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ANNALS OF THE

SOUTH AFRICAN MUSEUM

VOLUME XLI

aus aS"

SMaTiona WZ

ANNALS

OF THE

SOU AFRICAN MUSEUM

VOLUME XLI

: ~

ibang

2 ites SS) :

) o>

¢, o

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US nat

* a 3

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PRINTED FOR THE

TRUSTEES OF THE SOUTH AFRICAN MUSEUM

BY THE RUSTICA PRESS (PTY.) LIMITED, COURT ROAD, WYNBERG, CAPE

1952 = 1955

TRUSTEES OF THE SOUTH AFRICAN MUSEUM

Pror. R. S. ApAmson, F.R.S.S.Afr.

Pror. C. G. S. DE VILLIERS

Mr. CounciLLor H. E. GEARING

Pror. D. L. ScHOLTz

Mr. C. J. SIBBETT, J.P.

SCIENTIFIC STAFF OF THE SOUTH AFRICAN MUSEUM

KEPPEL HArcourT BARNARD, M.A., D.Sc., F.L.S., Director.

ALBERT JOHN Hesse, B.Sc., Ph.D., Assistant in Charge of the Entomological Department.

LizuweE Dirk BoonstrA, D.Sc., Assistant in Charge of the Palaeontological Department.

Miss E. MARGARET SHAW, B.A., Assistant in Charge of the Ethnological and Numismatic

Departments.

Miss G. Joyce Lewis, Ph.D., Assistant in Charge of the Botanical Department.

A. J. H. Goopwin, M.A., Honorary Keeper of the Archaeological Collections.

‘JUL 2 8 1955

LIST OF CONTRIBUTORS

H. ANDREAE

On South African Coccinellidae

The genus Anobium

K. H. BARNARD

South African Pycnogonida ‘

South African Parasitic Copepoda

S. BREUNING

Neue Lamiinae aus dem S.A. Museum..

G. J. BROEKHUYSEN

The breeding and growth of Hymenosoma orbiculare (Crustacea, Brachyura)

J. Forest

Les Paguristes des cétes occidentales et méridionales d’Afrique

A. D. Harrison

Four new species of Gregarines from mountain cockroaches of the Cape Peninsula

J. W. JAckson

A revision of some South African Brachiopoda, with descriptions of new species ..

Z. KaszAB

Studien tiber siidafrikanischen Meloiden

D. LEsTon

Notes on Ethiopian Pentatomidae. A new genus and species from the Cape Province

Notes on Ethiopian Pentatomoidea. Some specimens from Southern Africa in the

South African Museum

N. A. H. MILiarp

New species of Hydrozoa from South Africa

K. M. F. Scotr

Some new Caddis flies from the Western Cape Province

Bo ‘'TJEDER

Two new species of Bortomyia from South Africa

E. UHMANN

Finf Dactylispa-Typen Péringuey’s

W. WITTMER

Beitrag zur Kenntnis der Malacodermata Afrikas

PAGE

215

367

381

407

359

INDEX OF NEW GENERIC NAMES INTRODUCED IN THIS

VOLUME

Carvalhocoris (Insecta, Hemiptera, Pentatomidae) Leston

Falsotragiscus (Insecta, Coleoptera, Cerambycidae, Lamiinae) Breuning

Mimogmodera (Insecta, Coleoptera, Gerambycidae, Lamiinae) Breuning

Scapogoephanes (Insecta, Coleoptera, Cerambycidae, Lamiinae) Breuning

Tambusoides (Insecta, Coleoptera, Cerambycidae, Lamiinae) Breuning. .

DALE OF ISSUE, OF THE, PARTS

Part 1, October, 1952.

Part 2, October, 1953.

Part 3, April, 1954.

Part 4, June, 1954.

Part 5, February, 1955.

Part 6, April, 1955.

345

352

S7/

348

LIST OF PLATES

Plate I.

II. South African Brachiopoda.

III.

IV. Paguristes.

Acanthochondria

Achelia

Achtheinus

Achtheres

Aeptus

Aethemenes

Agonoscelis

Agulhasia

Ainigma

Alebion

Ammothea

Amphimachus

Anaphia

Anauxesis

Ancistrotos

Anobium ne

Anoplodactylus

Antestia

Antestiopsis

Anthosoma

Argulus

Atelocera

Attalus

Austroraptus ..

Blepisanis

Boerias

Bohmia

Bolbocoris

Boriomyia ..

Botryllophilus

Brachiella

Caligus

Callipallene

Cancerilla

Carbula

Cardiodectes

Carvalhocoris. .

Cecrops

Ceroplesis

Chaetonymphon

Charopinus

Chilophoxus

INDEX OF GENERA

PAGE

Chondracanthus

Chonopeltis

Clavella

Clavellopsis

Cloniocerus

Coccinella

Colotes

Colossendeis ..

Coponia

Coridius

Crania

Cryptacrus

Dactylispa

Dalsira

Dalterus

Dasytes

Decapotoma ..

Delegorguella. .

Deroplax

Dinematura

Dinemoura

Dinemurus

Dinometopus

Diploxys

Discoarachne

Dolichopsis

Dolops

Doropygus

Dorycoris

Durmia

Dysgamus

Echthrogaleus

Elvisura

Endeis

Enterocola

Entobius

Epilachna

Eubrachiella

Eunidia

Exochomus

Falsotragiscus. .

Farnya

445

Gelastaspis

Gregarina

Gryphus

Gunenotophorus

Gynenica

Halosoma

Halydicoris

Hannonia

Hatschekia

Hedybiinus

Hedybius

Heteropallene

Hydractinia

Hyllisia

Hymenosoma

Hyperaspis

Idactus

Iselma

Kraussina

Kyphomia

Lamproglena

Legnotus i

Lepeophtheirus

Lerida 3

Lernaea

Lernanthropus

Lerneopoda

Lingula

Lytta ..

Martinella

Medesicaste

Megathiris

Megerlina

Menida 5

Metapallene ..

Mimogmodera

Miihlfeldtia

Necrobia

Nemesis

Neococalus

Nesippus

Nogagus

Nogaus ee

Notodelphys ..

Notostenus

Nymphon

Nymphopsis

Obereopsis

Oeax ..

INDEX OF GENERA

126, 129

Olenecamptus

Orthagoriscicola

Paguristes

Pallene

Pallenoides

Pallenopsis

Pandarus

Pantellaria

Parabrachiella

Parapellene

Pausias. .

Pennella

Perissopus

Petroplax

Phanodemus

Philhedonus

Philorthragoriscus

Phoxichilidium

Phoxichilus

Phytoecia

Piezodorus

Pipetta

Procidella

Prosopocera

Prosymnus

Pseudochariesthes

Pseudopallene

Pycnogonum ..

Queubus

Rhopalorhynchus

Sabelliphilus

Scapogoephanes

Sitaris ‘

Sphaeronella .

Sphyrion

Stathmodera ..

Stenoria

Stenozygum ..

Strabax

Tambusoides

Tanystylum

Terebratella ..

Terebratulina

Tetraulax

Thriocera

Trebius

Veterna

Zanclopus

Zonitomorpha

Zygophylax

107,

PAGE

170

105

4, 21

108

age - oy Sf rae ae

ANNALS

_ OF THE

SOUTH AFRICAN MUSEUM ~

VOLUME XLI

PART I, iiiae: —

7 1. A Revision fs some South ie Brash toda with Dunne: of New

- Species—By J. WILFRED es D.Sc., F.S.A., F.G.S. (With

Plates I-III.)

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1. A revision of some South African Brachiopoda; with Descriptions of New Species.—

By J. WiLrrep Jackson, D.Sc., F.S.A., F.G.S. (Late of Manchester

Museum). (With Plates I-III.)

1. Introduction

More than twenty years ago I received through the courtesy of the Director

of the South African Museum a small collection of South African Brachiopoda

for report and description. Various unavoidable circumstances prevented me

from giving the necessary time and attention to the collection, and, in conse-

quence, the report on the same has been considerably delayed.*

The collection, though a small one, is of great interest as it contains several

hitherto undescribed species and thus adds materially to our knowledge of the

brachiopod fauna of the South African Province. ‘The specimens came from

several stations extending from the neighbourhood of Saldanha Bay on the

west coast to Delagoa Bay on the east. Those from the Saldanha Bay area

include a new species of Megathiris and one of Crana. Hitherto Megathiris has

not been reported from Cape waters. From False Bay there is also a strongly-

-costate Kraussina which differs sufficiently from the well-known Avaussina rubra

(Pallas) as to merit specific recognition. In addition there is a small species of

Gryphus from the south coast, off Cape St. Blaize and Cape St. Francis, which

appears to be undescribed. ‘The remaining species comprise a new Terebratu-

lina (formerly referred to Terebratulina septentrionalis Couth.), and Terebratulina

abyssicola Adams and Reeve, previously recorded from the Cape. To these

are to be added Lingula (two species) from Durban and Delagoa Bay, and

Agulhasia davidson from two new stations, one being off Durban.

The description of the above series provides an opportunity of discussing and

revising a number of other South African species preserved in the British

Museum (Natural History) and in other collections. I have to thank the

authorities of the above institution for the privilege of examining many of their

* Additional material was sent to the author in 1949, and his revised report was received

in 1950. Post-war difficulties have further delayed the publication of this report. Ed.

VOL. XEI. PART I.

pec 8 1952

2 ANNALS OF THE SOUTH AFRICAN MUSEUM

specimens; also Messrs. H. McClelland and J. R. le Brockton Tomlin, M.A.,

for the loan of material. My own private collection has also been fully

utilized. |

The most important results achieved through the close study of this material

is the placing of the Kraussina-series on a better basis, and the removal of certain

forms erroneously ascribed to that genus. For many years this series has been

in a state of confusion and a stumbling block to students of the Brachiopoda

through the erroneous identifications that have been made.

Some particulars with regard to punctation are added in the hope that these

may prove useful in future researches on this group.

2. Previous Research

The Brachiopod fauna of South Africa is not a very large one, but it includes

some interesting species. Very little research has been conducted upon these

in recent years, and nearly all the records date from the latter part of the

eighteenth century or the first two-thirds of the nineteenth century.

The first notice of Brachiopoda in Cape waters appears to be that of Pallas,

who, in 1766,!* described Anomia rubra (now Kraussina rubra). Chemnitz, in

1785,” described the same species as Anomia striata promontorit bonae sper, and

another species, also from the Cape, as Cognata Anomia craniolaris basi perforata.

Owing to Chemnitz not being a binomial writer, his species are not recognized.

G. B. Sowerby, in 1847,? redescribed Chemnitz’s second species as Terebratula

cognata, which is now recognized as AKraussina cognata (Sow.)

In 1819, Lamarck? described a species from Mauritius as Terebratula pisum.

This species has given rise to much confusion and has been misidentified on

several occasions. It is mentioned here as being an undoubted member of the

fauna of the Natal coast. ‘The form attributed to this species by Davidson® is

not that of Lamarck: it is referred to on a subsequent page.

G. B. Sowerby, in 1847,3 described a Terebratula algoensis from Algoa Bay,

coll. J. S. Bowerbank. The specimen, a bleached ventral valve, is in the

British Museum. It has been referred to Kraussina rubra by Reeve, while

Krauss regarded it as the same as his Terebratula natalensis. A discussion of this

species is given later.

Dr. F. Krauss, in 1848,° gave three species for South African waters, viz.

Terebratula capensis (Gmel.), Terebratula cognata (Chem.) Sow., and Terebratula

natalensis Krauss (Natal). ‘The 7. capensis (Gmel.) is Kraussina rubra (Pallas) ;

T. cognata (Chem.) Sow. is Kraussina cognata (Sow.); T. natalensis Krauss has

been referred to Kraussina and considered to be the same species as Terebratula

pisum Lamarck. ‘The question as to its generic and specific position is discussed

on a later page.

In 1850,7 A. Adams and Lovell Reeve described as new two species of

brachiopods as follows:— Terebratula capensis, dredged off the Cape of Good

Hope at a depth of 120 fathoms; and Terebratula abyssicola, Cape of Good Hope,

* Superior numbers refer to the Bibliography at end.

REVISION OF SOME SCUTH AFRICAN BRACHIOPODA 3

120 fathoms. The 7. capensis Ad. and Rve. is not that of Gmelin (= Kraussina

rubra): it was changed by Davidson in 18528? to Kraussia deshayesit (later Krauss-

sina) and regarded by him as nearly related to Terebratula pisum Lam. Like the

latter, it is not a Kraussina, but goes with pisum Lam. and pisum Dav. The

Terebratula abyssicola Ad. and Rve. is now Terebratulina abyssicola (Ad. and Rve.).

In 1870,° the late Dr. Dall described a Terebratella, which he later, 1871,1°

called Terebratella rubiginosa, from Simons Bay, Cape of Good Hope. The

single specimen is in the United States National Museum. Little can be said

concerning this species until more examples are forthcoming. I have been

unable to examine the original figure, but the description of the loop suggests

that it is of a peculiar type.

A rather remarkable species was described and figured by W. King in 187114

under the name of Agulhasia davidson, the examples having been obtained on

the Agulhas Bank, South Africa, from a depth of 45 to 60 fathoms. It is a very

small species with a curious and long beak to the ventral valve. ‘The genus

is said to have Cretaceous representatives in Europe.

During the ‘Challenger’ Expedition three species of brachiopods were met

with at the Cape. These were described by Davidson in 1880° as follows:

Terebratula vitrea var. minor Philippi (p. 29, pl. II, figs. 5-6); Terebratulina caput-

serpentis var. septentrionalis Couth. (p. 33, pl. I, figs. 6-9); and Avaussina pisum

(Val, apud Lam.) (p. 54, pl. IV, figs. 7-8). All were dredged at Station 142,

off the Cape of Good Hope, lat. 35°4'S., long. 18°37’E., December 1873.

150 fathoms. Terebratula vitrea var. minor is now known as Grpphus affinis

(Calcara) and is common in the Mediterranean. The specimens dredged at

the Cape are not that species. They are referred on a later page to a new form

of Gryphus. Terebratulina septentrionalis is a distinct species and not a variety of

T. caput-serpentis (now retusa L.). It occurs at many stations in the North

Atlantic, off Northern Europe and off the eastern states of America. The

Cape specimens are distinct from that form and are dealt with later in this

memoir. The examples referred by Davidson to Kraussina pisum are specifi-

cally distinct from Lamarck’s species. These two species are also generically

distinct from Kraussina.

In 1892,” Fischer and Oehlert gave a list of nine species as inhabiting Cape

waters and remarked on the great dissimilarity between this fauna and that of

the Magellanic Province. They called attention to the recorded presence of

two species of Kraussina in the Australian region, viz. K. lamarckiana and Kf.

atkinsom. These two species, however, do not belong to the genus Araussina:

the first is the genotype of Megerlina and the second may also belong to that

genus.

In the Journal of Conchology for 1901 E. A. Smith recorded Kraussina

atkinson. (YT. Woods)—a Tasmanian species—for Algoa Bay, Cape Colony

(Brit. Mus. presented by J. H. Ponsonby). I have examined the specimens in

question—four in number (B.M. 1900.6.13.5-8)—and find them to be un-

doubted juveniles of the Terebratula prsum Lam., which is not a true Araussina.

4 ANNALS OF THE SOUTH AFRICAN MUSEUM

In my opinion, the Tasmanian species also does not belong to the genus

Kraussina, as it possesses a different type of cardinalia and brachidium.

In 1908,14 Blochmann recorded the discovery by the ‘Valdivia’ Expedition

of a small species of Liothyrina (=Gryphus) in company with Kraussina rubra and

Terebratulina abyssicola (not septentrionalis) on the Agulhas Bank. He figured

the brachidium of the Liothyrina (pl. 39, fig. 31) and regarded it as distinct from

Liothyrina (now Gryphus) affins. It is possibly the same species as that referred

to Gryphus sp. nov. in this memoir.

Dr. J. Allan Thomson, in 1918,)° listed ten species as occurring at the Cape,

and called attention to the fact that no additions had been made to the

brachiopgd fauna of South Africa during the previous thirty years. He

remarked ‘especially on the absence of Crania, and discussed the relation of the

fauna with that of Australia and other places.

Dr. W. H. Dall, in 1920,!* recorded Pantellaria (olim Miihlfeldtia) echinata

(F. & O.) as from the Cape of Good Hope, 224 fathoms (Jeffreys coll.), one

specimen in the United States National Museum. The type locality of this

species is “Off Cape Bojador, Sudan [szc.] coast of West Africa, 407 fathoms’.

In a paper in 1921,1” the present author referred to the presence in the British

Museum (Natural History) of an immature example of Muihlfeldtia truncata

labelled ‘South Africa. J. H. Ponsonby coll., 1900. 6.13.4.’ ‘The occurrence at

the Cape of this and the previously mentioned species—both Northern forms,

one Mediterranean and east Atlantic, the other West African—is very remark-

able, but one feels somewhat doubtful as to the provenance of the specimens.

In 1932, Lt.-Col. W. H. Turton*™ published an account of the marine shells

of Port Alfred. These were largely dead and drifted specimens and included

a few brachiopods, one of which resembled a Cranza,

PLATE I

Figs. 1-4. Crania roseoradiata sp. nov. Syntypes, A. 5665. Dredged in 45 fathoms 7 miles

SSW. of Constable Hill (Saldanha Bay). Four dorsal valves. South African Museum.

Figs. 5-7. Terebratulina abyssicola (Adams and Reeve). A. 5617. Dredged in 85 fathoms,

33°6’S., 28°11’E., off SE. coast of the Cape. South African Museum. Fig. 7, exterior

view; figs. 5-6, interior views.

Fig. 8. Terebratulina meridionalis sp. nov. Holotype. Station 142, ‘Challenger’ Expedition

ere miles due south of Cape Point), 150 fathoms. British Museum (Nat. Hist.).

0. 78.6.15.3.

Fig. 2: Teta septentrionalis Couthouy. Casco Bay, Maine. British Museum (Nat.

ist.).

Figs. 10-13. Gryphus capensis sp. nov. Holotype, A. 5627. Dredged in 75 fathoms, 29 miles

SW. of Cape St. Francis. Paratype, A. 5626. Dredged in 125 fathoms, 73 miles S. by

W. of Cape St. Blaize. South African Museum. Figs. 10-11: Holotype. Fig. 12: Paratype.

Fig. 13: interior of Holotype.

Plate t.

Ann. S. Afr. Mus., Vol. XLI.

South African Brachiopoda.

REVISION OF SOME SOUTH APRICAN BRACHIOPODA 5

3. Description of Spectes

ORDER ATREMATA BEECHER, 1891

Superfamily LINGULACEA Waagen, 1885.

Family LINGULIDAE Gray, 1840.

Genus Lingula Bruguiere.

1797. Ency. Meths. Vers. I, p. 151, II, pl. 250.

Lingula sp. aff. murphiana (King MS.) Reeve.

Lingula murphiana (King MS.) L. Reeve, ‘Mon. of Lingula’, Conch. Icon.,

November 1859, pl. I, fig. 3. Davidson, ‘Mon. Rec. Brach.’, Trans. Linn.

Soc., ser. 2, vol. IV, Zool. pt. III, 1888, p. 215, pl. X XIX, fig. 11, and

pl. XXX, figs. 1-3.

In 1920, when studying the brachiopods in the British Museum (Natural

History), I noted a spirit specimen in the zoological department labelled:

‘Lingula anatina Lam. Durban. J. H. Ponsonby. 99.4.14.1.’

Many years ago, Mr. R. Cairns presented me with a large example of

Lingula with a label: “Durban, r900. J. P. Cregoe.’ On making inquiries

about this collector, Dr. K. H. Barnard tells me that Cregoe contributed quite

a lot to the Invertebrate collection of the South African Museum about that

period, chiefly from Natal and Lourenco Marques; but the Museum files

contain no biographical details.

The Lingula in question agrees closely in shape and size (viz. L. 58-5: W.

26:1 mm.) with the L. murphiana figured by Reeve (op. cit., pl. 1, fig. 3) and

has the same coppery-red colour with green border anteriorly. It also agrees

with two other large examples in my collection from New South Wales (precise

locality unknown).

Reeve’s description of his species is as follows: ‘Murphy’s Lingula. Shell

oblong-ovate, anteriorly square, posteriorly attenuated, umboes small, sharp,

yellowish-green, stained with coppery-red; valve closed throughout.

King MS. Mus. Cuming.

Hab. Moreton Bay, Australia; Strange.’

The specimen figured is, L. 57-8: W. 25:7 mm.

He further remarks: “Whether this should be regarded as an Australian

form of L. anatina or as a distinct species, it is certain that the differences are

obvious and constant. ‘The late Captain Phillip King, who was an excellent

conchologist . . . was struck by the peculiarities of this Lingula, and sent speci-

mens home to Mr. Cuming with the above name. More have been collected

by Mr. Strange in nearly the same locality, and all are distinguished from

L. anatina collected abundantly by Mr. Cuming in the Bay of Manila, by a

more square outline, and by a peculiar coppery-red tone of colour.’

Davidson (op cit.) figures two large examples from Moreton Bay in his

collection at the British Museum, and gives an amplified description and the

colour as coppery-red, with bands of different shades of green and brown. He

6 ANNALS OF THE SOUTH AFRICAN MUSEUM

gives the size as: L. 2 in. 6 lines2 W..c,1n. 1, dine (=L. 63:5; W227-7isasane

The size of the specimen in his plate XXX, figs. 1-3, 1s somewhat less, being:

L. 53°5: W. 24 mm.

Lingula murphiana, though first described from Queensland, appears to have

a wide distribution. According to W. H. Dall’s List bof 1920, 16 the United

States National Museum possesses the following: N.E. Australia, coll. Stearns,

2 specimens; Australia, coll. Cuming, 2 specimens; Fiji Islands, U.S. Ex. Exp.,

2 specimens; Vita Leva, near Rawa River, U.S. Ex. Exp., 2 specimens; and

off Shimbaya Gulf, Japan, E. S. Morse, 1 specimen.

One wonders how much reliance has been placed on colour in these determi-

nations. It seems to be a variable character in some forms.

I have in my possession four very large specimens of the same shape and

colouring as the Queensland form collected at Karachi by Lt.-Col. H. C.

Winckworth in September 1932 and given to me by his brother Mr. R. Winck-

worth. The sizes of these are: 64 X 28-1; 62-4 x 29; 60°8 X 29; and 59°3 X

27°7 mm. If these had been found at the Queensland locality, they would

have been referred to L. murphiana Rve.

From the same collector I have a smaller Lingula found at Trimenuaai

Ceylon, October 1929, also rather dark in colour with a green border. It

measures: L. 43:3; W. 18-8 mm. _ It might very well be a younger stage of the

above examples.

Dall!® cites Lingula translucida Dall—a ruddy-brown species—from aes

coll. Captain Shopland, 2 specimens and Fulton, 2 specimens, but does not

figure the species. He also cites the same form from Java, coll. Ward, 2 speci-

mens and Palmer, 1 specimen, and from Tataan Island, Philippines, 12 fms.,

coll. Bureau of Fisheries, 1 specimen. It is uncertain which is the type station.

Dall gives Java, but cites the catalogue number for Karachi.

With regard to the Lingula anatina of authors, this Eastern Seas species

(Type station, Amboyna, Moluccas) was referred to the Patella unguis of Linne

by Dall in 1920,%¢ but the shell to which Linné refers in Rumphius’ D’ Amboin-

ische Rariteitkamer, 1741, t. 40, fig. L, is a Parmophorus or Scutus and not a

brachiopod. Solander, in his Portland Catalogue, 1786, No. 1718, p. 77,

mentions Mytilus lingua or green Duck’s-bill Limpet, $., Humphrey's Conchology,

pl. 2, fig. 2, from Amboyna. This last work was published in 1770 and the

figured shell is certainly a Lingula. Though the name anatina has been in

use for a long time, the species should be known as Lingula lingua (Sol.).

Lingula sp. aff. exusta Reeve

Lingula exusta Reeve, ‘Mon. of Lingula’, Conch. Icon., November 1859, pl. II,

fig. 9. Davidson, ‘Mon. Rec. Brach.’, Trans. Linn. Soc., ser. 2, vol. IV,

Zool. pt. III, 1888, p. 217, pl. XXVIII, figs. 20-214.

Dr. K. H. Barnard has recently submitted to me two specimens of a Lingula

which he collected in 1912 at Delagoa Bay, Indian Ocean, S.E. Africa. The

specimens are rather small and measure: 20 x 8 and 17:2 X 64mm. Both

REVISION OF SOME SOUTH AFRICAN BRACHIOPODA yi

are light brown in colour and resemble the Lingula exusta of Reeve, though

about half the size of the type.

Reeve describes the species from a specimen in the Mus. Cuming as follows:

“The Swarthy Lingula. Shell oblong-ovate, rather thin, reddish-yellow,

deeply stained with brown towards the margin, shining, umboes rather sharply

beaked.

Hab. Moreton Bay, Australia; Strange.’

He further states: ‘If L. murphiana be an Australian form of L. anatina, this

might be regarded as the representative in the same locality of L. hians. Both

species exhibit a peculiar coppery-redness, heightened in this to a dark, shining,

swarthy tone of colour.’

Davidson (op. cit.) gives a fuller description mentioning its shining, darkish,

coppery yellow-brown colour. He reproduces Reeve’s figure in fig. 20;

fig. 21 being from a specimen from the same locality in his collection at the

British Museum. He gives the length as 1 in. 7 lines and breadth 8 lines

(10-6 < 17-1 mim.).

Dall, in his 1920 List (#*p. 265) gives Brammo Bay, Dunk Id., N. Queens-

land, coll. E. J. Banfield, 1 specimen, in the United States National Museum. °

This is much farther north than the original locality.

I possess some half-dozen specimens of a Lingula from Singapore with

similar colouring and shape to the Delagoa Bay examples, one or two being

only slightly larger.

There is need for further research in connection with the various forms of

Lingula, as there appears to be much confusion with regard to species.

The form known as L. anatina (now referred to lingua), though normally of a

light green colour (as in Reeve, 1841, pl. II, fig. 10), is at times suffused with a

bronzy colour, especially near the margins, or may be of a dark brown colour

all over the surface, as in specimens in my collection from Amboyna, Moluccas

(‘Siboga’), and Zamboanga, Philippines (coll. F. G. Pearcey, 1875).

OrDER NEOTREMATA BEEcHER, 1801.

Superfamily cRANIACEA Waagen, 1885.

Family CRANIIDAE King, 1846.

Genus Crania Retzius.

1781. Schrift. Berlin Ges. nat. Fr. 2 (4), p. 72.

Crania roseoradiata sp. nov.

(Pl. I, figs. 1-4.)

Description: Shell (upper, or dorsal, valve) small, depressed, somewhat ovate

in outline, with a short, almost straight posterior margin, apex low, subcentral,

directed posteriorly; surface eroded, strongly wrinkled by growth-lines, and

radiated by rose-coloured streaks or flames on a whitish background. Interior

very finely granulose, margin thin and narrow beyond the heavy submarginal

encircling ridge. Muscular impressions of posterior adductors well-marked,

8 ANNALS OF THE SOUTH AFRICAN MUSEUM

oval, obliquely disposed on low tubercles within the posterior angles of the

shell; imprints of protractors in small pits on external side and near the anterior

extremities of posterior adductors impressions; imprint of median odd muscle

very slight and situated on floor of valve between the posterior adductors;

imprints of anterior adductors very distinct in ovate pits with strong posterior

ridges, closer together than posterior pair and situated just behind the centre

of the valve; small pits for insertion of retractors of arms immediately adjacent

and external to the last; imprints of protractors of arms very slight on floor of

valve in front of anterior adductors. Anterior to the muscle impressions are

two shallow, ovate, cavities (one on either of the median line) showing indis- ©

tinctly the grooves of the pallial sinuses.

Dimensions: Syntypes, Length, 9 mm., breadth, to mm.

Mee yay soqueous 6 g mm.

Catalogue No. A5665, in the South African Museum.

Type-locality: 7 miles SSW. of Constable Hill, Saldanha Bay, dredged

in 45 fms., with Megathiris sp. nov.

Remarks: This is the first authentic Crania to be reported from South

Africa. In 1931, Lt.-Col. W. H. Turton submitted for my opinion a very

imperfect dead valve, 12 X 11 mm., from Port Alfred which I tentatively

referred to Crania. In 1932, Turton figured the specimen as Crania sp.

(pl. LXX, fig. 1843). Port Alfred lies nearly 500 miles to the east of Saldanha

Bay. The nearest record of the genus appears to be a species from West

Africa assigned by Reeve?’ to Crania rostrata Hoeninghaus. Four examples of

this form are in the Cuming Collection at the British Museum (Natural History).

Two are attached to fragments of dark-grey limestone and are the originals of

the two lower figures of Reeve (8pl. I, fig. 3). Davidson!® places Crania

rostrata among the synonyms of C. turbinata (Poli), but says ‘the so-termed

C. rostrata figured by Reeve in his Conch. Icon. does not agree with Hoening-

haus’s figures of his species’. He misquotes Reeve by giving ‘South Africa’

instead of ‘West Africa’ (vide Cuming) as the habitat. One of the British

Museum specimens shows interior details and the ventral valve agrees closely

with the figure of the interior given by Hoeninghaus.2® The position and

number of the pallial sinuses are the same and there is the same arrangement

PLATE II

Figs. 1, 2, 7, 8, 14, 15. Megathiris detruncata (Gmelin). Messina, Mediterranean (J. W. J.

coll.). Figs. 1-2: interior views of ventral and dorsal valves. Figs. 7-8: exterior views

of same. Fig. 14: interior view of dorsal valve (fig. 2). Fig. 15: interior view of ventral

valve (fig. 1).

Figs. 3-6, 9-13, 16,17. Megathiris capensis sp. nov. Holotype, A. 5616a. Paratype, A. 5616b.

Dredged in 45 fathoms, 7 miles SSW. of Constable Hill (Saldanha Bay). South African

Museum. Figs 3, 4: interior views of ventral and dorsal valves of Holotype. Figs. 5, 6:

interior views of ventral and dorsal valves of Paratype. Figs. 9, 10: exterior views of

ventral and dorsal valves of Holotype. Figs. 11, 12: exterior views of ventral and dorsal

valves of Paratype. Fig. 13: exterior view of dorsal valve of Holotype. Fig. 16: interior

view of dorsal valve of Holotype. Fig. 17: lophophore in spirit specimen.

Ann. S. Afr. Mus., Vol. XLI. Plate 1.

South African Brachiopoda.

REVISION OF SOME SOUTH AFRICAN BRACHIOPODA 9

with regard to the muscular impressions and prominent rostrellum. The

dorsal valve was not figured by Hoeninghaus so no comparison can be made

here. Until a much-desired revision is made of the various forms attributed

to either Crania anomala (Miill.) or Crania turbinata (Poli) little advance in

knowledge can be attained. Hoeninghaus’s type of Crania rostrata appears to

have come from the Mediterranean.

The absence of the ventral valve in the new Cape species renders it impossible

for a complete comparison to be made with the West African C. ‘rostrata’; but

a comparison of the dorsal valve shows that the two forms are sufficiently

distinct to merit a new name being given to the Cape species.

It is of some interest to compare C. roseoradiata with the C. suessi Reeve.1§

The latter species was founded on five specimens in the Cuming Collection

reported to have been collected by Mr. Strange at Sydney. It is described as

being faintly tinged with orange-red. ‘The five specimens are in the British

Museum and are now rather bleached: they possess somewhat yellowish rays.

They bear a label in German? handwriting (?Pfeiffer’s) ‘Sidney, M. Strange’

(note misspelling). In general form they somewhat resemble the West

African examples described and figured by Reeve as C. rostrata Hoen., and the

coloured rays of the upper valve suggest a relationship with the C. roseoradiata

of this memoir. Cyania suessi Reeve has also been recorded from Mast Head

Reef, Queensland, 17-20 fathoms.”4 This record is based upon a few worn

valves and there is some hesitation as to species.

The occurrence of Crania at the Cape is an interesting addition to the South

African fauna, but until a further study has been made it is difficult to ascertain

whether it is of northern or southern facies.

OrvEeR TELOTREMATA BeEEcHER, 1891.

Superfamily TEREBRATULACEA Waagen, 1883.

Family TEREBRATULIDAE Gray, 1840.

Subfamily CANCELLOTHYRINAE Thomson, 1926.

Genus Agulhasia King.

1871. Ann. Mag. Nat. Hist. (4), vil, p. 109.

Agulhasia davidsom King

Agulhasia davidson King, loc. cit., 1871, p. 111, pl. XI, figs. 1-7.

Terebratulina (Agulhasia) davidson’ King. Davidson, ‘Monograph of Recent

Brachiopoda’, Trans. Linn. Soc., ser. 2, vol. 1V, Zool. pt. I, 1886, pp. 36-7,

pl. 7, figs. 1-5.

Agulhasia davidson King. Thomson, Brachiopod Morphology and Genera (Recent

and Tertiary), 1927, pp. 182-3, fig. 52 (after Davidson).

Remarks: The original specimens of this interesting species were obtained

from 45 to 60 fathoms, on the Agulhas Bank, South Africa, and were described

by Professor W. King in 1871 (op. cit., supra). I have been unable to examine

the original examples, but the excellent description by Davidson (pp. 36-7)

10 ANNALS OF THE SOUTH AFRICAN MUSEUM

shows that the species is a remarkable one and sufficiently distinct from the

young of some species of Terebratulina, as was at one time thought.

This extremely small species is characterized by the beak being produced

into a tubular rostrum and by the possession of a long pedicle-collar (see

Davidson,” pl. 7, figs. 1a and 2). :

Through the kindness of Dr. K. H. Barnard I have been able to examine

three specimens of this species dredged by the Cape Government trawler

s.s. Pieter Faure from two new stations, one off the south-eastern Province, and

the other off Natal.

The details are as follows:

Lat. 34°S., long. 25°44’E. dredged (no depth given). 2 specimens.

A. 5633. South African Museum. Size: L. 4:2, W. 2°5 and L. 4:0, W. 2-3 mm.

Cape Natal (i.e. Durban), W. x N. 6 miles, 54 fathoms, 1 specimen. A. 5634.

South African Museum. Size: L. 4:2, W. 2:7 mm.

These specimens are smaller than the size given by Davidson, viz. 3 lines x

2 lines (= 6-3 X 4:6 mm.).

The specimens are interesting, not only in providing evidence of an extended

eastern range of the species, but also on account of the opportunity they offer

of studying the peculiar foraminal characters. As pointed out by Thomson

(8p. 74 and footnote), ‘the long beak is apicate, with a very long delthyrium

which is partly filled by lateral trigonal deltidial plates, leaving an elongate

pedicle-opening. ‘The pedicle-collar extends nearly to the cardinal margin,

and has the effect of restricting the pedicle mainly to an opening on the

cardinal margin.’ Davidson’s figures ('*pl. 7, figs. 1a and 2, repeated by

Thomson, fig. 52) show this quite clearly.

King (op. cit.) confused the deltidial plates with the area, and the pedicle-

collar with the deltidium.

Beecher (?°p. 390) referred to the deltidial plates as being obsolete; this

being probably due to the imperfect descriptions by King and Davidson.

Unfortunately details are lacking concerning the character of the animal,

though the cardinalia and brachidium have been figured.

The range of Agulhasia has been given as Cretaceous to Recent, but no

Cretaceous forms have been cited. Terebrirostra or Lyra from the Cretaceous

possesses a somewhat similar tubular beak, but with the foramen at the apex:

it belongs to an entirely different Family.

Genus Terebratulina d’Orbigny.

1847. C.R. Ac. Sc. Paris, XXV, p. 268.

Terebratulina abyssicola (Adams and Reeve)

(Pl. I, figs. 5-7)

Terebratula abyssicola Adams and Reeve, Voy. Samarang, Moll., 1850, p. 72,

pl. 21, fig. 5: Reeve, “Terebratula’, Conch. Icon., 1860, pl. 4, fig. 14.

Terebratula radiata Reeve, “Terebratula’, Conch. Icon., 1860, pl. 3, figs. 7a-b.

REVISION OF SOME SOUTH AFRICAN BRACHIOPODA II

Terebratulina abyssicola, Davidson, ‘Mon. Rec. Brach.’, Trans. Linn. Soc., ser. 2,

vol. IV, Zool. pt. I, 1886, pp. 37-8, pl. 5, fig. 54.

Terebratulina radiata, ibid., 1886, pp. 34-5, pl. 6, figs. 9-14.

Terebratulina africana Turton, The Marine Shells of Port Alfred, South Africa, 1932,

p. 260, pl. LXX, fig. 1842.

Non Terebratulina radiata Baily, Quart. Journ. Geol. Soc., vol. XIV, 1858, p. 136,

pl. VIII, figs. 3a-d (= Jurassic species from Crimea).

Non Terebratulina radiata Moore, The Geologist, vol. III, 1860, p. 444, pl. XIII,

figs. 11-14. (= Great Oolite species from Hampton Cliff, England).

Habitat: South-east coast of South Africa, lat. 33°6’S., long. 28°11’E., 85

fathoms (= opposite East London), 3 large examples A. 5617, South

African Museum: also 4 miles off Cape Morgan, 17 fathoms, 8 small

examples, A. 5618, South African Museum.

Remarks: ‘There has been much confusion in the past with regard to this

species, its type locality, and its relation to the T. radiata Reeve. The latter is

merely the adult form of 7. abyssicola Ad. and Rve., which antedates 7. radiata

by ten years. The type-locality is ‘Gape of Good Hope (dredged at the depth

of 120 fathoms); Belcher’, as given by Reeve. The reference to Corea in

Reeve’s ‘Corrigenda’ is evidently a mistake. In the Zoological Department

(Cuming Collection) of the British Museum are four specimens (three juveniles

and one large example attached to some extraneous object) of T. abyssicola,

with a legend on the bottom of the box ‘Hab. Cape of Good Hope, 120 fms.

M.C. ?Type’. The attached specimen looks like that figured in the ‘Samarang’

Report (pl 21, fic. 5) and in Conch. Icon. (pl. 4, fig. 14), but is smaller

- and more globular. It shows the beginning of a sulcus in the dorsal valve, and

the striae are like those on the umbonal part of a South African Museum

specimen (A. 5617). The British Museum also possesses specimens of T.

abyssicola from Port Alfred, Cape Colony (Coll. Lt.-Col. Turton, 1903, 12.19.

463-5). Davidson’s figure of T. abyssicola (pl. 5, fig. 54) is a poor copy of

that of Adams and Reeve.

As far as one can judge from a casual microscopic examination, the punctae

of the “Type-specimen’ of 7. abyssicola are the same as in Reeve’s type of T.

radiata, being small and slit-like and of the same density. ‘The type-specimen

of 7. radiata is preserved with two other examples in the Zoological Department

at the British Museum (Cuming Coll.), while the originals of Davidson’s

figures of the same species are in the Geological Department (Davidson Coll.)

of the same Institution. The original of jDavidson’s T. radiata (pl. 6,

fig. 11) has small, ovate punctae (slit-like) with the same density as Reeve’s

type.

The exact density of the punctae of the British Museum specimens has not

been counted, but examples of T. abyssicola from other sources have been

examined. Five somewhat dwarfed specimens from East London yielded the

following results: in one the density was 352-368 per sq. mm., with dimen-

sions of 30-40 X 15 »; in three others the density ranged from 448 to 512 per

12 ANNALS OF THE SOUTH AFRICAN MUSEUM

sq. mm., the dimensions being 20-30 X 15 pu; while the fifth example was very

densely punctate with 576 pores per sq. mm., with dimensions of 15-20 X 15 p.

Four others from East London yielded 384 per sq. mm., and two from Port

Elizabeth, 384-432 per sq. mm.

The three large examples, A. 5617 (pl. I, figs. 5-7), in the South African

Museum are larger than any figured by Davidson (pl. 6, figs. 9-14), their

dimensions being:

Length Breadth Thickness

No. 1 24°0 1Q°2 15;0 mm.

No. 2 27°5 19°3 LO-5) as

No. 3 28-4 29°0 16:0 E

The type of folding is sulciplicate (as Davidson, pl. 6, fig. 10), and all

are rayed with black flames or streaks. ‘The beaks are mesothyrid and the

deltidial plates disjunct.

The number of pores per sq. mm. in these three examples ranges from

308-400, the dimensions being 30-40 X 15 p, on the outer side, and 10 X I0 pn

on the inner. )

The eight small examples, A. 5618, in the South African Museum range in

length from 4:5 to 13-4 mm., the beaks are submesothyrid to mesothyrid and

the deltidial plates disjunct. They possess a uniplicate type of folding, the

larger ones showing the beginning of the sulciplicate stage. ‘Some are marked

with black streaks, and all agree closely with shallow water forms in my

collection from East London and Port Elizabeth. Some of these appear to be

stunted or dwarf examples and show irregularities of growth: the shell also is

much thickened.

This species under the name of 7. radiata was cited as a new brachiopod for

Western Australia by W. B. Alexander, in 1914.26 The solitary specimen

obtained had united deltidial plates, and this fact, together with its habitat,

seems to suggest the Terebratulina (olim Terebratula) cancellata (Koch), since

transferred to a new genus Cancellothyris by J. A. Thomson,?’ who also changed

Puiate III

Figs. 1, 2. Kraussina rubra (Pallas). Agulhas Bank, South Africa, 22 fathoms (J. W. J. coll.).

Fig. 1: exterior view. Fig. 2: interior view of dorsal valve.

Figs. 3-5. Kraussina crassicostata sp. nov. Holotype, A. 5615. Dredged in 23 fathoms, False

Bay, Cape. South African Museum. Fig. 3: exterior view. Figs. 4, 5: interior views

of dorsal and ventral valves.

Figs. 6 and 9. Megerlina striata sp. nov. Holotype, Station 142, ‘Challenger’ Expedition

(nearly 40 miles due south of Cape Point), 150 fathoms. British Museum (Nat. Hist.).

No. 78.6.15.27. Fig. 6: exterior view. Fig. 9: punctation.

Figs. 7 and 10. Megerlina pisum (Lam.). South Africa, J. H. Ponsonby coll. British Museum

(Nat. Hist.). No. 99.4.14.3765. Fig. 7: exterior view. Fig 10: punctation in specimen

in J. W. J. coll. (ex. Agnes Crane, from Cape, 150 fathoms).

Fig. 8. Megerlina capensis (Adams and Reeve). Cape. Lombe Taylor coll. British Museum

(Nat. Hist.). No. 74.12.11.386.

Ann. S. Afr. Mus., Vol. XLI. Plate 1.

- a fe»

e",a' vo

e¢? © + git,2n%s.s

Fen”) § |

PF Cette d. ahh

eo % 4 , § t; - -

as P a a &

South African Brachiopoda.

REVISION OF SOME SOUTH AFRICAN BRACHIOPODA 13

the specific name to australis on account of cancellata being preoccupied by Tere-

bratula cancellata Eichwald, 1829. But, unless the 7. cancellata Eichwald turns

out to belong to Cancellothyris also, there appears to be no reason for the change

of the specific name.

Turton, in 1932 (pl. LXX, fig. 1842), figured as Terebratulina africana sp.

nov. a small example from Port Alfred, near which place TJ. abyssicola is

common. His figure shows a shell without apparent folding, the ventral valve

having a submesothyrid foramen, disjoint deltidial plates and a pedicle-collar:

the dorsal valve has rather long crura and an almost completed loop. The

shell is thin and white with radiating striae and fine growth-lines, giving it a

cancellated appearance, and does not seem to show evanescence of the striae,

as in the next species. The absence of information regarding punctation

renders it difficult to decide whether Turton’s shell is to be referred to T.

abyssicola, which occurs in the near neighbourhood. Turton’s T. radiata

(*4p. 260) is a synonym of the last species.

Terebratulina meridionalis sp. nov.

(Pi le fie. 3)

Terebratulina caput-serpentis var. septentrionalis Davidson (non Couthouy), Voy.

Challenger, Zool. vol. I, 1880, pp. 33-6, pl. I, figs. 6-9.

Terebratulina septentrionalis Davidson (pars), ‘Mon. Rec. Brach.’, Trans. Linn.

bee ser: 2. vol. LV, Zool. pt. I, 1886, pp. 26-32.

Description: Shell of medium size, ovate, with curved sides and slightly

truncated front, widest at the middle. Valves regularly convex, ventral

slightly deeper than the dorsal and possessing a slight, narrow, distinct sulcus

anteriorly. Folding uniplicate. Surface of valves finely capillate, striae strong

and simple in umbonal region, increasing by bifurcation, and becoming

- evanescent later; finely punctate. Beak short, suberect, foramen large, almost

circular, submesothyrid, attrite; deltidial plates disjunct; pedicle-collar short.

Hinge teeth without dental plates or swollen bases. Cardinalia in dorsal

valve as is usual in Terebratulina, with socket-ridges fused to the crural bases and

forming prominent ridges which project a little behind the dorsal umbo;

small transverse cardinal process; no median hinge-plates. Crura long,

descending branches of loop short, transverse band arched ventrally, crural

processes united and transforming the loop into a ring. Muscular impressions

feeble.

Dimensions: Holotype, length 22 mm., breadth 18 mm., thickness 9:5 mm.

Holotype and Locality: Specimen in the British Museum (Nat. Hist.),

dredged by the ‘Challenger’ Expedition at Station 142, Cape of Good Hope

(= nearly 40 miles due south of Cape Point) in 150 fathoms.

Remarks: In his ‘Challenger’ Report, Davidson (p. 33, pl. I, figs. 6-9)

recorded the Terebratulina septentrionalis Couthouy (as a variety of T. caput-

serpentis = retusa) as abundant at Station 142, lat. 35°4’S., long. 18°37’E., off

the Cape of Good Hope, in 150 fathoms, together with ‘Terebratula vitrea var.

14 ANNALS OF THE SOUTH AFRICAN MUSEUM

minor’ and ‘Kraussina pisum’. In the same work he recorded the species from

lat. 46°40’S., long. 37°50’E. (= Marion Island) in 150 fathoms, associated

with ‘Platydia anomioides’ and *Waldheimia kerguelensis’ .

In his later work (p. 28), Davidson appears to cast some doubt upon the

authenticity of the Cape locality, but at the same time gives the Marion

Island latitude and longitude in error for the Cape.

By the courtesy of the British Museum authorities I have been able to

examine the ‘Challenger’ examples and to compare them with typical T.

septentrionalis Couthouy from Casco Bay, Maine, and other places. Some of

these are preserved in the Zoological Department of that Institution, and I was

surprised to find that one box, registered 78.6.15.28, contained two distinct

species, one with evanescent striae and the other (a large, solitary example)

agreeing with the true T. septentrionalis from Halifax, 83 fathoms (‘Challenger’

specimens, 78.6.15.24). I came to the conclusion that the true T. sepien-

trionalis had probably been used for comparison and accidentally left in the same

box with the others.

At the same Institution I also had the privilege of examining some specimens

preserved in spirit and coming from Station 142. These I found agreed

exactly with the examples mentioned above which showed evanescent striae;

and like them differed sufficiently from T. septentrionalis to merit the erection

of the present new species.

Compared with a spirit specimen of T. septentrionalis from Casco Bay, Maine,

of almost the same size, viz. length 22 mm., breadth 17 mm., thickness 11 mm.

(pl. I, fig. 9), the new species shows important differences in striation as well

as in punctation. The striation of 7. meridionalis is evanescent (pl. I, fig. 8)

and contrasts strongly with that of T. sepientrionalis (pl. I, fig. 9). The number

of striae is about the same in both species. On the outer surface of the ventral

valve, about the middle, 7. meridionalis has small ovate pores, 20-30 X 15 pw

in diameter, and 256-288 per sq. mm. In T. septentrionalis from Casco Bay,

the pores are small and rounded, 15-20 X 15-20 p, and 320-384 per sq. mm.

at the middle of the ventral valve. Anteriorly both species are very similar in

the possession of a slightly truncated front and low dorsal uniplication. The

foramen in each species is also submesothyrid and there are rudimentary

deltidia.

In addition to the British Museum material I have had the advantage of

comparing others in my own collection as well as examples in the South African

Museum. These are all from essentially the same neighbourhood as the

‘Challenger’ specimens and confirm the conclusion reached above. Specimens

from ‘Off Cape Point, S. Africa, 190 fathoms’ (in my own collection, ex South

African Museum) agree exactly with the ‘Challenger’ example from Station

142 (p. I, fig. 8). They are smaller in size, but low uniplication is visible. They

have small, ovate pores 256-320 per sq. mm., and 20-30 X 15 p in Size.

Three small specimens in the South African Museum, A. 5623, dredged in

85 fathoms, off Cape Point, 10 miles S. 16° W., belong to the same species, and

REVISION OF SOME SOUTH AFRICAN BRACHIOPODA 15

a very young example in the same Museum, A. 5632, dredged at the same depth

and place as the above, is probably the same.

The T. ‘septentrionalis’ and the two other species of brachiopods recorded by

Davidson from Marion Island require further investigation in the light of

present knowledge.

With regard to ‘Waldheimia kerguelensis’, the figures given by Davidson

(pl. III, figs. 3-5) suggest a somewhat different species from that under the same

name from Kerguelen Island (pl. III, fig. 1).

Terebratulina sp. indet.

Included in the South African Museum material are several juvenile examples

of a Terebratulina which must remain unidentified pending further material.

The specimens are all from the western side of the Cape—the opposite side

from which T. abyssicola was dredged. ‘Three lots are from the neighbourhood

of Table Bay, and two lots are from off Saldanha Bay. Some notes on these

may be useful in connection with further dredgings in these waters.

A. 5620. ‘Table Bay, 22 fathoms (South African Museum).

There are three juvenile examples from this locality which possess a rather

coarser and more distinct ornament than T. meridionalis. Near the umbo

there are 13 or more strong, rounded striae which soon break up into fascicules

of finer striae by bifurcation and trifurcation. ‘There are also a few inter-

polated fine striae. All extend to the anterior margin with occasionally a

second bifurcation at a late stage. The beak is short, suberect; foramen large

and almost circular, submesothyrid, attrite. The deltidia in two examples are

just touching at the points, but in the largest example they are disjunct

(perhaps broken or worn by the umbo of the dorsal valve). In the smallest

example the folding is uniplicate and in the two others slightly sulciplicate.

The species presents some resemblance to 7. abyssicola, but differs in punctation.

It has long oval pores on the outer surface ranging from 224 to 288 per sq. mm.,

the dimensions being 30 (-45) X 15-20 pw. On the inside of the valve they

are round and measure 15 X 15 wp.

A. 5621. 28 miles S. 76° W. of Lion’s Head, Cape Peninsula, 140 fathoms.

Four juvenile examples (South African Museum).

The punctation in these ranges from 272 to 288 per sq. mm., the diameters

being 20 X 15-20 p, on the outer side.

A. 5619. 34 miles S. 63° W. of Lion’s Head, Cape Peninsula, 154 fathoms.

Four juvenile examples (South African Museum).

The punctation ranges from 272 to 304 per sq. mm., with diameters of 15-20

X 10-15 p, on outer side.

These two lots seem to agree with A. 5620, but are too young for critical

comparison. ‘The punctae are near those of 7. meridionalis and less than in

T. abyssicola.

A. 5628. Off Saldanha Bay, 20 fathoms. Five juvenile examples (South

African Museum).

16 ANNALS OF THE SOUTH AFRICAN MUSEUM

The punctation ranges from 320 to 352 per sq. mm., with diameters of ?30 x

20 w (outer) and 10 X 10» (inner). One specimen has spicules in the genital

sinuses of the ventral valve near those of T. valdiviae BI.

A. 5622. 5 miles W. by S. of Constable Head (Saldanha Bay), 47 fathoms.

One juvenile example (South African Museum).

The punctation ranges from 308 to 320 per sq. mm., with diameters of 15 x

15 » (outer). ‘The striation is somewhat of the evanescent type and not that

of T. abyssicola.

The above two lots are puzzling, but are too immature for a definite con-

clusion to be reached.

Subfamily GRYPHINAE Sahni, 1929

(nom. nov., to replace Yerebratulinae Dall, 1870, which conflicts with the

genus Terebratulina).

Genus Gryphus Megerle von Mihlfeldt.

1811. Ges. nat. Fr. Berlin, Mag. V., p. 64.

Gryphus capensis sp. nov.

(Pl. I, figs. 10-13.)

Description: Shell small, longitudinally oval, with somewhat straight

sides and very slightly truncated front, widest at the middle and tapering

posteriorly. Valves regularly convex, ventral deeper than dorsal; anterior

commissure rectimarginate. Beak short, rounded and with no apparent ridges,

incurved dorsalwards and truncated by a small, circular, marginate foramen,

separated from the hinge-line by a symphytium: the foramen is also slightly

labiate and has a short, but distinct, pedicle-collar.

Interior of ventral valve smooth and with fairly strong muscle-impressions

in the umbonal region; teeth small, situated at the basal angles of the symphy-

tium: dorsal valve with cardinalia consisting of thin, concave hinge-plates

uniting the socket-ridges and crural bases; small accessory outer socket-ridges;

no median hinge-plates; cardinal process small and transverse; crural processes

some distance down the almost parallel descending branches of the loop;

transverse band narrow and arched ventrally; thin, thread-like septum separa-

ting the distinct muscle impressions in the umbonal region.

Dimensions: A. 5626 (Paratype), length 9-9 mm., breadth 6-5 mm.

A. 5627 (Holotype), length 13:5 mm., breadth 9-7 mm.

Type-specimens: As above; both in the South African Museum.

Type-localities: Holotype: 29 miles SW. of Cape St. Francis, 75 fathoms.

Paratype: 73 miles S. by W. of Cape St. Blaize, 125

fathoms.

Remarks: In his Annotated List of the Recent Brachiopoda, the late

Dr. W. H. Dall, in 1920,!* showed that the familiar generic name Liothyrina

Oehlert must give way to the earlier name of Gryphus of Megerle von Mihl-

feldt, 1811, the type species of which is the Anomia vitrea Born, 1778, a common

Mediterranean species.

two

REVISION OF SOME SOUTH AFRICAN BRACHIOPODA 17

The general characters of the new Cape species seem to place it in the

Gryphus group rather than in that of Liothyrella Thomson. As pointed out in a

previous paper,?® the Gryphus (olim Liothyrina) group comprises fairly large

shells with a somewhat truncated front, and, in some cases, broad dorsal

uniplication, together with a loop with long, almost parallel, descending

branches, with crural processes a little distance down, and a broad transverse

band, as in Gryphus vitreus, sphenoideus, cubensis, bartletti, and stearnsi: the Liothy-

rella group, on the other hand, comprises broadly dorsally uniplicate oval.

shells, usually of small size, with a rounded front, and possessing a loop of a

short triangular form, with crural processes close in, and a very narrow trans-

verse band, as in Liothyrella uva, notorcadensis, antarctica, etc. ‘The features in

these two groups are correlated with certain types of spiculation in the animal

(see Blochmann,!4” 1°),

Though Gryphus capensis is smaller than the other species assigned to the

genus, its loop bears considerable resemblance to that of G. stearnst or G. sphen-

oideus, as figured by Blochmann (1*°pl. 39, figs. 29 and 23a); the transverse

band is, however, much narrower.

A small Gryphus-like species was dredged by the ‘Challenger’ Expedition in

1873, off the Cape of Good Hope, at Station 142, lat. 35°4’S., long 18°37’E.,

at a depth of 150 fathoms. Davidson (*pp. 29-30, pl. II, figs. 5-6) referred

the species to Terebratula vitrea var. minor Philippi (now Liothyrella affinis Calcara).

The specimens are in the British Museum: they are not so elongate as G.

capensis, being slightly broader. A similar species was dredged by the ‘Valdivia’

Expedition in 1898, from the Agulhas Bank, and was described, but not named,

by Blochmann in 1906 ('#4p. 699) and in 1908 ('*°pp. 605, 613, and 630,

pl. 39, fig. 31). He compared it with the ‘Challenger’ specimen and concluded

that both were specifically distinct from L. affinis in the character of the loop

and in other features. In 1906 he referred it to the Gryphus (= Liothyrina)

series and ranged it with the larger species G. vitreus, etc.—a group without

certain spicules penetrating the bases of the cirri. In his distributional map

of 1908 (‘*°pl. 40), however, it is placed with the series containing Liothyrella

affinis, antartica, etc.

Unfortunately the ‘Valdivia’ example has not been fully figured, the loop

only having been illustrated by Blochmann (!*°pl. 39, fig. 31). Without a

further examination and a comparison of the ‘Challenger’ and ‘Valdivia’

examples with the present species it is not possible to be sure of their identity.

The habitat of the two former is some distance to the west of that of G. capensis,

but does not rule out the probability of the species being identical.

In connexion with the relationship of the South African brachiopod fauna,

mention might be made here of other occurrences of species of either Gryphus or

Liothyrella in localities not far removed from the South African region. Of

some interest is the record of Terebratula cernica Crosse,2* which was obtained

from the stomach of a fish taken at a depth of 80 fathoms off the Island of

Mauritius. It is a larger species than G. capensis but nothing is known of its

18 ANNALS OF THE SOUTH AFRICAN MUSEUM

loop or interior features. It was refigured by Davidson in 1886 ('’p. 16,

pl. I, fig. 19) as Liothyris cernica, and was later referred tentatively to the Gryphus

vitreus series by Blochmann ('*°p. 625, pl. XL). According to the latter,

Studer also mentions a fragment of a similar Terebratulid obtained at Mauri-

tius by the ‘Gazelle’ Expedition, 1874-6. Further, a Liothyrina (= Gryphus) sp.

indet. was recorded by Dall (**p. 439, pl. 26, figs. 1-2) from south of Saya

de Malha Banks, NNE. of Mauritius. In outward form it approached G.

bartlettz (West Indies) ; its loop showed affinity with G. sphenoideus (West Mediter-

ranean and East Atlantic, off Portugal, etc.); and the spiculae were closely

allied to G. vitreus (Mediterranean and East Atlantic) (!’p. 46).

Two examples of a further interesting species of Gryphus were dredged by

the ‘Challenger’ Expedition in 1876, off Ascension Island, in 420 fathoms, and

were regarded by Davidson (*p. 28, pl. II, figs. 10-11) as agreeing in every

respect with the G. cubensis (Pourtales) from 100 to 300 fathoms off the Florida

Reefs. Blochmann, in 1908 (!*°pp. 622-3, pl. 38, fig. 20) refigured one of

the ‘Challenger’ specimens, and remarked upon its somewhat asymmetric

outline. More examples are required in order to establish the exact status of

the Ascension Island form.

Family TEREBRATELLIDAE King, 1850.

Subfamily MEGATHYRINAE Dall, 1870.

Genus Megathiris d’Orbigny.

1847. C.R. Ac. Sc., Paris, xxv, p. 269.

Megathiris capensis sp. nov.

(Pl. II, figs. 3-6, 9-13, 16, 17.)

Description: Shell small, transversely ovate, with a long straight hinge-line,

almost or quite equal to the full width of the shell; rounded anteriorly; strongly

convex, the ventral valve being considerably deeper than the dorsal; surface

smooth except for rather strong growth-lines; test thick and visibly punctate;

cardinal area in each valve, that on the dorsal valve being small and visible

only at the cardinal extremities. Ventral valve very deep, with truncated

rostrum, large submesothyrid foramen, rudimentary deltidial plates separated

by linear grooves from the adjacent area which is high and triangular; interior

furnished with two short, lateral septa near the anterior border, and one long,

thin, median septum extending backwards from just within the anterior border

to the umbo where it is high-standing and supports a distinct pedicle-collar;

hinge-teeth fairly large, widely separated, and without dental plates; internal

surface covered with white, rounded, granulations like shagreen. Dorsal valve

shallow with slightly protuberant apex, furnished with three prominent septa

thickened and denticulated along their crests; the two lateral septa, which

commence within the anterior border, possess bulbous triangular apices with

points directed towards the lateral borders of the shell, they extend backwards

as low ridges to about the middle of the valve: the median septum, which has a

bulbous rounded apex, commences at the same distance from the anterior

REVISION OF SOME SOUTH AFRICAN BRACHIOPODA 19

border and extends backwards as a low ridge to the hinge platform: cardinalia

consisting of two prominent socket ridges, widely separated but united across

the umbonal cavity by two broad, excavate, hinge-plates resting on the posterior

prolongation of the median septum; no true cardinal process, the diductor

muscles being attached to a vertically striated apical portion of the interior of

the valve; crura very short, crural processes large, convergent; loop as in M.

detruncata: lophophore ptycholophous.

The shell shows no sign of folding, the anterior commissure being recti-

marginate. |

The lophophore is ptycholophous, as in Megathiris detruncata (Gmelin).

(See plate II, fig. 17.)

Dimensions: Holotype, A. 5616a, 7°71 X 5 mm.

Paratype, A. 5616b, 7°5 x 5 mm.

Type-specimens: As above, both in South African Museum.

Type-locality: Dredged in 45 fathoms, with Crania roseoradiata sp. nov.,

7 miles SSW. of Constable Hill (Saldanha Bay).

Remarks: ‘The genus has not hitherto been recorded from South Africa,

and its occurrence there is of extreme interest since it appears to be peculiarly

northern as regards both recent and fossil occurrences. A Tertiary fossil form

with some external resemblance to Megathiris has been recorded from Chile by

Dr. A. Philippi (“p. 218, pl. 49, fig. 11). The species is described as Terebra-

tula depressa and comes from Lebu, and the author suggests, from its extremely

small size, that it is possibly not fully adult. ‘It resembles’, he says, ‘the T.

detruncata of European Seas very much, which belongs to the genus Megathiris

d’Orbigny.’

D. P. Oehlert, in 1888 (*p. 814), referred T. depressa Phil., to Megathiris.

Von Ihering, in 1903 (*°p. 338), also placed it in the same genus along with

Argiope barretiana Davidson which occurs in the West Indies and Florida (43-

461 fathoms) and off Rio de Janeiro (7o fathoms). The latter is now regarded

as Argyrotheca barretiana. Von Thering (*p. 341) regarded the presence of a

species of Megathiris in the Tertiary of Chile as confirming the Atlantic

character of the Tertiary fauna of that country, the presence of this and other

forms being explained easily by the free communication of seas in Central

America, certainly in Eocene and probably in Miocene times.

Until other, and more adult, examples of the ‘Terebratula’ depressa Philippi

are forthcoming, there remains some doubt as to its generic relationship. It

may be pointed out that Philippi’s name is pre-occupied by the Terebratula

depressa Faujas 1799, a Cretaceous (Senonian) species from France and

Holland.*” This was referred to Megathiris by d’Orbigny in 1847 (*p. 149)

and to Cistella by de Morgan in 1883 (*p. 10 of reprint). Cistella is now

Arg yrotheca Dall.

The Cape species is very distinct from the genotype Megathiris detruncata

(Gmelin) (pl. II, figs. 1, 2, 7, 8, 14, 15), which occurs in the Mediterranean and

East Atlantic from Guernsey and the Scilly Is. to Madeira (16-100 fathoms),

20 ANNALS OF THE SOUTH AFRICAN MUSEUM

and is recorded as a fossil from the Eocene, Miocene and Pliocene of south and

eastern Europe.°? Megathiris capensis is larger and more transverse, in addition

to being entirely smooth externally and more rugose internally.

Jeffreys (*'p. 124, pl. 5, figs. ga-c) records a form of M. detruncata from

Guernsey which has weaker costation: it resembles a horse’s hoof in shape,

being longitudinally oval, instead of transversely oblong (as in Mediterranean

examples); the costae are much fainter and do not extend to the anterior

margin. He also refers to specimens in the M’Andrew collection from

Madeira, which, though smaller, have the same form and sculpture. These,

he considers, may therefore belong to a distinct and undescribed species.

Sacco, in 1902 (“pp. 30-1), figured and briefly described several varieties

of M. detruncata from the Italian Tertiary rocks. ‘These include a var. semi-

laevis (>*pl. 6, figs. 17-19) in which the costation does not reach the margin

which is smooth: it occurs frequently in the Middle Miocene and Lower

Pliocene, and suggests Jeffreys’s form previously mentioned and an Italian

Eocene species, referred to M. detruncata by Davidson (*pl. XXI, figs. 6-8).

Another of Sacco’s varieties is var. perlaevis (>pl. 6, fig. 20), in which the

valves are smooth or nearly so: it occurs somewhat rarely in the Middle

Miocene and Lower Pliocene. Sacco’s figure of this variety is an external

view of the dorsal valve: it strongly suggests the Cape species, but in the absence

of an interior view of this fossil a comparison of the cardinalia, etc., with those

of the present species cannot be made.

Subfamily KRAUSSININAE Dall, 1870.

The subfamily position of the genus Krausszna and some allied genera has

been the subject of much difference of opinion in the past. Dall in 1870

(?p. 138) created the subfamily Kraussininae for Kraussina rubra and four other

species including lamarckiana which later (in 1884) became the genotype of

Megerlina. The same subfamily was adopted by Davidson in 1887 ('p. 118),

and by Schuchert in March 1893 (*p. 160). But Beecher in 1893 (*’p. 391)

placed Kraussina and Megerlina in his new subfamily Magellaniunae along

with Magellama, Terebratella, and others, and at the same time created a new

subfamily Dallininae to embrace Dallina, Macandrevia, Terebratalia, Laqueus,

Miihlfeldiia, etc. At that time the genus Muiihlfeldtia included truncata and

what is now known as Frenulina sanguinolenta. ‘The latter species, under the

name M. sanguinea, was used by Beecher in illustration of one of the stages in

the ontogeny and morphology of the Dallininae, the stage being called the

‘Muhlfeldtiform’ stage. In studies made in 1916 (°p. 24) I found that

Miihlfeldiia truncata differed fundamentally from Frenulina sanguinolenta and I

transferred the former genus to the Magellaniinae and changed Beecher’s stage

name in the Dallininae from ‘Miuhlfeldtiform’ to ‘Frenuliniform’. This is now

generally accepted. My reasons for the transfer were based upon the absence

of dental plates in Miahlfeldtia truncata, as well as in the resemblance of the early

loops stage to a similar stage of Terebratella dorsata, and the appearance of the

REVISION OF SOME SOUTH AFRICAN BRACHIOPODA 2

secondary loop before the appearance of the primary lamellae. It was pointed

out for the first time in that paper that in the Dallininae the descending branches

of the loop (i.e. the primary loop) were united to the septum at an earlier stage

than in the Magellaniinae. In the same paper I also emphasized a hitherto

unrecognized feature in the Dallininae of the universal presence (except in

adults of the genus Dallina) of dental plates in the ventral valve.

Though possessing close affinities with the Magellaninae, I was not fully

satisfied in 1916 that Muhlfeldtia rightly belonged to that subfamily or to a new

one altogether. The matter was taken up later in the same year by J. A. Thom-

son (*!p. 498) and again in 1918 (p. 7). In these valuable papers he

foreshadowed a separate subfamily to include Araussina, Megerlina and Miihl-

feldtia, and in his later Manual (78p. 219) placed these three genera together

with Pantellaria and Aldingia in the emended but almost forgotten subfamily

Miihlfeldtiinae Oehlert 1887 (p. 1314). It is unfortunate that Oehlert in

proposing this subfamily was not more precise in specifying the genera he

proposed to assign to it, but one can assume, from the arrangement of the

genera on later pages (pp. 1322-3), that it comprised Muihlfeldtia (M.

truncata) [with Section Megerlina (M. lamarckiana) and Ismenia (I. pectunculus)],

Kraussina (K. rubra), and Platidia (P. anomiozdes).

The close relationship of Mihlfeldtiia, Megerlina and Kraussina as regards loop-

development and spiculation was clearly demonstrated by Deslongchamps in

1884 (778p. 122).

Davidson in 1887 ('’p. 118), placed Kraussina and Megerlina in the sub-

family Kraussininae, and Dall in 1920 (!*pp. 374-6), followed the same pro-

cedure, at the same time placing Muihlfeldtia and Pantellaria in the subfamily

Mihlfeldtiunae.

Though Kraussina, Megerlina and Miihlfeldtia are so closely related in loop

characters, there are certain other minor points in which they differ. Kraussina

does not possess a pedicle-collar in large adult shells: in these there is only a

narrow thickened band fused to the floor of the umbonal cavity (as in Copto-

thyris, Terebratalia, and other higher long-looped forms). In very young shells

about 4 mm. and 6 mm. in size the band is slightly free anteriorly, which

suggests that in still earlier stages there may be a free pedicle-collar. Megerlina

and Miihlfeldtia possess a free or true pedicle-collar up to the adult stage

(see 2°p. 25). Pending further detailed study of the animals I feel disposed at

present to place Kraussina, Megerlina and Miuihlfeldtia in Dall’s subfamily

Kraussininae which antedates Mihfeldtunae Oehlert by several years.

Genus Kraussina Davidson.

1859. SB. Ak. Wien. Math.-naturw., pl. xxxvui, p. 189, 210 (pro Kraussia

preocc.).

From a careful study of numerous specimens and of the literature upon the

subject, the only species which I consider as coming within the genus Kraussina

22 ANNALS OF THE SOUTH AFRICAN MUSEUM

proper are A. rubra (Pallas), K. cognata (Sowerby), K. gardineri Dall, and K.

crassicostata sp. nov., described in a later page.

The area of distribution of this genus is of peculiar interest. So far as is

known at present it ranges from the Cape of Good Hope to the Indian Ocean

south of the Saya de Malha Banks. The species restricted to the Cape itself

are three in number, viz. A. rubra, cognata and crassicostata. ‘The Indian Ocean

form is K. gardinerr. tis seen from the above that the distribution of the genus

is somewhat discontinuous, there being no record of its occurrence between

Port Alfred and the Saya de Malha Banks, except a reference by Davidson, in

1887 (1%p. 120), to some small specimens erroneously described by Gray in

1872 under the mistaken name of Terebratula truncata (= Miihlfeldtia truncata).

They were found attached to Ascidia, and to the stems of large algae, off the

coast of Natal. ‘These specimens are referred to Kraussina rubra by Davidson.

The occurrence of K. rubra here appears to me to be extremely doubtful in

view of the direction taken by the Mozambique Current, viz. towards the

Cape. Gray may have been correct in his attribution of the species to Miihl-

feldtia truncata. In 1921 (*p. 49) I referred to the presence in the British

Museum (Natural History) of an immature example of this typical Lusitanian

species, labelled ‘S. Africa. J. H. Ponsonby coll. 1900. 6.13.4’.

Unfortunately full details as to habitat and depth have not always been

recorded, hence it is impossible at present to give the bathymetric range of

Kraussina, but, from the available data and from the thick-shelled character of

the Cape species, the genus appears to be one of fairly shallow water.

It is also to be regretted that in most of the literature dealing with K. rubra

one finds no more precise locality than ‘Cape of Good Hope’. More data on

these points are badly needed.

The genotype of Araussina is the Anomia rubra Pallas 1766. ‘The original

generic title proposed by Davidson in 1852 (®°p. 369) for the reception of this

species was Araussia, but this name having been used by Dana for Crustacea

earlier in the same year, Kraussina was suggested by Suess in 1859 (*%p. 210)

in collaboration with Davidson, who confirmed it in 1861 (*4p. 39).

Davidson, in 1887 (}*p. 118), included seven species in the genus, five in

Kraussina proper (viz. rubra, cognata, deshayesi, pisum and atkinsom), and two in

the subgenus Megerlina (viz. lamarckiana and davidson). Only two of these are

strictly referable to Kraussina, viz. rubra and cognata. ‘The generic position of

the remaining five forms will be dealt with in the sequel.

Kraussina rubra (Pallas)

(Pl UT: figs. m7)

Anomia rubra Pallas, Misc. Zool., 1766, p. 182, pl. XIV, figs. 2-11.

Terebratula capensis Kuster (not Adams and Reeve), ‘Mart. and Chem.’, Conch.

Cab., 1848, p. 32, pl. 3, figs. 15-17.

Terebratula capensis Krauss (not Adams and Reeve), Siidafr. Moll., 1848, p. 32,

pl. Il, fig. 10.

REVISION OF SOME SOUTH AFRICAN BRACHIOPODA 23

Terebratula (Kraussia) rubra Reeve, ‘Mon. Terebratula’, Conch. Icon., 1861,

pl. IX, fig. 37.

Kraussina rubra Davidson, ‘Mon. Rec. Brach.’, Trans. Linn. Soc., ser. 2, vol. IV,

Zool. pt. II, 1887, pp. 119-21, pl. XX, figs. 19-23.

Remarks: This well-known species has been met with at several localities

around the Cape, but little is known as to its range in depth. It has been seen

by the writer from Port Elizabeth, Algoa Bay and Port Alfred, and from the

Agulhas Bank, 22 fathoms. A figure of a specimen from the last locality,

agreeing closely with Pallas’s original, is given here (pl. III, fig. 1) for com-

parison with the new species described in a later page.

Kraussina rubra has been described and figured by several authorities and

appears to present a certain amount of variation in outward form. ‘The Anomia

rubra, as figured by Pallas, is a transversely oval and costate shell, without

apparent fold or sinus; but many specimens of equal size which I have examined

show a strong ventral fold and dorsal sulcus: these features are well marked

from the umbonal region. In these the shells are ventrally uniplicate or

sulcate. Some forms tend to become more elongate than others. ‘There is

also some slight variation in the intensity of the costae.

It is not my intention to burden this paper with a full revised description of

Kraussina rubra, but some important details are necessary in order to emphasize

certain differences between this genus and that of Megerlina dealt with in a

later page.

In the interior of the dorsal valve of K. rubra the cardinalia consist of two

divergent socket-ridges bounding the dental-sockets. Lying between these

ridges is an umbonal callosity with two eye-like depressions on its surface.

There are no excavate hinge-plates, even in shells as small as 4 mm. in size.

At the apex a small cardinal process is present. A grooved median septum rises

from the umbonal callosity and extends some distance down the interior of the

valve. From its anterior extremity arise two short, deviating lamellae exten-

ding towards the ventral valve: these fork at their ends, giving rise to anterior

and posterior processes. ‘These processes are usually quite short, but in some

specimens the posterior processes are in the form of long narrow ribbons

extending backwards and slightly inwards towards each other: they possess

hook-like extremities. The loop is essentially adult in shells 6 mm. long.

The dorsal valve also possesses a slight, but distinct, cardinal area, a feature

of unusual occurrence, found also in the genera Miihlfeldtia and Megerlina

(see 1”). There are a few spines within the margin.

In the ventral valve the features of importance are the absence of dental-

plates and of a free pedicle-collar. As previously stated, in adult shells, there

is a thickened band in place of the latter, but in very small specimens this band

may be free anteriorly.

The dense character of the shells of this species renders it somewhat difficult

to make a study of the punctation. I have, however, succeeded in one or two

cases with interesting results. Adult shells have large oval pores on the

24 ANNALS OF THE SOUTH AFRICAN MUSEUM

exterior, these pores being denser and larger in the grooves than in the costae.

In specimens from Port Elizabeth I have counted as many as 256 pores per

sq. mm. in the grooves, as against 168 pores in the costae. In the former they

measure 60 X 30-35 » , and in the latter, 45-50 x 15-20 wu. On the inner

surface the dimensions are 15 X 15 p for all pores. The young specimens,

however, exhibit some interesting differences both as regards punctation and

interior details. ‘These are perhaps worth recording. In a specimen 6 mm.

long, on which fine costation is visible, the punctae appear to be evenly distri-

buted with no differentiation in the grooves or on the costae: they average 244

per sq. mm. at the middle of the ventral valve. Internally this valve has a

narrow band within the apex which is slightly free anteriorly. The dorsal

valve is like the adult, without excavate hinge-plates; and has a loop consisting

of two very broad divergent lamellae, slightly bifurcated at their extremities.

