliculi, and in

the shaft portion it contained, in addition, a series of anastomosing

channels in which lay blood-capillaries and some loose connective-

tissue. The mucous membrane covering a branchial arch was

prolonged over each gill-raker in the form of a complete investing

coat. The gill-rakers of Polydon were exoskeletal structures,

and they appeared to be composed of a substance closely allied

to osteo-dentine, and their presence on the branchial arches

might perhaps be accounted for on the supposition that the

sonata m forming the outer portions of the gill-clefts had migrated

on to the inner margins of the branchial arches. The food of

Polydon consisted of microscopical organisms, and the gill-rakers

function as a straining mechanism which effectually barred the

entry of such particles into the gill-cavities.

A paper was read by Dr. W. G. Ripewoop, F.Z.S., on the

“Cranial Osteology of the Hlopide and Albulide, with Remarks

on the Morphology of the Skull in the Lower Teleostean Fishes

generally.” The paper included a detailed account of the skulls

of Hlops saurus, Megalops cyprinoides, Albula conorhynchus, and

Bathythrissa dorsalis, together with remarks, generalisations,

and criticisms of published accounts of the skull-bones of fishes,

based mainly upon an examination of the skulls of 20 species of

fishes selected from among the most primitive families of the

existing Malacopterygii.

Among other features pointed out by Dr. Ridewood was the

fact that m Hlops and Albula, although the parietals met in the

middle line, the supraoccipital extended forward beneath them to

reach, and even to underlie, the frontal bones; in the former

genus and in J/egalops the supraoccipital also touched the ali-

sphenoid bones. ‘The posterior temporal fossee were so great in

Megalops that they reached the orbitosphenoid bone, and they

communicated freely above the brain-case. A careful comparison

of the skulls of Bathythrissa and Albula confirmed the close

relationship existing between these genera suggested by Boulenger

in 1898. Except that in some few respects the bones were

modified in correspondence with a deep-sea habit, the skull of

Bathyihrissa bore a very close resemblance to that of Albula.

In the second part of the paper Dr. Ridewood discussed the

relation of endosteal to ectosteal ossification, the eye-muscle

canal, the various fossx, fenestra, and the larger foramina of the

cranium, the bullz enclosing vesicles of the swim-bladder, the

union of a vertebral centrum with the back of the basicraniai

axis, the brush-like tendon-bones at the back of the skull (the

opisthotic limb of the post-temporal was shown to belong to this

series), the supratemporal and subtemporal bones, the sesamoid

articular (so-called “coronoid”), the modes of articulation of the

palatine and the hyomandibular bones with the cranium, the

opercular bones, the hyobranchial skeleton, and the “spicular

bone” (which Dr. Ridewood showed to be an ossified ligament,

and not the first pharyngobranchial as had been previously

supposed).

The next Meeting of the Society for Scientific Business will be

held on Tuesday, the 17th May, 1904, at half-past Hight o'clock P.m.,

when the following communications will be made :—

1. Sir Caarues Exrot, K.C.M.G.—On some Nudibranchs from

Kast Africa and Zanzibar.—Part V.

2. Mr. G. A. Boutencer, F.R.S.—Description of a new Tree-

Frog of the Genus Ayla, from British Guiana, carrying Eggs on

the Back.

3. Mr. F. E. Bepparp, F.R.S.—Notes upon the Anatomy of

certain Lowe.

The following Papers have been received :—

1. Dr. G. Srewarpson Brapy, F.R.S., C.M.Z.S.—On Entomo-

straca collected in Natal by Mr. James Gibson.

2. Lt.-Col. J. Matcoitm Fawcett.—On some new or little-known

Butterflies, mainly from high elevations in the N.H. Himalayas.

3. Dr. P. Cuaumers Muircurety.—On the Disposition and

Morphology of the Intestinal Coils in Mammals.

Communications intended for the Scientific Meetings of the

ZoOoLoGIcAL Society or Lonpon should be addressed to

P. CHALMERS MITCHELL, Secretary.

3 Hanover Square, Lonpon, W.

10th May, 1904.

No. 8.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON®*

May 17th, 1904.

Howarp Saunpers, Esq., F.L.S., Vice-President, in the Chair.

The Secretary read a report on the additions that had been

made to the Society's Menagerie during the month of April 1904,

and called special attention to a fine specimen of the Boatbill

(Canchroma cochlearia) obtained by purchase; to two Cheetahs

(Cynelurus jubatus), from the Soudan, presented byt Colla:

Mahon, C.B., F.Z.8.; and to two Keas (Nestor notabilis), presented

by Mr. T. E. Doune.

Dr. W. T. Catman, F.Z.S., exhibited a specimen of a blind

crustacean (Munidopsis polymorpha), from a subterranean lake in

the island of Lanzarote, Canaries, collected by Mr. Fairfax

Prevost. The species was described in 1892 by Dr. Koelbel, of

Vienna, from specimens collected by Prof. Simony, but appears

to have been overlooked by recent writers on the group to which

it belongs. The lake in which it is found is salt and communicates

with the sea, the water rising and falling with the tide. All the

other species of the genus, over 100 in number, are inhabitants

of the deep sea, none being recorded from a depth of less than

about 100 fathoms.

Mr. F. E. Bepparp, F.R.S., exhibited the body of an example

of Chlamydosaurus kingi which had died in the Menagerie some

years ago, and made remarks on the absence of femoral pores.

Mr. Ouprietp Tuomas, F.R.S., exhibited, on behalf of the

President, a sketch of a hind and fawn of Pére David’s Deer

(Llaphurus davidianus) from Hainan.

* This Abstract is published by the Society at 3 Hanover Square, London,

W., on the Tuesday following the date of Meeting to which it refers. It will

be issued, free of extra charge, to all Fellows who subscribe to the Publications,

along with the ‘Proceedings’; 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 Siz

Shillings per annum, payable in advance.

The fifth of a series of papers by Sir CuaruEs Extor, K.C.M.G.,

on Nudibranchs from Zanzibar and East Africa, was read.

Twenty species of the genera Notodoris, Trevelyana, Nembrotha,

Marionia, Bornella, and Pleuroleura were treated of in the paper,

of which eleven were described as new.

Mr. G. A. Boutencsr, F.R.S., described a new species of Tree-

Frog of the genus Hyla, from British Guiana, carrying eggs on

the back.

Mr. F. E. Bepparp, F.R.S., read a paper containing notes on the

anatomy of certain species of Snakes of the family Borde.

A communication from Dr. G. Stewarpson Brapy, F.R.S.,

C.M.Z.S., contained an account of a collection of Entomostraca made

in Natal by Mr. James Gibson. Eleven species were enumerated

in the paper, of which nine were described as new, one being made

the type of a new genus.

The next Meeting of the Society for Scientific Business (closing

the Session 1903-04) will be held on Tuesday, the 7th June, 1904,

at half-past Eight o’clock p.m., when the following communications

will be made :—

1. Lt.-Col. J. Matcorm Fawcrrr.—On some new or little-

known Butterflies, mainly from high elevations in the N.E.

Himalayas.

2. Dr. A. G. Burier, F.Z.8.—On Seasonal Phases in Butterflies.

3. Mr. F. E. Bepparp, F.R.S.— Note on an apparently

Abnormal Position of the “‘ Brephos ” within the body of a Skink

(Chalcides lineatus). ;

4. Dr. KE. A. Gortp1, C.M.Z.8.—On the rare Rodent Dinomys

branichi Peters.

5. Mr. C. Satuntn, C.M.Z.S.—On the Black Wild Cat of

Transcaucasia.

6. Mr. R. LypeKkrer.—On a Buffalo Skull from Hast Central

Africa.

7. Dr. A. Smrra Woopwarp, F.R.S.—On Two new Labyrintho-

dont Skulls.

Communications intended for the Scientific Meetings of the

ZooLoGicaAL Sociery oF Lonpon should be addressed to

P. CHALMERS MITCHELL, Secretary.

3 Hanover Square, Lonpon, W.

24th May, 1904.

No. 9.

ABSTRACT OF THE PROCEEDINGS

ZOOLOGICAL SOCIETY OF LONDON.

June 7th, 1904.

F. DuCane Gopman, Esq., D.C.L., F.R.S., Vice-President,

in the Chair.

The SucrEerARy read a report on the additions—276 in number

—that had been made to the Society’s Menagerie during May

1904, and called special attention to the following specimens, all

new to the collection :—Three Andaman Banded Crakes (Rallina

canningt), presented by the Government of India; a Yellow-

handed Howler (Mycetes beelzebul), an Antilopine Kangaroo

(Macropus antilopinus), and a Grey Solitaire (Myiadectes

wnicolor), deposited.

The Secrerary also made some remarks on two fully-adult

specimens of the Orang-Utan which he had recently seen in Paris.