Widely-spaced spinules occur just within the margin of the valve. In a

smaller specimen 4 mm. long, the punctae are moderately round with no

differentiation, being 188 per sq. mm. _ Internally it is much the same as the

other, but the extremities of the divergent lamellae are without bifurcations.

The spiculation of the mantle of Kraussina rubra was studied by Deslong-

champs in 1864 (*p. 25, pl. II, figs. 10-12), and again in 1884 (3*°pp. 121

et seq. and 160, pl. XIX, fig. 7). The spicules are of a very special form and

very small. ‘Two systems exist, one in the mantle, the other in the pallial

sinuses. Spicules also exist in the walls of the visceral chamber and in the

arms.

Kraussina cognata (Sowerby).

Terebratula cognata Sowerby, Thes. Conch., vol. 1, 1847, p. 346, pl. 68, figs. 12-14.

Terebratula cognata Kiister, in ‘Mart. and Chemn.’, Conch. Cab., Bd. VII, I,

1848, p. 46, pl. 4, figs. 5-6 (as figs. 3-4 in text).

Terebratula (Kraussia) cognata Reeve, ‘Monogr. Tereb.’, Conch. Icon., 1861,

pl. IX, figs. 38a, b.

Kraussina cognata Davidson, ‘Mon. Rec. Brach.’, Trans. Linn. Soc., ser. 2, vol.

IV, Zool. pt. II, 1887, pp. 121-2, pl. XX, figs. 24-6 (var.? 27-30).

Remarks: According to figures and descriptions this species is somewhat

subtrapezoidal or elongated, with a nearly straight hinge-line almost as long

as the width of the shell. The colour is pale yellow and the surface covered

with radiating costae. Chemnitz, in 1785 (#p. 78, pl. 76, figs. 688a, b)

described and figured a shell from the Cape as ‘Cognata Anomiae craniolaris basi

perforata’, but from his poor description and illustrations and from the fact of

his not being a binomial writer, the Terebratula cognata of Sowerby is regarded

as the type.

Davidson gives copies of the figures of Sowerby and Reeve which show the

type of folding as sulcate.

Unlike Kraussina rubra, this species increases in length but retains the mega-

thyrid type of cardinal margin.

REVISION OF SOME SOUTH AFRICAN BRACHIOPODA 25

This species requires more careful study and more material. I have only

been able to examine one or two indifferent specimens which appear to agree

with the description of the species. In these the cardinalia, etc., agree essen-

tially with K. rubra, and the interior of the dorsal valve is very spinous just

within the margin. Spicules are present in the ventral and dorsal mantles.

Owing to the almost smooth condition of the surface of the valves, there is no

differentiation in the punctation. ‘The pores about the middle of the ventral

valve are small, even, and widely-spaced, and number 160-176 per sq. mm.

They are 15 X. 15 p» in size, but a few reach 20 X 20 wp.

Habitat: South Africa, near the Cape of Good Hope.

Note. Dr. K. H. Barnard has recently submitted a large example of a

Kraussina containing the dried animal dredged in 10-14 fathoms at Saldanha

Bay by the Preter Faure. Its registered number in the South African Museum

is A. 5607. It is said to have been identified as Araussina cognata by G. B.

Sowerby and may be this species. The dimensions are: L. 32:2, W. 29:5,

D. 18-3 mm. The earlier growth-lines suggest a shell wider than long, as in

Kraussina rubra. ‘The shell is hardly sulcate and of a pale yellow colour, and

possesses moderately strong radiating costae.

The occurrence of the genus at Saldanha Bay is of extreme interest: it

probably reached the South Atlantic coast by the’aid of the Benguela Current.

Two other interesting brachiopods are recorded in this memoir from this

neighbourhood, viz. Crania roseoradiata sp. nov. and Megathiris capensis sp. nov.

Kraussina gardinert Dall

Kraussina gardinert Dall, “Brach. of the Sea Lark Exped.’, Trans. Linn. Soc. Lond.,

ser. 2, Zool. vol. 13, 1910, p. 440, pl. 26, figs. 3-6.

Description: Shell rude and solid, greyish white, similar in shape to Muhl-

Jfeldtia truncata when young with shell 12 x 13 mm. in size. During growth,

the outline elongates proportionally more that it widens, and an adult measures

24°5 mm. in length by 23 mm. in width at the broadest part. Outline then

roughly rhombic. Dorsal valve slightly sulcate; ventral valve with corre-

sponding convexity. Latter valve slightly more convex than dorsal: total

thickness of adult about halfits length. Beak short and wide; peduncle short;

foramen wide and incomplete, margins more or less eroded or defective; wide

flattened area of irregular shape on each side of foramen.

Surface of valves with coarse, rounded costae, strongest mesially, with sub-

equal roundly excavated interspaces: costae mostly continuous from beaks,

and increasing rather by bifurcation than by intercalation: about forty costae

in adult, the laterals being finer and closer-set than the others.

Interior of valves strongly calcified; spinules within margin.

Habitat: Station C1, Indian Ocean, south of the Saya de Malha Banks, in

123-153 fathoms (dredged by J. Stanley Gardiner, after whom it is named).

26 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks: Judging from Dall’s figures, the cardinalia and brachidium of

this species are essentially of the Araussina rubra type, but there appears to be

more excessive calcification in the umbonal cavity.

The occurrence of Araussina in the Indian Ocean is of great interest. Asso-

ciated with the species was a curious intraplicate Rhynchonellid (Hemithyris

sladem Dall) which is tentatively referred to the Early Tertiary New Zealand

genus Aetheia by Thomson (8p. 157); and a Liothyrina sp. indet., having some

affinities with Mediterranean forms (see also Jackson,’ p. 46).

Kraussina crassicostata sp. nov.

(Pl. III, figs. 3-5.)

Description: Shell solid, subcircular, about as broad as long, with a broad,

straight hinge-line somewhat shorter than the greatest width of the shell

(= submegathyrid). Valves convex, the ventral much more so than the

dorsal; sulcate (i.e. with a shallow sulcus in the dorsal valve opposed to a fold

in the ventral valve); margins waved by alternate multicostation. Cardinal

area in both valves. Surface covered with 10-12 coarse, angular, radial

ridges, strongest mesially and becoming obsolete near the cardinal angles,

separated by angular interspaces: ridges continuous from beak, broadening

anteriorly, and increasing by bifurcation: they are crossed by fairly strong

growth lines. Colour of surface reddish; punctate. Beak well-truncated,

foramen very large (due to wear), and incomplete anteriorly,* slight deltidial

plates, narrow sessile pedicle collar (i.e. thickened band) within the umbonal

cavity. Interior of ventral valve fairly smooth, with distinct muscle impressions

posteriorly; bluntly ridged round the margin (= reverse of external ornament) ;

row of small spines just within the margin; teeth small, without dental plates,

situated at anterior angles of inner margins of the two widely-separated portions

of the transversely-grooved area. Interior of dorsal valve smooth, with well-

defined muscle impressions, bluntly ridged margin, and row of spinules, as in

ventral valve: cardinalia and brachidium essentially of the Kraussina rubra type:

no hinge-plates proper, their place being occupied by two eye-shaped pits (for

insertion of the pedicle-muscles) in a thick callus in the umbonal cavity between

the distant and thick socket-ridges. Cardinal process small and prominent,

united laterally to the posterior bases of the socket-ridges by narrow, vertically

striated, muscular impressions, within the posterior margin of the umbonal

cavity; rudimentary outer socket-ridges bounding the widely separated portions

of the area of this valve. Mesial septum distinct, extending from the umbonal

callus (where it appears to be grooved on its upper surface) to about the centre

of the valve, where it supports two, short and narrow, divergent and ventrally

directed lamellae.+

Dimensions: A. 5615 (Holotype), length 16:2 mm., breadth 16-2 mm.,

thickness 11:3 mm.

* Owing to wear it is not possible to define the type of foramen.

+ In the holotype the extremities are broken, but in another example the extremities are

flattened and forked (as in K. rubra).

REVISION OF SOME SOUTH AFRICAN BRACHIOPODA 237)

Type-specimens: Holotype and two metatypes, No. A. 5615, in the South

African Museum, Cape Town.

Type-locality: False Bay, 23 fathoms.

Remarks: This species differs from the genotype, Kraussina rubra, in posses-

sing much coarser and less numerous costae which tend to become obsolete

near the cardinal angles; the ventral valve is also deeper. Avaussina cognata

(Sow.) is a longer species with a flattened dorsal valve, and of a pale yellow

colour: Kraussina gardineri Dall is greyish in colour and possesses a more

calcified cardinalia, etc.

The external surface of the ventral valve has large ovate punctae in the

grooves with diameters of 45 < 30 », and about 256 per sq. mm., while the

costae have smaller ovate punctae of less density: internally the punctae are

very small and round, about 10 X I0 wp.

Spicules are present in fragments of the ventral mantle adhering to the

interior of the shell, especially in the sinuses.

In some specimens in the Manchester Museum of apparently the same

species and dredged with Allopora nobilis Kent near the Cape of Good Hope,

in 26 fathoms, the punctae are similarly diversified, but are somewhat larger

and rather less in relative density than in A. 5615.

Genus Megerlina Deslongchamps.

1859. Ktud. crit. Brachiop., p. 159.

1884. Bull. Soc. Linn. Normand., viii, pp. 210, 242.

Having dealt with the species appertaining to the genus Kvaussina proper,

attention may be directed to other forms which in the past have been errone-

ously referred to that genus. Three related South African species are included

in this group which may be termed the ‘Terebratula pisum series’.

Davidson, in his description of the ‘Challenger’ Brachiopoda (°p. 54),

committed a most remarkable error in attributing to the Terebratula pisum

Lamarck an altogether different species. I have examined the Cape specimens

figured and described by Davidson as Kraussina pisum (Lam.), in the ‘Challenger’

Report (°pl. IV, figs. 7, 7a, 7b and 8) and later in his “Monograph of Recent

Brachiopoda’ ('%p. 123, pl. XXI, figs. 1-4). The figure 7a of pl. IV of the

former report is reversed in printing from the stone (as are some others in the

same work): that in the latter report (pl. XXI, fig. 2) is not reversed. The

latter, and adjacent figures, however, are not so correct in detail as those in the

earlier work, as they do not show the strong ventral plication of the species in

question. Davidson’s specimens are preserved in the British Museum: figs. 7,

ya and 7b, are the same shell drawn in different views and repeated in ‘Recent

Brachiopoda’, pl. X XI, fig. 2; this shell is in the Zoological Department of the

above Institution and is registered No. 78.6.15.27. Figures 1, 1a, 1b, 3 and ga

of pl. X XI are| drawn from other specimens now preserved in the Geological

Department of the same Institution (Reg. No. B. 12405). Figure 8 (interior

28 ANNALS OF THE SOUTH AFRICAN MUSEUM

view) of pl. IV (‘Challenger’ Report) is repeated as fig. 4 of pl. X XI, in the

Monograph. !

The above species is certainly not the Terebratula pisum of Lamarck, 18109,

as, according to the original description (*pp. 245-6), this is a small, smooth,

subglobose red shell, resembling a cherry stone, and 9 mm. wide. The habitat

given is ‘Isle de France’ (= Mauritius). Davidson’s ‘Kraussina pisum’ bears

costae, somewhat like Megerlina lamarckiana (Dav.), but finer, and presents no

resemblance to a cherry stone. It is also of a uniform light yellow colour,

with no red markings at all. This species does not possess the Kraussina-type of

cardinalia and brachial support, but shows a greater resemblance to Megerlina

in these features. What I regard as the true Terebratula pisum Lamk. possesses

essentially the same type of cardinalia and brachial support and must be placed

in the same genus. A new specific name, therefore, is required for Davidson’s

shell. One other species, viz. T. capensis Ad. and Rve. = deshayest Dav., also

comes into the same genus, viz. Megerlina.

Megerlina striata sp. nov.

(Pl. 111; figs. 6; 9.)

Kraussina pisum (non Lamk.) Davidson, Voy. Challenger, Zool. vol. I, 1880,

p- 54, pl. IV, figs. 7-8. ‘Mon. Rec. Brach.’, Trans Linn. Soc., ser. 2, vol. IV,

Zool. pt. II, 1887, p. 123, pl. XXI, figs. 1-4.

Description: Shell small, transversely oval (except in young); colour

yellowish-white. Dorsal valve slightly convex, with a distinct central longi-

tudinal depression extending from a prominent umbonal swelling to the

anterior margin and increasing in width rather rapidly. Hinge line nearly

straight, about two-thirds the width of the valve. Ventral valve deeper than

dorsal, longitudinally keeled. Folding sulcate. Beak slightly incurved, with

a rather large incomplete foramen, bounded laterally by two small deltidial

plates which curve upwards towards the dorsal umbo but fail to meet; beak-

ridges very distinct, leaving a small flattened cardinal area between them and

the deltidial plates. Foramen submesothyrid. Surface of valves covered with

numerous distinct radii which increase by bifurcation and interpolation:

cardinal angles smooth (i.e. radii absent on these portions) ; concentric growth-

lines present at variable intervals. Interior of ventral valve with traces of

radii like the exterior but reversed; spinous within margin. Pedicle-collar

sessile, closely appressed to apex of umbonal cavity but slightly free in front

especially near bases of teeth. The latter are comparatively small and not

supported by dental plates. Interior of dorsal valve covered with rows of

pustules radiating from apex; spinous within margin. Rudimentary area

present in this valve.

The cardinalia (as seen in immature shell) consist of two rather stout diver-

gent socket-ridges, excavate below, and supported on inner sides anteriorly by

two spurs which extend to the median septum and enclose two eye-shaped

muscular pits. Cardinal process feeble. Median septum extending from near

REVISION OF SOME SOUTH AFRICAN BRACHIOPODA 29

apex to centre of valve where it gives off two broad, thin diverging lamellae,

the upper extremities of which are slightly forked, the posterior forks curving

inwards to some extent. Midway down each lamella, on the exterior or

dorsal face, an accessory process, or ledge, is present, analogous to that in

Megerlina lamarckiana, but less developed.

Shell with large subcircular punctae.

Dimensions : Length Breadth Thickness

Holotype 1. (pl. III, fig. 6) I4°1 16-2 5°9 mm.

Paratype 2. 13°5 14°9 Boles.

Ny 3. (interior details) 9°3 8:5 Ce

NaeweiNo. 1 — pl. IV, fig. 7 of ‘Challenger’ Report, and pl. XXI1, fig. 2 of

Recent Brachiopoda.

Habitat: Station 142, lat. 35°4’S., long. 18°37’E., off Gape of Good Hope,

150 fathoms (‘Challenger’ Expedition). Type in the British Museum (Nat.

Hist.) Registered No. 78.6.15.27.

Remarks: ‘The above description is based upon three examples in the

Zoological Department of the British Museum, the largest of which was figured

by Davidson, as stated above. Owing to the risk of damage in opening the

two large examples, the details of the cardinalia and brachidium are taken

from the smallest specimen (N. 3), but these have been checked to some extent

by No. 2 which was partly opened. ‘There is reason to believe that No. 1

would exhibit a slightly more adult condition of the loop, etc.

Two other examples of this species were also figured by Davidson (**pl. 21,

figs. 1 and 3) and are now in the Geological Department of the above Institu-

tion. These have also been examined externally and found to agree with those

above.

The shells of this species have much larger punctae than those of M. pisum

and M. capensis. The pores are subcircular in outline and range from 208 to 240

per sq. mm. about the middle of the ventral valve. The size of the pores in

No. 1 is 45-50 X 45-50 » (outer surface) (pl. III, fig. 9). They appear to be

evenly distributed over the radii and grooves.

The South African Museum possesses the following specimens which seem

to belong here:

A. 6410. 1 specimen (broken), dredged in 40 fathoms, St. Sebastian Bay,

Agulhas Bank (K. H. Barnard, 1922).

A. 5663. 5 specimens dredged off East London in 32 fathoms, by the

Preter Faure.

A. 7690. 2 specimens (locality and depth unknown). Preter Faure coll.

A. 5605. 1 specimen (broken) dredged off Cape St. Blaize (depth unknown).

Pieter Faure coll.

The shells are mainly transversely oval, sulcate, and of a light yellowish

colour, except one shell of A. 7690 which is white, and possess rather strong

radii. In A. 5605 the radii are evanescent.

30 ANNALS OF THE SOUTH AFRICAN MUSEUM

The four larger shells of A. 5663 measure: 10 X 10°7, 9°9 X 10, 10 X 9°83

and 8-5 x 8-7 mm.

Megerlina pisum (Lamarck)

(PIM fos) 7, 10.)

Terebratula pisum Lamarck, Animaux sans Vert., vol. VI, 1819, p. 245. (Text

written by Valenciennes owing to Lamarck’s blindness.)

Terebratula natalensis Krauss in Kiister, Conch. Cab., von Mart. u. Chem., Bd.

VII, I, 1844 and 1848, p. 36, pl. 2b, figs. 4-7. (Plate published, 1844,

text, 1848.)

Terebratula pisum Lamk. Sowerby, Thes. Conch., I, 1846, p. 345, pl. 69, figs.

37-9-

Terebratula algoensis Sowerby, Thes. Conch., 1, 1847, p. 362, pl. 71, figs. g1-2.

Terebratula natalensis Krauss, Siidafrikanischen Mollusken, 1848, p. 33, pl. 2, figs.

Ila-c.

? Terebratula (Kraussia) pisum Lamk. Reeve, Conch. Icon., vol. XIII, 1861, pl. 9,

fig. 36a (non 36b). 7

Kraussina atkinsont (non ‘T. Woods), E. A. Smith, Journ. Conch., vol. X, 1901,

p. 116.

Description: Lamarck’s description (*pp. 245-6) is as follows:

‘Térébratule pois. Terebratula pisum.

T. testa minuta, subglobosa, laevi, subantiquata, rubella margine integro

antice valde sinuato.

Habite a ’Ue-de-France. Par. M. Mathieu. Mus. no. Petite coquille semblable

a un noyau de cerise, ne le surpassant pas en grosseur. Elle a 9 millimétres de

largeur.

The following is an amplified description, based upon South African

examples.

Shell small, transversely oval, with a subterebratulid type of cardinal mar-

gin; valves about equal in depth; dorsal valve sulcate, ventral broadly carinate;

type of folding sulcate. Surface nearly smooth, with faint traces of costation;

test finely punctate. Colour milk-white or slightly suffused with red. Beak

suberect; foramen large, submesothyrid, incomplete; deltidial plates discrete,

small, trigonal. Pedicle-collar sessile, slightly free in front. Hinge-teeth

without dental plates. In the dorsal valve the cardinalia consist of two promi-

nent, divergent, socket-ridges curling over the dental sockets posteriorly, and

supported at their anterior corners by rudimentary excavate hinge-plates in

the form of two spurs extending inwards towards the centre line of the valve:

these enclose an imperfect hinge-trough; cardinal process small. The brachi-

dium rises from the floor of the valve near the middle as two ventrally directed

deviating lamellae which bifurcate slightly at their extremities: outer sides of

the lamellae with two short accessory processes, representing the anterior

portions of the descending branches: the lamellae extend backwards as low

convergent plates forming a pseudo-septum as far as the imperfect hinge-

trough. .

REVISION OF SOME SOUTH AFRICAN BRACHIOPODA 31

Dimensions: Length Breadth Thickness

Figured specimen (pl. III, fig. 7) 10°3 11 5°6 mm.

Others. South Africa 10-7 10°5 Baha sae

Umkomaas Q°4. 10°2 ee

» 9°3 LOFT 4°4 »

8:6 9°3 4°3

Habitat: Umkomaas, Natal; Port Elizabeth, Algoa Bay; Durban, Natal;

East London, etc.

Remarks: ‘Though originally described from Mauritius, there are several

specimens from such places as Port Elizabeth, Durban, etc., both in the

British Museum (Nat. Hist.) and in private hands, which conform closely with

Lamarck’s description. As Lamarck’s type was never figured, it seems

desirable that illustrations should be given here of the South African specimens

upon which my conclusions as to identity are based. Among the specimens

preserved in the Zoological Department of the British Museum are seven from

Umkomaas, Natal (J. H. Ponsonby coll. 1901.9.23.61-7), unfortunately without

precise details as to depth, etc. ‘I'wo other and larger specimens together with

several juveniles are also in the same Institution, and are labelled simply

‘S. Africa. J. H. Ponsonby. 99.4.14.3765-3771'. These two sets, as well as

numerous others in private collections, including specimens in my own

collection from Durban, Natal, and Port Elizabeth, Algoa Bay (both ex. H.

McClelland, 1922), have been used in the study of the species. A specimen

from the British Museum set marked ‘S. Africa’ has been selected to illustrate

the exterior features (pl. III, fig. 7) and both this and others have been used

for the interior details.

The punctae in the test of this species differ from those of M. striata in being

more oval and on the whole less numerous (pl. III, fig. 10). Judging from the

examination of about 15 specimens the number ranges 120-220 per sq. mm.

at the middle of the ventral valve. ‘The most prevalent numbers, however,

are 160-190. The dimensions of the punctae are a little variable according to

the presence or absence of strong growth-lines: they average 40-45 25-30 p

on outer surface, and 15-20 X 15-20 w on inner surface.

Large spicules occur in the pallial sinuses, but are scanty in the cirri.

Victor Sganzin (*%p. 12) in his reference to the Terebratula pisum Lamk.

says it is very small and extremely rare: it is found at a great depth in Tombeau

Bay, Mauritius, and dredging is necessary in order to obtain it.

The Terebratula natalensis of Krauss (®p. 33, pl. 2, figs. 11a-c) appear to

belong to the species under review. His figure (fig. 11b) of the hinge-processes

and brachial-support is remarkably accurate except that he appears to have

overlooked the accessory ledges on the outer sides of the deviating lamellae.

In his description he speaks of the shells being generally white in colour, more

rarely flushed with red, and notes the sulcate character as well as the finely

striated or almost entirely smooth condition of the surface of the valves. He

records the species as living in great numbers at Natal Point (= Durban) in a

depth of some fathoms, on stones, Cardita variegata, Arca kraussi, etc.

32 ANNALS OF THE SOUTH AFRICAN MUSEUM

With regard to the species described and figured by Kiister (**p. 36, pl. 2b,

figs. 4-7) as the 7. natalensis of Krauss, there is a little uncertainty as to absolute

identity. His figures seem to suggest that he may have had before him an

example of the Terebratula capensis Ad. and Rve. (= Kraussina deshayesi Dav.).

The costation shown in the figures is stronger than in M. pisum (Lamk.).

Sowerby (*°pp. 345-6), in his remarks on the species, says, ‘also found at

Sydney by Mr. Jukes’. This is undoubtedly an error: the Sydney citation

probably refers to the Megerlina lamarckiana (Dav.), which was unknown at that

time.

The Terebratula algoensis G. B. Sow., was described and figured in 1847

(7p. 362, pl. 71, figs. g1-2) from a single bleached ventral valve in the

British Museum (N. H.), Zoological Department, labelled ‘Algoa Bay; J. S.

Bowerbank’. I have examined the specimen, which bears a registration

number (rather indistinct), viz. 42.12.19.26, on the interior of the valve, and

consider it to be a somewhat irregularly-grown valve of M. pisum (Lamk.).

The number of punctae per sq. mm. is about 160, and the dimensions are;

outer, 45-50 X 25-30 w; inner, 15-20 X 15-50 p. |

In the Journal of Conchology for 1901, E. A. Smith (*%p. 116) records

Kraussina atkinson (T. Woods)—a Tasmanian species—for Algoa Bay, Cape

Colony (Brit. Mus., J. H. Ponsonby). I have examined the specimens in

question—four in mumber B.M. 1900.6.13.5-8)—and find them to be un-

doubted juveniles of M. pisum (Lamk.). The number of punctae per sq. mm.

ranges from 160 to 182, and the size of the pores on the outer surface is 30-40 X

25-30 p. In the ‘Kraussina atkinson (T. Woods), judging from three specimens

in my own collection from Long Bay, S. Tasmania (the type locality), the pore-

density per sq. mm. is much greater, being 255-264; and the pores are practi-

cally circular and measure: outer, 30-35 X 30-35 »; Inner, 25-30 X 25-30 u.

They are distinctly visible under a lens. I do not consider atkinsom a true

Kraussina as it differs in shell-characters, and in its cardinalia and brachidium.

Davidson’s figures (%pl. 21, figs. 5-6) are, unfortunately, not quite accurate.

The shells in my possession show the sulcate type of folding and two of them

have rather indistinct costae on their outer surface. ‘The species seems to be

passing from a costate to a smooth stage. The ventral valve has a submeso-

thyrid beak, a fairly large foramen, incomplete, and bordered anteriorly by

small triangular deltidial plates; no dental plates; pedicle-collar deep,

sessile, vertically striated, and slightly free anteriorly. In the dorsal valve the

cardinalia consists of two upstanding socket-ridges on the inner sides of which

are slight plates or buttresses which descend to the floor of the valve without

meeting in the median line. These descending buttresses extend forward and

converge about half-way down the valve, leaving a triangular trough below

the apex in which is seen the two scars of the dorsal pedicle muscles. ‘There is

little or no cardinal process. Under each of the dental-socket brackets a

slight cavity is present. About the centre of the valve arises the rudimentary

brachidium in the form of two lamellae directed outwards and ventrally. The

REVISION OF SOME SOUTH AFRICAN BRACHIOPODA 33

conjunct bases of these lamellae extend backwards to a point which is em-

braced by the ends of the converging buttresses of the cardinalia, and the whole

process presents the appearance of a bifurcated septum. ‘The upper extremi-

ties of the lamellae are slightly forked, but bear no accessory processes on their

external faces. With the exception of the latter, the whole structure is essen-

tially that of Megerlina lamarckiana, or at least an early stage thereof, but

whether atkinsoni should be placed in Megerlina or not is doubtful until more

specimens are examined. It should certainly be removed from Kvaussina.

I have recently received from Dr. Barnard a very small, smooth, white,

sulcate shell, showing rather strong growth-halts. The foramen is incomplete:

the deltidial-plates imperfect.

The specimen was dredged off Cape Natal (Durban) in 62 fathoms by the

Pieter Faure, and was identified by G. B. Sowerby as Araussina atkinson. Its

museum registered number is A. 5604.

For the present, I am inclined to regard it as a juvenile Megerlina pisum.

Note. The interior details of the Terebratula pisum Lamk. for Mauritius are

unknown and I have based my conclusions as to the identity of the South

African specimens on outward appearance only. If further examples are

obtained at Mauritius showing differences in cardinalia and brachidium the

South African species should be known as Megerlina natalensis Krauss.

Megerlina capensis (Adams and Reeve)

(Pl. III, fig. 8).

? Terebratula natalensis Krauss in Kiister, Conch. Cab., von Mart. and Chem.,

Bd. VII, I, 1844 and 1848, p. 36, pl. 2b, figs. 4-7. (Plate published 1844;

text 1848.)

Terebratula capensis Adams and Reeve (non Gmelin), Voyage of H.M.S. ‘Sama-

(anoeLO50; p. 71, pl. 21, fig. 4 Gn colour).

Kraussia deshayesu Davidson, Proc. Zool. Soc., 1852, p. 80, pl. 14, figs. 20-1 (in

colour).

Terebratula (Kraussia) deshayesu Dav. L. Reeve, Conch Icon., vol. XIII, 1861,

pl. 9, figs. 35a, b.

? Terebratula (Kraussia) pisum Lamk. L. Reeve, Conch. Icon., vol. XIII, 1861,

pl. 9, fig. 36b (non 36a).

Kraussina deshayestt Davidson, ‘Mon. Rec. Brach.’, Trans. Linn. Soc., ser. 2, vol.

IV, Zool. pt. II, 1887, p. 122, pl. 20, figs. 31, 31a and 31b (= same figures

asin, 1652, but in black: fig. 31a — fig. 20; fig. 31b,= fig. 21):

Description: Shell small, subovate, valves almost equal in depth; dorsal

valve sulcate, ventral broadly cardinate, type of folding sulcate. Surface of

both valves costate, some bifurcated and intercolated costae; concentric growth-

lines moderate: test finely punctate. Colour yellowish, suffused with crimson.

Beak suberect: foramen large, submesothyrid, incomplete; deltidial plates

discrete, small, trigonal. In ventral valve, pedicle-collar sessile, slightly free

in front; hinge-teeth without dental plates; interior sparsely tuberculate, with

34. ANNALS OF THE SOUTH AFRICAN MUSEUM

a fringe of stronger tubercles just within the margin. In the dorsal valve the

cardinalia and brachidium are essentially the same as in M. striata and pisum,

but the accessory processes on outer sides of divergent lamellae are represented

by slight curved ridges; interior of valve with rows of pustules radiating from

the beak, increasing in size progressively, and ending in strong spines just

within the margin.

Dimensions : Length Breadth Thickness

Figured specimen (1) 7-4 6-7 3-1 mm.

(Pl. III, fig. 8).

Others (2) 8-3 7°6 Bannan

(3) 8-5 8-2 38,

Habitat: Cape of Good Hope, 120 fathoms.

Remarks: I have seen four specimens of this species from the Cape, all of

which are in the British Museum (three in the Zoological Department and one

in the Geological Department). One of these specimens (No. 1) is now refigured

(pl. III, fig. 8). It was originally figured by Reeve (*8pl. IX, fig. 35b) and is

part of the Lombe Taylor collection in the above Institution, registered as

74.12.11.386. Reeve also figured another specimen (No. 2 above) from the

Cuming collection in the British Museum (*8pl. IX, fig. 35a).

The species was first described as Terebratula capensis in 1850 by Adams and

Reeve (’pl. X XI, fig. 4) from a specimen dredged by the ‘Samarang’ in 120

fathoms at the Cape of Good Hope. The figure (presumably natural size)

shows a shell 13:3 X 14:2 mm. in size. I have not been able to trace the

original example. Davidson, in 1852 (pl. XIV, figs. 20-1) figured and

described the species as Kraussia deshayestt as there was already a K. capensis

(Gmelin) (a synonym of K. rubra Pallas). He gave the locality as “Korea.

Coll. Cuming’. The figure is repeated by Davidson in ‘Recent Brachiopoda’

(pl. XX, figs. 31, 31a, 31b, in black), with the habitat corrected as follows:

‘Dredged by Sir Edward Belcher off the Cape of Good Hope, in a depth of 120

fathoms.’ This specimen is in the Davidson Collection, Geological Depart-

ment, British Museum (N.H.) registered as B. 12402, and measures, according

to the figure (fig. 31) 10°6 X 9°5 mm.

The shells of this species have rather large ovate pores, evenly spread over

the costae and grooves, and ranging from 224 to 280 per sq. mm. about the

middle of the ventral valve. The size of the pores (externally) is 40-50 xX

30-50 py.

The South African Museum has one perfect shell and a dorsal valve which I

attribute to this species. ‘The specimens were dredged with M. striata off East

London in 32 fathoms by the Pieter Faure and are registered as A. 5663.

The perfect shell is longitudinally oval and measures: 8-3 X 7-6 mm. It

agrees closely with Davidson’s specimen (pl. 20, fig. 31) in the geological

department of the British Museum (Nat. Hist.) and that of pl. III, fig. 8 of this

memoir. It is suffused with red. ‘The interior of the odd dorsal valve is

papillose: the brachium is broken.

REVISION OF SOME SOUTH AFRICAN BRACHIOPODA 35

Subfamily MAGELLANINAE Beecher, 1893.

Genus Terebratella d’Orbigny.

neazg. Cok. Ac. Sc. Paris, XXV, p. 260:

Terebratella rubiginosa Dall

Terebratella sp. Dall, Amer. Journ. Conch., vol. 6, 1870, p. 122, pl. 6, fig. 4.

Terebratella suffusa (non Reeve) Dall, Amer. Journ. Conch., vol. 7, 1871, p. 65.

Terebratella rubiginosa Dall, Amer. fourn. Conch., vol. 7, 1871, p. 65. Dall, Proc.

Acad. Nat. Sct. Philad., 1873, p. 135.

Terebratella(?) rubiginosa Dall. Davidson, ‘Mon. Rec. Brach.’, pt. 2, 1887,

puoi pl. 16, fic. 19 (after Dall).

Terebratella rubiginosa Dall, ‘Annot. List of Recent Brach.’, Proc. U.S. Nat. Mus.,

vol. 57, 1920, pp. 372-3.

Remarks: According to Dall (p. 372) the type locality for this species

is Simons Bay, Cape of Good Hope. ‘The solitary specimen is in the United

States National Museum. Dall reports that, ‘The species is entered in the

early Smithsonian register with a large number of mollusks collected by

Stimpson at the above locality during the Ringgold and Rodgers exploring

expedition’, and he thought there was no reason to doubt its having formed

part of that collection.

It is unfortunate that more is not known of this important member of a

typically southern genus. ‘The original specimen has been well described by

Dall and Davidson and from their statements it would appear that there is

some peculiarity in the septal attachment of the loop. It is to be hoped that

more specimens will be obtained in dredgings in Cape waters.

The nearest allied species seems to be the Terebratella enzenspergert Blochmann

from Kerguelen Is. (“Gauss’ and ‘Challenger’ Expeditions). ‘This species was

originally referred to Terebratella dorsata (Gmelin) by Davidson (°pp. 44-5,

pl. 4, fig. 4), a South American species.

In a joint report on the ‘Siboga’ brachiopods published in 1937°8 reference

was made to ? Terebratalia sp. from the region of the Sulu Islands, Moluccas,

dredged at 275 metres. Some time afterwards I received a number of other

species which had been mislaid and among them I found further and more

adult examples of the species in question and coming from the same station and

from another near by at 522 metres. They had been referred to Terebratella

dorsata by a previous worker, but are not that species, nor Terebratella sanguinea

(Leach) a New Zealand form, though coming near to the latter in general

appearance, but smaller. They appear to be a new species of Terebratella.

This occurrence of a member of the essentially Southern Hemisphere

Magellaninae in this neighbourhood is of particular interest from the point of

view of distribution. The form is now being closely studied.

36 ANNALS OF THE SOUTH AFRICAN MUSEUM

4. Conclusions

In this memoir the following six new species are described from South

African waters: Crania roseoradiata, Gryphus capensis, Terebratulina meridionalis,

Megathiris capensis, Kraussina crassicostata and Megerlina striata. ‘Vhree of these

forms have been misidentified previously, three are quite new to the South

African marine fauna-list, Megathiris being the most noteworthy. No doubt

further dredgings would add others.

Amplified descriptions are given of some of the previously known South

African brachiopods.

The South African brachiopod fauna is not a large one, but is, nevertheless,

of particular interest. ‘There are fifteen species belonging to nine genera, viz.

Crania (1 sp.), Lingula (2 spp.), Agulhasia (1 sp.), Terebratulina (2 spp.), Gryphus

(1 sp.), Megathiris (1 sp.), Kraussina (3 spp.), Megerlina (3 spp.), and Terebratella

(1 sp.). There are also a few rather doubtful records of other forms.

The peculiar genus Agulhasia is confined to South African waters, though it

is said to have Cretaceous representatives in Europe. Kvaussina, as delimited

in this memoir, is almost restricted to the Cape, the only exception being a

species—Kraussina gardineri Dall—found in the northern Indian Ocean, in

123-153 fathoms on the Saya de Malha Banks. This primitive genus is probably

an old one, but nothing is known of its geological history. The related genus,

Megerlina, has three species at the Cape, viz. striata (150 fathoms), pisum (150

fathoms), and capensis (120 fathoms), one of which, pisum, is also recorded from

a great depth at Mauritius; one representative, the genotype, M. lamarckiana

(17-110 fathoms), in south-east Australia; one, M. atkinsoni, from Tasmania

(Long Bay, 10 fathoms); and one species, M. davidsoni, found in the crater at

St. Paul Island, south Indian Ocean (tide level to 10 metres), where also

occurred Liothyrella wintert (Blochmann).

From Mauritius are also recorded Lacazella mauritiana Dall, allied to the

northern L. mediterranea (Risso); ?Gryphus cernica (Crosse) from the stomach of

a fish dredged at 80 fathoms; and Liothyrina (?Gryphus) sp. Blochmann.

At the Island of Réunion a single shell resembling the Mediterranean-East

Atlantic Mihlfeldtia truncata was found on a crustacean at 200 fathoms: it was

described under the name of Morrisca gigantea by Deshayes in 1863. Dall'é

places this tentatively in the genus Pantellaria and as possibly synonymous with

the West African (407 fathoms) Pantellaria (olim Miihlfeldiia) echinata (F. &

O.), which he also cites for the Cape of Good Hope, 224 fathoms (Jeffreys’s

coll. U.S. Nat. Mus.), as well as New South Wales (Angas coll. U.S. Nat.

Mus.). ‘This form requires further study. In an earlier part of this memoir

I referred to the presence in the British Museum (Nat. Hist.) of an immature

example of Miihlfeldtia truncata labelled ‘South Africa. J. H. Ponsonby coll.

1900’.

From the Saya de Malha Banks (123-153 fathoms) are recorded Gryphus sp.

indet. Dall, which has affinities with the northern forms G. vitreus and G.

sphenoideus; Kraussina gardinerr Dall (as stated above) and ?Aetheza (olim Hemt-

REVISION OF SOME SOUTH AFRICAN BRACHIOPODA 37

thyris) sladeni (Dall). The genotype of Aetheza is Terebratula gualtert Morris from

the Oamaruian—Eocene to Miocene—of New Zealand. The recent Hemi-

thyris colurnus Hedley from the east coast of Australia (100-250 fathoms) may

belong to this genus.

It may be of interest to refer briefly to the known brachiopod fauna from

certain places to the south-east of South Africa, especially the Marion Island

district including Prince Edward Island and Crozet Island; Kerguelen Island

and Heard Island; and St. Paul Island.

The Marion Island group is situated on an elongated submarine ridge of

less than 1,000 fathoms in depth and is separated from South Africa by sea

less than 2,000 fathoms deep. From this neighbourhood are recorded the

following species: ‘Waldheima’ (?Magellania) kerguelenensis Dav., with two

specimens of Platidia anomioides (Sc. & Ph.) attached, off Marion Is. 100 fathoms;

Terebratulina septentrionalis Dav. (?Couth.) off Marion Is. 150 fathoms; Platidia

anomioides, five examples, off Prince Edward Is., depth ?; and Liothyrella

moseleyi (Dav.), off Crozet Is. 210 fathoms.

Kerguelen and Heard Islands are situated on a similar submarine elevation

of less than 1,000 fathoms deep and separated from the Marion Island group

by sea of over 2,000 fathoms deep. From here are recorded Tereoratella

enzenspergeri Bloch. (= T. dorsata Dav. non Gmelin), Royal Sound, Kerguelen,

20-30 fathoms and Observatory Bay, Kerguelen; ‘Waldheimia’ (? Magellania)

kerguelenensis, Balfour Bay, Kerguelen, 20-60 fathoms, and Observatory Bay,

Kerguelen, 5-6 fathoms; and the same species with three examples of Tegu-

lorhynchia pyxidata (Dav.) and ?Gryphus sp. (= Terebratula uva Dav. non Brod.),

two fragments of a dead shell, off Heard Island 150 fathoms.

St. Paul Island lies on a small elevation of the sea-floor less than 1,000 fathoms

deep and separated from the Kerguelen group by sea over 1,000 fathoms deep.

The species recorded from here are Liothyrella wintert (Bloch.), 371 fathoms, and

Megerlina davidson (V élain), low water in crater.

The new observations in this memoir provide further material for the study

of former land connections in southern regions. Little advance, however, can

be made in this study until further researches are carried out in the case of the

Tertiary brachiopods of many areas. The absence of Tertiary marine rocks

in South Africa adds to the difficulty.

There is still much to be done in connexion with the clearer definition of

certain genera and species recorded from the island groups named above.

The ‘Waldheima’ kerguelenensis Dav., for example, possesses cardinalia of a

peculiar type, judging from Davidson’s figures, assuming these to be correct

Reelolil, fie. 85° pl. +X, fig.,16).. Pichler (**pl. XLIII;, fig..16) shows a

different type of cardinalia in a somewhat younger Kerguelen example of

Magellania presumed by him to be Davidson’s species.

The ?Gryphus sp. is probably a new species and not related to the Liothyrella

uva (Brod.) of South America and the Falklands. It may indeed have close

affinity with the Gryphus capensis Jackson of the Cape.

38 ANNALS OF THE SOUTH AFRICAN MUSEUM

The Terebratulina septentrionalis of Davidson from Marion Island may be

closely allied to the 7. merzdionalis Jackson of the present memoir.

The Platidia anomioides from Marion Island and Prince Edward Island is

another problem. It is apparently a wide-spread northern form (fossil and

recent) of the ancient Tethys and has spread west through the ancient Strait

of Panama to California. ‘The occurrence off Marion Island, if authentic,

would suggest a derivation from the Eastern Tethys, via the East African coast

and so to Marion Island, when the latter was connected by shallower seas

with South Africa, probably in Tertiary times. In connexion with the above

there is the interesting occurrence of the Tethyian genus Megathiris at the Cape,

and Lacazella at Mauritius and Gryphus sp. at Saya de Malha Banks.

With regard to the genera Araussina and Megerlina in South African waters,

these may have spread from the ancient India-Africa isthmus across the

Indian Ocean and have been distributed by the Mozambique and Agulhas

currents from the east coast round the Cape.

There is also to be considered the occurrence of Megerlina at St. Paul ielend

South Indian Ocean and off south-east Australia.

There are many other problems, but until the various points raised above are

clarified, it would be hazardous to dogmatize on the origin of the South African

brachiopod fauna.

So far as present evidence is concerned, it might be pointed out that the

peculiar brachiopod fauna of South Africa seems to show little affinity with

that of Antarctica, though the presence of Crania and Lingula seem to link it

with the Australian region.

More information is required regarding the duration of the free-swimming

larval stage of these marine sedentary animals before we are in a position to

answer questions as to former connections between the different oceans and

different land-masses.

If the free-swimming stage is long enough, it is possible for species to be

transported fair distances by the aid of ocean currents.

The inarticulate brachiopods are interesting from this point of view.

Brachiopoda are essentially dwellers on the continental shelves and the

majority inhabit waters less than 166 fathoms in depth: they are also influenced

by the nature of the sea-bottom; Lingulids, especially, being mud and sand

dwellers and restricted to the littoral region in warm waters. Lingula is rare

below 7 fathoms, and is usually found living in burrows in low water in estuaries

and bays, indicating a preference for habitats more or less freshened by river

waters.

The pelagic larvae of Lingula, possessing a mouth and functional stomach

and therefore self-sustaining, have been met with occasionally. ‘They have thus

considerable capacity for transportation over deep oceans in surface currents.

Yatsu, in his memoir of 1902,°° says larvae of Lingula ‘anatina of Japan may

live in the free-swimming stage in aquaria for not longer than two months.

J. H. Ashworth, in 1915,°’ gave an interesting account of the larvae of

REVISION OF SOME SOUTH AFRICAN BRACHIOPODA 39

Lingula (presumed to be anatina) obtained by him in the southern part of the

Red Sea and in the Indian Ocean about 4° south of Colombo. He also refers

to earlier records of Lingula larvae near Zamboanga, Mindanao, Philippines;

off Japan; and off South Burma. In addition, he cites the occurrence of

larvae, presumed to be of Lingula, off the west coast of Africa. There is a

species recorded from here, the Lingula parva E. A. Smith 1871, taken at

Whydah, Dahomey.

The researches of Yatsu and Ashworth seem to suggest that the free-swimming

stage of Lingula is about five or six weeks.

5. Bibliography

LIST OF PRINCIPAL REFERENCES

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XXVII, p. 20.

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XII. 1913-1924 Palaeontology ind Cee oe .. (excl. Part 7)

XIII... 1913-1923. Archaeology and Zoology ..—_—.. 2 eee

XIV. 1915-1924 Zoology .. a Be oh a 7 ae eee

XV. 1914-1916 Zoology .. se cs cece So apes a

MOVIL 9919-1999 . Botany. 3. pe oS 5 a eee

XVII. 1917-1920 Zoology: .. ... «. ats nec ee

XVIII 1921 Zoology oa oe

XIX. 1924-1925 Zoology .. a ae as ne ae ‘

XX. 1924-1926 Zoology .. ae ig aes = Ne ae:

XXI. 1925-1927 Zoology .. He Se ere ee ae ee

XXII. 1925-1926 Zoology ae oe 5 aos

XXIV. 1929-1938 ‘Anthropology and Ethnology ses ~ » (excl. Part 2);

XXV. 1927-1928 Zoology .. ae wets: Gla 292 Giguere

XXVI.- 1928 Zoology .. ve wes pie ee a eae se ae

XXVII. 1929 Anthropology .. fe We eho

XXVIII. 1929-1932 Palaeontology .. Se SE es

XXIX. 1929-1931 Zoology .. =t

XXX. 1931-1935 Zoology .. aie

INDEX of papers, authors, and subjects, published j in Vols L-Xxx. See

XXXII. 1934-1950 Palaeontology a foes a (Pare 4 ‘cams

XXXII. 1935-1940 Zoology .. na en ee : : - ees

XXXII. . 1939 Zoology .. .. he en me Pent tii

XXXIV. 1938 Zoology .. a ne

XXXV. Reserved for conclusion of i broeeaph i in Vol. XXXIV.

XXXVI. 1942-1948 Zoology ia. oe ae BSN ec ae

XXXVII. 1947- ~— Archaeology Sit. i? a ee

XXXVIII. 1950 Zoology .. Spe ae a Cobre

MAMIX:.f952 4; Zoology’ 2:00) ue. 4 at wie ine Gee ie

XL. 1952- Botany... se a4 ce ae (Part 1) —

| Copies may be obtained from—

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ny Pw 0 OW ND KD ww wD ww OO 1) U9 eB OD G9 oA GO OO OO OO oe 09 mt OO oR ip

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cere the Geological and Palaeontological parts, which are obtainable from the ge

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ANNALS

OF THE

SOUTH ‘AFRICAN MUSEUM

VOLUME XLI

PART EH, containing : —

. On South African Coccinellidae species incertae. By H. ANDREAE, D.Sc.

The genus Anobium Thunb. By H. ANDREAE, D.Sc. —

. Notes on the Ethiopian Pentatomidae. IV. A new genus and species from the

Cape Province. By D. Leston, F.Z.S., F.R.E.S. (With one text-figure.)

. Notes on the Ethiopian Pentatomoidea. X. Some specimens from southern Africa

in the South African Museum, with a note on the remarkable pygophore of

Elvisura irrorata Spin. and description of a new species of Piezodorus

Fieber. By D. Leston, F.Z.S., F.R.E.S. (With 17 text-figures.)

. Studien tiber siidafrikanischen Meloiden. By Dr. Z. Kaszas.

DEC 1 4 1953

)

LIBRARN

ISSUED OCTOBER 1953 PRICE 7s. 6d.

PRINTED FOR THE

TRUSTEES OF THE SOUTH AFRICAN MUSEUM

BY THE RUSTICA PRESS LIMITED, COURT ROAD, WYNBERG, CAPE

timate ot,

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wis

SAGES TS

py ast

pan

eS re

2. On South African Coccinellidae, species incertae. By H. ANDREAE, D.Sc., Hon.

Curator of Coleoptera, South African Museum.

Coccinella gibba ‘Thunb., Nov. Spec. Ins. 1, 1781, p. 13, f. 14, from the Cape, is

cited in W. Junk’s Coleopterorum Catalogus, pars. 118, p. 67, as Epilachna incertae

sedis. Crotch has already remarked (Revis. Cocc. 1874, p. 9): ‘Little or no

reliance can be placed on the localities’ in this publication. Yet there is a South

African species which agrees with Thunberg’s description in so far that all the

characters given are within its limits of variation, although exactly the same

combination is not found in the specimens in the South African Museum

collection. ‘The main points are:

Light brownish red, metasternum black, prothorax with a median black spot,

elytra with six black spots each, two basal, three median, forming an arcuate

band, one towards the apex near the suture; 5-8 mm. The figure shows no

spot on the prothorax and a straight band narrowed in two places, widened in

three.

Out of sixteen specimens which I refer to gibba, six have a spot on the pro-

thorax, one has the median spots (3, 4, 5) connected, one has the normal spots

6 and 8 missing (8 alone is missing in 8), the metasternum alone is black or

fuscous in 7, in the others either some abdominal segments are infuscated at the

base or the whole underside is red. 5°5-7-5 mm. Cape: Douglas; Transvaal:

Potchefstroom, Pretoria, Pietersburg, Lydenburg, Zoutpansberg.

In the Transvaal specimens, spots 1, 2 or I, 2, 5, sometimes also 6, 7 or 6, 7, 8,

are connected, the elytra showing a design like EL. hirta ab. caesarea Sic., but head

and prothorax are red.

J. Weise describes (Deutsche Ent. Keitschr. 1888, p. 83) a form from the

Kwango, northern Angola, in which spots 1 and 2 are also connected, forming

a dentate band from one humeral callus to the other; he stated that EL. hirta

Thunb. var. invalida Muls. gradually changes into var. insidiosa Muls., therefore

he gave no names to the intermediate forms. FE. gibba ‘Thunb. apparently

differs from var. invalida Muls. only by elytral spots 3, 4, 5 connected, 6 and 8

missing, central spot on the prothorax and metasternum black. As it is a very

variable species, the definition should be amended.

Light brownish red, prothorax with or without median black spot, elytra red

witha black spots (2, 3,2, 1), 1, 2 or 1, 2, 5 or 3, 4, 5, sometimes also’6, 7 or

6, 7, 8, connected, 8 or 6 and 8 may be missing. Underside red, metasternum

mostly black or fuscous, abdominal segments often infuscate at the base.

VOL. XLI. PART 2.

DEC 9

1993

42 ANNALS OF THE SOUTH AFRICAN MUSEUM

E. gibba Thunb. therefore comprises the specimens with red head and pro-

thorax so far placed in ab. discors Muls. and caesarea Sic. (both have head and

prothorax black), and znvalida Muls. with connected elytral spots and dark

metasternum.

As E. gibba (p. 13) has preference, E. hirta (p. 23) becomes a subspecies.

Key to the South African forms

1” Head and prothorax red. subsp. gibba

2” Elytra red with black markings (2, 3, 2, 1), (2, 3, 2) or (2, 3, I).

3” Some elytral spots connected, metasternum mostly black or fuscous,

abdominal segments often infuscate at base. gibba f. typ.

3’ All the spots free, underside red, only sides of metasternum and spots

on abdominal segments sometimes infuscate. ab. invalida

2’ Elytra black with free or connected yellowish markings (1, 2, 2, 1),

basal and apical lunulate, others rounded or subquadrate.

4” Markings 1, 2 or I, 2, 3 or I, 2, 3, 1 connected. ab. peringueyt

4’ All the markings free. ab. insidiosa

1’ Head and prothorax black. subsp. hirta

5’ Elytra like 2’.

6” Markings 1, 2, 3 free or connected, 4, 5, 6 free. hirta f. typ.

(Ab. 12-verrucata F. may belong here; description not available.)

6’ Markings 4, 5 forming a transverse band, the others free. ab. guttatofasciata

5’ Elytra like 2”.

7” Spots 1, 2, 5 connected, the others free, 8 (and 6) sometimes missing. _ab. caesarea

7’ All the spots free, or others than 2 and 5 connected. ab. discors

Of the other species described from the Cape in the same paper and not

identified so far, C. trinotata, p. 11, fig. 11, may be identical with or closely allied

to Martinella justitiae Gorh. from Natal; the descriptions agree fairly well,

except that in the type of justztvae the suture is partly black and the three spots

are united into one patch. Of the two specimens in the South African Museum,

the male agrees completely with the description while the female has no black

suture, only a triangular patch with rounded angles and concave sides indicating

that it is formed from three spots. So far no specimen with free spots has been

recorded.

C. oculata, p. 14, fig. 18, and C. lunata, p. 19, fig. 28, are probably not from

South Africa; Junata might be a Cyrtocaria (Madagascar).

In Germ. Mag. Ent., IV, 1821, p. 183, Wiedemann described C. erythrochila

from the Cape, collected by the Rev. Hesse, Gape Town, which has not been

recorded again. The author compares it with two species of Exochomus, auritus

(flavipes) and haemorrhoidalis, but points out that the scutellum is much larger

and the fine punctation of the elytra almost striate. The coloration is very

peculiar: aeneous, sides of prothorax and elytra with a broad, reddish-yellow

margin.

To this species I refer a specimen in the collection of the South African

Museum from Jakkalswater, Bushmanland, coll. R. M. Lightfoot, Oct. 1911,

ON SOUTH AFRICAN COCCINELLIDAE 43

although it is smaller (3 mm. instead of 2 lines) and the last abdominal segment

as well as the legs (except the posterior side of the femora) is testaceous. These

differences I take to be sexual, the type being a female and the present specimen

a male. It is a Hyperaspis allied to H. pumila Muls. but without yellow spot on

the disc of the elytra, the testaceous margin twice as broad at the shoulder,

widened towards the apex and terminated by the apical patch which is just

indicated by a slight widening of the rounded end.

3. The genus Anobium Thunb. By H. ANDREAE, D.Sc., Hon. Curator of

Coleoptera, South African Museum.

In Novae Insectorum Species, 1, 1781, p. 8, Thunberg established his genus Anobium

with six species, all recorded from South Africa, three from other countries also;

this caused some trouble and was not accepted by other entomologists. Fabricius

in Mantissa Insectorum, 1787, p. 35, mentions only two as synonyms, the others

he could obviously not identify, and in the Junk catalogue, part 23, 1910, three

are mentioned, the others remain species dubiae.

Thunberg’s description: “Antennae clava perfoliata, triarticulata. Thorax

marginatus, rotundatus. Corpus convexum, lineari-oblongum. Elytra flexilia.’

looks rather vague, but it actually applies only to some Cleridae, subfam.

Corynetinae,* and even here the genera Yenerus Cast., Tarsostenus Spin., Cory-

netinus Reitt. and Opetiopalpus Spin. are excluded; only seven genera with

together twenty-three species described from South Africa and some unnamed

species in the South African Museum’s collection were left. Under these

circumstances it was not difficult to identify all Thunberg’s species.

1. A. ruficolle Vhunb., l.c., p. 8, is now Necrobia ruficollis F., as stated by Fabricius

and Junk, Catalogus; a cosmopolitan.

2. A. capense Thunb., l.c., p. 9, is now Prosymnus capensis (Thunb.), so far

unknown. Three unnamed specimens in the museum’s collection agree

with the description, except the base of the antennae, joints 1-6, which

are rufo-testaceous; this is unimportant, the rufous or testaceous base of

antennae has also been overlooked in No. 4 and No. 6. The fasciae of

the elytra are formed by white appressed hairs, elsewhere the upper side

bears unusually long and strong black bristles. The terminal joints of

maxillary and labial palpi are elongate triangular, and the femora are

deeply grooved on the under side. The species shows all the important

characters of Prosymnus Cast., the different vestiture of the elytra alone

would not justify the establishing of a new genus. Length 3-5 to 5 mm.

Apparently rare. Cape: Cape Town, coll. J. C. Bridwell; Kalk Bay, coll.

R. M. Lightfoot; Bredasdorp, coll. H. Fry.

* In Nova Acta Soc. Sc. Upsala, VII, 1821, p. 174, Thunberg described Dasytes opacus and D. rufipes,

and transferred his Anobium coeruleum and A. viride to the genus Dasyies, citing the transfer of

coeruleum to Dasytes by Fabricius (Syst. Eleuth., II, 1801, p. 75) and accepting it, although

both species are really Cleridae and have nothing to do with Dasyies. Dasytes rufipes Thunb.,

so far not identified, is according to the description certainly different from Anobium rufipes

Thunb. and probably a Dasytes.

THE GENUS ANOBIUM THUNB. 45

3. A. bifasciatum Thunb., l.c., p. 9. Now Thricera bifasciata Thunb., syn. T.

bifasciata Hintz. The two descriptions agree very well, but Thunberg’s

figure (rather poor and not agreeing with the description) has probably so

far prevented the identification. Cape: widely distributed but rare.

Described by Hintz (Deutsche Ent. Zeitschr., 1902, p. 403) as T. bifasciata

from Dunbrody, two spec., probably coll. Father O’Neil; by Gorham

(Proc. Zool. Soc. Lond., 1905, 2, p. 274) as T. bicinctella from Port Elizabeth,

two spec., coll. Dr. H. Brauns. In South African Museum coll., one spec.

from Dunbrody, apparently coll. by Father O’Neil, no date, and two

from Knysna, Oct. 1916, coll. L. Peringuey. I found one at Parow near

Cape Town, 28/8/47, and one on the farm Tierhoek, Piquetberg Mts.,

19/10/47. The var. tricolor Hintz is recorded from Dunbrody (Hintz), 2

spec.), Port Elizabeth (Gorham, 2 spec.) and East London (South African

Museum coll., Oct. 1912, R. M. Lightfoot, one spec.).

viride Thunb., l.c., p. 9, now type of the genus Notostenus Spin. (Clerites IT,

1844, p. 89). A well-known species, frequent on flowers of the arum lily.

Thunberg’s statement: ‘Habitat in Africa & India Oriental’ is a mistake,

the species is South African. One hundred and ten specimens in the South

African Museum coll., mostly from the western Cape, but also from Port

St. Johns and Durban.

5. A. rufipes Thunb., l.c., p. 10, is Necrobia rufipes De Geer. This was stated by

Fabricius (Mant. Ins., 1787, p. 35), but not taken over by Schenkling in

Junk, Catalogus, part 23; a cosmopolitan.

6. A. coeruleum Thunb., l.c., p. 10, is a Dolichopsis, not a Notostenus, as stated in

Junk, Catalogus, part 23, p. 136. A very variable species, colour from

golden green to violaceous blue, elytra different from head and prothorax

or upper side unicolorous, antennae with articles 2 to 4 (sometimes 5 also)

testaceous, rufous or dark fuscous above, rufous below, prothorax fairly

convex or rather flat, punctation uniform and rather close or sparse on

the disc, especially in the anterior half, length 2-5 to 5mm. The extreme

forms may easily be mistaken for different species, but the intermediate

forms make any division impossible. Frequent from Cape Town to the

Cedarbergen and Zwartbergen, on flowers of various plants, especially

Ficoideae, Selaginaceae and Compositae. Dolichopsis cyanella Gorh. (Trans.

Ent. Soc. Lond., 1878, p. 155) is the same species and therefore a synonym.

4. Notes on the Ethiopian Pentatomidae. IV. A new genus and species from the Cape

Province. By D. Leston, F.Z.S., F.R.E.S. (With one text-figure.)

Carvalhocoris gen. nov.

MobDERATE size, oval; base of venter tuberculate; mesosternum with a low

but distinct carina of even height throughout; odoriferous aperture large and

oblique, continued by a long, smooth and gently curving ‘horn’ towards and

almost reaching the anterior border of the mesosternum at its furthermost point;

evaporating area distinct, its lateral border raised; rostrum reaching to posterior

trochanters, second segment longest, third widest and subequal to the fourth.

2 Carvalhocoris nigra gen. et sp. nov. §

NOTES ON THE ETHIOPIAN PENTATOMIDAE 47

Head breadth to length ratio as 3: 2; greatly deflected, sides sinuate, apex

widely rounded, margins not reflexed; interlobar sutures straight; upper surface

slightly convex.

Pronotum with anterior margin broadly emarginate, anterior border between

and behind the eyes smooth and impunctate; lateral margins reflexed, more so

anteriorly, rounded gradually but slightly, entire; postero-lateral angles

rounded, obtuse, rather raised; posterior border gently emarginate, postero-

internal angles about 150°.

Scutellum long, broadly rounded at apex, sides narrowing just before the

middle; disc raised slightly, posterior portion deflected gradually downwards.

Apex of corium reaching caudally as far as the apex of the scutellum; membrane

dark, almost obscuring the few parallel veins; reaching to apex of abdomen.

The whole upper surface densely punctate, rugosely so on the scutellum,

connexivum punctate. Tibiae flat and canaliculate externally. Antennae short,

ratios of segments 6: 10: 11: 14: 13; fourth and fifth segments slightly thicker

than second and third; ‘first not surpassing apex of head. Head, thorax and

abdomen beneath punctate but less so than the upper surface.

Type: Carvalhocoris nigra sp. nov.

This genus should be placed in Pentatominae, tribe Pentatomini; it pertains to

most of the genera placed in Menidaria Distant. From Amphimachus Stal it can

be distinguished by the shorter scutellum, from Aegaleus Stal by the absence of

impunctate fasciae on the scutellum and from Menida Motsch. by the smaller

ventral spine. This last genus has been used as a dumping ground for many

species and is a composite one as at present used; it is possible that certain

species included in it should be placed within Carvathocoris.

Carvalhocoris nigra sp. nov.

Shining black except for the antennae, these flavous, darker apically. Head

above with a fine narrow flavous margin anteriorly; anterior margin of pro-

notum narrowly flavous but irregularly so in the centre; lateral margins of the

pronotum narrowly flavous; corio-pronotal junction externally flavous.

Connexivum flavous with contiguous black spots either side of the sutures.

Under side flavous with dark brown punctures; tibiae and femora flavous

spotted with brown, heavily so on the latter. A linear series of three large black

spots, one on each thoracic pleurite centrally. Ventral spiracles brown. Length,

26mm.; maximum pronotal width, 9 4 mm.

Type: @ Prieska, Cape of Good Hope, October 1887 (now in South

African Museum, Cape Town).

Paratypes: 4 Aliwal North, Cape Province, December 1922. @Q same,

4,350 ft., January 1923. (Both collected by Turner and now in British Museum

I am happy to be able to name this genus in honour of Dr. J. C. M. Carvalho,

the eminent Brazilian hemipterist and explorer. My thanks are tendered to

the Director, and Dr. Hesse, of the South African Museum, for submitting a

small collection of their specimens to me for determination.

5. Notes on the Ethiopian Pentatomoidea. X. Some specimens from southern Africa in

the South African Museum, with a note on the remarkable pygophore of Elvisura

irrorata Spinola and description of a new species of Piezodorus Fieber. By

D. Lesron, F.Z.S., F.R.E.S. (With 17 text-figures.)

By courtesy of the Director and Dr. A. J. Hesse, I have examined a collection

of previously undetermined Shieldbugs belonging to the South African Museum,

Cape Town. Most of the specimens were taken at localities within, or adjacent

to, the Union.

The literature contains not a few references to the fauna of South Africa; the

pioneer workers were Palisot de Beauvois, Thunberg and Germar. The first

important work is that of Stal (1865); in the first volume of Hemiptera Africana

he collated all the earlier records and placed them in recognizable modern

genera. Later, Stal (1876) brought his classification up to date and the broad

outlines of generic and higher taxonomy in use to-day are entirely due to this

great Swedish entomologist.

It is only in later works that localities are given with sufficient precision to

form a basis for zoogeography, and these works include Wallengren (1875) ;

Distant (1892, 1898); Schouteden (1912); Schumacher (1913); Hesse (1925,

1935) and Leston (1952a, 19525). Besides these, new species from the area are

described in papers by Bergroth, Distant, Montandon, Jensen-Haarup,

Schouteden and Leston, as well as in the catalogues of Westwood, Dallas and

Walker. Solely to aid further knowledge of the distribution of species, the

following list is given. |

BRACHYPLATIDAE Lest., 1952a (Plataspidae)

Gelastaspis brown Kirk.

1902. Kirkaldy. Entomologist. xxxv, p. 166

Umtali, Southern Rhodesia (A. Bodong)

CypnipAE Billberg, 1820

Legnotus tibialis Stal

1853. Stal, Ofv. Vet. Ak. Forh, x, p. 222

M’fongosi, Zululand, iv—v, 1934, xii-li, 1934-5, v, 1935 (W. E. Jones)

Legnotus melaleucus (Thunb.)

1783. Thunberg, Nov. Ins. Sp., ii, p. 50

Kamieskroon, Namaqualand, ix, 1930 (South African Museum staff)

NOTES ON THE ETHIOPIAN PENTATOMOIDEA 49

PENTATOMIDAE Leach, 1815

SCUTELLERINAE Leach, 1815

SCUTELLERINI Leach, 1815

The nomenclature and classification of the Scutellerinae have recently been

investigated (Leston, 1953a, 19530); as a result, the groups Scutellerin: Leach,

Sphaerocorini Stal and Elvisurini Stal are now treated as subtribes within Scutel-

lerini. All three subtribes occur in the Union of South Africa.

Elvisura trrorata Spinola

1837. Spinola, Essai Hem. p. 359

Karkloof (now: Kloof), Natal, x11, 1915 (Bell Marley)

This was a female, but through the courtesy of the British Museum (Nat.

Hist.) authorities I have been able to examine the genitalia of a male of this

rarity. The genus is distinct from all other Scuiellerinae in that the scutellum is

distinctly keeled and the bugs have a marked resemblance to Buprestid beetles.

The male genitalia are shown in figures 1-4. The pygophore is large with a

well-developed caudal lip: the cephalad margin is remarkable for the presence

of a pair of large, flat, rectangular and freely movable flaps attached immediately

within the margin; they appear to be adherent throughout their length but

with a root-like attachment at their outer corner. The flaps are very thin but

sclerotized, with a pale band along the proximal border; the distal border is

fringed by a few fine hairs (not shown in the figure).

In work now in progress on the Angolan Pentatomid fauna a pair of movable

pieces has been found on the internal-lateral margins of the pygophore in

Crollius Dist.; they were triangular and large so that, when brought together in

the resting position, they completely covered the pygophoral opening. Dr. R. I.

Sailer of the U.S. National Museum has informed me, in Uitt., that he has

discovered similar processes on the pygophore of other Podofini, including

Podops Lap.; he proposes to call them hypopygeal appendages. It is probable that

the hypopygeal appendages are homologous in the Podopid genera with the

flaps noted in Eluisura irrorata; thus it is proposed to adopt this term for the

structures noted in FE. zrrorata, but the difference in position of the appendages

in the two groups must, of course, be stressed; it is not a major difference

presenting insuperable difficulties in homology. A difficulty is met, however,

in assessing its taxonomic import; is it a primitive character? The little known

of the life-histories of Podops and of Elvisura suggests quite different ecological

roles so that structural convergence is hardly to be considered seriously.

The theca of E. irrorata (figs. 2 and 3) is strongly sclerotized and short; it is

quite impossible for the vesica and conjunctival appendages to be retracted (a

constant character in the Scutellerint). Ventrally there projects a pair of appen-

dages, third conjunctival appendages in Singh-Pruthi’s terminology; they are

sclerotic except towards the base, where they fuse, and are thus freely movable.

On the dorsal surface of the theca project the fused first and second conjunctival

50 ANNALS OF THE SOUTH AFRICAN MUSEUM

appendages. As in all other Scutellerini, the second is more sclerotized while the

first is membraneous except for the tips. The first conjunctival appendages are

apically bifid and terminally sharp. The principal difference to be noted in the

Elvisura trrorata Spinola

1. Pygophore from above. H, hypopygeal appendages; p, posterior margin.

2. Aedeagus, ventral. B, basal plates; T, theca; V, vesica; a, b, c, first, second and

third conjunctival appendages.

3. Aedeagus, lateral. S, supravesical process.

4. Paramere, lateral.

Scale = 0:5 mm.

appendages of Elvisura from those of other genera is the large size of the common

basal area of the first and second conjunctival appendages. The third pair of

appendages are covered, sparsely, by minute points.

The vesica exhibits the usual basal convolutions and well-developed ejacula-

tory reservoir while the gonopore is oblique and large: these are all common

characteristics of the Scutellerint but, in addition, the external part of the vesica

NOTES ON THE ETHIOPIAN PENTATOMOIDEA i

supports a rigid, densely sclerotized, thin and rough-edged vertical lamina. This

structure has also been noted in Steganocerus multipunctatus and (Leston, 19536)

termed the supravesical process. It is flecked in E. zrrorata by a few minute

broad-based spines and whereas in S. multipunctatus the supravesical process is

somewhat thick and terminated at its highest point forwards by a bifurcation,

in £. irrorata it is thin and simple anteriorly, albeit very ragged at the

margins.

As in most Scutellerint, the actual path taken by the ductus seminis remains

untraced due to the density and complexity of the vesica; the basal plates have

not been studied nor figured except merely to show their attachment.

The parameres are simple hook-like pieces with a ridge of hairs; they take a

form common to many Scutellerint.

Thus the entire male genitalia of FE. zrrorata is of the usual Scutellerinine

pattern as described elsewhere (Leston, 1953); it differs in possessing (1) a

supravesical process similar to that found in Steganocerus and (2) a pair of movable

processes on the pygophore which are tentatively homologized with the hypo-

pygeal processes of certain genera of Podopini.

Cryptacrus comes (Fabr.)

1803. Fabricius, Syst. Rhyngotorum, p. 130

Amatongas Forest (between Umtali and Beira); the specimen is of

var. apicalis Dist.

PacHycorin! Dallas, 1851 (Tetyrinz)

Deroplax circumducta (Germar)

1837. Germar, Rev. Ent. (Silbermann), v, p. 190

Vryburg, Bechuanaland, 1904 (Jones); Otjitundua (160 miles north-west of

Outjo), South West Africa, 11, 1926 (S. Afr. Mus. Exp.). The South West

African specimen is close to D. nigropunctata (Stal), but Iam unable to distinguish

between these two species. ‘They must remain distinct pending an examination

of Stal’s type.

Deroplax silphoides (Thunb.)

1783. ‘Thunberg, Nov. Ins. Sp., ii, p. 29

Pretoria, Transvaal, x, 1944 (R. Wilson). This specimen is similarly marked

to a long series in the British Museum from Nyasaland. It is referred to the

variable D. silphoides. Figs. 5-8 depict the range in variation; it is considered

undesirable to add further to the number of named varieties of this species.

The male genitalia have been examined (figs. 9-12); they are similar to the

genitalia of D. circumducta figured elsewhere (Leston, 1953) and quite unlike

the genitalia of Scutellerin. ‘The pygophore encloses a wide proctiger which is

marked by a pale band near the base. The aedeagus, shown laterally in fig. 9,

encloses a long filimentous vesica of the ‘penisfilum’ type in Baker’s terminology

(Baker, 1931). Surrounding the vesica is a long, finger-like extenstion of the

conjunctiva. The first and second conjunctival appendages only are present

52 ANNALS OF THE SOUTH AFRICAN MUSEUM

(the third is always pre-

sent in Scutellerini); they

are, as usual, joined

basally. The first pair is

long and membraneous,

terminating in a lightly

sclerotized point; the

second is biramous,

sclerotized, with one arm

produced into a filiform

process extending to the

apex of the vesica. The

parameres are shown in

two views (figs. 11, 12);

the head bifurcates and is

closely spinose. On the

dorsum of the parameres

is a tubercle carrying an

armature of bristles while

the basal area of the

tubercle is similarly

armed. It shouldbe

noted that figures g and

10 are somewhat dia-

grammatic.

Deroplax silphoides (Thunb.). 5-8. Variation

in a variety from Nyasaland.

AMYOTINAE Lest., 1953a (Asopinae)

Dorycoris pavoninus (Westw.)

1837. Westwood, Cat. Hemipt., coll. Hope, i, p. 39

Stellenbosch, iv, 1932 (P. v. Heerden). The specimen is of a form near to

var. fuscosus.

PopoPiNnaE Dallas, 1851 (Graphosomatinae)

This subfamily includes the two tribes Podopint Dallas and Tarisim Stal

(Graphosomin auctt.). How far it is a natural assemblage remains for future

investigation to elucidate, but while the Podopini are reasonably homogeneous

the Tarisini are certainly not so and probably many of the species are Pentatominae.

Bolbocoris rufus (Westw.)

1837. Westwood, op. cit., i, p. 12

Kamanyab, S.W.A., ii, 1925 (S. Afr. Mus. Exp.); Kaross (30 miles NW.

of Kamanyab), S.W.A., ii, 1925 (S. Afr. Mus. Exp.). Despite the widespread

NOTES ON THE ETHIOPIAN PENTATOMOIDEA 53

distribution of two of its species the genus Bolbocoris is centred in South Africa,

with at least five species in the Transvaal.

PENTATOMINAE Leach, 1815

This enormous subfamily is overdue for revision, but until this has been

attempted the Stalian groups are retained.

co a

Deroplax silphoides (Thunb.)

9. Aedeagus, lateral. t, theca; co, conjunctiva; a, b, first and second conjunctival

appendages; d, ductus seminis; v, vesica; d.e, ductus ejaculatorius.

10. Pygophore, terminal. p, proctiger; r, right paramere.

11, 12. Two views of right paramere.

PENTATOMINI Leach, 1815

Menida lythrodes (Germar)

1837. Germar, Rev. Ent. (Silbermann), v, p. 175

Letaba, Transvaal, v and xii, 1945 (E. C. G. Bedford). ‘From orange-tree

—fed on red-scale and on orange.’

54 ANNALS OF THE SOUTH AFRICAN MUSEUM

Menida decoratula (Stal)

1853. Stal, Ofv. Vet. Ak. Forh., x, p. 222

Kaoko Otavi, S.W.A., iii, 1926 (S. Afr. Mus. Exp.).

Menida distanti Horvath

1892. Horvath, Term. Fiizet, xv, p. 258

Louis Trichardt, ‘Transvaal, i, 1928 (R. F. Lawrence); Bulawayo, Southern

Rhodesia, v, 1917 (R. W. E. Tucker); Mtunzini, Zululand, vi, 1940.

Agonoscelis puberula Stal

1853. Stal, op. cit., x, p. 216

Outjo, S.W.A. (S. Afr. Mus. Exp.); Potgietersrust, Transvaal, iv, 1934

(R. F. Lawrence); Upington, Cape Province, vii, 1936 (S. Afr. Mus. staff).

Agonoscelis odendaal Dist.

1910. Distant, Ann. Mag. Nat. Hist., (8), vi, p. 95

Salisbury, Southern Rhodesia, v, 1913.

Agonoscelis erosa (Westw.)

1837. Westwood, op. cit., i, p. 33

Outjo, S.W.A., 1, 1926 (S. Afr. Mus. Exp.).

7 Amphimachus circumflexus (Stal)

1855. Stal, op. cit., xi, p. 182

Florida, Transvaal, x, 1918 (R. W. E. Tucker). No locality was given by

Stal; the majority of specimens seen come from the Transvaal.

Antestia lymphata Kirk.

1909. Kirkaldy, Cat. Hemiptera, i, p. 129

Salisbury, Southern Rhodesia. This specimen has been reported upon else-

where (Leston, 1952c); it is distinct from the coffee-bugs and these have been

removed to Antestiopsis Lest. A. lymphata Kirk. appears to be extremely rare;

the British Museum possesses only Dallas’s original type and paratype, described

by him as Pentatoma maculata. ‘The South African Museum’s specimen is the

first to have a precise locality.

Gen. Piezodorus Fieber

1861. Fieber, Europ. Hem., \xx, p. 329

1905. Jakovlev, Rev. russe Ent.., v, p. 142 (Pausias)

Pausias, type Piezodorus martin Put., is only distinguishable from Pzezodorus in

that the spiracles are pale, whereas in the latter they are dark; P. hessez has pale

spiracles, but in all other respects it is a true Piezodorus. ‘These conditions are

best satisfied, as are the zoogeographical factors, by sinking Pauszas (Syn. Nov.).

When dealing with a large number of specimens from many parts of Africa

it has been found that the usual taxonomic characters: length of ventral spine,

shape of mesosternal carina, antennal ratios, etc., give little help in separating

species. The separation of P. pallescens, purus, and hybnert can probably only be

based on the genitalia of the males, but the degree of variation in the parameres

NOTES ON THE ETHIOPIAN PENTATOMOIDEA 55

needs further investigation. It appears that the three might best be considered

as facets of a single polytypic species; the zoogeography has not helped in this

group and ecological factors are probably the cause of the different ‘species’.

Key to the African (south of the Equator) species

The only other species reported from the area is P. bequaerti Schout. from the

Belgian Congo.

1. Spiracles pale. Length over 11 mm. hesset

Spiracles dark. Length less than 11 mm. 2

2. Posterior tibiae usually flattened. Second antennal segment always longer

than third. purus

Posterior tibiae not flattened. Second antennal segment equal to or shorter

than third. 3

3. Parameres as in fig. 16. hybnert

Parameres more elaborate. pallescens

Prezodorus purus (Stal)

MORAG Sel, Clos Gtis, 3G 0, OI

Kaoko Otavi, $.W.A., ii, 1926; Warmbad (12 miles SE. of Zesfontein),

Kaokoveld, S.W.A., ii, 1925; Outjo, S.W.A., i, 1926; Kamanyab, $.W.A.,

iii, 1925; Kaross (30 miles NW. of Kamanyab), S.W.A., ii, 1925 (all S. Afr.

Mus. Exp.).

Keimoes and Riemvasmak, Gordonia, Cape Province, vii, 1925 (K. H.

Barnard); Aughrabies Falls, Kenhardt Div., Cape Province, v, 1934 (R. F.

Lawrence); Murraysburg District, Cape Province, ili, 1931 (Mus. staff).

The determination of Prezodorus species is a matter of some difficulty; the

species are variable and the Oriental hybnert (Gmel.), now spread over much of

East and Central Africa, has been confused with purus by authors. The genitalia

are very similar throughout the genus.

Fig. 13 shows the aedeagus of purus laterally; the conjunctival appendages

are very large and almost entirely membraneous; it has been found difficult

to homologize them with the appendages in other groups. The head of the

paramere (fig. 14) enables this species to be distinguished from both Aybneri and

pallescens (Germ.). The latter is also a common species in South Africa.

Piezodorus hesset sp. nov.

®. Pale yellow-brown, a red band across the pronotum between the postero-

external angles, ante-ocular area of head red; a black metallic line just within

and parallel with the lateral paraclypeal margins. Hemielytra with a brown

speck on the apex of the median margin of the clavus. Antennae indefinitely

reddish. Venter pale yellow-brown, spiracular eminences pale dirty-yellow.

Pronotum and head punctate, the former with its anterior margin raised and

rounded, its lateral margins straight, reflexed and yellow. Elytral texture thin,

semi-transparent. Membrane glassy, colourless. Antennal segments 0-42, 1-00,

56 ANNALS OF THE SOUTH AFRICAN MUSEUM

1:16, 1-11 and 1-01 mm. long respectively. Ventral spine stout, reaching to

before the intermediate coxae.

@ Length 12-4 mm. Maximum pronotal breadth 6-2 mm.

6 Length 11-8 mm. Maximum pronotal breadth 6:2 mm.

Holotype 9: Otjikondo (40 miles WNW. of Outjo, S.W.A., 1, 1925 (S. Afr.

Mus. Exp.).

Piezodorus purus Stal. 13, aedeagus, lateral. 14, head of paramere, lateral.

Piezodorus hessei sp. nov. 15, metathoracic gland opening, and evaporatorium.

Piezodorus hybneri (Gmel.). 16, head of paramere, lateral.

Piezodorus hesset sp. nov. 17, head of paramere, lateral.

Paratypes: Three ¢ ¢ same locality as holotype; one 9 Kaoko Otavi,

Kaokoveld, S.W.A., ili, 1926; one ¢ Otshu, Hoarusib River (50 miles W. of

Kaoko Otavi), S.W.A., ii, 1926; one 3 Kaross (30 miles NW. of Kamanyab),

S.W.A., ii, 1925 (all S. Afr. Mus. Exped.).

Holotype and paratypes in the South African Museum except for one para-

type presented to the British Museum (Nat. Hist.), and two retained in the

author’s collection.

NOTES ON THE ETHIOPIAN PENTATOMOIDEA 57

Gynenica capenert Lest.

1953. Leston, Rev. zool. bot. Afr. (in press)

M’fongosi, Zululand, i, 1935 (W. E. Jones). ‘This species appears to be con-

fined to Natal and Zululand.

Gynenica marginella Dallas

1851. Dallas, List. Hemipt. Brit. Mus., i, p. 181

Resolution, Grahamstown, Cape Province, i-iv, 1928 (Miss Walton).

Stenozygum alienatum (Fabr.)

1803. Fabricius, Syst. Rhyngotorum, p. 173

Wankie, Southern Rhodesia, xi, 1923 (C. W. Tyler).

Boerias brunnea Jensen-H.

1931. Jensen-Haarup, Ent. Medd., xvii, p. 325

Knysna, Cape Province, 1, 1931 (K. H. Barnard). The previously known

localities for this species are given by Leston (19526); it is confined to the south-

west of the Cape Province.

Boerias maculata ( Dist.)

1910. Distant, op. cit., (8), vi, p. 87

Warrenton, Cape Province, iv, 1931 (J. T’.). Not previously recorded from

the Cape Province, but known from Southern Rhodesia, the Transvaal, and

Basutoland.

Boerias rubrocincta (Dist.)

1910. Distant, op. cit., (8), vi, p. 87

Pretoria, Transvaal, v, 1927 (S.M.); Smithfield, O.F.S., 1910 (Kanne-

meyer), ‘On orangia’. ‘These two records considerably extend the published

distribution. Distant described the species from Natal.

Boerias victorin (Stal)

1856. Stal, op. cit., xiii, p. 194

[nec Jeannel, 1913]

Keurbooms River, Knysna, Cape Province, i, 1931 (K. H. Barnard).

Durmia haedula (Stal)

1865. Stal, Hemipt. Afric., i, p. 149

Louis Trichardt, Transvaal, i-1i, 1928 (R. F. Lawrence).

Durmia tomentwentris (Germ.)

1837. Germar, Rev. Ent. (Silbermann), v, p. 168

M’fongosi, Zululand, i-ii, 1935 (W. E. Jones).

Carbula litigatrix Kirk.

1909. Kirkaldy, Cat. Hemipt., 1, p. 88

M’fongosi, Zululand, 1, 1935 (W. E. Jones); Cayimaeis (Caimaiais, 25 miles

SE. of Zesfontein), Kaokoveld, $.W.A., iii, 1925 (S. Afr. Mus. Exp.).

53 ANNALS OF THE SOUTH AFRICAN MUSEUM

Veterna sanguineirostris (Thunb.)

1822. Thunberg, Hemipt. Rostr. Cap., 11, p. 4

1890. Distant, C.R. Soc. ent. Belge, xxxiv, p. lv (mimica)

1892. id., Naturalist in Transvaal, p. 250 (patula)

I can find no constant characters upon which to separate V. patula Dist. and

V. mimica Dist. from this species; they are weak but geographical subspecies;

sanguineirostris centred in the Cape Province, patula in the Transvaal, Natal, and

Southern Rhodesia, and mimica in the Belgian Congo. (Syn. Nov.)

Subspecies sanguineirostris

Somerset West, Cape Province, vili-ix (A. J. Hesse).

Subspecies patula

Pretoria, Transvaal, xi, 1930.

Diploxys fallax Stal

1865. Stal, Hemipt. Afric., i, p. 129

M’fongosi, Zululand, i, 1935 (W. E. Jones).

Coponia waterbergensis (Dist.)

1902. Distant, Ann. S. Afr. Mus., ii, p. 252

Louis Trichardt, ‘Transvaal, i-ii, 1928 (R. F. Lawrence). The specific distinc-

tions are obscure in this genus but the specimen before me has been compared

with Distant’s type. Undoubtedly C. cornuta (Dist.), C. thoracica (Dist.) and

C. waterbergensis are very closely allied, and their differences, based on the pro-

notal angles, are scarcely specific.

| Lerida punctata (P. de Beauv.)

1805. Palisot de Beauvois, Ins. Afr. Amer., p. 84

M’fongosi, Zululand, 1, 1935 (W. E. Jones).

Farnya versicolor (Dist.)

1881. Distant, Proc. <ool. Soc. Lond., p. 271

Junction of Marico and Limpopo Rivers, Transvaal, ii, 1887. (A. W.

Eriksson.) ‘This was collected by the celebrated explorer-collector; it is the

first Transvaal record. Elsewhere (Leston, 1952a) I have reported its capture

in Natal.

Aethemenes stalianus Kirk.

1909. Kirkaldy, Cat. Hemipt., i, p. 47

Bindura, Southern Rhodesia (D. Coghill).

Antestiopsis orbitalis (Westw.) (Comb. Nov.)

1837. Westwood, Cat. Hemipt. coll. Hope, i, p. 35

Michell’s Pass, Ceres, Gape Province, x, 1934 (Museum staff).

Halydicoris corticinus (Germ.)

1837. Germar, op. cit., v, p. 178

Pretoria, Transvaal, ii, 1930 (1.B.K.); Port St. Johns, Cape Province (G.

Shortridge); Schuiverberg, Transvaal.

NOTES ON THE ETHIOPIAN PENTATOMOIDEA 59

PHYLLOCEPHALINI Dallas 1851

Amyot and Serville introduced the name Phyllocephalides for this group,

but being vernacular it is invalid. The first Latin group name is Dallas’s

Phyllocephalidae.

Dalsira projecta (Dist.)

1898. Distant, Ann. Mag. Nat. Hist. (7), 1, p. 303

‘Transvaal’.

Dalsira subtruncata (Walker)

1868. Walker, Cat. Heteropt. Hemipt. Brit. Mus., iii, p. 491

Letaba, Transvaal, xi, 1948.

Hatyini Spinola, 1850

Atelocera notatipennis Stal

1858. Stal, Ofv. Vet. Ak. Forh., xv, p. 312

Moroqueng (Morokwen), Bechuanaland, 1905; Vryburg, Bechuanaland, x,

1939 (Mus. staff).

Atelocera natalensis Stal

TOHGe Ocal) OPmeles xen: 2116

Northern Damaraland, i, 1888 (A. W. Eriksson).

AEPTINI Stal, 1876

Aeptus singularis Dallas

1851. Dallas, List. Hemipt. coll. Brit. Mus., i, p. 146

Louis ‘Trichardt, Transvaal, 1, 1928 (R. F. Lawrence).

Myrocueini Stal, 1876

Delegorguella atomaria (Dallas)

Hope Dallas op Acie, op. 130

Vryburg, Bechuanaland, 1-ii, 1930 (C. G. Stone).

Neococalus clausus (Walker)

1867. Walker, Cat. Heteropt. Hemipt. coll. Brit. Mus., i, p. 175

Acornhoek, East Transvaal, xi, 1918 (R. W. Tucker); Port St. Johns, Cape

Province; Willowvale, Eastern Cape Province, 1, 1917.

DInIDoRINAE Stal, 1870

Coridius nubilus (Westw.) (Comb. Nov.)

1837. Westwood, op. cit., I, p. 25

Pretoria, Transvaal, i1, 1932; Vredefort, O.F.S., 1902; Port St. Johns, Cape

Province, 1902 (G. Shortridge); Kakamas, Kenhardt Div. v, 1934 (R. F.

Lawrence).

60 ANNALS OF THE SOUTH AFRICAN MUSEUM

REFERENCES

Distant, W. L., 1892, A naturalist in the Transvaal, London.

Loe 1898, ‘Rhynchota from the Transvaal, Mashonaland and British Nyasaland’,

Ann. Mag. nat. Hist., (7) 2: 294-316.

Hesse, A. J., 1925, ‘Contributions to a knowledge of the fauna of S.W. Africa. iv. A list of the

Heteropterous and Homopterous Hemiptera of S.W. Africa’, Ann. S. Afr. Mus., xxiii: 1-190.

id., 1935, ‘Scientific results of the Vernay-Lang Kalahari expedition, March to Sep-

tember, 1930’, Ann. Transvaal Mus., xvi: 581-603.

Leston, D., 1952a, ‘Notes on the Ethiopian Pentatomoidea, v. On the specimens collected by

Mr. A. L. Capener, mainly in Natal, Ann. Mag. nat. Hist., (12) 5: 512-20.

id., 19526, ‘Notes on the Ethiopian Pentatomoidea. vi. Some insects in the Hope

Department, Oxford’, Ann. Mag. nat. Hist., (12) 5: 893-904.

ide 1952c, ‘Notes on the Ethiopian Pentatomoidea. i. The genotype of Antestia’, Rev.

zool. bot. Afr., xlv: 268-70.

itl 1953a, “Che suprageneric nomenclature of the British Pentatomoidea’, Ent. Gaz.,

iv: in press.

id., 19530, ‘Notes on the Ethiopian Pentatomoidea. viii. Scutellerinae Leach of Angola,

with remarks upon the male genitalia and classification of the subfamily’, Publ. Cult. Comp.

Diam. Angola: in press.

Schouteden, H., 1912, ‘Cimicidae, Coreidae, Reduviidae de la région du Zoutpansberg

(Transvaal)’, Rev. zool. Afr., ii: 101-14.

Schumacher, F., 1913, ‘Ein Beitrag zur Kenntnis der Rhynchoten-Fauna Siidafrikas, insbeson-

dere von Deutsch-Stidwestafrika, Klein-Namaland und dem Kalaharigebiet’, Schultze,

Reise, v (Denkschr. med. Ges. Fena, xvii: 49-88).

Stal, C., 1865, Hemiptera Africana, i, Stockholm.

id., 1876, ‘Enumeratio Hemipterorum, v’, K. Svensk Vet.-Ak. Handl., xiv (4): 1-162.

Wallengren, H. D. J., 1875, ‘Insecta Transvaaliensia—Bidrag till Transvaalska Republikens i

Sodra Afrika insektfauna’, Ofv. K. Vet.-Ak. Forh., xxxii (1): 83-137.

©. Studien tiber stidafrikanischen Meloiden (Coleoptera). Von Dr. Z. Kaszas,

Ungarisches Naturwissenschaftliches Museum, Budapest.

Herr A. J. HEssE war so liebenswiirdig und schickte mir einige von Péringuey

beschriebene Meloiden, sowie die undeterminierten, meist neuen Arten der

Sammlung des South African Museums in Cape Town, zur Untersuchung.

Aus diesem Material beschreibe ich im folgenden 12 neue Arten, ausserdem

wurde es mir moglich die Gattung Jselma Haag-R. naher zu studieren und statt

der vollkommen unbrauchbaren Bestimmungstabelle von Péringuey, einen

neuen, auf Grund morphologischer Merkmale aufgebauten Bestimmungs-

schliissel zu publizieren. Diese Arbeit soll nur eine Vorstudie zu einer grésseren

monographischen Aufarbeitung der afrikanischen Meloiden sein, an welcher

ich gegenwa4rtig arbeite.

Gen. Jselma Haag-R.

1879. Haag-Rutenberg, D. Ent. Z., xxiii, p. 402.

Iselma lanuginosa sp. nov.

Einfarbig glanzend schwarz, Ober- und Unterseite, besonders aber der

Vorderk6érper sehr lang und dicht, aufstehend schwarz behaart. Kopf breit

und rundlich, Stirn gewolbt, Scheitel breit abgerundet. Schlafen lang, Stirn

zwischen den Augen hinten bis zum Scheitel in der Mittellinie unpunktiert und

etwas aufgebogen, die Punktierung sehr fein und dicht. Halsschild deutlich

breiter als der Kopf, etwa so lang wie breit, vor der Mitte am breitesten, nach

vorne gerade und pl6otzlich, hinten ebenfalls ziemlich stark verschmalert, Basis

nur schwach gerandet, die Mitte breit und flach eingedriickt, Scheibe beider-

seits neben der Mittellinie etwas weiter nach hinten grubenartig vertieft. Die

Punktierung auf der Scheibe sparlich, an den Seiten aber dicht, dazwischen ist

der Grund glatt. Die Behaarung einfach aufstehend, ohne anliegende Behaa-

rung. Fliigeldecken an der Basis sehr breit, nach hinten verschmalernd, die Basis

an der Innenseite der Schultern breit und stark eingedriickt, dieser Eindruck

reicht weiter auf der Scheibe der Fliigeldecken nach hinten und die Spuren

dieser sind noch weit hinter der Mitte erkennbar. Schultern breit abgerundet,

stark vortretend. Nahtwinkel der Fliigeldecken am Ende abgerundet. Die

Skulptur der Fliigeldecken besteht vorne aus ziemlich grossen und spdarlichen

Punkten, nach hinten wird die Punktierung immer feiner und auch etwas

raspelartig. Die Behaarung dicht, lang und schrag nach hinten gerichtet, ohne

anliegende Haare. Fiihler beim Mannchen lang und fadenférmig, die beiden

62 ANNALS OF THE SOUTH AFRICAN MUSEUM

ersten Glieder dicht, lang behaart, glanzend, die iitbrigen Glieder matt und mit

ausserst feinen, kurzen, seidenartigen Haaren bedeckt. Die Glieder 3-10 haben

am Ende je einige lange Haare. Das 3. Glied langer als das 4., dann werden die

Glieder stufenweise langer und diinner, das Endglied mehr als 1, 5-mal so lang

wie das 10. und auch viel diinner, etwas gebogen. Bezne einfach, die Tarsen diinn,

Vordertarsen ebenso lang wie die Schienen, die einzelnen Glieder lang, Hinter-

schienen mit gleichgrossen, kurzen, am Ende schrag abgestutzten Enddornen.

Hinterschienen am Ende mit rotlichen Haaren. Samtliche Schenkel unten sehr

lang behaart, Schienen, besonders die Vorder- und Mittelschienen anliegend

behaart. Unterseite ebenfalls lang, schwarz, abstehend behaart, sehr fein und

dicht etwas raspelartig punktiert, das 5. freiliegende Abdominalsegment unten

am Ende beim Mannchen beiderseits abgerundet, die Mitte kurz ausgeschnitten,

das 6. in der Mitte tief V-formig ausgerandet und beiderseits spitzig. Ldnge:

15mm. Breite: 5 mm.

1 Mannchen aus NW. Cape Province, Nieuwoudtville, Sept. 1936, leg.

C. W. Thorne & K. H. Barnard in der Sammlung des South African Museum

(Monotypus). |

Diese Art ist durch die sehr lange, dichte Behaarung des Korpers, den langen,

schmalen Eindruck auf den Fliigeldecken, die Form und Eindriicke des Hals-

schildes sehr gut ausgezeichnet. Sie ist nur mit den Arten J. hirsuta Thunb.,

I. ursus Thunb. und den neuen Arten J. analis sp. nov., sowie I. semillima sp. nov.

verwandt. Alle diese Arten besitzen aber kiirzere und sparlichere Behaarung,

sowie abweichende Skulptur und Form der Oberseite.

Iselma simillima sp. nov.

Sie steht J. ursus Thunb. sehr nahe, so dass ich verzichte eine ausftihrliche

Beschreibung zu bringen. J. simillima unterscheidet sich von J. ursus ‘Thunb.

durch die Kopfform, der Kopf ist kleiner, flacher, hinten langer und schméler,

am Scheitel abgerundet, Schlafen kurz und nach hinten verschméalert, die

Oberseite grob punktiert, Halsschild langer als breit, Seiten abgerundet, nach

hinten nur wenig verschmalert, Scheibe beiderseits flach gedriickt und dicht

punktiert. Fligeldecken mit Erzschein, die Punktierung dicht und fein, hie und

da in Querrichtung zusammenfliessend. Letztes Abdominalsegment am Ende

fast gerade abgeschnitten, beiderseits mit abgerundet stumpfwinkligen, etwas

mehr behaarten und gewolbten Ecken. Die Behaarung der Oberseite einfarbig

schwarz, die lange Behaarung aufstehend, dazwischen befindet sich kurze,

anliegende, oder schrag abstehende, dichte, ebenfalls schwarze Behaarung.

Lange: 13 mm. Breite: 4, 5 mm.

2 Mannchen aus Cape Province, Giftberg, siidlich von Van Rhynsdorp, 1911,

leg. R. M. Lightfoot (Holo- und Paratypus) in der Sammlung des South African

Museum. Die Paratype wurde fiir die Sammlung des Ungarischen Natur-

wissenschaftlichen Museums in Budapest iiberlassen.

STUDIEN UBER SUDAFRIKANISCHEN MELOIDEN 63.

Iselma analis sp. nov.

Glanzend schwarz, die Fligeldecken mit schwachem Metallglanz, schwarz

behaart. Kopf schmal und lang, Stirn etwas gewolbt, Schlafen nach hinten am

Scheitel breit abgerundet, die Punktierung grob aber sparlich, die Mittellinie

hinten am Scheitel sogar unpunktiert, Oberseite sehr lang abstehend behaart,

ohne anliegende Behaarung. Halsschild lang eiformig, vor der Mitte am breites-

ten, stark gewélbt, nach hinten abgerundet und nur wenig verschmalert, nach

vorne gerade, die Scheibe kaum merkbar verflacht. Oberflache grob und sehr

dicht fast einander beriihrend punktiert, die Punktierung in der Mitte der

Scheibe etwas sparlicher. Die Behaarung sehr lang abstehend, sparlich und

schwarz, dazwischen sind aber kurze, anliegende, verschiedenartig gerichtete,

rotliche Haare vorhanden. Basalrand schmal, aber ziemlich scharf. Fliigeldecken

parallelseitig, grob und dicht, in Querrichtung zusammenfliessend punktiert,

Oberflache neben der Naht etwas flach. Die Innenseite der Schultern nur leicht

eingedriickt, der Eindruck reicht aber nicht nach hinten. Das Ende der Fligel-

decken einzeln abgerundet. Die Behaarung lang aufstehend, sehr sparlich,

dazwischen mit schrag aufstehenden, kurzen, schwarzen und anliegenden

Haaren. Fiihler ziemlich dick, die Glieder 3—5 zylindrisch, mit geraden Seiten,

aber an der Basis viel schmaler als am Ende, die Glieder 6—7 an der dusseren

Seite merklich konkav, an der inneren Seite aber konvex, so dass das Ende

etwas sdgeartig erscheint, die Glieder 7—11 diinner, langer, das Endglied faden-

formig. Bene einfach, Tarsen sehr lang und diinn, die Enddorne der Hinter-

schienen kurz und gleichlang, am Ende schrag abgestutzt. Unterseite fein und

raspelartig punktiert, das Analsegment beim Mannchen am Ende in der Mitte

etwas ausgeschnitten, seitlich vollkommen abgerundet und die Oberflache am

Ende breit verflacht. Lange: 7-g mm. Breite: 3-3,9 mm.

2 Mannchen und 1 Weibchen aus Cape Province, Giftberg, siidlich von Van

Rhynsdorp, Sept. 1911, leg. R. M. Lightfoot (Holo- und Paratypen) in der

Sammlung des South African Museum; die Paratype wurde fiir die Sammlung

des Ungarischen Naturwissenschaftlichen Museums in Budapest iiberlassen.

Diese Art ist unter den Verwandten von J. ursus ‘Thunb. die kleinste. Sie

unterscheidet sich von JI. ursus ‘Thunb. ausser der Grésse noch durch die Kopf-

form, welche bei J. ursus ‘Thunb. breiter, flacher und am Scheitel abgestutzt

erscheint. Von der nachst verwandten J. simillima sp. nov. unterscheidet sie

sich durch die sekundaren Geschlechtsmerkmale des Mannchens, die abwei-

chende Skulptur der Ober- und Unterseite und durch den viel kleineren

Ko6rper.

Iselma hessei sp. nov.

Gross und breit, schwarz, glanzend. Fliigeldecken von den Schultern an

parallel dem Seitenrand bis zum Ende mit einer sehr breiten, roten Binde. Kopf

langoval, mit langen, etwas parallelen Schlafen, Scheitel hinten halbkreis-

formig breit abgerundet, Stirn ziemlich flach, in der Mitte etwas buckelig, sehr

grob und dicht punktiert, die Mittellinie aber unregelmassig, hie und da

64 ANNALS OF THE SOUTH AFRICAN MUSEUM

unpunktiert. Die Behaarung abstehend und lang, schwarz, ziemlich sparlich,

ohne anliegende Haare. Halsschild langer als breit, etwas vor der Mitte am

breitesten, nach vorne gerade verjiingt, nach hinten ziemlich parallel und nur

vor der Basis eingeschniirt, Basalrand breit und scharf. Oberflache in der Mitte

und beiderseits auf der Scheibe undeutlich verflacht. Die ganze Oberseite

ausserst grob und sparlich mit tiefen, rundlichen Punkten besetzt, die Punk-

tierung an den Seiten feiner und dichter, die Scheibe hie und da unpunktiert,

glatt. Die Behaarung aufstehend, lang und schwarz, dazwischen auch mit

feineren, diimneren, etwas schrag abstehenden schwarzen Haaren. Fliigeldecken

breit parallelseitig, Schultern abgerundet und stark vorstehend, die Innenseite

breit abgeflacht, Oberflache gegen die Naht breit, scheinbar flach. Die Punk-

tierung sehr grob und dicht, in Querrichtung unregelmassig zusammenfliessend,

die breite rote Binde aber nur sehr sparlich und auch feiner punktiert. Die

ganze Punktierung wird gegen das Ende feiner. Aus den Punkten wachsen

ziemlich kurze und anliegende, dunkle Haare, dazwischen nur hie und da einige

lange, schrag abstehende Haare. Fiihler lang, beim Mannchen ziemlich stark

gesagt, die einzelnen Glieder am Ende nach aussen etwas spitzwinklig vorge-

zogen. Beine einfach, ohne besondere Kennzeichen. Hinterschienen mit

kurzen, am Ende schrag abgestutzten Enddornen. Unterseite fein und dicht,

raspelartig punktiert, schwarz behaart; letztes Abdominalsegment am Ende

gerundet ausgeschnitten, seitlich beiderseits mit stumpfen Ecken. Lédnge:

15 mm. Bredie > 5); ram

1 Mannchen aus NW. Cape Province, Namaqualand, Bowesdorp (zwischen

Garies und Springbok), Sept. 1941, leg. Museum Staff (Monotypus) in der

Sammlung des South African Museum.

Diese Art ist wegen ihrer ganz eigenartigen Farbe der Fliigeldecken sofort zu

erkennen und sieht keiner anderen Jselma-Art ahnlich. Morphologisch steht die

neue Art in der Gruppe von JI. flavipennis Haag-R. und JI. namaqua Pér., von

welchen sie aber durch die vollkommen abweichende Farbe der Fliigeldecken,

die sehr grobe Skulptur des Vorderkorpers und durch die Fihlerform des

Mannchens leicht zu unterscheiden ist.

Ich benenne diese interessante neue Art zu Ehren des Herrn Kollegen Dr.

A. J. Hesse im S.A. Museum, Cape Town.

Iselma csikw sp. nov.

Gestreckt, Vorderkérper braunschwarz, Fliigeldecken in der Mitte heller, an

den Seiten und neben der Naht breit dunkler braun, die ganze Ober- und

Unterseite mit anliegenden, sehr dichten, grauweissen Haaren bedeckt, so dass

der Koérper grau erscheint. Kopf langoval, stark gewolbt, Schlafen lang und

parallel, hinten halbkreisf6rmig abgerundet; die Punktierung gleichmassig,

ziemlich fein und dicht, die Zwischenraume zwischen den Punkten etwa so

gross wie die Punkte selbst. Die Behaarung kurz, hinten nach vorne und vorne

nach innen gerichtet. Halsschild viel langer als breit, in der Mitte am breitesten,

STUDIEN UBER SUDAFRIKANISCHEN MELOIDEN e 65

nach vorne starker und fast gerade verjiingt, nach hinten ziemlich parallelseitig

und nur vor dem sehr scharfen und dicken Basalrand ausgeschweift. Die Mitte

nur leicht und die Scheibe beiderseits kaum merkbar verflacht. Die Skulptur

fein aber sehr dicht, die Behaarung anliegend und verschiedenartig gerichtet.

Fltigeldecken lang parallelseitig, Schultern abgerundet etwas vortretend, die

Skulptur besteht aus dusserst dicht stehenden, raspelartigen, in der Quere stel-

lenweise zusammenfliessenden Punkten. Die Behaarung sehr dicht und anlie-

gend, schrag nach aussen gerichtet. Fihler fadenformig, die Glieder ziemlich

gleichformig, am Ende kaum dinner, das 3. Glied das langste, doppelt so lang

wie das 4. und auch langer als das Endglied. Die Glieder von 4. an werden

stufenweise kiirzer, so dass das 10. merklich kiirzer ist als das 4. Bezne einfach,

Tarsen mit langen Gliedern, Schienen am Ende mit langen goldgelben Haaren,

Tarsen unten ebenfalls goldgelb behaart. Die Enddorne der Hinterschienen

kurz und dick, ihr Ende breit und schrag abgestutzt, der innere Enddorn etwas

langer. Unterseite sehr dicht mit raspelartigen Punkten besetzt, vollkommen

anliegend grauweiss behaart. Lange: g mm. Breite: 3 mm.

1 Weibchen aus Ost-Afrika, Tanagebiet, Witu (Monotypus) in der Samm-

lung des Ungarischen Naturwissenschaftlichen Museums in Budapest.

Diese Art ist durch mehrere Eigenschaften sehr charakteristisch. Sie besitzt

das nordlichste Vorkommen in der Gattung und morphologisch steht sie in der

Nahe von I. quadrimaculata Borchm., doch steht sie weit von den bekannten

Arten. Eine ahnliche Farbung und Behaarung des Korpers, sowie eine ahnliche

Fihlerform kommt bei den ubrigen Arten nicht vor.

Ich erlaube mir diese neue Art Herrn Direktor E. Csiki, dem verdienstvollen

ungarischen Coleopterologen zu widmen.

Im folgenden gebe ich eine Bestimmungstabelle der bisher bekannten Arten

der Gattung Iselma Haag-R.

1 (40) Die Enddorne der Hinterschiener viel kiirzer als das 1. Glied der Hintertarsen.

2(15) Der ganze Korper schwarz, selten mit metallischem Schimmer.

3 (12) Der ganze Korper tiefschwarz, héchstens die Fliigeldecken mit schwachem Metall-

schimmer.

4 (5) Fliigeldecken am Ende zugespitzt, schwarz behaart, Kopf und Halsschild eingedriickt

(nach Haag-Rutenberg). 1. hirsuta Thunb.

5 (4) Fliigeldecken am Ende abgerundet.

6 (7) Ober- und Unterseite mit ausserst langen und dichten, vollkommen schwarzen Haaren

besetzt, die Behaarung ist doppelt, d.h. es sind langere, mehr aufstehende und

dazwischen kiirzere aber kaum anliegendere Haare vorhanden. Fliigeldecken an

der Basis neben den Schultern stark eingedriickt, der Eindruck reicht schrag weit

nach hinten. Halsschild in der Mitte und beiderseits neben der Mitte stark

eingedriickt. L.: 15 mm. NW. Cape Province (Nieuwoudtville).

. 2. lanuginosa sp. nov.

7 (6) Die Behaarung, besonders die anliegenderen Haare viel kiirzer, die langeren Haare

viel sparlicher, der Eindruck nur an der Schulter deutlich, nicht nach hinten

verlangert, die Eindriicke des Halsschildes viel schwacher, oder sogar vollkommen

fehlend.

8 (9) Kopf flach und breit, hinten abgestutzt, Schlafen mit gerundet rechtwinkligen Beulen,

ohne Mandibeln scheinbar quadratisch. Halsschild beiderseits neben der Mitte

sehr schwach eingedriickt, grob und dicht punktiert. Fliigeldecken sehr dicht,

9 (8)

10 (11)

II (10)

12 (3)

13 (14)

14 (13)

15 (2)

16 (23)

17 (18)

148) (7)

19 (20)

O (19)

21 (22)

ANNALS OF THE SOUTH AFRICAN MUSEUM

vorne etwas zusammenfliessend punktiert, die anliegende Behaarung dunkelbraun

und neben der Naht etwas seidenglanzend. L.: 9-12 mm. Cape Province.

-» . 3. Ursus Thunb:

Kopf langlich, Stirn mehr gewolbt, hinten vollkommen breit abgerundet, ohne

Mandibeln langer als breit. Fliigeldecken mit deutlicherem Erzschein, dichter

behaart, auch hinten mit in der Querrichtung zusammenfliessender Punktierung.

Grosser; letztes Abdominalsegment beim Mannchen in der Mitte etwas ausgeschnit-

ten und die Seiten beiderseits etwas eckig aufgebogen. 3. Glied der Fiihler beim

Mannchen breiter und auch etwas langer als das 4. Kopf breiter und flacher,

hinten weniger abgerundet. L.: 13 mm. Cape Province (Giftberg, Van Rhyns-

dorp). . . . 4. simillima sp. nov.

Kleiner; letztes Abdominalsegment beim Mannchen abgestutzt, Oberflache breit

verflacht, Seiten beiderseits ohne Ecke. Das 3. Fihlerglied beim Mannchen

schmaler und etwas ktirzer als das 4. Kopf schmaler und langer, hinten breiter

abgerundet, Stirn gewolbter. L.: 7-9 mm. Cape Province (Giftberg, Van Rhyns-

dorp). . . . 5. analis sp. nov.

Fliigeldecken mit ausgesprochenem Metallschimmer, die Unterseite, besonders aber

das Abdomen ebenfalls metallisch. Kopf und Halsschild, sowie die Brust meist auch

mit metallischem Schimmer.

Halsschild langer als breit, mit scharfem und dickem Basalrand, beiderseits neben der

Mitte der Scheibe mit je einem flachen, langlichen Eindruck; Kopf ziemlich gross,

viel breiter als die Halfte des Halsschildes. Die anliegende Behaarung des Korpers

braungelb, die aufstehenden Haare besonders am Vorderkorper sehr lang und

schwarz, die Fliigeldecken mit Bronzeglanz. Oberflache der Fliigeldecken schwach

eingeflacht. L.: 10-11, 5 mm. Cape Province (Clanwilliam).

. 6. planidorsis Pér.

Halsschild kugelig, nach allen Richtungen gewolbt, so breit wie lang, die Basis nur

schwach gerandet, Scheibe in der Mitte und beiderseits neben der Mitte ein-

gedriickt. Kopf sehr klein und flach, nur halb so breit wie der Halsschild. Die

anliegende Behaarung dunkelbraun, die aufstehende schwarz. Fligeldecken

blaugriin, Unterseite blaulich, Kopf und Halsschild mit griinlich-kupferigem

Schimmer. L.: 7-9, 5 mm. Cape Province (Clanwilliam, Worcester).

. 7. metallescens Per.

Korper schwarz, Fliigeldecken aber einfarbig rotgelb, aelbror braungelb, oder gelbrot

mit schwarzen Flecken, oder schwarz mit gelbroten Makeln.

Fliigeldecken mit gelber, oder rotlicher Zeichnung, oder gelb mit schwarzen Flecken.

Fliigeldecken schwarz, die Seiten von den Schultern bis zum Nahtwinkel aber sehr

breit rot gesAumt. Kopf und Halsschild ausserordentlich grob punktiert, die

Punktierung der Fliigeldecken auch sehr grob und viel dichter als die des Hals-

schildes. Halsschild ohne anliegende Behaarung, nur sehr lang schwarz behaart,

Fliigeldecken mit schwarzer, anliegender Behaarung und nur hie und da mit

einigen langen, aufstehenden Haaren. Letztes Abdominalsegment des Mannchens

ausgeschnitten. L.: 15 mm. N.W. Cape Province Ne ae Bowesdorp.)

. 8. hessei sp. nov.

Fliigeldecken gelbrot mit schwarzen Flecken, oder schwarz mit seibed Makeln.

Fliigeldecken gelbrot mit dunkler, breiter Naht, welche sich nach hinten erweitert,

oder die dunkle Farbung erweitert sich oft so stark, dass nur die Schultern und je

eine lange Makel am Seitenrand hinter den Schultern gelb bleibt. Halsschild sehr

lang, fast zylindrisch gewolbt, sehr sparlich und ziemlich fein punktiert. Die

anliegende Behaarung der Fliigeldecken gelbrot. L.: 6-11 mm. S.W. Afrika

(Abachaus, 48 km. NW. von Otjiwarongo). . . . g. hobohmi Kasz.

Fliigeldecken gelbrot mit schwarzen Flecken, die Naht héchstens am Ende dunkel.

Beine und Fiihler dunkel, héchstens die Hinterschienen gelbrot. Fliigeldecken gelbrot,

mit je 2 grossen, der Lange nach miteinander verbundenen schwarzbraunen

Flecken; nur die Basis, das Ende, die schmale Naht und der Seitenrand, sowie die

Mitte an den Seiten zwischen beiden Flecken gelb. Halsschild sehr dicht und

ziemlich fein punktiert, die Mitte stark eingedriickt. Kopf breit und gewolbt. Die

22 (21)

23 (16)

24 (25)

25 (24)

26 (35)

27 (28)

28 (27)

29 (34)

30 (33)

31 (32)

32 (31)

33 (30)

34 (29)

35 (26)

36 (37)

37 (36)

38 (39)

STUDIEN UBER SUDAFRIKANISCHEN MELOIDEN 67

Oberflache hellgelb dicht behaart. Fiihler des Mannchens am Ende des 4. und 5.

Gliedes dick. L.: 12 mm. Ost-Afrika (Lukuledi).

. 10. qguadrimaculata Borchm.

Schienen und Tarsen, sowie die Fihler gelbrot. Fliigeldecken gelbrot, am Ende aber

mit je einem grossen, ovalen, schwarzen Fleck, so dass die Naht im hinteren

Finftel auch schwarz ist. Kopf viel schmaler, Halsschild viel sparlicher punktiert,

Fliigeldecken hellgelb, Vorderkorper braun und Unterseite ganz anliegend gelbgrau

behaart. L.: 12 mm. Ost-Afrika (Barikiva). . . . It. ttbialis Borchm.

Fliigeldecken einfarbig gelbrot, rotgelb, hdchstens mit ausserst schmaler dunkler

Naht, oder der ganze Korper einfarbig graubraun.

Der ganze Korper einfarbig graubraun. Die Behaarung sehr dicht und anliegend

grauweiss, Halsschild lang, mit scharfem Basalrand, Kopf schmal und lang, mit

parallelen und langen Schlafen, hinten aber breit abgerundet. Das 3. Fiihlerglied

sehr langgestreckt, fast doppelt so lang wie das 4. L.: 9g mm. Ost-Afrika (Tana-

gebiet, Witu). Neil ie L2H CSUKI SD. MON:

Korper schwarz, die Fliigeldecken aber gelbrot, oder rotgelb.

Fliigeldecken nur mit einfarbiger, schwarzer Behaarung. Meist grosse Arten.

Fliigeldecken mit dunkler Naht. Fihler stark und lang, etwas gesagt, Halsschild oval,

dicht und grob punktiert. Die schwarze Behaarung der Fliigeldecken kurz. L.:

11-14 mm. (nach Haag-Rutenberg und Péringuey). Cape Province, Cape Division.

. . . 13. rufipennis Haag-R.

Fligeldecken einfarbig gelb oder rot, die Naht nicht dunkel.

Grosser, 11-15 mm.

Fliigeldecken gelb, Halsschild breit, stark gewolbt, kaum langer als die grésste Breite.

Die Mitte, sowie die Scheibe beiderseits neben der Mitte des Halsschildes flach

eingedriickt. Oberflache des Halsschildes sehr grob punktiert. L.: 13-14 mm. Cap

der guten Hoffnung. . . 14. flavipennis Haag-R.

Halsschild ausserst grob punktiert, die Mittellinie der Scheie unpunktiert und etwas

aufgewolbt, die Scheibe beiderseits hie und da mit einigen (oder nur mit je einem)

Spiegelfleck. Fliigeldecken starker punktiert und an den Seiten lederartig gerunzelt.

L.: 14-16 mm. NW. Cape Province (Namaqualand). 15. namaqua Pér.

Fligeldecken rot. Halsschild schmaler und langer, sehr grob aber sparlich punktiert,

Kopf ebenfalls schmal und sehr lang, Wangen parallel, Kopf hinten halbkreis-

formig abgerundet. Fliigeldecken grob punktiert und die Punkte meist in Quer-

richtung zusammenfliessend. L.: 11-15 mm. N.W. Cape Province (Namaqua-

land). . . . 16. brunneipes Haag-R.

Kleiner, 9 mm. nicht tiberragend. Halsschild ziemlich rundlich, stark punktiert,

dazwischen fein gerunzelt, die Mitte und beiderseits die Scheibe schwach verflacht.

Kopf ziemlich breit und kurz, Schlafen hinten sich plotzlich verschmalernd.

Fliigeldecken sehr dicht und grob punktiert, zusammenfliessend. L.: 7-9 mm.

Cape Province (Clanwilliam). . . . 17. boschimana Pér.

Fliigeldecken hell behaart, oder die Haare am Grunde schwarz, am Ende aber

gelb, so dass die Behaarung von oben gesehen hell erscheint, oder zwischen den

gelben Haaren sind auch schwarze Haare untermischt.

Fuhler stark gesagt. Oberseite doppelt behaart, die lange, aufstehende Behaarung ist

schwarz, die kiirzere einfarbig gelbrot, die Basis der Haare auch gelbrot; Kopf und

Halsschild mit dichter und anliegender, gelber Behaarung. Halsschild langlich und

die Scheibe beiderseits flach eingedriickt. Die Punktierung der Fliigeldecken

ziemlich sparlich. L.: 10-12 mm. Cape Province (Grahamstown, Willowmore).

. 18. optata Pér.

Fihler einfach, fadenformig, nicht gesagt, Halsschild kiirzer, meist rundlich. Die

kurzere Behaarung der Fliigeldecken am Grunde schwarz, am Ende aber gelb, so

dass die Behaarung von oben gesehen einfarbig gelb erscheint.

Halsschild in der Mitte der Scheibe und auch beiderseits eingedriickt. Kopf

verhaltnissmassig klein und langlich, die Punktierung der Fliigeldecken sehr dicht.

Grosser, die Behaarung mehr braunlich. L.: 8-g mm. Cape Province (Cape Town,

Van Rhynsdorp). . . . 1g. rubripennis Lap.

68 ANNALS OF THE SOUTH AFRICAN MUSEUM

39 (38) Halsschild scheibenformig, die Mitte und die Scheibe beiderseits meist ebenfalls flach,

der Kopf viel kiirzer, rundlich und breiter. Die Punktierung der Fliigeldecken etwas

feiner und dichter. Die Behaarung mehr gelblich. Kleinere Art. L.: 6-8 mm.

N.W. Cape Province (Namaqualand). . . . 20. pallidipennis Haag-R.

40 (1) Der innere Enddorn der Hinterschienen etwas langer als das lange 1. Hintertarsen-

glied. Der aussere Enddorn kaum halb so lang als der innere. K6rper schwarz,

Flugeldecken gelb.

41 (42) Halsschild langlich oval, die Mittellinie erhaben und lang, Kopf mit anliegender

gelber Behaarung. L.: 7 mm. (nach Péringuey). NW. Cape Province (Namaqua-

land). - » . 21. okiepana Pér.

42 (41) Halsschild rundlich, kugelig, die Mittellinie nur in der Mitte etwas erhaben, oder

vollkommen flach. Kopf ohne gelbe Behaarung. Fliigeldecken blassgelb, gelb

behaart, hie und da mit einigen schwarzen, aufrecht stehenden Haaren. L.: 6-8

mm. N.W. Cape Province (Bushmanland). . . . 22. longispina Kasz.

Gen. <onitomorpha Per.

1909. Péringuey, Trans. Roy. Soc. S. Afr., 1, p. 272.

Konitomorpha costata sp. nov.

Einfarbig gelbrot, die Fiihler von 3. Glied an schwarz, die Basis des 3. Gliedes

aber noch gelbrot, Fliigeldecken am Ende bis zum hinteren Drittel schwarz,

vorne hinter dem Schildchen mit einer breiten, gemeinsamen Binde, welche

die Basis mit den Schultern und die Seiten frei lasst. Beine und Unterseite

gelbrot, die Tarsen aber zweifarbig, d. h. die Glieder 2-4 der Vorder- und

Mitteltarsen, sowie die Basis des Krallengliedes schwarz, an den Hintertarsen

aber das 2., 3. und die Basis des 4. Gliedes schwarz. Kopf oval, ohne Mandibeln

langer als breit, Stirn zwischen den Augen schmal und flach, etwa nur so breit

wie der Querdurchmesser eines Auges an der schmalsten Stelle, Clypeus eben-

falls flach, Schlafen lang und ziemlich parallel, hinten schrag abgestutzt und

in der Mitte gerade, Oberflache hinter den Augen breit verflacht, die Mitte

kaum buckelig. Die Punktierung vorne dicht und fein, hinter den Augen und

am Scheitel sehr sparlich, aber etwas gréber. Halsschild sehr lang und trapez-

formig, vorne so breit wie der Kopf, die Vorderecken breit abgerundet, nach

hinten fast gerade erweitert, etwa 14-mal so lang wie an der Basis breit; in der

Quere stark gewoélbt, weit vor der Mitte beiderseits grubig vertieft, die Basis in

der Mitte etwas verflacht, der Basalrand fein, aber scharf, die Mitte etwas

ausgeschnitten. Die Punktierung ist unregelmassig, die Mittellinie hie und da

unpunktiert, die Scheibe beiderseits dicht und etwas feiner punktiert, im iibrigen

ist die Punktierung sehr sparlich. Fliigeldecken lang und parallel, hinter der

Mitte nur leicht gewélbt, Seitenrand von oben tiberall gut sichtbar. Schultern

durch einen flachen Eindruck abgegrenzt, breit verrundet. Oberflache mit

Spuren von Langsnerven, von welchen der innere und dussere weit hinter der

Mitte, der mittlere aber nur vor dem Ende erloschen ist. Die Punktierung ist

fein und sparlich, gegen das Ende noch feiner und vor dem Ende sogar mikro-

skopisch, etwas lederartig. Fiihler die Basis des Halsschildes mit 3 Gliedern

iiberragend, die Glieder sind gleichdick, das 1. Glied lang und dick, das 2.

langer als breit, das 3. breiter, die Glieder vom 3. an fast gleichlang und gleich-

breit, etwa doppelt so lang wie breit. Beime lang und schlank, das 1. Glied

STUDIEN UBER SUDAFRIKANISCHEN MELOIDEN 69

samtlicher Tarsen langer als die beiden folgenden Glieder zusammen. Die

beiden Enddorne der Hinterschienen sind sehr kurz und dick, zugespitzt.

Krallen sehr stark gekammt. Unterseite A4usserst fein raspelartig punktiert und

anliegend kurz, gelb behaart. Lange: 14 mm. Breite: 4, 6 mm.

1 Weibchen aus S.W. Afrika, Kaross (ungefahr 100 km. SO. von Zesfontein,

Kaokoveld), Febr. 1925, leg. Mus. Exped. (Monotypus) in der Sammlung des

South African Museum.

Diese Art steht der <. seminigra Pic noch am nachsten, ist aber von ihr durch

die vollkommen abweichende Skulptur der Decken, sowie die verschiedene

Farbung des Korpers leicht zu unterscheiden.

Konitomorpha sellata Fahraeus

Diese Art hat in Siid-Rhodesien, ‘Transvaal und Natal eine weite Verbreitung;

sie ist eine der haufigsten <onitomorpha-Arten der Gegend. Die Untersuchung

eines groésseren Materials ergab, dass die nomenklatorische forma typica eine

sehr seltene Form sein muss, da ich in dem von mir bestimmten, relativ reichen

Material kein einziges Stiick gefunden habe, welches ich mit der Beschreibung

von Fahraeus vollkommen identifizieren konnte. Die Art ist sehr veranderlich

und ich fand eine ganze Reihe bisher unbekannter Formen, welche ich hier im

Rahmen einer Bestimmungstabelle publiziere.

1 (22) Fliigeldecken gelbrot mit schwarzer Binde oder mit schwarzer Zeichnung oder mit

schwarzem Ende.

2 (5) Das Ende der Fliigeldecken schwarz, der schwarze Fleck erreicht oft die vordere

Halfte der Fliigeldecken.

3 (4) Beine gelb, nur Schenkel und Schienenende, sowie die Tarsen dunkel. 1. Fiihlerglied

bis zur Mitte rotlich. Das Ende der Fliigeldecken bis zum hinteren Drittel schwarz.

13. ab. stevensoni m.

4 (3) Beine dunkel, nur die Schienen gelb, Schienenende auch dunkel. Fiihler schwarz.

Fliigeldecken schwarz, nur das vordere Drittel gelbrot.

. 14. ab. seminigripennis m.

5 (2) Das Ende der Fliigeldecken gelbrot.

6 (13) Basis der Fiihler schwarz.

7 (12) Fliigeldecken mit einer mehr-weniger breiten Querbinde vor der Mitte; die Basis der

Fligeldecken quer gelbrot. Schenkel meist gelbrot.

8 (9) Hinterbrust rot, nur die Pleuren dunkel. Schenkel gelbrot, das Ende aber dunkel.

. g. ab. pectoralis m.

9 (8) Hinterbrust schwarz, selten die Pleuren hell.

10 (11) Schenkel dunkel. . . . 10. ab. nigricornis m.

11 (10) Schenkel rot, nur das Ende dunkel. wep) Wie ab. fanraeusy ml.

m2(7) Fliigeldecken bis zum hinteren Drittel schwarz, nur die Schultern und das Ende

gelbrot. Beine dunkel, nur die Basis der Schienen rot. 12. ab. nigricolor m.

13 (6) Basis der Fiihler gelbrot.

14(15) Fliigeldecken gelbrot, hinter der Mitte mit einem gemeinsamen, kleinen, runden,

schwarzen Fleck an der Naht. Beine gelb, Schenkel und Schienenende, sowie die

Tarsen dunkel. Die beiden ersten Glieder der Fiihler gelb.

: . 5. ab. unipunctata m.

15 (14) Fliigeldecken mit schwarzem Ende oder mit einer schwarzen vorderen Querbinde,

oder mit 2 kleinen Flecken an den Seiten.

16 (21) -Fliigeldecken mit einer schwarzen Querbinde vor der Mitte.

17(18) Nur das 1. Fihlerglied rot. Schenkel dunkel. Basis der Fliigeldecken rot.

. . « 6. forma typica Fahr.

7O ANNALS OF THE SOUTH AFRICAN MUSEUM

18 (17) Fuhler mit 2 gelben Basalgliedern, Schenkel rotgelb, nur das Ende dunkel.

19 (20) _ Fltigeldecken schwarz, nur die Schultern und hinter der Mitte gelbrot. Brust schwarz.

. 7. ab. rhodesiana m.

20 (19) Fliigeldecken mit einer breiten schwarzen Querbinde vorne, die die Basis frei lasst.

Brust dunkelbraun, die Pleuren aber gelbrot. . . . 8. ab. testaceicornis m.

21 (16) Flugeldecken vorne beiderseits neben der Mitte mit je einem dunklen Fleck.

. . 4. ab. brunneonotata Pic

22 (1) Fligeldecken einfarbig gelbrot.

23 (24) Basis der Fihler schwarz. Beine gelbrot, Schienen- und Schenkelende, sowie die

Tarsen aber dunkel. Brust schwarz. . . . 3. ab. bang-haasi Pic

24. (23) Fiuhlerbasis gelbrot.

25 (26) Die beiden ersten Glieder der Fiihler gelbrot. Beine gelb, Tarsen braun, Brust

schwarz. . . . 2. ab. testaceipes Pic

26 (25) Die 3 ersten Glieder der Fiihler gelbrot. Beine gelb, Tarsen braun und die Brust gelb.

. I. ab. unicolor m.

1. sellata ab. unicolor ab. nov.: Die ersten 3 Glieder der Fiihler gelb. Ober- und Unterseite,

sowie die Beine gelbrot, Tarsen aber dunkel. Siid-Rhodesia, Gwanda, 1906, leg. D. Dods

(Monotypus, Coll. S.A. Mus. Cape Town).

2. sellata ab. testaceipes Pic: Die ersten zwei Fihlerglieder rot, Beine gelb, Tarsen braun,

Fligeldecken ohne Binde, Abdomen rot, Brust schwarz.

3. sellata ab. bang-haasi Pic: Fiihler schwarz, Schenkelbasis und Fliigeldecken rotlich,

Abdomen rot, Brust schwarz.

4. sellata ab. brunneonotata Pic: Fihlerbasis rot, Beine rotlich, Fliigeldecken mit je einer

kleinen Makel an den Seiten (nach Pic).

5. sellata ab. unipunctata ab. nov.: Die beiden Basalglieder der Fihler gelb, Schenkel und

Schienen rot mit dunklen Enden, Tarsen braun, Brust dunkel, in der Mitte braun, Fliigeldecken

gelbrot, hinter der Mitte mit einem gemeinsamen runden Fleck an der Naht. Siid-Rhodesien,

Sawmills, Febr. 1923, leg. R. H. R. Stevenson (Monotypus, Coll. Mus. Budapest).

6. sellata forma typica: Das 1. Fihlerglied rotlich, Schenkel, Schienenende und Tarsen

schwarz, Fliigeldecken vor der Mitte mit einer schwarzen Querbinde, Abdomen rot, Brust

schwarz (nach Fahraeus, aus der Original-beschreibung).

7. sellata ab. rhodesiana ab. nov.: Die beiden ersten Glieder der Filer rot, Fliigeldecken

schwarz, nur die Schultern und hinter der Mitte rot, Brust schwarz, Beine wie ab. Fahraeusi

m., Schenkel rot, héchstens das Ende schwarzlich, Schienen rot, Tarsen braun. S. Rhodesia,

Bulawayo, leg. G. Arnold (Monotypus, Coll. S.A. Mus. Cape Town).

8. sellata ab. testaceicornis ab. nov.: Die beiden Basalglieder der Fiihler gelb, Beine rot,

Schenkel- und Schienenende, sowie die Tarsen dunkel, Hinterbrust schwarz, die Pleuren aber

gelbrot, Fliigeldecken gelbrot mit einer breiten schwarzen vorderen Querbinde. Siid-Rhodesien,

Gwanda, 1906, leg. D. Dods (Monotypus, Coll. S.A. Mus. Cape Town).

g. sellata ab. pectoralis ab. nov.: Fiihler schwarz, Schenkel rétlich, Hinterbrust schwarz, nur

die Pleuren dunkel, Fliigeldecken mit schwarzer Querbinde. Siid-Rhodesien, Gwanda, 1906,

leg. D. Dods (Monotypus, Coll. S.A. Mus., Cape Town).

10. sellata ab. nigricornis ab nov.: Fiihler schwarz, Schenkel, Schienenende und Tarsen

schwarz, Fliigeldecken vor der Mitte mit einer Querbinde, Abdomen rot, Brust schwarz. Natal,

Durban, 1891, leg. N. Barker (Monotypus, Coll. S.A. Mus., Cape Town).

11. sellata ab. fahraeusi ab. nov.: Fiihler schwarz, Schenkel rot, héchstens das Ende schwarz-

lich, Hinterbrust schwarz, Fliigeldecken wie bei der Stammform, mit einer schwarzen Querbinde.

Mfongosi, Zululand, leg. W. E. Jones (Holotypus, Coll. S.A. Mus., Cape Town); Natal,

Durban, 1891, leg. N. Barker (2 Paratypen Coll. S.A. Mus., Cape Town, 1 Paratype Coll.

Transv. Mus., Pretoria); Siid-Rhodesia, Gwanda, 1906, leg. D. Dods (1 Paratypus Coll. S.A.

Mus., Cape Town); Rhodesia, Bulawayo, leg. G. Arnold (1 Paratypus Coll. Transv. Mus.,

Pretoria); Transvaal (2 Paratypen Coll. Mus. Budapest).

12. sellata ab. nigricolor ab. nov.: Fihler schwarz, Beine auch schwarz, nur die Basis der

Schienen rétlich, Brust schwarz, Fliigeldecken schwarz, nur die Schultern und hinter der Mitte

rotlich. Siid-Rhodesien, Penkridge, 7 Dec. 1927, leg. R. H. R. Stevenson (Holotypus, Coll.

S.A. Mus., Cape Town); S.W. Afrika, Ovamboland, Onolonga, Febr. 1923, leg. Mus. Exped.

(1 Paratypus Coll. S.A. Mus., Cape Town).

STUDIEN UBER SUDAFRIKANISCHEN MELOIDEN 71

13. sellata ab. stevensoni ab. nov.: Das 1. Glied der Fiihler bis zur Mitte rot, Fliigeldecken

gelbrot, das Ende aber bis zum hinteren Drittel schwarz, Brust schwarz, Beine rot, das Ende der

Schenkel und Schienen, sowie die Tarsen braun. Siid-Rhodesien, Penkridge, 15 XII 1927, leg.

R. H. R. Stevenson (Monotypus, Coll. S.A. Mus., Cape Town).

14. sellata ab. seminigripennis ab. nov.: Fiihler schwarz, Schenkel, Schienenende und Tarsen

schwarz, Fliigeldecken bis zum vorderen Drittel breit schwarz, vorne gelbrot, Brust schwarz.

Zululand, Mfongosi, leg. W. E. Jones (Monotypus, Coll. S.A. Mus., Cape Town).

Gen. Sitaris Latr.

Tao2) natreille pists Orust Insel. p. TO]:

Sitaris bushmanica sp. nov.

Kleine, breite, robuste Art. Kopf samt den Fiihlern und Palpen, die Mittel-

und Hinterbrust, sowie die Schenkel samtlicher Beine schwarzbraun, Hals-

schild, Schienen, Tarsen und Abdomen gelbrot, die Seiten der Segmente aber

mit je einem dunklen Fleck, Fliigeldecken gelbrot, in der Mitte aber mit einer

queren Makel, sowie das Ende breit schwarz, Schildchen und das Tergit der

Hinterbrust schwarz. Aopf breit, an den kurzen und nach hinten erweiterten

Schlafen am breitesten, diese seitlich abgerundet und hinten beiderseits in der

Mitte auch etwas leicht ausgebuchtet, so dass sie doppelt leicht gebuckelt

erscheinen. Augen flach-gewolbt, nierenformig. Stirn zwischen den Augen sehr

breit und flach, so breit wie die ersten 4 Fiihlerglieder zusammen lang, Clypeus

mit einem queren, ziemlich scharfen Kiel. Oberseite dicht und fein punktiert,

die Punkte sind zwischen den Augen und neben den Augen stark zusammen-

gedriickt, die Buckel aber hinten kaum punktiert. Sehr fein, dunkel behaart.

Halsschild breit, Seiten ziemlich parallel, vorne an den breit abgerundeten

Vorderecken am breitesten, nach hinten kaum verjiingt, Vorderrand in der

Mitte etwas flach ausgebuchtet, Hinterrand in breitem Bogen verrundet.

Scheibe beiderseits neben der Mittellinie etwas vor der Mitte sehr flach ein-

gedriickt. Oberflache dusserst sparlich, aber gréber punktiert als der Kopf.

Schildchen sehr gross und breit, glanzend, die Mitte breit unpunktiert, das Ende

abgestutzt. Filiigeldecken an der Basis breit und nach hinten verschmalert, innen

auch am Schildchen einander nicht beriihrend, die Innenseite nur sehr leicht

ausgeschnitten, aber sehr breit klaffend, aussen ebenfalls nur wenig, aber doch

starker ausgerandet, das letzte Drittel der Fliigeldecken ziemlich schmal und

parallelseitig und nur halb so breit wie vorne an der breitesten Stelle, das Ende

ziemlich scharf zugespitzt. Die Skulptur ist sehr fein lederartig gerunzelt.

Fiihler kurz und etwas flach, die Basis des Halsschildes kaum iiberragend; die

ersten zwei Glieder glanzend, die itibrigen matt. Das 2. Glied breiter als lang,

das 3. nicht genau 1, 5-mal so lang wie breit, das 4. Glied kaum etwas langer,

die darauf folgenden aber stufenweise kiirzer und auch etwas schmaler, so dass

das 10. Glied so lang wie das 3., aber fast doppelt so lang wie breit ist. Beine

diinn und ziemlich kurz, Vorder- und Mitteltarsen sehr diinn, deutlich langer

als die Schienen, das Klauenglied viel langer als das dickere 1. Glied, Hinter-

schienen im Querschnitt flach, mit ganz kurzen und dicken, abgestutzten

Enddornen. Krallen sehr fein gekammt. Unterseite glanzend, Hinterbrust an

72 ANNALS OF THE SOUTH AFRICAN MUSEUM

den Seiten sehr dicht, fast zusammenfliessend, die Mitte dagegen viel feiner und

sehr sparlich punktiert. Ldnge: 7 mm. (bis zum Ende der Fliigeldecken).

- Breite: 3, 9 mm. (an den Schultern).

1 Exemplar aus N.W. Cape Province, Bushmanland, Jackals Water (zwischen

Steinkopf und Orange Fluss), R. M. Lightfoot, Oct. 1911 (Monotypus) in der

Sammlung des South African Museum.

Sie ist von den bekannten Arten der Gattung Sztaris aus Siid-Afrika, uzw.

S. capensis Pér., S. notaticollis Pér. und S. fitzsimonst Kasz. weit entfernt. Alle diese

Arten besitzen viel langere und deutlicher ausgeschweifte Fligeldecken, ausser-

dem eine vollkommen abweichende Farbung und Skulptur. Sie sieht einer

Stenoria Muls. sehr ahnlich, doch besitzen die Stenoria-Arten nicht so stark ein-

geschniirte Fliigeldecken und unter den afrikanischen Arten gibt es keine ahnlich

gefarbte und skulptierte Form.

Gen. Stenoria Muls. & Rey

1857. Mulsant & Rey, Ann. Soc. Linn. Lyon., iv, p. 394.

Stenoria discomaculata sp. nov.

Kopf schwarz, der Scheitel hinten in der Mitte und der Hals aber oben

rotlich, Fiihler und Palpen braun, die Mundteile aber unten hell, Halsschild

rotgelb, vor der Mitte beiderseits mit je einer grossen, runden Makel auf der

Scheibe, Schildchen, Mittel- und Hinterbrust, die ersten 3 Abdominalsegmente,

sowie das 4. an den Seiten schwarz, die beiden letzten Segmente gelbrot, Beine

schwarz. Fligeldecken braun mit je zwei sehr grossen, queren, gelbroten

Makeln, uzw. eine hinter den Schultern und eine hinter der Mitte, beide Flecke

erreichen den Seitenrand, die Naht bleibt aber schmal dunkel. Kopf breit und

flach, Schlafen hinter den Augen kurz und erweitert, breit verrundet, Scheitel

in der Mitte gebuckelt und die innere Seite durch einen langlichen Eindruck

vom Hinterrand der Augen nach hinten begrenzt, Stirn zwischen den Augen

vorne in der Mitte etwas grubig eingedriickt. Clypeus auch eingedriickt. Die

Punktierung dusserst dicht, fast zusammenfliessend, Scheitel hinten sparlicher.

Halsschild breit scheibenf6rmig, im vorderen Drittel am breitesten, nach vorne

in sehr breitem und starkem Bogen verjiingt, nach hinten in leichtem Bogen stark

verschmalert, die hintere Ecke fast abgerundet, stumpfwinklig, Hinterrand

kaum gerundet, Vorderrand in der Mitte kurz und ziemlich plotzlich aus-

geschnitten. Oberflache etwas vor der Mitte beiderseits schwach verflacht. Die

Punktierung dusserst sparlich und viel gréber als die des Kopfes. Fltigeldecken

lang, am Ende gleichmdssig verschmalert, hinten deutlich schmAler als an der

Basis, Seiten- und Innenrand gerade, Seitenrand nur im hinteren Viertel etwas

ausgebuchtet, fein gerandet, Innenrand aber breit klaffend, das Ende abgerun-

det spitzwinklig. Oberflache ohne Punktierung, lederartig sehr fein gerunzelt.

Fiihler diinn und etwas abgeflacht, die Basis des Halsschildes weit iberragend,

die einzelnen Glieder gleichbreit. Das 2. Glied etwas langer als breit, das 3.

doppelt so lang wie das 2. und etwas mehr als zweimal so lang wie breit, die

STUDIEN UBER SUDAFRIKANISCHEN MELODIEN 73

Glieder vom 4. an kaum merklich etwas kiirzer, das Endglied aber wieder lang,

mehr als 1, 5-mal so lang wie das 10. Bene diinn, Schienen ziemlich abgeflacht,

die beiden Enddorne der Hinterschienen sehr klein, gleichformig und spitzig;

Tarsen lang und diinn, viel langer als die Schienen, 1. Glied der Vorder- und

Mitteltarsen so lang, der Hintertarsen deutlich langer als das Klauenglied. Die

Krallen sind sehr fein gekammt. Unterseite anliegend, sparlich hell behaart,

Hinterbrust an den Seiten und die Pleuren sehr dicht, fast runzelig punktiert,

die Mitte dagegen kaum punktiert, glatt, das Abdomen dicht, etwas raspelartig

punktiert. Ldnge: 8,5 mm. Breite: 3 mm.

1 Exemplar aus N.W. Cape Province, Bushmanland, Jackals Water (zwischen

Steinkopf und Orange Fluss), R. M. Lightfoot, Oct. 1911 (Monotypus) in der

Sammlung des South African Museum. |

Eine sehr charakteristische neue Art, welche in der Gattung keine ahnliche

besitzt. Die einzige, bisher aus Ost-Afrika bekannte Art: S. gzbdzcollis Borchm.

besitzt nur am Ende schwarze Fliigeldecken, einfarbig schwarzen Kopf und

Halsschild, vollkommen abweichende Form und Skulptur. Die andere hier

beschriebene neue Art der Gattung: S. hessez sp. nov. steht noch am nachsten,

sie besitzt aber ausser der abweichenden Farbung noch sehr grob skulptierte

Fliigeldecken, viel dickere Fiihler und eine ganz abweichende Kopf- und Hals-

schildform.

Stenoria hessei sp. nov.

Kopf, Fiihler und Beine, sowie die Palpen tiefschwarz, die Unterlippe aber

gelb, Halsschild glanzend rotgelb mit einem breiten, queren schwarzen Fleck,

welcher durch Verschmelzung von zwei Makeln entstanden ist, Fliigeldecken

schwarz, hinter der Mitte mit je einem ziemlich gut begrenzten rundlichen,

gelbweissen Fleck und vor der Mitte mit je einer kleineren, oft schwach begrenz-

ten und verwischten Makel; Abdomen glanzend schwarz, die ersten 4

Segmente schwarz, die Mitte des 4. und die beiden letzten Segmente aber

gelbrot. Kopf sehr breit und gewolbt, Schlafen kurz, mit breit abgerundeten

Ecken, Scheitel hinten in seiner ganzen Breite im Bogen flach gerundet, Stirn

zwischen den Augen vorne beiderseits leicht grubig vertieft, die Mittellinie etwas

aufgebogen, Clypealsutur etwas eingedriickt. Oberflache sehr grob und dicht

punktiert, die Punkte eng aneinander stossend, am Scheitel aber etwas sp4rlich.

Halsschild fast doppelt so breit wie lang, im vorderen Viertel am breitesten, nach

vorne plotzlich und gerade verjiingt, nach hinten aber stark und etwas gebogen

verengt, die Seiten gehen in den Basalrand an den Hinterecken in eine breit

verrundete Ecke ttber. Vorderrand in der Mitte nur leicht und kurz aus-

geschnitten, Hinterrand fast gerade. Oberflache einfach, ziemlich flach, ohne

Findriicke, ausserst grob und sehr sparlich punktiert, zwischen den groben

Punkten aber mit feinerer, dichter Punktierung, die groben Punkte an den Seiten

viel dichter. Die Behaarung sparlich und dunkel. Schildchen breit und glanzend,

dicht und fein punktiert, hinten aber glatt und glanzend. Fligeldecken nicht

punktiert, aber dusserst grob und dicht gerunzelt, die Runzelung ist vollkommen

74 ANNALS OF THE SOUTH AFRICAN MUSEUM

unregelmassig. Die Naht ist gerade, aber klaffend, der Seitenrand im hinteren

Drittel leicht ausgeschnitten, scharf gerandet, das Ende einzeln abgerundet.

Die Behaarung fein und anliegend, sparlich und dunkel. FMihler dick und lang,

fast die Mitte des KGrpers erreichend, die beiden Basalglieder sind kurz, das

2. Glied kugelig, das 3. so lang wie die ersten zwei Glieder zusammen und viel

breiter, etwa doppelt so lang wie breit, flach abgeplattet, das 4. etwas kiirzer und

schmaler, das 5. wieder etwas langer, aber noch schméaler, die folgenden Glieder

etwa gleichlang und gleichbreit, das letzte Glied aber diinn. Bezne diinn, ohne

besondere Kennzeichen, Tarsen ebenfalls sehr diinn und lang, viel langer als

die Schienen, die beiden Enddorne der Hinterschienen kurz und spitzig. Das

1. Glied der Tarsen nur am Hinterbein etwas langer als das Klauenglied. Die

Krallen sind fein kammartig gezahnt. Lange: 6-6,5 mm. Breite: 2-2, 5 mm.

3 Exemplare aus Cape Province, Graafwater, Oct. 1947, leg. C. W. Thorne

(Holo- und Paratypen) in der Sammlung des South African Museum; eine

Paratype wurde fiir die Sammlung des Ungarischen Naturwissenschaftlichen

Museums in Budapest tberlassen. ,

Das Tier wurde aus dem Nest von Osmza globicola St. (Hymenoptera, Apidae)

gezogen. |

Eine sehr interessante und leicht kenntliche Art, welche unter den Arten der

Gattung keine nahere Verwandte besitzt. Sie zeigt nur mit S. discomaculata

sp. nov. eine gewisse morphologische Ubereinstimmung, doch hat S. hessei sp.

nov. vollkommen abweichende Farbung, andere Skulptur der Oberseite,

besonders der Fliigeldecken, usw.

Ich erlaube mir diese hochinteressante neue Art Herrn Dr. A. J. Hesse in

Cape Town zu widmen.

Gen. Lytta Fabr.

1775. Fabricius, Syst. Ent., p. 260.

Uber die mit Lyitta spilotella Pér. verwandten Arten aus Siidafrika.

In diese Gruppe gehoren von beschriebenen Arten L. spilotella Pér., L. elegan-

tula Pér., L. enona Pér. und L. vellicata Er. Jetzt kommen noch 3 weitere neue

Arten dazu, so dass es nicht tiberflissig erscheint fiir die Arten der Gruppe eine

Bestimmungstabelle zu verfertigen, wie folgt:

I (4) Beine, oder wenigstens die Schenkel rotgelb.

2 (3) Schenkel und Schienen gelb. Kopf rot, nur vorne am Clypeus, unten in der Mitte,

dann eine sehr schmale und lange Makel zwischen den Augen auf der Stirn

schwarz, Halsschild rotgelb, die Seiten aber ringsum schmal schwarz, Fliigel-

decken gelbrot mit unregelmassigen, sparlich stehenden, kleinen schwarzen

Flecken und mit 4 schwach erhabenen, heller gefarbten Nerven, Unterseite

schwarz, Hinterbrust aber breit rotgelb, die Trochanteren und Coxen, sowie die

Tarsen und das Ende der Schienen dunkel, Fiihler schwarz, das 1. Glied fast bis

zur Spitze und die Basis des 2. und 3. Gliedes ebenfalls gelbrot. Die Behaarung

anliegend, sehr fein und weissgrau. Halsschild mit scharfer, schmaler Langsmittel-

furche, welche von der Basis nach vorne bis zum vorderen Viertel reicht. L.: 14-18

mm. Transvaal (Waterberg, Zoutpansberg). . . « 2. Spilotella Pér.

3 (2) Nur die Schenkel hell, die Kniee und Schienen, sowie die Tarsen dunkel. Kopf

schwarz, Halsschild gelbrot mit schwarzen Flecken, uzw. eine gréssere Makel in

STUDIEN UBER SUDAFRIKANISCHEN MELOIDEN 73

der Mitte der Scheibe und beiderseits mit je einem kleineren langlichen Fleck.

Flugeldecken gelbrot. L.: 10 mm. (nach Péringuey). Cape Province (Clanwilliam).

: 1. elegantula Pér.

4 (1) Beine schwarz, hdchstens die Schienen gelb, Schenkel immer schwarz. Kopf schwarz,

mit je einem gelbroten Fleck neben der Innenseite der Augen.

5 (6) Schienen gelb. Unterseite, Halsschild, Schildchen, Fiihler und Schenkel schwarz.

Halsschild fast so breit wie der Kopf, mit parallelen Seiten, die Langsmittelfurche

scharf und von der Basis bis zur Mitte reichend. Fliigeldecken gelbrot. mit dunkel-

braunen, unregelmassigen und auch unbehaarten Flecken, das Ende der Fliigel-

decken etwas breiter braun gesdumt. Die Naht, dann ein Langsnerv in der Mitte,

einer an der Innenseite der Schulterbeulen, welcher fast bis zum Ende reicht und

der Seitenrand erhaben aufgewolbt. Kopf sehr fein und sparlich, Halsschild

dichter punktiert, glatt. L.: 14mm. S.W. Afrika (Bethanie).

. . 3. namaqua sp. nov.

6 (5) Schienen, sowie die ganze Unterseite schwarz, Beine und Filer ebenfalls schwarz.

7 (8) Halsschild schwarz, mit je einem grossen, ovalen, gelbroten Fleck etwas vor der Mitte

der Scheibe. Fliigeldecken dunkel, die Naht, der Seitenrand, sowie je drei Langs-

nerven auf der Scheibe gelbrot. Kopf und Halsschild zwischen den Punkten

lederartig gerunzelt und matt. Halsschild etwas langer als breit, schmaler als der

Kopf, die Langsmittelfurche bis zum vorderen Viertel reichend, Fliigeldecken

dicht und fein, fast runzelig punktiert. Die Behaarung anliegend, weissgelb.

L.: 13-19 mm. S.W. Afrika (Onolonga, Nagusib). . . . 4. rubrolineata sp. nov.

8 (7) Halsschild einfarbig schwarz.

g (10) Fliigeldecken gelbrot mit regelmdssigen braunen, unbehaarten Flecken. Die Naht,

der Seitenrand und ein Langsnerv auf der Scheibe an der Innenseite der Schulter-

beulen heller. Halsschild mit je einem Spiegelfleck auf der Scheibe beiderseits

neben der Mitte und auch die Mittellinie unpunktiert, zwischen der groben Punk-

tierung sehr fein und dicht punktiert, dazwischen der Grund aber glanzend.

Fligeldecken auf mattem Grund dicht und etwas erloschen punktiert. L.: 13-14

mm. S.W. Afrika (Kaross, Zesfontein). . . 5. székessyi sp. nov.

10 (g) Fliigeldecken dunkelbraun oder schwarz, nur die Naht, der Seitenrand und die

aufgewolbten Langsnerven gelbrot.

11 (12) Fliigeldecken auf der Scheibe mit je zwei Langsnerven, welche gelbrot sind. Fihler

kurz, die einzelnen Glieder fast konisch, fast so breit wie lang. Kopf dicht und stark

punktiert, Scheitel eingedrtickt, die Punkte sind nebeneinandergeriickt. L.: 15 mm.

(nach Péringuey) ; Cape Province (Uitenhage). . . 6. enona Pér.

12 (11) Fligeldecken nur mit einem dorsalen Langsnerv an der Inenceite der Schulter-

beulen, welcher breit gelbrot ist. Fiihler kraftig und lang, die einzelnen Glieder

sind deutlich langer als breit und ziemlich zylindrisch. Kopf fein und sehr sparlich

punktiert, Scheitel nicht eingedriickt. Halsschild mit breiter Langsmittelfurche.

L.: 13 mm. Angola (Benguella). gees, vellicala Lt.

Lytta namaqua sp. nov.

Langgestreckt. Kopf schwarzbraun, mit je einem grossen, gelbroten Fleck

an der Innenseite der Augen, Halsschild, die ganze Unterseite, Beine, excl.