Dr. Gintuer, F.R.S., Vice-Pres.Z.8., on behalf of the PRESIDENT,

exhibited a series of hybrid Pheasants killed at various times in

the coverts at Woburn, where many distinct species had been

turned out into the open. He stated that nothing of a definite

nature was known as to their parentage, but proposed tentative

determinations.

Dr. F. D. Drewirt, F.Z.S., exhibited and made remarks upon

two fine antlers of the North-African Red Deer (Cervus elaphus

barbarus) which he had brought from the high forest land

extending far inland on the borders of Tunis and Algeria.—

Dr. Drewirr also exhibited a pair of horns of Loder’s Gazelle

(Gazella leptoceros) from South Algeria.

A communication from Dr. GraHaAm RensHAw, F.Z.8., con-

tained notes, illustrated by photographs, on a pair of Short-

horned Buffaloes in the Antwerp Zoological Gardens.

* This Abstract is published by the Society at 3 Hanover Square, London,

W., on the Tuesday following the date of Meeting to which it refers. It will

be issued, free of extra charge, to all Fellows who subscribe to the Publications,

along with the ‘ Proceedings’; butit may be obtained on the day of publication

at the price of Sixpence, or, if desired, sent post-free for the sum of Siv

Shillings per annum, payable in advance. ,

Mr. F. KE. Bepparp, F.R.S., exhibited and made remarks upon

a skull of the Cape Crowned Crane (Balearica chrysopelargus)

showing paired lateral and a single median bony bosses, suggestive

of those of the horn-bearing vertebrates.

Mr. R. E. Houprne exhibited and made remarks upon a series

of 12 photographs of the horns of the Wapiti Stag living in the

Society’s Gardens, taken in fortnightly intervals from the shedding

of the horns on March 30th, 1903, to the first week in October,

when they were clear of velvet.—Mr. Houprine also exhibited a

fine pair of the shed horns of the Irish Red Deer, having 14 points

and weighing 8 lbs.

Mr. R. I. Pocock, the Superintendent of the Gardens, exhibited

and made remarks upon living specimens of hairless varieties of

the Common House-Mouseé (J/us musculus) and the Brown Rat

(If. decumanus).—Mr. Pocock also exhibited and made remarks

upon some young examples of the Egyptian Fat-tailed Gerbille

(Pachyuromys duprest), born in the Menagerie.

A communication from Lt.-Col. J. Maucoum Fawcrrt con-

tained descriptions of ten species of Butterflies, mainly from high

elevations in the N.E. Himalayas. Hight of them were new

species or varieties and were diagnosed as follows :—

MELIT#A TIBETANA, Sp. n.

Hab. Khamba Jong, Thibet, 15,000 ft. elevation.

Allied to the var. of JZ. sindwra Moore (described by Elwes in

Trans. Ent. Soc. 1888, pt. i. p. 336), which = I. sikkimensis

Moore, but differs from it in its much paler coloration, and the

presence on the upperside of pale ochreous bands and spots, and

more prominent fuscous markings.

ARGYNNIS CLAUDIA, Sp. n.

Hab. Khamba Jong, Thibet, 15,000 ft. elevation.

Allied to A. clara Blanchard, but differs from it in its rounder

wings, smaller size, and paler coloration on upperside; the outer

margins of both wings are broadly fuscous, containing a sub-

marginal series of pale spots, without the submarginal and

marginal fine black lines beyond them as in A. clara.

Lyc@NA ARIANA Moore, var. ARENE, var. n.

Hab. Khamba Jong, Thibet, 15,000 ft. elevation.

Differs from L. ariana Moore, from the N.W. Himalaya, in

being smaller, and in the discal row of black spots on the under-

side bemg minute, fewer in number, and obsolete in some

specimens in the hind wing, and in the discal white spots at apex

of cell of hind wing being larger.

Lyc@®NA PHERETES Hiibner, var. PHARIS, var. n.

fab. Khamba Jong, Thibet, 15,000 ft. elevation.

This form differs from the var. asiatica of Elwes in having

fewer discal black spots on fore wing, and in the size and length

of the pale cellular spot on hind wing underside.

Liyc#naA (ZizmRA) ZEBRA, sp. n.

Hab. Tounghoo, Burma (June).

Nearly allied to Z. (7.) marginata from China, but differs in

the discal row of fuscous spots of the fore wing being fainter and

more regular on the underside, and in the black border of the

upperside being more even in its inner edge and bending inwards

on the costa.

LycaNA (NIPHANDA) MARCIA, sp. 0.

Hab. Tounghoo, Burma (June).

Allied to JV. tessellata Moore, from Penang, but differs in its

much less heavy markings on the underside.

COLIAS BERYLLA, sp. n.

Hab. Khamba Jong, Thibet, 15,000 ft. elevation.

This species belongs to the C. hyale section of Colias, and is

nearest to C. erschoffi Alphéraky, from Thian Shan Mts., but

differs in the greater melanism of hind wing.

COLIAS NINA, sp. n.

Hab. Khamba Jong, Thihet, 15,000 feet elevation.

This species appears to be a transition between the C. hyale and

C’. edusa sections of Colias.

Dr. A. G. Burier, F.Z.8., contributed a paper “On Seasonal

Phases in Butterflies,” in which he stated that so-called “seasonal

forms” were only the variations of polymorphic species adapted

to seasons, by the gradual elimination of those varieties which

were unsuited to the environment of thoseseasons. This adapta-

tion being incomplete in the case of many species, it was absurd

to express astonishment if forms characteristic of the wet and dry

seasons sometimesappearedsimultaneously. Because they did some-

times appear together, the author had some years since proposed

the term “seasonal phase” to supersede that of ‘seasonal form.”

Capt. RicHarp Crawsuay, F.Z.S., contributed a note on the

Prey of the Lion, showing that the popular belief of the Lion

being a fastidious feeder was incorrect, but that he ate carrion not

killed by himself in all stages of putrefaction, and was not averse

to preying on such very small game as the smaller rodents. The

tips of a Porcupine’s quills found buried in a Lion’s fore paw were

exhibited, together with the skull and complete skin of the Lion.

Attention was drawn to the filthy state of this Lion’s claws,

which enabled it to be realised how it was that septic poisoning

so often terminated fatally with persons mauled by Lions.

Mr. F. E. Bepparp, F.R.S., read the following three papers

based on observations he had made in the Society’s Prosectorium :

(1) “ Note on an apparently Abnormal Position of the ‘ Brephos’

within the Body of a Skink (Chaletdes lineatus)” ; (2) ‘“Contribu-

tions to the Knowledge of the Visceral Anatomy of the Pelagic

Serpents Hydrus platyurus and Platyurus columbinus” ; and (3)

‘‘On the Presence of a Parasternum in the Lacertilian Genus

Tiliqua and on the Poststernal Ribs in that Genus.”

A communication from Dr. HK. A. Gorupr, C.M.Z.8., contained

a description and an account of the habits of the rare Rodent

Dinomys branickit Peters, living specimens of which had recently

been received at the Goeldi Museum, Para. Dr. Goeldi was un-

certain as to the exact habitat of this animal, but he was of

opinion that it was in the almost unexplored regions of the eastern

slopes and tablelands of Bolivia and Peru bordering on Brazil.

A communication from Dr. C. Satunin, C.M.Z.8., contained

an account of the Black Wild Cat of Transcaucasia, which he

described as a new species.

Mr. R. LypexKer, F.Z.8., described the skull of a Buffalo from

East Central Africa as the type of a new race, with the name

Bos caffer maithewsi. This Buffalo appeared to connect the

typical B. caffer with B. c. nanus, having horns approaching those

of the latter, but its skin and hair black. A white tip to the tail

seemed a special peculiarity.

Mr. LypkkeER also described from Ichang a new species of

Tufted Deer (Zlaphodus ichangensis) differing from H. michianus

in its darker colour, whiter tail, smaller antlers, longer tusks,

shorter nasals, and certain other features in the s«ull.

Dr. A. SmrrH Woopwaprp, F.R.S., read a paper on two new

Labyrinthodont skulls acquired by the British Museum. They

were the skull of a new species of Capitosawrus from the Trias of

N. Staffordshire, which showed the occiput more clearly than any

Labyrinthodont skull hitherto described, and demonstrated its

Amphibian character; and the skull of a new genus allied to Trema-

tosaurus, from the Trias of Spitzbergen, which was the first satis-

factory Labyrinthodont fossil described from the Arctic Regions,

This Meeting closes the Session 1903-04. The next Session

(1904-05) begins in November.

Communications intended for the Scientific Meetings of the

ZooLocrcaL Society oF Lonpon should be addressed to

P. CHALMERS MITCHELL, Secretary.

3 Hanover Square, Lonpon, W.

14th June, 1904,

ContENTS (continued).