Schienen schwarzbraun, Schienen gelbrot, Schildchen dunkel, nur das Ende

gelb, Fliigeldecken gelbrot mit einigen sparlich stehenden und unregelmassigen

braunen Flecken, die Schulterbeule und das Ende der Fliigeldecken braun, die

Naht, der Seitenrand, sowie 2 dorsale Langsnerven heller. Kopf gross und rund-

lich, mit stark gewélbten, grossen Augen, Stirn einfach, kaum gewélbt, Scheitel

gerundet, Schlafen etwas parallelseitig, dann in breitem Bogen vollkommen

verrundet. Clypealsutur in der Quere scharf eingeschnitten. Oberflache sehr

sparlich und fein punktiert, der Grund ist ziemlich glatt, glanzend, die Mittel-

linie zwischen den Augen kaum angedeutet. Halsschild sehr breit und quer,

76 ANNALS OF THE SOUTH AFRICAN MUSEUM

kaum etwas schmaler als der Kopf, mit parallelen Seiten und ziemlich flacher

Oberseite, vom vorderen Viertel an stark abgeschniirt, Vorderrand gerade

abgestutzt, Hinterrand mit breit verrundeten Hinterecken und mit ziemlich

gerader Mitte. Die Langsmittelfurche sehr scharf und tief, nach vorne bis zur

Mitte reichend. Scheibe beiderseits vor der Basis bis zur Mitte leicht und lang-

lich eingedriickt. Die Skulptur besteht aus ziemlich feiner und ungleichartiger

Punktierung, dazwischen ist der Grund mikroskopisch punktiert, doch leicht

glanzend. Fligeldecken sehr dicht und fein, runzelig punktiert, gegen das Ende

aber wird die Punktierung immer feiner, so dass das Ende nur dusserst fein

lederartig ist. Der Grund ist zwischen den Punkten fettglanzend. Filer die

Basis des Halsschildes erreichend, das 3. Glied etwas mehr als doppelt so lang

wie breit, die folgenden Glieder sind kirzer und etwa gleichlang und gleichbreit,

das Endglied wieder etwas langer, oval. Beine ohne besondere Kennzeichen, die

Enddorne der Hinterschienen vollkommen gleich gebildet, kurz und am Ende

schrag abgesetzt. Das 1. Glied der Hintertarsen sehr lang, fast so lang wie die

beiden folgenden Glieder zusammen, das Endglied viel kiirzer. Unterseite sehr

fein und sparlich mit raspelartigen Punkten besetzt, anliegend weissgrau

behaart. Ldnge: 14mm. Breite: 4 mm. |

1 Mannchen aus $.W. Afrika, Gr. Namaqualand, Bethanie, 1 Feb. 1885

(Monotypus) in der Sammlung des Ungarischen Naturwissenschaftlichen

Museums in Budapest.

Sie ist allein mit L. székessy2 sp. nov. verwandt, welche Art aber viel schmaler

ist, nicht paralleles Halsschild, vollkommen abweichende Skulptur und ein-

farbig dunkle Beine besitzt.

Lytta rubrolineata sp. nov.

Gross und langgestreckt; Kopf schwarz, mit je einem grossen gelbroten Fleck

beiderseits neben der Innenseite der Augen, Halsschild ebenfalls schwarz, die

Scheibe aber beiderseits etwas naher dem Vorderrand als dem Hinterrand mit

je einer grossen, ovalen Makel an der Mittellinie, Fliigelddecken schwarz, die

Naht, der Seitenrand und die Scheibe mit je 3 langen Langsnerven, welche

gelbrot gefarbt sind und vor dem Ende zusammenfliessen, Unterseite, Beine und

Fiihler schwarz, die Behaarung der Ober- und Unterseite weissgrau, anliegend.

Kopf gross und rundlich, mit stark gew6lbten und grossen Augen, Stirn zwischen

den Augen ziemlich flach, Schlafen nach hinten verjiingt und breit abgerundet,

die Punktierung fein und sparlich, hie und da etwas dichter, der Grund ist

dazwischen mikroskopisch sehr dicht punktiert und kaum glanzend. Halsschild

deutlich langer als breit, in Querrichtung stark gewélbt, etwas vor der Mitte

am breitesten, nach vorne konisch und gerade verengt, nach hinten fast parallel.

Die Langsmittelfurche tief und scharf, fast bis zum vorderen Viertel reichend.

Oberflache ungleichartig und runzelig dicht punktiert, der Grund ist sehr dicht

lederartig gerunzelt, matt. Die Behaarung anliegend, etwas schuppenartig, die

roten Makeln sind in der Mitte unbehaart. Fliigeldecken mit 3 erhabenen Langs-

nerven, welche unbehaart sind, die Zwischenraume dicht und fein, runzelig

STUDIEN UBER SUDAFRIKANISCHEN MELOIDEN 77

punktiert, dazwischen ist der Grund chagriniert und matt. Die Behaarung der

Zwischenraume etwas fleckenartig, auf einigen Flecken fehlt die Behaarung

vollkommen. Fihler die Basis des Halsschildes erreichend, das 3. Glied doppelt

so lang wie breit, die Glieder vom 4. an stufenweise kiirzer, aber gleichbreit, so

dass das 10. kaum etwas langer als breit ist, das Endglied langoval, zugespitzt.

Beine einfach, ohne besondere Kennzeichen, Mittel- und Hinterschienen aussen

abgeflacht, die beiden Enddorne der Hinterschienen dick, ziemlich kurz, das

Ende schrag abgestutzt, das 1. Glied der Hintertarsen fast so lang wie die beiden

folgenden Glieder zusammen und 1, 5-mal so lang wie das Klauenglied. Unter-

seite Ausserst fein und sparlich raspelartig punktiert. Lange: 13-19 mm. Breite:

5-7 mm.

1 Exemplar aus 8.W. Afrika, Nagusib (ungefahr 24 km. SO. von Namutoni),

Jan. 1923, leg. Mus. Exped. (Holotypus) in der Sammlung des South African

Museum und 1 Exemplar aus 8.W. Afrika, Onolonga (ungefahr 50 km. SW.

von Ondongua, Ovamboland), Febr. 1923, Mus. Exped. (Paratypus), welches

fur die Sammlung des Ungarischen Naturwissenschaftlichen Museums in

Budapest iiberlassen wurde.

Diese Art ist wegen ihrer Farbung leicht kenntlich und kann mit Sicherheit

auf Grund der Farbung von den nachst verwandten Arten unterschieden

werden. Ausser der Farbung ist die neue Art auch durch die Skulptur von den

bekannten Arten verschieden. L. spilotella Pér. besitzt roten Kopf und rotes,

schwarz gesdumtes Halsschild, weiter gelbrote, und braun gefleckte Fliigel-

decken, rote Beine, etc., L. namaqua sp. nov. gelbe Schienen, einfarbig schwarzes

Halsschild und ausserdem ganz abweichende Form und Skulptur des Vorder-

kérpers, die Arten L. vellicata Er. und L. enona Pér. haben beide einfarbig

schwarzes Halsschild und abweichende Fligeldeckenzeichnung, usw.

Lytta székessyt sp. nov.

Breit und gross; Kopf schwarz mit je einem gelbroten Fleck beiderseits neben

der Innenseite der Augen, Halsschild, die ganze Unterseite, Fiihler und Beine

schwarz, Fliigeldecken gelbrot mit spdrlich stehenden, unregelmdssigen und

unbehaarten braunen Flecken, die Schulterbeulen und das Ende breiter braun.

Kopf gross und quer, stark gewdélbt, Schlafen kaum parallel, abgerundet,

Scheitel gewdlbt, ohne eingedriickte Mittellinie, Stirn zwischen den Augen

vorne in der Mitte etwas aufgewolbt. Die Punktierung ziemlich fein und dicht,

dazwischen ist der Grund schwach punktiert, ziemlich glanzend. Halsschild so

lang wie breit, schmaler als der Kopf, vor der Mitte am breitesten, nach vorne

plotzlich und gerade verjiingt, nach hinten gerundet etwas verengt. Hinterrand

fein, aber scharf gerandet. Die Langsmittelfurche schwach vertieft, etwas vor

der Mitte erloschen, breit und unpunktiert. Scheibe beiderseits neben der

Mitte mit je einer grésseren unpunktierten, etwas aufgewolbten Stelle. Die

Punktierung dicht und fein, der Grund dazwischen mit mikroskopischen

Punkten dicht besetzt, fettglanzend. Fiiigeldecken auf der Scheibe mit je einem

Langsnerv an der Innenseite der Schulterbeulen, weiter die Naht, sowie der

78 ANNALS OF THE SOUTH AFRICAN MUSEUM

Seitenrand scharf nervenartig aufgebogen. Die Oberseite dicht und runzelig

punktiert, dazwischen chagriniert und matt. Fiihler kraftig, die Basis des Hals-

schildes etwas iiberragend, mit ziemlich gleichartigen Gliedern. Das 3. und 4.

Glied etwas schmaler, scheinbar langer, die Glieder vom 5. an dicker und mehr

rundlich, das Endglied lang und zugespitzt. Beine ohne besondere Kennzeichen,

die beiden Enddorne der Hinterschienen gleichdick und kurz, schrag abgestutzt.

Mittel- und Hinterschienen im Querschnitt langoval, Aussenseite abgeplattet.

Unterseite sehr dicht und fein mit raspelartigen Punkten besetzt, die Punktierung

lasst eine breite Langsbinde an der Hinterbrust spiegelglatt frei. Lange: 13-14

mm. Breite: 4, 5 mm.

1 Exemplar aus S.W. Afrika, Kaokoveld, Zesfontein, Febr. 1925, leg. Mus.

Exped. (Holotypus) in der Sammlung des South African Museum und 1

Exemplar aus 8.W. Afrika, Kaross (ungefahr 100 km. SO. von Zesfontein),

Febr. 1925, leg. Mus. Exped. (Paratypus), welches fir die Sammlung des

Ungarischen Naturwissenschaftlichen Museums in Budapest iiberlassen wurde.

Die Art steht L. namaqua sp. nov. sehr nahe, sie besitzt eine 4hnliche Farbung,

doch besitzt L. namaqua sp. nov. helle Schienen, breites und paralleles Halsschild,

sowie abweichende Skulptur. |

Zu Ehren meines lieben Freunden, Direktor Dr. V. Székessy benannt.

Gen. Decapotoma Voigts

1902. Voigts, Wien. Entom. Keit., Xxi, p. 177.

(pro Decatoma Dejean 1821, preocc.)

Decapotoma csiki sp. nov.

Kleine, breite, einfarbig schwarze, glanzende Art, Fiihler auch schwarz,

Fliigeldecken mit je 8 kleinen, rundlichen Makeln, uzw.: eine an den Seiten

neben den Schultern, eine an den Seiten im vorderen Drittel, eine neben den

Seiten im hinteren Viertel, eine an der Naht hinter dem Schildchen, eine neben

der Naht in der Mitte, eine neben der Naht im hinteren Viertel, weiter eine

ausserst kleine an der Innenseite der Schulterbeulen und schliesslich eine runde

Makel etwas vor der Mitte der Scheibe, so dass endgiiltig vorne 3, mitten 3 und

hinten 2 Flecke vorhanden sind. Kopf sehr breit, mit grossen, stark gewodlbten

Augen, Schlafen ganz kurz und nach hinten abgerundet, Stirn leicht gewolbt,

die Mittellinie unpunktiert, Oberflache grob und dicht mit nabelartigen Punk-

ten besetzt, aus welchen aufrecht stehende, lange, schwarze Haare wachsen,

dazwischen ist der Grund glatt und glanzend. Halsschild schmdéler als der

Kopf, Seiten bis zum vorderen Viertel parallel, vorne gerundet stark ver-

schmalert, in Querrichtung stark gewolbt, Oberflache ohne Eindriicke, nur vor

der Basis in der Mitte etwas verflacht. Die Punktierung sehr grob und sparlich,

hie und da etwas dichter, der Grund ist dazwischen vollkommen glatt und

glanzend. Fltigeldecken von den Schultern an nach hinten breit erweitert, das

Ende einzeln vollkommen abgerundet, Oberflache sehr dicht und ziemlich

stark punktiert, die Punkte sind stark nebeneinandergeriickt, an der Basis ist

STUDIEN UBER SUDAFRIKANISCHEN MELOIDEN 79

die Spur nur einer Langsrippe erkennbar; glanzend, anliegend sehr fein

behaart und sparlich mit langen aufstehenden schwarzen, dickeren Haaren

besetzt. Die gelben Flecke sind viel sparlicher punktiert. Fuhler kurz, das Ende

sehr dick, das 2. Glied kugelig, das 3. schmal und nur kaum doppelt so lang wie

breit, das 4. und 5. ebenso breit, aber kiirzer, das 6. kaum merklich breiter, das

7. aber trapezformig, nur so lang wie das 6., das 8. langer und auch breiter

trapezformig als das 7., das 9. noch erweitert, quer, zylindrisch und das Endglied

sehr dick und lang, langer als die beiden vorhergehenden Glieder zusammen.

Beine einfach, ohne besondere Kennzeichen, Schienen deutlich langer als die

Tarsen, schmal und langgestreckt, die beiden Enddorne der Hinterschienen

sehr klein und gleichspitzig, diinn. Lange: g mm. Bretie: 3, 5 mm.

3 Exemplare aus S.W. Afrika, Kaross (ungefahr 100 km. SO. von Zesfontein,

Kaokoveld), Febr. 1925, leg. Mus. Exped. (Holo- und Paratypen) in der

Sammlung des South African Museum, von welchen 1 Paratype fiir die Samm-

lung des Ungarischen Naturwissenschaftlichen Museums in Budapest iiber-

lassen wurde.

Ich kenne keine nahere Verwandte innerhalb der Gattung. Sie ist durch die

einfarbig schwarzen Fihler, sowie durch die Fliigeldeckenzeichnung sehr

charakteristisch und kann mit keiner beschriebenen Art verwechselt werden.

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: | PART III, containing a 7

3. South African Pycnogonida. By K. H. Barnarp, D.Sc., F.L.S. (With

34. text-figures.) : |

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ISSUED APRIL 1954 PRICE 15s.

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BY THE RUSTICA PRESS (PTY.) LIMITED, COURT ROAD, WYNBERG, CAPE

7. South African Pycnogonda. By K. H. Barnarp, D.Sc., F.L.S. (With 34

text-figures. )

This paper is the result of an examination of the South African Museum

material comprising mainly the collection made by the Cape Government

trawler s.s. Pieter Faure, together with a moderate number ‘of specimens

obtained from time to time by shore-collecting.

The history of our knowledge of South African Pycnogonids has been given

by Flynn (1928). Gordon (1932) has redescribed three species from the type

material. Since then Helfer (1937), Barnard (1946) and Stock (1951) have

described material from South Africa south of lat. 15° S.

Flynn (1928) gave a list of the then-known South African species, including

a Mozambique record. The list omits Parapallene merstraszi which Flynn

himself records in his paper; but on the other hand it includes Pallenopsis

crossland: which neither Flynn nor any other author has recorded from South

Africa. Not counting ‘Pallenopsis sp. Loman’ (1923) the list contains thirty-six

species.

A second paper by Loman, also published in 1923, and containing two new

records (one from Angola), was not mentioned by Flynn. In 1937 Helfer

described three new genera and species; in 1946 Barnard one new genus and

ten new species; and in 1951 Stock one new genus and two new species. In

the present paper three of Loman’s species and records, one of Flynn’s species,

and one of Helfer’s are regarded as synonyms. The list is appreciably longer

than Flynn’s, and contains fifty-three species.

All the accepted families are represented in the fauna. Of the newly

recorded constituents it is interesting to note the occurrence of the genera

Pipetta and Austrorapius. The genus Queubus Brnrd., of uncertain affinities, is

placed near but not in the family Pycnogonidae.

No examples of ‘extra-legged’ (10 or 12 pairs) species have been found

in South African waters. These remarkable Pycnogonids are mostly inhabitants

of the Antarctic, though examples have also been found in the Caribbean Sea

(see: Hedgpeth, 1947, Smiths. Misc. Coll., cvi, pp. 9 sqq.).

The early stages of some species are parasitic in the zooids of Hydroids,

causing the formation of galls. An example from South Africa has been noted

by Warren (1908, Ann. Natal Mus., i, p. 293) and referred to by Flynn (1928,

P- 5):

Collectors and others should be warned against the use of cotton wool as a

plug in tubes or bottles to prevent shaking while in transit, because no other

animals, not even Crustacea, seem to get their claws so inextricably entangled

mor, XLi, PART 3.

AUC Iso

82 ANNALS OF THE SOUTH AFRICAN MUSEUM

in it. A plug of soft paper should be interposed between the animals and the

cotton-wool plug in a tube.

List of South African Species

Colossendeidae

Colossendeis colossea Wilson

, macerrima Wilson

Pipetia capensis Brnrd.

Rhopalorhynchus kréyert. Wood-Mason

Nymphonidae

Nymphon capense Hodgson ? = phasmatodes Bohm

5, australe Hodgson

5, signatum Mobius

», adistensum Mobius

5, ptlosum Mobius syn. bipunctatum Flynn

» comes Flynn

» affins Stock

5, natalense Flynn

5, macroctenatus Brnrd.

5 erenatiunguis Brnrd.

» settmanus Brnrd.

Pallenidae

Callipallene sp.

Pseudopallene gilchristt Flynn

Parapallene spinosus (Mobius)

af nierstraszi Loman

a calmam Flynn

e hodgson: Brnrd.

¥ algoae Brnrd.

Metapallene dubitans (Hodgson) syn. Procidella gibber Helfer

Pallenoides magnicollis Stock

Pallenopsis oscitans (Hoek)

5, brevidigitata Mobius

5 intermedia Flynn syn. Pallenopsis sp. Loman, 1923, P. fluminensis

Loman, 1923, non Kroyer

5 capensis Brnrd.

», (Rigona) crossland: Carp.*

be » ovalis Loman

Hannoma typica Hoek

* See p. 116,

SOUTH AFRICAN PYCNOGONIDA 83

Phoxochilidiudae

Phoxochilidium capense Flynn

Anoplodactylus aculeatus Mobius

ms pelagicus Flynn

‘Pallene’ lappa Bohm

Endeidae

Endeis clipeatus (Mo6bius)

5, mollis (Carp.)

Ammotheidae

Béhmia chelata (Bohm)

5, tuberosa Mobius

Achelia quadridentata (Hodgson)

5, brevicauda (Loman)

Nymphopsis cuspidata (Hodgson) syn. abstrusus Loman

Kyphomia setacea Helfer ? Ammothella setacea

Ascorhynchus ornatum (Helfer)

Austroraptus thermophilus Brnrd.

T anystylidae

Tanystylum ornatum Flynn

Discoarachne brevipes Hoek

Pycnogonidae

Pycnogonum cataphractum Mobius

$5 forte Flynn

si microps Loman

45 portus Brnrd.

3 pusillum Dohrn

Incertae sedis

FT.

Queubus jamesanus Brnrd.

Key to the South African Familtes

Body extended, crurigers distinctly separated (except Pallenopsis subg.

Rigona, where they are contiguous). Proboscis directed forward (except

Hannonia).

A. No chelifers (in adult). No auxiliary claws. Colossendeidae

B. Chelifers present.

1. Ovigers present in both sexes, with compound spines (fig. 3d).

Ocular tubercle not, or not far, in front of 1st crurigers.

a. Palps present, 5-jointed. Nymphonidae

b. Palps absent, or knob-like, or 3-4-jointed. Pallenidae

2. Ovigers present in g only, with simple spines. Phoxochilidiidae

C. Chelifers and palps absent. Auxiliary claws present. Endeidae

Body more or less compact, segmented or the segmentation suppressed,

crurigers usually narrowly separated. Proboscis directed forwards, or more

or less downwards, or bent under body.

84 ANNALS OF THE SOUTH AFRICAN MUSEUM

A. Palps present. Ovigers present in both sexes, without apical claw.

1. Palps 6-10-jointed. Ammotheidae

2. Palps 4-6-jointed Tanystylidae

B. Palps absent. Ovigers sometimes present in § only, with apical claw Pycnogonidae

Fam. CoLOSssENDEIDAE

1908. Loman, Siboga Exp. monogr., xl, p. 21 and conspectus facing p. 19

(subfam. Colossendeinae.)

1913. Schimkewitsch, Zool. Anz., xli, p. 614.

1913. Bouvier, 2me Exp. Antarct. Franc., p. 37.

1933. Calman and Gordon, Proc. Roy. Soc. Lond., B, cxili, p. 113.

Legs 8-12. Body of 4-6 segments, fused or free. Proboscis large, immovable.

Chelifers absent (usually) in adult. Palps long, usually 8-10-jointed, arising

from knob-like processes. Ovigers in both sexes, arising from knob-like

processes, 10-jointed, with apical claw, last 4 joints with non-serrate (usually)

spines. Bases of palps and ovigers approximate or contiguous. Legs long,

tarsus usually of fair length, tarsus and propodus unarmed or not armed

with strong spines, without auxiliary claws. Genital apertures on all legs

(2nd coxal joint) in both sexes.

Key to the South African Genera

Octopodous

1. Body segments fused. Crurigers not widely separated. Very large

species. Colossendeis

2. Body segmented. Crurigers widely separated.

a. Abdomen elongate. 2nd coxa longer than Ist or 3rd. Pipetta

b. Abdomen minute. All three coxae subequal Rhopalorhynchus

Gen. Colossendeis Jarzinsky

1881. Hoek, Rep. H.M.S. ‘Challenger’, iii; pp. 28, 61.

1919. | Bouvier, loc: cit.) p, 53:

1917. id., Res. Scz. Camp. Monaco, fasc. 51, p. 7.

1932. Gordon, “Discovery Rep., vi, p. 11.

1936s.) id Austral Antarct. Exp. (C2 ee. ps 7

1944.) 1d! Bb eAUNE ZZ. Antarct: Hap... B. ve pte lor

Octopodous. Proboscis more or less evenly cylindrical, or constricted

basally.

Key to the South African Species

Last 3 joints of the legs together much shorter than two-thirds length of 2nd tibial

joint (brevitarsal group).

1. Proboscis stout, constricted basally. Palps much longer than proboscis, 4th

joint shorter than 2nd. colossea

2. Proboscis slender, evenly cylindrical. Palps a little longer than proboscis, 4th

joint longer than end. macerrima

a ——

SOUTH AFRICAN PYCNOGONIDA 85

Colossendeis colossea Wilson

1881. Wilson, Bull. Mus. Comp. Zool. Harv., viii, p. 244, pl. 1, fig. 1., pl. 3,

figs. 5-7.

Seateettioek, loc. cit., pp. 61, 147, pls, figs. 1, 2;+pl.°10, figs. 1-5 (gigas)...

1917. Bouvier, loc. cit., p. 13, pl. 1, fig. 2, pl. 2, fig. 1 (coloured) (references).

1923. Calman, Rec. Ind. Mus., xxv, p. 266.

1928. Flynn, Fish. Mar. Biol. Surv. Rep., vi, Spec. Rep. 1, p. 7.

1944. Gordon, loc. cit, p. 11.

1953. Stock, Temminckia, ix, p. 311.

Proboscis stout, constricted in basal third. Palp much longer than proboscis,

4th joint shorter than and, 6th about twice as long as 5th; 7th, 8th and oth

subequal. Femur of legs subequal to ist tibia, much longer than 2nd tibia.

Length of body (excl. proboscis) 21 mm., proboscis 33 mm., legs 240 mm.

(Hoek: 32°5, 47-5 and 301 mm. resp.). Reddish (see Bouvier’s coloured figure).

Localities: off Cape Point, 660-755 fathoms (Flynn); off Cape Point, 650-

moeagniath. (S. Afr. Mus.); off Gneka River, 43 fath., and 29° 44 S.

31° 20’ E., 46 fath. (Flynn).

Distribution. Southern Indian Ocean, East Indian Sea, off west coast of

South America, Japan, North and Central Atlantic.

Colossendeis macerrima Wilson

foam Visor, loc: cit-)'p. 246, pl. 1; fig. 2, pl. 3, figs: 9-12, pl. 5, fig. 32:

1881. Hoek, loc. cit., pp. 64, 147, pl. 8, figs. 3-7 (leptorhynchus)

far ye0) Bouvier, loc. cit., p. 10, pl. 1, fig. 1 (coloured), pl: 3, figs. 1) 2.

maze Calman, Toc. cit.,‘p: 267." '

1928. Flynn, loc. cit., p. 7 (macerrime, and p. 6 macerriama, typ. err.).

1953. Stock, Temminckia, ix, p. 307, fig 17 e-h.

Proboscis slender, nearly evenly cylindrical, often slightly upturned in

distal third. Palps only a little longer than proboscis, 4th joint much longer

than 2nd, 5th and 6th subequal, 7th and 8th subequal, gth usually a little

longer than 8th. Femur of legs subequal to Ist tibia, much longer than and

tibia.

Length: body 17 mm., proboscis 34 mm., legs 155 mm. Reddish.

Localities: off Cape Point, 660 fath. (Flynn); off Cape Point, 700-800

fath. (S. Afr. Mus.); off Gneka River, 43 fath. (Flynn).

Distribution. As for colossea.

Remarks. Much less abundant than colossea in Cape waters.

Flynn has remarked that the shallow-water records of both species need

confirmation. Gneka* River is about midway between Keiskamma Point and

East London.

* Spelt Nieca in the s.s. Pieter Faure log-books and Rep. Gov. Biologist 1901 chart; Ncera

on sheet 5 of the topographical map issued by the Irrigation Dept. 1939; also sheet SE 35/24

Port Elizabeth, 1:500,000. 1950.

86 ANNALS OF THE SOUTH AFRICAN MUSEUM

Gen. Pipetta Loman

1904. Loman, Tijdschr. Ned. Dierk. Ver. (2) viii, p. 264.

TQOG,) 1d loc vent piney.

1914. Hodgson, Zool. Anz., xlv, p. 159.

1915. id., Ann. Mag. Nat. Hist. (8) xv, p. 141 (English version of 1914 paper).

1927. id., D. Stdpol. Exp., xix (zool. xi), p. 314.

Octopodous. Body segmented, very slender. Crurigers slender, very

widely separated. Ocular tubercle conical, no eyes. Proboscis elongate, very

slender. Abdomen elongate, slender. Palps slender, 8-jointed. Ovigers

10-jointed, with small apical claw. ‘Legs slender, no auxiliary claws.

Two species hitherto known: P. weberi Loman 1904 from the East Indies,

2,081 metres; and P. australis Hodgson 1914 from the Antarctic (‘Gauss’

winter quarters), 2,450 metres.

Calman and Gordon (1933, loc. cit., p. 114) exclude this genus from the

Colossenderdae; Hedgpeth (1947, Smiths. Misc. Coll., cvi, p. 5) includes it with

a question mark.

Pipetta capensis Brnrd.

Bic.

1946. Barnard, Ann. Mag. Nat. Hist. (xi) 13, p. 60.

Body rod-like, slender, with scattered microscopic granules. Crurigers

separated by increasingly wide intervals, the end pair slightly the longest,

4th pair shortest; 1st with a low conical elevation on dorsal apex, 2nd-4th

each with a strong upstanding spiniform process. Ocular tubercle high,

slender, conical, no eyes. Proboscis proximally slightly swollen, distally very

slender. Palps slightly longer than proboscis, 1st joint short, swollen, 2nd

longest, apically swollen, with a few setae, 4th long, about two-thirds length

of 2nd, with a few setae, 3 spine-setae and a longer one on apex, 5th-8th joints

short, sparsely setose. Oviger 1st joint short, swollen, 2nd a little more than

twice the grd, 4th one and a half times the 2nd, 5th short, 6th longest, apically

clavate, 7th-1oth joints short, more or less swollen, each with a few (2-4)

blunt denticles on inner margins, apical claw small.

Legs very slender, 3rd coxa with a strong upstanding slender spiniform

process; femur with a few long setae and a spine-seta on dorsal apex; Ist

tibia slightly shorter than femur, 2nd tibia slightly shorter than ist tibia, both

with scattered setae; tarsus about half length of propodus, the latter slender,

slightly curved, with fine setae on lower margin, claw feebly denticulate on

lower margin.

Body, tip of proboscis to tip of abdomen, about 7-5 mm. (specimen broken

in half); proboscis 3:5 mm., abdomen 1:75 mm., femur 2 mm., Ist tibia

1°75 mm., 2nd tibia 1-5 mm.

Locality: off Cape Point, N. 81° E. 32 miles, 460 fath. 1 specimen (S. Afr.

Mus.).

SOUTH AFRICAN PYCNOGONIDA 8 7

Fig. 1. Pipetta capensis Brnrd. a. dorsal view. 5. lateral view of ocular tubercle, from left side.

c. palp. d. oviger. e. 2nd tibia, tarsus and propodus of leg.

88 . ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks. Resembles weber: in having the tarsi long proportionately to the

propodi (not very short as in australis), but differing from both species in having

spiniform processes on the 2nd-4th crurigers.

Gen. Rhopalorhynchus Wood-Mason

1873. Wood-Mason, 7. Asiat. Soc. Beng., xlii, p. 171.

1908. Loman, loc. cit., p. 23.

Octopodous. Body segmented, slender, rod-like. Crurigers short, stout,

widely separated. Ocular tubercle short, conical, eyes present. Proboscis

elongate, fusiform. Abdomen very short. Palps slender, 9-jointed. Ovigers

slender, 10-jointed, with small but stout unguiform apical claw. Legs slender,

without auxiliary claws.

Rhopalorhynchus kréyeri Wood-Mason

igus?

1873. Wood-Mason, loc. cit., p. 171, pl. 13.

1923. Calman, Rec. Ind. Mus., xxv, p. 268, fig. 1 (references and synonymy;

the date of publication of Haswell’s tenuissima is 1885).

1953. Stock, Temminckia, ix, pp. 279, 280, fig. 1b (chart).

Fig. 2. Rhopalorhynchus kréyeri Wood-Mason. Dorsal view, with lateral view of ocular tubercle

from right side.

Proboscis with a sharp denticle medio-dorsally at about the middle of its

length. Ocular tubercle rather abruptly constricted above the eyes (cf.

Carpenter, 1907, fig. 28). Oviger inner margins of 7th-1oth joints with

numerous feebly biserrate spines (in several rows), roth joint in addition with

a stout spine distally opposing the small unguiform claw (as in Calman’s

figure; apparently overlooked by Loman, 1908, pl. 15, fig. 216). Legs with

coxal joints subequal, femur distally clavate in 9, in ¢ more slender and less

conspicuously clavate, 1st tibia slightly shorter than femur, 2nd tibia slightly

shorter than Ist, tarsus about two-thirds propodus, claw strong, shorter than

tarsus (cf. Loman, pl. 15, fig. 220).

SOUTH AFRICAN PYCNOGONIDA 89

Proboscis 7:5, femur g, Ist tibia 8-5, and tibia 7-5 mm.

Localities: off Hood Point, near East London, 42 fath.; off Port Shepstone,

Natal, 24 fath.; and off Cape Natal (Durban), 85 fath. 399 (S. Afr. Mus.).

Distribution. Maldive Is., Ceylon, Indian Seas, Andaman Is., East Indies,

Torres Straits, Queensland.

Remarks. ‘The position of the denticle on the proboscis corresponds with

Loman’s fig. 215, not with Calman’s fig. 1 a, 6. Calman refers to the variation

in the claw of the legs, but does not suggest that slender and strong claws are

characteristic of the g and Q respectively.

Fam. NyYMPHONIDAE

1913. Schimkewitsch, Zool. Anz., xli, p. 607.

1913. Bouvier, 2me Exp. Antarct. Franc., p. 65.

1932. Gordon, ‘Discovery’ Rep, vi, p. 24.

Legs 8 or 10. Body segmented. Proboscis immovable. Chelifers well

developed, usually conspicuously chelate. Palps 5-jointed (7-jointed in

Paranymphon Caullery). Ovigers in both sexes, 10-jointed, with apical claw,

last 4 joints usually with denticulate spines (8-jointed, without denticulate

spines in Paranymphon). Legs with or without auxiliary claws. Genital apertures

on all legs in 9, on last 3 in g.

Gen. Nymphon Fabr.

1794. Fabricius, Ent. Syst., 1v, p. 417.

1888. Sars, Arch. Naturv. Christ., xii, p. 352 (Chaetonymphon).

1908. Loman, Szboga Exp., monogr., xl, p. 37.

1915. Calman, Brit. Antarct. (Terra Nova) Exp., zool. III, p. 28.

1923. Loman, Swed. Antarct. Exp., 1, 2, p. 10.

1932. Gordon, loc. cit., p. 26 (revision of Antarctic species).

1937. Giltay, Mem. Mus. Roy. Hist. Nat. Belge. (2) fasc. 9, p. 83 (key to 7

species on west coast of Africa, Morocco to Cape of Good Hope).

1944. Gordon, B.A.N.Z. Antarct. Res. Exp., B, v, pt. 1, p. 17 (key to Antarctic

and Subantarctic species).

1951. Stock, Mem. Inst. Roy. Sci. Nat. Belge. (2) fasc. 43, p. 5.

1953. id., Temminckia, ix, p. 282.

Octopodous. Proboscis subcylindrical. Ocular tubercle at base of neck,

never in front of bases of ovigers, eyes usually present. Finger and thumb of

chelifer with more or less slender spiniform denticles. Oviger with apical

claw, last 4 joints with denticulate spines. Legs with or without auxiliary

claws.

Remarks. A large genus, the Antarctic and Subantarctic species of which

have been usefully analysed by Gordon.

gO ANNALS OF THE SOUTH AFRICAN MUSEUM

Key to the South African Species

I. Auxiliary claws vestigial or absent.

A. Auxiliary claws vestigial. Main claw slender. Oviger modified; denticu-

late spines on last 4 joints 23-37.

B. Auxiliary claws absent.

1. Finger and thumb of chelifer apically crossing, each with 50-65

spines. Palp and joint longest.

a. Neck short, crurigers narrowly separated.

i. Crurigers less than half their own width apart. 3 oviger

modified, spines 48-50.

ii. Crurigers about their own width apart. g oviger not modified.

a. Spines on oviger 54-63(66).

B. Spines on oviger 44.

b. Neck long, crurigers widely separated. Spines on oviger 25.

2. Finger and thumb of chelifer not crossing, with very numerous and

fine spines (300 +). and, 3rd and 4th joints of palp subequal.

Spines on oviger 46.

II. Auxiliary claws well developed.

A. ‘Tarsus at least half length of propodus, sometimes a little longer than

propodus.

1. Finger and thumb of chelifer apically crossing; former with many

more (twice as many) spines than latter.

a. Legs feebly setose.

i. Neck short. Tarsus shorter than propodus. Oviger with 36-38

denticulate spines.

ii. Neck long. Tarsus longer than propodus. Oviger with 40-42

denticulate spines

b. Legs strongly setose. Neck short. Tarsus shorter than propodus.

Oviger with 23-26 spines.

2. Finger and thumb not crossing, an approximately equal number of

spines on both.

a. Finger and thumb each with less than 25 spines. Oviger with

34-36 spines.

6. Finger and thumb each with 45 or more spines. Oviger with 39

spines.

B. ‘Tarsus only one-third length of propodus. Finger and thumb of chelifer

with an equal and moderate number of long spines, and long curved

setae on the setose pad. Oviger with 51-55 spines.

Nymphon australe Hodgson

australe

capense

comes

affinis

natalense

microctenatus

signatum

distensum

pilosum

crenatiunguis

angolense

setimanus

1902. Hodgson, br. Mus. Rep. ‘Southern Cross’ Crust., p. 257, pl. 40.

1902. Mobius, D. Tiefsee Exp. ii, p. 181, pl. 26, figs. 1-6. (Chaetonymphon

altioculum.)

1913. Bouvier, loc. cit., p. 73, figs. 25-31 (stylops).

1915. Calman, loc. cit., p. 36 (synonymy).

1920) )) Elymmn, loc. jet. .p.) 16:

1932. Gordon, loc. cit., p. 59, figs. 25 d, 26 b.

1938. id., Austral. Antarct. Exp: @) AUG. p.i13.

1944. id., loc. cit., p. 24 and var. caecum, p. 25, fig. 5 a-d.

Body compact, neck short, crurigers separated by a distance varying from

a quarter to equality with their own width. Proboscis subcylindrical. Ocular

SOUTH AFRICAN PYCNOGONIDA QI

tubercle high, conical, eyes subterminal. Abdomen reaching to end of 1st

coxa of last leg, clavate or pyriform. Sparsely setose on cephalic lobe and

crurigers.

Chelifer scape longer than proboscis, hand subequal to scape, finger and

thumb longer than palm, nearly straight, apically curved and crossing, each

with 38-46 closely set spines. Palp with and joint longest, 4th and 5th joints

subequal. Oviger in § modified, 5th joint longest, distal half strongly expanded,

6th joint also swollen, last 4 joints with 23-37 denticulate spines. Legs rather

short, setose, 2nd coxa slightly longer than 1st + 3rd, femur shorter than either

tibia, Ist tibia longer than and, tarsus longer than propodus, claw slender,

half length of propodus, auxiliary claws vestigial.

Body including abdomen 7-8 mm.

Locality: off Cape Point, 1,190 fath. (Flynn). The locality of another

specimen, from a depth of only 46 fathoms, is regarded by Flynn as doubtful.

Distribution. Circumpolar (antarctic), nearest locality to South Africa:

Bouvet Island.

Remarks. The s.s. Pieter Faure obtained no specimens referable to this species.

Nymphon capense Hodgson

? 1879. Bohm, MB. Ak. Wiss. Berlin, p. 173 (phasmatodes).

1908. Hodgson, Tr. Roy. Soc. Edinb., xlvi, p. 169, pl. 1, figs. 2, 2a.

1928. Flynn, loc. cit., p. 4 (remarks on synonymy).

fez Gordon: Ann. Mas. Nat. Hist.) (10), 1x,\p: 117, figs. 11, 12 (redes-

cription).

ne32,) id., loc. Cli., pp. 27-32, 34 (in key), 62.

Body stout, neck short, base of oviger large, occupying most of the space

between Ist cruriger and anterior cephalic expansion. Crurigers distinctly but

narrowly separated. Ocular tubercle stout, low, rounded, eyes distinct.

Proboscis bottle-shaped, apically rounded. Abdomen slightly longer than

last crurigers, oblique (at about 45°).

Chelifer scape a little longer than proboscis, hand longer than scape, finger

and thumb apically curved and crossing, with 60-65 closely set spines, in 2 or 3

alternating sizes. Palp with 2nd joint longest, 3rd, 4th and 5th progressively

shorter. Oviger ¢ with 3rd joint inflated, 4th and 5th subequal, rather short

and stout, 5th apically dilated, 6th deeply excavate on one side with long setae

protruding from the cavity, last 4 joints with 48-50 denticulate spines (Hodgson:

Byea2, 00,024, Gordon, (12,,12, 11, 19), claw: with 13-15, spinules. In; 2

normal (see: comes). Legs slender, feebly setose, 2nd coxa equal to 1st + 3rd,

femur shorter than Ist tibia, and tibia longer than 1st (7, 8 and 10-7), tarsus

and propodus equal or latter a little longer, claw two-thirds to three-quarters

the length of propodus, no auxiliary claws.

Body (¢ 2) 2:7-2°8, proboscis 1-3-1-6, 3rd leg femur 2-6-3, Ist tibia 3-4-4,

2nd tibia 4-4-6 mm. (Gordon).

Q2 ANNALS OF THE SOUTH AFRICAN MUSEUM

Locality: off Dassen Island, 35 fath. 99 and ovig. 3d (Hodesos South-

west of Cape of Good Hope, 50 fath. (Béhm).

Remarks. Flynn thought that Hodgson’s species was synonymous with

Boéhm’s. Béhm’s specimen was a @ with possibly abnormal ovigers. Hodgson’s

description was based on ovigerous gj as well as 99, and the type material

has been re-examined and figured by Gordon, who does not, however, discuss

the possible synonymy. Giltay (1937) and Stock (1951) both accept the

synonymy.

The species is a small one.

Nymphon comes Flynn

Figs. 3, 5¢

1929. Flynn, loc. cit., p. 14, figs. 4-6.

Body stout, neck well marked, of medium length, cephalic segment subequal

to rest of body (incl. abdomen). Crurigers separated by distances not quite

equal to their own width, their length subequal to median width of segments.

Ocular tubercle shortly conical, in juvenile (body 5 mm.) with 2 apical points

(as seen from in front) which gradually disappear leaving a blunt point in

adult, eyes distinct. Proboscis stout, expanded in middle, apically rounded.

Abdomen about as long as last crurigers, slightly clavate, horizontal. A few

minute setules on hind margins of segments, and on crurigers.

Chelifer scape slightly longer than proboscis, hand about equal to scape,

finger and thumb longer than palm, apices curved and crossing, inner margins

with more or less regularly alternating long and short spine-teeth, about 50

in number. Palp rather stout, 2nd joint longest, 3rd, 4th and 5th progres-

sively shorter. Oviger in ¢ 4th joint slightly longer than 5th, slightly curved,

5th slightly curved at base, apically expanded on ventral surface with a semi-

circlet of strong spines, last 4 joints with 54-63 (66) denticulate spines resp.

17-21, 14-16, 11-12 (Flynn: 15), and 12-14. In 9 4th joint slightly longer than

5th, both nearly straight. Legs moderately slender, ist tibia a little longer

than femur, 2nd tibia one and a half to one and a third times as long as femur

(Flynn says 2nd tibia is twice the femur and one and a half times Ist tibia, his

measurements are nearly in agreement, but his figures are not), tarsus sub-

equal to, or slightly shorter than propodus, claw half to two-thirds as long as

propodus, no auxiliary claws. Femur and Ist tibia in 3 not appreciably swollen

distally. Femoral cement glands in ¢ (ventral) minute, very numerous, at

least 70. Feebly setulose, more numerous setae on Ist and and tibiae.

Body @ incl. abdomen 12, abdomen 1°5, proboscis 6, 2nd leg 67 mm. (femur

15, Ist tibia 17, 2nd tibia 20 mm.). Body dg incl. abdomen 10-5, abdomen 1°5,

proboscis 5, 2nd leg 59 mm. (femur 12, Ist tibia 14:5, 2nd tibia 18 mm.).

Localities: False Bay, 22 fath. 1 g (Flynn); off Cape Infanta, 46 fath.

4 99 2 non-ovig $d; off Cape Seal, 80 fath. 1 juv.; off Cove Rock (East

London), 43 fath. 1 non-ovig ¢ (S. Afr. Mus.).

SOUTH AFRICAN PYCNOGONIDA 93

Remarks. In addition to the discrepancy in the leg shown in fig. 6, Flynn

figures the oviger as g-jointed (fig. 5).

Although reaching a larger size than Flynn’s § specimen (the East London

g is approximately the same size as the type), these specimens seem to belong

to comes. There is one point of difference: the length of the claw of the legs.

a Cc d b

Fig. 3. Nymphon comes Flynn. a. dorsal view. 5b. front views of ocular tubercle of juvenile, 5 mm.

in length (top), intermediate stage, and adult (bottom). c¢. oviger g 5th joint, ventral view

above, external view below. d. oviger g distal joints with spine and claw further enlarged.

In the type it is about one-third the propodus, but in all the present specimens,

including the juvenile, it is at least a half, usually two-thirds the propodus.

This is a much larger species than capense (phasmatodes), even the type being

nearly three times as big; but it is difficult to find (from Hodgson’s description)

morphological differences. ‘The denticulate spines on the last 4 joints of the

oviger are given by Hodgson as resp. 15, 12, 11, 12 in number; and he describes

the teeth on the finger and thumb of the chelifer as being of three sizes.

94 ANNALS OF THE SOUTH AFRICAN MUSEUM

Nymphon affinis Stock

1951. Stock, loc. cit., p. 5, figs. 1-6.

This species, described from a single 2 5-8 mm. in length, is very close to

comes, and Stock himself suggests that it may be the @ of the latter species of

which only the ¢ has hitherto been known.

Stock relies on the following differences in establishing a separate species.

Finger and thumb of chelifer with alternating larger and smaller spines;

Stock considers that Flynn would certainly have mentioned this feature if it

had been present in comes. The 5th joint of palp is relatively longer: five-

sevenths the length of 4th, instead of half. The different number of spines on

the oviger: 15, 11, 10, 8 = 44, as against 66.

Little reliance can be placed on the first difference, as Flynn’s descriptions

are not always very detailed. Variation would probably be found in the relative

lengths of the joints of the palp and legs if a large number of specimens of

different sizes were available.

For the present, affinis may stand, characterized by the 44 spines on the

oviger. The 5 mm. juvenile specimen of comes has 54 spines on the oviger.

Locality: WLiideritzbucht, 8 fath. (Stock).

Nymphon natalense Flynn

1926) Blymn; loc. jcit., p. 11s fig.9:

Body slender, neck long and narrow, cephalic segment a little longer than

rest of body (incl. abdomen). Crurigers separated by distances about twice

their own width, their length considerably greater than median width of

segments. Ocular tubercle shortly conical, eyes distinct. Proboscis rather

stout, expanded in middle, apically rounded. Abdomen shorter than last

crurigers.

Chelifer scape slightly shorter than proboscis, hand longer than scape by

nearly one-third (details of finger and thumb not given). Palp 2nd joint

longest, 3rd, 4th and 5th progressively decreasing. Oviger 5th joint longest,

apically expanded in g, denticulate spines on last 4 joints resp. 9, 6, 5, 5.

Legs long and slender, 2nd coxa very long, a little more than twice as long as

ist and about half the length of ist tibia, 1st and 2nd tibiae subequal, tarsus

shorter than propodus, claw about three-quarters the length of propodus, no

auxiliary claws. Feebly setose.

Body 3:6, proboscis 1-2, leg 18-7 mm. (femur 3:2, rst and 2nd tibiae 4 mm.).

Locality: off Port Natal (Durban), pelagic, 1 damaged 3 (Flynn).

Remarks. ‘There are several discrepancies between the description of the

legs, the measurements, and the figure.

I have seen no specimens.

- SOUTH AFRICAN PYCNOGONIDA

Nymphon microctenatus Brnrd.

Fig. 4

1946. Barnard, Ann. Mag. Nat. Hist., (xi), 13, p. 60.

9—Body moderately stout, neck moderate, base of oviger separated from

ist cruriger and occupying the whole of the rest of the neck behind the anterior

cephalic expansion. Crurigers separated by about their own width, or rather

less, their length subequal to median width of segments. Proboscis very stout,

ee reomrars

Fig. 4. Nymphon microctenatus Brnrd. a. dorsal view of cephalic segment with proboscis and

basal joint of chelifer. 5. lateral view of ocular tubercle. c. 4th and 5th joints of palp. d. inner

view of chela (spines much more numerous than shown).

nearly evenly cylindrical, apex truncate. Ocular tubercle low and rounded,

eyes distinct. Abdomen not extending beyond last crurigers, clavate, hori-

zontal. Glabrous.

Chelifers widely separated at base, scape slightly longer than proboscis,

hand about as long as scape, palm short, finger and thumb both slightly curved

downwards, their apices following the same curve, i.e. not crossing, margins

with an enormous number (estimated at 300-400) of slender closely set spinules

96 ANNALS OF THE SOUTH AFRICAN MUSEUM

like a very fine comb. Palp long and very slender, almost glabrous, 2nd, 3rd

and 4th joints subequal, 5th a little more than half 4th joint, gently curved.

Oviger 5th joint longer than 4th (subequal to 3rd + 4th), 4th with slight

swelling proximally on inner side, 5th straight, 6th with a rather prominent

spine on inner apex, last 4 joints with 46 denticulate spines (14, 11, 10, 11) on

inner margin, spines with 6 (4 major) denticulations, claw with 18-19 spines.

Legs, coxae and femur moderately stout, tibiae distinctly more slender, 2nd

coxa a little longer than 1st + 3rd, femur a little shorter than Ist tibia, and

tibia considerably longer than Ist, tarsus distinctly but not much shorter than

propodus, claw about half the length of propodus, no auxiliary claws. Coxae

and femur glabrous, Ist tibia very sparsely setose, 2nd tibia sparsely and

finely setulose.

Body incl. abdomen 7-5, abdomen 1, proboscis 3, 2nd leg coxa 3:5,

femur 10, Ist tibia 11-5, 2nd tibia 20 mm.

Locality: off Buffalo River (East London), 195 fath. 1 9 (S. Afr. Mus.).

Remarks. ‘The specimen is broken in pieces, but all essential parts are present.

Nymphon signatum Mobius

Fig. 54

1902. Mobius, D. Tiefsee Exp., ii, p. 178, pl. 24, figs. 1-7.

1932. Gordon, Ann. Mag. Nat. Hist. (10) ix, p. 104, fig. 5 (redescribed).

1932. id., loc. cit., pp. 28-9 (in Table iii), 35 (in key).

Body moderately stout, neck stout and moderately long, base of oviger

occupying two-thirds of its length, cephalic segment widely expanded.

Crurigers separated by about half their own width, their length less than

median width of segments. Proboscis stout. Ocular tubercle higher than wide,

apex bluntly pointed or feebly bifid. Abdomen extending slightly beyond last

crurigers, horizontal or slightly oblique. Glabrous.

Chelifer with a few setae dorsally and apically, chela at least as long as

scape, finger and thumb apically curved and crossing, former with 60-64 spines,

latter with about 35 longer spines, the two series of spines approximately

the same length. Palp 2nd joint longest, 3rd-5th progressively shorter, 4th ++

5th subequal to 3rd. Oviger in ¢ 4th joint slightly shorter than 5th, both

curved, 5th with apical lobe bearing a semicirclet of spines (cf. comes); in 9

Ath joint longest, 4th and 5th joints straight; last 4 joints with 36-38 (11-13,

g-11, 7-9 and 8-10) denticulate spines, each with 5 major denticulations,

claw two-thirds roth joint, with 10 spinules.

Legs moderately long and slender, femur and Ist tibia distally swollen in 3,

femur shorter than Ist tibia, 2nd tibia longer than Ist tibia, with spines on its

apex, tarsus nearly as long as propodus, latter with numerous spines on lower

margin, claw nearly half length of propodus, auxiliary claws well developed.

Feebly setose, especially in 9, scattered setae on femur, more numerous on Ist

SOUTH AFRICAN PYCNOGONIDA 97

and and tibiae. Femoral cement glands in ¢ (ventral) 17-20, the distal ones

much smaller than the proximal ones.

Body incl. abdomen 8:5, proboscis 4:5, abdomen 1-5, 2nd leg femur 9g,

ist tibia 11, 2nd tibia 15 mm.

an py a ON aag iy

Le. as stn MEER Ly, hy

Lae:

enn PH Hp

Pippen rm Desh

se iulaniauusaner es g

Fig. 5. Chela of: a. Nymphon signatum Mobius, juv. 5. Nymphon distensum Mobius. c. Nymphon

comes Flynn.

Localities: Agulhas Bank, 100 metres (Mobius); False Bay, 19-27 fath.

I ovig. 6; 10 miles off Cape St. Blaize, 1 9; off Gt. Fish Point, 49 fath. 3

ovig. gg, I non-ovig 3, 1 immature J; off Glendower Beacon (Port Alfred),

66 fath. 1 non-ovig. ¢ (S. Afr. Mus.).

Remarks. With the holotypes of this species and distensum before her, Gordon

pointed out certain small differences, but remarked that intermediate forms

98 ANNALS OF THE SOUTH AFRICAN MUSEUM

might be found to unite them. The small amount of material in my hands

does not confirm this, although two of the differential characters mentioned

by Gordon must be discarded. Spines are present in all cases on the apex of

and tibia; evidently in the holotype they had been rubbed off. Differences

in the number of spines on ventral margin of propodus are not reliable.

The two species are distinguished by their build, length of neck, and

relative lengths of tarsus and propodus (tarsus < propodus in szgnatum,

tarsus > propodus in distensum).

The 3 from False Bay is particularly heavily built, spines on finger and

thumb of chelifer resp. 50 and 30, denticulate spines on oviger (only one is

complete) 45.

A small specimen, body incl. abdomen 2°75 mm., from off Umhlangakulu

River (Natal), 50 fath., may belong to this species. The neck and distance

separating the crurigers as in signatum, but abdomen oblique, ocular tubercle

rather high and distinctly bifid apically. Second coxa a little longer than

ist + 3rd, distinctly swollen apically. Oviger with 32 denticulate spines

(10, 8, 6, 8). Chelifer hand with finger strongly curved, with fine apical point

curving in the opposite direction, with 38 spines, thumb with 28 spines.

Tarsus shorter than propodus, claw scarcely one-third length of propodus,

auxiliary claws two-thirds main claw.

Nymphon distensum Mobius

Fig. 50

1902. Mobius, loc. cit., p. 179, pl. 25, figs. 1-6.

1928. Flynn, loc. cit., p. 10.

1932. Gordon, loc. cit., p. 105, fig. 6 (redescribed).

1932. id., loc. cit., pp. 28-9 (in Table iii), 35 (in key).

Body rather slender, neck slender and rather long, base of oviger occupying

less than half its length, cephalic segment not widely expanded. Crurigers

separated by distances less than their own width (except the last 2 pairs),

their length subequal to median width of segments. Proboscis stout. Ocular

tubercle rather high, conical, more or less constricted above eyes, apex more

or less distinctly bifid. Abdomen very slightly longer than last cruriger, clavate,

horizontal. Glabrous.

Chelifer scape subequal to proboscis, hand as long as scape, finger and

thumb apically curved and crossing, former with 60-70 (holotype 9, fide

Gordon: 60-64), latter with 30-33 longer spines, the two series of spines of

approximately equal length. Palp 2nd joint longest, 3rd-5th progressively

shorter, 4th + 5th subequal to 3rd. Oviger in ¢ 4th and 5th joints subequal,

both curved, 5th with apical lobe bearing semicirclet of strong spines (cf.

comes); in @ 4th and 5th joints subequal, both straight; last 4 joints with

40-42 (13-16, 9-11, 8-9 and g-10) denticulate spines, each with 5 major denti-

culations, claw two-thirds to three-quarters length of 1oth joint, with about

15 spines.

SOUTH AFRICAN PYCNOGONIDA 99

Legs long and slender, femur and 1st tibia distally swollen in 3, 2nd coxa

equal to or a little longer than 1st + 3rd, femur shorter than 1st tibia, 2nd tibia

much longer than (one and one-third) 1st tibia, apex with spines, tarsus a little

longer than propodus (Gordon: tarsus three-quarter length of propodus in

4th leg, but figure of 1st leg shows tarsus a trifle longer than propodus), claw

about half length of propodus, auxiliary claws one-third, or a little more,

main claw. Feebly setose, especially in 9, a few scattered setae on 1st and 2nd

tibiae. Femoral cement glands in 3 (ventral) 12-15.

Fig. 6. Nymphon pilosum Mobius. a. ocular tubercle, front (above) and lateral (below) views.

b. femur g. ¢. oviger § 5th joint. d. spines from distal joints of oviger, normal and abnormal (or

worn).

Body incl. abdomen ovig. g 11, 2 13:5, abdomen J 1°5, 2 2, proboscis 3 5,

2 6, and leg 3 64, 9 92 mm. (femur J 11°5, 9 20, Ist tibia J 15, 9 23, 2nd tibia

3 25°5, 2 37°5 mm.).

Localities: Agulhas Bank, 154 metres, 1 2 (Mobius); off Gneka River, 43

fath. 1 2 (Flynn); off Buffalo River (East London), 195 fath. 3 gd (1 ovig.),

12; off Cape Morgan, 87 fath. 1 2; off Cape Seal, 37 miles, 80 fath. 1 juv.;

Cape Point N. 71° E. 18 miles, 230 fath. 1 9 (S. Afr. Mus.).

Remarks. Holotype @ redescribed by Gordon. The few specimens I have

examined all agree with distensum as described by Gordon, and for the present

this species may be retained as distinct from szgnatum.

100 ANNALS OF THE SOUTH AFRICAN MUSEUM

The largest 9, whose measurements are given above, came from the Cape

Point locality.

Flynn’s specimen appears to be aberrant in having the tarsus shorter than

propodus (cf. szgnatum) as in Mobius’s figure, which, as Flynn says, cannot be

relied upon for strict accuracy.

N. arabicum Calman 1938 seems to be very similar to distensum.

Nymphon pilosum Mobius

Fig. 6

1902. Mobius, loc. cit., p. 179, pl. 24, figs. 8-12.

1928. Flynn, loc. cit., p. 8, figs. 1, 2 (bzpunctatum).

Body moderately stout, neck short and stout, base of oviger occupying whole

space between Ist cruriger and cephalic expansion. Crurigers separated by

about their own width or rather less, their length a little less than median

width of segments. Proboscis moderately stout, usually widest in proximal half

and slightly tapering to a rounded apex. Ocular tubercle rather high,

columnar, slightly expanded above owing to the large eyes, surmounted by a

transverse ridge which usually shows two little points. Abdomen distinctly

longer than last crurigers, cylindrical or slightly clavate, horizontal. A few

setae on hind margins of cephalic and following two segments.

Chelifer sparsely setose, subequal to or slightly longer than proboscis, hand

subequal to scape, finger and thumb apically curved and crossing, former

with about 40 spines, latter with about 26 larger spines, the two series of spines

of approximately equal length. Palp 2nd joint longest, 4th and 5th sub-

equal or the latter slightly the shorter, 4th + 5th subequal to 3rd. Oviger

in ¢ 4th joint slightly curved, with slight swelling proximally on inner side,

5th slightly longer than 4th, curved, apically swollen, setose, especially on

apex ventrally; in 9 4th and 5th joints subequal, straight; last 4 joints with

23-26 (g, 5-6, 4-5 and 5-6) (one specimen with 10, 6, 5, 6 = 27), stout and

strongly denticulate spines, with 3 major denticulations, claw with 6-7 spines.

Legs moderately stout, femora and 1st tibiae not distally swollen in d,

2nd coxa subequal to, or in the juv., slightly longer than, 1st + 3rd coxae,

femur shorter than Ist tibia, which is shorter than 2nd, tarsus distinctly though

not much shorter than propodus, claw not quite half length of propodus,

auxiliary claws about half length of main claw. A few setae on the coxae,

femur with short and longer outstanding setae, tibiae with numerous long

outstanding setae. Femoral cement glands in ¢ large, about 5 or 6 (4-7) on

low knobs or ridges on ventral surface.

Body incl. abdomen 7, abdomen 1-75, proboscis 3, 2nd leg 38 mm. (femur

6, 1st tibia 7, 2nd tibia 9-5 mm.). The non-ovigerous ¢¢ are a little smaller

than the largest 9.

Localities: Agulhas Bank, 154 metres 1 9 (Mobius); off Port Natal (Durban),

tow-net, 5 gd; 4 9° (Flynn); off Cape St. Blaize, 85-90 fath., 4 non-ovig. 3d,

SOUTH AFRICAN PYCNOGONIDA IOI

6 99; off Cape Seal, 80 fath. 1 9; off Cape St. Francis, 75 fath. 1 non-ovig. 3;

Someon oy 2 20.1 1) 200math, 12) (S. Ate, Mus.) 20% )567 S..919 11 1, 339

metres (Fisheries Survey, 1948).

Remarks. There would seem to be little doubt that bipunctatum is synonymous;

at least it is strange that Flynn did not compare his specimens with pilosum;

possibly being taken in a tow-net and therefore presumed to be pelagic uncon-

sciously created an impression of specific distinctness.

The ocular tubercle is characteristic. he stout spines on the last joints of

the oviger are also very striking.

EAA

Fig. 7. Nymphon crenatiunguis Brnrd. a. chela with denticles further enlarged. 6. apex of claw

of oviger 69. c. chela of a g 2°25 mm. in length.

Owing to their pilosity the legs are often much obscured by foreign matter;

some of the specimens have examples of the stalked barnacle Scalpellum

agulhense Brnrd. attached to the legs; and one specimen has one of the barnacles

standing upright immediately behind the ocular tubercle.

Nymphon crenatiunguis Brnrd.

Bic. 7

1946. Barnard, loc. cit., p. 60.

Body moderately stout, neck short, base of oviger contiguous with first

cruriger and extending nearly to beginning of cephalic expansion. Crurigers

separated by less than their own width, their length subequal to (a trifle

102 . ANNALS OF THE SOUTH AFRICAN MUSEUM

shorter than) median width of segments. Proboscis nearly cylindrical, apex

rounded. Ocular tubercle rather low, conical, in front view with 2 little

points surmounting the eyes. Abdomen slightly longer than last crurigers,

clavate, oblique (at about 45°). Glabrous.

Chelifer scape subequal to proboscis, hand shorter than scape, palm oblong,

widening slightly distally, finger and thumb relatively short and stout, former

subequal to palm, apices not crossing, finger with 20(9)-22(¢) closely set

spiniform denticles, thumb with 18(Q)-17(¢) more slender and slightly more

widely spaced denticles. Palp and joint longest, 4th + 5th joints a little longer

than grd, 5th longer than 4th. Oviger in ¢ 5th joint longer than 4th, gently

curved, with apical lobe bearing strong recurved spines; in 2 4th and 5th

subequal; last 4 joints with 34-36 denticulate spines (Jj 13, 7, 7,7; 214, 9, 6, 6

and 13, 9, 7, 7), Spines stout with 3 major denticulations. Claw with about

15 denticles, which distally become successively shorter and form a rounded

and minutely crenulate (apparent) apex, the true apex being acute and minute.

This form is found on both ovigers of the g and of the 9, though in one oviger

of the latter the distal denticles are sharper and more prominent than in the

figure (the apex thus appears more serrate than crenulate).

Legs moderately stout, 2nd coxa subequal to ist + grd, femur shorter than

ist tibia, 2nd tibia longer than Ist, tarsus and propodus subequal, claws about

one-third propodus, auxiliary claws strong, at least half main claw. Glabrous

except the 2nd tibia which is sparsely and minutely setulose. Femoral cement

glands in ¢ 4-5 low knobs or tubercles on ventral surface.

Body incl. abdomen ¢ 4, 2 3, abdomen 3:75, 9-5, proboscis 3 1°75, 9 1°5,

leg femur ¢ 4°5, 2 4, Ist tibia ¢ 5, 9 5, 2nd tibia ¢ 7-5, 9 6-5 mm.

Locality: off East London, 85 fath. 1 non-ovig, g, 1 9 (S. Afr. Mus.).

Remarks. Distinguished by the hand of chelifer, which resembles in general

that of grossipes Hoek 1881 and multtdens Gordon 1932. As in these two species

the 5th joint of the palp is longer than 4th; but the tarsus and propodus of the

legs are subequal, and the spines on the last joints of oviger are different (in

shape from those of grossipes, and in number from those of multidens). The

apex of claw of oviger seems to be unusual if not peculiar.

One non-ovigerous ¢, body incl. abdomen 2:25 mm., from off Durnford

Point, Zululand, go fath., agrees with the above described specimens in most

characters, including the peculiar claw of the oviger, but has the hand of the

chelifer sightly different. The finger is subequal to palm, but both it and the

thumb are more slender, the former with 15, the latter with 14 denticles

(fig. 7c). The femur has only 3, but very distinct, cement gland knobs. The

5th joint of oviger has the apical lobe with recurved spines, and the denticulate

spines on last 4 joints number 12, 9, 7, 6.

Nymphon angolense Gordon

? 1923. Loman, Goteb. K. Vet. Samh. Handl., xxvi 6 (Medd. Goteb. Mus.

Zool. Avd. no. 22), p. 5. (N. gracillimum, non Calman.)

SOUTH AFRICAN PYCNOGONIDA 103

19p220 Gordon, loc. ieit., (ps 77, figs. 30, 37.

no374)/) Giltay, loc, cif., p:. 88.

LOG.) tock) loc. cit.,\p.) 7.

Neck long and slender, base of oviger contiguous with rst cruriger. Crurigers

separated by more than their own width, their length greater than median

width of segments. Proboscis cylindrical, apex rounded. Ocular tubercle low,

rounded. Abdomen slightly longer than last crurigers, elevated at about 45°.

Chelifer scape subequal to proboscis, hand slightly longer than scape, palm

slender, finger and thumb slightly longer than palm, slender, apices crossed,

about 55 spines on finger, 45 on thumb. Palp 2nd joint longest, 3rd three-fifths

end, 4th half 3rd and 5th slightly longer than 4th. Oviger in ¢ slender, 5th

joint longer than 4th, straight, with a few hook-like spines on apex; in 2 4th

and 5th joints relatively short; last 4 joints with 39 denticulate spines (13, 10,

8, 8) (Stock: ¢ 13, 9, 7, 7). Claw with 14 spinules.

Legs slender, 2nd coxa slightly longer than 1st + grd, femur shorter than

Ist tibia, 2nd tibia longer than ist, propodus slightly longer than tarsus,

auxiliary claws at least two-thirds length of main claw.

Body incl. abdomen 7, proboscis 1-8-1-9, femur ¢ 6-8, 2 7, ist tibia ¢ 9-2,

9 8-4, and tibia 12 mm.

Localities: off Elephant Bay, Angola (approx. 13° 10’ S. 12° 45’ E.), 73-91

metres (Gordon); Tiger Bay, and Elephant Bay, 100-110 metres (Stock) ;

off Port Alexander, 60 fath. (Loman).

Remarks. Gordon set out several differences between angolense and the

holotype of gracillamum, but not the number of spines on the oviger. This is not

included in Calman’s original description but is given, on Calman’s authority,

Dylon (loc) cit. p. 5, tootnote)) as'39 (12) 846; 7) and) 30 (12,7, 5,16).

Momams specimens showed 31 (11, 6, 6, 8). and 28 (10, 7, 5, 6). All these

numbers are less than those in angolense.

Gordon made no comment on Loman’s record, but it seems more probable

that the Port Alexander specimen (‘verstitimmelten’, Loman) should be

identified with angolense than with an Antarctic species, in spite of the difference

in number of the spines on the oviger.

Port Alexander lies between the two localities recorded by Stock.

Nymphon setimanus Brnrd.

Fig. 8

1G46.,, Barnard: loc: (cit... OL.

Body moderately stout, neck long (cf. distensum), base of oviger contiguous

with first cruriger and occupying one-third to nearly one-half of neck. Crurigers

separated by rather less than their own width, their length subequal to median

width of segments. Proboscis stout, nearly cylindrical, apex rounded-truncate.

Ocular tubercle low, scarcely if at all higher than basal width, in lateral view

104. ANNALS OF THE SOUTH AFRICAN MUSEUM

conical, in front view apically truncate, with two minute points above the eyes.

Abdomen extending a little beyond last crurigers, somewhat elongate, oblique

(at about 45°). Glabrous.

Chelifer scape a little shorter than (or scarcely as long as) proboscis, hand

subequal to scape, finger and thumb much longer than palm, slender, curved,

apically crossing, each with 13-15 equally long spines, setose pad well developed

with numerous very long, almost spiniform setae on inner surface. Palp and

and 3rd joints subequal, or the latter a trifle the longer, 5th shorter than 4th,

4th + 5th longer than 3rd. Oviger 4th and 5th joints subequal, last 4 joints

eae

~ > 7

Fig. 8. Nymphon setimanus Brnrd. Inner view of chela 9.

with 51-55 denticulate spines (15, 13, 12, 11 or 18, 12, 12, II or IQ, 12, IT, 13),

each with about 6 major denticulations, claw with about 13 denticles, the

distal ones separated by crenulations, the apical point scarcely extending

beyond last denticle (cf. crenatiunguis).

Legs moderately stout, 2nd coxa equal to or a trifle longer than 1st + 3rd

coxae, femur only a little shorter than ist tibia, and tibia not much longer

than ist, tarsus short, about one-third propodus, claw short, subequal to

tarsus, auxiliary claws two-thirds main claw. Almost glabrous, even on 2nd

tibia only a few minute and scattered setules.

Body incl. abdomen 6, proboscis 2, abdomen 1-25, 2nd leg 2nd coxa 2:5,

Ist + 3rd coxae together 5, femur 6, Ist tibia 6-5, 2nd tibia 7 mm. Pinkish,

the palps and ovigers and a band around apex of femur pale whitish

(Pt. Elizabeth Mus. and U.C.T. specimens).

Localities: Bird Island Passage (Algoa Bay), 10-16 fath. 1 9; off East London,

32 fath. 1 9; off Port Shepstone (Natal), 24 fath. 1 9 (S. Afr. Mus.); shelly

beach, Cape Recife (Port Elizabeth Mus.); main channel, Knysna harbour;

Bushmans River mouth (Algoa Bay) (Univ. Cape Town Ecol. Surv. 1947

and 1950).

SOUTH AFRICAN PYCNOGONIDA 105

Remarks. Distinguished from all the other South African species by the

short tarsus.

Neck shorter than in andamanense Calman 1923. ‘The hand of chelifer

bears some resemblance to that of maculatum Carp. 1910.

Fam. PALLENIDAE

1908. Loman, Szboga Exp. monogr., xl, p. 40, and conspectus facing p. 19

(subfam. Palleninae).

1909. Schimkewitsch, ool. Anz., xxxiv, p. 6.

mons) ic, ibid., xli, p. 610.

1944. Gordon, B.A.N.Z. Antarct. Res. Exp., B, v, pt. 1, p. 36.

1947. Hedgpeth, Smiths. Misc. Coll., cvi, p. 4.

Octopodous. Body segmented (or last two segments fused). Proboscis

immovable. Chelifers present, chelate. Palps absent, or represented in ¢ by

a I-4-jointed rudiment. Ovigers in both sexes, 10-jointed, with or without

apical claw, distal joints with a single row of serrate, acute, or oval spines.

Legs with or without auxiliary claws. Genital pores on all legs in 9, on last

two legs in g. Eggs in several masses.

Key to the South African Genera

I. Ocular tubercle at hinder end of cephalic segment.

A. Legs with auxiliary claws. Palps absent.

1. Oviger without apical claw. Finger and thumb of chelifer serrate

or spinose. Callipallene

2. Oviger with apical claw. Finger and thumb of chelifer smooth,

with small dentiform lobe. Pseudopallene*

B. Legs without auxiliary claws.

1. Oviger with apical claw. Palps absent. Parapallene

2. Oviger without apical claw.

a. Palps reduced to a single knob-like joint. Metapallene

b. Palps absent. Pallenoides

II. Ocular tubercle at front end of cephalic segment. Palps absent or reduced

to tubercles.

A. Legs with auxiliary claws. Oviger without apical claw. Pallenopsis

B. Legs without auxiliary claws. Oviger with apical claw. Hannonia

Gen. Callipallene Flynn

1836. Johnston, Mag. Xool. Bot., i (4), p. 380. (Pallene; nom. preocc.

Megerle, 1823.)

1908. Loman, loc. cit., p. 42. (Pallene.) |

1929. Flynn, Mem. Queensl. Mus., 1x, p. 252, footnote.

1948. Correa, Pap. Avulsos Dept. Zool. S. Paulo, ix, 1, p. 1 (key to species).

*The presence or absence of auxiliary claws is not specifically mentioned in the single

South African species referred to this genus. See p. 107.

106 ANNALS OF THE SOUTH AFRICAN MUSEUM

1952. Stock, Beaufortia, no. 13, pp. 1-14, figs. 1-27. (European species. )

1953. id., Boll. Mus. Civ. Venezia, vi, 2, p. 179.

Cephalic segment with neck. Last 2 body segments often fused. Proboscis

short, without setae around mouth. No trace of palps. Chelifer scape I-jointed,

finger and thumb finely denticulate. Oviger 10-jointed, without (usually)

apical claw, spines on distal joints oval or round (not pointed), smooth or

finely serrated, 5th joint in ¢ with apical process. Legs with auxiliary claws.

Callipallene sp.

Fig. g

General appearance, length and shape of neck similar to fig. 7 of Stock

1952; all sutures between body segments distinct. Crurigers separated by

Q b | Cc d

Fig. 9. Callipallene sp. a. hand of chelifer. 6. propodus and claw of leg. c. 5th joint of oviger g.

d. distal joints of oviger 3.

intervals subequal to their own width; length slightly less than median width

of segments. Ocular tubercle conical. Abdomen obliquely upturned at

45-60°. Body and crurigers glabrous.

Chelifer scape stout, hand turned downwards, inner margin of finger

minutely and indistinctly serrulate, of thumb with about 10 conspicuous blunt

spiniform teeth. Oviger 5th joint longest, in ¢ with outstanding apical process

bearing a seta, the 4 distal joints with 38-46 (10, 10, 9, 9,—12, 12, 10, 12)

bluntly rounded spines, the margins of which appear to be quite smooth, no

apical claw.

SOUTH AFRICAN PYCNOGONIDA 107

Legs, femur in @ swollen, rst tibia shorter than femur and shorter than and

tibia, which is slightly longer than femur, propodus gently curved, with 4

large spines proximally, auxiliary claws strong, two-thirds length of claw,

simple (not pectinate at base); glabrous except for one or two fine setae on

each joint.

Body incl. abdomen 1°5, 2nd leg 7-5 mm.

Localities: off Walker Point, 47 fath., and off Cape St. Blaize, 40 fath.

(S. Afr. Mus.). Three specimens (2 jg, 1 mutilated 9 with developing ova).

Remarks. As these specimens show no outstanding features by which they

can be separated from the other species of the genus, I think no specific name

should be given to them until further material is available.

Gen. Pseudopallene Wilson

1804. Latreille, Nouv. Dict. Hist. Nat., xxiv, p. 137. (Phoxichilus.)

1878. Wilson, Trans. Connect. Ac., v.

1902. Stebbing, Knowledge, xxv, p. 187. (Phoxichilus, non. Latr.)

1909. Schimkewitsch, loc. cit., pp. 6, 7, 8.

1910. Hodgson, Schultze Reise, iv, p. 225 (in key).

1917. Bouvier, Res. Sci. Camp. Monaco, li, p. 28.

nego. Elynn, loc. 'cit..p. 23:

1951. Hedgpeth, Smiths. Misc. Coll., cvi, p. 4.

1953. Stock, Temmuinckia, ix, p. 204.

[not Pseudopallene Bouvier 1913 = Austropallene Hodgson 1914]

Proboscis. with setae around mouth. No trace of palps. Chelifer scape

1-jointed, hand short and swollen, finger and thumb not serrated or spinose,

but with dentiform projections. Oviger 10-jointed, with serrated apical claw,

5th joint in g with apical projection. Legs without auxiliary claws.

Remarks. After considerable argument as to the validity of Latreille’s name,

the name Pseudopallene seems to have been generally accepted.

Pseudopallene gilchristt Flynn

1Q2O ge ly iilWlOC- Cit. 4) 29 pe. 113.

Body slender, segmented. Crurigers widely separated. Neck elongate.

Ocular tubercie low, rounded. Abdomen shorter than last crurigers.

Chelifer scape shorter than proboscis, opposable margins of finger and

thumb each with a small rounded lobe.

Legs slender, elongate, minute spines on Ist and 2nd tibiae, claw more than

half length of propodus (no auxiliary claws ?).

Body 3:6-3:9, femur 3°8-4°8, 1st tibia 4°5-5°6, 2nd tibia 6-6-8 mm.

Locality: off Port Natal (Durban), surface (tow-net) (Flynn).

Remarks. Flynn makes no mention of auxiliary claws; presumably they are

absent, in conflict with the generic definition.

No examples of this species are in the South African Museum collection.

108 ANNALS OF THE SOUTH AFRICAN MUSEUM

Gen. Parapallene Carp.

1892. Carpenter, Sc. Proc. R. Dublin Soc., n.s., vii, p. 553-

1908. Loman, loc. cit., pp. 40, 42.

1909. Schimkewitsch, loc. cit., pp. 7, 9.

1929. Flynn, Mem. Queensl. Mus., ix, p. 258.

1937. Calman, Ann. Mag. Nat. His. (10), xx, p. 530.

1938. id., John Murray Exp., v, p. 156.

1953. Stock, Temmuinckia, ix, pp. 282, 297, fig. 2b (chart).

Body segmented. Proboscis constricted in middle. Chelifer scape 1-jointed,

hand swollen, finger and thumb fitting closely together when closed, margins

not denticulate. Palps completely absent. Oviger 10-jointed, in 9 4th joint

longest and 5th without apical process, in g 5th longest and with or without

apical process, distal 3 or 4 joints with a row of biserrate (usually, sometimes

obscurely serrate or simple) spines, with apical claw. Legs without auxiliary

claws (except in challengeri). Genital pores on last 2 legs in 3, on all legs in 9.

Key to the South African Species

1. Each cruriger with conspicuous pointed process dorsally.

a. 3rd coxa with conical process of ventral apex. spinosus

b. 3rd coxa with semicirclet of strong spines on ventral apex. calmani

2. Crurigers without processes (or only a minute tubercle on each).

a. Neck long.

i. 3rd coxa with a single spine on ventral apex. Feebly serrate spines on

7th-ioth joints of oviger 19, 16, 16, 16. algoae

ii. Strongly serrate spines on 7th-1oth joints of oviger 10, 9, 7, 9. nierstrazi

b. Neck short. 3rd coxa with semicirclet of fine spinules on ventral apex.

Feebly serrate spines on 7th-1oth joints of oviger 8-9, 8-9, 7, 7-6. hodgsoni

Parapallene spinosus (Mobius)

Fig. 10

1902. Mobius, D. Tiefsee Exp., ili, p. 188, pl. 28, figs. 8-12. (Anoplodactylus s.)

Crurigers well separated by about their own width, length subequal to or

slightly longer than median width of segments; each with an upstanding

conical process on dorsal apex. Neck moderately constricted, not defined by a

collar at base. Proboscis cylindrical, apically truncate. Ocular tubercle high,

conical, apex recurved and shortly bifid or narrowly truncate (acute in juv.

with body length 7-8 mm.), eyes of uniform size. Abdomen in 9 not longer

than last crurigers, in ¢ slightly longer, rather stout, nearly erect. Glabrous.

Chelifer scape stout, with a spine on upper apex and a fringe of spine-setae

on inner surface, hand curved downwards and inwards so that finger lies

ventral to thumb, palm subglobular, setose on inner surface around base of

finger and thumb, finger longer than thumb, inner margins of both entire.

Oviger 4th joint longest, in 2 8th-1oth joints with 7, 4-6, and 6 spines on inner

margin, spines moderate, simple or sometimes obscurely notched, or apically

somewhat hooked, claw with a few (8-9) spines on inner margin and 3 on both

SOUTH AFRICAN PYCNOGONIDA 109

inner and outer margins apically, the last pair adpressed to base of spiniform

apex (but these somewhat variable); in g 7th-roth joints with the spines

slightly stronger than in Q but not stout (claw missing in the only adult 3

specimen).

Legs rather stout, 3rd coxa with a conspicuous conical process on ventral

apex, femur about one and a half times 1st tibia, with 3 conical tubercles on

dorsal apex, the medio-dorsal one much larger than the dorso-lateral ones, Ist

tibia with 4 conical tubercles on dorsal apex, the dorsal pair larger than the

Fig. 10. Parapallene spinosus (Mobius). a. inner and front views of left chela. b. lateral view of

3rd coxa of leg. c. propodus and claw of leg. d. gth and toth joints with claw of oviger Q.

e. oviger of juvenile, body length 8 mm.

lateral ones (but not as large as the medio-dorsal one on femur), also a small

medio-dorsal tubercle near base, and one near middle on each lateral surface;

ond tibia a little longer than femur; propodus without heel, with 4 strong

spines proximally on lower margin, followed by smaller ones, upper apex

projecting over base of the stout claw. Very short and fine spinules ventrally

on apices of 2nd and 3rd coxae; on the other joints in these spinules become

more numerous distally, especially on 2nd tibia, but to the naked eye the legs

appear glabrous or the tibiae very finely hispid; in ¢ however there are a few

distinct but slender spines on femur, several on Ist tibia, and more on 2nd tibia.

Base of chelifers to base of abdomen 2 10, abdomen 2-2:25, proboscis 3,

and leg 38 mm. (femur 10°5, Ist tibia 6-5, 2nd tibia 12).

Localities: St. Francis Bay, Agulhas Bank, 10 metres (Mobius); off East

London, 52 fath. 1 9, 1 juv.; off Glendower Beacon (Port Alfred), 66 fath.

I not quite adult g; False Bay, 17-33 fath. 2 99, 2 juv.; Table Bay, 22 fath.

1g (S. Afr. Mus.).

I10 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks. The conical process of the 3rd coxa is very conspicuous and forms

an easy mark of identification. See further under calmanz.

Loman (1908, loc. cit., p. 64) suspected that this species should not be

included in Anoplodactylus on account of the number (Qg) of joints in the oviger.

The South African Museum specimens confirm this. They are obviously

examples of spinosus, although Mébius seems to have ignored the position of

e ‘ .

Fig. 11. Parapallene calmani Flynn. a, 6. lateral and ventral views of 3rd coxa of leg. c¢. distal

joints of oviger 3 with roth joint and claw further enlarged. d. distal joints of oviger 2. e. femur

and 1st and 2nd tibiae of leg ¢.

the ocular tubercle and makes no mention of it as he does in the case of

Anoplodactylus aculeatus. He had only one specimen and rashly assumed that

the 2 would be without ovigers.

One of the South African Museum specimens (body length 8 mm.) is

particularly interesting because it has an oviger closely resembling M6bius’s

figure 12; a figure is given here showing that it is not quite fully developed,

with the distal joints not completely demarcated and without spines on inner

margins.

P. spinosus is a larger species than calmant.

Parapallene calmani Flynn

Big 00

1928. Flynn, loc. cit., p. 16, figs. 7,8.

In general the description given for spinosus will apply to calmani. Chelifer

with spine on upper apex. The 3rd coxa is not conically produced, but has a

SOUTH AFRICAN PYCNOGONIDA Tete)

semicirclet of about 8-9 strong spines on ventral distal margin. The oviger,

both in ¢ and 9, has stronger spines on the inner margins of 7th-r1oth joints.

Localities: off East London, 47 fath. (Flynn); Algoa Bay, 10-17 fath. 3

non-ovig. §g, 1 9; off Glendower Beacon (Port Alfred), 66 fath., 1 non-ovig.

6 (S. Afr. Mus.).

Remarks. The presence in South African waters of two species both with

conical projections on the crurigers makes the identification of Flynn’s species

a little doubtful. Flynn might have overlooked the apical spines on the 3rd

coxa, and his fig. 8 gives no indication of them; on the other hand he could

scarcely have overlooked conspicuous conical processes and his figure, from

the viewpoint from which it is drawn, would surely have given some indication

Fig. 12. Parapallene algoae Brnrd. a. cephalic segment with chelifers. 6. lateral view of ocular

tubercle. c. distal joints of oviger 9.

of projections if they had been present. I therefore assign the specimens with

a semicirclet of spines on 3rd coxa to calmani; and the species with conical

processes on 3rd coxa is obviously spinosus (Mobius).

Flynn says the 4th joint of the 10-jointed oviger is the longest; his fig. 7

shows seven joints beyond the longest joint, making an 11-jointed oviger. This

is probably a draughtsman’s error.

Parapallene algoae Brnrd.

Bic

1946. Barnard, loc. cit., p. 61.

Body rather slender. Crurigers without processes, well separated by at

least their own width, length greater than median width of segments. Neck

well marked, only half as wide as apical width of cephalon, not defined by a

ie? ANNALS OF THE SOUTH AFRICAN MUSEUM

collar. Proboscis cylindrical, apically truncate. Ocular tubercle high, conical,

apex, acute, eyes of uniform size. Abdomen shorter than last crurigers, nearly

vertical. Body and crurigers microscopically hispid.

Chelifer as in calman, but without spine on upper apex of scape. Oviger

(2) 4th joint longest, 7th-1oth joints with numerous spines: 19, 16, 16, 16,

spines biserrate apically, claw finely serrate on inner margin except at base.

Legs (2) 3rd coxa with a single short but prominent spine on ventral apex,

dorsal apex of femur and ist tibia each with a semicirclet of 4-6 spines, Ist

tibia distinctly shorter than femur, 2nd tibia one and a half to one and three-

quarter times as long as Ist tibia, propodus as in calmam but lower margin

with 7-8 large spines, 3 or 4 of the proximal ones being the largest, claw

relatively longer than in calmam. Femur minutely hispid, ist tibia finely

spinulose-setose, 2nd tibia more strongly so, the longest (but none as long as

width of tibia) spine-setae forming a row or fringe along each side.

Base of chelifers to base of abdomen 15, abdomen 2, proboscis 4:5 mm.;

in S. Afr. Mus. specimens all legs severed from bodies, longest 80 mm. (femur

19'5, Ist tibia 14, 2nd tibia 20).

Localities: Algoa Bay, off Gt. Fish Point, and off Cape Morgan, 32-87

fath. 4 99 (S. Afr. Mus.); Plettenberg Bay, 30 fath. (Port Elizabeth Mus.).

Parapallene hodgsont Brnrd.

Fig. 13

1946. Barnard, loc. cit., p. 61.

Crurigers well separated by about their own width, length subequal to

median width of segments, each with a very small tubercle on upper surface

near apex. Neck constricted, but short, without collar at base. Proboscis

cylindrical, apically truncate. Ocular tubercle high, conical apex acute or

bifid, eyes of uniform size. Abdomen as long as last crurigers, oblique. Body

and crurigers glabrous.

Chelifer as in calmani but without spine on apex of scape. Oviger (9) 4th

joint longest, 7th-10th joints with rather stout spines resp. 8-9, 8-9, 7, 7-6,

spines simple or feebly serrate subapically, claw very feebly serrulate on inner

margin.

Legs (2) 3rd coxa with semicirclet of fine spinules on ventral apical margin,

femur longer than Ist tibia, 2nd tibia longer (about one and one-third) than

femur, propodus as in calmani but spines on distal part of lower margin stronger.

Femur slightly hispid, 1st and 2nd tibiae more conspicuously so, 2 (or 3) small

spiniform tubercles on upper apex of femur and st tibia.

Base of chelifers to base of abdomen 8-5, abdomen 1-5, proboscis 3 mm.;

and leg 48 mm. (femur 12, Ist tibia 10, and tibia 16-5 mm.).

Localities: off East London and Hood Point, 47-52 fath. 3 99, 1 juv. (S. Afr.

Mus.).

SOUTH AFRICAN PYCNOGONIDA 113

Remarks. The feebly serrate or simple spines on 7th-10th joints of oviger

prevent these specimens from being identified with merstraszi; also the neck is

shorter.

Parapallene merstrasz1 Loman

1908. Loman, loc. cit., p. 44, pl. 9, figs. 122-127.

1928. Flynn, loc. cit., p. 18.

1938. Calman, loc cit., p. 158, fig. 7.

1953. Stock, loc. cit., pp. 297, 299, fig. 3 a (chart).

Fig. 13. Parapallene hodgsoni Brnrd. a. posterior view of cruriger. 5b. lateral view of ocular

tubercle, with hind view of apex. c. distal joints of oviger 9.

Locality: 29° 35’ S., 31° 14’ E. (Natal), 25 fath. (Flynn). —

Distribution. East Indies.

Remarks. Calman examined a syntype which had the ocular tubercle

truncate with 2 points (fig. 7 A); this might be regarded as a malformation.

He also described the spines on the coxae and following joints but without

stating whether they were dorsal or ventral; presumably they were dorsal.

The spines on 7th-1oth joints of oviger are strongly serrate (Loman, fig. 124).

Gen. Metapallene Schimk.

1909. Schimkewitsch, loc. cit., pp. 7, 11, and table I.

1910. Hodgson, loc. cit., p. 235 (Heteropallene).

1938. Helfer, SB. Ges. naturf. Fr. Berlin, 1937, p. 172 (Procidella).

1952. Stock, Bull. Inst. Roy. Sci. Nat. Belge, xxviii, 14, p. 4 (remarks on

Procidella) .

1953. id., Beaufortia, 1V, No. 35, p. 38 (remarks on Procidella).

It4 ANNALS OF THE SOUTH AFRICAN MUSEUM

Body robust, segmented, crurigers not widely separated. Cephalic segment

broad, distally expanded, neck short. Proboscis short, without setae around

mouth. Abdomen short. Ocular tubercle at base of neck. Chelifer scape

1-jointed. Palp reduced to a single joint. Oviger 10-jointed, 5th joint with or

- without apical process in 3, distal joints with a single row of denticulate spines,

no apical claw. Legs without auxiliary claws.

Remarks. At Liideritzbucht, S.W.A., three specimens of a small Pycnogonid

have been collected and each has been made the type of a new species by three

separate authors. Although one suspects that these three specimens all belong

to one and the same species, the question is into what genus this species should be

put. Two new genera have been proposed.

Heteropallene Hodgson must fall into synonymy, because both Hodgson and,

a year earlier, Schimkewitsch named Pallene languida Hoek (1881, p. 79, pl. 12,

figs. 1-5) as the genotype of their genera.

M. languida is based on a 3 with the palps reduced to mere knobs; Hodgson’s

specimen, sex not stated, had rudiments of palps ‘each a slightly curved joint’ ;

in Procidella based on a 9 Helfer said each palp was reduced to a single conical

joint, but this is really the undeveloped oviger (see his figure) which he stated

was absent. In Hodgson’s specimen the 5th joint of the oviger ¢ has no apical

process, whereas in M. languida it has.

In view of these ambiguities, the two earlier Liideritzbucht specimens can

be only provisionally included in Metapallene, with Procidella as a synonym.

The third specimen has recently been adequately described as Pallenozdes

magnicollis (see infra).

Stock (1953) considers that Procidella gibber may be identical with M.

dubitans, but that as it was founded on a juvenile specimen it should be ignored.

Metapallene dubttans (Hodgson)

Fig. 14a

1910. Hodgson, loc. cit., p. 226, fig. 4 (distal joints of oviger) (Heteropallene d.).

1938. Helfer, loc. cit., p. 172, fig. 5 (Procidella gibber).

1953. Stock, loc. cit., p. 38, fig. 3 (Procidella gibber).

Body very robust, crurigers narrowly separated, with a few small setae

distally, the posterior pair very short and almost completely fused, with the

short abdomen embedded between them (Hodgson) or freely projecting

(Helfer’s figure). Ocular tubercle low, broad, near hind margin of cephalic

segment (Helfer’s figure); eyes small (Hodgson), without pigment (Helfer).

Chelifer stout, scape and hand setose, finger and thumb denticulate. Oviger

4th and 5th joints longest, 5th without apical process (Hodgson’s figure),

7th-ioth joints with 45 (12, 10, 11, 12 in Hodgson’s figure) serrate spines;

no apical claw.

Legs sparsely setose; propodus with 2 large spines proximally, claw very

stout, no auxiliary claws.

SOUTH AFRICAN PYCNOGONIDA

Length 1-5 mm. (Hodgson), 4:4 mm. (Helfer).

Locality: Liideritzbucht (Hodgson, Helfer). Hodgson’s specimen was

collected by Dr. L. Schultze; Helfer’s specimen was collected 18 Dec. 1903,

therefore presumably also by Dr. Schultze who was in Liideritzbucht in

December 1903 (1908. Schultze, Reise, i, Introduction, p. vi).

Fig. 14. a. Procidella gibber Helfer 2 (after Helfer). 5, c. Pallenoides magnicollis Stock, 3 with

propodus of leg and distal joints of oviger (spines omitted) further enlarged (after Stock).

Gen. Pallenoides Stock

1951. Stock, Mem. Inst. Roy. Sct. Nat. Belge, (2) fasc. 43, p. 8.

1952. id., Bull. Inst. Roy. Sct. Nat. Belge, xxvili, 14, p. 4.

Differs from Metapallene in having no trace of palps. Oviger without apical

Proboscis with a constriction near base, and setae

process on 5th joint J.

around mouth.

Pallenoides magnicollis Stock

Fig. 14), ¢

195 Us otock, loc. eit. p. 11, figs..7-13:

Compared with Metapallene dubiians the following points may be noted.

Abdomen short, embedded between the last crurigers as in Hodgson’s

description (thus differing from Helfer’s Procidella). Ocular tubercle farther

forward than in Helfer’s figure; eyes distinct. Oviger with 48 (14, 11, 10, 13)

serrate spines on 7th-1oth joints.

Length 1-75 mm. (3).

Locality: Liideritzbucht, 8 fathoms. ‘Mercator’ (Stock).

116 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks. ‘The locality and the close resemblance of this specimen to the

specimens described by Hodgson and Helfer, lead one to suspect that all three

specimens belong to one species. However, Stock has given a good recognizable

description of a species, whereas it must be admitted that dubitans is a species

inquirenda. Provisionally therefore I follow Stock. Re-examination of Hodgson’s

and Helfer’s specimens may help, but abundant fresh material is really required

to solve the problem satisfactorily.

Gen. Pallenopsis Wilson

1881. Wilson, Bull. Mus. Comp. Zool. Harv., viii, p. 250.

1902. Mobius, D. Tiefsee Exp., iti, p. 184.

1908. Loman, loc. cit, p. 65.

1913. Bouvier, 2me Exp. Antarct. Fr., p. 107.

1915. Calman, Terra Nova Exp., iii, p. 41.

1916. Loman, Zool. Med., ii, p. 15.

1923. Calman, Rec. Ind. Mus., xxv, p. 279.

1927. Hodgson, D. Stidpol Exp., xix, p. 334.

1932. Gordon, ‘Discovery’ Rep., vi, p. 87.

1933. Stephensen, Medd. om Gronland, \xxix, 6, p. 21.

1947. Hedgpeth, Smiths. Misc. Coll., cvi, pp. 3, 4, 6.

1953. Stock, Temminckia, ix, pp. 281, 288, fig. 4 (chart).

Body segmented, or segments fused. Ocular tubercle on front margin of

cephalic segment overhanging base of proboscis. Chelifer with scape 2-jointed,

or undivided, finger and thumb moderate, when closed meeting or gaping.

Palps reduced to tubercles. Oviger 10-jointed in g, in 9 rudimentary, some

of the joints sometimes fused; apical joint with numerous spine-setae, or a

single row of non-serrate spines, no apical claw. Legs with auxiliary claws

large or small, occasionally absent. Genital pores on last two legs in g, on all

legs in 9.

Key to the South African Species

A. Finger and thumb of chelifer meeting when closed.

1. Body segmented.

a. Auxiliary claws strong.

i. 2nd tibia one and one-sixth to one and a quarter as long as Ist

tibia. Legs (at least in 3) conspicuously setose. intermedia

ii. 2nd tibia one and one-third to one and a half as long as Ist

tibia. Legs not conspicuously setose. capensis

6. Auxiliary claws weak. brevidigitata

2. Body unsegmented or partly segmented (Rigona).

a. All segments fused. tst tibia with a few spine-setae, some arising

from small tubercles. ovalis

b. 3rd and 4th segments fused. 1st tibia with numerous digitiform

processes bearing spine-setae. [crosslandi|*

B. Finger and thumb of chelifer slender, curved, gaping when closed. oscitans

* P. crosslandi Carpenter (1910. 7. Linn. Soc. Lond., xxi, p. 257, pl. 27, figs. 10-20. Sudanese

Red Sea) is included because it was listed by Flynn (1928, p. 6), although neither Flynn nor,

so far as I can discover, any other author has recorded it from South African waters.

- — _ — = . a aE

SOUTH AFRICAN PYCNOGONIDA Lyle].

Pallenopsis intermedia Flynn

Figs. 15a, 16

? 1923. Loman, Ark. Zool., xv, 9, p. 10. (Pallenopsis sp.)

1923. id., Medd. Goteb. Mus., no. 22 (Goteb. K. Vet. Handl., xxvi), p. 3

( fluminensis, non Kréyer).

1928. Flynn, loc. cit., p. 20, figs. 10-12.

Body segmented. Crurigers well separated, but by spaces less than their

own width. Ocular tubercle conical, ending in a sharp point, anterior eyes

Fig. 15. a. Pallenopsis intermedia Flynn. b, c. Pallenopsis brevidigitata Mobius, with toth joint of

oviger. d. Pallenopsis ovalis Loman.

larger than posterior pair. Proboscis cylindrical, apically rounded. Abdomen

directed more or less obliquely upwards. Spiniform setae on dorsal surface of

body, scattered or more or less in paired groups of 1 or 2 on each segment,

shorter and more numerous along ends of crurigers, 9 usually less setose than

dg; abdomen with 2 rows of spines (often rubbed off).

Chelifer scape divided into 2 subequal portions, with spiniform setae on

distal margin of each; finger and thumb simple, closely fitting, finger longer

than thumb, glabrous (without spinose pad). Palpal tubercles distinct.

Oviger 10th joint as long as gth, oval, apically blunt, with numerous spine-

setae; in g 5th joint distinctly longer than 4th, and 6th conspicuously swollen.

Legs 2nd coxa subequal to 1st + 3rd, femur and Ist tibia subequal or the

latter slightly the shorter, 2nd tibia slightly longer than femur; propodus

cylindrical, without heel and not projecting over base of claw; auxiliary

claws large, at least half length of main claw; short spiniform setae on all

118 ANNALS OF THE SOUTH AFRICAN MUSEUM

joints, longer setae on 1st and and tibiae arranged chiefly in 3 fringes (one

mid-dorsal and one along each side), some of the longest setae slightly pinnate,

dg more conspicuously setose than 9. Cement duct on ventral surface of femur

short but distinct in ovig. ¢.

Body incl. abdomen ¢ 13, 2 15, proboscis 35, 2 6, 2ndleg g 56 (femur 13,

Ist tibia 12, and tibia 14), 9 67 mm. (resp. 17, 15, 18). Amber-coloured,

darker lines on sides of legs.

Localities: Sebastian Bay, Walker Bay, Cape Barracouta and Cape Infanta,

24-40 fath. (Loman); False Bay, off Cape Infanta, and off Gneka River (see

footnote, p. 85), 23-43 fath. (Flynn); False Bay and Agulhas Bank as far east

as Cape St. Francis, and in the East London area, 20-90 fath. (S. Afr. Mus.).

Remarks. In the s.s. Pieter Faure collection there are no specimens from the

area between Cape St. Francis and the East London area (of which the most

westerly locality is Gneka River mouth) in spite of the very considerable

amount of trawling done in and around Algoa Bay. On the trawling grounds

on the Agulhas Bank between Cape Infanta and Cape St. Francis this is the

commonest Pycnogonid. For this reason it seems certain that the specimens

identified as fluminensis Kroyer (a Brazilian species) by Loman (1923) should

really be assigned to intermedia. |

Ovigerous gg were found in September, October, December, February,

April, May and July; thus breeding probably occurs throughout the year.

Apparently very like vanhéffen Hodgson (1927. loc. cit., p. 336, fig. 9) from

the Antarctic, but the latter has a slender proboscis.

Loman (1923. Ark. Zool.) did not give the size of his specimen from Cape

Point Lighthouse; presumably it was collected in the littoral zone like

Discoarachne brevipes and Nymphopsis abstrusus (= cuspidata). It may perhaps be

a specimen of this species. |

Pallenopsts capensis Brnrd.

1946. Barnard, loc. cit., p. 62.

Body segmented. Crurigers widely separated (cf. brevidigitata, fig. 155), each

longer than median width of segment and about one and a half times as long

as their own width. Ocular tubercle high, bluntly conical, anterior eyes

slightly larger than posterior pair. Proboscis cylindrical, apically rounded.

Abdomen horizontal or slightly oblique, about as long as last cruriger. Body

with 1 or 2 short spine-setae in middle of each segment, and a few similar ones

on ends of crurigers.

Chelifer scape longer than proboscis, divided into 2 subequal portions,

hand as in intermedia, finger glabrous. Palpal tubercles conspicuous. Oviger as

in intermedia, in g 5th joint only slightly longer than 4th, 6th conspicuously

swollen.

Legs 2nd coxa subequal to 1st + 3rd, femur longer than rst tibia, end tibia

longer than femur, propodus without heel and not projecting over base of

SOUTH AFRICAN PYCNOGONIDA 119g

claw, lower margin with 4 or 5 large spines proximally and 4-6 shorter ones

distally, auxiliary claws strong, half length of main claw. Fine and short

spine-setae on apices of coxal joints, scattered over femur and Ist tibia, more

numerous on 2nd tibia. Cement duct on ventral surface of femur inconspicuous

in ovig. 3. |

Body incl. abdomen ¢ 18, 2 16, proboscis g 6:5, 2 6, 2nd leg g 98 (and

coxa 9, femur 25, Ist tibia 23, 2nd tibia 33), 9 75 mm. (resp. 7°5, 19, 17, 23).

Fig. 16. Pallenopsis intermedia Flynn. a. distal joints of oviger g. 0b. chelifer. c. one of the longest

tibial setae. d. propodus and claws of leg.

Localities: ‘Table Mountain 8. X E#E., 58 miles, 190 fath. 1 ovig. ¢; Cape

Point N. 16° E. 10 miles, 85 fath. 1 2; Cape St. Blaize N. X E. 73 miles, 125

fath. 1 non-ovig. ¢ (2nd leg 84 mm.). (S. Afr. Mus.)

Remarks. A 9 specimen, off Tugela River mouth, 200 fath., resembles the

above specimens in all characters mentioned, but is much more slender,

resembling in fact the 9 brevidigitata with which it was taken. Body incl.

abdomen 10:5, 2nd leg 72 mm. (femur 17, Ist tibia 17, 2nd tibia 21); width

of femur and 2nd tibia resp. 1 and -75 mm., as compared with 2 and 1-5 mm.

in the above @ capensis. Except for the auxiliary claws and the terminal joint

of oviger the specimen would be identified as brevidigitata.

120 ANNALS OF THE SOUTH AFRICAN MUSEUM

Pallenopsis brevidigitata Mobius

Fig. 150, c

1902. Mobius, loc. cit., p. 185, pl. 27, figs. 7-13.

1928. Flynn, loc. cit., p. 19, fig. 9.

1938. Calman, John Murray Exp., v, p. 160.

Body segmented. Crurigers widely separated, at least twice as long as their

own width, width across 2nd pair subequal to body length (excl. abdomen).

Ocular tubercle short, conical, on extreme front of cephalon. Proboscis

cylindrical, apically rounded. Abdomen horizontal, not extending as far as

end of last crurigers. Dorsal surface glabrous.

Chelifer scape divided, 1st portion longer than (but not ‘nearly twice as

long as’) and, hand as in intermedia, finger glabrous. Palpal tubercles small.

Oviger in 9 4th and 5th joints subequal, 6th slightly expanded, more so in J,

1oth joint slender, apically tapering, with a single row of large spines on

lower margin.

Legs 2nd coxa at least twice as long as 3rd, femur and ist tibia subequal,

and tibia longer, propodus without heel and not projecting over base of claw,

with large spines proximally on lower margin, followed by a comb-like series

of smaller spines, claw three-quarters length of propodus, slender, auxiliary

claws very short. Fine and short spine-setae, chiefly in 3 rows, on femur, Ist

and 2nd tibiae. Cement duct on ventral surface of femur inconspicuous in

ovigerous ¢.

Body incl. abdomen @ 9:5, proboscis 5, 2nd coxa 5, femur 16, Ist tibia 15,

and tibia 18 mm.

Localities: 29° 44'S. 31° 20’ E. (off Durban), 46 fath. (Flynn); off Gwayang

River, Mossel Bay, 31 fath. 1 damaged 9; off Tugela River mouth, 200 fath.

1 damaged 2; off Cape Natal, 54 fath. 2 ovig. 3d, 2 99 (S. Afr. Mus.).

Distribution. Off Dar-es-Salaam, 404 metres; Zanzibar area, 421-457 metres.

Remarks. All the South African Museum specimens are more or less

damaged.

Calman refers to several discrepancies in M6bius’s figures. Another

inaccuracy occurs in fig. 7 which shows an extra (tibial) joint in the 2nd legs.

Pallenopsis (Rigona) ovalis Loman

Fig. 15d

1908. Loman, loc. cit., p. 68, pl. 10, figs. 137,)138:

1923. Calman, loc. cit., p. 284, fig. 11.

1928. Flynn, loc. cit., p. 23.

1953. Stock, loc. cit., fig. 3 @ (chart).

Body unsegmented. Crurigers contiguous at their bases. A more or less

distinct median longitudinal rib on cephalon. Ocular tubercle short, conical,

SOUTH AFRICAN PYCNOGONIDA 121

anterior eyes scarcely larger than posterior pair. Proboscis cylindrical.

Abdomen pointing obliquely upwards or nearly vertical. Dorsal surface

glabrous.

Chelifer with scape undivided, hand as in intermedia, finger with a spinose

pad on basal half (cf. alcocki Calman, loc. cit., fig. 9 c). Palpal tubercles

distinct. Oviger in ¢ 5th joint slightly shorter than 4th, 6th slightly swollen,

10th joint as in intermedia.

Legs 1st coxa with 2 small spiniferous projections on upper apex, 2nd coxa

distinctly longer than 3rd, femur and 1st tibia subequal or the latter slightly

€

Fig. 17. Pallenopsis oscitans (Hoek). a. tarsus, propodus and claw of leg. 6. hand of chelifer.

c. distal joints of oviger @.

shorter, both with a small spiniferous process on upper apex, 2nd tibia longer

than femur or Ist tibia and more slender; propodus, claw and auxiliary claws

as in intermedia. Numerous short spine-setae on joints, and some longer ones

arising from small tubercles on ventral surface of femur conspicuous in

ovigerous <.

Body incl. abdomen 6, proboscis 3, femur 5°5, Ist tibia 5, 2nd tibia 6 mm.

Another specimen, of which the body is damaged, has femur 7:5, rst tibia 7,

and and tibia 8 mm.

Localities: 10 miles SE. of East London, 47 fath. 1 ovig. ¢; Algoa Bay, 25

fatto ous. atc. Mus):

Distribution. East Indies, Andaman Is., Ceylon.

122 ANNALS OF THE SOUTH AFRICAN MUSEUM

Pallenopsis oscitans (Hoek)

Fig. 17

1881. Hoek, Rep. H.M.S. ‘Challenger’, iii, p. 89, pl. 13, figs. 1-5. (Phoxichili-

dium o.)

Body segmented. Crurigers well separated (by about their own width),

each slightly longer than median width of body. Ocular tubercle low and

rounded, eyes not traceable. Proboscis cylindrical, slightly tapering distally,

apically rounded. Abdomen elongate, slender, slightly clavate, oblique. A

few fine and rather long setae on the crurigers and abdomen.

Chelifer with scape a little longer than proboscis, 2nd joint slightly shorter

than ist, hand nearly as long as 2nd joint, palm short, finger and thumb

distally slender, curved, when closed gaping, apices crossed. Palpal tubercles

moderately large but inconspicuous. Oviger in 9 5th joint a little longer than

4th, 6th not swollen, apical joint slender, with long fine setae.

Legs 2nd coxa subequal to ist + 3rd, femur very slightly longer than 1st

tibia, 2nd tibia longer than femur, propodus without heel but with a very

large spine basally, followed by smaller ones on lower margin, apex not pro-

jecting over claw, claw strong, reaching almost to base of propodus and

forming with the large spine on the latter a prehensile ‘chela’, auxiliary claws

short and weak. Fine setae on all joints, becoming more numerous on femur

and ist and 2nd tibiae but not forming dense fringes as in intermedia, nor a

thick fur.

Body incl. abdomen 11°5, abdomen 4:75, proboscis 5, 2nd leg 47 mm.

(2nd coxa 4, femur 11, Ist tibia 10, 2nd tibia 14 mm.).

Locality: off Gape Point, N. 86° E., 43 miles, go0-1,000 fath. 1 9 (S. Afr.

Mus.).

Distribution. 38° 25’ N., 35° 80’ W. (Azores), 1,675 fath.

Remarks. Although the proboscis is more like that of pzlosa as described by

Hoek, and the legs are somewhat more hairy, there is little doubt that this

specimen should be identified as oscztans; the apical joint of the oviger and the

propodus of the legs, with its enormous basal spine, correspond with Hoek’s

figures. The ocular tubercle shows no trace of eyes, but the specimen has

suffered desiccation.

Differs from longirostris Wilson in the apical joint of the oviger, and propodus;

and from tritonis Hoek (syn. holti Carp.) in the same features and also in the

relative lengths of palm and fingers of chelifer (see: Carpenter, Fish. Irel. Sct.

Invest., 1905, pl. 1, figs. 3, 4, 6, holti; I have not seen Hoek’s 1883 paper).

P. calcanea Steph. 1933 differs in having the ovigers closer together, a much

larger ocular tubercle, a shorter abdomen, a different apical joint on the

oviger, propodus of legs with 2 moderate-sized basal spines on a distinct heel,

and no auxiliary claws.

SOUTH AFRICAN PYCNOGONIDA 123

Pallenopsis sp. Gordon

1932. Gordon, loc. cit., p. 91, fig. 45.

A young specimen from 35° 14’ S., 6° 49’ E., pelagic, which Stephensen

(loc. cit., p. 24) suggests may be the same as his calcanea (loc. cit., p. 21, fig. 5)

from Greenland. Gordon’s record scarcely comes within the South African

area.

Gen. Hannonia Hoek

1881. Hoek, Rep. H.M.S. ‘Challenger’, iii, p. 92.

1891. Sars, Norw. N. Ail. Exp. Pycnogonida, p. 6.

1902. Pocock in Lankester, Encycl. Brit., 1oth ed. Arachnida (type of a

separate family).

1904. id., Q. 7. Micr. Sci., n.s. xlviii, p. 225 (reprint of 1902 article).

1904. Loman, Zool. Jahrb. Abt. Syst., xx, p. 385 (systematic position).

1905. Cole, Ann. Mag. Nat. Hist., (7) xv, pp. 408, 410 (systematic position).

1908. Loman, loc. cit., pp. 15, 16, synopsis facing p. 19 (systematic position).

1909. ‘Thompson, Cambr. Nat. Hist., iv, p. 533 (systematic position).

1927. Calman, Trans. Zool. Soc. Lond., xxii, p. 410 (systematic position).

1947. Hedgpeth, Smiths. Misc. Coll., cvi, p. 4. |

Fig. 18. Hannonia typica Hoek. a. dorsal view. 6. lateral view, with tubercle (? vestigial palp)

further enlarged. c. ocular tubercle of adult, from right side. d. chelifer of juvenile (above)

and adult (below). e. oviger g with distal joints further enlarged. f. spine from leg.

Body stout, segmented, crurigers narrowly separated. Proboscis on narrow

stalk, swollen (sac-like), deflexed and bent beneath body. Chelifers rudi-

mentary, 2-jointed, more or less chelate. Palps absent (see remarks). Ovigers

124 ANNALS OF THE SOUTH AFRICAN MUSEUM

in both sexes, 10-jointed, not modified in g, with apical claw. Legs stout,

tarsus very short, no auxiliary claws. Genital pores on last 2 legs in 4, on all

legs in 9. Eggs in a single cake-like mass held by both ovigers. Femoral

cement glands 3 seemingly absent.

Remarks. An endemic South African genus with one species, whose

systematic position has been the subject of much discussion.

The presence of the minute papilla, between the bases of the chelifers and

ovigers, discovered by Calman (1927) is herewith confirmed. I have not,

however, seen any indication in my specimens of its being 2-jointed. It is

present in both sexes, lies just below a small spiniferous process on the side

of the cephalic segment, and has a subapical spinule. It occurs in a juvenile

specimen, 2-5 mm. body length (as measured below), but is neither more nor

less feebly developed than in adults.

Calman’s suggestion that this papilla represents the last vestige of a palp

would seem to bring the genus into an intermediate position in the series

Béhmia-Rhynchothorax-Pycnogonum, where Loman in his conspectus (1908)

placed it. Calman did not accept Bouvier’s suggestion that Hannonia was an

Ammotheid. Hedgpeth (1947) places it in the Pallenidae.

Hannonia typica Hoek

Fig. 18

1691. Eloek; loc. Jeit..4p. 92, pli 14. fiesme-ise

1904. Loman, loc. cit., p. 383, pl. 14, figs. 12-15.

1910. Hodgson, Schultze. Reise, iv, p. 227.

1O23 ss Momany Aghw00l-..%V. Os Ong

1927. Calman, loc. cit., p. 410.

Body compact, the posterior margins of the segments forming transverse

arched ridges, each with a median boss. Crurigers separated by less than

half their own width, their length subequal to or a little longer than median

width of segments, each with a knob on dorsal apex. Ocular tubercle

moderately high, bluntly conical, eyes distinct. Proboscis very much swollen

beyond the comparatively narrow basal stalk. Abdomen extending beyond

last crurigers, clavate, bent slightly downwards. Short spinules on the trans-

verse ridges and bosses, crurigers, abdomen, top of ocular tubercle, and front

and sides of cephalic segment.

Chelifers never quite vestigial, but variable, the chela sometimes well

developed and apparently with mobile finger. Ovigers similar in both sexes,

but in ¢ the distal joints rather stouter and more spinose, the spines minutely

setulose.

Legs stout, 2nd coxa slightly longer than either ist or 3rd, on last 2 legs in 3

with a process on ventral apex, more conspicuous on 3rd than on 4th leg,

femur slightly longer than either of the tibiae, which are subequal, tarsus

narrower than 2nd tibia, propodus without heel, with numerous small but no

SOUTH AFRICAN PYCNOGONIDA 125

strong spines on lower margin, claw about one-third, or a little less, length of

propodus. Numerous spines on the joints, arising from slightly raised tubercles,

particularly distinct on dorsal surface of 2nd tibia, and especially in juveniles.

Femoral cement glands § seem to be absent; neither Loman (1904, p. 384)

nor myself have found any.

Body (base of chelifers to base of abdomen) 5, proboscis 4, abdomen 2, 2nd

leg femur 2-5, 1st and 2nd tibiae 2 mm.

Localities: Cape Town (Hoek, Loman); Liideritzbucht (Hodgson); Port

Natal (Durban) (Loman); Saldanha Bay and Melkbos Strand (Table Bay)

littoral, Mossel Bay, 20 fath. (S. Afr. Mus.).

Remarks. Not nearly so common as Discoarachne brevipes; in the course of a

considerable amount of shore-collecting at Sea Point (Table Bay) I have

never found a specimen there, although on the other side of Table Bay (Melk-

bos Strand) it is moderately common.

Two ovigerous $4 were collected at Saldanha Bay in September.

Fam. PHOXICHILIDIIDAE

1908. Loman, Szboga Exp. monogr., xl, p. 62, conspectus facing p. 19.

(Phoxichilidae.)

1913. Schimkewitsch, Zool. Anz., xli, p. 611.

1947. Hedgpeth, Smiths. Misc. Coll., cvi, p. 4.

Octopodous. Body segments free or fused. Chelifers well developed,

chelate. Palps rudimentary, reduced to tubercules or absent. Ovigers in 3

only, 5-9-jointed, with simple spines. Legs usually with auxiliary claws.

Key to South African Genera

1. Cephalic segment short, without neck. Oviger 5-jointed. Phoxichilidium

2. Cephalic segment with distinct neck; ocular tubercle near front margin.

Oviger 6-jointed. Anoplodactylus

Gen. Phoxichilidium M.-Edw.

1836. Johnston, Miscell. Zool., i, Mag. Zool. & Bot., i, p. 378. (Orithyia,

preocc. Fabr. 1708.)

1840. Milne-Edwards, Hist. Nat. Crust., (Roret’s Suite a Buffon. Crust.),

lll, Pp. 535.

1881. Hoek, Rep. H.M.S. ‘Challenger’, iii, p. 31 (part).

1908. Loman, Szboga Exp. monogr., xl. pp. 63, 64.

* Body segments free. Ocular tubercle on anterior portion of cephalic segment,

in advance of Ist crurigers. Chelifers with scape undivided, finger and thumb

gaping when closed. Palps completely absent. Ovigers ($ only) 5-jointed.

Legs with minute auxiliary claws. Genital pores on last 2 legs in 3, on all

legs in 9.

126 ANNALS OF THE SOUTH AFRICAN MUSEUM

Phoxichilidium capense Flynn.

1928. Flynn, loc. cit., p. 27, figs. 15, 16.

Body stout, especially the cephalic segment and ist pair of crurigers.

Crurigers narrowly separated (the 3rd and 4th pairs more widely separated).

Ocular tubercle obtusely conical, eyes distinct. Abdomen rather short and

stout, erect. Proboscis stout, apically truncate.

Chelifer rather stout, as long as proboscis, hand (palm) longer than broad,

finger and thumb curved, inner margins entire.

Legs (2) stout, 2nd coxa with strong conical projection (bearing the genital

pores) on ventral apex, femur longer than either of the tibiae, which are sub-

equal (text; according to the measurements and figure the 1st tibia is slightly

the longer), propodus with strong spines on heel, lower margin with about 12

spines, claw strong, auxiliary claws minute.

Body 3°75, proboscis 3:18, femur 5:17, Ist tibia 4°31, 2nd tibia 3-75 mm.

(Flynn).

Locality: Hout Bay, west coast of Cape Peninsula, depth not given (Flynn).

Known from a single 9.

Gen. Anoplodactylus Wilson

1821. Say, 7. Ac. Nat. Sct. Philad., ii, p. 59. (Anaphia.)

1878. Wilson, Amer. 7. Sct. (3), xv, p. 200.

1908. Loman, loc. cit., p. 71.

1908. Norman, 7. Linn. Soc. Lond., xxx, p. 202. (Anaphia.)

1912. Loman, Bull. Inst. ocean. Monaco, no. 238, p. 7. (subgen. Halosoma

Cole).

1923. Calman, Rec. Ind. Mus., xxv, p. 285.

1927. id., Trans. Zool. Soc. Lond., xxii, pp. 405, 407.

Body segments free. Ocular tubercle on front margin of cephalic segment

overhanging base of proboscis. Chelifer with scape undivided, finger and

thumb either stout and meeting when closed, or slender and gaping. Palps

completely absent. Oviger 6-jointed, ultimate and penultimate joints feebly

developed, without strong spines, no apical claw. Legs with or without

auxiliary claws. Genital pores on last 2 legs in 4g, on all legs in Q (but see

Calman, 1927). Femoral cement glands ¢ on femur usually single, tubular

(numerous and cribriform in cribellatus Calman, 1923).

Remarks. Wilson’s genus is said to be the same as Anaphia Say, but this

synonymy has not been generally adopted.

Mobius’s South African species spinosus has been transferred to Parapellene.

Key to the South African Species

1. Hand of chelifer stout, finger and thumb short, meeting when closed. aculeatus

2. Hand of chelifer slender, finger and thumb long, slender, gaping when closed. _pelagicus

SOUTH AFRICAN PYCNOGONIDA | edly |

Anoplodactylus aculeatus Mobius

1902. Mébius, D. Tiefsee Exp., ili, p. 188, pl. 28, figs. 1-7.

Body slender. Crurigers separated by intervals not greater than their width,

their length subequal to median width of segments. Ocular tubercle conical,

sharply pointed, eyes distinct. Abdomen short, vertical.

Chelifer scape stout, longer than proboscis, with conical process on dorsal

apex, hand stout, finger and thumb short, stout, meeting when closed.

Fig. 19. Anoplodactylus pelagicus Flynn. a. dorsal view. 6. lateral view of cephalic segment.

c. hand of chelifer. d. 3rd-6th joints of oviger g with 5th and 6th joints further enlarged.

e. 2nd coxa of leg with genital pore. f. cement gland on femur. g. tarsus, propodus and claw

of leg.

Legs long, 2nd and 3rd coxae with a spiniform process on ventral apex,

femur longer than ist tibia, both joints with a spiniform process on dorsal apex,

ist tibia also with a spine at end of basal third of its length, 2nd tibia longer

than femur (subequal in fig. 1), sparsely setose, propodus with 4 large spines

on basal heel and 8 smaller ones on lower margin, claw strong, no auxiliary

claws. ;

Body, excl. abdomen, about 8 mm. (according to Mébius’s fig. 1).

128 ANNALS OF THE SOUTH AFRICAN MUSEUM

Locality: Agulhas Bank, 126 metres, 2 99 (Mobius).

Remarks. ‘This species, based on the female only, is very like insignis (Hoek)

1881, based on the g, from Bahia, S. America. Both have projections on the

chelifer scape, 2nd and 3rd coxae, femur and Ist tibia; but insignis has also a

process on the Ist coxa, and the crurigers are a little more widely separated.

Cf. also insignis subsp. bermudensis Cole (1904. Proc. Boston. Soc. Nat. Hist., xxxi,

p- 325, pl. 20, and pl. 22, figs. 21-29).

Anoplodactylus pelagicus Flynn

Fig. 19

1920. Flynn; loc, cits, ps25, fie. 4e

Body slender. Crurigers separated by intervals greater than their width,

their length greater than median width of segments. Ocular tubercle low and

flattened or rounded on top, eyes small, unpigmented. Abdomen not very

long, erect.

Chelifer scape slender, extending a little beyond tip of proboscis, hand with

long slender finger and thumb, curved and gaping when closed, inner margins

with a few spine-setae. Oviger 3rd joint longer than 2nd, both elongate, 5th

somewhat swollen, with a patch of setae distally and 3 small recurved denticles

on inner margin, 6th shorter than 5th, ovate, setose on inner margin.

Legs long, slender, ventral apex (with genital pore) of 2nd coxa of last 2

legs ¢ slightly more prominent than on first 2 legs; femur and 1st and 2nd

tibiae subequal, 2nd tibia in the $.A.M. specimen not so much more slender

than Ist tibia as is shown in Flynn’s figure, and the propodus not so noticeably

thicker, 2 strong spines on basal heel and about 18 small spines on inner

margin, claw strong, auxiliary claws very small (scarcely visible except when

claw is seen in dorsal or ventral view); tibiae finely and sparsely setose.

Cement gland on dorsal surface of femur tubular, not very long.

Body, excl. abdomen, ¢g 2°5, proboscis 1-5, femur (also 1st and 2nd tibiae)

2:75 mm. Flynn’s measurements are slightly greater, except the leg measure-

ments of ¢.

Localities: off Port Natal (Durban), pelagic (Flynn); Cape Point NE. X

E 2 E. 28 miles, 300 fath. 1 ¢ (S. Afr. Mus.).

Remarks. Flynn refers (loc. cit., p. 3) to the capture of 7 specimens in a tow-

net, but in the absence of further data it does not follow that they were from

the surface or even pelagic: the tow-net may have been attached to the beam

of the trawl. The single specimen in the South African Museum was taken

together with “‘Alcyonarians’ (s.s. Preter Faure log-book, P.F. no. 18159).

‘Pallene’ lappa Bohm

1879. Bohm, MB. K. Ak. Wiss. Berlin, p. 182. Ibo, Mozambique.

1881. Hoek, loc. cit, p. 31.

SOUTH AFRICAN PYCNOGONIDA 129

1910. Hodgson, loc. cit., p. 225.

1912. Loman, Bull. Inst. ocean. Monaco, no. 238, p. 6.

1926. Flynn, loc. cit. p. 4.

Hoek and Hodgson considered that B6hm’s specimen was immature, with

not fully developed ovigers. Loman, with whom Flynn agrees, included it in

Halosoma, a subgenus of Anoplodactylus. Loman gives as one of the characters

of Halosoma: ovigers 6-jointed; but Flynn says B6hm’s specimen has 7-jointed

ovigers.

The locality is, strictly speaking, outside (12° S.) the South African area,

but the record is included here as Flynn has referred to it.

Fam. ENDEIDAE

1908. Norman, 7. Linn. Soc. Lond., xxx, p. 231.

1932. Gordon, ‘Discovery’ Rep., iii, p. 93.

See generic characters.

Gen. Endeis Phil.

1843. Philippi, Arch. Naturg. Jahrg., ix, p. 175.

1902. Mobius, D. Tiefsee Exp., iii, p. 195. (Phoxichilus, non Latr.)

1902. Stebbing, Knowledge, xxv, p. 187. (Chilophoxus.)

1908. Loman, Siboga Exp. monogr., xl, p. 77. (Phoxichilus, non Latr.)

1908. Norman, loc. cit., p. 231.

1915. Calman, ‘Terra Nova’ Exp. Rep., iii, p. 48.

no2gid-. iec. Ind. Mus., xxv, p. 200:

Octopodous. Body segmented.. No chelifers. No palps. Ovigers only in J,

7-8-jointed, without apical claw, with simple spines. Legs with auxiliary claws.

Key to the South African Species

1. Crurigers narrowly separated. Propodus of legs apically produced over base of

claw. clipeatus

2. Crurigers well separated. Propodus of legs not apically produced. mollis

Endets clipeatus (Mo6bius)

Fig. 20

1902. Mobius, loc. cit., p. 196, pl. 30, figs. 6-10. (Phoxichilus clipeatus [sic];

clypeatus on plate.)

1920. 5 Elynn, loc. cit., p. 20:

Body rather stout, cephalic segment with front margin feebly bilobed, with

a minute chitinous point on each lobe in the ¢ specimen (possibly representing

the remnants of the chelifers). Crurigers narrowly separated, about as long as

median width of segments. Ocular tubercle short, conical. Abdomen about

as long as last crurigers, apically notched. Proboscis stout, swollen in middle,

apex truncate. Body and crurigers glabrous. Whole surface of body and

appendages closely and minutely ‘pitted’ (? glandular).

130 ANNALS OF THE SOUTH AFRICAN MUSEUM

Oviger 8-jointed, but articulation between 7th and 8th joints obscure,

6th joint with a variable number of more or less recurved spines on inner

margin.

Legs, femur and 2nd tibia subequal, 1st tibia slightly shorter, propodus

apically produced in a cylindrical process over base of the claw, claw and

auxiliary claws strong. Glabrous. Femoral cement glands about 40 in a

single row.

a F b

Fig. 20. Endeis clipeatus (Mobius). a. tarsus, propodus and claw of leg. b. oviger 3. c. 6th and

7th—8th joints of same further enlarged.

Body (3) incl. abdomen 4, proboscis 3, 2nd leg 20 mm. (femur 5, Ist tibia

4°25, 2nd tibia 5:25 mm.). Yellowish, a red line down middle of body and red

longitudinal lines on legs.

Localities: St. Francis Bay, shallow water (Mébius); off west coast of Cape

Peninsula (Flynn); St. James, False Bay, littoral. 1 juv.; west coast of Cape

Peninsula, littoral; False Bay, 10 fath. and Algoa Bay, 52-63 fath. 2 99,1 ¢

(S. Afr. Mus.). |

Endeis mollis (Carp.)

Rig om

1894. Carpenter in Herdman’s Ceylon Pearl Fish. Rep. Suppl. Rep., xiii, p. 182,

pl. figs. 1-7. :

1907. id., Trans. Linn. Soc. Lond. (2) zool. xii, p. 98.

1923. Calman, loc. cit., p;,203,, fig.. 16.

1927. id., Trans. Kool. Soc. Lond., xxii, p. 408.

1938. id., John Murray Exp., v. p, 160.

1951. Stock, Mem. Inst. Roy. Sct. Nat. Belge. (2) fasc. 43, p. 17, figs. 23, 24.

SOUTH AFRICAN PYCNOGONIDA 131

Body rather slender, cephalic segment with 2 rounded lobes in front over

base of proboscis (possibly representing the chelifers). Crurigers separated by

more than their own width, slightly longer than median width of segments.

Ocular tubercle short, conical. Abdomen slightly longer than last crurigers,

oblique. Proboscis stout, swollen in middle, apex truncate. Body and crurigers

glabrous. Whole surface of body and appendages closely ‘pitted’ (? glandular).

Fig. 21. Endeis mollis (Carp.). a. dorsal view. 6. inner view of 4th—7th joints of oviger 3.

c. outer view of 6th joint of same. d. propodus and claw of leg.

Oviger 7-jointed, on a basal knob, penultimate joint swollen, inner margin

on inner side (as coiled up in situ) with 2 recurved spinules, outer surface with

a broad lobe bearing 3 spinules, apical joint with 1 spinule on inner margin

and 2-3 on apex.

Legs, femur slightly longer than Ist tibia, smooth, cylindrical, with one

fairly strong spine-seta and 2 smaller setae on dorsal apex, 2nd tibia slightly

longer than femur, 1st and 2nd tibiae with a few scattered setae, propodus

apically truncate, not produced over base of claw. Femoral cement glands

about 23, in a single row.

Body (tip of proboscis to tip of abdomen) ¢ 7:5, 2nd leg 18 mm. (femur

4°5, Ist tibia 4, 2nd tibia 4-75 mm.).

132 ANNALS OF THE SOUTH AFRICAN MUSEUM

Locality: off Durnford Point, Zululand, 13 fath. 1 non-ovig. 3g, 1 9 (S. Afr.

Mus.).

Distribution. Ceylon, Maldives, Arabian coast, Indian coasts, Nicobars,

Christmas Island, Tonga Is.

Remarks. Apparently resembles the typical form. Calman records some

variations in the shape of the femur, number of cement glands, etc.

Fam. AMMOTHEIDAE

1908. Loman, Szboga Exp. monogr., xl, conspectus facing p. 19 and

Pp. 49:

1909. ‘Thompson, Cambr. Nat. Hist., iv, p. 534.

1913. Schimkewitsch, Zool. Anz., xli, p. 612.

1947. Hedgpeth, Smiths. Misc. Coll., cvi, p. 4.

Octopodous. Body extended or compact, segmented or the segmentation

more or less obsolete. Proboscis more or less movable, directed forwards or

downwards. Chelifers reduced, chelate in juvenile, but rudimentary in adult.

Palps 6-10-jointed (Nymphonella 17-20). Ovigers in both sexes, (9)-10-jointed,

with or without apical claw, with simple or serrate spines. Legs usually with

auxiliary claws. Genital pores on all legs in 9, on last 2 legs in g. Eggs in

several masses. |

Key to the South African Genera

I. Proboscis fusiform or subcylindrical, directed forwards or downwards.

A. Palp 7-jointed. Béohmia

B. Palp 8-jointed. Achelia

C. Palp g-jointed.

1. All body segments fused. Crurigers narrowly separated. Nymphopsis

2. Body segments, at least the anterior ones, distinct. Crurigers widely

separated.

a. Legs with auxiliary claws. Ocular tubercle very elongate. Kyphomia

6b. Legs without auxiliary claws. Ainigma

II. Proboscis flask-shaped, curving downwards. Austroraptus

Gen. Béhmia Hoek

1881. Hoek, Rep. H.M.S. ‘Challenger’, ii, p. 24.

1902. Mobius, D. Tiefsee Exp., iii, p. 189.

1908. Loman, loc. cit., conspectus facing p. 19.

1947. Hedgpeth, loc. cit., p. 4 (in list of genera).

Body stout, segmented, arched. Crurigers narrowly separated. Proboscis

elongate, conical, inserted ventrally and bent under cephalic segment.

Chelifers rudimentary, 2-jointed, chelate, inserted ventrally and invisible from

above. Palps 7-jointed. Ovigers 10-jointed, with apical claw, not modified

in g. Legs stout, tarsus very short, no auxiliary claws. Genital pores on all

legs in 9, on last leg in g. Pale non-spinose (or almost so) areas on dorsal

surface of all femora in g. Eggs carried in a single cake-like mass.

SOUTH AFRICAN PYCNOGONIDA 133

Remarks. An endemic South African genus, with two species.

Flynn (loc. cit.) claimed to have found genital pores on the dorsal surface

of the 2nd coxa of all the legs; this position would be distinctly unusual, in

fact unique, if it were correct. My own observations disprove this; the genital

pores are in the normal ventral position, and on the last leg only (as in

Pycnogonum).

Key to the South African Species

1. Smaller species. Legs not more than twice as long as body. Tubercles distinct, but

moderate. Cephalic segment parallel-sided or widening in front. Femur 2 (2) or

3 (g) times as long as wide. chelata

2. Larger species. Legs about four times as long as body. Cephalic segment narrowing

in front. Femur (¢) four and a half to five times as long as broad. tuberosa

Béhmia chelata (Bohm)

Higes22

1879. Bohm, MB. Ak. Wiss. Berlin, p. 192, pl. 2, figs. 5-5d. (Pycnogonum c.)

Roar © Eloek, loc. cit., p. 24.

1902. Mobius, loc. cit., p. 189, pl. 28, fig. 15 (front of cephalic segment).

1909. Schimkewitsch, Xool. Anz., xxxiv, p. 3, fig. 1 A and C (chelifer and

palp).

1915. Calman. ‘Terra Nova’ Exp. Rep. ool., iii, p. 13 (oviger mentioned).

ig208 2 blynun, loc. cit., p. 30, fig. 17.

Body strongly arched, median portion of each segment raised into a rounded

boss, similar to the ocular tubercle, the boss on 4th segment smaller than the

others, all sharper and more pronounced in ¢ than 92; cephalic segment

nearly parallel-sided in 2 with somewhat prominent rounded antero-lateral

corners, in ¢ sides diverging forwards, antero-lateral corners prominently and

subacutely produced. Crurigers narrowly separated, the last pair a little more

widely separated; dorsal apices tending, especially in 4g, to form small

nodiform bosses. Ocular tubercle conical apically rounded in Q, subacute in 4,

eyes not conspicuous. Proboscis inserted ventrally, base wide, quickly tapering

to a subacute apex, projecting downwards or bent under cephalic segment.

Abdomen projecting beyond last crurigers, clavate, deflexed. Whole surface

reticulate, with short spinules arising from conical bases, including ocular

tubercle.

Chelifer scape stout, spinose, chela rather slender, but well developed and

apparently functional (finger mobile). Palp 4th joint much the longest, 5th

small, 6th and 7th rather slender, more so in ¢ than Q, spinose.

Oviger 5th joint longer than 4th, last 4 joints with rather strong spines on

inner margins, mostly in pairs, resp. about 8, 5, 4, 5 pairs, sometimes fewer

in g, inner apex of 10th joint in g with a single unguiform process resembling

but smaller than the apical claw; alike in both sexes except for this latter

feature.

Legs stout, stouter in 9 than 3, decreasing in length backwards, the 3 coxae

subequal in length, but 2nd somewhat stouter in 2 owing to the genital pores,

134. ANNALS OF THE SOUTH AFRICAN MUSEUM

on last leg in § a little longer than either of the other two; femur longer than

1st tibia on all legs in 9, subequal to it in g or on the Ist leg slightly shorter,

about twice as long as wide in 9, about 3 times in g, 2nd tibia subequal to

femur in Q, slightly longer in 4, tarsus and propodus together a little shorter

than Ist tibia, propodus without heel, with several short but stout spines along

lower margin, claw about half length of propodus. Surface reticulate like that

of the body, and spinulose. At dorsal apex of 2nd coxa of all legs in both sexes

Fig. 22. Béhmia chelata (B6hm). a. abnormal double ocular tubercle. 6. chelifer. c. palp.

d. oviger g with apex of roth joint and claw further enlarged. e. apex of 10th joint and claw of

oviger 2 f. dorsal apex of 2nd coxa ($9) of leg. g. ventral apex of 2nd coxa of last leg ¢.

h. dorsal surface of femur of 3 with pale patch (‘cement gland’).

a small pale spot. Femoral cement gland in ¢ an elongate oval pale spot on

dorsal surface on each leg.

Eggs in a single cake-like mass.

Body (from front margin of cephalic segment to base of abdomen) g@ 5:5,

proboscis 2°5, abdomen 3 2, @ 1°5, 1st leg (without claw) approx. g 10,

2 8-5 mm. Amber-coloured, or reddish-brown, sometimes bicoloured dark

and light, with dark bands on legs (see under Remarks).

Localities: (loc. ? Bohm); St. Francis Bay, shallow water (Mobius); off

Ball Point, Cape Town (Flynn); False Bay, littoral to 12 fath.; off Cape St.

Blaize, off Knysna, and off Cape St. Francis, 25-70 fath; off Gt. Fish Point,

Keiskamma, East London and Cape Morgan, 33-77 fath.; off Umtwalumi and

Umhloti River mouths (Natal), 25-40 fath.; off Tugela River and Durnford

Point (Zululand), 13-47 fath. 3¢, 29, juv. (S. Afr. Mus.).

SOUTH AFRICAN PYCNOGONIDA 135

Remarks. On two points my observations differ from those of Flynn. The

more important is the position of the § genital pores. Although Flynn had

both gg and 99 he did not notice that the pale spot on dorsal apex of 2nd coxa

occurred in both sexes. It is obviously not a genital pore. When the integument

of this joint is treated with a clearing reagent the pale spot is seen to be due to

feeble chitinization; elsewhere the integument is strongly but not uniformly

chitinized, producing a reticulate pattern (fig. 22f). Further proof that these

pale spots are not openings is shown by the fact that spines may be present on

them as elsewhere on the integument.

The true genital pore in g is a very definite opening, in the normal position

on the ventral surface of the 2nd coxa, and it occurs only on the last leg.

On the upper surface of the femur of all the legs is another pale patch, oval

in shape and very conspicuous, but which occurs only in the g. Flynn calls

this the cement gland, and it probably is; but I have failed to find the longitu-

dinal slit-like opening described by him (cf. also under tuberosa). The paleness of

the patch is due to lack of chitinization, as in the case of the spot on 2nd coxa;

and there are minute spinules and even a few spines on its surface, if not

centrally at least marginally. Some small circular structures (fig. 22h) may be

the actual cement glands, but I cannot detect any individual or communal

pores. P

The chelate termination of the oviger was mentioned incidentally by Calman

(1915), but he did not state that it is found in the male only.

This species is common on branching Alcyonarians (Gorgonia, Melitodes,

Spongioderma, Villogorgia, Wrightella, etc.) but I have also found it in the littoral

zone among Hydroids and Polyzoa.

With such a typically warm-water distribution (Zululand southwards),

Flynn’s record ‘Ball Point, Cape Town’ is a little surprising, if this locality ©

really is in Table Bay. I have not been able to trace Ball Point on any chart.

I have not taken Béhmia chelata either at Sea Point or at Melkbos Strand

(both in Table Bay), but it is fairly common at low tide at St. James, False

Bay. It is, however, a more frequent inhabitant of Alcyonarians, and nearly

every piece of ‘Fan-coral’ (Gorgonia flammea) which I have examined in a fresh

state, has one or more of these little Pycnogonids clasping its branches.

Coloration. As a rule the animal is uniformly coloured, irrespective of the

colour (white, yellow, pink, red) of the Alcyonarian on which it is found.

Two lots, however, from the Keiskamma-Cape Morgan area are bicoloured

and very striking. The ground colour is the usual amber or yellowish-brown

with dark umber-brown on parts of the body and legs as follows: on the body

the dark colour occupies the whole upper surface except the antero-lateral

corners of the cephalic segment, ocular tubercle, and a patch behind it, the

4 dorsal tubercles, and the apical bosses on the crurigers; the abdomen has a

dark streak on each side; on the legs the 1st coxa is dark, dorsally only in

juvenile, but also ventrally in adult. The 2nd coxa is dark dorsally with a

slight suffusion ventrally in adult; the femur and Ist tibia each with a dark

136 ANNALS OF THE SOUTH AFRICAN MUSEUM

band in distal half on dorsal surface only. The pale spots on the 2nd coxae

($2) and the cement gland (¢) remain pale.

One of these lots is recorded as being found on a ‘yellow Alcyonarian’, and

the piece preserved with the Pycnogonid is Wrightella fragilis. In the preserved

state this Alcyonarian is white with the nodes yellow or brownish, but whether

the two colours are distinct in life is not recorded.

Aberration. One ¢ (Somerset Strand, False Bay) has the ocular tubercle

divided into two tubercles placed transversely; one of the eyes seems to be

se pre nerqh CHAM abyAMEEUL At 4 MMe phe veWeiiertyatyye geen

SUE ECT SP PEE er pre iyrtre ties

Fig. 23. Béhmia tuberosa Mobius. Dorsal surface of femur of g showing pale patch (‘cement

gland’), with a portion of the latter further enlarged; semidiagrammatic, minor reticulation

omitted.

present, but the others could not be traced. For rarity of abnormalities in

Pycnogonids see Gordon, 1932. ‘Discovery’, Rep., vi, p. 131.

Bohmia tuberosa Mobius

Fig. 23

19025 J Mobius: loc) cit... p. 160, ple 2o esanlo ml Ae

1909. Schimkewitsch, loc. cit., p. 3, fig. 1 B and D (chelifer and palp).

g—similar to chelata, but the ocular tubercle, antero-lateral processes of

cephalic segment, median bosses on segments and those on apices of crurigers

all very prominent and sharply conical, the median boss on 4th segment lower

than the others. Sides of cephalic segment somewhat convergent. Surface

reticulate and spinulose as in chelata.

Chelifer and palp as in chelata, but in the latter the 7th joint is shorter than

the 6th. Oviger as in chelata, but the pairs of spines on last 4 joints more

numerous, resp. about 10, 8, 6,9; and roth joint without an unguiform process

on lower apex.

SOUTH AFRICAN PYCNOGONIDA 137

Legs longer and more slender than in chelata; 2nd coxa a little longer than

either 1st or 3rd, femur a little longer than Ist tibia, about five times as long

as broad, 2nd tibia a little longer than femur, rather slender and tapering so

that tarsus is very little narrower than its apex, tarsus and propodus two and a

half to three times in length of and tibia, propodus without heel, with numerous

spines on lower margin, claw a little more than one-third (but not quite half)

length of propodus (Mobius, fig. 14: half length of propodus). Surface

reticulate and spinulose, with pale spot on dorsal apex of 2nd coxa of all legs.

Femoral cement gland an elongate-oval pale patch on dorsal surface of femur

in all legs. Genital pore on ventral apex of 2nd coxa of last leg only.

Body (front margin of cephalic segment to base of abdomen) 6, proboscis 4,

abdomen 3, 2nd leg (without claw) approx. 23 mm. (femur 6, Ist tibia 5,

and tibia 7 mm.). Reddish-brown.

Localities: Agulhas Bank, 154 metres 1 g (Mobius); off East London,

250-300 fath. 1 non-ovig. ¢ (S. Afr. Mus.).

Remarks. The ‘Valdivia’ specimen was a ¢ (dorsal patches) 7 mm. in length

including abdomen. There is an obvious discrepancy in the length of the claw

on the legs in Mobius’s two figures; as fig. 14 shows a claw half length of

propodus and therefore not so very different from that of the specimen above

described, one must regard the long slender claws in the picture of the whole

animal (fig. 13) as fanciful.

With a body length of 7 mm. incl. abdomen, the 2nd leg of the ‘Valdivia’

specimen was 13 mm. long; the present specimen measuring 9 mm. incl.

abdomen has the 2nd leg 23 mm. long.

The remarks on the pale coxal spot and femoral patches and the position

of the ¢ genital pore made under chelata apply here also. With the greater

length of femur, the femoral patch is more elongate. The patch itself is paler

than the rest of the integument, but there is a still paler line down the centre

of it, as seen under a moderate magnification. This is not due to a slit-like

opening, but to a double series of minute transverse ‘marks’ (fig. 23). Their

real structure is doubtful, but they appear to be in the substance of the chitin

as they are not obliterated by rubbing a needle-point over them either on

the external or internal surface. (Schimkewitsch, loc. cit., p. 4, called them:

Internal trabeculae.) Scattered over the pale area are a number of minute

‘glands’, smalier and more numerous than those in chelata. These can be

removed by rubbing the internal surface.

Gen. Achelia Hodge.

? 1838. Costa, Fauna Regn. Napoli. Arachn. Trach., p. 7. (Phanodemus.)

1864. Hodge, Rep. Brit. Assoc., xxxii, Not. misc., p. 102 and Ann. Mag.

Nat. Hist. (3) xiii, p. 114.

1881. Hoek, Rep. H.M.S. ‘Challenger’, iii, p. 26.

1881. Dohrn, F. Fl. Golf. Neapel., iii, pp. 133, 225, 227, 228 (Ammothea, non

Leach).

138 ANNALS OF THE SOUTH AFRICAN MUSEUM

1891. Sars, Norw. N. Atl. Exp. Pycnogonidae, p. 120. (Ammothea, non Leach.)

1908. Loman, Szboga Exp. monogr., xl, pp. 10, 11 and synopsis facing p. 19.

(Ammothea, non Leach.)

1913. Bouvier, 2me Exp. Antarct. Franc., pp. 45, 46, 138.

1915. Calman, ‘Terra Nova’ Exp. Zool., ii, p. 56.

1917. Bouvier, Res. Sct. Camp. Monaco, li, p. 38. (Ammothea subg. Achelia.)

1927. Hodgson, D. Stidpol Exp., xix (zool. xi), p. 344.

1932. Gordon, ‘Discovery’ Rep., vi, p. 110.

1938. Calman, John Murray Exp., v, p. 160.

1938. Gordon, Austral. Antarct. Exp., C II, 8, p. 22.

Body compact, segmented, but segments more or less fused. Crurigers

contiguous or narrowly separated. Ocular tubercle near front margin of

cephalic segment. Proboscis large, fusiform, directed forwards. Chelifer with

single-jointed scape, and terminal more or less bifid joint, which in juvenile

forms a small but functional chela. Palp 8-jointed, more or less geniculate,

and and 4th joints longest. Oviger 10-jointed, in both sexes, 7th-1oth joints

each with 1 (or 2) pinnate or doubly-serrate spines, without apical claw.

Legs moderate, tarsus small, propodus strongly spinose on lower margin,

claw strong, auxiliary claws moderate or strong. Genital pores on all legs in 9,

on last two legs in @, at apices of conical processes. Femoral cement gland ¢

single, opening near dorsal apex.

Remarks. Since Leionympha Mobius was shown to be the same as Ammothea

Leach, the small, compact, shallow-water species formerly included in the

latter genus, are now put into Achelia. Bouvier (1913 and 1917), however,

uttered a warning against regarding this name as stable, suggesting (as did

Dohrn) that Phanodemus was the earliest generic name for these forms.

Key to the South African Species

1. Abdomen extending beyond last crurigers. quadridentata

2. Abdomen not extending beyond last crurigers brevicauda

Achelia quadridentata (Hodgson)

Fig. 24

1910. Hodgson, Schultze. Reise, iv, p. 223, figs. 2, 3 (palp, oviger).

Body compact, widest in front, narrowing posteriorly, cephalic segment

large, segments 3 and 4 fused, and abdomen apparently not marked off by a

suture. Crurigers contiguous, each with a conical tubercle on upper apex.

Proboscis as long as body with abdomen, broadly fusiform. Ocular tubercle

on front margin of cephalic segment, conical, eyes distinct. Abdomen extending

to end of 1st coxa of last leg.

Chelifer scape undivided, the terminal joint representing the chela feebly

bifid. Palp geniculate, 2nd and 4th joints longest, distal joints setose. Oviger

SOUTH AFRICAN PYCNOGONIDA 139

5th joint longest, 7th, 8th and gth joints each with one doubly-serrate (pinnate)

spine, 10th joint with 2 similar spines, one laterally and one apical; alike in

the two sexes, but that of 3 rather stronger.

Legs without processes, 1st coxa with a small setiferous tubercle on posterior

part of distal margin dorsally, with a smaller one below it, propodus with 3

strong spines proximally on lower margin, and 2 moderately strong ones

distally, auxiliary claws about half length of main claw. Genital pores ¢ at

apices of rounded processes on ventral apices of 2nd coxae of last 2 legs; in 9

Fig. 24. Achelia quadridentata (Hodgson). a. dorsal view (4th joint of palp foreshortened).

b. lateral view of 2nd coxa of last leg g. c. apex of femur with cement gland g. d. propodus and

claws of leg. e. oviger g with distal joints further enlarged.

on all legs, but not on projections. Femoral cement gland 3 small, behind the

dorsal apical spine.

Body incl. abdomen 1-5, proboscis 1-5 mm.

Localities: Liideritzbucht (Hodgson); St. James, False Bay, littoral. 1 ovig.

6, 1 9 (K. H. Barnard, 1912); Table Bay, 5 fath. (University Cape Town

Ecol. Surv.).

Remarks. ‘The above description of the St. James’s specimens covers that

given by Hodgson, and I think the identification of these specimens is correct.

Hodgson’s specimens had 4 proximal spines on the propodus; the present

specimens have only 3.

14.0 ANNALS OF THE SOUTH AFRICAN MUSEUM

Achelia brevicauda (Loman)

1904. Loman, Zool. Fahrb. Abt. Syst., xx, p. 376, pl. 14, figs. 1-4 (3)

Body compact, widest in front (but not so wide as in quadridentata), antero-

lateral angles of cephalic segment subacutely angular (Loman: a strong

conical tubercle), segments 3 and 4 fused, abdomen not marked off by a

suture. Crurigers contiguous, a double setiferous tubercle (or two separate

tubercles) at each antero-distal and postero-distal corner dorsally. Proboscis

nearly as long as body, fusiform. Ocular tubercle on front margin of cephalic

segment, rounded or bluntly pointed apically, ies distinct. Abdomen short,

extending to end of last crurigers.

Chelifer, palp and oviger as in quadridentata (cf. fig. 24). The 6th joint of

oviger has a strong recurved spine proximally, as in A. Jangi Dohrn (1881.

pl. 5, fig. 2).

Fig. 25. Nymphopsis cuspidata (Hodgson).

Legs without prominent processes, Ist coxa with 2-3 setiferous conical

tubercles on both antero- and postero-distal corners dorsally, distal corners

of 2nd and 3rd coxae each with 1-2 much smaller and inconspicuous tubercles,

femur with 2 major spines and an apical one on upper margin, propodus with

3 strong spines on lower margin proximally and 2-3 less strong ones distally,

auxiliary claws about half length of main claw. Genital pores ¢ on prominent

ventral processes on last 2 legs. Femoral cement gland ¢ small, opening behind

the apical spine (cf. fig. 24 c).

Body incl. abdomen 1-25, proboscis 1, 3rd leg without claw 3:75 mm.

Localities: Port Elizabeth. 1 non-ovig. 3 (Loman); Sea Point, Cape Town.

1 non-ovig. ¢ (K. H. Barnard, 1914).

Remarks. ‘The pinnate spines on the distal joints of the oviger are described

as ‘oak-leaf spines’ (Loman, et auct.) and are figured by Loman as more or

less of this shape. Both in this and the previous species, however, the general

outline is ovate, the basal serrations or pinnae being much longer than the

distal ones. Even Dohrn’s figures (1881) do not do full justice to these spines.

SOUTH AFRICAN PYCNOGONIDA I4!I

Gen. Nymphopsis Hasw.

1885. Haswell, Proc. Linn. Soc., N.S.W., ix, p. 1025.

1908. Loman, loc. cit., p. 49.

1920. Flynn, Pap. Proc. Roy. Soc. Tasman. for 1919, p. 83.

1O22 Loman, Ak... <00l-,, xvA.Oy De 7.

1928. id., Tijdschr. Ned. Dierk. Ver. (3), i, Afl. 2, p. 39.

1929. Flynn, Mem. Queensl. Mus., ix, p. 256.

no92))) Gordon, loc: cit. p: 129:

Body compact, segments completely fused (the cephalic segment distinct in

denticulata Gordon 1932). Crurigers separate. Ocular tubercle a short distance

Fig. 26. Nymphopsis cuspidata (Hodgson). a. lateral view of ocular tubercle and dorsal spiniferous

processes. b. chelifer of immature example. c. apex of scape and degenerate chela of adult.

d. palp. e. oviger .

behind front margin of cephalic segment. Proboscis large, fusiform, directed

forwards. Abdomen with 2 pairs (genotype armata) or 4 pairs of spinose

processes. Chelifer with 1-jointed (genotype armata) or 2-jointed scape, apex

more or less cup-like, into which the small terminal joint can be withdrawn,

in adult the latter is a feebly bifid joint, but in juvenile is a functional chela.