June 7, 1904,

The Secretary. Report on the Additions to the Society’s Menagerie in May 1904........ 129

The Secretary. Note on two specimens of the Orang-utan he had seen in Paris.......... 129

Dr. Ginther, F.R.S. Exhibition of, on behalf of the President, and remarks upon, a series

of mounted specimens of hybrid Pheasants ......0e-cceceeererececeeccssnsseee - 129

Dr. F. D. Drewitt, F.Z.8. Exhibition of, and Ponee upon, some antlers of the North-

African Red Deer ..... Ieper ebro Bote La Eick, UMBO™ Atma enan Se a ordeal “creat 130

Dr. F. D. Drewitt, FZS. Exhibition of a pair of horns of Loder’s Gazelle from Southern

rem A Oe Leap male Scarbin"a tyeisi wfesiats.c wiuinicteiverstajol ape staletelstaraisteztis ais'a's) <(chetskee SHeoseue 130

Dr. Graham Renshaw, ¥.Z.S. Exhibition of photographs of, and remarks upon, a pair

of Short-horned Buffaloes in the Antwerp Zoological Gardens ........-+....++--+- 130

Mr. F. HE. Beddard, F.R.S. _ Exhibition of, and remarks upon, the skull of a Cape Crowned

Granei se: ).). eee aiefeiaras sit SEO CAUE BROCE MU COD OE Eee (Dee Ra Hes cis ot Aaroa ace eee 131

Mr. R. E. Holding. Exhibition of photographs of, and remarks upon, the antlers of the

Wapiti in various stages of growth .......0e0.cee cece cece cece ee teeter ec ce cere ce 131

Mr. R. E. Holding. Exhibition of a pair of antlers of the Irish Red Deer ........... - 133

Mr. R. I. Pocock. Exhibition of, and remarks upon, living hairless specimens of the House- >

Mr. R. I. Pocock. Exhibition of, and remarks upon, young examples of the Egyptian

Bae-tanedsGerbille =. sca o fs:s%s luni o;0/alo a sielets's oon “lone Sremeeiphtttain = « siokets: srojeter ste cietersts tars 133

1. On some New and Little-known Butterflies, mainly from high elevations in the

N.E. Himalayas. By Lt.-Col. J. Maucoum Fawcerr. (Plate IX.) ........-.....-. 134

2%. On Seasonal Phases in Butterflies. By A. G. Boren Ph.D., F.LS., FZS., &lviccesss.) 149

. The Prey of the Lion. By Capt. Ricnarp Crawsway, F.Z.8. ......-0c0-.teceeeees. 144 —

. Note on an apparently Abnormal Position of the “ Brephos” within the Body of a Skink

(Chaleides lineatus). By Franx KE, Brpparp, M.A., F.R.S., Prosector to the Society.. 145

5. Contributions to the Knowledge of the Visceral Anatomy of the Pelagic Serpents Hydrus

platyurus and Platyurus colubrinus. By FRANK EK. Bupparp, M.A., F.R.S., Prosector

fo the Society sc. -jiacceteinleei 1s cintecie SPAinecataas OOD CONS Chea an OBOR EA Beane Omen ee 147

6. On the Presence of a Parasternum in the Lacertilian Genus Tiliqgua, and on the Post-

sternal Ribs in that Genus. By Frank H. Bepparp, M.A., F.R.S., Prosector to the

ISO CIGLY? 0) oie, ons arop alain nt <hevefater tm aolei obo orelegsNaltrielletataleneteiefoes C-iciai fer e cheid) «ice sianareleratcleterecamie 154

7. On the rare Rodent Dinomys branichii Peters. By Dr, Emm A. Gortpr, 0.M.ZS.,

Director of the Goeldi Museum, Pard. (Plate X.) 0.2.0.2 cee se cee e ees ns cese ne 158=

8. The Black Wild Cat of Wonsceaccaias By CO. Saruniy, of Tiflis C.M.Z.8. .......... 162

9, Ona Buffalo-Skull from East Central Africa. By R. LyprkKer...... SOUt ree nn 163

10. The Ichang Tufted Deer. By R. LyDEKKER Bra ET. ah maton Meare nr tah ge aif taba ae 166

11. On Two New Labyrinthodont Skulls of the Genera Capitosauwrus and Aphaneramma.

By A, Smiru Woopwarp, LL.D., F.R.S., F.Z.8. (Plates XI. & XIL).............. 170

LIS? OF PLATES:

1904—V OL, II.

PART ‘1,

Plate Page

I. Myzopoda aurita .ivececssacccvetceceee foie ER TEN sincerntoeeae 2

Ii. Gill-rakers of Polyodon ........+- Sie atr ans wacom ULAe aon eels 22

III. 1. Notodoris minor. 2. Trevelyana coccinea. 3. T. ceylonica.\

4. T. crocea..... Uhlan etetatehe reac te ce ate @ eicrole ole felsha\e eiete enets folnianere |

IV. 1. Trevelyana bicolor. 2. Nembrotha cristata. 3. N. afin 83

4, Marionia levis, 5. Teeth of (a) Bornella digitata, (b) B.

excepta, and (c) B. simpler ..00..sesoseee MARA RCC mRES )

Nica OEULNSE onc on cutee ayn aces selwe hae test ste eee -» 106

VI. .

VII. } Entomostraca from Natal ......... HEROS BSG we hie ade wei 12)

VIII

IX. Butterflies from the N.H. Himalayas.....-..s2.sececeeeeseee 134

X. Dinomys branickii...... Wasa eee old aricept cate wy aie em cet eye - 158

KI, Capitosaurus stantonensis ..ceresccsecceceercsecerersscsases } 170

XIL., Aphaneramma rostratum oo... .cccrssscesecessccccrserres os

NOTICE.

The * Proceedings ’ for the year are issued in four parts, forming two volumes,

as follows:—

VOL. I.

Part I. containing papers read in January and February, in June.

Ii. = ay »» -Mareh and April, in August.

VOL. II.

Part I. containing papers read in May and June, in October,

II. 5 54 »» | November and December, in April.

——.

. ‘Proceedings,’ 1904, Vol. I. Part II. was published on August 2nd, 1904.

The Abstracts of the papers read at the Scientific Meetings

in May and June are contained in this Part.

PROCEEDINGS —

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

- OF THE.

ZOOLOGICAL SOCIETY

OF LONDON.

1904, vol. I.

PARE: Pe

CONTAINING PAPERS READ IN

NOVEMBER ann DECEMBER.

ee cai inst tute,

G/ eS

APRIL 1905. ( ” WAY 26190

. 1 ean’

“ ational Nuses

ee oa

PRINTED FOR THE SOCIETY; ive

SOLD AT THEIR HOUSE IN HANOVER SQUARE.

| LONDON:

MESSRS. LONGMANS, GREEN, AND CO.,

PATERNOSTHR-ROW.

[Price Twelve Shillings.) el

LIST OF - CONTENTS.

1904.—Vot. II.

Part IL.

November 15, 1904.

The Secretary. Report on the Additions to the Society’s Menagerie in June, July, August, —

September, and October, 1904

Cec ee ee Ce

Mr. R. Lydekker. Note ona sketch of a Deer from Hainan......ceeessssceeeecucsees

et ceoee reece ow ee ee ee

Mr. F. EH. Beddard, F.R.S. Note on the voice of a young Kangaroo

Mr. Frederick Gillett, F.Z.8. Exhibition of, and remarks upon, some Antlers of the Altai

Stag, shed in the Society’ 8 Meso

eo ec Phe ote oer resem sete ee eree eee eee ee OFF e ee ee

‘Dr. P. L. Sclater, F.R.S. Note on the specimens of the Okapi in the Congo Free State

AVIVA Lh os) ec se ae Sa ietev eleln: eleva) ae tcis ie ean) Sheialwt eal aja le eaten cates} ltsy/aiqe elfen Cun\ehminte iets

Mr. W. B. Tegetmeier, F.Z.8. Exhibition of a specimen of an Asiatic King-Crab picked

up alive off the Isle of Wight .............ece cere wees OR Ap ea arer aaa: A

Page

18)

Prof. J. C. Ewart, F.R.S8. Bshibison of skins and description of a new species of Zebra

from Hast Africa

ies ievelaLelie’, e.elle @iaelele © 6.6.0, © 8 eee ee et pin) wie wile elelehe ile 21010 e0r lee \ecive s. ete ese inie 1S 6

1. On Mammals from the Island of Fernando Po, collected by Mr. E, Seimund. By

Omni ebaOMwAS, HRS. sees. (Plate XUN). a oi) \5 a apaierelarstey arererttal= i SAA pene

On Hylocherus, the Forest-Pig of Central Africa. By Ouprinip Tuomas, F.R.S., F.Z.8.