Palp g-jointed, and and 4th joints longest. Ovyiger 10-jointed, in both sexes,

distal joints with long spine-setae, but no serrate spines (except denticulata

Gordon), without apical claw.

Legs moderate, tarsus small, propodus strongly spinose on lower margin,

claw strong, auxiliary claws well developed, or feebly, or absent. Genital pores

on all legs in 9 on short projections; in ¢ on long curved processes on last

142 ANNALS OF THE SOUTM AFRICAN MUSEUM

2 legs. Femoral cement gland ¢g single, opening near dorsal apex. Eggs in

several packets.

Remarks. ‘The distribution of this curious and easily recognized genus

embraces Australia, East Indies, Japan, Venezuela, Falkland Islands and

South Africa.

The body, abdomen, chelifers and legs are armed with spiniferous processes

and/or branched or pinnate spines.

Nymphopsis cuspidata (Hodgson)

Figs. 25-27

1910. Hodgson, Schultze. Reise, iv, p. 221, fig. 1 (juv.). (Ammothea c.)

1923. Loman, loc. cit., p. 7, fig. D (g). (N. abstrusus.)

Body together with the crurigers broadly oval in outline, in mid-dorsal line

3 elevated spiniferous processes. Cephalic segment with a spinose tubercle,

sometimes more or less divided, on each antero-lateral corner. Crurigers

longer than median width of body, contiguous, each with 2 spinose tubercles

dorsally, one on antero-distal corner and a much larger and upstanding one

on postero-distal corner. Proboscis shortly fusiform. Ocular tubercle high,

cylindrical, tapering above the eyes to a conical point. Abdomen long, slender,

with 4 pairs of spiniferous processes dorso-laterally, of which the gnd and 4th

pairs are the largest, 2 small spines on apex, and 1-2 laterally.

Chelifer scape 2-jointed, 1st joint with a spiniferous process on dorsal apex,

end joint with 3 such processes of which the middle one is the largest, and 2-3

on distal margin, hand of chela in juv. with 2 spines, finger and thumb curved,

inner margins with minute spaced denticles, in adult chela degenerate,

represented by a feebly bifid non-spinose joint. Palp with a few long spines

on 4th joint, and numerous long spines on 5th-gth joints. Oviger with long

spines on 6th-8th joints, 9th joint with a single spine, roth joint with 2 spines.

Legs with spiniferous processes arranged mostly in pairs or 2 series dorso-

laterally, most numerous on ist and 2nd tibiae; Ist coxa with a stout spiniferous

upstanding conical tubercle (like the postero-distal one on cruriger) medio-

dorsally; 2nd coxa with ventral apex moderately protuberant on all legs in 9, ©

produced in a cylindrical process on first 2 legs in $ with long spine-setae

all around its apex; produced in a long curved process on last 2 legs in § with

long spine-setae only on the surface facing away from the body, the apex (with

genital pore) minutely spinulose; femur slightly longer than 1st tibia, and

latter slightly longer than and tibia; propodus slightly curved, with spaced

single spines on upper margin, lower margin with stout spines of which the 3

proximal ones are very strong and prominent, claw strong, auxiliary claws

stout, about half length of main claw. Femoral gland ¢ lying along dorsal

surface of femur immediately under the cuticle, and opening between 2 simple

spines near apex.

SOUTH AFRICAN PYCNOGONIDA 143

Body and abdomen each about 1:5, proboscis 3, leg (without claw) 7-5 mm.

Localities: Liideritzbucht (Hodgson); Cape Point Lighthouse (Loman);

Sea Point, Cape Town, littoral (K. H. Barnard, 1912, 1914, 1919); False Bay,

12 fath.; Port Elizabeth, East London (S. Afr. Mus.).

Remarks. ‘There can be no doubt that Loman’s specimen is the adult of

Hodgson’s species, who stated that both his specimens were immature. As

Hodgson said, the spinose nature of the animal was sufficiently characteristic

to justify describing a new species even on such material.

Fig. 27. Nymphopsts cuspidata (Hodgson). a. spine from cruriger and dorsal surface of Ist coxa.

6. pedunculate spine from leg. c¢. lateral view of 2nd coxa of leg 9. d. apex of 2nd coxa of Ist

and and legs g. e. 2nd and 3rd coxae and femur of 4th leg g, with apex of 2nd coxa, and

femoral cement gland further enlarged. £. propodus and claw of leg.

He correctly remarked that the ‘branched’ spines were really cuticular

prolongations bearing one or more movable spines. They should not be called

‘compound spines’ (Loman) or “pinnate spines’ (Flynn), at least not in this

species; in korotnewi Schimk. the armature of the legs does appear (Loman,

1908. loc. cit., pl. 13, fig. 179) to be composed of pinnate spines.

The spiny armature of the animals collects much foreign matter; unencum-

bered specimens are rare, and dirty ones are difficult to clean.

Loman described the proboscis as ‘slender’, but this adjective is scarcely

applicable to the present specimens.

The 9 from False Bay has the proboscis shorter than the chelifers and quite

invisible in dorsal view; also the medio-dorsal processes on the body are very

much elevated, their height (excluding the apical spine) almost equal to the

length of the abdomen.

144 ANNALS OF THE SOUTH AFRICAN MUSEUM

Recent collecting (1946, 1947) by the Zoology Department of the University

of Cape Town shows that this species is fairly common in shallow water in

some places in False Bay.

Gen. Kyphomia Helfer

1938. Helfer, SB. Ges. Naturf. Fr. Berlin, 1937, p. 179.

1953. Stock, Beaufortia, iv, no. 35, p. 41.

Body compact, last two segments fused. Crurigers widely separated. Ocular

tubercle near front margin of cephalic segment, slender, elongate. Chelifer

with well-developed scape, and rudimentary 2nd joint. Palp g-jointed, 2nd

and 4th joints longest. Oviger 10-jointed, with a few serrate spines, no apical

claw. Legs moderate, tarsus small, propodus spinose on lower margin, claw

strong, auxiliary claws as long as main claw.

Remarks. Stock (1953) states that Helfer’s types of this species have been

lost, but from its likeness to an East Indian species he is convinced that Helfer’s

species should be included in the genus Ammothella (1900. Verrill, Tr. Conn. Ac.

Sci., X, p. 581, as subgen. of Ammothea: palp 1o- instead of 8-jointed).

Kyphomia setacea Helfer

Fig. 28a

1938. Helfer, loc. cit., p. 179, fig. 9 a-c.

Body covered with small warts. Crurigers about equal to median width of

body. Proboscis ovoid. Ocular tubercle longer in 9 than in 4, eyes pigmented.

Abdomen slender, clavate.

Chelifer with 1-jointed scape, 2nd joint knob-like with 2 apical points.

Oviger with a few setae and serrate spines on last 4 joints (2-3 serrate spines on

last joint).

Legs with numerous long and short setae, femur in 2 swollen, in 3 nearly

cylindrical, auxiliary claws as long as main claw. Femoral gland ¢ opening

at apex of a slender process at end of femur.

Length 3 mm., span 6-7 mm.

Locality: Agulhas Bank, 35° 19’ S. 20° 12’ E., 126 metres (Helfer).

Gen. Ainigma Helfer

1938. Helfer, loc. cit., p. 181

Body compact, segmented. Crurigers widely separated. Ocular tubercle

near front margin of cephalic segment, high. Chelifer with well-developed

scape and rudimentary 2nd joint. Palp g-jointed, 2nd and 4th joints longest.

Oviger 10-jointed, apical joints with small serrate spines, terminal joint non-

serrate. Legs moderate, tarsus one-third length of propodus, which is non-

spinose, claw as long as propodus, no auxiliary claws.

SOUTH AFRICAN PYCNOGONIDA 145

Remarks. Hedgpeth (1947. Smiths. Misc. Coll., cvi, p. 4) accepts and places

this genus in the Ammotheidae. Stock (1952. Bull. Inst. Roy. Sci. Nat. Belge,

XxVili, 14, p. 5; and 1953. loc. cit. infra) considers it a synonym of Ascorhynchus.

A species of Ascorhynchus (A. glaber Hoek, 1881) was collected by H.M.S.

Challenger between the Cape and Kerguelen (46° 46’ S. 45° 31’ E., 1,375 fath.).

a b

Fig. 28. a. Kyphomia setacea Helfer, 2 with distal joints of leg. 6b. Aznigma ornatum Helfer, lateral

view, dorsal view of cephalic segment, distal joints of oviger, and distal joints of leg.

(After Helfer.)

Ainigma ornatum Helfer

Fig. 285

1938. Helfer, loc. cit., p. 181, fig. 10 a-f.

1953. Stock, Beaufortia, vi, no. 35, p. 41, fig. 6. (Ascorhynchus 0.)

1953. id., Temminckia, ix, p. 304 (in key to species).

Body covered with small warts, each segment with a conical medio-dorsal

process. Crurigers about equal to median width of body, each with a conical

upstanding process. Proboscis ovoid. Abdomen slender, clavate. Chelifer

with 1-jointed scape, 2nd joint knob-like.

Length 2:88 mm.

Locality: Agulhas Bank, 35° 19’ 8. 20° 12’ E., 126 metres (Helfer).

146 ANNALS OF THE SOUTH AFRICAN MUSEUM

Gen. Austroraptus Hodgson

1907. Hodgson, Nat. Antarct. Exp., 11, p. 54.

1915. Calman, ‘Terra Nova’ Exp. Rep., iii, p. 62.

1927. Hodgson, D. Stidpol Exp., xix (zool. xi), p. 349.

1932. Gordon, ‘Discovery Rep., vi, p. 114.

1944. id., B.A.N.Z. Aniarct. Res. Exp., v, p. 57-

Body compact, segments fused or only indistinctly demarcated. Crurigers

narrowly separated or contiguous. Proboscis flask-shaped, stout at base,

narrowing to a point, directed or curved downwards. Chelifers small, chelate,

rudimentary. Palp 6-jointed, the apical joint inserted laterally on the 5th and

Fig. 29. Austroraptus thermophilus Brnrd. a. dorsal view (palp bent downwards). 5b. lateral view.

c. ventral view of proboscis. d. 1st and 2nd coxae of 4th leg ¢.

projecting forwards or upwards (the apical joint: may be tripartite). Oviger

10-jointed, in both sexes, apical joints with simple non-serrate spines, no apical

claw. Legs moderate, tarsus short, auxiliary claws moderate, small, or minute.

Remarks. Previously known only from the Antarctic (3 species); the dis-

covery of a species in the considerably warmer waters of South Africa is

noteworthy.

Appears to differ from the species of Tanystylum with 6-jointed palps only in

the shape of the proboscis.

Austroraptus thermophilus Brnrd.

Figs. 29, 30

1946. Barnard, Ann. Mag. Nat. Hist. (xi) 13, p. 62.

Body compact, all the segments fused, smooth; cephalic segment broad

with a tubercle at each antero-lateral corner. Crurigérs contiguous, their

SOUTH AFRICAN PYCNOGONIDA 147

length greater than median width of segments, each with a tubercle on antero-

distal and postero-distal corners dorsally, a little more prominent in g than 9.

Ocular tubercle conical, rather broad and low, eyes distinct. Proboscis

markedly flask-shaped, broad proximally, rapidly narrowing to a downwardly

curving apex. Abdomen elongate, extending beyond end of Ist coxa of last

leg, horizontal.

Chelifers vestigial, composed of a single conical projection with 2 apical

setules, less prominent in 9 than in g. Palps robust, curving downwards, 4th

Fig. 30. Austroraptus thermophilus Brnrd. a. ocular tubercle from right side. 6. palp. c. oviger g.

d. femur and tst tibia of leg g. ¢. apex of femur with cement gland ¢. f/f. distal joints of leg.

joint longest, angular in middle of upper margin, a pale or semitransparent

oval spot indicating a gland (but orifice not traceable), 5th joint conical,

setose, 6th ovate, inserted laterally on 5th and projecting forwards and upwards,

setose. Oviger in § with 4th and 5th joints gently curved, distal joints feebly

armed, 8th and oth each with a simple spine on inner margin, toth joint with

2 apical spines; in 9 similar but shorter and weaker, 4th and 5th joints straight.

Legs 1st coxa with the antero-distal and postero-distal corners produced in a

tubercle, the anterior one larger than the posterior and carrying I or 2 spines,

both tubercles are more prominent in J than in 9; 2nd coxa with lower apex

slightly but not prominently gibbous, genital pores in @ on all legs, in 3 only on

last two; femur with upper apex produced, Ist and 2nd tibiae subequal (and

subequal to femur) with low rounded spiniferous tubercles arranged more or

less in 2 longitudinal dorso-lateral series; propodus with 3 large spines proxi-

mally on lower margin, auxiliary claws well developed, nearly half length of

148 ANNALS OF THE SOUTH AFRICAN MUSEUM

main claw. Femoral cement gland ¢ single, elongate, lying immediately below

the cuticle, opening on distal surface of the apical projection.

All the tubercles on the cephalon, crurigers and Ist coxae, and the

nodulosities on the legs are microscopically hispid.

Body incl. abdomen 3 (trunk and abdomen subequal), proboscis 1-25, leg

without claw 8 mm.

Localities: False Bay, 23 fath. 1 9; off Cape St. Blaize, 34 fath. 1 ovig. 3g;

off Nanquas Peak (east of Algoa Bay), 63 fath. 2 non-ovig. g¢ (S. Afr. Mus.).

Remarks. Distinguished from the genotype polaris by the contiguous crurigers,

and from all the three Antarctic species by the longer abdomen and more

markedly flask-shaped proboscis, shorter ocular tubercle, shorter tubercles on

Ist coxae, and the distinct though not prominent nodulosities on the tibiae;

the auxiliary claws also are stronger.

Fam. TANYSTYLIDAE

1913. Schimkewitsch, ool. Anz., xli, p. 613.

1947. Hedgpeth, Smiths. Misc. Coll., cvi, p. 4.

Octopodous. Body compact, segments fused or segmentation visible only

laterally. Crurigers narrowly separated or contiguous, or fused. Proboscis

large, fusiform, directed forwards. Chelifers absent in adult, sometimes

rudiments in juvenile. Palps 4-6-jointed. Ovigers in both sexes, 10-jointed,

without compound spines. Legs stout, tarsus very short, auxiliary claws present.

Genital pores on all legs in 9, on last 2 in g. Eggs in several masses.

Key to the South African Genera

1. Palp 6-jointed. Tanystylum

2. Palp 5-jointed. Discoarachne

Gen. 7 anystylum Miers

1879. Miers, Philos. Trans. Roy. Soc. Lond., clxviii, p. 213.

1932. Gordon, Ann. Mag. Nat. Hist. (10), x, p. 87.

1Q924) Wd. Discovery mivep.. vino lle

Body subcircular, unsegmented. Crurigers narrowly separated or contiguous,

or fused. Proboscis stout, movable, directed forwards. Palp 6-jointed.

Key to the South African [incl. Tristan d’ Acunha] species

1. Crurigers fused. ornatum

2. Crurigers separate. [ pfeffert |

Tanystylum ornatum Flynn

19262) Plynn) loc, cit),;p. 33) es. 20, om.

Crurigers fused, the divisions between them indicated by superficial markings;

each cruriger with a pair of knobs distally on dorsal side. Ocular tubercle

moderately high, rounded, surmounted by a short pointed process. Proboscis

ovoid. Abdomen clavate, oblique.

SOUTH AFRICAN PYCNOGONIDA 149

Chelifers represented by a pair of small setose processes. Palp 4th joint

longest, 2nd and 6th subequal. Oviger in ¢ 4th joint curved, 5th longest, 8th

at right angles to 7th, toth with 2 apical spines; in @ (as figured by Flynn)

Ath and 5th joints subequal, 6th-9th joints each with 1 spine on outer apex,

gth also with 2-3 spines on inner apex, 10th joint with 2 apical spines.

Legs 1st coxae ending in 2 or more lobes dorsally, the three coxae subequal,

femur and tibiae subequal, propodus without heel.

Body with proboscis, excl. abdomen 1-8 mm.

Locality: off west coast of Cape Peninsula, 25-30 fath. (Flynn).

Remarks. ‘The oviger as represented in fig. 21a as that of a 9 seems rather

remarkable, especially as regards the 10th joint with its 2 stout opposing spines

(cf. Discoarachne brevipes 3). It may be noted that the latter species was taken

in the same locality together with the TYanystylum specimens.

Gen. Discoarachne Hoek

1881. Hoek, Rep. H.M.S. ‘Challenger’, iii, p. 74.

1901. Cole, ool. Jahrb. Abt. Syst., xv, pp. 243 sqq.

1904. Woman, ibid., xx, p. 383.

1908. id., Siboga Exp. monogr., xl, pp. 14 sqq. and synopsis facing p. 19.

Resembling Yanystylum but palp 5-jointed. Genital pores in ¢ on last 2 legs,

in 2 on all legs. Femoral cement gland ¢ a single pore near dorsal apex.

Remarks. Loman (1904) erroneously stated that the crurigers were fused,

and that the ¢ oviger was g-jointed. The latter statement was corrected by

him in 1923 (loc. cit. infra).

The genus is sometimes regarded (1909. Thompson, Cambr. Nat. Hist., iv,

p- 535) as a subgenus of Tanystylum.

Discoarachne brevipes Hoek

Fig. 314, b

leo) wiloek, loc. cit. pp. 74, 105, pl. 7, figs. 8-12.

TOO Cole, loc. cit. p:) 243, pl..13.

1904. Loman, loc. cit., p. 379, pl. 14, figs. 7-11.

1908. id., loc. cit., p. 46 (incidentally mentions ‘commensals’ [? Rotifers] on

the palps).

1910. Hodgson, Schultze. Reise, iv, p. 227.

nee welkoman, Ark. Zool, xv79, p- 6, fig. C.

nOZee Hibynn loc: cit...p4)35-

1951. Stock, Mem. Inst. Roy. Sci. Nat. Belge (2) fasc. 43, p. 3.

Body oval, crurigers distinct and not fused though separated only by clefts.

Proboscis somewhat fusiform. Ocular tubercle low, rounded, eyes distinct.

Abdomen extending to end of last crurigers, rather slender, clavate, horizontal.

Chelifers represented in young specimens by 2 small setiferous processes,

sometimes 2-jointed (Loman, 1904, fig. 7a); in older specimens these tiny

150 ANNALS OF THE SOUTH AFRICAN MUSEUM

stumps may be retained, but as often as not no trace of the chelifers is found.

Palp slender, with first 2 joints short, 3rd longest, often swollen, 4th short.

Oviger in § with 8th joint attached at right angles to 7th, roth with 2 apical

and rather stout spines simulating a minute chela; in 9 normal, roth joint

longer than oth, with several apical spines.

b Gu a

Fig. 31. a, b. Discoarachne brevipes Hoek, dorsal view, and goth and roth joints of oviger <.

c, d. Pycnogonum forte Flynn, dorsal view with 3rd right leg to same scale (coxae in dorsal view,

remainder of leg in postero-lateral view).

Legs stout, the three coxae subequal, femur somewhat contorted, tarsus

much narrower than end tibia, propodus with 3 stout spines at base but no

distinct heel. Very minutely hispid. Femoral cement gland ¢ single, near the

apex on dorsal surface.

Body (tip of proboscis to tip of abdomen) 4-4°25 mm.

Localities: Cape ‘Town, littoral (Hoek, Cole, Loman, Flynn); off west coast

of Cape Peninsula, 25-30 fath. (Flynn); Cape Point Lighthouse, littoral

(Loman); False Bay (probably littoral) (Hodgson); Port Natal (Durban)

(Loman); Liideritzbucht (Stock); Port Nolloth, Saldanha Bay, Table Bay,

west and east coasts of Cape Peninsula, Kleinmond (mouth of the Palmiet River,

Cape Province), Plettenberg Bay, East London, all littoral (S. Afr. Mus.).

Remarks. This is a very common Pycnogonid on both sides of the Cape

Peninsula among weeds, Hydroids, anemones, etc., between tide marks.

SOUTH AFRICAN PYCNOGONIDA I51

Ovigerous jg have been collected in February, July and September to

November; probably breeding occurs throughout the year.

Stock’s record is presumably based on the ‘Mercator’ collections, although

he makes no mention of this species in the systematic part of his paper. The

specimens collected by Dr. Schultze, and recorded by Hodgson, came from

False Bay, not from Liideritzbucht. ;

Fam. PycNOGONIDAE

1908. Loman, Siboga Exp. monogr., xl, conspectus facing p. 19.

1909. Thompson, Cambr. Nat. Hist., iv. p. 539.

1913. Bouvier, 2me Exp. Antarct. Franc., pp. 46, 149.

1927. Calman, Trans. Zool. Soc. Lond., xxii, pp. 408, 410.

1947. Hedgpeth, Smiths. Misc. Coll., cvi, p. 5 (in list of families and genera).

Decapodous (Pentapycnon) or octopodous. Body segmented or the segmen-

tation more or less suppressed. Proboscis projecting forwards or more or less

bent under body. Chelifers reduced or absent, if present either chelate in adult

or only injuvenile. Palps usually absent. Ovigers present in ¢ only, 6-9-jointed,

with apical claw, but no spines. Legs without auxiliary claws (sometimes

vestiges). Genital pores on last leg only in both sexes. Eggs in a single cake-

like mass.

Remarks. Several genera which are in one respect or another aberrant have

been associated with the genus Pycnogonum in this family.

Gen. Pycnogonum Briin.

1764. Briinnich, Entomologia, p. 84.

1881. Dohrn, F. Fl. Golf. Neapel, ii, p. 202.

1902. Mobius, D. Tiefsee Exp., ii, p. 193.

1908. Loman, loc. cit., pp. 5-9, conspectus facing p. 19 and p. 34.

1913. Bouvier, loc. cit., pp. 150, 156.

coz one ice Ant...Scls Nat Zool. (10), Vs ps 113:

Octopodous. Body with segments 3 and 4 often close together and junction

difficult to trace, or fused. Crurigers separate or contiguous. Proboscis

movable, stout. Chelifers absent (but see: P (?) claudum Loman, 1908). Palps

absent. Ovigers in § only, sometimes absent even in ovigerous specimens.

Legs with auxiliary claws vestigial or absent. Genital pores on 2nd coxa of

last leg in both sexes, in 9 on dorsal surface, in ¢ on ventral surface.

Remarks. Dohrn (loc. cit., p. 203) stated that the genital pores in 9 were

dorsal, and this applies to his puszllum; but in his nodulosum he described them

as being ventral, albeit near the posterior margin.

Key to the South African Species

I. No auxiliary claws.

A. Junction of 3rd and 4th segments not evident, these two segments more

or less fused. Crurigers contiguous.

152 ANNALS OF THE SOUTH AFRICAN MUSEUM

1. 2nd tibia at least half length of 1st tibia. Proboscis cylindrical.

a. 4 medio-dorsal tubercles (only 1 on cephalic segment). cataphractum

b. 5 medio-dorsal tubercles (2 on cephalic segment). Sorte

2. and tibia scarcely one-third length of 1st tibia. Proboscis tapering. portus

B. 3rd and 4th segments quite distinct, and all crurigers separate. Ocular

tubercle very small. microps

II. Small auxiliary claws present. pusillum

Pycnogonum cataphractum Mobius

1902. ) Mobius; loc cit, ps 194, plaso, teenie

Integument reticulate. Body stout, cephalic segment wider than following

segments, all of which are (according to the figure) distinct; 4 medio-dorsal

conical tubercles, one on each segment. Crurigers contiguous, each with a

conical boss or process on dorsal apex. Ocular tubercle conical, rounded,

shorter than the tubercle behind it, eyes distinct. Proboscis stout, somewhat

narrowed in distal half, apex truncate. Abdomen extending slightly beyond

last crurigers, apically truncate.

Legs with conical or digitiform processes, one on dorsal apex of Ist coxa,

4-5 on femur, 1-2 on ist tibia, 1 on 2nd tibia, tibiae subequal, each slightly

shorter than femur, claw one-third length of propodus (drawn too long in the

figure), no auxiliary claws.

Smaller specimen: body 9, proboscis 4, abdomen 1 mm.; larger (incom-

plete) specimen; proboscis 5 mm.

Locality: St. Francis Bay, shallow water (Mo6bius).

Remarks. P. tumulosum Loman, 1908, East Indies, has somewhat similar

nodose legs, but has 2 medio-dorsal tubercles on cephalic segment and none

on 4th segment.

The smaller of the ‘Valdivia’ specimens had eggs attached to the ventral

surface but no ovigers.

Pycnogonum forte Flynn

Ries ouch

19205) Flyin loc. (cits) ps Sila ese To ml@n(eyE

Very close to cataphractum. Third and 4th segments fused, last pair of

crurigers fused; 5 medio-dorsal, more or less digitiform, tubercles of which 2

are on cephalic segment and one on each of the other segments. Abdomen

longer, extending considerably beyond last crurigers.

Legs in holotype less conspicuously tubercular than in cataphractum. In the

present specimen the legs are conspicuously tubercular; a conical tubercle on

upper apex of Ist coxa, 1 or 2 low rounded tubercles dorsally on both 2nd and

grd coxae; on femur 2 rows of rounded tubercles increasing in size distally,

with a large conical tubercle between the apical pair; Ist tibia with a large

tubercle near base followed by 2 rows of rounded tubercles; 2nd tibia similar,

but the tubercles less prominent; propodus feebly granulose on upper surface,

finely setulose on lower margin, claw about one-third length of propodus,

SOUTH AFRICAN PYCNOGONIDA 153

auxiliary claws absent. Genital pores in present specimen on dorsal surface of

2nd coxa of last leg.

Total length 10:5 mm., proboscis 5 mm., abdomen 2 mm. (Flynn’s specimen

resp. about 10°3, 3°8, 1°5).

Localities: off Gneka River*, 1 2 (Flynn); off Great Fish Point, 49 fath.

1 9 (S. Afr. Mus.).

Remarks. ‘This species is so extraordinarily like cataphractum that, when more

material is available, it will probably be found to be conspecific.

Fig. 32. Pyncnogonum portus Brnrd. Dorsal view; lateral view, with process further enlarged;

lateral view of leg, with dorsal surface of femur, and tarsus and propodus further enlarged.

Mobius may possibly have overlooked the 2nd tubercle on the cephalic

segment, but however that may be the longer abdomen in forte seems to be a

definite differential character, unless it be sexual.

Flynn’s figure gives the impression of shallow depressions between the

crurigers and the body proper; in the present specimen they are much deeper.

And though the 4th segment is fused with the grd the suture is visible dorsally.

The present specimen has a very small and indistinct granule on the hinder

slope of the ocular tubercle.

Pycnogonum portus Brnrd.

Fig..92

1946. Barnard, Ann. Mag. Nat. Hist., (xi) 13, p. 62.

Integument reticulate. Body stout, cephalic segment very broad, following

segments rapidly decreasing in width, junction of 3rd and 4th difficult to trace

dorsally; 5 medio-dorsal tubercles, 2 on cephalic segment, one on each of the

other segments, each tubercle ornamented with a number of minute conical

* See p. 85 footnote.

154. ANNALS OF THE SOUTH AFRICAN MUSEUM

or digitiform processes. Near hind margin of cephalic and of and segment

2 conical tubercles on either side of the medio-dorsal tubercle, and on 3rd

segment one similar tubercle flanking the medio-dorsal one; several additional

minute granules or projections, resembling those on the medio-dorsal tubercles,

scattered over the integument. Crurigers contiguous, each with an ornamental

tubercle on upper apex. Ocular tubercle rather low, rounded, eyes distinct.

Proboscis stout at base, rapidly tapering to a narrowly truncate apex. Abdomen

extending beyond last crurigers, ornamented like the medio-dorsal tubercles

and with a small tubercle on dorsal apex.

Legs stout, nodulose and granulose with granular and digitiform outgrowths

similar to those on body, Ist coxa with a tubercle on dorsal apex, femur rather

strongly convex on proximal ventral surface and on distal dorsal surface, on

the latter several outgrowths of various shapes, some bearing a spinule, and

also a large prominent tubercle on apex, Ist tibia a little shorter than femur,

with a low tubercle or boss proximally on dorsal surface, and numerous out-

growths, 2nd tibia very short, scarcely one-third length of Ist tibia, tarsus

spinulose on lower surface, propodus slightly tapering, granulose, with spinules

on lower margin, claw about one-third length of propodus, no auxiliary claws.

Genital pores on dorsal surface of 2nd coxa of last leg in 9; not traceable on

the ovigerous specimen which is presumably a 6.

Total length 3-5 mm., body plus abdomen 2:75 mm.

Locality: Port Elizabeth, littoral. 1 9, 1 ovigerous § (University of Cape

Town Ecological Survey, 1936).

Remarks. ‘The integumentary ornamentation, and the shortness of the 2nd

tibia are the noteworthy features of this species.

Remarkable also is the complete absence of ovigers in the specimen which

carries a flat cake-like mass of eggs. The eggs are loosely cemented to the ventral

surface of the body, and the specimen is presumably a dg, especially as no

distinct and large genital pores, as are easily observed in the second specimen,

can be traced.

So far as I am aware the absence of ovigers in ovigerous specimens had not

been observed except by Mobius (1902, loc. cit.) in his description of cata-

phractum. Are the ovigers never developed in some species? Or does a male

before he is fully developed (structurally and perhaps also sexually) have to

carry as best he can the packet of eggs which a female may dump upon him?

Pycnogonum microps Loman

1904. Loman, Zool. Jahrb. Abt. Syst., xx, p. 378, pl. 14, figs. 5, 6 (9)

Integument reticulate and granulose. Body stout, the segments especially

3rd and 4th distinct, cephalic segment not greatly wider than the others; 4

medio-dorsal tubercles, two (a smaller anterior and a larger posterior one) on

cephalic segment, and one on each of segments 2 and 3. Crurigers almost as

long as their segments, not contiguous, each with a small tubercle near dorsal

SOUTH AFRICAN PYCNOGONIDA 155

apex. Ocular tubercle very small, eyes minute. Proboscis very stout, cylindrical,

apex truncate. Abdomen extending slightly beyond ends of Ist coxae of last

legs, apically truncate.

Legs stout, granulose but without tubercles or bosses; 2nd tibia nearly as

long as Ist, no auxiliary claws.

Body 3, proboscis 1:5, abdomen -5 mm.

Locality: IUlovo or Isipingo (Natal), littoral. 1 Q (Loman).

Remarks. Quite different from the other South African species in the form of

the body.

Loman’s figure 6, showing the body in profile, does not seem to correspond

with fig. 5, as it shows only the smaller medio-dorsal tubercle on the cephalic

Fig. 33. Pycnogonum pusillum Dohrn. Lateral view; portion of integument; rst leg.

segment, and a transverse ridge culminating in the median boss on each of the

following segments. The intersegmental sutures seem to be wrongly indicated.

Pycnogonum pusillum Dohrn

Fig. 33

1881. Dohrn, Fauna Flora Golf. Neapel., iii, p. 207, pl. 16, figs. 4-8.

1953. Stock, Bull. Inst. Roy. Sci. Nat. Belge, xxviill 14, pp. 1-3, and 5, fig. 1.

Integument with numerous, but scattered granules crowned with minute

spinules. Body stout, segments distinct, cephalic segment not wider than the

others; 3 medio-dorsal prominences, one each on the cephalic and the two

following segments. Crurigers not contiguous, but angle of divergence between

and and grd greater than that between the others; each with a prominence

near dorsal apex. Ocular tubercle as large as the body prominences, eyes well

developed. Proboscis very stout, slightly tapering to the truncate apex.

Abdomen extending slightly beyond end of 1st coxae of last legs, apically

rounded. No ovigers.

I 56. ANNALS OF THE SOUTH AFRICAN MUSEUM

Legs stout, granulose like the rest of the integument, without tubercles or

bosses, but dorsal apex of femur somewhat gibbous; and tibia nearly as long

as Ist; auxiliary claws small but distinct. Genital pore not traceable.

Body 2, proboscis -75, abdomen -5 mm.

Locality: False Bay (Cape), 8 fath. (Univ. Cape Town Ecol. Surv., Aug.

1952, one specimen).

Distribution. Gulf of Naples (Dohrn); Santa Maria Bay, Angola (13° 25’ S.).

(Stock.)

Remarks. After its original description seventy years elapsed before this

species was rediscovered; and then far away from its previously known habitat.

It is strange that a couple of years later another example should be collected,

and even farther afield. That is assuming that the identification of the present

specimen is correct. ‘There is no justification for a n.sp.; the single specimen

can be compared only with pusillum, and only one feature requires comment.

Dohrn used the words ‘betrachtlicher Hocker’ and ‘betrachtlicher Buckel’

in describing the projections on the segments and crurigers, but unfortunately

omitted to indicate these projections in his figure (as Stock pointed out). I would

use a similar term, boss, projection or prominence, in preference to ‘tubercle’

for the present specimen. Stock, however, used the word ‘tubercle’, and his

figure gives the impression that these prominences are in fact small and well-

defined tubercles.

Dr. Stock, to whom a sketch of the Cape specimen was sent, considers that

it should be identified as pusillum; and states that the ocular tubercle and

body prominences are subject to considerable variation in the genus Pycnogonum.

INCERTAE SEDIS

Gen. Queubus Brnrd.

1946. Barnard, Ann. Mag. Nat. Hist. (xi) 13, p. 63.

Octopodous. Body distinctly segmented, anterior part of cephalic segment

produced considerably in front of first crurigers. Proboscis movable, elongate,

slender, tapering. Chelifers and palps absent. Oviger ¢ 10-jointed, with apical

claw (2 unknown). Legs strong, but of good length proportionately to body,

end coxa longer than either ist or 3rd, tarsus short, propodus stout, with heel

and strong spines on lower margin, claw strong, no auxiliary claws. Femoral

pores on ventral apex of and coxa of last 2 legs. Eggs carried in a single mass.

Remarks. Resembling Pycnogonum in the absence of chelifers and palps, but

differing in the armature and subchelate shape of the propodus of the legs.

The absence of the 2 makes the position and relationships of the genus

uncertain.

Pigrogromitus Calman, 1927, is another genus with some remarkable features

and of uncertain position. ‘The name of the present genus is taken from the

same passage in Shakespear.

SOUTH AFRICAN PYCNOGONIDA 157

Queubus jamesanus Brnrd.

Fig. 34

1946. Barnard, loc. cit., p. 63.

g — In general resembling Pycnogonum crassirostre, but legs relatively longer.

Integument smooth. Hind part of cephalic and 4th segments, and anterior

part of 2nd and 3rd segments raised into a prominent conical boss. Crurigers

Fig. 34. Queubus jamesanus Brnrd. Dorsal view 3 (gap between 3rd and 4th crurigers drawn too

wide, abdomen foreshortened and drawn in true length alongside); lateral view, with oviger,

6th and roth joints of latter further enlarged; tarsus, propodus and claw of leg.

separated by half to three-quarters their own width (gap between 3rd and

4th in the figure is too wide), their length less than median width of segments.

Ocular tubercle large, conical, not very high, eyes distinct. Proboscis long,

narrowing rapidly, the distal two-thirds slender, cylindrical. Abdomen deflexed,

clavate, with pointed apex.

Oviger 10-jointed, 4th and 5th joints longest, 5th curved, 6th ovately

expanded, inner margin with a strong bifid spine and a simple spine, 7th-1oth

joints with minute spinules, apical claw short.

Legs smooth, 2nd coxa longer than either 1st or 3rd, but not quite equal to

the other two combined, femur subequal to the three coxae combined, Ist and

and tibiae subequal, each shorter than femur, tarsus strongly spinulose on

158 ANNALS OF THE SOUTH AFRICAN MUSEUM —

lower surface, propodus curved, with heel carrying 6-7 strong spines, with

other spines on lower margin decreasing in size, lower apex angularly promi-

nent, spinulose, claw strong, reaching to the heel on propodus, no auxiliary

claws.

Base of proboscis to base of abdomen 4 mm., proboscis 3 mm., leg without

claw 9 mm.

Locality: St. James, False Bay, low-tide. 1 ovig. 3 (K. H. Barnard, 1912).

Remarks. The single mass of eggs was carried by the right oviger only, the

left being folded up alongside.

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ANNALS

SOUTH AFRICAN MUSEUM

- VOLUME XLI

- PART IV, containing :-—

8. Les Paguristes des cétes occidentales et méridionales d’ Afrique. Par JACQUES

Forest. (Avec planche IV et 70 figures dans le texte.)

ISSUED JUNE 1954. PRICE 155.

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TRUSTEES OF THE SOUTH AFRICAN MUSEUM

BY THE RUSTICA PRESS (PTY.) LIMITED, COURT ROAD, WYNBERG, CAPE

aee Me

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Ra aid

8. Les Paguristes des cotes occidentales et méridionales d’ Afrique.

par JAcquEs ForREstT

Muséum National d’Histoire Naturelle, Paris

(Avec planche IV et 70 figures dans le texte)

Ie INTRODUCTION

Sur les cinquante-et-une espéces ou varietés de Paguristes mentionnées par

Alcock dans son inventaire des Pagurides du monde (1905), trois seulement appar-

tenaient a la faune ouest-africaine et il n’y en avait qu’une qui fut signalée pour

PAfrique du Sud. Encore dans ces quatre cas, ne connaissait-on que les types! On

aurait pu croire a cette époque que le genre Paguristes, florissant dans lIndo-

Pacifique et dans la mer des Antilles, n’était que sporadiquement et fort pauvrement

représenté dans |’Atlantique africain. La description, depuis lors, de trois nouvelles

espéces ne modifiait guere les conclusions précédentes: il ne s’agissait toujours que

de quelques individus.

Aujourd’hui, aprés avoir dépouillé un matériel abondant et d’origine variée, il

faut bien constater que la pauvreté de la faune ouest et sud-africaine en Paguristes

n’était qu’apparente: il y a peut-étre en certain point de ce littoral plus d’espéces et

plus d’individus appartenant a ce genre qu’en aucune autre région du monde. Aux

sept espéces connues, s’ajoutent maintenant neuf especes nouvelles. Paguristes

mauritanicus dont on ne connaissait que le type a été récolté en plus de vingt-cing

stations et les collections en renferment des centaines d’exemplaires.

La liste des espéces n’est d’ailleurs certainement pas close, car bien des régions

et bien des niveaux restent inexplorés. A considérer la carte de repartition (fig. 69)

il semblerait que c’est autour de Dakar que les Paguristes sont les plus nombreux et

les plus variés; il est peu probable en effet que des dragages opérés en d’autres points

puissent ramener un échantillonnage de faune pagurienne plus intéressant que

celui du 18 janvier 1952, en Baie de Rufisque: les quelques centaines de Pagures

récoltés—de petite taille puisqu’ils occupaient un volume total de moins d’un litre-

appartenaient a quatorze espéeces différentes dont quatre de Paguristes. On doit

cependant tenir compte du fait que cette région de Dakar, depuis la cote jusqu’a

une profondeur de 100 a 120 métres, est l’une des mieux connues du littoral ouest-

africain; il ne faudrait pas conclure, au vu des vastes blancs qui figurent sur notre

carte de répartition, que les Paguristes sont totalement absents des cotes du Cameroun

ou du Sud-ouest africain par exemple. Ces blancs signifient surtout que l’on se

trouve en présence de zones pratiquement inexplorées ou des recherches ultérieures

permettront sans doute de retrouver des espéces signalées ailleurs ou révéleront des

formes nouvelles. Si on considére d’autre part que beaucoup de Paguristes vivent a

159

VOL. XLI. PART 4.

AUR S

1994

160 ANNALS OF THE SOUTH AFRICAN MUSEUM

des profondeurs assez grandes—Alcock estimait que 30% des espéces de ce genre se

trouvent a un niveau inférieur a 100 brasses (180 métres environ)—on peut s’attendre

a en découvrir de nouvelles dans |’Atlantique africain lorsqu’on aura la possibilité

de draguer en-dessous de 120 meétres, limite de la plupart des récoltes effectuées a ce

jour. Le matériel provenant de profondeurs plus grandes est fort peu abondant et

provient pour la plus grande part des campagnes du ““Travailleur’”’ et du ““Talisman”’.

La grande majorité des Paguristes intertropicaux d’Afrique occidentale ont un

point commun: a l’exception de P. oculatus var. rubropictus, variété d’une espéce

méditerranéenne et de P. marocanus qui vit a une profondeur plus grande que les

autres, tous présentent une anomalie de lappareil génital femelle; lorifice sexuel

est unique et situé sur la coxa gauche; il n’y a pas trace d’orifice droit. Dans le

genre Paguristes nous n’avons pour l’instant retrouvé cette disposition de l’appareil

génital femelle que chez des espéces de la Mer Rouge; ceci n’est pas le seul point

commun entre les Paguristes d’Afrique occidentale et ceux de Mer Rouge dont la

systématique embrouillée par Nobili est entierement a revoir: P. hispidus, P. micro-

phthalmus et P. rubrodiscus—les trois espéces aux écailles oculaires trés allongées—

sont plus proches de P. jousseaumei, décrit par Bouvier d’Aden et de Suez, que

d’aucune autre espéce ouest-africaine et cette constatation s’ajoute a celles que

nous avons eu l’occasion de faire a propos d’autres Paguridae. Certains Diogenes du

Golfe de Guinée existent aussi en Mer Rouge, ou y sont représentés par des formes ©

voisines (Forest, 19524, p. 13).

Chez d’autres Paguridae appartenant ceux-la a la sous-famille des Eupagurinae,

il n’existe aussi qu’un orifice génital femelle, situé sur la coxa de la troisieme patte

thoracique gauche comme chez les Paguristes; il s’agit des genres Sympagurus et

Parapagurus. Il est intéressant de noter que dans ces deux genres, que lon considére

en général comme bien éloigné des Paguristes, le male posséde également deux

paires de pléopodes en général modifiés en gonopodes (voir p. 163). |

| Au point de vue biogéographique on remarque qu’il existe sur la cOte occi-

dentale d’Afrique, entre les Tropiques, un groupe de Paguristes dont on ne trouve

aucun représentant ni au nord ni au sud, ni en une autre région du monde. _L’examen

de la carte de répartition (fig. 69) montre que le genre est représenté par dix espéces

entre !’Equateur et le Tropique du Cancer. Parmi ces dix espéces il en est quatre

que I’on retrouve au sud de l Equateur, P. oculatus var. rubro-pictus (si la détermination

d’Odhner est exacte), P. mauritamicus, P. hispidus et P. virilis. Une cinquieme espéce,

la seule qui serait propre a cette région, est fort proche de P. virilis: c’est P. skoogt.

Ainsi, dans le cas ot |’abondance et la diversité des Paguristes au large de Dakar ne

seraient pas seulement dues a une meilleure connaissance de ce point du littoral,

on pourrait admettre que la céte sénégalaise constitue le centre de dispersion des

espéces intertropicales ou tout au moins la région ou le genre a rencontré les con-

ditions de vie les plus favorables.

Les Paguristes d’Afrique occidentale paraissent rechercher les fonds coquilliers,

caillouteux ou vaseux, de préférence aux fonds de sable fin. On ne les rencontre

guére dans la z6ne intercotidale, mais surtout a partir de quelques métres jusqu’a

cinquante métres environ. I] semble que, pour une espéce au moins, il y ait une

PAGURISTES D’AFRIQUE L’OUEST ET DU SUD 161

relation entre la latitude et la profondeur a laquelle on la trouve: Paguristes mauri-

tanicus a été capturé entre 5 et 30 métres en A.O.F. et entre 50 et 100 métres au

Congo. P. marocanus qui n’appartient pas au groupe intertropical proprement dit a

été dragué entre 115 en 290 metres, mais il n’est connu que par un trés petit nombre

d’exemplaires et il est possible qu’on le rencontre a d’autres niveaux.

Pour l’Atlantique africain intertropical, nous avons disposé d’un matériel

abondant dont la base était constituée par la collection du Muséum d’Histoire

Naturelle, avec notamment les spécimens recueillis par le ““Travailleur’’ et par le

“Talisman”. Nous avons également trouvé de nombreux Paguristes parmi les

Pagures rassemblés au cours des derniéres années par les zoologistes de l'Institut

Frangais d’Afrique Noire et dans la collection qui nous a été confi¢e par I’Institut

Royal des Sciences Naturelles de Belgique (Croisiére du “Mercator” et Expédition

“M’Bizi’’). D’autres échantillons ont été récoltés par MM. E. Postel et R. Sourie de

Dakar. La répartition géographique des espéces a été établie en tenant compte

également des spécimens dragués au large de la Gold Coast par R. Bassindale,

lesquels seront étudiés en détail avec les autres Pagures de méme provenance.

Nous n’avons pas rencontré de difficulté particuliére dans l’étude des Paguristes

de cette région et ceci surtout parce que nous avions un assez grand nombre de

specimens de la plupart des espéces. Toutes—a l’exception de P. difficilis Forest

qu'il est assez malaisé de distinguer de P. mauritanicus Bouvier sans examiner les

pléopodes sexuels du male—sont relativement faciles a déterminer a l’aide de

caractéres bien apparents. Notons cependant qu’ils sont presque tous revétus d’une

abondante pilosité constituée par de longs poils plumeux; ces poils retiennent de

fines particules de vase entre leurs barbules et cachent les régions du corps qui sont

précisément les plus caractéristiques; aussi est-il nécessaire le plus souvent de nettoyer

les spécimens a déterminer a l’aide d’une brosse fine.

Nous avons été embarrassé par le cas de Paguristes skoogi Odhner: c’est la seule

espéce déja décrite dont nous n’ayons pas examiné de spécimens. Le type, contraire-

ment a ce que nous espérions, ne se trouverait ni a Stockolm ni a Géteborg. Notre

P. virilis en est certainement fort proche, mais ne répond pas exactement a la

description d’Odhner, ce qui nous interdit d’identifier les deux espéces pour

Pinstant.

Les Paguristes d’Afrique du sud—a l’exception du type de P. gamianus de H.

Milne-Edwards conservé au Muséum de Paris—nous ont été communiqués par le

Dr. K. H. Barnard, et appartiennent au South African Museum ou au Département

de Zoologie de l’Université de Cape Town. Ces espéces nous réservaient des

difficultés bien plus grandes que les précédentes.

Nous avons tout d’abord hésité a établir l’espéce nouvelle décrite ici sous le

nom de P. barnardi, en raison de limpossibilité de la distinguer du P. gamianus H.

Milne-Edwards d’aprés l’aspect de la carapace et des appendices céphaliques.

Cependant des différences marquées dans la pilosité et l’ornementation des pattes

ambulatoires et dans la taille, ainsi que le nombre assez grand de spécimens dont

162 ANNALS OF THE SOUTH AFRICAN MUSEUM

nous disposions et que l’absence de formes intermédiaires, nous ont finalement

amené a penser qu'il s’agissait réellement de deux espéces distinctes.

Une seconde difficulté s’est présentée avec le spécimen décrit a la suite de

P. gamianus sous le nom de Paguristes sp. C’est un individu male d’une taille supér-

ieure a celle des plus grands P. gamianus, dont il différe sur un certain nombre de

points. Ces différences sont-elles liées a l’Age du spécimen en question? Quoi qu’il en

soit il ne nous a pas paru opportun d’en faire une nouvelle espéce.

Enfin, si nous décrivons sans hésitation P. macrotrichus sp. nov. d’aprés un spéci-

men femelle déterminé—a tort—par Stebbing comme P. gamianus, nous ne sommes

pas absolument certains que le Paguristes male plus petit, et d’aspect assez différent,

qui figure ici sous le nom de P. agulhasensis sp. nov., n’appartienne pas a la méme

espece.

I] subsiste ainsi, dans la systématique des Paguristes sud-africains, quelques

points douteux, qui concernent des spécimens isolés. Ils ne seront éclaircis que

lorsqu’on aura récolté un matériel plus nombreux, condition essentielle d’une

systématique valable, surtout lorsqu’on a affaire a des animaux aussi variables que

des Paguridae.

Ces espéces ne présentent guere d’affinités avec celles que l’on trouve entre les

Tropiques. En ce qui concerne leur répartition géographique le fait le plus frappant

qui apparait lorsqu’on considere la figure 70, c’est la distribution des deux especes

parentes, P. gamianus et P. barnardi: la premiére est surtout representée sur la cote

occidentale d’Afrique du Sud ou s’exerce l’influence des courants froids, alors que la

seconde n’a été trouvée que sur la céte sud, baignée par le courant chaud des

Aiguilles.

On posséde peu de renseignements écologiques sur les espéces sud-africaines ;

la plupart des échantillons proviennent de la région de la zone intercotidale ou des

faibles profondeurs, sauf le P. agulhasensis dragué par 55 metres et le P. macrotrichus,

ramené de 90 meétres.

En raison de importance que présentent—comme nous lavons indiqué plus

haut—les pléopodes sexuels males dans la systématique du genre Paguristes nous

avons fait précéder l’étude proprement dite des espéces ouest-et sud africaines d’un

chapitre consacré a la morphologie et a la physiologie de cet appareil génital externe

que nous avons comparé a celui de quelques Anomoures. La courte diagnose du

genre Paguristes est suivie d’un tableau de détermination valable pour les espéces

qui nous intéressent. La description de chacune d’elles est suivie de quelques

remarques sur sa position systématique et sur sa répartition. Elles ont été classées

suivant l’ordre géographique nord-sud et ouest-est.

Les tailles minima et maxima ont été données pour chaque échantillon: nous

avons choisi comme dimension la longueur de la carapace (l.c.) mesurée depuis la

pointe du rostre jusqu’au bord postérieur de la carapace, ce qui permet des

comparaisons beaucoup plus significatives que la longueur totale.

Les abréviations utilisées sont p. 1, p. 2, p. 3, etc. . . . pour les chélipédes et les

appendices thoraciques suivants, et pl. 1, pl. 2, pl. 3, etc. . . . pour les appendices

abdominaux.

PAGURISTES D’AFRIQUE L’OUEST ET DU SUD 163

II. Les APPENDICES SEXUELS DES Paguristes MALES

L’un des caractéres les plus remarquables des Paguridae est la régression accen-

tuée des appendices abdominaux, uropodes exceptés. Chez la plupart cette régres-

sion se traduit par la réduction, plus forte chez les males que chez les femelles, des

appendices situés sur le coté gauche et par la disparition complete de ceux de droite.

A cété de ces formes chez lesquelles est réalisé le type pagurien parfait, il en existe

d’autres qui sont encore pourvues d’une ou de deux paires d’appendices dans la

région antérieure de abdomen. Les plus primitifs a cet égard sont des Pagurinae,

les Paguristes, dont il est plus particuliérement question ici, et les Paguropsis. Dans ces

deux genres les femelles ont conservé des pl. 1 pairs et les males des pl. 1 et des pl. 2

pairs. Aucun autre représentant de la sous-famille ne présente plus de pléopodes

symétriques, mais chez une espéce australienne, T7rizopagurus strigimanus (Miers) 11

existe, sur le deuxieme somite abdominal du male, un petit appendice droit différant

seulement du gauche par la taille plus réduite et par la plus grande variabilité de

lexopodite (Forest, 1952c, p. 8, fig. 8).

Chez les Eupagurinae la persistance d’appendices pairs sur l’abdomen peut

revétir divers aspects. Dans certains genres le male posséde des pl. 1 pairs, alors que

la femelle en est dépourvue, c’est le cas de Tomopaguropsis. La formule inverse est

observée chez Nematopagurus, Pylopagurus, Pagurodes, Ceratopagurus chez lesquels la

femelle seule a des pléopodes symétriques, ceux de la premiere paire. Enfin chez

Parapagurus, Sympagurus et Xylopagurus, si la femelle n’a que des pléopodes impairs

ou si méme ceux-ci ont disparu, le male est encore pourvu de pl. 1 et de pl. 2.

Lorsque les pl. 1 subsistent seuls, que ce soit chez le male ou chez la femelle,

ce sont des appendices d’un type assez simple et peu varié: un article basilaire

assez fort, un article distal en général foliacé, bordé de longues soies; leur role dans

la reproduction n’apparait pas nettement, bien qu’on les désigne souvent sous le

nom de pléopodes sexuels. Par contre lorsqu’il existe a la fois des pl. 1 et des pl. 2

pairs chez le male, il s’agit, a quelques exceptions prés,! d’organes tres différenciés

constituant les éléments d’un appareil copulateur dont la structure permet d’inter-

préter le fonctionnement et le rdle.

Les Paguristes, objets de la présente étude, ont des appendices males batis sur le

méme type, mais cette uniformité recouvre une grande diversité dans les proportions

des régions homologues et dans ’ornementation. On est ainsi amené, en systémati-

que, a accorder aux pl. 1 et aux pl. 2 des Paguristes une valeur identique a celle

qu’on attribue aux premiers appendices sexuels des Brachyures.

Morphologie et Physiologie

Nous avons choisi comme type d’appareil génital externe male de Paguristes

celui de P. oculatus Fabricius, espéce bien connue, d’assez grande taille et tout a fait

représentative du genre au point de vue qui nous intéresse maintenant (fig. 1).

1 Quelques espéces appartenant au genre Sympagurus présentent une variabilité extréme des

pléopodes pairs. Chez Sympagurus bicristatus A. Milne-Edwards par exemple en rencontre des males

pourvus des deux pl. 2 ou du gauche seulement. D’autres n’ont plus de pl. 2, mais deux pl. 1. Les

cas extrémes sont l’absence compléte de pléopodes pairs et trés rarement la présence simultanée des

pl. 1 et des pl 2. Ces appendices sont constitués par deux articles simples au plus, ils ne sont parfois

représentés que par un bourgeon.

164 ANNALS OF THE SOUTH AFRICAN MUSEUM

La premiére paire de pléopodes s’insére immédiatement en arriére du dernier

segment thoracique. Les deux appendices sont assez rapprochés, rabattus en avant

sur la région sternale thoracique, logés entre les coxae de la derniére paire de

péréiopodes. Ils se composent (fig. 2) de deux articles 4 demi-soudés: T’article

basilaire assez court a section sub-triangulaire et l’article distal foliacé, enroulé de

telle fagon qu’il forme une profonde gouttiére ouverte vers le plan sagittal. Lorsque

Fig. 1. Paguristes oculatus Fabricius: ¢ coxae des p. 5 et gonopodes.

animal est examiné par la face ventrale cet article distal parait donc constitué par

deux lames: l’une, que nous qualifierons d’inférieure (/.2.), visible en totalité au

premier plan; la lame supérieure, en contact avec le thorax, est en partie cachée par

la précédente. La lame inférieure est allongée, épaissie et plus fortement calcifiée

dans le voisinage du bord libre qui est garni de longs poils dirigés vers avant,

légérement spatulée dans la région antérieure; son bord distal est armé de nom-

breux crochets cornés recourbés vers l’extérieur. La lame supérieure est découpée

en deux lobes: un lobe distal triangulaire (/.d.) et un lobe interne (J. znt.) trés arrondi

couvert de longs poils.

PAGURISTES D’AFRIQUE L’OUEST ET DU SUD 165

On observe chez les adultes, au moment de la reproduction, a l’intérieur méme

des deux appendices, de nombreuses masses arrondies qui ont Vaspect de petites

oranges épluchées. Chacune de ces masses est constituée par un agrégat de cellules

a gros noyaux qui représentent sans doute des glandes dont la sécrétion intervient

dans la fixation des spermatophores sur la femelle.

Les pl. 2 (fig. 6) sont plus longs et plus gréles que les pl. 1; ils s’insérent beau-

coup plus loin sur abdomen et bien plus latéralement. Ils se composent d’un trés

court article basilaire (coxa rudimentaire), d’un second article long et mince,

presque rectiligne, obliquement rabattu vers le plan sagittal (basis) et d’un article

allongé, foliacé, enroulé sur lui-méme et formant ainsi une gouttiére ouverte vers

Pextérieur. Cet article parait résulter de la soudure partielle de endopodite pro-

prement dit et d’une piéce homologuable a V’appendix masculina de Thalassina et

peut-étre a appendix masculina et au stylamblys des Natantia supérieurs (Eucyphidea).

La région antérieure de chaque appendice a un bord libre €paissi, couvert de longs

poils dirigés vers l’avant; elle est logée a lintérieur de la gouttiere du pl. 1 corres-

pondant chez la plupart des Paguristes examinés.

D’apres la structure de ses constituants, on peut imaginer que le fonctionnement

de l’appareil génital externe des Paguristes est le suivant: au moment de la copulation

qui n’a jamais été observée dans ce genre” mais qui est sans doute comparable a

celle des Ecrevisses, décrite par plusieurs auteurs, la face ventrale du male viendrait

en contact avec la face ventrale de la femelle. Les premiers pléopodes du male

sappliquent contre les coxae de la quatrieme paire de pattes thoraciques de la

femelle. C’est ici qu’interviendraient les minuscules crochets qui arment le bord

antérieur des premiers pléopodes sexuels: fixés aux coxae de la femelle, dont la

région antérieure forme une créte anguleuse garnies de denticules, ces crochets

maintiendraient les appendices males en position. Les spermatophores sont émis

au niveau de l’ouverture postérieure de la gouttiére des pl. 1; Varticle distal des

pl. 2, jouant le rdle de balai, les pousse a l’intérieur de la gouttiére puis les expulse

au niveau des orifices de la femelle ou ils adhérent aux longs poils qui couvrent les

pieces sternales et la base des appendices thoraciques.

Le dispositif décrit ci-dessus est moins parfait mais comparable a celui qu’on

observe chez le Cancer pagurus § dont l’article distal des pl. 1 enroulé sur lui-méme

ne forme plus simplement une gouttiére mais un canal complet, a l’intérieur duquel

se déplace l’extrémité antérieure du second pléopode comme un piston dans un

cylindre, pour reprendre la comparaison de Balss (1944).

L’appareil genital externe chez quelques Anomoures

L’anatomie comparée de l’appareil génital externe male ne serait pas un

élément négligeable dans l’étude de |’évolution des Reptantia, mais ne permettrait

pas a elle seule de situer les divers groupes les uns par rapport aux autres. Ce n’est

pas dans le but de placer exactement les Paguristes parmi les Paguridae, ni les Paguridae

parmi les Anomoures que nous allons comparer les appendices sexuels males de

? Cavolini (1787) a décrit, mais de facon extrément peu précise, la copulation chez des Pagures

qui étaient peut-étre des P. oculatus.

ANNALS OF THE SOUTH AFRICAN MUSEUM

166

“EAS

HHATEN

Ne (( Un

Pléopodes sexuels droits chez quelques Anomoures g: en haut, pl. 1, en bas, pl. 2.

2, 6, Paguristes oculatus Fabricius; 3, 7, Sympagurus gracilipes A. M.-Edw.; 4, 8, Galathea squamifera

Leach; 5, 9, Thalassina anomala Herbst.

l.i., lame inférieure; L.int., lobe interne; 1.d., lobe distal; ex., exopodite; end, endo-

Figs. 2-9.

podite; a.m., appendix masculina; a.i., appendix interna.

PAGURISTES D’AFRIQUE L’OUEST ET DU SUD 167

Paguristes oculatus a ceux d’autres formes. Cette comparaison montrera simplement

les rapports qui existent a ce point de vue entre les Paguristes, les Eupagurinae, les

Pylochelidae et les Galatheidae, et a quel point un autre Anomoure, Thalassina, peut-

étre considéré comme primitif en ce qui concerne la structure de l’appareil génital

externe.

Les figures 3 et 7 représentent les pl. 1 et pl. 2 droits d’un Eupagurinae, Sympagurus

gracilipes A. Milne-Edwards. Les premiers appendices se composent comme chez

P. oculatus de deux articles. Le distal est foliacé mais ne présente qu’un léger enroule-

ment si bien qu’il n’est plus possible de parler de lames supérieure et inférieure. Par

ailleurs, les bords de l’article forment une courbe continue, si bien qu’il n’y a plus

de limite précise entre les régions que nous avons désigné sous le nom de lobe distal

et de lobe interne chez Paguristes oculatus. Quant aux pl. 2 ils sont assez trapus et

formés de trois articles comme chez les Paguristes. On observe en plus un exopodite

réduit a un bourgeon qui est d’ailleurs absent chez un autre Eupagurinae, Parapagurus

pilosimanus.

Les Pylochelidae dont nous n’avons pas de spécimens males en collection semblent

presenter une assez grande hétérogénéité des pléopodes pairs males. Les figures

données par Boas (1926, fig. 12 D, E et fig. 13 D, E) montrent qu’il existe de notables

différences entre les Mixtopagurus et les Pylocheles. Si les pl. 1 sont dans les deux cas

assez voisins et nettement plus petits que les pl. 2, ceux-ci sont pourvu chez les

premiers d’un exopodite qui manque chez les seconds. D’autre part l’article distal

des pl. 2 présente chez Pylocheles trois lobes séparés par de profondes encoches qui

paraissent correspondre a celles, moins marquées, qu’on observe sur le pourtour de

Particle distal chez P. oculatus.

Chez Galathea squamifera Leach (Galatheidae) on retrouve Penroulement accentué

de la région distale des pl. 1 (fig. 4). Les pl. 2 (fig. 8), plus longs et plus forts que les

pl. 1, ont une région distale trés élargie, légerement enroulée, et résultant aussi de la

fusion partielle de deux articles. Il existe un exopodite rudimentaire comme chez

Sympagurus gracilipes.

Si les différences les plus importantes entre les pléopodes sexuels males de P.

oculatus et ceux des quelques Anomoures que nous avons jusqu’a maintenant passés

en revue, concernent surtout les proportions des différentes régions des appendices,

la fusion plus ou moins complete de certains lobes et la présence chez certains d’un

court exopodite, dans tous les cas ’appareil destineé a faciliter la fixation des spermato-

phores sur la femelle fonctionne a peu prés dans les mémes conditions. I] se compose

d’une partie droite et d’une partie gauche, qui interviennent sans doute simultané-

ment mais indépendamment lune de l’autre: les spermatophores issus de chacun

des deux orifices sexuels male sont véhiculés par les appendices situés du cété corres-

pondant. Chez Thalassina anomala (Herbst) les deux premiéres paires d’appendices

abdominaux sont modifiés en gonopodes chez le male, mais on n’a plus ici un

appareil double: les quatre pléopodes forment un appareil unique. Chacun des

pl. 1 (fig. 5) est constitué par une piéce résultant de la cohalescence de plusieurs

articles dont les limites sont encore visibles. I1 comprend une lame principale

bordée de longues soies sur son bord externe, un lobe distal saillant et arrondi, un

168 ANNALS OF THE SOUTH AFRICAN MUSEUM

lobe antéro-interne couvert de petits poils crochus constituant un rétinacle. Les

pl. 2 (fig. 9) ont une structure plus compliquée que chez les formes précédentes:

endopodite comprend une piéce basilaire allongée, déprimée, sur laquelle

s'articule, en avant, un flagelle sub-cylindrique constitué par de courts articles plus

ou moins cohalescents, et une piéce de méme longueur que la portion basilaire et

également déprimée, que Boas désigne sous le nom d’appendix masculina. Un lobe

interne situé dans la région proximale de l’appendix masculina et qui forme rétinacle

serait suivant le méme auteur a homologuer a l’appendix interna des Eucyphidea.

L’exopodite multi-articulé est sub-cylindrique et égal aux deux tiers environs de

lendopodite; son article proximal est beaucoup plus long que les suivants. Dans

leur complexité les pl. 2 de Yhalassina anomala male paraissent bien plus proches

de ceux de certains Astacura, Astacus astacus L. par exemple (voir Balss 1941, p. 172,

fig. 230) que de ceux des Paguridae.

Nous avons dit qu’il s’agissait d’un appareil unique: en effet, au moment de la

copulation, les deux pl. 1 maintenus en contact par les rétinacles forment une seule

gouttiére dont le bord postérieur s’ouvre au niveau des orifices sexuels qui sont trés

rapprochés. Les spermatophores sont poussés a l’intérieur, puis a l’extérieur de

cette gouttiére par les extrémités des pl. 2 accolés eux aussi l’un a l’autre par leurs

rétinacles.

Utilisation des appendices sexuels males pour la détermination des Paguristes

Chez les Brachyures, les premiers pléopodes des males adultes se sont révélés

comme d’excellents caractéres et permettent souvent de distinguer avec une facilité

relative des espéces par ailleurs fort voisines. Nos observations nous ont montré

qu’il en était de méme pour les Paguristes: dans ce genre, les premiers et aussi les

seconds appendices abdominaux présentent, d’une part, une grande constance de

forme, de proportions et d’ornementation chez les males adultes d’une espéce

donnée, et d’autre part, des differences caractéristiques d’une espéce a lautre dans

de nombreux cas.

La technique d’utilisation des pl. 1 et des pl. 2 males dans la systématique des

Paguristes est trés simple. Les appendices sont désarticulés a la base et détachés

sous la loupe binoculaire a l’aide d’une aiguille lancéolée; ils sont ensuite immergés

pendant quelques instants dans l’alcool absolu, qu’ils proviennent d’animaux con-

servés dans l’alcool a 70° ou dans l’eau formolée. On les monte enfin a la résine

mastic dans la position qu’ils occupent sur le Pagure lorsqu’on |’examine par la face

ventrale. Dans beaucoup d’espéces les pl. 1 présentent un enroulement trés accentué

de l’article distal si bien que la lame inférieure est enticrement par dessus et cache

une partie de la lame supérieure; il est donc nécessaire de retourner la préparation

pour voir la totalité du lobe distal et du lobe interne. Lorsque l’enroulement est

plus faible; ce qui est le cas pour plusieurs espéces sud-africaines, l’article distal

s’étale entre lame et lamelle et son contour est entiérement visible du méme cété de

la préparation.

Les seconds pléopodes ont également une région distale plus ou moins enroulée

en hélice. I] n’est pas toujours possible d’étaler parfaitement cette région en pressant

PAGURISTES D’AFRIQUE L’OUEST ET DU SUD 169

sur la lamelle, et il arrive que les différents articles des appendices n’occupent pas

une position absolument comparable d’une préparation a l’autre. Les dessins ont

été exécutés avec la face la plus pileuse de la région distale par dessus.

Les figures 42 a 54 qui représentent les pl. 1 des Paguristes males des cétes occi-

dentales et méridionales d’Afrique montrent la diversité d’aspect de ces appendices

et les points sur lesquels ils different: présence ou absence de crochets sur le bord

Figs. 10-12. Trois états du développement

du premier pléopode sexuel chez Paguristes

gamianus H. Milne-Edwards: 10, jeune ¢ a

carapace de 4 mm.; jeune ¢ a carapace de

5°5 mm.; 12, g adulte 4 carapace de 12 mm.

antérieur de la lame inférieure, proportions de cette lame, forme du lobe interne.

Les pl. 2 sont sans doute moins différenciés mais dans de nombreux cas la portion

distale de l’endopodite a une forme caractéristique. Il faut noter que les pléopodes

sexuels ne sont pas toujours de bons caractéres, et que P. gamianus ne parait guére

pouvoir étre distingué de P. barnardi par les pl. 1.

Pour que les comparaisons prennent toute leur valeur il est nécessaire qu’elles

portent sur des appendices complétement formés; il est probable que la longueur

par rapport a celle du corps, la forme et l’ornementation des pléopodes ne sont

acquises qu’a la maturité sexuelle. C’est ce que montrent les figures 10 4 12 qui

représentent, au méme grossissement, trois états de développement du pl. 1 droit

170 ANNALS OF THE SOUTH AFRICAN MUSEUM

chez P. gamianus. Le premier appartient a un individu a carapace de 4 mm., le

second est celui d’un spécimen a carapace de 5:5 mm., le dernier enfin est celui

d’un adulte a carapace de 12 mm. Entre les deux extrémes, alors que le rapport des

tailles est de 3 a 1, le rapport des longueurs des appendices est de 5 a 1.

III. SysTEMATIQUE

PAGuRIDAE Dana

Pagurinae Ortmann

Gen. Paguristes Dana

Rostre variable, parfois trés réduit mais toujours présent. Ecailles oculaires

bien développées, plus ou moins écartées. Ecailles antennaires fortes, denticulées.

Flagelle antennaire de longueur moyenne ou court. Endopodite des maxillules avec

un petit appendice latéral externe. Flagelle de ?endopodite des premiers maxilli-

pédes dirigé vers l’intérieur et non vers l’avant. Maxillipedes externes rapprochés

ala base. Chélipedes de méme forme, le plus souvent sub-égaux, a ongles générale-

ment cornés, a doigts mobiles dans un plan horizontal. Dactyle des quatriéme

pattes thoraciques terminal, ne formant pas une pince avec le propode.

13 paires de branchies (pas de branchies sur le dernier segment thoracique).

Chez le male une paire de pléopodes, modifiés en gonopodes, sur chacun des

deux premiers segments abdominaux. Sur chacun des trois segments suivants un

seul pléopode—le gauche—dont la rame externe est bien développée et la rame

interne rudimentaire.

Chez la femelle, une paire de pléopodes uniramés sur le premier segment

abdominal; sur les trois segments suivants, un pléopode gauche aux deux rames bien

développées, plus fort que chez le male; sur le cinquiéme segment un pléopode

gauche de méme type que ceux du male. Souvent un repli de Pabdomen prenant

naissance en arriére du dernier pléopode biramé et formant chambre incubatrice.

Les Paguristes ont une trés large répartition; on en rencontre dans la plupart

des mers chaudes ou tempérées, depuis la zone littorale jusqu’a des profondeurs de

plusieurs centaines de metres.

Si les Paguristes d’Afrique du sud paraissent former un groupe a part, dont les

représentants n’ont pas, jusqu’a present, été signalés en d’autre régions, rien ne

prouve cependant quil y ait un cloisonnement géographique et que certains

d’entre eux ne seront pas retrouvés dans des localités beaucoup plus septentrionales.

C’est la raison pour laquelle nous avons rassemblé les espéces des cdtes occidentales

et méridionales d’Afrique dans un tableau unique qui permettra de les distinguer

les unes des autres.

Tableau de détermination des Paguristes des cétes occidentales et méridionales d’ Afrique.

1. Ecailles oculaires uni—ou, rarement, bidentées .. os

Ecailles oculaires a bord antérieur armé de 3 dents au moins. .. :

2. Flagelles antennaires de méme longueur ou plus courts que les atiloawles semilenese

Flagelles antennaires nettement plus longs que les pédoncules oculaires

Hes eo Out

10.

Palys

12.

13.

14.

15.

PAGURISTES D’AFRIQUE L’OUEST ET DU SUD 171

. Diamétres des cornées sensiblement égal a la moitié du diameétre maximum des pédoncules

oculaires (fig. 25). Afrique du Sud, littoral. .. a i Paguristes engyops (p. 204).

Diamétre des cornées légérement inférieur au diamétre maximum des pédoncules oculaires (fig.

14). Cdtes du Sahara, 115-290 metres. ore oH a Paguristes marocanus (p. 175).

Chélipédes paraissant finement granuleux; le gauche beaucoup plus fort que le droit. Une large

tache rouge sur la face interne de la main. Maroc, A.O.F., jusqu’a 150 metres. Angola (?).

Paguristes oculatus var. rubro-pictus (p. 172).

Chélipédes tuberculés ou épineux, sub-égaux. .. ay. Ss sk ih Be sais Sige

Face interne du propode des pattes ambulatoire revétue de poils plumeux trés serrés dissimulant

compléetement l’ornementation de cette face. Région supérieure du dactyle des mémes appen-

dices armée seulement de fines épines cornées, cachées sous une frange de poils extrémement

épaisse (fig. 39, et pl. IV). Afrique du Sud, littoral... ae Paguristes barnardi (p. 208).

Pilosité des pattes ambulatoires beaucoup moins forte, laissant apparaitre l’ornementation du

tégument. .. sie a: at Ba Sc Ae a aye = 33 is, Os

Ecailles oculaires modérément écartées. Bord supérieur du carpe, du propode et du dactyle des

p. 2 avec de fortes dents a base calcifiée, a pointe cornée. Des epines cornées sur la face interne

du propode de ces appendices (fig. 38, et pl. IV). Afrique du Sud, jusqu’a 24 metres.

Paguristes gamianus (p. 200).

Ecailles oculaires trés écartées. Des dents beaucoup moins fortes sur les p. 2. Pas d’épines cornées

sur la face interne du propode de ces appendices. .. i on oH ¥: Sesuke ny

Ecailles antennaires avec 4 a 5 dents sur le bord interne (fig. 27). Afrique du Sud, 90 métres.

Paguristes macrotrichus (p. 211).

Ecailles antennaires avec une seule dent sur le bord interne (fig. 26). Afrique du Sud, 55 métres.

Paguristes agulhasensis (p. 205).

Ecailles oculaires beaucoup plus longues que larges._ .. a aE aE ang Sigs

Ecailles oculaires sensiblement aussi longues que larges. Se uss Ay ig Bay pli

Pédoncules antennulaires a peine plus ihe que les a oa oculaires (fig. 19). Sénégal,

6-15 metres. sin é Be Paguristes rubrodiscus (p. 189).

Pédoncules antennulaires Lyne tn les yeux de ie moitié de la longueur de leur dernier article

au moins. wi ie ay ig ahs oe sh ua as Be sc SLO:

Pédoncules antennaires a peu pres de méme longueur que les pédoncules oculaires (fig. 20).

Guinée frangaise, 8-10 metres. .. ; : os . .Paguristes microphthalmus (p. 191).

Pédoncules antennaires dépassant les yeux de la moitié de ia longueur de leur dernier article.

(fig. 22). Liberia, Congo, 12-15 metres. si bie Se . .Paguristes hispidus (p. 196).

Ecusson céphalothoracique sensiblement aussi long que large. aH Mi ae ‘ot SOR

Ecusson céphalothoracique nettement plus long que large. Be si an be Be LG:

Pédoncules antennaires et oculaires de méme longueur. Angola, 72 métres.

. Paguristes staaee e 197).

Pédoncules antennaires n’atteignant pas tout a fait la base des cornées (fig. 21). Guinée fran-

caise, Congo, 8-36 metres. Le m si ae oe .. Paguristes virilis (p. 193).

Rostre dépassant nettement les pointes latérales (fig. 15). Sénégal, Mauritanie, 5-50 metres.

reser Saget (p. 176).

Rostre atteignant au plus l’alignement des pointes latérales. .. spe se an

Propode des p. 2 a peine deux fois plus long que haut et a bord supérieur trés fortement denté

(fig. 33). Sénégal, 8-40 metres. i bes .. Paguristes oxyacanthus (p. 186).

Propode des p. 2 au moins deux fois et demie athe long que haut et a bord supérieur armé de

dents aigués mais petites. aie oe Be ae whe me ae se AiG ti

Main des chélipédes ovale, bord antérieur des pléopodes 1 du ¢ inerme (fig. 44). Mauritanie,

Sénégal, Congo, 5-85 metres. .. .. Paguristes mauritanicus (p. 179).

Main sub-triangulaire, bord antérieur ais pléopore I di na armé de crochets (fig. 45). Sénégal,

8-10 metres. ae ; 2 Paguristes difficilis (p. 183).

172 ANNALS OF THE SOUTH AFRICAN MUSEUM

Paguristes oculatus Fabricius

var. rubro-picitus A. Milne-Edwards et E. L. Bouvier

Figs. 13 et 28

Paguristes maculatus var. rubro-picta, A. Milne-Edwards et E. L. Bouvier, 1892, p. 207.

Paguristes oculatus var. brunneo-pictus, A Milne-Edwards et E. L. Bouvier, 1900, p. 165,

pl. VI, fig. 1.

Paguristes oculatus var. brunneo-pictus, ‘T. Odhner, 1923, p. 6.

Materiel examiné: 1°. Les spécimens récoltés par le ‘““Travailleur’ et par le

‘““Talisman”’ et décrits par A. Milne-Edwards et E. L. Bouvier (types de la variété).