CRIS G SPAMS Do NV er. over uteea tbe Gusto shout ce sane ais deta heuiolep tenure oh epeNefeatan char fel ects plein eets

On the Species of Crowned Cranes. By P. Cuatmers Mircanit, M.A., D.Sc., Seoretary

to the Society

oe eC op mo ok ee OP Oe HH eee ecw weet toe tH oere ee *Foun tt esos eese t Foe oe eo

4. On the Mouse-Hares of the Genus Ochotona. By J. Luwts Bonnors, M.A., F.Z.8. ....

5, On some Edible and other New Species of Earthworms from the North Island of New

Zealand. By W. B. Bunuam, D.Sc. M.A, we Z8., Professor of Biology in the

University of Otago, New Zealand

eo ee ee eo ee Oe BE or ee ee HEHE ae eo ee OO eZ ores e Sanne

November 29, 1904.

Dr. Walter Kidd, F.Z.S. Exhibition of a drawing of, and ae upon, the extensor

surface of the hand of a Chimpanzee

er ee eC ee

1. Some Observations on the Field ‘Natural ae of the Lion. By oe Browarp

Orawsuay, F.Z.8.

181 |

183

220

263

Contents continued on page 3 of Wrapper.

THE ZOOLOGICAL SOCIETY OF LONDON.

Tars Society was founded in 1826 by Sir Sramrorp Rarrtzs,

Mr. J. Sasrve, 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.

COUNCIL.

HIS GRACE THE DUKE OF BEDFORD, K.G., President.

Str ALExanDER Barro, Br.

Wittum T. Branrorp, Esa., ||

C.L.E;, LL.D., F.B.S., Vice-

President.

Grorce A. Bovtenerr, KEsea.,

F.R.S., Vice-President.

Tuomas H. Burroveues, Esa.

Freperic G. D. Drewirr, Ese.,

M.D., F.R.C.P.

Hersert Drucz, Ese, F.LS.,

Vice-President.

DroumMonp,

CHARLES Esa.,

Treasurer.

Freperick Gittert, Esa. |

F. Du Cane Gopman, Hse,

D.C.L., F.RAS., Vice-President. |

| Arperr Ginruer, Ese., M.D.,

Pa.D., F.RS., Vice-President.

Sir Epmunp Gites Loprr, Br.

FE. G. B. Meape-Watpo, Esa.

P. CHatmers Mrrcnett, Ese.,

M.A., D.Se., Secretary.

Tre Hon. L.Watrer RoruscHiLp,

D.Sc., M.P.

Howarp Saunpers, Esa., Vice-

President.

Davin Saarpe, Hse., M.D., F.R.S.

OuprieLp Tomas, Esa., F.R.S.

Surron Timmis, Esa.

Cuartis §. Tours, Ese., M.A.,

F.R.S.

Aveustus F, Wiener, Esa.

The Society consists of Fellows, and Honorary, Foreign, and

Corresponding Members, elected according to the By-Laws.

The Gardens in the Regent’s Park are open from Nine o’clock a.m.

till Sunset.

The Offices (3 Hanover Square, W.), where all communications

should be addressed, are open from Ten till Five, except on Satur-

days, when they are closed at Two o’clock p.m.

The Library (3 Hanover Square), under the superintendence of

Mr. F H. Warernouss, Librarian, is open from 10 a.m. to 5 P.M.,

on Saturdays to 2 p.m. It is closed in the month of September.

The Meetings of the Socicty for General Business are held at the

Office on the Thursday following the third Wednesday in every

month of the year, except in September and October, at Four p.m.

The Meetings for Scientific Business are held at the Office twice

a month on Tuesdays, except in July, August, September, and

October, at half-past Hight o’clock p.m.

The Anniversary Meeting is held on the 29th April, at Four p.m.,

or the nearest convenient day (April 28, 1905).

_ TERMS FOR THE ADMISSION OF FELLOWS.

Frrirows pay an Admission Fee of £5, and an annual Contri-

bution of £3, due on the 1st of January, and payable in advance,

or a Composition of £30 in lieu thereof; the whole payment,

including the Admission Fee, being £35.

No person can become a Frttow until his Admission Fee and

First Annual Subscription have been paid, or the annual payments

have been compounded for.

Frtrows elected after the 30th of September are not liable for

the Subscriptions for the year in which they are elected.

PRIVILEGES OF FELLOWS.

Frttows have Personal Admission to the Gardens with Two

Companions daily, upon signing their names in the book at the

entrance gate.

Frtrows receive a Book of Saturday and a Book of Sunday Orders

every year. These Orders admit two persons to the Gardens on each

Saturday and fwo on each Sunday in the year. But the Saturday

Orders are not available if the Frntow shall have used his privilege

of personally introducing two companions on the same day.

Fstnows also receive every year Twenty Free Tickets (Green),

each valid for the admission of one adult any day of the week,

including Sunday. Children’s Tickets (Buff) can be had in lieu of

Green Tickets in the proportion of two Children’s Tickets to one

Adult's. These Tickets, if not made use of in the year of issue, aro

available for following years.

In no case can two children be passed through the gates as

one adult.

Fettows, if they wish it, can exchange the Book of Saturday

Orders for Twenty Green Tickets available for any day. The Book

of Sunday Orders can also be exchanged for a similar packet of

Twenty Tickets. These books must, however, be returned entire,

and the exchange can only be made during the year of their issue.

The annual supply of Tickets will be sent to each Fettow on the

Ist of January in every year, on his filling up a form of Standing

Order stating in what way they should be made up, and to what

address they should be sent. Forms for this purpose are supplied

on application.

The Wire of a FrLtow can exercise all these privileges in his

absence.

Ferttows have the privilege of receiving the Society’s Publications

on payment of the additional Subscription of One Guinea every

year. 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. Frriows are likewise entitled to purchase the

Transactions and other Publications of the Society at 25 per cent.

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 1871, if above the value of Five pounds.

Frrtows also have the privilege of subscribing to the Annual

Volume of the Zoological Record for a sum of £1, payable on the

1st July in each year, but this privilege is forfeited unless the

subscription be paid before the 1st of December following.

Frttows may obtain a TransreraBtn 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 Frtrow who intends to be absent from the United Kingdom

during the space of one year or more may, upon giving to the

Secretary notice in writing, have his name placed upon the

“ dormant list,” and will be thereupon exempt from the payment of

his annual contribution during such absence.

Any Frtxow, having paid all fees due to the Society, is at liberty to

withdraw his 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, M.A., D.8c.,

Secretary.

3 Hanover Square, London, W.,

April, 1905.

MEETINGS

OF THE

ZOOLOGICAL SOCIETY OF, LONDON

FOR

SCRE NAGE 1 Cy SB USS NE RYSIS:

(AT 83 HANOVER SQUARE, W.)

Session 1904-1905.

1904.

Tunspay, Novemprr1l5and29 | ‘Turspay, DecemBer 13

1905.

Tourspay, JANuaRY 17 Turspay, Aprin ..18

an Frsruary 7 and 21 s5 May oo... Zandill6é

3 MER CHangei// aes) lk es URI sooo &

The Chair will be taken at half-past Eight o'clock in the Evening

precisely.

LIST OF THE PUBLICATIONS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

Tue scientific publications of the Zoological Society of London

are of two kinds—“ Proceedings,” published im 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 attached to each annual volume of

the “ Proceedings,” to illustrate the new or otherwise remark-

able species of animals describedin them. 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,

on the first of the months of June, August, October, and

April, the part published in April completing the volume

for the last half of the preceding year. From January 1901

they have been issued as two half-yearly volumes.

The “‘ Transactions” contain such of the more important

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 25 per

cent. less than the price charged for them to the Public. A

further reduction of 25 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 £1

(which includes delivery in the United Kingdom only),

payable on the Ist July in each year; but this privilege

is forfeited unless the subscription be paid before the Ist of

December following.

The following is a complete list of the publications of the

Society already issued.

[ April, 1905. ]

TRANSACTIONS OF THE ZOOLOGICAL SOCIETY OF LONDON.

4to. 16 vols. and Index. pee

Vol. I., containing 59 Plates.... (1833-385) .... £318 6.

> Le 63 etalon ese ees

ey Sk ee /(issiiesy <0 (eo aon

5 ME A SORT kn Ok gS

BTV, « -,) tj) Die har ee en S660) me Tea me

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BTV LLL coos eat me ae CRED) io 8 8

TL eal wOGG oot Shee 7 67 7\aeeene mo ei eoRe

CR ee et age LOSES) Las WO 0 8.

index, WViolorilP Neue. ra sean. Us) sso, OY 7 6.

Vol Xi contaiming 07 Plates. (1880-35). eo 2h Om

xen Pe Une SSO aman Oa.

sxc, SS de Seas) ee G8

oO AMG a VSOIEO I de eis yf).

LEN. Se CEG) 6 BBG

POV L, Oo 88 SMe) ., & @ 0.

SCM SEIN orn Sibu ecm IIS enn Te 6.

Jil (une LCe ICO) GWT: 6”

op ee les yp gp UD pe (UH LSE). 2

PROCEEDINGS OF

THE a OF

CORRESPONDENCE OF THE ZOOLOGICAL

LONDON. 8vo. 2 vols. (Letterpress only).

Part I.

1830-31.

1 vol. 8yo.

JOU, Ite

”

eocoeeteeeece st toe

eee cece ee tw oe

Price to

Fellows.

As.

6d.

Price to the

Public.

HOH~ATAI~IO~N

SCIENCE AND

SOCIETY OF

Price to the

Public.

638.7

6s.

PROCEEDINGS OF THE ZOOLOGICAL SCCIETY OF LONDON.

8vo. 15 vols. (Letterpress only) and Index.

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an oul s34e » As Oda) ai OS: - X. 1842. - 4s, 6d. .. 6s.

oo, UU WEBS), 5 Ase Od. Os: 56 XI. 1843. 9 4s. 6d. .» Gs.

ele S36: 3 4s. 6d. .. Gs. » IL 144. s 4s. Od. .. 6s.

oy Aoulltsse y Asani OS: 5) WcllialS45: 5 As Od a eos:

og Wk ANSE), i As. Gd. .. 6s. yy) CNIVE S46. 5 4s. Od. 2. 68.F

VIL RIS39: a As. Od. 3 OS; XV, 1847, ‘ As, Gd... 6s.

gy WADE, ISO) ce ASO eos: Index 1830-1847. e 4s. 6d... 6s.

8vo. 13 vols. and Index. (Sccond Series.)

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Part XVI. 1848. 1 vol. 8vo. 4s. 6d. GSE doped tien 2 emOMecn te ell (hr

Fs XVII. 1849. $ 4s. 6d. G5: Sirens EO :8.2%.0) Seon

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oy 0 IDG Spall, 4s. 6d, Osan tiene, ete 015.9). Oe

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» AXAVIT. 1859. ‘ 4s. 6d. OB adoe terns le, 8 een Oi

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Index 1848-1860. . 4s, 6d, 6s.

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NGO ees xo) a Coen eae omen aoan wo onbOmaaGO Paco oes og 18s. eee ees:

‘ pee of merpenren Weethy oc ROM ihc snbcoor sy areas tial mde weed NS Shee nears 24s

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a jy ta IR ee Stee hala ca ae AIR areca ON oN NSA Adar ao 24'S.

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ATES SRR, Genes ee eet, era esc core aR gla es BN as Seta oe Ac DSI NG satieet. G6 24s

7) 7

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. (Highth Edition.) 8vo.

1883. Cloth, 4s. 6d.

List of the Vertebrated Animals now or lately Living in the Gardens

of the Zoological Society of London. (Ninth Edition.) 8vo.

1896. Cloth, 6s.; Paper, 5s.

Catalogue of the Library of the Zoological Society of London.

(Fifth Edition.) 8yvo. 1902. Cloth, 6s.; Paper, 5s.

These publications may be obtained at the Soctnry’s OFrricr

(3 Hanover Square, W.), at Messrs. Lonemans’ (Paternoster Row,

EL.C.), or through any bookseller.

Tit ZOOLOGICA SRBCOR D.

——203e300—

({\HE object of the Zvonogicat Recorp 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,’ after Vol. 40, will be published by

the Society at the price of 40s. per volume. But all Members of the

Zoological Society of London will have the privilege of receiving it,

including the cost of delivery, at a subscription price of 30s. per

annum. This Subscription is due on the Ist of July in every year,

and the privilege of Subscription is forfeited unless the amount be

paid before the Ist of December following.

The Zoological Society, having purchased the entire stock of

the ‘Zoological Record,’ is able to supply complete sets. The

thirty-seven Volumes to the end of the nineteenth century, and the

Index-Volume (1880-1900) in addition, will be supplied for £15

net (or without the Index-Volume, for £14 10s. net). Volumes of

any single year (exclusive of the last five volumes and Vol. 6) can

likewise be supplied at 10s. per volume net.

The price of the Index Zoologicus (Index-Volume 1880-1900)

is 20s., to Fellows 18s.

Members of the Society wishing to subscribe to the ‘ Record’

are requested to apply at this office for a Form, to be returned

when filled up and signed by the subscriber. In order to facilitate

the payment of the subscription, a Banker’s Order Form is also

supplied to those who prefer that mode of payment, This order,

when filled up and signed, should be sent to the Society’s office for

registration ; it will then he sent to the Agents named therein.

Learned Societies and Institutions and members of the former

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SEPARATE DIVISIONS OF THE ZOOLOGICAL RECORD.

At present each Volume of the Zootocican Record consists of

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XVI.

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Ceallemrerawa 55 oo oo

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Orders should be made payable to ‘‘The Zoological Society,” and

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April, 1905.

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LIST OF VOLUMES or rue ‘ZOOLOGICAL RECORD.’

The Record of Zoological Literature, 1864-1868, Vols. 1.—v.

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The Zoological Record, Volume the Thirty-sixth ; being Records

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Brown, D. Sharp, F. A. Bather, A. Willey, and R. v. Lendenfeld.

Edited (for the Zoclogical Society of London) by Davin Suarp, M.A.,

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The Zoological Record, Volume the Thirty-seventh; being

Records of Zoological Literature relating chiefly to the year 1900.

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Brown, D. Sharp, F. A. Bather, A. Willey, and E. A. Minchin.

Edited (for the Zoological Society of London) by Davin Saarp,

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The Zoological Record, Volume the Thirty-eighth; being

Records of Zoological Literature relating chiefly to the year 1901.

By J. A. Thomson, R. Lydekker, R. Bowdler Sharpe, G. A. Boulenger,

Alice L. Embleton, E. R. Sykes, E. A. Smith, 8S. Pace, Albert

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the Zoological Society of London) by Davip Suarp, M.A., Ba R asi.

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The Zoological Record, Volume the Thirty-ninth ; being Records

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Calman, E. R. Sykes, E. A. Smith, Alice L. Embleton, F. A. Bather,

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Calman, E. R. Sykes, E. A. Smith, Alice L. Embleton, F. A. Bather,

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Index Zoologicus. An alphabetical list of names of genera

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Zoological Record, 1880-1900; together with other names not

included in the ‘ Nomenclator zoologicus’ of 8. H. Scudder. Com-

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These publications may be obtained at the Socrrty’s OFrFIce

(3 Hanover Square, W.).

No. 10.

ABSTRACT OF THE PROCEEDINGS

ZOOLOGICAL SOCIETY OF LONDON?

November 15th, 1904.

Dr. W. T. Buanrorp, C.I.E., F.R.8., Vice-President,

in the Chair.

The SECRETARY read a report on the additions that had been

made to the Society's Menagerie during the months of June,

July, August, September, and October, 1904. Amongst these

several were of great interest.

Prof. J. C. Hwart, F.R.S., exhibited some skins and a series of

lantern-slides of Zebras from East Africa. He pointed out that a

variety of Crawshay’s Zebra occurred in Hast Africa in the vicinity

of Lake Nakuru. The Kast African variety differed from the

typical Crawshay’s Zebra of British Central Africa mainly in

having fewer vertical body-stripes. Prof. Hwart also pointed out

that, in addition to the Grant and Crawshay varieties, Hast Africa

had a Zebra which, in certain points, resembled the Common or

Mountain Zebra of South Africa. This new Zebra had, like the

Mountain Zebra, very long ears, a short mane, a well-marked

gridiron over the croup, and narrow hoofs. It differed from the

true Mountain Zebra in having a broad dorsal band and very

decidedly smaller chestnuts.

Mr. R. LyprxKer, F.R.S., communicated the fact that the

sketch exhibited to the Society last Session as that of a Pére

David’s Deer from Hainan was drawn by the artist from memory

and not direct from a specimen, and it could, therefore, not be

taken as evidence of the occurrence of this Deer in that country.

Mr. F. HE. Bepparp, F.R.8., described the sound emitted by a

) Wy

1 s, 0 i!

young Kangaroo that had been found in its dead mother’s pouch.

Mr. Freperick GiuLEert, F.Z.8., exhibited some antlers of the

Altai Stag (Cervus eustephanus), and made remarks upon their

growth as observed by him in the Society’s Gardens.