2°. 1 g, l.c. 13°5 mm., Cap de Naze (région de Dakar), 28-30 m., févr. 52, Cremoux

coll. 3°. 1 9 lc. 4 mm., Ile Bissagos, 120 m., “‘Président Théodore Tissier’, 1936,

Station 718.

Description: L’écusson céphalothoracique (fig. 13) est toujours notablement plus

long que large et représente plus de la moitié de la longueur totale de la carapace.

Le rostre long et aigu dépasse de beaucoup les dents latérales et atteint a peu prés le

milieu des écailles oculaires. Les pédoncules oculaires sont légerement renflés a la

base et au sommet; le gauche, en général un peu plus grand que le droit, a une

longueur comprise entre les 3/4 et les 4/5 de celle de la région pré-cervicale. Les

cornées sont échancrées postérieurement. Les écailles oculaires sont largement

écartées, leur moitié distale est triangulaire, l’extrémité est acuminée et le bord

antéro-latéral entier. Les pédoncules antennulaires atteignent ou dépassent légeére-

ment le bord postérieur des cornées. Les pédoncules antennaires arrivent au tiers ou

au quart distal des pédoncules oculaires; le deuxiéme article est armé d’une épine

au bord antéro-interne et de deux longues épines a l’angle antéro-externe: le

sommet bidenté de l’écaille antennaire n’atteint pas tout a fait l’extrémite distale

du dernier article; son bord interne présente quatre a neuf dents acérées, et son

bord externe, deux ou trois dents.

Le chélipéde gauche est en général plus long et plus gros que le droit: la

largeur de la main gauche est égale aux 3/4 de sa longueur environ, alors que la main

droite est deux fois plus longue que large. La face supérieure du carpe, du propode

et du dactyle des deux chélipédes est couverte de nombreux petits tubercules coni-

ques a pointe cornée, uniformément répartis, et qui, sur les bords des articles,

prennent laspect de faibles dents cornées.

Les pattes ambulatoires dépassent légérement les chélipédes. Les p. 2 (fig. 28)

ont un mérus a bord inférieur armé de denticules aigus; le bord supérieur du carpe

est défini par une rangée de sept 4 neuf dents cornées, celui du propode par sept a

dix dents, et celui du dactyle par une ligne des denticules dont la taille diminue

réguliérement de l’arriére vers l’avant; le bord inférieur et la face interne du pro-

pode sont faiblement spinuleux et le bord inférieur du dactyle est armé de longues

et fines épines cornées. Les p. 3 présentent, aux mémes endroits que les précédentes,

des dents beaucoup plus réduites.

PAGURISTES D’ AFRIQUE L’OUEST ET DU SUD

173

—

=m | Cro

Figs. 13-17.

Région antérieure de la carapace et appendices céphaliques antérieurs:

Paguristes oculatus var. rubro-pictus A. M.-Edw. et E. L. Bouvier; 14, P. marocanus A. M.-Edw. et

13;

E. L. Bouvier; 15, P. fagei Forest; 16, P. mauritanicus Bouvier; 17, P. difficilis Forest.

174. ANNALS OF THE SOUTH AFRICAN MUSEUM

Les femelles possédent un orifice sexuel sur chacune des coxae des p. 3; un

vaste repli de abdomen qui prend naissance en arriére du troisiéme pléopode

impair, recouvre la plus grande partie de le ponte.

La lame principale des pl. 1 du male s’élargit distalement en spatule; son bord

antérieur est armé de nombreuses et fortes épines en crochets, disposées en plusieurs

rangées dans la région médiane. Le lobe interne est séparé par une profonde

encoche aigué du lobe distal qui n’atteint pas tout a fait l’apex de la lame

inférieure.

La coloration des régions calcifiées est d’un blanc-rosé plus ou moins maculé

de rouge. Les principales marques persistantes sont des taches d’un rouge pourpre

mélé de violet, situées sur les faces interne et externe du mérus des chélipédes prés

de l’articulation du carpe; il y a aussi une large tache sur la face interne de la

région palmaire, en arriére du doigt mobile. Les p. 2 et les p. 3 présentent deux

larges anneaux rouge, l’un sub-distal, ’autre proximal, sur le dactyle, et un anneau

proximal sur le propode.

La pilosité de cette variété est plus forte que celle de la forme typique. Les

chélipédes sont recouverts d’une pubescence qui prend laspect d’une courte frange

sur les bords externes de la main.

Remarques: Le Paguristes oculatus Fabricius est avant tout une espéce méditer-

ranéenne, mais on |’a signalé aussi dans |’Atlantique entre la céte méridionale du

Portugal et le Gap Mazagan. Les spécimens récoltés au large de la cote occidentale

du Maroc et du Soudan par le “Travailleur’”’ et par le ““Talisman”’ ont été décrits

sous le nom de P. oculatus var. brunneo-pictus par A. Milne-Edwards et E. L. Bouvier,

mais huit ans auparavant, dans des observations préliminaires sur ce matériel, ces

auteurs avaient propose pour la variété le nom de rubro-picta qui—la loi de priorité

s’appliquant ici sans qucun doute—doit étre substitué a celui de brunneo-pictus.

Un male, beaucoup plus grand que ceux du “Travailleur” et du ““Talisman’’,

dragué au sud de Dakar et vivement coloré, répond a la description de la variété,

ainsi qu’une petite femelle capturée au large des [les Bissagos. Ces spécimens ne

présentent que peu de differences morphologiques avec ceux de Méditerranée. Les

pléopodes sexuels en particulier, qui permettent de distinguer facilement des

Paguristes fort voisins, sont ici identiques dans les deux cas. La distinction ne repose

en fait que sur la pigmentation (et encore ne s’agit-il que de taches colorées supple-

mentaires ou plus intenses), sur la pilosité plus forte, sur la taille plus faible, et sur

une gracilité plus grande des p. 2 et p. 3 dans la variété. Ces caractéres sont pro-

bablement étroitement liés aux facteurs externes; profondeur, conditions physico-

chimiques du milieu, etc. ... Gependant nous n’avons pas rencontre d’intermédiaires

entre la forme typique et la variété, et nous continuerons a les distinguer tout au

moins provisoirement. |

Si les spécimens signalés par Odhner sous le nom de P. oculatus var. brunneo-

pictus appartiennent bien a cette espéce, elle s’étendrait vers le sud jusqu’a l’Angola.

PAGURISTES D’AFRIQUE L’ OUEST ET DU SUD 175

Paguristes marocanus A. Milne-Edwards et E. L. Bouvier

Figs. 14, 29, 42, 55

Paguristes marocanus, A. Milne-Edwards et E. L. Bouvier, 1891, p. 152.

Paguristes marocanus, A. Milne-Edwards et EK. L. Bouvier, 1892, p. 207.

Paguristes marocanus, A. Milne-Edwards et E. L. Bouvier, 1900, p. 167—70, pl. XXIII,

fig. 1-6.

Matériel examiné: 1 3, l.c. 3°5 mm., “Talisman’’, 8.7.83, devant le Cap Bojador,

PouNect 17. 6, W., 130 m. (type dessinc). 1 g,1.c. 4 mm., “Talisman” 13.7.83, au

nord du banc d’Arguin, 21° 51’ N. et 19° 48’ W., 115-140 m. (type). Les autres

spécimens mentionnés par les auteurs ne figurent pas dans la collection.

I 4g, Le. 3 mm., cdtes du Sahara, 1884, de Cuverville coll.

Description: L’écusson céphalothoracique (fig. 14) est tres allongé et deux fois

plus long que la région postérieure de la carapace; sa largeur représente les 4/5

environ de sa longueur. Le rostre aigu est aussi saillant que les dents latérales dont

il est séparé par deux profondes concavités. Les pédoncules oculaires sont assez

larges et renflés aux extrémités; leur longueur représente les 5/8 environ de celle de

Pécusson céphalothoraciaque. Les cornées sont tres grandes. Les écailles oculaires

sont écartées, longuement lancéolées, et ne présentent aucune indentation sur le

bord antéro-latéral. Les pédoncules antennulaires dépassent les yeux de la moitié

de leur dernier article environ. Les pédoncules antennaires atteignent les cornées;

le deuxieme article est armé d’une seule épine a chacun de ses angles antérieurs.

L’écaille antennaire atteint le milieu du dernier article; son extrémité est bidentée,

et elle présente deux dents sur son bord interne qui est concave et trois dents sur son

bord externe; le flagelle est a peine plus long que les pédoncules oculaires.

Les chélipédes sont sub-égaux. Le carpe est aussi long que le dactyle et plus

long que le bord palmaire interne. La main est deux fois plus longue que large. Les

bords interne et externe de la face supérieure du carpe sont marqués, le premier par

quatre dents assez fortes, le second par une rangée irréguliere de petits tubercules.

I] existe aussi des tubercules sur la face supérieure de la main, trés arrondis et peu

nombreux dans la région médiane, un peu plus denses et un peu plus saillants sur les

cotés et notamment sur le bord palmaire interne qui est armé de quatre ou cing

petites dents.

Les pattes ambulatoires p. 2 (fig. 29) et p. 3 dépassent largement les chélipedes;

elles sont gréles et leurs deux derniers articles sont peu déprimés latéralement. Les

bords supérieur et inférieur du mérus des p. 2 sont spinuleux; le bord supérieur du

carpe est armé de six dents acérées et celui du propode de cing dents; le rapport des

longueurs du propode et du dactyle est légerement supérieur a 2/3. Les p. 3 sont

inermes a l’exception du bord supérieur du carpe qui présente l’épine distale

habituelle; la coxa est nettement plus longue et le mérus nettement plus court que

pour p. 2-

L’article proximal des pléopodes 1 du male (fig. 42) n’est que légérement plus

court que l’article distal. La lame inférieure de ce dernier est prés de quatre fois

176 ANNALS OF THE SOUTH AFRICAN MUSEUM

plus longue que large, le bord antérieur est armé de huit ou neuf courts crochets;

le lobe interne atteint et le lobe distal dépasse apex de la lame inférieure.

Nous n’avons pas pu examiner de femelle de cette espéce.

Les spécimens étudiés sont tous décolorés et d’une teinte blanchatre uniforme.

D’aprés A. Milne-Edwards et E. L. Bouvier “la couleur dans l’alcool tire sur le

rose-chair, avec quelques taches blanchatres’’.

La pilosité générale est assez faible; cependant de trés longs poils garnissent

le bord antérieur des écailles oculaires et ’ornementation des chélipédes est en

grande partie dissimulée par les poils qui les recouvrent et qui sont salis par la vase.

Remarques: Cette espéce au nom assez mal choisi puisque tous les spécimens

connus ont été capturés bien au sud des cétes marocaines? ne risque d’étre confondue

avec aucun des autres Paguristes de la cote occidentale d’Afrique. La région pré-

cervicale beaucoup plus longue que la région postérieure, le bord frontal aux trois

dents aigués aligneées, les écailles oculaires aigués, a bord antérieur entier et frangé

de longs poils, la briéveté des flagelles antennaires permettent de lidentifier rapide-

ment. A en juger par les spécimens examinés, et qui sont des males adultes dont le

plus grand a une carapace de 4 mm., c’est une espéce de petite taille, Pune des plus

petites du genre. C’est aussi une espéce qui vit a une assez grande profondeur,

puisque le ““Talisman”’ l’a capturée sur des fonds de 115 a 290 m. Le fait qu’on ne

ait signalée qu’au large des c6tes du Sahara ne signifie pas que sa répartition

eéographique est aussi restreinte. Le matériel récolté au-dela des fonds de 120 m.

est en effet pour linstant relativement peu abondant. Le Paguristes qui parait

présenter le plus d’affinités avec P. marocanus appartient a la faune sud-africaine:

c’est P. engyops Barnard qui vit dans la zone littorale. Nous verrons les rapports qui

existent entre les deux espéces a la suite de la description de la seconde.

Paguristes fager Forest

Figs. 15, 30, 43, 57

Paguristes fager Forest, 1952, p. 260, fig. 3.

Matériel examiné: De nombreux individus § et 9 de 1:5 a 7 mm. (l.c.) devant

Thiaroye, baie de Rufisque, dragage sur fond coquillier par 10 a 15 m., 18-1-52,

Paraiso coll. (Syntypes).

2 det 3 9, lic. 3-5 a 5-5 mm.: Cap Blanc, 50 m., 1907, Gruvel coll.

1 9, Le. 1-8 mm.: chalutier “Cabellou’’, au large du Sénégal, 12 a 15 m.,

10-1-41, Monod coll.

3 6 et 16 9, lc. 3a 6 mm.: chalutier “Vers l’Horizon’’, au large du Sénégal,

8 m., fond caillouteux a grosses Ascidies, 21-1-41, Monod coll.

6 g et 2 9 ovigeres, l.c. 2 4 5 mm.: Gorée, filets a langouste, 23, 24-8-50,

Paraiso coll.

6 g et 9, M’Bao (Sénégal), dragage a un mille de la céte, 5-14 m., 7-2-51,

Delais coll.

3 Milne-Edwards et E. L. Bouvier l’ont sans doute ainsi baptisée parce que l’un des spécimens se

trouvait dans une coquille de Stnistralia marocana,

PAGURISTES D’AFRIQUE L’OUEST ET DU SUD 177

Figs. 18-22. Région antérieure de la carapace et appendices céphaliques antérieurs: 18,

Paguristes oxyacanthus Forest; 19, P. rubrodiscus Forest; 20, P. microphthalmus Forest; 21, P. virilis

Forest; 22, P. hispidus A. M.-Edw. et E. L. Bouvier.

178 ANNALS OF THE SOUTH AFRICAN MUSEUM

10 get 12 2 dont 8 ovigéres, l.c. 2 4 7 mm., chalutier “Maid Honour’, 8° 38’ N.,

a 8° 42’ N., 8 a 12 m., 1 au 8-3-48, Cadenat coll.

Description: Les régions pré-et post-cervicale de la carapace sont sensiblement

de méme longueur; la premiere (fig. 15) est un peu plus longue que large, le rapport

de ses dimensions étant de 5/4 environ. Le rostre, large et en angle obtus, dépasse

légérement Valignement des deux dents latérales. Les pédoncules oculaires, un

peu plus courts que le bord frontal, sont renflés aux extrémités, et tout particuliére-

ment dans la région proximale chez les jeunes individus. Les cornées atteignent le

milieu du dernier article du pédoncule antennulaire et l’extrémité du pédoncule

antennaire. Les écailles oculaires sont grandes, larges, trés rapprochées et leur bord

antérieur est armé de cing a huit denticules. L’angle externe du deuxiéme article

des antennes est uni-ou bidenté. Les écailles antennaires atteignent au maximum le

milieu du dernier article du pédoncule; leur extrémité est plus ou moins nettement

bidentée, et elles sont armées de deux ou trois épines sur le bord interne légérement

concave et d’une épine sur le bord externe. Les chélipédes sont égaux et de méme

forme. Le bord inférieur du mérus forme une créte denticulée. La main est deux

fois plus longue que large, le doigt mobile est un peu plus long que le

bord palmaire interne. Un léger hiatus subsiste entre les doigts lorsque ceux-ci sont

fermés. La face supérieure des trois derniers articles porte de nombreux tubercules

coniques, peu élevés et a pointe cornée, plus aigus vers les bords latéraux. Sur la

face supérieure de la main, en arriere de l’articulation du dactyle, existe une pro-

tubérance tuberculée plus ou moins développée. Les pattes ambulatoires, p. 2 et

p. 3 dépassent les chélipédes. Les p. 2 (fig. 30) ont un mérus a bord inférieur non

denticulé, un carpe a bord supérieur armé d’une rangée irréguliére de dents cornées

assez fortes qui se prolonge sur le propode sous la forme d’une ligne de dix a douze

dents cornées plus petites. Le dactyle a une section circulaire et son diamétre a la

base est compris quatre a cing fois dans sa longueur; le bord supérieur et le bord

inférieur sont marqués par une rangée d’épines minuscules largement espacees.

Cet article est a peu pres de méme longueur que le précédent. Les p. 3 sont peu

différentes des p. 2 dans leur forme générale, mais elles sont inermes a l’exception

d’une épine distale sur le bord supérieur du carpe et d’une rangeée de tres petites

épines sur le bord inférieur du dactyle.

La femelle ne posséde qu’un orifice sexuel situé sur la coxa de la p. 3 gauche,

et ne présente pas de repli de abdomen en arriére du troisieme pléopode impair.

Les pléopodes 1 du male (fig. 43) ont une lame inférieure a moitié distale élargie en

spatule et bordée de fines épines recourbées vers l’extérieur. Le lobe interne est

séparé du lobe distal qui dépasse l’apex de la lame inférieure par une encoche

arrondie.

La coloration de fond des régions calcifiées est d’un blanc jaunatre ou orange.

Les écailles oculaires, la base des pédoncules oculaires et une étroite région sous les

cornées sont rouge-orange, alors que la partie moyenne des pédoncules oculaires,

les pédoncules antennulaires et antennaires, et l’endopodite des maxillipédes externes,

sont d’un bleu plus ou moins intense. Les chélipédes sont maculés de rouge-orange.

La face interne du mérus présente une coloration constante: la région antérieure

PAGURISTES D’AFRIQUE L’OUEST ET DU SUD 179

est blanche, le reste d’un rouge-orange présentant un maximum d’intensité a la

limite des deux zones.. Les deux premieres paires de pattes ambulatoires présentent

également de grandes zones rouge-orange qui couvrent en particulier la moitié

proximale du propode et du dactyle.

Remarques: Il n’y a pas de raisons de comparer Paguristes fagei a l'une plutét

qu’a autre des espéces de la cote occidentale d’Afrique. I] posséde tout un ensemble

de caractéres, et notamment une pigmentation qui le distinguent de tous les autres

représentants du genre. C’est ainsi que le bleu intense des pédoncules oculaires, des

antennules et des antennes permet d’identifier au premier coup d’oeil les exem-

plaires frais, méme de trés petite taille. Au bout de quelques années dans l’alcool, il

se produit une décoloration presque totale, mais on peut, dans la plupart des cas,

reconnaitre la ligne brisée qui sépare la zone foncée de la zone claire sur la face

interne du meérus des chélipédes. D’autres caractéres stables et d’observation facile

sont utilisables par ailleurs: la forme de la carapace et surtout celle du bord frontal

avec le rostre qui dépasse légérement les dents latérales dont il est séparé par des

concavités peu profondes, les écailles oculaires courtes, larges et rapprochées, plutét

quadrangulaires que triangulaires.

Les pléopodes 1 du male sont aussi d’un type trés particulier et ne présentent

quelque ressemblance qu’avec ceux du P. oculatus.

P. fagei est un espece de petite taille: les plus grands exemplaires ont une

carapace de 7 mm., et chez les plus petites 2 ovigéres cette région du corps n’a pas

plus de 2-5 mm. Des ¢ de 2 mm. de carapace ont déja des pléopodes sexuels de

méme forme que chez les individus plus agés. Une partie des spécimens observés

étaient a l’intérieur de coquilles de Gastéropodes: Nassa, Fusus, Turritella, Tritonalia,

etc. ..., mais la plupart étaient logés a Pintérieur de colonies de Bryozoaires dont

chacune présente un aspect bien particulier: la masse centrale abritant le Pagure

se prolonge latéralement par deux longues cornes formant balancier et présente

dorsalement une créte digitée.

La répartition géographique de la nouvelle espéce, telle qu’on peut se la repré-

senter actuellement, ‘s’*étend du Cap Blanc (22° N. environ), au Sud de Dakar

(8° 38’ N.). Elle a été récoltée principalement entre 5 et 15 m. de profondeur, mais

Péchantillon du Cape Blanc a été dragué a 50 m.

Paguristes mauritanicus Bouvier 1906

Figs. 16, 31, 44, 56

Paguristes mauritanicus Bouvier, 1906 a, p. 186, fig. 1; 1906 b, p. 96, fig. 1; 1906 ¢,

p- 199.

Matériel examiné: 1 3, \.c. 7 mm.: au large de Novak Chott, 16-24 m., Gruvel

coll. (Type).

I 9, lc. 4°5 mm., 1 g l.c. 6-5 mm. et 1 9 l.c. 4 mm.: Baie de Hann (Dakar),

6-1o m., sable coquillier grossier, 1950, R. Sourie coll.

180 ANNALS OF THE SOUTH AFRICAN MUSEUM

a7),

Figs. 23-27. Région antérieure de la carapace et appendices céphaliques antérieurs: 23,

Paguristes gamianus H. M.-Edw.; 24, Paguristes sp. (Table Bay); 25, P. engyops Barnard;

26, P. agulhasensis sp. nov.; 27, P. macrotrichus sp. nov.

PAGURISTES D’AFRIQUE L’OUEST ET DU SUD 181

Nombreux individus 3 et 2 de 3 a 8 mm. (l.c.) dont plusieurs ovigéres: devant

Thiaroye, baie de Rufisque, 10-15 m., sable coquillier, 18-1-52, Paraiso coll.

1 9, l.c. 3 mm., 1 J, lc. 5 mm., M’Bao (Sénégal), dragage a un mille de la céte,

5-14 m., 7-2-52, Delais coll.

1 g, l.c. 6 mm.: “‘Mercator’’, dragage entre les Iles Tamara et Roume (Iles de

Los), 8-10 m., 10-12-36.

ieee, 4mm... 9 les 2-5;mimt 1) 2) less imm:, et 29 ovigeres, ics 4) et 5 mm.,

“M’Bizi’, St. 144, 26 milles SSW. Cap Lopez, 95-100 m., 10-3-49.

1 2 ovigere, l.c..5 mm.:. “M’Bizi’, St. 142,:29 milles'S.. Cap: Lopez, 50 m.;

9-3-49-

2 6, lc. 4:5 et 8-5 mm.: “M’Bizi’, St. 26, 25 milles SW. Pointe de Banda,

85 m., 6-9-48.

I 9 juv., l.c. 2°5 mm., 1 9, l.c. 6 mm.: “M’Bizi’, St. 8,15 milles SW. Moita Seca,

50 m., 3-8-48.

48 spécimens récoltés en 15 stations entre 14 et 44 m. par R. Bassindale au

large de la Gold Coast. Le nombre, la taille et le sexe des individus provenant

de chaque station figureront dans |’étude d’ensemble des Pagures récoltés par

R. Bassindale.

Description: La région postérieure de la carapace est de méme longueur ou un

peu plus courte que l’écusson céphalothoracique; celui-ci (fig. 16) a une largeur

sensiblement égale aux 6/7 de sa longueur; les aires latérales sont faiblement spinu-

leuses. Le rostre, en angle obtus, n’atteint pas l’alignement des dents latérales. Les

pédoncules oculaires sont renflés aux extrémités; le rapport de leur longueur a

celle de la région précervicale est compris entre 3/4 et 2/3. Les écailles oculaires sont

largement écartées; leur région distale est obliquement tronquée, et le bord antéro-

latéral est armé d’une épine apicale et de deux ou trois petites dents plus petites. Les

pédoncules antennulaires dépassent les yeux de la moitié de la longueur de leur

dernier article au moins. Les pédoncules antennaires atteignent le milieu des cornées;

le deuxieme article est armé de deux trois épines a l’angle antéro-externe qui est assez

saillant; l’écaille antennaire, dont l’extrémité bidentée arrive au quart ou au tiers

proximal du dernier article, est armée de une ou deux dents sur son bord interne

légérement concave et de deux ou trois dents sur son bord externe.

Le chélipéde droit est en général un peu plus fort que le gauche, mais de méme

forme. Le carpe, de méme longueur que le dactyle, est plus long que le bord

palmaire interne. La main est a peu prés deux fois plus longue que large, son

épaisseur maximum est comprise 2-4 fois environ dans sa longueur. La face supéri-

eure du carpe est armée de cing a six fortes dents sur le bord interne et de dents

plus petites irréguli¢érement disposées le long du bord externe. La face supérieure du

propode et du dactyle est couverte de tubercules coniques peu élevés qui ont plutét

l’aspect de gros granules chez les grands spécimens. Ces tubercules sont un peu plus

aigus sur le bord externe de la main et ce sont cing ou six fortes dents cornées qui

marquent le bord palmaire interne.

Les pattes ambulatoires de la premiere paire (fig. 31) dépassent largement les

chélipédes. Le bord supérieur du carpe est armé de cing ou six dents, Le bord

182 ANNALS OF THE SOUTH AFRICAN MUSEUM

supérieur du propode d’une douzaine de dents; cet article est a peu prés trois fois

plus long que haut, alors que la longueur du dactyie représente a peu pres sept fois

sa hauteur a la base. Le rapport des longueurs du dactyle et du propode, mesurés

le long de leur bord supérieur, est de 5/3 environ. Les p. 3 sont un peu plus gréles

et un peu plus longues que les p. 2; elles sont inermes a l’exception d’une épine

distale au bord supérieur du carpe.

Les femelles n’ont qu’un orifice sexuel qui s’ouvre sur la coxa de la p. 3 gauche.

Chez les femelles adultes, un large repli de Pabdomen qui prend naissance en

arriére du troisiéme et dernier pléopode biramé recouvre cet appendice et le précédent

et’ éventuellement, une grande partie de la ponte.

Les pléopodes 1 du ¢ (fig. 44) ont une lame inférieure quatre fois plus longue

que large, a bord interne légérement concave, et dépourvue de crochets sur le bord

antérieur; sur les préparations, le lobe interne, trés large et trés arrondi déborde

largement la lame inférieure et se rattache au lobe distal trés saillant par une

concavité réguliere et peu profonde.

Des exemplaires qui n’ont séjourné qu’un mois dans l’alcool, présentent la

coloration suivante: les régions calcifiées ont un fond blanchatre; il y a une large

tache brun-rouge en avant de la carapace, tout prés du bord frontal; les écailles

oculaires, la base des pédoncules antennaires, et les pédoncules oculaires tout entiers

sont d’un brun-rouge intense. Les chélipédes sont brun-violacé, les doigts et les

tubercules blancs. Les deux premiére pattes ambulatoires portent de larges anneaux

alternativement clairs et foncés. Sur les échantillons qui ont séjourné plus longtemps

dans l’alcool, les marques colorées s’éclaircissent et tendent a devenir rose. C’est la

coloration des pédoncules oculaires et des pattes ambulatoires qui subsiste le plus

longtemps.

Les poils plumeux trés denses qui couvrent la région frontale dissimulent

lornementation de cette région, mais laissent voir une grande partie des pédoncules

oculaires. Les appendices thoraciques sont aussi fortement pileux notamment sur

les faces supérieure et inférieure, cependant les doigts des chelipédes restent

toujours visibles.

Remarques: Le Paguristes mauritanicus n’était connu jusqu’a présent que par

Punique individu ¢ décrit par Bouvier. Les nombreux spécimens dont nous dis-

posons maintenant montrent qu’il ne s’agit pas d’une espéce rare. C’est de la région

de Dakar que proviennent la plus grande partie des échantillons, dragués depuis la

cote jusqu’a une profondeur de 20 4 25 m. Les individus capturés au sud de cette

région et jusqu’a l’embouchure du Congo, proviennent d’un niveau inférieur, de

50 a 100 m., et présentent une différence de coloration dans les pédoncules oculaires,

lesquelles ont une moitié proximale bleutée, et une moitié distale orange. Les autres

caractéres, et notamment la forme des pléopodes du 3, ne permettent pas de con-

sidérer ces spécimens comme appartenant a une forme différente.

L’espéce la plus voisine est le P. difficilis sp. nov. décrit plus loin, dont Paspect

est fort peu différent, et qui ne peut guére étre distingué que par la forme des

chélipédes et surtout par celle des pléopodes 1 du mle, trés caractéristiques.

PAGURISTES D’AFRIQUE L’OUEST ET DU SUD 183

Les coquilles abritant les P. mauritanicus appartiennent aux genres les plus divers,

Bullia, Cancellaria, Clavatula, Gibbula, Marginella, Mesalia, Nassa, Terebra, Trophon,

Turritella, etc. C’est en Baie de Rufisque, devant Thiaroye, et par 10 a 15 m. qu’ont

été capturés les plus nombreux spécimens de P. mauritanicus. Ce dragage a ramené

en outre d’autres espéces de Paguridae: Petrochirus pustulatus juv., Pagurus pectinatus

juv., Pseudopagurus granulimanus, Diogenes ovatus, D. pugilator var. intermedius, Eupagurus

sculptimanus et E. souriei, et enfin des spécimens de trois autres espéces nouvelles

de Paguristes: P. faget, P. rubrodiscus et P. oxyacanthus.

Paguristes difficilis Forest.

Figs. 17, 45, 61

Paguristes difficilis Forest, 1952, p. 259, fig. 1.

Matériel examiné: 8 § de 4 a7 mm., 2 9 de 7 mm.: Baie de Hann, 8-10 m.,

sable coquillier grossier, Janvier 1950, Sourie coll. Type.

Description: L’écusson céphalothoracique (fig. 17) est a peu prés de méme

longueur que la région postérieure de la carapace et un peu plus long que

large; ses aires latérales sont faiblement spinuleuses. Le rostre obtus et arrondi

n’atteint pas tout a fait Palignement des dents latérales. Les pédoncules oculaires

renflés aux extrémités, ont une longueur égale aux 5/6 de la région pré-cervicale;

les cornées sont profondément échancrées postérieurement. Les écailles oculaires

sont largement écartées; leur moitié distale est triangulaire; l’épine distale est

suivie de deux ou trois petites dents sur le bord antéro-latéral. Les pédoncules

antennulaires dépassent les yeux de la moitié de la longueur de leur dernier article.

Les pédoncules antennaires atteignent le milieu des cornées; le deuxieme article

est armé de trois dents a angle entéro-externe qui est assez saillant; la pointe de

Pécaille antennaire atteint le tiers proximal du dernier article, ses bords latéraux

sont armés de deux ou trois dents; le flagelle antennaire est égal aux 3/4 de la

longueur de la carapace.

Les chélipedes sont égaux. Le carpe est un peu plus court que le dactyle dont

la longueur représente les 2/3 environ de la longueur totale de la main. Celle-ci est

sub-triangulaire et présente son maximum de largeur pres de la base; son épaisseur

maximum est comprise 2-7 fois dans la longueur. Le bord interne de la face supéri-

eure de carpe est défini par une rangée de quatre fortes dents cornées, et le bord

externe par des dents plus nombreuses mais plus petites. Le propode et le dactyle

ont une face supérieure couverte de nombreux tubercules arrondis pourvus d’une

trés petite pointe cornée. Le bord interne de la main porte une rangée de quatre

fortes dents.

Le propode des pattes ambulatoires de la premiére paire est armé de douze

a quinze fortes dents cornées sur son bord supérieur; le bord supérieur du dactyle

porte également des petites dents cornées sur toute sa longueur. Le rapport des

longueurs du dactyle et du propode est de 3/2 environ. Les p. 3 sont inermes,

exception faite de la petite dent distale au bord supérieur du carpe.

184 ANNALS OF THE SOUTH AFRICAN MUSEUM

Figs. 28-33. Deuxiéme patte thoracique gauche, face interne, dénudée: 28, Paguristes oculatus

var. rubro-pictus A. M.-Edw. et E. L. Bouvier; 29, P. marocanus A. M.-Edw., et E. L. Bouvier;

30, P. faget Forest; 31, P. mauritanicus Bouvier; 32, P. rubrodiscus Forest; 33, P. oxyacanthus

Forest.

PAGURISTES D’AFRIQUE L’OUEST ET DU SUD 185

Les femelles n’ont qu’un orifice sexuel sur la coxa de p. 3 gauche. Un large

repli abdominal qui prend naissance en arriére du troisitme pléopode impair

recouvre les appendices précédent, et, éventuellement, une partie de la ponte. Les

pléopodes 1 du male (fig. 45) ont une lame inférieure a peine deux fois et demie plus

longue que large qui, sur les préparations, recouvre presque complétement le lobe

interne. Chez les adultes le bord antérieur de cette lame est armé de petites épines

recourbées vers l’extérieur. Le lobe distal est étroit et séparé du lobe interne par une

profonde concavité.

La région antérieure de la carapace, les écailles oculaires, la base des pédoncules

antennaires sont roses. Les pédoncules oculaires sont orange-clair. Les chélipédes

sont de la méme teinte avec les doigts blancs. Les pattes ambulatoires portent de

larges anneaux d’un rose plus foncé.

Les poils qui couvrent la région frontale et les appendices thoraciques sont

assez denses mais ne dissimulent pas completement l’ornementation du tégument.

Remarques: Cette espéce est la plus difficile a caractériser parmi les Paguristes

d’Afrique occidentale. En effet, elle présente une grande ressemblance dans |’aspect

de la carapace, des appendices céphaliques antérieurs et des pattes ambulatoires,

avec. P. mauritanicus. Si la région pré-cervicale est nettement moins allongée chez

P. difficilis, si P. mauritanicus a de plus des pédoncules oculaires un peu plus gréles, ce

sont la des caractéres assez variables et sur lesquels il serait dangereux de baser la

détermination. Par contre la forme des chélipédes différe sensiblement d’une espéce

a autre; chez P. mauritanicus, le main, subovale, présente son maximum de largeur

vers le milieu de l’article et le rapport de |’épaisseur maximum sur la longueur a une

valeur de 2-4 environ. Chez P. difficilis, la main plus finement granulée est sub-

triangulaire et présente son maximum de largeur un peu en avant de l’articulation du

carpe; elle est aussi moins renflée: le rapport de son épaisseur maximum sur sa

largeur ayant une valeur de 2-7 environ. Lorsqu’il s’agit de spécimens frais la dis-

tinction peut se faire d’aprés la pigmentation des pédoncules oculaires: de teinte claire

chez P. difficilis, d’un rouge-brun tres foncé chez P. mauritanicus. Mais pour les males,

le meilleur moyen de distinguer les deux espéces est d’examiner les pl. 1. Chez P.

difficilis, d’un rouge-brun trés foncé chez P. mauritanicus. Mais pour les males, le

meilleur moyen de distinguer les deux espéces est d’examiner les pl. 1. Chez P.

difficilis la lame inferieure est beaucoup plus large et, chez l’adulte, armée de crochets

sur le bord antérieur. Chez les plus petits exemplaires étudiés les crochets sont

absents, mais la forme générale est le méme que chez les individus plus Aagés et

Paspect bien différent de ce qu’on observe chez les P. mauritanicus au méme stade.

Les dix spécimens que nous avons entre les mains nous sont parvenus sans

coquilles, mais quelques-uns présentent l’aplatissement dorso-ventral et l’étirement

des plaques sternales thoraciques qui caractérisent les pagures vivant dans des

coquilles a étroite ouverture, Conus ou Marginella par exemple.

Paguristes difficilis n’a été récolté que dans une seule localité, au large de Dakar,

sur les mémes fonds que P. mauritanicus: le méme dragage a ramené trois spécimens

de cette derniére espéce.

186 ANNALS OF THE SOUTH AFRICAN MUSEUM

Paguristes oxyacanthus Forest

Figs. 18, 33, 47, 58

Paguristes oxyacanthus Forest, 1952, p. 261, fig. 5.

Matériel examiné: 1 9 juv., l.c. 7 mm.: Baie de Hann (Dakar), 8-19 m., sables

coquilliers grossiers, 7-1-50, Sourie coll.

1 Q juv., lc. 6-5 mm.: Baie de Hann, 12 m., cailloux et coquilles, 17-6-50,

Sourie coll.

3 9, lc. 9°5 a 11 mm., 2 G juv., lc. 5:5 et 7 mm.: devant Thiaroye, Baie de

Rufisque, dragage sur fond coquillier, 10-15 m., 18-1-52, Paraiso coll. Type.

I g, l.c. 12°5 mm.: sud de Gorée, 40 m., Février 52, Postel coll.

Description: L’écusson céphalothoracique est a peu prés aussi long que la région

postérieure de la carapace; le rapport de sa longueur a sa largeur est de 7/6 environ;

sa surface présente un relief tourmenté, avec une dépression transversale trés

marquee en arriére du bord frontal, avec des tubercules saillants et de nombreuses

spinules sur les aires antérieures et latérales; le rostre en angle obtus n’atteint pas

tout a fait Palignement des dents latérales.

Les pédoncules oculaires sont renflés a la base et, plus faiblement, au niveau

des cornées qui présentent une échancrure postérieure aigué; leur longueur est

égale aux 4/5 de celle de la région pré-cervicale. Les écailles oculaires sont écartées;

elles sont armées d’une €pine terminale et de trois petites dents sur le bord antéro-

latéral. Les pédoncules antennulaires dépassent les yeux de la moitié de leur dernier

article. Les pédoncules antennaires n’atteignent pas la base des cornées; l’angle

antéro-externe saillant du deuxiéme article est pourvu de quatre dents groupées par

deux. L’écaille antennaire atteint le tiers proximal du dernier article, son extrémité

est bifide, et ses bords latéraux portent trois fortes dents, celles du bord interne étant

groupées dans la moitié proximale; le flagelle est égal aux 2/3 de la carapace.

Les chélipédes sont é¢gaux et fortement épineux; le bord inférieur de la face

interne du meérus est défini par une douzaine de denticulations assez fortes. Le

carpe est plus long que le dactyle et celui-ci un peu plus long que le bord palmaire

interne. La face supérieure du carpe et du propode est réguli¢érement convexe de

avant vers l’arriére, mais parait plutét concave que convexe en section trans-

versale: en effet, les bords latéraux sont armés de trés fortes dents a pointes cornées

légérement inclinées vers avant et vers lextérieur: le bord externe du carpe est

armé de huit dents, le bord interne de quatre dents trés fortes; le bord externe du

propode est défini par une rangée de dents assez nombreuses, mais plus petites; par

contre, le bord interne ne porte que trois dents trés développées. Le reste de la face

supérieure de ces articles est couvert de tubercules coniques de plus petites tailles.

La face externe du dactyle c’est-a-dire celle qui est orientée vers le plan sagittal de

lanimal—est large et plate; elle est limitée en haut par une ligne de dents cornées

assez forte, en bas, par des tubercules coniques cornés peu saillants; il existe une

troisieme ligne longitudinale intermédiaire constituée par de petites épines cornées. —

Les pattes ambulatoires p. 2 et p. 3 dépassent largement les chélipédes. Les

p. 2 (fig. 33) ont un merus dont le bord inférieur est spinuleux. La région supérieure

PAGURISTES D’ AFRIQUE L’OUEST ET DU SUD 187

du carpe est armée de dix fortes dents cornées irréguli¢érement disposées. Le bord

supérieur du propode est défini par une ligne de neuf dents un peu plus petites.

Toute la face interne de cet article est couverte de petits tubercules coniques a

pointe cornée. La face interne du dactyle porte une rangée supérieure et une rangée

inférieure de petites spinules cornées. Les deux articles distaux sont assez fortement

déprimés latéralement; le propode n’est pas beaucoup plus de deux fois plus long

que haut et le rapport des longueurs du dactyle et du propode est de 5/4 environ.

Les p. 3 different peu des p. 2 dans la forme de leurs articles, mais le bord supérieur

du carpe ne porte qu’une épine distale; il n’y a pas de fortes dents sur le bord

supérieur du propode mais toute la face interne de cet article est couverte de

tubercules coniques moins forts que sur p. 2.

Les femelles n’ont qu’un orifice sexuel, lequel s’ouvre sur la coxa de p. 3 gauche;

un vaste repli de la peau qui prend naissance en arricre du troisieme pléopode

impair recouvre tous les pléopodes antérieurs et par conséquent la totalité de la

ponte.

Les pléopodes 1 du male ont une lame inférieure pres de quatre fois plus longue

que large, a bord latéraux sinueux et paralléles a bord antérieur faiblement denticulé.

Le lobe interne plus large que la lame principale est séparé du lobe distal trés proémi-

nent par une encoche arrondie peu profonde.

Tous les spécimens observés présentent une coloration caractéristique. Les

regions calcifiées sont d’un blanc-rosé, maculé de rouge carmin. Cette teinte est

particulierement intense sur la carapace, ou elle forme deux taches en arriére des

concavités du bord frontal, sur la base des pédoncules antennaires, dans les régions

proximales des quatre derniers articles des p. 2 et des p. 3, et dans la région antérieure

de la face interne du mérus. Les pédoncules oculaires et antennulaires sont blancs

avec des bandes longitudinales vermillons.

La région antérieure de la carapace, les appendices céphaliques antérieurs, et

les pattes thoraciques sont fortement pileux. L’extrémité antérieure des pédoncules

oculaires émerge d’un revétement assez dense de poils plumeux.

Remarques: Cette espéce se rapproche du P. mauritanicus Bouvier et du P.

difficilis Forest par la forme et les proportions de la carapace et des appendices

céphaliques antérieurs. Elle s’en distingue par la coloration, par la taille et Pacuité

des dents et épines des chélipédes et des pattes ambulatoires p. 2 et p. 3, par la

proportion des articles de ces appendices: le rapport de la hauteur a la longueur du

propode des p. 2 en particulier est 4 peine inférieur a 1/2 alors qu’1il est au plus égal

a 1/3 dans les deux autres especes; les plépodes 1 du ¢ ont une lame principale

beaucoup moins large que chez P. difficilis, et a bord antérieur denticulé, alors que

cette région est inerme chez P. mauritanicus.

P. oxyacanthus est une espéce assez grande si on la compare aux autres Paguristes

de cette région: les 2 de 5:5 et 7 mm. (I.c.), draguées en Baie de Rufisque le 18-1-52,

présentent des caractéres juvéniles: la premiére paire de pléopodes (pl. 1) est peu

développée, les pléopodes impairs sont petits et les trois premiers ont une rame

interne réduite, et il n’y a pas de repli abdominal en arriére de pl. 3. Au contraire

188 ANNALS OF THE SOUTH AFRICAN MUSEUM

chez les 2 de plus grande taille, ce repli est trés développé et recouvre non seulement

la région antérieure de l’abdomen mais les coxae des pattes ambulatoires. 2 2 de

Figs. 34-36. Deuxieme patte thoracique gauche, face interne, dénudée: 34, P. microphthalmus

Forest; 35, P. hispidus A. M.-Edw. et E. L. Bouvier; 36, P. virilis Forest.

6-5 et 7 mm. (l.c.), présentent des caractéres intermédiaires et notamment un repli

abdominal peu développé.

Les spécimens qui nous sont parvenus avec leurs coquilles étaient logés dans des

Mesalia, des Murex, et des Turritella. ‘Tous proviennent de la région de Dakar, d’une

profondeur comprise entre 8 et 40 m.

PAGURISTES D’AFRIQUE L’OUEST ET DU SUD 189

Paguristes rubrodiscus Forest

Figs. 19, 32, 46, 59

Paguristes rubrodiscus Forest, 1952, p. 261, fig. 6.

Matériel examiné: 1 9, 1.c. 7 mm., et 1 9 ovigeére, l.c. 7-5 mm.: Gorée, Aott 1946.

Type.

I g, lc. 9 mm.: Anse Bernard (Dakar), sable et cailloux, 6-10 m., 6-5-50,

Sourie coll.

1 dg, lc. 10 mm.: Baie de Hann, vers le banc de Bel Air, 12 m., cailloux et

coquilles, 17-6-50, Sourie coll.

I 9 ovigére, l.c. 7 mm.: Gorée, filets a langoustes, 23,24-8-50.

I individu adulte (non extrait de sa coquille): Gorée, dragage sur fond rocheux,

15 m., 28-12-51, Delais coll.

peeeee 45a Gimm.. 209 Veo 5-5cet 6smm~.,) 1 2 juve, lc. 4, mm: ; devant

Thiaroye, Baie de Rufisque, dragage sur fond coquillier, 10-15 m., 18-1-52, Paraiso

coll.

Description: L’écusson céphalothoracique (fig. 19) a une longueur sensiblement

égale aux 5/4 de sa largeur et aux 4/7 de la longueur totale de la carapace; elle est

faiblement spinuleuse en arriere des pédoncules antennaires. Le rostre est aigu et

atteint l’alignement des dents latérales dont il est séparé par deux profondes con-

cavités. Les pédoncules oculaires sont renflés a la base et s’élargissent au niveau

des cornées qui sont assez petites et ne sont pas échancrées postérieurement.

Les écailles oculaires sont plus longues que larges, rapprochées et armées de

quatre a cing indentations dans la région antérieure. Les pédoncules antennulaires

dépassent les pédoncules oculaires du quart ou du tiers de la longueur de leur

dernier article. Les pédoncules antennaires sont plus courts et n’atteignent pas les

cornées; leur deuxiéme article présente deux ou trois épines a chacun des angles

antérieurs; les écailles antennaires sont longues, elles arrivent au tiers distal du

dernier article pédonculaire, elles ont une extrémité bidentée et présentent trois ou

quatre dents irréguliérement disposées sur le bord interne qui est légérement con-

cave et deux ou trois dents au bord externe. Les flagelles antennaires ont une

longueur égale au 3/4 de la région antérieure de la carapace.

Les chélipédes sont égaux et de méme forme. Le carpe est allongé et sa face

supérieure est armée de dents cornées acérées dont les plus fortes sont disposées en

deux rangées longitudinales, une externe de six, l’autre interne de quatre. La main

est deux fois plus longue que large, le dactyle est nettement plus long que la région

palmaire. La face supérieure porte des tubercules coniques peu proéminents sauf

sur le bord palmaire interne qui est marqué par quatre trés fortes dents cornées

crochues dirigées vers l’avant. Il n’y a pratiquement pas de hiatus entre les doigts

lorsqu’ils sont en contact par leurs ongles cornés.

Les deux premiéres paires de pattes ambulatoires dépassent légérement les

chélipédes. Le mérus des p. 2 (fig. 32) est armé de cing 4a six petites dents sur son

bord inférieur. Le bord supérieur du carpe est défini par une rangée irréguliére de

neuf fortes dents cornées, le bord supérieur du propode par six a huit dents cornées

190 ANNALS OF THE SOUTH AFRICAN MUSEUM

un peu plus petites. Les p. 3 sont un peu plus longues que les p. 2, leurs articles sont

inermes, a l’exception du carpe dont le bord supérieur présente une ou deux dents

distales. Pour ces deux paires d’appendices le dactyle est un peu plus long que le

propode et a une section circulaire.

La femelle ne posséde qu’un orifice sexuel situé sur la coxa de la troisiéme

patte thoracique gauche. Immédiatement en arriére du troisieme pléopode impair

on observe chez les femelles adultes un petit repli du tegument qui est frangé de

longs poils et recouvre la base de cet appendice.

Les pléopodes 1 du male (fig. 46) ont une lame inferieure trés allongée, a bords

latéraux rectilignes et paralléles sur presque toute leur longueur, a bord antérieur

arrondi et inerme. Le lobe interne beaucoup plus large que la lame inférieure est

séparé du lobe distal par une encoche étroite et profonde.

Les régions calcifiées sont d’un blanc-rosé, maculé de rouge. Il y a en particulier

deux taches rouges persistantes de chaque cété de la région pré-cervicale de la

carapace, en arriere des zones épineuses. Les pédoncules antennulaires portent une

bande longitudinale rouge sur toute la longueur de leur face supérieure et de leur

face inférieure. La pigmentation la plus caractéristique est celle de la face interne

du mérus des chélipédes qui est marquée dans sa région antérieure par une tache

rouge arrondie, toujours visible apres un séjour de plusieurs années dans Valcool.

La plus grande partie de la région antérieure du corps disparait sous un revéte-

ment de longs poils plumeux, particulicrement denses sur les aires latérales de la

carapace, sur les pédoncules et les écailles oculaires, sur les écailles antennaires, et

sur les face supérieure et inférieure des pattes thoraciques.

Remarques: Le Paguristes rubrodiscus présente quelques affinités avec P. hispidus

A. Milne-Edwards et E. L. Bouvier et avec P. microphthalmus Forest qui ont comme

lui des écailles oculaires plus longues que large et contigués, des cornees petites et

peu ou pas échancrées en arriére; cependant tout un ensemble de caracteres ne

laisse guere la possibilité de le confondre avec lune ou l’autre de ces formes: région

antérieure de la carapace bien plus longue que large, rostre étroit et aigu, pédoncules

antennulaires a peine plus longs que les pédoncules oculaires, alors que dans les

deux autres espéces, ces appendices dépassent les cornées des 2/3 de leur dernier

article. La présence de la tache circulaire rouge sur la face interne du merus des

chélipédes permet d’ailleurs de Videntifier rapidement. Les trois espéces désignées

ci-dessus sont apparentées aux Paguristes de la Mer Rouge et l’aspect général du

P. perspicax Nobili (décrit comme variété de P. jousseaume: Bouvier) est fort voisin de

celui de P. rubrodiscus, les différences portant sur des caractéres nombreux mais,

dans l’ensemble, peu marqués.

La taille des P. rubrodiscus adultes examines est comprise entre 4:5 et Io mm.

(l.c.); les Q adultes ne présentent qu’un court repli de l’abdomen en arriére de

pl. 4. Quelques-uns des spécimens étaient logés dans des coquilles de Clavatula ou de

Turnitella, et un d’eux dans une coquille enti¢érement recouverte de Balanes; tous

ont été récoltés dans la région de Dakar, entre 6 et 15 m. de profondeur.

PAGURISTES D’AFRIQUE L’OUEST ET DU SUD 1QI

Paguristes microphthalmus Forest

Figs. 20, 34, 48, 60

Paguristes microphthalmus Forest, 1952, p. 260, fig. 4.

Matériel examiné: 3 3, l.c. 4:5 a 8 mm., 1 @ ovigere, l.c. 6 mm.: “‘Mercator’’,

dragage entre les iles Tamara et Roume (Iles de Los), 8-10 m., 10-12-36. Type.

Description: La région pré-cervicale représente prés des 2/3 de la longueur

totale de la carapace; le rapport de sa largeur sur sa longueur est de 5/6 environ.

Le rostre, en angle obtus, atteint presque l’alignement des deux dents latérales. Les

pédoncules oculaires sont deux fois plus larges a la base qu’au niveau des cornées

dont l’échancrure postérieure n’est qu’a peine indiquée. Leur longueur est légére-

ment inférieure aux 3/4 de celle de la région pré-cervicale. Les écailles oculaires sont

longues, assez étroites, et contigués; leur bord antérieur est tronqué et armé de

trois longues épines suivies d’une petite dent sur le bord externe. Les pédoncules

antennulaires dépassent les cornées des 2/3 de leur dernier article. Les pédoncules

antennaires sont a peu pres de méme longueur que les pédoncules oculaires; le

deuxiéme article a un angle antéro-externe peu saillant armé de deux ou trois dents.

L’écaille antennaire a une extrémité bidentée qui dépasse de peu la base du dernier

_article; son bord interne, concave, porte cinq ou six dents, le bord externe deux

ou trois dents.

Les chélipédes sont égaux et assez courts; la face interne du mérus dont le

bord inférieur est armé de six ou sept dents a une largeur maximum sensiblement

égale au 3/4 de la longueur. Le carpe est un peu plus long que le bord palmaire

interne et celui-ci est de méme longueur que le doigt fixe. La face supérieure du

carpe présente de fortes dents cornées dont la plupart sont disposées en deux rangées:

Pune de six le long du bord externe, l’autre de quatre ou cing le long du bord interne.

La main est couverte de tubercules coniques, peu saillants, sauf sur le bord palmaire

interne marqué par cing dents cornées.

Les deux paires de pattes ambulatoires sont un peu plus longues que les chéli-

pedes. Les p. 2 (fig. 34) ont un mérus dont le bord inférieur est marqué par quelques

spinules. La région supérieure du carpe porte une rangée de six ou sept fortes dents,

et une seconde rangée plus externe de six autres dents qui n’occupent que la moitié

distale; le bord supérieur du propode est défini par une rangée de douze dents

assez fortes, réguli¢rement espacées, qui se prolonge jusqu’ a la moitié distale du

dactyle sous la forme de denticules de taille décroissante. Le rapport des longueurs

du dactyle et du propode, mesurées le long du bord supérieur, est de 4/3 environ.

Les p. 3 sont inermes a l’exception d’une faible denticulation sur le bord

supérieur du mérus et d’une dent distale au bord supérieur du carpe. Le rapport des

longueurs du dactyle et du propode est voisin de 3/2.

Les femelles n’ont qu’un orifice sexuel, qui s’ouvre sur la coxa de p. 3 gauche.

Le repli de abdomen qu’on observe chez elles en arriére du troisitme pléopode

impair, est peu développé.

Les pléopodes 1 du male (fig. 48) ont une lame inférieure quatre fois et demie

a cing fois plus longue que large, a bord antérieur arrondi. Une rangée de minus-

192 ANNALS OF THE SOUTH AFRICAN MUSEUM

cules épines assez espacées s’étend du milieu du bord latéral externe au tiers interne

du bord antérieur. Le lobe interne trés développé est s¢paré par une encoche aigué

du lobe distal, lequel dépasse largement la lame inférieure.

Les exemplaires observés sont d’une teinte blanchatre a peu prés uniforme.

Une tache plus foncée sans doute rouge a Vorigine subsiste cependant sur la face

interne du mérus des chélipédes, dans sa région antérieure.

Gaillerol.m

Figs. 37-41. Deuxiéme patte thoracique gauche, face interne, dénudée: 37, Paguristes agul-

hasensis sp. nov.; 38, P. gamianus H. M.-Edw.; 39, P. barnardi sp. nov.: 40, P. engyops Barnard;

41, P. macrotrichus sp. nov.

PAGURISTES D’ AFRIQUE L’OUEST ET DU SUD 193

Les poils qui couvrent les régions latérales de la carapace et les appendices

céphaliques antérieurs ne sont pas assez denses pour cacher complétement le

tégument. Par contre les chélipédes et les deux paires de pattes ambulatoires p. 2

et p. 3 disparaissent en grande partie sous un épais revétement de poils plumeux.

Remarques: Par Vaspect général de la région antérieure du corps, par la forme

allongée et la contiguité des écailles oculaires, P. microphthalmus Forest se rapproche

de P. hispidus A. Milne-Edwards et E. L. Bouvier et de P. rubrodiscus Forest. La

longueur relative des pédoncules antennaires et oculaires permet de distinguer

facilement les trois espéces: dans la premiére, les pédoncules antennaires et ocu-

laires sont a peu prés de méme longueur, dans la seconde, les pédoncules antennaires

dépassent les yeux de la moitié de la longueur de leur dernier article et dans la

troisiéme, ils n’atteignent pas les cornées. Par l’aspect de la région antérieure du

corps P. microphthalmus est aussi apparenté a P. jousseaume: de la Mer Rouge; les

différences portent sur la forme et l’ornementation des chélipédes et des deux

premieres pattes ambulatoires.

Les quatre spécimens qui constituent le type de cette espéce ont été dragués

dans les parages des iles de Los, par 8 a 10 m. de profondeur.

Paguristes virilis Forest

Figs. 21, 36, 49, 63

Paguristes virilis Forest, 1952, p. 262, fig. 7.

Matériel examiné: 1 g et 2 9, l.c. 5-5 mm.: Le Roume (I. de Los), sable vasard,

BOC URUS

ig eca 7.5 mim. et 1 P ovigere, lc. 5-5 mm.: Mercator’, dragage entre les

tiles Tamara et Roume (I. de Los), 8-10 m., 10-12-36. ‘Type.

I g, Lc. 6 mm. et 1 9 ovigére de 7 mm.: “M’Bizi”’, station 174,13 milles W. de

Banana, vase, 30-35 Mm., 3-4-49.

12 spécimens provenant de 5 stations entre 11 et 28 m. au large de la Gold

Coast. Ces spécimens seront étudiés avec l’ensemble de la collection R. Bassindale.

Description: L’écusson céphalothoracique est a peu prés de méme longueur que

la région postérieure de la carapace, sensiblement aussi large que long et couvert de

petites saillies spinuleuses, plus nombreuses sur les aires latérales. Le rostre obtus

est nettement plus court que les dents latérales. I] n’y a pas de limite définie entre

le bord frontal et les cotés de la carapace.

Les pédoncules oculaires sont assez étroits dans leur région moyenne mais

renflés aux extrémités; ils sont un peu plus courts que la région antérieure de la

carapace. Les écailles oculaires sont aussi longues que larges, assez écartées et leur

bord antérieur est quadridenté. Les pédoncules antennulaires dépassent les yeux

de la moitié de leur dernier article. Les pédoncules antennaires plus courts n’atteig-

nent pas les cornées, angle antéro-externe saillant de leur deuxiéme article est armé

de quatre dents, leur bord interne de deux dents; |’écaille antennaire, qui atteint le

milieu du dernier article pédonculaire, a des bords latéraux rectilignes dont chacun

194 ANNALS OF THE SOUTH AFRICAN MUSEUM

est armé de cinq ou six fortes dents régulicrement espacées. La longueur du flagelle

est égale aux 3/4 de celle de la carapace environ.

Les chélipédes sont égaux. Le carpe et le dactyle sont de méme longueur et

plus long que le bord palmaire interne. Le rapport de la largeur a la longueur de la

main est de 3/5 environ. La face supérieure de ces articles est couverte de tubercules

coniques peu saillants sauf sur les bords latéraux du carpe et sur le bord palmaire

interne ou ils prennent l’aspect de fortes dents a pointe plus ou moins émoussée.

Les doigts entrent en contact sur toute leur longueur lorsqu’ils sont fermés.

Les deux paires de pattes ambulatoires p. 2 et p. 3 dépassent les chélipédes de

toute la longueur de leur dactyle. Les p. 2 (fig. 36) ont un mérus dont le bord

supérieur seul présente quelques denticulations; le bord supérieur du carpe est

armé d’un nombre trés variable de dents a pointes cornées; il y a sur l’article suivant

sept a onze dents plus faibles. Le dactyle est trés gréle, son plus grand diamétre

est compris neuf fois environ dans sa longueur; il se termine par un ongle trés petit.

Le rapport des longueurs du dactyle et du propode est de 5/3 environ.

Les p. 3 different des p. 2 en ce que leur mérus est réguli€rement denticulé le

long du bord supérieur alors que les autres articles sont inermes. Les pattes sont

plus gréles encore que les précédentes: le dactyle est dix fois plus long que large et

pres de deux fois plus long que le propode.

La femelle ne posséde qu’un orifice génital, sur la coxa de p. 3 gauche. En

arriére du troisieme pléopode impair biramé, un court repli de ’abdomen coiffe la

partie postérieure de la ponte.

Les pléopodes 1 du male (fig. 49) sont trés développés. Le lobe interne n’est

séparé du lobe distal que par une simple fente et la lame supérieure prend ainsi

Paspect d’un triangle tres allongé.

Les specimens observés ne préesentent pas de coloration caractéristique.

La pilosité est assez forte mais les longs poils plumeux qui couvrent les bords de

la carapace, les appendices céphaliques antérieurs et les pattes ambulatoires ne sont

pas assez denses pour dissimuler complétement le tegument. Par contre les chélipédes

disparaissent sous un épais manchon de poils.

Remarques: Paguristes virilis appartient au groupe de Paguristes dont les écailles

oculaires sont courtes et ont un bord antérieur denticulé. Quelques caractéres

stables et bien apparents permettent de le distinguer des espéces déja deécrites:

région antérieure de la carapace aussi large que longue, rostre trés bas, écailles

antennaires a bords rectilignes avec des dents aigués nombreuses et réguli¢rement

espacées. C’est sans doute avec Paguristes skoogi qu’il présente les plus grandes

affinités; nous n’avons malheureusement pas pu savoir ou sont conserves les

spécimens décrits sous ce nom par T. Odhner en 1923. D’aprés la description

originale et la figure qui l’accompagne, deux caractéres permettraient de distinguer

les deux espéces: chez P. skoogi les pédoncules antennaires sont aussi longs que les

pédoncules oculaires et les écailles antennaires sont armées de trois dents au bord

externe, et de quatre a cinq dents au bord interne, alors que, chez P. virilis, les

pédoncules antennaires n’atteignent pas les cornées et les bords externe et interne

des écailles antennaires sont armés de cing ou six dents. On peut encore relever des

PAGURISTES D’ AFRIQUE L’OUEST ET DU SUD 195

Figs. 42-54. Premier pléopode sexuel droit chez les Paguristes g des cdtes occidentales et

méridionales d’Afrique: 42, P. marocanus A. M.-Edw. et E. L. Bouvier; 43, P. fagei Forest;

44, P. mauritanicus Bouvier; 45, P. difficilis Forest; 46, P. rubrodiscus Forest; 47, P. oxyacanthus

Forest; 48, P. microphthalmus Forest; 49, P. virilis Forest; 50, P. hispidus A. M.-Edw. et E. L.

Bouvier; 51, P. engyops Barnard; 52, Paguristes sp. (Table Bay); 53, P. agulhasensis sp. nov.;

54, P. barnardi sp. nov.

ie 42 0) /0051147./40)< 205) 52.) X) 1551 54.) 6 22), ues autres, X95.

196 ANNALS OF THE SOUTH AFRICAN MUSEUM

différences dans la taille des pédoncules oculaires, dans l’ornementation et les pro-

portions des chélipédes et des deux premiéres paires de pattes ambulatoires.

Il faut noter que les pléopodes 1 du male de P. virilis ont une forme bien

caractéristique et sont prés de deux fois plus longs que chez les autres Paguristes de

méme taille. Chez les femelles ovigéres examinées le repli abdominal est peu

développé et ne recouvre que la base du troisieme pléopode biramé.

Les coquilles a Pintérieur desquelles étaient logés quelques-uns des spécimens

examinés appartiennent aux genres Murex et Clavatula.

P. virilis a été décrit d’aprés du matériel provenant d’une part des iles de Los,

d’autre part, d’un point beaucoup plus méridional, puisque situé au large de

lembouchure du Congo, mais ces deux localités, géographiquement fort éloignées,

présentent un caractére commun: les eaux renferment une grande quantité de vase

en suspension et les fonds sont vaseux. En ce qui concerne la profondeur, remarquons

que les P. virzlts du Congo ont été pris par 30 a 35 m., alors que ceux des I. de Los

ont été dragués sur des fonds de 8 a 10 m.

Paguristes hispidus A. Milne-Edwards et E. L. Bouvier

Figs. 22, 35, 50, 62

Paguristes hispidus, A. Milne-Edwards et E. L. Bouvier, 1892, p. 208; 1900, p. 170,

pl. XXIII, fig. 7-10. |

Matériel examiné: 1 3, 4 mm (type) et 1 2 3-5 mm., “Commandant Parfait’,

Monrovia (Liberia).

1g, 6 mm., “M’Bizi’, St. 38, 4 milles W. Moita Seca, sable et vase, 12-15 m.,

5-10-48.

Description: A. Milne-Edwards et E. L. Bouvier ont donné une description

détaillée du type, un male dont la carapace mesure 4 mm. Comme le spécimen de

M’Bizi est légerement différent, probablement en raison de sa taille plus grande, il

est utile de donner une description le l’espéce qui, portant sur plusieurs spécimens,

aura ainsi une validité plus grande. |

La région antérieure de la carapace n’est que légérement plus longue que la

région postérieure: le rapport de la largeur de l’écusson céphalothoracique a sa

longueur est de 7/8 environ; ses aires latérales sont spinuleuses. Le rostre est beau-

coup moins saillant que les dents latérales. Les pédoncules oculaires sont gréles,

fortement renflés a la base et légérement au niveau des cornées; celles-ci sont

petites et ne présentent pas d’échancrure postérieure; la longueur des pedoncules

représente les 2/3 environ de celle de Vécusson céphalothoracique. Les écailles

oculaires sont contigués et trés allongées; leur bord antérieur est armeé de quatre

longues dents aigués. Le bord antérieur des cornées atteint le quart proximal du

dernier article du pédoncule antennulaire et le milieu du dernier article du pédoncule

antennaire. Le deuxiéme article de ce dernier présente deux ou trois dents aigués sur

la saillie antéro-externe. L’écaille antennaire dont l’extrémité atteint la base du

dernier article, est armée, en plus des deux dents apicales, de trois ou quatre dents

sur le bord interne et d’une dent sur le bord externe.

PAGURISTES D’AFRIQUE L’ OUEST ET DU SUD 197

Les chélipédes sont égaux et de méme forme. La face supérieure de la main est

sub-triangulaire; elle présente sa plus grande largeur—soit les 5/8 de sa longueur—

au niveau du quart proximal et se rétrécit progressivement jusqu’a l’extrémité des

doigts. Le dactyle est nettement plus long que le bord palmaire interne. Un large

hiatus subsiste entre les doigts lorsque les ongles sont en contact. La face supérieure

du carpe, peu bombée, est limitée vers l’intérieur par une rangée de six dents cornées

et vers l’extérieur par quelques dents moins fortes. La face supérieure de la main

est couverte de tubercules coniques peu serrés dont la plupart sont disposés en lignes

longitudinales. Le bord palmaire est marqué par quatre ou cing dents un peu plus

aigués. Les pattes ambulatoires, trés gréles, dépassent largement les chélipedes. Le

bord supérieur du carpe de p. 2 (fig. 35) est armé de sept épines. Le propode et le

dactyle sont inermes; la longueur du premier ne représente que les 2/3 de celle du

second. Les p. 3 sont également inermes, a l’exception du bord supérieur du carpe

pourvu d’une épine distale; leurs articles et surtout le propode et le dactyle, sont

plus gréles encore que ceux des p. 2.

La femelle examinée (a carapace de 3°5 mm.) n’a qu’un orifice sexuel sur la

coxa de p. 3 gauche. Les trois premiers pléopodes impairs de ce spécimen présentent

une disposition particulicre. Ils sont rapprochés les uns des autres et groupés dans

le quart antérieur de Pabdomen. Un quatriéme pléopode, uriramé, est situé loin en

arricre.

Les pléopodes I du male (fig. 50) ont une lame inférieure un peu plus étroite a

Pextrémité antérieure qu’a la base, le bord antérieur de cette lame est inerme. Le

lobe interne est trés développé et séparé par une profonde encoche du lobe distal fort

saillant.

Les spécimens examinés présentent une coloration générale blanchatre. La

pilosité est assez peu développée si on la compare a celle des autres Paguristes de la

cote occidentale d’Afrique. Les poils, bruns, assez longs, mais peu serrés, retiennent

cependant la vase et cachent en partie le tégument.

Remarques: Par leurs pédoncules antennaires qui dépassent les cornées de la

moitié de leur dernier article, par la gracilité de leurs pattes ambulatoires et l’absence

de dents ou d’épines sur leurs deux derniers articles, ces Paguristes de petite taille (le

3S de 4 mm. et la 2 de 3:5 mm. sont adultes) ne risquent pas d’étre confondus avec

une autre espéce. On ne peut guére leur trouver d’affinités qu’avec P. microphthalmus

et P. rubrodiscus comme nous l’avons signalé a la suite de la description de ces deux

formes. L’espéce n’était connue que par les deux spécimens capturés par le ‘““Com-

mandant Parfait” au large du Liberia a une profondeur non précisée, décrits par

A. Milne-Edwards et E. L. Bouvier et par des spécimens de Gold Coast et du

Dahomey signalés par Balss. Le male dragué par l’Expédition ‘“‘M’Bizi” provient

d’une localité beaucoup plus méridionale, située au large du Congo, et d’une

profondeur de 12 a 15 m.

Paguristes skoogt Odhner

Paguristes skoogt, Odhner, 1923, p. 6. pl. 1.

Parmi les Pagurisies des cOtes occidentales ou méridionales d’Afrique que nous

avons examinés, aucun n’était identifiable a cette espéce, tout au moins si l’on se

198 ANNALS OF THE SOUTH AFRICAN MUSEUM

rapporte a la description qui en a été faite par Odhner. D’autre part, il ne nous a

pas été possible de découvrir ou étaient conservés les spécimens en constituant le

type. La description qui figure ci-dessous a été établie d’apres celle d’Odhner.

Description: L’écusson céphalothoracique est un peu plus long que large et

représente un peu plus de la moitié de la longueur totale de la carapace. Le rostre,

court et obtus, n’atteint pas l’alignement des dents latérales. Les pédoncules ocu-

laires sont cylindriques: leur longueur est égale aux 3/4 de celle de la région anté-

rieure de la carapace. Les écailles oculaires sont assez écartées, un peu plus longues

que larges et armées de trois dents sur le bord antéro-latéral, en arriére de l’épine

distale. Les pédoncules antennulaires dépassent les cornées de la moitié de leur

dernier article; les pédoncules antennaires sont aussi longs que les pédoncules

oculaires. Les écailles antennaires atteignent le milieu du dernier article des pédon-

cules; elles présentent quatre ou cinq dents sur le bord interne et trois dents sur le

bord externe. La longueur du flagelle est légerement supérieure a la moitié de celle

de la carapace.

Le chélipéde droit est un peu plus long que le gauche. Le carpe et le propode

sont recouverts de petites épines sans pointes cornées; le bord interne de chacun de

ces articles est armé de quatre a cinq dents plus fortes. Les doigts sont en contact

sur toute leur longueur. 3

Les pattes ambulatoires p. 2 et p. 3 dépassent les chélipédes; le rapport des

longueurs du propode et du dactyle est égal a 2/3 environ. Les pattes de la premiére

paire (p. 2) sont armées de dents sur le bord supérieur du carpe et du propode et de

petites épines localisées sur les bords supérieur et inférieur du dactyle. Les pattes

suivantes (p. 3) ne présentent qu’une épine distale sur le bord supérieur du carpe.

Paguristes skoogt, comme la plupart des espéces ouest-africaines est assez fortement

pileux. Les régions ou les poils sont les plus longs et les plus denses sont les aires

branchiales de la carapace, la face supérieure des chélipédes, les faces supérieure et

inférieure des pattes ambulatoires.

Remarques: Le dessin et la description de Paguristes skoogi donnés par Odhner,

suffsants pour distinguer cette forme des seules espéces d’Afrique occidentale con-

nues jusque la, P. hispidus, P. oculatus et P. mauritanicus, présentent quelques impréci-

sions qui rendent difficile la comparaison avec les autres Paguristes recemment deécrits.

L’espéce la plus proche est sans doute Pagauristes virilis Forest, caractérisée par un

écusson céphalothoracique trés court, des écailles antennaires a bords latéraux

réguliérement dentés, des pattes ambulatoires a dactyle long et gréle, et des pleopodes

1 trés allongés chez le male.

Les caractéres suivants, décrits ou figurés par Odhner, nous obligent a con-

sidérer pour l’instant, P. skoogi comme une espéce distincte de P. wirilis: Pécusson

céphalothoracique est un peu plus long que large, les pédoncules oculaires ont une

longueur égale aux trois quarts de celle de la région antérieure de la carapace, les

écailles antennaires sont armées de trois dents sur le bord externe et non de cing ou

six, les pédoncules antennaires sont aussi longs que les pédoncules oculaires et non

nettement plus courts.

PAGURISTES D’ AFRIQUE L’OUEST ET DU SUD 199

Ik yy y Uy, 59

Figs. 55-68. Deuxiéme pléopode sexuel droit chez les Paguristes $ des cétes occidentales et

méridionales d’Afrique: 55, P. marocanus A. M.-Edw. et E. L. Bouvier; 56, P. mauritanicus

Bouvier; 57, P. fagei Forest; 58, P. oxyacanthus Forest; 59, P. rubrodiscus Forest; 60, P. micro-

phthalmus Forest; 61, P. difficilis Forest; 62, P. hispidus A. M.-Edw. et E. L. Bouvier; 63, P.

virilis Forest; 64, P. engyops Barnard; 65, P. gamianus H. M.-Edw.; 66, P. barnardi sp. nov.

67, P. agulhasensis sp. nov.; 68, Paguristes sp. (Table Bay).

Figs. 55, 60, 62, 67, x 35; 58, 65, 68, x 22. Les autres, x 28.

200 ANNALS OF THE SOUTH AFRICAN MUSEUM

L’examen des six exemplaires d’Odhner, s’il est possible de les retrouver, ou de

spécimens provenant de la méme région que le type (Port Alexander, Angola, 72 m.

de profondeur) et la comparaison des pléopodes du male, qui, chez P. wirilis, sont si

caractéristiques, confirmera la validité de cette derniére espéce ou permettra de

Pidentifier a Paguristes skoogi.

Paguristes gamianus (H. Milne-Edwards)

Figs) if 14 125/29. 90)05 et ply ave

Pagurus gamianus, H. Milne-Edwards, 1836, p. 283; 1837, p. 235.

Paguristes gamianus, Stimpson, 1858, p. 74.

Paguristes rosaceus, Barnard, 1947, Pp. 3753 1950, p. 420, fig. 78 e-g.

et non Paguristes gamianus, Stebbing, 1910, p. 351.

29 > oo Balss}) TO12)) pl On, tiga o:

yr) 29 oy) Stebbing, 1920, p.- 257.

9 9 », Barnard, 1950, p. 418, fig. 78 a—d.

Matériel examiné: 2 3, l.c. 12 et 15 mm.; 2 9, Lc. 11 et 14.mm.: Cap de Bonne-

Esperance, Raynaud coll. (Type).

1 9, lc. 10 mm.; 2 9 ovigéres, l.c. 11 et 11-5 mm.: Buffels River, 23-2-40.

3 9, lc. 10 42 14 mm.; 1 9 ovigere, l.c. 11 mm.: Hondeklip Bay, 8-2-40. —

2 9, lic. 10 et 11 mm.: Groen River, 7-3-40

1 9, lc. 10 mm.: Zout River, 19-4-40.

1 Q juv., lc. 6 mm.: Langebaan Lagoon, 16-7-46; 1 9 ovigere, l.c. 11 mm.:

Langebaan Lagoon, 26-4-48.

1 9, lc. 7 mm.; 1 9 ovigére, l.c. 10 mm.: Langebaan Lagoon.

1 9, lc. 13°5 mm.: Table Bay, dragage par 17 m.

1g, l.c. 8 mm.: St. James (False Bay), 23-2-33.

1 9, lc. 16 mm.: False Bay, dragage par 24 m., 22-2-52.

1g, 1l.c.8mm., 1 9, l.c. 16 mm.: False Bay, dragage par 11-12 m., 5-3-52.

I g juv., l.c. 6 mm. (avec un orifice sur la coxa de la troisieme patte thoracique

droite); 4 9 juv., Le. 4°5.45:°5mm.; 2 9, l.c. 6:5 et 7 mm.: Cape Hangklip, 8-1-40.

4 2 ovigeres, l.c. 7 a 11°5 mm.: Hermanus, 30-6-39.

2 9 juv., l.c. 4 et 5 mm.; 2 9 juv., l.c. 5 mm.: Danger Point, 5-7-39.

2 Ole: 7et 7.5 mm) e325 mm.) Dancertone

2 9, l.c. 7 et 8 mm., 1 9, I.c. 7-5 mm.: Keurbooms River, Plettenberg Bay (Type

de P. rosaceus Barnard).

Tous ces échantillons—a l’exception du premier qui est le type de H. Milne-

Edwards et qui est conservé au Muséum de Paris—proviennent des collections du

Département de Zoologie de |’Université de Cape Town et du South African

Museum. Ce matériel nous a été aimablement communiqué par le Dr. K. H.

Barnard, Directeur du South African Museum.

Description: L’écusson céphalothoracique est un peu plus long que la région

postérieure de la carapace et le rapport de sa largeur sur sa longueur est égal a 4/5

environ. Le rostre aigu dépasse nettement les dents latérales. Les pédoncules

PAGURISTES D’ AFRIQUE L’OUEST ET DU SUD 201

oculaires sont renflés a la base et plus faiblement au niveau des cornées qui sont

assez petites et présentent une échancrure postérieure. Leur longueur représente les

5/9 environ de celle de la région pré-cervicale. Les écailles oculaires largement

écartées, sont lancéolées et ne présentent pas d’indentations sur le bord antéro-

latéral. Les pédoncules antennulaires sont a peu pres de méme taille que les pédon-

cules oculaires. Les pédoncules antennaires n’arrivent pas tout a fait a la base des

cornées; leur second article est armé de trois dents aigués a langle antéro-externe

qui est assez saillant. L’extrémité bidentée de l’écaille antennaire atteint ou dépasse

de peu le milieu du dernier article; son bord interne, sensiblement rectiligne, est

armé de trois dents assez peu saillantes; son bord externe présente au moins une

dent dans la région antérieure. Le flagelle est presque aussi long que la région

antérieure de la carapace.