* his Abstract is published by the Society at 3 Hanover Square, London,

W., on the Tuesday following the date of Meeting to which it refers. It will

be issued, free of extra charge, to all Fellows who subscribe to the Publications,

along with the ‘ Proceedings’; but it may be obtained on the day of publieation

at the price of Sixpence, or, if desired, sent post-free for the sum of Sic

Shillings per annum, payable in advance.

Dr. P. L. Scuarer, F.R.S., made some remarks on the specimens

of the Okapi lately inspected by him in the Congo Free State

Museum, at Tervueren, near Brussels.

Mr. W. B. Tecermerer, F.Z.8., exhibited a specimen of the

Asiatic King Crab which had been discovered in a living condition

off the Isle of Wight.

Mr. Otprienp Tromas, F.R.S., read a paper on the Mammals

obtained in the island of Fernando Po by Mr. E.Seimund during

an expedition supported by the Society's President (the Duke of

Bedford), Mrs. Percy Sladen, and the Hon. Walter Rothschild.

Twenty-four species were enumerated, and the two following new

subspecies were described :-—

GALAGO DEMIDOFFI POENSIS, subsp. n.

Similar to true G. demidoffi, but the under surface much paler,

and the inner faces of the limbs pure white. Size as in the typical

form.

Hab. Bantabiri, Fernando Po, alt. 1800 m.

Type. B.M. no. 4.7, 1.8.

ANOMALURUS FRASERI NIGRENSIS, subsp. n.

Similar to true 4. fraseri, but size smaller and colour paler.

Length of hind foot 57 mm. Skull: tip of nasals to back of

parietals 53 mm.; length of wpper tooth series 11-9 mm.

Hab, Abutschi, Lower Niger.

Type. B.M. no, 2.11.10. 5.

A special genus, Sylvisorew, with type S. morio, Gray, was formed

to include the Shrews hitherto put with JZ, varius in Myosorex, and

another, Wimetillus, for the Bat described as Vesperugo (Vesperus)

moloneyt Thos.

A list was appended of the Mammals, 36 in number, now

known to inhabit the island.

Mr. Ouprienp THomas also read a paper, entitled “On Hylo-

cherus, the Forest-Pig of Central Africa,” and exhibited the

skulls and portions of skin of this animal obtained by Mr. R.

Meinertzhagen in the Nandi Forest, British East Africa.

This Pig had been heard of by the late Sir H. Stanley, Sir H. H.

Johnston, Mr. F. J. Jackson, and other naturalists, but Mr. Mein-

ertzhagen was the first to obtain specimens of it, and these he had

presented to the National Museum.

Hylocherus meinertzhageni, as it was proposed to be called,

proved to be a remarkably interesting type, annectant in many

vespects between the ordinary Pigs and the Wart-Hog (Phaco-

cheerus), to which latter on the whole it was most nearly allied. It

agreed with that genus in the number of its incisors, the presence

of large basisphenoid pits, the great thickness of the upper

canines (though not their length), and in the direction and shape

of the lower ones. Its molars were not hypsodont, but were of a

type very different to that found in Sus, and one from which the

peculiar hypsodont molars of Phacocherus might conceivably have

been evolved.

‘he general shape of the skull was more normal than in Phaco-

cheerus, while in the breadth of the crown there was a peculiar

resemblance to the Sus erymanthius of the Pikermi beds, though

in its tooth-structure this latter animal was a typical Sus.

Hylocherus showed no relationship to Potamocharus either in

the structure of the nasal region or the character of the lower

canines, these being of the more primitive shape found in Phaco-

cherus and the Sus verrucosus group, while in Pofamocherus they

were of the more specialized type characteristic of Sus scrofa and

its allies.

Dr. P. Caters Mircnenn read a paper on the African

Crowned Cranes, giving notes on the distribution and characters

of the species. He recorded the extension of the 1ange of the

Cape form, Balearica regulorwm, to Kavirondo, in Uganda, and

he described as new a species founded on specimens obtained by

Lady William Cecil at Khartoum and presented to the Society’s

Gardens :—

BALEARICA CECILIA, Sp. nov.

Similar to B. pavonina, and hitherto confused with it. Smaller

and darker, beak shorter and black to the tip; cheek-patch red

below, white above, but the white area very small.

Mr. J. Lewis Bonnotsr, F.Z.S., read a paper on the “ Mouse-

Hares” of the genus Ochotona inhabiting the Palearctic Region.

It was pointed out that the various species might be easily divided

into three divisions, from a study of their skulls, the differences

lying chiefly in the incisive and palatal foramina. In the first of

these, the Ladacensis group, the two sets of foramina were

entirely separated by the bony palate; in the next division,

called the Rufescens group, there was only one large foramen

slightly constricted in the centre; and in the last division, called

the Curzonie group, the large foramen was triangular in shape.

The paper dealt fully with the various species, their seasonal

changes, and distribution. One new species belonging to the

Rufescens group was described under the name 0. wardt, after

Col. A. E. Ward, who had collected it in Kashmir. It was of

moderate size, and of a uniform iron-grey in winter, becoming red,

especially about the head and neck, in summer. Since the paper

was written, Mr. Lyon, jun., in a paper on the whole of the

Duplicidenta, subdivided the genus Ochotona into three subgenera,

viz. Ochotona, Conothoa, and Pika; these subgenera coincided with

the three groups mentioned above, viz: Curzoniw, Rufescens, and

Ladacensis respectively.

A communication from Prof. W. Buaxtanp Benuam, F.Z.8.,

contained descriptions of twelve new species of Karthworms from

the North Island of New Zealand.

The next Meeting of the Society for Scientific Business will be

held on Tuesday, the 29th November, 1904, at halt-past Hight

o'clock P.M., when the following communications will be made :—

1. Capt. Ricwarp CrawsHay, F.Z.8.—Some Observations on

the Field Natural History of the Lion.

2. Sir Guartes Exior, K.C.M.G.—On some Nudibranchs from

East Africa and Zanzibar.—Part VI.

3. Mr. R. Lypexxer, F.R.S.—The Altai Lynx.

4, Mr. R. Lypexxer, F.R.S.—On old Pictures of Giraffes and

Zebras.

5. Dr. H. J. Hansen.—On the Morphology and Classification

of the Asellota Group of Crustaceans, with Descriptions of the

renus Stenetrium and its Species.

6. Mr. G. A. Bounencrr, F.R.S.—On the Lacerta depressa of

Camerano.

The following papers have been received :—

1. Mr. Roperr Gurney, F.Z.8.—On a small Collection of

Freshwater Entomostraca from South Africa.

2. Mr. W. F. Lancuesrer, M.A.—On a Collection of Sipunculids

made at Singapore and Malacca.

3. Mr. W. F. Lancuzsrur, M.A.—On a Collection of Gephyrea

from Zanzibar.

4, Mr. W. F. Lancursrer, M.A.—On the Sipunculids and

Echiurids collected during the “Skeat Expedition” to the Malay

Peninsula.

5. Mr. R. Lypexxerr, F.R.S.—On two Lorises.

6. Mr. Percy I. Laray, F.Z.8.—Descriptions of new Species of

Peruvian Rhopalocera.

7. Mr. A. D. Iums, B.Sec—On the Oral and Pharyngeal

Denticles of Elasmobranchs.

8. Dr, W.'T. Buanrorp, C.I.H., F.R.S.—Descriptions of Indian

and Burmese Land-Shells referred to the Genera Macrochlamys,

Bensonia, Taphrospira (gen. noy.), Microcystina, Huplecta, and

Polita.

9. Mr. F. E. Bepparp, F.R.8S.—Some Notes on the Cranial

Osteology of Uromastia.

10. Mr. F. E. Bepparp, F.R.8.—On the Deaths in the Mena-

gerie from January to June, 1904.

11. Dr. W. G. Rivewoop, F.Z.8.—On the Cranial Osteology of

the Clupeoid Fishes.

12. Mr. Martin A. C. Hinron.—On some Abnormal Remains

of Cervus elaphus from the Post-Pliocene Deposits of the South

of England.

13. Prof. EK. A. Muincuin, F.Z.S.— The Characters and

Synonymy of the British Species.of Lewcosolenia.

Communications intended for the Scientific Meetings of the

ZooLoGicAL Society or Lonxpon should be addressed to

P. CHALMERS MITCHELL, Secretary

3 HANovER SQuaRE, Lonpon, W

22nd November, 1904.

No. 11.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

November 29th, 1904.

G. A. Boutencer, Hsq., F.R.S., Vice-President, in the Chair.

Dr. WAtrER Kipp, F.Z.S., exhibited a drawing of, and made

some remarks upon, the extensor surface of the hand of a

Chimpanzee.

A communication was read from Capt. RicHarp CrAwsHAyY,

F.Z.S., containing some observations on the field Natural History

of the Lion made during 17 years of travel and residence in

Central Africa.

The sixth part of Sir Cartes Hxiot’s contributions “ On some

Nudibranchs from East Africa and Zanzibar” was read. It

contained an account of thirty species and varieties, of which

eight of the former and one of the latter were described as new.

Mr. R. LyprxKer exhibited and read a paper on some photo-

graphs of Giraffes and a Zebra taken from pictures in the Art

Collection at Windsor Castle, and an old print of a Zebra dated

1762. Mr. Lydekker was of opinion that the picture and print

of the Zebra had been taken from the same animal.

Tn a second communication Mr. LypExkKerr described two speci-

mens of Lorises, one a Slow Loris (WVycticebus) and the other a

Slender Loris (Loris), which had recently been acquired by the

British Museum. The latter was pointed out to be sufliciently

different from the typical Z. gracilis to be entitled to subspecific

rank,

* This Abstract is published by the Society at 3 Hanover Square, London,

W., on the Tuesday following the date of Meeting to which it refers. It will

be issued, free of extra charge, to all Fellows who subscribe to the Publications,

along with the ‘ Proceedings’ ; 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.

A paper was read from Dr. H. J. Hansen, of Copenhagen,

dealing with the Morphology and Classification of the Asellota

group of Crustaceans, and giving descriptions of the genus Stene-

ériwm and its species.

Mr. G. A. Boutencsr, F.R.S8., V.P., read a paper on the Lizard

Lacerta depressa of Camerano and its varieties.

A communication from Mr. Roperr Gurney, F.Z.8., giving an

account of a small collection of Freshwater Hntomostraca from

South Africa, was read. The collection comprised examples of

five species, three of which were described as new.

Mr. F. EH. Brepparp, F.R.8., read a paper on the Cranial

Osteology of the Egyptian Mastigure (Uromastix spinipes), based

on observations made in the Society’s Prosectorium.

The next Meeting of the Society for Scientific Business will be

held on Tuesday, the 13th of December, 1904, at half-past Hight

o'clock p.m., when the following communications will be made :—

1. The Hon. Water Roruscuitp, M.P., ¥.Z.5.—Some Notes

on Anthropoid Apes.

2. Dr. W. G. Rrpewoop, F.Z.S.—On the Cranial Osteology of

the Clupeoid Fishes.

3. Prof. EH. A. Mrncaty, F.Z.8S.—The Characters and Synonymy

of the British Species of Lewcosolenia.

The following Papers have been received :—

1. Mr. W. F. Lancurster, M.A.—On a Collection of Sipun-

culids made at Singapore and Malacca.

2. Mr. W. F. Lancunster, M.A.—On a Collection of Gephyrea

from Zanzibar.

3. Mr. W. F. Lancurster, M.A.—On the Sipunculids and

Kchiurids collected during the “‘Skeat Expedition” to the Malay

Peninsula.

4. Mr. Percy I. Laray, F.Z.8.—Descriptions of new Species

of Peruvian Rhopalocera.

5. Mr. A. D. ius, B.Sc.—On the Oral and Pharyngeal

Denticles of Elasmobranchs.

6. Dr. W. T. Buanrorp, C.I.E., F.R.S.—Descriptions of Indian

and Burmese Land-Shells referred to the Genera Macrochlamys,

Bensoma, Taphrospira (gen. nov.), Microcystina, Ewplecta, and

Polita.

7. Mr. Martin A. C. Hivron.—On some Abnormal Remains

of Cervus elaphus from the Post-Pliocene Deposits of the South

of England.

8. Mr. G. A. BovuLencrer, F.R.S.—A Contribution to our

Knowledge of the Varieties of Lacerta muralis in Western

Hurope and North Africa.

9. Mr. M. Jacopy.—Descriptions of Thirty-two new Species of

Halticince (Phytophagous Coleoptera) from South and Central

America.

10. Mr. G. 'T. Bernunr Baxer.—Notes on a small Collection

of Heterocera from the Fiji Islands, with Descriptions of some

new Species.

ll. Mr. F. EK. Bepparp, F.R.S.—A Contribution to the

Anatomy of Chlamydosaurus and some other Agamide.

12. Mr. F. E. Bepparp, F.R.S.—A Note on the Brain of

Cynopithecus niger.

Communications intended for the Scientific Meetings of the

ZOoLoGIcAL Sociery oF Lonpon should be addressed to

P. CHALMERS MITCHELL, Secretary.

3 HANovER Square, Lonpon, W.

6th December, 1904.

No. 12.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

December 13th, 1904.

Herpert Druce, Esq., F.L.8., Vice-President, in the Chair.

The SECRETARY read a report on the additions that had been

made to the Society’s Menagerie during November 1904, and

called special attention to a Ferret-Badger (Helictis personata)

from Burmah; to two Chimpanzees (Anthropopithecus schwein-

furthi) from the Bagomo Forest, Uganda; to a male specimen of

the Senegal race of the African Buffalo (Bubalus centralis) ; and

to three Mouse-Hares (Ochotona roylei) from Baluchistan. The

total number of additions during the month was 150.

Mr. OLprietD THomas, F.R.S., exhibited specimens of a pair of

Gazelles from Jerusalem which had been obtaimed and presented

to the British Museum by Dr. Selah Merrill. They represented

a new species allied to the Atlas Gazelle, and were proposed to be

named

GAZELLA MERRILLI, sp. n.

Allied to G. cwvieri, but smaller, with shorter horns, these

being more strongly curved backwards below and recurved

forwards at their tips. Premaxille not reaching nasals.

Extreme length of the typical male skull 194 mm.; horn,

length over curves anteriorly 241.

Hab. Hizmeh, N. of Jerusalem.

The Hon. Waiter RoruscuiLp, Ph.D., F.Z.8., read a paper on

Anthropoid Apes. He characterised four forms of Gorilla (Gorilla),

twelve of Chimpanzee (Simia), and four of Orang-Utan (Pongo),

* This Abstract is published by the Society at 3 Hanover Square, London,

W., on the Tuesday following the date of Meeting to which it refers. It will

be issued, free of extra charge, to all Fellows who subscribe to the Publications,

along with the ‘ Proceedings’ ; 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 Sic

Shilliugs per annum, payable in advance.

each of the last with a dimorphic phase. Of these the South

Cameroon race of Gorilla gorilla and the Gaboon White-faced

Chimpanzee were described as new. In illustration of the paper

Mr. Rothschild exhibited seven stuffed Gorillas, eight stuffed

Chimpanzees, twenty-six skulls of Gorillas, two skulls of Chim-

panzees, two entire skeletons of Gorillas, some casts of a Gorilla

and Gorilla skulls, a number of photographs and drawings of

Chimpanzees, and two, drawings of Orang-Utans. The bulk of

the mounted specimens, all but three Gorillas and two Chimpanzees,

were mounted under Mr. Rothschild’s personal supervision, from

photographs and sketches from life, by Rowland Ward; the five

exceptions were set up by Gerrard.

Dr. W. G. RipEewoop, F.Z.8., read a paper on the Cranial

Osteology of the Clupeoid Fishes, a sequel to two papers dealing

with the skull in the Hlopoid and Mormyroid Fishes which had

already been published. The skulls of Chanos, Chirocentrus,

Chatoéssus, Clupea, and seven other genera were described in

greater or less detail, and in a Summary the relative values of

such ‘“ Clupeoid” characters as the temporal foramen, auditory

fenestra, pre-epiotic fossa, pro-otic and squamosal bulle, and

posterior wings of the parasphenoid bone were discussed.

Dr. Ridewood arrived at the conclusion that Chanos had no close

affinity with the Clupeide; and he pointed out that the skull of

Chirocentrus so closely resembled that of the Clupeidee that it was

only the strongly marked differences to be found in other parts of

the body which justified the retention of the Chirocentride as a

family distinct from the Clupeide.

Professor E. A. Mincutn, F.Z.8., read a paper on the characters

and synonymy of the British species of Sponges of the genus

Leucosolenia, based on examination of the type specimens in the

British Museum and elsewhere. Three species were recognised,

namely :—JL. botryoides (Hil. & Sol.), synonyms Ascandra botrys

Heckel and A. mitida H.; LZ. complicata (Mont.), synonyms

Ascandra pinus H., Ascortis fabriciti H., Asculmis armata H.,

and possibly Ascyssa acufera H.; and L. variabilis H., synonyms

Ascortis corallorrhiza H., Ascandra tenuis Schuffner, and Leuconia

somesii Bowerbank. The paper aimed at giving an exhaustive

account of the characters and variations of the three species.

Dr. W. T. BuanForp, C.I.E., F.R.S., read a paper containing

descriptions of 18 species of Land-Mollusca belonging to the genus

Macrochlamys and its allies. All were from the British Indian

Empire, and about half were collected by the describer; some

were from Colonel Beddome’s or other collections. Of the species

described, twelve were referred to Macrochlamys, one to Taphro-

spira (nov. gen., type Helix convallata Benson), one to Bensonia,

one to Polita, two to Microcystina, and one to Huplecta.

A communication from Mr. Martin JAcopy contained de-

scriptions of a new genus and thirty-two new species of Phyto-

phagous Coleoptera of the family Halticidee from South and

Central America.

The next Meeting of the Society for Scientific Business will be

held on Tuesday, the 17th January, 1905, at half-past Hight

o'clock p.m., when the following communications will be made :—

1. Mr. W. F. Lancurster, M.A.—On a Collection of Sipun-

culids made at Singapore and Malacca.

2. Mr. W. F. Lancuuster, M.A.—On a Collection of Gephyrea

from Zanzibar.

3. Mr. W. F. Lancuestsr, M.A.—On the Sipunculids and

Echiurids collected during the ‘“ Skeat Expedition” to the Malay

Peninsula.

4. Mr. A. D. Imus, B.Sc.—On the Oral and Pharyngeal

Denticles of Klasmobranchs.

5. Mr. F. EK. Bepparp, F.R.S.—A Contribution to the Anatomy

of Chlamydosaurus and some other Agamide.

6. Mr. F. E. Brepparp, F.R.S.—A Note on the Brain of

Cynopithecus niger.

The following Papers have been received :—

1. Mr. Percy I. Laray, F.Z.S8.—Descriptions of new Species

of Peruvian Rhopalocera.

2. Mr. Marvin A. C. Hinron.—On some Abnormal Remains

of Cervus elaphus from the Post-Phocene Deposits of the South

of England.

3. Mr. G. A. Bovunencrer, F.R.S.—A Contribution to our

Knowledge of the Varieties of Lacerta muralis in Western

Europe and North Africa.

4. Mr. G. T. Beruune Baker, F.Z.8.—Notes on a small

Collection of Heterocera from the Fiji Islands, with Descriptions

of some new Species.

5. Mr. F. EK. Bepparp, F.R.S.—Some Notes on the Cranial

Osteology of Uromastin.

Communications intended for the Scientific Meetings of the

ZooLoGIcAL Soctety or Lonpon should be addressed to

P. CHALMERS MITCHELL, Secretary.

3 HANovER Square, Lonpon, W.

20th December, 1904.

peat stoi artaen ete) ? ’

Pe aa ContEyts (continued).

November 29, 1904 (dontouieaies

Pag

. On some Nudibranchs from East Africa and Zanzibar.—Part VI. By Sir C. Extor,

‘K.C.M.G., late H.M. Commissioner for the East African Protectorate, F.Z.S.

(Plates XVI. & Ba Veistatedal a alatoraheteuh Orseiatane ois apeyerrele die lartnae a) am Beatty ee ane tar evepaetals 268

3. On a small Collection of Freshwater Entomostraca from South Africa. By eee

i Gomnnys BA B.S s (Plate K VELL) 050 ir ale ea hae aie ee eal ey ee OS

4. On the Morphology and Classification of the Asellota-Group of Crustaceans, with

' Descriptions of thé Genus Stenetrium Hasw. and its Species. By H. J. Hansun,

oS SEM Le Sue (Plates: X EX Ty ee te OS ei ee ie a rn a ee ce ae 302

5. On the Lacerta depressa of Camerano. By G. A. Bovutunenr, F.R.S., V.P.2Z.8.

(Plate XXIT.) oes eee cece cette cette cette eet ee eee cette ee 332

Pere lier ch Cicitr, and Zebms, “By Ri trum 28 01 Oe 339

te On Two Lorises. By R. pies (Plate XXII.) Sh i eeiahob Neues ik le vale cone oan eae 345

December 13, 1904.

The Secretary. Report on the Additions to the Society's Menagerie in November 1904 .. 347

y P My g

Mr. Oldfield Thomas, F.R.S. Exhibition of specimens and description of a new Gazelle

from Palestine | 347

Be ee ee ec ce ee ee BS ee See ee ee tees sees serene eeerese OH etse sence Sess

1. The Characters and Synonymy of the British Species of Sponges of the Genus

‘ Leucoselenia. By H. A. Mincum, F.Z.8., University College, London ............ 349

2. Descriptions of Thirty-two new Species of Halticine (Phytophagous Coleoptera) from

South and Central America. By Martin Jacosy, F.ES. ...-......-. eee eee eens 396

3. Notes on Anthropoid Apes. By the Hon. Wattmr Rotuscuiyp, Ph.D., F.Z.8. (Plate

PNONSTIV ED) hapa sere eye Pate trey hate TRL GE lawl ic 10 agftn w So biahay Gla Ne aMeRe vaseeatoaie al aeate yoy seats Aan 415

4. Descriptions of Indian and Burmese Land-Shells referred to the Genera Macrochlamys,

Bensonia, Taphrospira (gen. nov.), Microcystina, Huplecta, and Polita. By W. T.

‘BuANKorDs C.RB ee xcH RS.) Vise iZibs (lake eNO Va aie epeeaela rete emcaaemet sata 441

5. On the Cranial Osteology of the Clupeoid Fishes. By W. G. Ripmwoop, D.Sce., F.Z.S., :

Lecturer on Biology at St. Mary’s Hospital Medical School, London .............. 448

AUN cee ane ia Sas ese Aes a RIT TUS ek tre tae) UN acon ee Comet anak ee ath a a 495

Titlepage Pee Ass a Bayesian caus Semen rete Ra ene Seen neocon nent cin ine nn a AY i

Shot Council and Omcerser. sce ctdperree ee yee rae eect ae te sere cttesact hee Ute CE li

MISO CONLSMUB seca eles orci Ge ahe, Su ciad eeHR aeeaeak arable tetra open Oma tense Tcl ERNE Sy austen ay Use teas ea iii

Alphabetical Listof Contributors 20. s aac a paves neato) sae alan ae i iyi Ge aca eee et ace ix

feet lattes = aac on stetet ele ey. 7 aie eran diem at ena se Ree eS XVil

nist ote xt OMe sii. 5 a1 Dei sine «lerelsie/ sc! ls,a aitiaveany Rar sa 5 Mack TNS i Ane a feed Latah De re dees & Xvili

istobNer Gonerio Denne ey... c's x ooo eke soir tamre gc ale Wate aie ne whe Kise Ae san aap XXIii

LIST OF PLATES. 99

1904.—V OL. ‘TI. ne

PART OT.

Plate Page

XIII. Scotonycterts bedfordi ..0+..-..sseeeeees ciel aegis misiean ne petaete 183

ae \ Hylocherus meinertzhagent ..... Wop irate rer Cae » acta ape aaa 193

XVI. 1. Phyllidia nobilis. 2, 8. Hervia lineata. 4, 5. Facelina\ *

lineata. 6. Stiliger varians. 7, 8. Hlysia marginata....+. |

XVII. 9. Madrella ferruginosa. 10, 11. Facelina lineata. Pye Stiliger \ 268

* irregularis. 13,180. Placobranchus ocellatus. 14-17. Elysia |

dubia. 18. EH. Marginatd.....veerceees cece neene BEI ae e

XVIII. South-African Freshwater Entomostraca ........--++..+-- . 298

XIX. 1. Stenetrium armatum Hasw. 2. S. mediterranewm, nu. sp.

3. S. serratum, D. SP. se cece ce cy cee tenes cece ce esenwe

KX. 1. Stenetrium serratum, n. sp. 2. S. occidentale, nu. sp. 3. S. +302

AUEULENSE, D. BPoieicce vs cecccecaz sce sence tteenracs even I

XXI. 1. Stenetrium antillense, un. sp. 2. S. siamense, n. sp. 3-6, |

Various :Asellotan. cs «sis th. wisce ale «nye als bie iminrala (ws wie ole cients )

XXII. Lacerta depressa Camerano ....- Riaithakeoaramaeraletetatens Oak Asc 332

NOK Maces Of Liorisedee ce nics cise Sx etletta eee steer tereets cic ateeetete 345

XXIV. Stmia vellerosus (Gray) (very old male) isi eserrenjitenateluer shee . 415

‘XXV. Macrochlamys and similar Land-Shells from India........ «- 441

NOTICE.

The ‘ Proceedings’ for the year are issued in fowr parts, forming two volumes,

as follows:— :

VOL. I.

Part I. containing papers read in January and February, in June. ie

Ty), bs % », March and April, in August.

VOL. II.

Part I. containing papers read in May and June, in October.

mie ri - »» November and December, in April.

‘ Proceedings,’ 1904, Vol. II. Part I. was published on October Ist, 1904.

‘The Abstracts of the papers read at ‘he Scientific Dieetines in

November and December are contained in ae ae mith,

a one

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