Les chélipédes sont égaux et de méme forme. Le carpe est de méme longueur

ou légérement plus court que le dactyle et plus long que la région palmaire. La

main est un peu plus de deux fois et demie plus longue que large. Le carpe, le pro-

pode et le dactyle, sont couverts par dessus de dents épineuses a pointe cornée,

particuliérement développées sur le bord interne des articles.

Les pattes ambulatoires sont assez courtes et trapues: elles dépassent légére-

ment les chélipedes. Les p. 2 (fig. 38 et pl. ITV) ont un meérus a bord inférieur spinu-

leux; le bord supérieur du carpe porte des dents cornées qui existent aussi sur toute

la face interne du propode et sont particulicerement développées sur le bord supérieur

de cet article ou on observe une rangée de huit dents a trés longue pointe cornée,

plus fortes que sur le carpe. Cette rangée se prolonge sur le dactyle sous la forme de

dents de plus en plus petites qui se réduisent a des épines cornées dans le voisinage

de l’ongle. Le propode est a peine deux fois et demie plus long que haut. Le dactyle

a une hauteur a la base comprise cing fois environ dans sa longueur. Le rapport des

longueurs du premier et du second de ces articles est sensiblement égal a 3/4.

Les p. 3 different des pattes précédentes en ce que leurs dents épineuses, qui

occupent les mémes emplacements, sont beaucoup moins fortes.

Les femelles possédent un orifice sexuel sur chacune des coxae des p. 3; un

repli de abdomen qui prend naissance en arriére du troisieme pléopode impair

biramé recouvre les appendices ovigéres.

Les pl. 1 du male (fig. 12) ont un article distal en général peu enroulé; la

région désignée sous le nom de lame inférieure chez P. oculatus n’est pas nettement

séparée du lobe distal. Les pl. 2 (fig. 65) ont une portion distale a peine plus courte

que la portion proximale.

La région antérieure de la carapace, les pédoncules oculaires, les antennes, les

chélipédes et les pattes ambulatoires sont rouge-rosé ou carminé et souvent maculés

de pourpre; la moitié distale des dactyles des p. 2 et p. 3 est blanche. Cette colora-

tion disparait rapidement dans l’alcool et la plus grande partie des spécimens

examinés sont uniformément jaunatres.

La pilosité est assez forte sur les chélipédes et sur les deux paires.d’appendices

suivantes, mais les poils plumeux fasciculés ne forment pas de revétement continu

et ne dissimulent pas complétement l’ornementation du tégument.

202 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarques: Le type de Paguristes gamianus H. Milne-Edwards est représenté par

2 get 2 2 adultes provenant du Cap de Bonne-Espérance. Le spécimen signalé par

Balss (1912), et ceux de Stebbing redécrits par K. H. Barnard, different considérable-

ment du type et appartiennent beaucoup plus probablement a l’espéce décrite plus

loin sous le nom de P. macrotrichus. Par contre le Paguristes rosaceus que K. H. Barnard

a décrit sans avoir pu le comparer au type de l’espece de H. Milne-Edwards est bien

a mettre en synonymie avec celle-ci.

Nous décrirons plus loin sous le nom de Paguristes sp. un spécimen apparenté a

Paguristes gamianus mais en différant cependant sur quelques points assez importants:

extrémités bidentées des écailles oculaires, forme des chélipédes, proportions des

deux derniers articles des pattes ambulatoires, ornementation des pléopodes sexuels.

Nous indiquerons a la suite de la description de ce Paguristes sp. les raisons qui nous

font hésiter a le considérer comme appartenant a une nouvelle espéce.

Parmi les Paguristes provenant d’Afrique du Sud et que nous avions déterminés

comme P. gamianus figuraient un certain nombre de spécimens qu’il était impossible

de distinguer de la forme typique par l’aspect de la carapace et des appendices

céphaliques antérieurs mais qui présentaient une pilosité fort différente des chéli-

pedes et des deux paires de pattes ambulatoires. Aprés épilation, la forme et

l’ornementation des dactyles de celles-ci se sont également révélées nettement

différentes; nous avons été ainsi amené a séparer ces individus de Vespeéce de

H. Milne-Edwards et a les décrire sous le nom de Paguristes barnard: sp. nov.

Les échantillons de P. gamianus proviennent surtout de la zone intercotidale des

cétes ouest et sud-ouest d’Afrique du Sud, depuis Buffels River, jusqu’a la région

de Cape Town. L’un d’entre eux provient cependant d’un point situé beaucoup

plus a Vest, de Plettenberg Bay. Quelques spécimens parmi les plus grands ont été

dragués entre 11 et 24 m. de profondeur.

Paguristes sp.

Figs. 24, 52, 68

Matériel examiné: 1 3, l.c. 20 mm., Table Bay (céte occidentale d’Afrique du

Sud, dragage par 17 m.

Description: L’écusson céphalothoracique est un peu plus long que la région

postérieure de la carapace; le rapport de sa largeur et de sa longueur est égal a 6/7.

Le rostre ne dépasse pas l’alignement des dents latérales. Les pédoncules oculaires,

renflés 4 la base ont une longueur égale aux 2/3 environ de celle de la région pré-

cervicale. Les cornées sont petites et légérement échancrées postérieurement. Les

écailles oculaires sont triangulaires et ont un sommet bidenté. Les pédoncules

antennulaires sont un peu moins longs que les pédoncules oculaires. Les pédoncules

antennaires sont plus courts encore; le second article a un angle antéro-externe

fortement saillant et armé de trois dents aigués. L’écaille antennaire atteint le tiers

distal du dernier article pédonculaire, son bord interne est armé de quatre fortes

dents, son bord externe de trois dents. Le flagelle est aussi long que la région

antérieure de la carapace.

PAGURISTES D’AFRIQUE L’OUEST ET DU SUD 203

Les chélipédes sont égaux et de méme forme. Le carpe est un peu plus court

que le dactyle dont la longueur représente plus des 2/3 de celle de la main; celle-ci

est sub-triangulaire et plus de deux fois plus longue que large. La face supérieure

des trois derniers articles est couverte de fortes dents a pointe cornée, moins nom-

breuses sur les doigts ou on observe deux bandes longitudinales inermes. Les pattes

ambulatoires sont un peu plus longues que les chélipedes. Les trois derniers articles

des p. 2 sont armés par dessus de fortes dents cornées acérées. La face interne des

deux derniers est également épineuse, on observe en particulier une rangée longitu-

dinale sub-médiane de sept dents sur la face interne du propode. Cet article est a

peu prés trois fois plus long que haut et d’un tiers plus court que le dactyle. Celui-ci

a une hauteur maximum comprise six fois dans sa longueur. Les pattes de la paire

suivante présentent des proportions voisines et des dents cornées implantées dans les

mémes régions mais moins fortes.

Les pl. 1 du male (fig. 52) ont une lame inférieure a bord antérieur arrondi

armé de six ou sept crochets, séparée du processus distal trés long par une incisure

bien marquee.

L’unique exemplaire ne présente pas de coloration caractéristique.

Les trois premieres paires de pattes thoraciques présentent une pilosité assez

forte constituée par des touffes de longs poils, plumeux a l’extrémité, particuliére-

ment nombreux sur les régions de la face supérieure des chélipedes proches du bord

externe, sur la région supérieure du carpe et sur les régions supérieure et inférieure

du propode et du dactyle des deux paires ambulatoires. I] n’y a pas de revétement

continu de poils plumeux trés serrés sur la face interne du propode de ces appen-

dices et les dents épineuses situées sur cette face sont toujours bien dégagées.

Remarques: Le spécimen deécrit ci-dessus présente des affinités avec Paguristes

gamianus, tout en en différant sur un certain nombre de points. Comme il s’agit d’un

individu de taille nettement supérieure a celle des plus grands P. gamianus observés, il

est possible que ces différences soient liées a Age et nous avons préféré ne pas le

décrire comme espéce nouvelle. Les ressemblances avec Vespece de H. Milne-

Edwards portent sur Vaspect général, sur les proportions de la carapace et sur

lornementation des trois premiéres paires de péréiopodes. Paguristes sp. se distin-

gue de P. gamianus par la briéveté du rostre, par les écailles oculaires dont l’extré-

mité antérieure est bidentée, par la forme des chélipedes dont la main est sub-

triangulaire et dont le dactyle est trés long, par les proportions des deux derniers

articles des pattes ambulatoires p. 2 et p. 3: le propode de ces appendices a une

hauteur comprise trois fois environ dans sa longueur: le dactyle une hauteur

maximum comprise six fois environ dans sa longueur, alors que ces rapports valent

respectivement 2°5 et 5 chez P. gamianus.

Un dernier caractére morphologique rend probable l’appartenance de Paguristes

sp. a une espéce distincte: les pl. 1 g présentent, sur le bord antérieur de la lame

inférieure des crochets bien développées qui n’existent pas chez P. gamianus. Quant

aux pl. 2, la forme de leur région distale est particuliére (fig. 68).

I] faut noter que ce Paguristes a été capturé en méme temps qu’une femelle de

P. gamianus typique dans la région de Table Bay.

204 ANNALS OF THE SOUTH AFRICAN MUSEUM

Paguristes engyops Barnard

Figs. 25, 40, 51, 64

Paguristes engyops, Barnard, 1947, p. 375; 1950, p. 421, fig. 78 h-z.

Materiel examiné: 3 g, l.c.2 a3 mm., 6 Q ovigeres, l.c. 2:25 a 3 mm., Buffels Bay,

False Bay, littoral, dans des coquilles de Trochidae (Type).

I 4, l.c. 7 mm., Langebaan Lagoon, 26-4-48.

4 9, le. 3 a 7-5 mm., 1 @ ovigére, l.c. 5-5 mm., Langebaan Lagoon, récolte

intercotidale.

Description: La longueur de l’écusson céphalothoracique est égale aux 7/6

environ de sa largeur et aux 3/5 de la longueur totale de la carapace. Le rostre aigu

atteint ’alignement des deux pointes latérales dont il est séparé par deux fortes

concavités. Les pédoncules oculaires larges a la base s’amincissent progressivement

jusqu’a la région cornéenne qui est légérement renflée; leur longueur représente les

4/7 environ de celle de la région pré-cervicale; les cornées ne sont pas échancrées

postérieurement. Les écailles oculaires sont rapprochées, assez grandes, lancéolees,

uni- ou, rarement, bidentées a l’extrémité avec parfois une épine vers le tiers distal du

bord interne qui est fortement convexe. Les pédoncules antennulaires dépassent les

yeux de la moitié environ de la longueur de leur dernier article. Les pédoncules

antennaires atteignent la base des cornées, leur second article a un angle antéro-

externe aigu mais peu saillant. L’écaille antennaire courte et large atteint la base du

dernier article pédonculaire, son bord interne est rectiligne et inerme ou faiblement

denticulé, son bord externe convexe et armé de deux ou trois fortes dents. Le

flagelle antennaire est un peu plus court que les pédoncules oculaires.

Les chélipédes sont sub-égaux chez les femelles et chez les jeunes males. Chez

les males adultes le droit est un peu plus fort que le gauche. Le carpe est un peu

plus long que le dactyle. La longueur de ce dernier représente les 2/3 environ de la

longueur totale de la main. Il y a quelques tubercules peu saillants sur la face

supérieure du carpe, plus forts sur le bord interne. La région palmaire est aussi

pourvue de quelques tubercules, plus développés également sur le bord interne. Les

doigts sont couverts de petits tubercules arrondis. Une forte dent au milieu de la

face interne du doigt fixe se loge dans une concavité bien marquée de la face

correspondante du doigt mobile.

Les pattes ambulatoires sont trapues; elles sont nettement plus courtes que les

chélipédes et comme eux sont faiblement armées: des dents peu nombreuses et

émoussées existent sur le bord supérieur du carpe et du propode des p. 2 (fig. 40),

alors que le dactyle n’est armé par dessous que de quelques spinules cornées placées

en arriére de l’ongle. Les p. 3 sont inermes a l’exception d’une dent distale sur le bord

supérieur du carpe et de quelques spinules sur les faces interne et inférieure du

dactyle. Dactyle et propode des deux paires sont sub-égaux.

Les femelles possédent une paire d’orifices génitaux, et un trés court repli

abdominal en arriére du troisiéme pléopode impair. Les pl. 1 du male (fig. 51) ont

une lame inférieure spatulée dans la région distale et armée sur le bord antérieur de

PAGURISTES D’AFRIQUE L’OUEST ET DU SUD 205

huit a dix crochets. Le lobe interne est presque aussi proéminent que le processus

distal dont il est séparé par une profonde encoche.

La pilosité est constituée par des touffes de poils plumeux trés denses sur les

écailles oculaires, plus ou moins abondants sur les trois premiéres paires d’appendices

thoraciques.

Remarques: Si la description originale du Paguristes engyops différe quelque peu

de celle que nous donnons ci-dessus, c’est que le type est constitué par des individus

de trés petite taille. Les individus plus agés ont en particulier une région antérieure de

la carapace moins allongée, des pédoncules oculaires plus gréles, des cornées plus

renflées que les jeunes.

I] est intéressant de noter que l’espéce la plus voisine de P. engyops est P. maro-

canus des cétes marocaines. Tous deux sont de petite taille, ont une carapace et un

bord frontal d’aspect voisin, des cornées non échancrées postérieurement et dépig-

mentées dans l|’alcool, des écailles oculaires 4 peu prés de méme forme, des flagelles

antennaires courts et épais, des pléopodes males de méme type. Elles sont cependant

bien distinctes: la réduction des cornées chez P. engyops, la forme des chélipédes, les

proportions des pattes ambulatoires sont autant de caractéres qui interdisent toute

confusion.

I] existe aussi entre les deux espéces des différences écologiques considérables.

Les spécimens connus de P. marocanus, malheureusement fort peu nombreux, ont été

dragués sur des fonds de 115 a 290 m., alors que l’un des spécimens de P. engyops au

moins provient de la zone intercotidale, les autres étant seulement qualifiés de

“‘littoraux”’ sur les étiquettes.

Les plus petits males observés (2 mm. de carapace) ont déja des pléopodes du

type adulte, et les plus petites femelles (2:25 mm.) sont déja ovigéres, portant 3 ou

4 oeufs dont le plus grand diamétre est compris entre o-8 et 1-2 mm.

Paguristes agulhasensis sp. nov.

Figs. 26, 37, 53, 67

Matériel examiné: 1 3, l.c. 5-5 mm., Agulhas Bank, dans une coquille de Fusus

rubrolineatus, 55 metres. (Type.)

Description: La longueur de Vlécusson céphalothoracique est légérement

supérieure a la largeur et sensiblement égale aux deux tiers de la longueur de la

carapace. Le rostre, large et aigu, dépasse de peu |’alignement des pointes latérales.

Les pédoncules oculaires, épais et sub-cylindriques, sont égaux aux deux tiers de la

région antérieure de la carapace. Les écailles oculaires qui sont trés écartées ont

une région antérieure triangulaire se terminant par une pointe trés aigués. Les

pédoncules antennulaires sont de méme longueur que les pédoncules oculaires. Les

pédoncules antennaires n’arrivent pas tout a fait au niveau des cornées, langle

antéro-externe du second article est fort saillant et armé de trois dents aigués.

L’écaille antennaire dont la pointe dépasse le milieu du dernier article pédonculaire

a un bord interne concave armé d’une dent proximale, et trois dents aigués sur le

_ bord externe. Le flagelle est un peu plus court que l’écusson céphalothoracique.

206 ANNALS OF THE SOUTH AFRICAN MUSEUM

Les chélipédes sont sub-égaux. La longueur du dactyle est sensiblement la

méme que celle du carpe et représente les 2/3 environ de la longueur totale de la

main. La face supérieure du carpe et du propode est ornée de quelques tubercules

a sommet corné mais peu aigu sauf sur le bord palmaire interne, lequel est marqué

par quatre fortes dents. Sur la moitié proximale du dactyle dans le prolongement

du bord palmaire existent des dents cornées aigués.

Les p. 2 (fig. 37) portent sept ou huit petites dents cornées irréguliérement

implantées dans la région supérieure du carpe et une rangée de sept dents un peu plus

fortes sur le bord supérieur du propode; on observe sur le bord supérieur du dactyle

des denticules de taille décroissante, réduits a leur pointe cornée dans le voisinage de

Vongle, et, sur le bord inférieur, une douzaine de spinules cornées. Le rapport des

longueurs du propode et du dactyle est sensiblement égal a 3/4, et la hauteur maxi-

mum du dernier article est comprise cinq fois et demie dans sa longueur. Les p. 3

sont inermes a l’exception du bord supérieur du carpe pourvu d’une dent distale et

des régions supérieure et inférieure du dactyle armées de spinules cornées.

Les régions les plus pileuses sont la région rostrale, les écailles antennaires, les

aires branchiales de la carapace et les appendices thoraciques. Les poils présents

sur le propode et sur le dactyle des trois premieres paires de péréiopodes sont

dépourvus de barbules, alors que sur les autres régions on observe une forte propor-

tion de soies plumeuses.

Remarques: La description ci-dessus est celle d’un unique spécimen male, de

petite taille mais possédant des pléopodes sexuels bien développés. I présente

quelque ressemblance dans l’aspect général avec Pagurisies gamianus et avec le

spécimen décrit plus loin sous le nom de P. macrotrichus sp. nov.

Paguristes agulhasensis se distingue des Paguristes gamianus de méme taille par la

forme du rostre plus large, moins aigu et moins saillant, par les pédoncules oculaires

plus épais, par les écailles oculaires plus écartées, par les écailles antennaires armées

d’une seule dent proximale sur le bord interne et non de trois ou quatre, et enfin,

caractére qui est peut-étre le plus net, par le présence de dents moins nombreuses et

beaucoup moins fortes sur le propode et le dactyle des pattes ambulatoires; ces deux

articles présentent d’ailleurs une pilosité moins dense et constituée par des poils

simples alors qu’on observe une forte proportion de poils plumeux chez P. gamianus.

I] faut noter que les premiers pléopodes sexuels du spécimen étudié (fig. 53)

semblent avoir atteint leur taille et leur forme définitive ce qui permet de croire que

nous avons affaire a un adulte, alors que chez les P. gamianus de méme taille, soit

5°5 mm. de carapace, ils sont plus courts et n’ont pas acquis leur plein développe-

ment. La structure de ces appendices ne permet cependant pas de distinguer les

deux espéces.

La ressemblance de P. agulhasensis et du spécimen femelle décrit plus loin sous le

nom de Paguristes macrotrichus nov. sp. est sans doute plus grande notamment en ce

qui concerne les denticules implantés sur les deux derniers articles des pattes

ambulatoires p. 2 et p. 3. P. macrotrichus présente cependant plusieurs caractéres qui

ne permettent guére de l’identifier 4 P. agulhasensis pour l’instant et en particulier

un rostre plus court et arrondi, des écailles oculaires plus petites et plus écartées

PAGURISTES D’AFRIQUE L’OUEST ET DU SUD 207

Paguristes oculatus type

P.oculatus rubro-=pictus

P,marocanus

“Pp. fagei Vid

P,mauritanicua

Pecitiiciiitis

P,oxyacanthus

P.,rubrodiscus

P.microphthalmus

P.virilis

P,hispidus

P,skoogi

P.gamianus

P,engyops

P,agulhasensis

P,barnardi no fe O. .

P,macrotrichus O =) &

ss)

eal

r |

Fig. 69. Le genre Haguristes sur les cOtes occidentales et méridionales d’Afrique. Les espéces

chez lesquelles les femelles ne possédent qu’un orifice génital sont représentées par des signes

carrés.

208 ANNALS OF THE SOUTH AFRICAN MUSEUM

encore, des écailles antennaires armées de plusieurs dents sur leur bord interne, une

zone allongée dépourvue de tubercules ou de dents sur les doigts des chélipédes et

enfin une pilosité plus forte, qui sur le propode et le dactyle des pattes ambulatoires

est constituée par de longs poils plumeux.

Paguristes barnardi sp. nov.

Figs. 39, 54, 66, et pl. iv.

Paguristes rosaceus (pars), Barnard, 1947, p. 375.

Matériel examiné. 1 3, l.c. 6 mm., 1 ¢ juv., le. 4 mm., 1 9 juv., Le. 4°5 mm.:

Kleinmond, 19-3-309.

I g, l.c. 10 mm.: Kleinmond, 20-3-39.

5 d, lc. 8a 13 mm., 1 Q, lc. 5-5 mm.: Cap Agulhas, 29-9-39. (Syntypes.)

4 4, le. 5:5 a 7°5 mm., 1 Q ovigere, l.c. 6 mm.: Arniston, 23-11-39.

5 6, lc.6a10mm.: Still Bay, 10-11-39. (Syntypes.)

I 4, l.c. 6-5 mm., 3 d juv., l.c. 3a 4 mm., 1 9 juv., l.c. 4 mm.: Knysna, 6-4-30.

2 4, l.c. 7 et 10 mm., 1 ¢ juv., l.c. 5 mm., 1 9 ovigere, |.c. 5 mmesoy Sune

2:5 et 3-5 mm.: Storms River, 2-4-39. (Syntypes.)

I Q, lc. 5 mm., 1 Q, l.c. 6-5 mm.: Jeffries Bay, 1-6-30.

1 Q, l.c. 7 mm.: Port-Elisabeth.

Tous les échantillons appartiennent a |’Université de Cape Town ou au South

African Museum; ils ont été récoltés dans la zone intercotidale.

Description: En ce qui concerne la carapace et les appendices céphaliques

antérieurs la description de P. gamianus peut s’appliquer a P. barnardi. Les différences

entre les deux espéces portent surtout sur les trois premiéres paires de péréiopodes

et sur la pilosité. Dans l’espéce nouvelle on observe les caractéres suivants:

Le carpe des chélipédes est un peu plus long que le dactyle et sa face supérieure est

armée de dents cornées plus nombreuses que chez P. gamianus. Le carpe des p. 2

(fig. 39) est aussi plus épineux. Le dactyle des p. 2 et des p. 3, un peu plus long que

le propode, est peu déprimé latéralement; son bord supérieur ne porte pas de fortes

dents a base calcifiée, mais de nombreuses spinules cornées de tres petite taille.

Comme chez P. gamianus les pl. 1 ¢ (fig. 54) sont peu enroulés, l’encoche qui

sépare le lobe interne du lobe distal est en général plus profonde que dans l’espéce

de H. Milne-Edwards, mais ce caractére n’est pas assez net pour qu’on puisse lui

accorder une valeur discriminatoire. Par contre la portion distale du dernier article

des pl. 2 ¢ (fig. 66) est bien plus courte que la portion proximale, alors que les deux

régions sont sub-égales chez P. gamianus.

Les régions calcifiées des spécimens les moins décolorés sont teintées de rose.

Le carpe et le propode des chélipédes ont une face supérieure bordée latérale-

ment d’une frange trés dense de longs poils plumeux et les dents cornées présentes

sur cette face sont en grande partie dissimulées par un revétement trés épais de poils

plumeux plus courts. La face supérieure du carpe, les faces supérieure et inférieure

du propode et du dactyle des deux paires d’appendices suivantes sont également

garnies d’une épaisse frange de longs poils plumeux qui cachent les dents et spinules

PAGURISTES D’AFRIQUE L’OUEST ET DU SUD 209

présentes sur ces régions (pl. iv). Comme sur les chélipédes, il y a sur la face interne

du propode un feutrage continu de poils courts.

Remarques: L’examen des Paguristes figurant dans les collections du South

African Museum et de |’Université de Cape ‘Town sous le nom de P. gamianus nous

a amené tout d’abord a en séparer un spécimen qui sera décrit sous le nom de P.

macrotichus sp. nov. Les autres peuvent étre rassemblés en deux groupes: le premier

comprend les individus identifiables au véritable P. gamianus de H. Milne-Edwards

et en particulier le type de P. rosaceus Barnard. Le second est constitué par d’assez

nombreux spécimens fort proches de P. gamianus par l’aspect de la carapace et des

appendices céphaliques mais présentant des différences marquées dans l’ornemen-

tation des dactyles des pattes ambulatoires et surtout dans la pilositeé.

La constance de ces différences, absence de formes intermédiaires et certaines

considérations biologiques nous ont décidé a faire du second groupe une espéce

nouvelle que nous sommes heureux de dédier au Dr. K. H. Barnard. Les caractéres

permettant de distinguer P. barnardi de P. gamianus ont été indiqués ci-dessus.

L’aspect du dactyle de la premiére ou de la seconde patte ambulatoire montre

immédiatement a quelle espéce on a affaire: chez P. gamianus cet article est déprimé

latéralement et armé par dessus de fortes dents a base calcifiée, bien visibles en

raison du peu de développement de la pilosité (pl. 1V en bas); chez P. barnard:, sa

section est sub-circulaire et il présente par dessus et par dessous une frange trés

fournie qui couvre complétement les régions supérieure et inférieure et cache la

totalité des spinules cornées (pl. IV en haut).

I] faut noter que P. gamianus est de plus grande taille que P. barnardi. Le tableau

ci-dessous donne la taille moyenne (I.c.) en millimétres pour les différentes catégories

dindividus: 3 juvéniles, ¢ adultes, 2 juvéniles, 2 adultes (ovigéres incluses), et 9

ovigeres.

gd juv. adultes Q juv. Qadultes Qovig. g¢ + Q adultes

P. gamianus 3) 10-7 5:2 9:7 9-9 9-8

P. barnardi 3:8 6-4 39 Oyen a) 5:8 7-8

Le nombre d’individus examinés, quarante-quatre pour la premiére espéce,

trente-quatre pour la seconde, est suffisant pour que les chiffres obtenus aient une

signification; il faut retenir en particulier la différence de taille 4 la maturité

sexuelle: la plus petite 9 ovigére mesure 7 mm. chez P. gamianus et 5 mm. seulement

chez P. barnardt. :

Dans l’ensemble la répartition géographique des deux espéces présente aussi des

différences: les échantillons de P. gamianus proviennent pour la plupart des cétes

ouest de l’Afrique du Sud alors que les P. barnardi ont été capturés sur la céte sud.

Sur les 13 localités ou ont été récoltés les P. gamianus, 12 se trouvent a l’ouest et au

nord de Danger Point, cap situé dans la région occidentale de la céte sud. Sur les

8 localités de P. barnardi, 7 sont a est de ce cap. Sur la figure 70 les courants marins

ont été schématiquement représentés, ainsi que les points ou ont été récoltés P.

gamianus et P. barnardi. La premiere espéce a surtout été récoltée sur les cdtes baignées

par les courants froids (courant du Benguela) et la seconde, sur la céte sud, la ou le

210 ANNALS OF THE SOUTH AFRICAN MUSEUM

courant chaud des Aiguilles exerce son influence. La région ou les deux espéces se

rencontrent l’une et l’autre est celle du Gap et de Danger Point qui voit aussi la

confluence des eaux chaudes et froides. En ce qui concerne le niveau de récolte,

notons que tous les P. barnard: se trouvaient dans la zone intercotidale comme beau-

coup de P. gamianus mais que quelques-uns de ceux-ci ont été dragués entre 11 et

24 m.

CAPE TOWN

™~ NG

Fig. 70. Détail de la distribution des Paguristes sur les cotes d’Afrique du Sud. Pour l’expli-

cation des signes voir fig. 69. Courants chauds — — -— et courants froids ———.

PAGURISTES D’AFRIQUE L’OUEST ET DU SUD QI

Paguristes macrotrichus sp. nov.

Figs. 27, 41

Paguristes gamianus, Stebbing, 1910, p. 351.

.F ah Balss, 1912, p. 91, fig. 2.

s i Stebbines 19205 py 2577.

y se Barnard, 1950, p. 418, fig. 78, a-d.

Matériel examiné: 1 9, l.c. 7 mm., région de Natal, 90 metres, identifiée par

Stebbing comme P. gamianus H. Milne-Edwards. (Type.)

Description: La longueur de lécusson céphalothoracique est légerement supér-

ieure a sa largeur et représente les deux tiers environ de la longueur totale de la

carapace. Le rostre arrondi atteint ’alignement des pointes latérales. Les pédon-

cules oculaires sont sub-cylindriques, assez épais, faiblement dilatés au niveau des

cornées qui présentent une échancrure postérieure. Leur longueur est égale aux

3 de celle de la région pré-cervicale. Les écailles oculaires sont petites, triangu-

laires dans la région distale, et trés écartées. Les pédoncules antennulaires sont un

peu plus longs que les pédoncules oculaires, ceux-ci dépassant les pédoncules

antennaires du tiers de leur longueur. Les écailles antennaires sont longues; leurs

bords latéraux sont armés de trois a cing dents irrégulicrement développées. Les

flagelles antennaires ont la méme longueur que l’écusson céphalothoracique.

Les chélipédes sont égaux. Le carpe est plus court que le dactyle et celui-ci a

une longueur égale aux deux tiers de la longueur totale de la main. La face supér-

ieure de ces trois articles porte des tubercules sauf dans les régions voisines des bords

de contact des doigts. Il y a cing dents corneées sur le bord interne du carpe, quatre

sur le bord palmaire interne, et d’autres plus nombreuses mais plus petites et plus

aigués sur les régions externes des doigts. Les p. 2 (fig. 41) sont armées de quelques

dents cornées sur la face supérieure du carpe, de sept dents plus réguliéres sur le

bord supérieur du propode et d’une rangée de denticules de taille décroissante sur

le bord supérieur du dactyle: le bord inférieur de cet article est également armée

d’épines cornées assez longues. Le rapport des longueurs du propode et du dactyle

est sensiblement égal a trois quarts.

Les p. 3 sont inermes a l’exception du bord supérieur du carpe armé d’une

épine distale et de la face interne du dactyle qui présente deux rangées longitudinales

de spinules cornées, lune prés du bord supérieur l’autre prés du bord inférieur.

Un repli abdominal recouvre les trois premiers pléopodes du spécimen étudié

qui est une femelle non ovigére.

La pilosité est surtout constituée par de longs poils fins et plumeux assez denses

mais ne dissimulant pas l’ornementation du tégument. Les régions les plus pileuses

sont: le bord frontal, les écailles oculaires et antennaires, les trois premiéres paires

de péréiopodes et particulicerement les régions supérieures de leurs trois derniers

articles ou l’on observe des faisceaux de longs poils insérés a la base des dents et des

tubercules ou dans des dépressions du tégument.

Remarques: Le spécimen étudié ci-dessus a été déterminé par Stebbing, décrit

et figuré par Barnard sous le nom de Paguristes gamianus. 11 appartient probablement

DP. ANNALS OF THE SOUTH AFRICAN MUSEUM

a la méme espéce que les spécimens décrits par Balss sous le méme nom. En fait, il

s'agit d’une forme bien différente de l’espéce de Milne-Edwards, comme en témoigne

la comparaison des figures 23 et 27 pour la région antérieure de la carapace et les

appendices céphaliques antérieurs, et 38 et 41 pour les pattes ambulatoires. L’éco-

logie des deux espéces est aussi bien différentes: P. macrotrichus a été capturé par go m.

(Stebbing) et 150 m. (Balss) de profondeur, alors que les véritables P. gamianus

proviennent de la zone intercotidale ou d’une profondeur n’excédant pas 24 m.

P. macrotrichus parait plus proche du spécimen deécrit plus haut sous le nom de

Paguristes sp. qui est soit un P. gamianus trés agé, soit une espéce nouvelle. A ne

considérer que l’aspect de la région antérieure du corps, 11 parait hasardeux d’identi-

fier les deux spécimens a une méme espéce. Les pédoncules oculaires sont bien plus

eréles chez Paguristes sp. qui, par ailleurs, a des écailles oculaires bidentées, mais il

faut faire preuve de beaucoup de prudence dans l’utilisation de caractéres présentés

par des individus de taille trés dissemblable et, qui plus est, de sexes différents. La

comparaison des autres régions du corps ne peut trancher la question. En effet si

les chélipédes sont d’une forme voisine, les pattes ambulatoires paraissent moins

fortement armées chez P. macrotrichus: chez celui-ci on observe seulement sur la face

interne du propode de peu nombreuses et minuscules spinules alors qu’il existe en

cet endroit de fortes dents cornées chez Paguristes sp. La différence dans la denticu-

lation est ici bien plus accentuée qu’entre les plus petits et les plus grands spécimens

de P. gamianus.

La comparaison entre P. macrotrichus et P. agulhasensis sp. nov. figure a la suite

de la description de ce dernier.

Parmi tous les Paguristes d’ Afrique du Sud que nous avons eu l’occasion d’étudier,

Punique spécimen femelle de P. macrotrichus est celui qui provient de la localité la

plus orientale (Natal) et de la plus grande profondeur.

BIBLIOGRAPHIE

Alcock, A., 1905. Catalogue of the Indian Decapod Crustacea in the collection of the Indian Museum.

Pt. 2. Anomura, fasc. 1. Pagurides, i-xi, 1-197, 16 pl.

Balss, H., 1912. ‘Paguriden.’ Wiss. Ergebnisse der deutsche Tiefsee-Exp. “Valdivia”, Iena, 20, no. 2,

87-124, pl. 7-11.

, 1921. ‘Crustacea VI: Decapoda Anomura und Brachyura.’ Beitrége zur Kenntnis der Meeres-

fauna Westafrikas, Hamburg, 3, no. 2, 13-67.

, 1940-44. “Decapoda: Morphologischer Teil.” (H. G. Bronns: Klassen und Ordnungen des

Tierreichs, Leipzig, 5 [1], 7. Buch, 22-660).

Barnard, K. H., 1947. ‘Description of new species of South African Decapod Crustacea with notes on

synonymy and new records.’ Ann. Mag. Nat. Hist. (11) 13, 361-92.

» 1950. ‘Descriptive Catalogue of South African Decapod Crustacea (Crabs and Shrimps).’

Ann. S. Afr. Mus., 38, 1-837, 154 figs.

Boas, J. E. V., 1880. ‘Studier over Decapodernes Slaegtskabsforhold.’ K.D. Vid. Selsk. Skr. 6 R. nat. og.

math. Afd., Kobenhavn, J, 2, 26-210, 7 pl.

, 1926. ‘Zur Kenntnis Symmetrischer Paguriden.’ Biol. Meddel. K. Danske Vid. Selsk, 5, 6, 1-52,

25 fig.

PAGURISTES D’ AFRIQUE L’ OUEST ET DU SUD 213

Bouvier, E. L., 1892 ‘Etude sur les Paguriens recueillis par le Dr. Jousseaume sur les cétes de la Mer

Rouge.’ Bull. Soc. Philom. Paris, 1891-92, sér. 8, 4, 50-5.

, 1906 a. ‘Sur les Crustacés Décapodes marins recueillis par M. Gruvel en Mauritanie.’ Bull.

Mus. Nat. Hist. Nat., Paris, 12, no. 4, 185-7, fig. 1.

, 1906 b. Mission des Pécheries de la céte occidentale d’ Afrique, no. 7, 95-97-

, 1906 c. Act. Soc. Linn., Bordeaux, J (61), 198-200.

Brocchi, M., 1875. ‘Recherches sur les organes génitaux males des Crustacés Décapodes.’ Ann. Sc.

Nat. Kool. (6), 2, 1-131, pl. 4-10.

Cavolini, 1787. Memoria sulla generazione dei Pesci e dei Granchi, Naples.

Chevreux, M. et Bouvier, E. L., 1892. “Voyage de la Goélette “‘Melita’’ aux Canaries et au Sénégal,

1889-90, Paguriens.’ Mem. Soc. Zool. France, 5, 83-144, pl. I-IV.

Fabricius, J., 1775. Systema Entomologiae.

Forest, J.. 1952 a. ‘Remarques sur les genres Diogenes Dana et Tr Aalnennts Henderson... .’ Bull. Inst.

Roy. Sci. Nat. Belgique, 28, no. 11, 1-15, 15 fig.

, 1952 6. ‘Notes préliminaires sur les Paguridae des cétes occidentales d’Afrique. II. Diagnose

sommiaire de six espéces nouvelles appartenant au genre Paguristes Dana.’ Bull. Mus. Nat. Hist.

Nat., Paris, 2e sér., 24, no. 3, 257-62, figs.

» 1952 c. ‘Contributions a la revision des Crustacés Paguridae. 1. Le genre Trizopagurus.’

Meém. Mus. Nat. Hist. Nat., Paris. sér. A, Zool. 5, no. 1, 1-40, 25 figs.

Krauss, F., 1843. Die Siidafrikanischen Crustaceen, Stuttgart, 1-68, pl. 1-4.

Miers, E. J., 1881. ‘On a collection of Crustacea made by Baron Hermann Maltzam (sic = Maltzan)

at Gorée Island, Senegambia.’ Ann. Mag. Nat. Hist., London, (5) 8, no. 45-47, 204-20, 259-81,

364-77, pl. 13-16.

Milne-Edwards, A., et Bouvier, E. L., 1891. ‘Sur les modifications que présentent les Pagures suivant

le sens de l’enroulement de la coquille qu’ils habitent.’ Bull. Soc. Philomath. Paris, sér. 8, 3, 151-3.

———, 1892. “Observations préliminaires sur les Paguriens recueillis par les Expéditions du ‘““Travail-

leur” et du “Talisman”’.’ Ann. Sci. Nat. Zool., sér. 7, 13, 185-226.

, 1900. “Expéditions du ““Travailleur” et du ‘““Talisman’’.’ Crustacés Décapodes 1. Paris, 1-396

B2nple

Milne-Edwards, H., 1836. “Observations zoologiques....’ Ann. Sci. Nat. Zool., Paris, sér. 2, 6, 257-88,

2 pl.

Monod, Th., 1933. “Sur quelques Crustacés de l’Afrique occidentale.’ Bull. Com. Et. Hist. Scient.

A.O.F., 15, no. 2-3, 456-548, figs.

Nobili, G., 1906 a. ‘Faune carcinologique de la Mer Rouge. Décapodes et Stomatopodes.’ Ann. Sc.

Nat. Zool., Paris, 1, 1-347, pl. I-XI.

, 1906 6. ‘Mission J. Bonnier et Ch. Pérez (Golfe Persique 1901). Crustacés Décapodes et

Stomatopodes.’ Bull. Sci. France-Belgique, Paris, 40, 13-159, pl. II-VII.

Odhner, Th., 1923. ‘Marine Crustacea Podophthalmata aus Angola und Siid-Afrika gesammelt von

H. Skoog 1912.’ Géteborg K. Vet. Samh. Handl., 27, no. 5, 1-39, figs.

Rathbun, M. J., 1900. “The Decapod Crustaceans of West Africa.’ Proc. U.S. Nat. Mus., Washington,

22, no. 1199, 271-316.

Schmitt, W., 1926. ‘The Macruran, Anomuran, and Stomatopod Crustaceans collected by the

American Museum Congo Expedition, 1909-1915.’ Bull. Amer. Mus. N. H., New York, 53, 1-67,

g pl.

Stebbing, T. R. R., 1g10. ‘General catalogue of South African Crustacea.’ Ann. S. Afr. Mus., 6,

281-593, pl. XV—-XXII.

, 1920. “South African Crustacea.’ IX. ibid., 17, pt. 4, 231-72, pl. XVIII-XXVII.

Stimpson, W., 1858. ‘Prodromus descriptionis animalium evertebratorum, quae ... VII. Crustacea

Anomoura.’ Proc. Acad. Sci. Philadelphia, 63-90.

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Deuxiéme patte thoracique gauche, face interne: en bas, Paguristes gamianus H. M.-Edw.;

en haut, P. barnardi sp. nov.

The ANNALS OF THE SOUTH AFRICAN M USEUM are issued 1 in se at ee.

intervals as material becomes available. As far as possible each volume is devoted cxchineliy

to a particular subject (Zoology, Botany, etc.).. Two or more volumes may be in course of

publication concurrently.

Most of the Geological and ON papers are vues in conjunction. with the — 1

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Some volumes and parts are out of print, and others are Snly sold as paris ofa set, or volume, — if

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Out of print:

XXIV (Part 2), XXXI ice WT} ee

Zoology oe ae aN ee ee

Palaeontology

Geology, Palaeontology, Zooey, Anthropology

sLoology i.e. ic 2

Botany (era Part eu

Zoology

Palaeontology ag Golo .. (excl. Part 7)

prea ine and Zoology —.... ae cc ade

Zoology ee a Me

Zoology

Botany .. as “* BSE! ee Seager

Zoology .. Bee ey ae oe eather):

Zoology .. ste Bee ea Sy se

Zoology .. a a ie wet), Oy are eee

Zoology ae

Zoology

Anthropology dnd: iiaoleey

Zoology eis

Zoology

Anthropology.

Palaeontology ve 24 Bae

Zoology .. a He wat ie LO eae

Zoology oe

INDEX of papers, authors, and subjects, published i in Vols. I-XXX Bs

- Vol.

TIT. 1903-1905

IV. 1903-1908

V. 1906-1910

VI. 1908-1910

IX. aigi1i1—1918

X. IQ1I-1914 ~

XI. rg11—1918

XII. 1913-1924

XIII. 1913-1923

XIV. 1915-1924

XV. 1914-1916

XVI. 1917-1933

XVII. 1917-1920 |

XVIII 1921

XIX. 1924-1925

XX. 1924-1926

XXI. | 1925-1927

XXIII. 1925-1926

XXIV. 1929-1938

XXV. 1927-1928

XXVI. 1928

XXVIII. 1929

XXVIII. 1929-1932

XXIX. 1929-1931

XXX. 1931-1935-

XXXI. 1934-1950

XXXIT. 1935-1940

XXXITI. 1939

XXXIV. 1938

XXXV~.

XXXVI. 1942-1948

XXXVIT. 1947-1952

XXXVIII. 1950

XXXIX. 1952

XL. 1952-

XLII. rg952-

XLII. 1953-

except the Geological and Palaeontological parts, which are obtainable from the

Palaeontology Sistem akg ac 4 .. (Part 4 only) =

Zoology .. ne ie is wis: Veeigtean aes

Zoology re

Zoology

Reserved for conclusion of monograph i in tel XXXIV.

TOONOR YS KS oe =e ae neat Es

Archaeology ce = a = a eeat ne

Zoology

Zoology : a6

Botany f Part Esc “Pak 2

Zoology oe is Be 6d. Part 2, 7s.6d. Part 3

Palaeontology Part 1, 125. 6d. Part 2

Copies may be obtained from—

The LIBRARIAN, Soutu Arrican Museum, Care Town,

ERNMENT PRINTER, PRETOR

Gov. P , PRETORIA.

. (excl. Part 2) |

PNW OOW NN & w wD ODO te els ch sia bs Gs Gigs moo om th

Cc ONW HD

a eS ‘ 4 . a

Vols. I, II, V (Parts 1, 2, 9), VII, VIII, IX (Part 1), XII (Part a XXII,

Whee

LO O° SO r.0 6 6 oomo00 CMOCORMOM0O0M000®&

2 O2NDMOOCO

ANNALS

SOUTH AFRICAN MUSEUM

: - VOLUME XLI

PART V, containing :— : ae

. 9. New Species of Hydrozoa from South Africa. By N. A. H. Mivrarp,

Department of Zoology, University of Cape Town. (With three

text-figures.) —

10. South African Parasitic Copepoda. By K. H. Barnarp, D.Sc., F.L.S.

(With 33 text-figures.) |

11. The Breeding and Growth of Hymenosoma orbiculare Desm, (Crustacea,

Brachyura). By G. J. BRoEKHUYSEN, PH.D., Department of Zoology,

University of Cape Town. (With 13 text-figures.)

ISSUED FEBRUARY 1955 PRICE 20s. —

PRINTED FOR THE

TRUSTEES OF THE SOUTH AFRICAN MUSEUM .

BY THE RUSTICA PRESS (PTY.) LIMITED, COURT ROAD, WYNBERG, CAPE

Me; ai ER

by y,

9g. New Species of Hydrozoa from South Africa.*—By N. A. H. Mixvarp,

| Department of Zoology, University of Cape Town. (With three text-

| figures. )

INTRODUCTION

The species described were collected by members of the Zoology Department

during the course of an ecological survey of the estuaries and shallow coastal

waters of South Africa. ‘The survey has received financial support from the

Carnegie Corporation, the South African Council for Scientific and Industrial

Research, the Staff Research Fund of the University of Cape Town, and the

Natal Provincial Council. Specimens bearing the reference letters FAL were

collected from False Bay, Cape, by Mr. J. C. Morgans, who is investigating

the bottom fauna of this area by diving and by dredging.

Fam. HyDRACTINIDAE

Aydractima altispina n. sp.

Fig. 1

Description. Colonies living on the shell of the gastropod Thais (Purpura)

squamosa. Hydrorhiza covered with a layer of free coenosarc, from which the

zooids arise direct. Spines long, smooth, hollow and horn-coloured.

Gastrozooids creamy-white in colour with a pink area below origin of

tentacles, in preserved specimens short and fat, generally shorter than the

spines. Tentacles 5-12 in number, but usually 6 or 8, with long and short ones

alternating, about four times the length of manubrium in living specimens,

short and lumpy when preserved. Several bifurcating tentacles observed. No

spiral zooids or tentacular filaments.

Gonozooids much smaller than gastrozooids, with mouth and 3-5 very short

tentacles. Sporosacs borne on short stalks below tentacles and near base of

gonozooid, male and female on separate colonies. Male sporosacs spherical,

with width slightly exceeding length; with four radial canals and a circular

* The South African Museum collection of Hydrozoa consists mainly of the material

collected many years ago by the Cape Government trawler s.s. Pieter Faure. Samples from this

collection were submitted to an overseas specialist, who for various reasons was prevented from

reporting on them. The collection has now been submitted to Dr. Millard who will study it in

conjunction with the collection made in recent years by the Zoology Department of the

University of Cape Town. This is Dr. Millard’s first report on these collections. [Ed.]

215

VOL. XLI. PART 5.

MAR 2 D108e

216 ANNALS OF THE SOUTH AFRICAN MUSEUM

one; containing four groups of spermatogenic cells arranged around a central

spadix, which is solid except just at the base; no sign of tentacles. Female

sporosacs oval; with four radial canals and a circular one; containing approxi-

mately 32 eggs (about 8 eggs visible in transverse section, about 4 on each side

in longitudinal section), arranged around a central, hollow spadix.

Fig. 1. Hydractinia altispina n. sp.

A. Portion of preserved colony on shell of host, showing an extra large gastrozooid, a small

gastrozooid, a male gonozooid and a number of spines.

B. Transverse section of male sporosac, showing the 4 groups of spermatogenic cells surrounding

the spadix, and the 4 radial canals.

C. Longitudinal section of female sporosac, passing through a radial canal on one side.

Measurements. Spines reaching 0:98 mm. in length. Gastrozooids reaching

4. mm. in length to origin of tentacles when expanded, up to 1:03 mm.

preserved.

Gonozooids reaching 0:44 mm. in length to origin of tentacles (preserved).

Male sporosacs, max. length 0:42 mm., max. width 0-51 mm.

Female sporosacs, max. length 0-46 mm., max. width 0:36 mm.

Cotypes in Zoology Department, University of Cape Town, nos. F 274,

CP 258, B 92, FAL 7 Z. CP 258 also in S.A. Museum.

Localities. St. James (intertidal), Lambert’s Bay (intertidal), False Bay

(dredged from 24 metres), |

NEW SPECIES OF HYDROZOA FROM SOUTH AFRICA P49

Remarks. ‘This species possibly includes the specimens from South West

Africa described by Broch, 1914, as Hydractinia sp. The size of the spines and

gastrozooids, and the number of tentacles on the latter, are within range, but

the name of the host-snail was not given and no gonozooids were described.

Apart from this, only two species of Hydractinia have previously been recorded

from South Africa. Of these H. pacifica Hartlaub 1905 is without spines, and

H. parvispina Hartlaub 1905 has very short spines.

Ripe sporosacs have been found in March, July and August, and sterile

colonies in February and April.

The host-snail, Thais squamosa, typically inhabits the lower parts of the inter-

tidal area in the Cape Province, and is also brought up occasionally in dredgings

in False Bay.

Hydractima kaffraria, n. sp.

Fig. 2

Description. Colonies living on shell of Nassarius krausstanus. Wydrorhiza

covered with a layer of free coenosarc. Zooids arising direct from hydrorhiza.

No spines.

Gastrozooids with 8-15 tentacles arranged in two close-set, alternating

verticils. Manubrium conical when contracted, generally club-shaped when

expanded.

No spiral zooids at lip of shell, but a few tentacular filaments sparsely and

irregularly scattered amongst the zooids.

Gonozooids smaller than gastrozooids, with 5-12 tentacles arranged in two

close-set, alternating verticils. Mouth apparently present, though very small.

Sporosacs spherical, borne immediately below tentacles, each on a short stalk,

male and female on separate colonies. Male sporosac with four radial canals

and a circular, and with indications of a velum and rudimentary tentacles on

inner edge of umbrella margin. Female sporosac with four radial canals and a

circular, containing 21-32 ova arranged around a central spadix.

Measurements (preserved). Tentacular filaments up to 3:0 mm. long.

Gastrozooids reaching 2:0 mm. in length to origin of tentacles.

Gonozooids reaching 1-1 mm. in length to origin of tentacles, usually about

0-7'mm. Stalk of sporosac about 0-05 mm. long.

Male sporosac reaching 0:55 mm. in length by 0-50 mm. in diameter.

Female sporosac reaching 0-45 mm. in length by 0-40 mm. in diameter.

Cotypes in Zoology Department, University of Cape Town, nos. BRE 111 A,

female colony, from the Breede River Estuary; HAM 3 Q, male and female

colonies, from the Keiskamma River Estuary. Paratype SUN 3 N in S.A.

Museum.

Localities. Breede River Estuary, Knysna Estuary, Sunday’s River Estuary,

Kleinmond Estuary (Bathurst Division), Bushman’s River Estuary, Keiskamma

River Estuary, Durban Bay.

218 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks. ‘This species was recorded by Day, Millard and Harrison (1952)

from Knysna Estuary as Hydractinia sp. It is closely related to H. pacifica

Hartlaub, which species, however, has no radial canals and only one ovum in

the female sporosac. It is also close to H. parvispina Hartlaub, from which it

differs in the absence of spines.

Fig. 2. Hydractinia kaffraria n. sp.

A. Three individuals of a colony: gastrozooid, tentacular filament and female gonozooid

bearing sporosacs.

B. Longitudinal section of female sporosac.

C. Longitudinal section of male sporosac.

Ripe sporosacs have been found in January, February, July and September.

Since no empty sporosacs have been observed there is a possibility that these

structures have a short free-living existence before the discharge of the sexual

products.

The host snail, Nassarius kraussianus, is common in the Zostera beds of the

estuaries of the Cape Province and Natal. The hydroid has its centre of distri-

bution in the Transkei area of the Cape Province, extending as far north as

Durban on the east coast and as far west as the Breede River on the south coast.

NEW SPECIES OF HYDROZOA FROM SOUTH AFRICA 219

Fam. LAFOEIDAE

K.ygophylax cornucopia n. sp.

Fig. 3

Description. WHydrorhiza creeping on the posterior surface of the stem of

Antenella africana Broch, divided irregularly by slightly oblique nodes.

Colony stolonial, with hydrothecae arising directly from the hydrorhiza,

each on a short stem. Stems directed alternately to right and left, forming two

rows more or less at right angles to one another. Stems also directed upwards

forming an angle of roughly 45° with the hydrorhiza. Stem borne on apophysis

Fig. 3. <ygophylax cornucopia n. sp.

A. Normal colony growing on Antenella africana.

B. A single hydrotheca with female gonotheca.

C and D. Two forms of branching which occur in the paratype FAL 217 N.

220 ANNALS OF THE SOUTH AFRICAN MUSEUM

of hydrorhiza, and consisting usually of one basal segment and a hydrothecal

pedicel which is not sharply demarcated from the hydrotheca. Sometimes there

is no node separating the basal segment from the apophysis. Apophysis and

basal segment devoid of nematothecae.

Pedicel and hydrotheca together forming a cornucopia-shaped figure,

curving evenly outwards and then upwards, widening towards mouth. Pedicel

more than twice length of hydrotheca, and bearing one nematotheca about

half-way along its adcauline side. Nematotheca barrel-shaped, slightly longer

on side away from pedicel, mounted on short stalk.

Hydrotheca bilaterally symmetrical, aperture oval and facing upwards,

margin smooth and more or less perpendicular to hydrorhiza. Diaphragm

oblique and bilaterally symmetrical, sloping downwards and inwards towards

its aperture which is on the adcauline side.

Hydranth with abcauline blind pouch, conical proboscis and about 18

tentacles. Can be completely retracted into hydrotheca.

Gonothecae not aggregated into coppinia, but borne singly upon hydrothecal

pedicels, at more or less the same level as nematothecae. Female gonotheca

long and slender, with no distinct stalk, tapering at proximal end to its attach-

ment, truncated at distal end, oval in section. Containing numerous eggs

arranged in a mass around a central blastostyle, which extends for about

3 length. Male gonotheca similar in shape but slightly shorter, containing a

dense mass of spermatogenic cells. Male and female on separate hosts.

Notes

In one of the paratypes (FAL 217 N), in addition to the normal form, some

branching individuals occur, in which secondary stems (including basal seg-

ment, pedicel and hydrotheca) may arise from a primary one. The secondary

stems arise from the sides of the primary hydrothecal pedicel at about the same

level as the nematotheca. As many as 3 secondary stems have been seen to

arise from one primary in this way, or a primary stem may give rise to a

secondary, and this to a tertiary.

In another paratype (FAL 78 S) an additional very short segment is

occasionally interpolated between the normal basal segment of the stem and

the apophysis of the hydrorhiza.

Gonophores have been found in June, August and December.

Measurements (Holotype)

Diameter of hydrorhiza: 0°03 — 0:05 mm.

Length of basal segment: 0:06 — o-12 mm.

Length of hydrothecal pedicel, abcauline: 0°22 — 0-29 mm.

a He a ,, to diaphragm aperture: o-18 — 0-25 mm.

Length of hydrotheca, abcauline: 0:06 — 0:10 mm.

5 se 5 adcauline: 0:05 — 0:09 mm.

Max. width of hydrotheca, at margin: 0-10 — 0-13 mm.

> 99 » at diaphragm aperture: 0:06 — 0:08 mm.

NEW SPECIES OF HYDROZOA FROM SOUTH AFRICA PEPE

Length of nematotheca plus stalk: 0:05 — 0:06 mm.

Female gonotheca, length: 0-50 — 0°83 mm.

- ss max. width: 0-14 — 0:29 mm.

Male gonotheca, length: 0-49 — 0°56 mm.

. i max. width: 0-18 — 0-24 mm.

In the paratypes some of the measurements show a slightly larger variation,

as follows:

Length of hydrothecal pedicel, abcauline: 0-20 — 0:29 mm.

ee 55 fy ,, to diaphragm aperture: 0-15 — 0:25 mm.

Length of hydrotheca, abcauline: 0:06 — 0-12 mm.

Max. width of hydrotheca, at margin: 0-10 — 0-15 mm.

Length of nematotheca plus stalk: 0:03 — 0:06 mm.

Holotype in Zoology Department, University of Cape Town, no. FB 131 B,

from dredgings in False Bay, at 5-8 metres. Numerous colonies on Antenella

africana, some bearing gonophores.

Paratypes: TB 1 B, FAL 78S, FAL 169 X, FAL 217 N, in Zoology Depart-

ment; FAL 78 S also in S.A. Museum.

Localities. False Bay, from 4 to 18 metres. Table Bay, from 19 to 20 metres.

Remarks. Kygophylax cornucopia differs from other members of the genus in the

stolonial nature of the colony, absence of coppinia, position of nematophores,

length of hydrothecal pedicel and asymmetry of diaphragm.

It is included in the genus <ygophylax for the following reasons. Levinsen,

1913, maintained that among the hydroids the arrangement of the hydrothecae

in the colony is not a good diagnostic character. In the genus <ygophylax this

is borne out by the fact that in %. valdiviae Stechow 1923 a, hydrothecae may

arise either from upright stems (which are unfascicled) or direct from the

hydrorhiza.

Further, Levinsen and also Broch, 1918, maintain that the presence of

coppinia in the Lafoedae is not a generic character. They are present, for

instance, in some species of Lafoea and not in others. Totton, 1930, has pointed

out that although coppinia are present in &. szbogae Billard 1918, some of the

gonothecae arise separately.

In shape the hydrotheca and its pedicel resemble most those of Z. szbogae

Billard 1918, where also the double curvature is apparent and the hydrothecal

pedicel is long.

XK. africana Stechow 1923 b is the only other species so far reported from

South Africa. No reproductive structures were described, but otherwise it

differs from <. cornucopia in the features listed above.

REFERENCES

Billard, A., 1918. ‘Notes sur quelques espéces d’hydroides de l’expédition du “Siboga’”’.’ Arch.

Kool. exp. gén. Paris, vol. LVII, pp. 21-7.

Broch, H., 1914. ‘Hydrozoa benthonica.’ Beitr. Kennt. Meeresfauna Westafrikas von W. Michael-

sen, vol. I, pp. 19-50.

222 ANNALS OF THE SOUTH AFRICAN MUSEUM

Broch, H., 1918. ‘Hydroida II.’ Danish Ingolf Exped., vol. V, pp. 1-205.

Day, J. H., Millard, N. A. H. and Harrison, A. D., 1952. “The Ecology of South African

Estuaries. Part III. Knysna: A Clear Open Estuary.’ Trans. Roy. Soc. S. Afr.,

vol. XXXIII, pp. 367-413.

Hartlaub, C., 1905. ‘Die Hydroiden der magalhaensischen Region und chilenischen Kiiste.’

Fauna Chilensis, vol. III, pp. 497-714.

Levinsen, G. M. R., 1913. ‘Systematic Studies on the Sertulariidae.’ Vidensk. Medd. naturh

Foren. Kjob., vol. LXIV, pp. 249-323.

Stechow, E., 1923 a. ‘Neue Hydroiden der Deutschen Tiefsee-Expedition, nebst Bemerkungen

uber einige andre Formen.’ ool. Anz., vol. LVI, pp. 1-20.

Stechow, E., 1923 b. ‘Ueber Hydroiden der Deutschen Tiefsee-Expedition, nebst Bemerkungen

uber einige andre Formen.’ Zool. Anz., vol. LVI, pp. 97-119.

Totton, A. K., 1930. ‘Coelenterata. Part V. Hydroida.’ Brit. Antarct. (“Terra Nova’) Exped.,

1910. Nat. Hist. Rep., Zool., vol. V, pp. 131-252.

10. South African Parasitic Copepoda—By K. H. Barnarp, D.Sc., F.L.S.

(With 33 text-figures.)

This paper includes the species of South African Parasitic GCopepods which

have been already recorded in the literature, together with the additional

species of which I have been able to examine specimens.

Most of this material is in the South African Museum collections, but I wish

to thank Dr. Fitzsimons, Director of the Transvaal Museum; Dr. Hewitt,

Director of the Albany Museum, Grahamstown; Dr. Lawrence of the Natal

Museum, and Dr. van Hille and Mr. Macnae of Rhodes University, Grahams-

town, for submitting specimens for identification. Mr. A. C. Harrison of the

Cape Provincial Inland Fisheries has obtained specimens from fresh-water

fishes; and Mr. C. L. Biden has collected very useful material while investi-

gating sharks, including species hitherto unrecorded from South Africa. I wish

to thank also the Librarian of the Medical Library, University of Cape Town,

for access to periodicals not available in the Museum.

The greater part of the South African Museum collection has been derived

from the collection of marine fishes made by the Cape Government trawler

s.s. Pieter Faure. My examination of several hundreds of fishes, however, pro-

duced only a rather surprisingly small number of parasites, especially in the

case of permanently fixed parasites. Undoubtedly a considerable number of

species have yet to be discovered when South African fishes are specially

examined for parasites as soon as possible after capture.

Although only a preliminary study, the present compilation may be useful

for future work. A list of the South African hosts with their parasites is given

at the end of this paper.

Suborder ARGULOIDA

1932. Wilson, Bull. U.S. Nat. Mus., no. 158, p. 11.

Fam. ARGULIDAE

1902. Wilson, Proc. U.S. Nat. Mus., xxv, pp. 635 sqq. (review of family).

1904. Thiele, Mitt. Zool. Mus. Berlin., ii, pp. 5 sqq.

1928. Monod, Rev. Zool. Bot. Afric., xvi, pp. 242 sqq. (African genera

and species, with keys).

1943. Ringuelet, Rev. Mus. la Plata, n.s. iii, no. 19, pp. 43 sqq. (neo-

tropical species).

1947. Brian, An. Mus. Argent., xlii, pp. 353 sqq. (S. American species).

1948. Ringuelet, Rev. Mus. la Plata, v, no. 33, pp. 281 sqq. (neotropical

Species).

223

224 ANNALS OF THE SOUTH AFRICAN MUSEUM

Sexes alike. Body flattened; cephalothorax shield-like, consisting of fused

head and 1st thoracic segment; 3 free thoracic segments; abdomen unseg-

mented, bilobed. Eyes distinct. Four pairs of biramous swimming legs, with

plumose setae. No ovisacs; eggs attached to stones and other objects.

Marine, estuarine, and fresh water. On the skin, fins, and in the branchial

chamber of fishes. Also on tadpoles. Both sexes leave the host at the breeding

season, and the eggs are laid on any suitable surface, in single rows, each row

containing 3-25 eggs. Monod gives a figure of eggs scattered more or less

singly on a fish scale. ‘The larvae when hatched are in general shape similar

to the adult. Accessory sexual structures in male present on the basal joints of

the posterior legs.

Although fresh-water members of this family are numerous in Central

Africa, their occurrence in southern Africa, especially the extreme south,

appears to be rare. In parts of Rhodesia, however, e.g. the Matopo Dam, a

heavy infestation has been recorded (Hey, ‘Inland Fish. Dept., Cape, Rep.

no. 3.’ 1946, p. 34).

Key to African genera

1. No ventral sucking discs. Dolops

2. Ventral sucking discs.

a. One pair of antennae. Cephalothorax constricted anteriorly. Thorax

elongate. Chonopeltis

b. Two pairs of antennae. Cephalothorax subcircular or elliptical. Thorax

short. Argulus

Gen. Dolops Audouin

1837. Audouin, Ann. Soc. ent. Fr., vi, p. xii (‘nom. nud.’ apud Neave.

Nomencl. Zool.).

1857. Heller, SB. Ak. Wiss. Wien. math. nat. kl., xxv, p. 89 (Gyropeltis).

1891. Stuhlmann, Zool. Jahrb. Abt. syst., vi, p. 152 (Gyropeltis).

1899. Bouvier, Bull. Soc. Philom. Paris, ser. 8, x, p. 53 and ser. 9, 1, p. 12.

1902. Wilson, Proc. U.S. Nat. Mus., xxv, pp. 638, 701 (in key), 730.

r904. | Mhiele) locucit. p47:

1911. Maidl, Arb. Zool. Inst. Univ. Wien., xix, p. 317 (anatomy).

1928. Monod, loc. cit., p. 243 (in key).

1943. Ringuelet, loc. cit., p. 75 (S. American species).

Cephalothorax subcircular. Anterior maxillipeds hook-like, stout, with

strong falciform claws, not transformed into sucking discs. Apical joint of

posterior maxillipeds without claws, but with spiniferous processes. No preoral

stylet.

Neave quotes Audouin’s name as nomen nudum, but Wilson (p. 638) says

Audouin’s description ‘establishes with sufficient accuracy the characteristics

of the genus’, and Bouvier and later writers accept it.

Several species in South America, one species in Africa.

Dolops ranarum (Stuhlmann)

Fig. 1 a—d

1891. Stuhlmann, loc. cit., p. 154, fig. (Gyropeltis r.).

1902, Wilson, loc. cit., p. 737, pl. 27, figs. 89 (2), 90 (dg).

SOUTH AFRICAN PARASITIC COPEPODA 225

1904. ‘Thiele, loc. cit., p. 17, pl. 6, figs. 26-31 (appendages).

1913. Cunnington, Proc. Zool. Soc. Lond., p. 263.

1928. Monod, loc. cit., p. 248.

1940. Brian, Rev. Zool. Bot. Afric., xxxiii, p. 78, figs 1-3 (Q).

Cephalothorax subcircular. Abdominal lobes narrow, 2—24 times as long as

wide, with notch on inner margin. Flagellum (an elongate setose process

attached at end of 2nd basal joint of leg on its dorsal surface) on 1st and and

legs only. Basal joint of 3rd leg 3 with a scabrous papilla pointing dorsally and

forwards.

Q11mm., 9 4°5 mm. (total length). Greyish or greenish.

Fig. 1. Dolops ranarum (Stuhlmann). a, dorsal view; b, anterior maxilliped; c, posterior maxil-

liped; d, basal joint of 3rd leg g. Chonopeltis inermis Thiele. e, adult (copy after Thiele and

Brian); jf, juvenile, transformation of anterior maxilliped into sucking disc (only some of the

supporting rods shown).

Localities. Wartebeest Poort dam, Pretoria, mouth and gills of Tilapia

mossambica (‘Transvaal and Albany Museums); Aapies River, near Pretoria,

on Barbus gunning: (Transv. Mus.); Olifants River, Middelburg, on Barbus

swierstrae (Transv. Mus.); Matopo dam, Bulawayo, on introduced Black Bass

(Huro salmonoides) (per A. C. Harrison, May 1950).

Distribution. Lake Victoria Nyanza (Stuhlmann, Cunnington), Lake Tan-

ganyika (Cunnington), Lake Nyasa (Cunnington), White Nile (Cunnington),

Lake Moero (Mweru) (Monod), Lake Albert (Brian).

On various fishes: Protopterus, Silurids, Eutropius, Lates. ‘The original

specimens were found on tadpoles (hence ranarum),

226 ANNALS OF THE SOUTH AFRICAN MUSEUM

Gen. Chonopeltis Thiele

1900. Thiele, Zool. Anz., xxiii, no. 606, p. 46.

1902. Wilson, loc. cit., pp. 701 (in key), 729.

1904. Thiele, Mitt. <ool. Mus. Berlin, ii, p. 43.

1912. Calman, Proc. Zool. Soc. Lond., p. 766.

1928. Monod, Rev. Zool. Bot. Afric., xvi, p. 244 (in key).

Front portion of cephalothorax constricted, so that the sucking discs are

partially visible in dorsal view; posterior portion forming oval lateral lobes

not extending to end of thorax and thus covering only the anterior 2 pairs of

legs. Free segments of thorax elongate. Abdominal lobes pointed. Antenna 1

absent (see Calman, loc. cit.); antenna 2 simple, 4-jointed. Anterior maxilli-

peds transformed into sucking discs. Posterior maxillipeds as in Argulus. No

preoral stylet. Secondary sexual characters in ¢ on and and 4th legs, the

latter with only one natatory ramus.

One species in Africa, one in Japan.

Chonopeltis inermis ‘Thiele

ES Ies fa

1o00% | harcle loc citsps ago)

1902. Wilson, loc. cit., p. 720:

1904. Thiele, loc. cit., p. 44, pl. 9, figs. 110-16 (9).

1913. Cunnington, Proc. Zool. Soc. Lond., pp. 263, 280 (locality only).

1928. Monod, loc. cit., p. 260, pl. 1, fig. C, pl. 7, figs) GD ae

1940. Brian, Rev. Zool. Bot. Afric., xxxiil, p. 80, figs. 1-14 (2, g, juv.)

(var. schoutedent).

1951. WDartevelle, Zooleo., n.s. no. 9, p. 12, figs. 1, 2)(juv.', juvenag van.

schoutedent) .

Front portion of cephalothorax subquadrangular, with more or less developed

median indent on front margin; ventrally front margin with chitinous border

from which 2 pairs of ribs converge backwards; the antenna arises between

the inner and outer rib of each pair. The length of the abdominal lobes in

relation to basal portion variable (see Brian’s figures). In 2 mm. juveniles

(S. Afr. Mus.) the lobes are shorter than the basal portion.

Up to 9 1omm., ¢ 7 mm. (Brian).

Locality. Okawango River, at the bases of the barbels on chin, in folds of

lower lip, and in axil of pectoral fin of Synodontis melanostictus. (S. Afr. Mus.

3 juv.)

Distribution. East Africa, on ‘Chromis’ (Cichlidae) (Thiele); Lake Nyasa

(Cunnington); Congo, in branchial cavity of Gnathonemus and Marcusenius

(Monod, Brian).

Remarks. ‘The 3 specimens in the S. African Museum are juveniles 1-75-2 mm.

in length, smaller than any in Brian’s material. They show the transformation

of the anterior maxillipeds from a chelate appendage into a sucking disc

(fig. tf) (cf, Wilson, rgoa, loc. cit., fig. 11, after Claus).

SOUTH AFRICAN PARASITIC COPEPODA 227]

Dartevelle claims schoutedent as a separate species characterized by

lengthening of the hinder part of body, different shape of appendages, and

particularly the smallness of the 4th pair of legs.

Gen. Argulus O. F. Miller

1785. Miiller, Entomostr., p. 121.

1902. Wilson, loc. cit., pp. 701 (key to species), 704.

1913. Cunnington, Proc. Zool. Soc. Lond., p. 265 (key to African species).

1928. Monod, loc. cit., pp. 244 (key to African species), 249.

1931. id., Rev. Zool. Bot. Afric., xxi, p. 18 (emendation to key).

1940. Brian, ibid., xxxiii, p. 87.

1940. Meehean, Proc. U.S. Nat. Mus., |xxxviil, pp. 459 sqq.

1943. Ringuelet, loc. cit., p. 54 (S. American species).

1944. Wilson, Proc. U.S. Nat. Mus., xciv, pp. 551 sqq.

Cephalothorax subcircular or elliptical. Free segments of thorax short

(normally). T'wo pairs of antennae, the anterior pair armed with stout hooks.

Anterior maxillipeds transformed into sucking discs. Preoral stylet present.

Secondary sexual characters in § on 2nd—zth legs.

Marine and fresh water. Very few species have been obtained from marine

fishes around the African coast. Monod (1928) records 4 species from the

whole of the Mediterranean and western coast of Africa. Two more species

are recorded below, occurring on the south-east side of the continent.

The genus is common on fresh-water fishes in Central Africa, but hitherto

none have been recorded from the southern region. One species is herein

described; it is evidently very rare; in the course of revising the south-west

Cape fishes I have examined many thousands of specimens, both in the field

and in the Museum collections, but without finding a single specimen of

Argulus.

The following species have been recorded from localities sufficiently close to

our South African area to raise a reasonable expectation that they may be

found to occur actually within our boundaries. They are included in the key,

but not described here.

Marine

A. alexandrensis Wilson 1923 (g). Port Alexander, on Zeus. (syn. otolith

Brian 1940 (g). Cameroons, on Pseudotolithus).

A. darieveller Brian 1940 (9). Congo, on Polynemus.

Fresh water

A. africanus Thiele 1900 (2 g). Lakes Nyasa and Moeru. Widely distributed

in Central Africa on various fishes.

. reticulatus Wilson 1920 (2 3). Congo River, on Hydrocyon.

. wilsonit Brian 1940 (¢). Congo River, on Hydrocyon.

. rhipidiophorus Monod 1931 (@ ¢). Lake Albert, on ? Hydrocyon.

. schoutedent Monod 1928 (9 g). Katanga.

. ryckmansu Brian 1940 (2). Matadi, River Congo.

Cunnington (1913) criticized Wilson’s 1902 identification key on the ground

that the relative length of the carapace was an unsatisfactory and inconstant

hs AS AR A A

228 ANNALS OF THE SOUTH AFRICAN MUSEUM

character. Wilson (1923) counter-criticized. Where the carapace is manifestly

so short as to leave the last two thoracic segments completely uncovered, one

may assume this condition to be natural and not produced by contraction or

distortion in preservative (e.g. schouteden). Where, however, ‘species’ are

distinguished and keys constructed according as the carapace just reaches, or

just does not reach, to the base of the abdomen, or overlaps the abdomen to a

greater or less extent, one does not feel on such sure ground. The varying

action of different methods of preservation, or of similar preservative on

different individuals, does not appear to be fully realized by all investigators.

Key to South African species (and some other African species)

Marine

I. Abdominal lobes pointed.

a. Carapace evenly elliptical, not extending beyond base of abdomen;

antero-lateral sinuses slight; suckers not visible in dorsal view. alexandrensis

b. Carapace elliptical, extending to middle of caudal lobes, antero-

lateral sinuses deep; suckers partly visible in dorsal view. multipocula

c. Carapace narrowed in front, but without definite sinuses, extending

to base of abdomen; suckers partly visible in dorsal view. belones

II. Abdominal lobes rounded. dartevellei

Fresh Water

I. Last 2 thoracic segments entirely free and visible. Abdominal lobes

pointed. schoutedent

II. Carapace covering more or less all the thoracic segments and extending

at least to base of abdomen.

A. Abdominal lobes rounded.

St : : rhipidiophorus

1. Basal plate of maxilliped with 3 spines (2 at least acute). { Peenan

2. Basal plate with 3 lobes.

a. Lobes close together. africanus

: : reticulatus

b. Lobes widely separated by triangular spaces. { Bratt

B. Abdominal lobes pointed (bluntly). capensis

Wilson in 1902, and in the descriptions and explanation to figures in 1944

used the term ‘Maxilliped’, but in his 1944 Key followed Meehean 1940 in

terming these appendages (Second) Maxillae.

Argulus belones van Kampen

Fig. 2

1909. Wan Kampen, ool. Anz., xxxiv, p. 443, figs. 1-4 (9).

@—Carapace elliptical, narrowed in front, posterior lobes broadly rounded,

almost reaching the abdomen. Eyes moderate, far forward, their distance

apart about twice the distance of eye from lateral margin. Anterior respiratory

area subsemicircular (inner margin nearly straight), well separated from the

elongate reniform posterior area. Abdomen broadly ovate, a little longer than

wide (5:4), sinus extending 2 length, narrowly triangular, lobes pointed, anal

lobes minute. Antenna 1 with small claw on anterior margin, strong apical

claw, and a small projection on hind margin. Antenna 2 4-jointed, sparsely

setose. Suckers situated well forward, partially visible in dorsal view where the

carapace narrows, completely hiding the eyes in ventral view; rods supporting

the margin composed of a number of short discs or cups, each wider than long.

Accessory spines between maxillipeds apparently absent. Maxilliped robust,

SOUTH AFRICAN PARASITIC COPEPODA 229

basal plate with large scabrous area, hind margin trilobed, the lobes rounded,

the middle one smaller than the others; 2nd and grd joints with apical scabrous

areas, 4th and 5th joints abruptly narrower than 3rd, 4th with a few scabrosities

on outer margin, 5th tipped with 2 points and a spine. Basal joint of 4th leg

with trapezoidal projection on hind margin, the outer posterior corner pointed.

No flagella on any of the legs.

One 2 15 mm., two 99 also with full ovaries 8 and 9 mm., one immature

5mm. Greenish.

Fig. 2. Argulus belones van Kampen. a, 9, respiratory areas

dotted (not visible in dorsal view); 6, antennae 1 and 2.

¢, portion of margin of sucking disc; d, posterior maxilliped;

eé, basal joint of 4th leg.

Locality. Natal, on side of head of Barracuda (Sphyraena commersoni) (collected

by the late H. W. Bell-Marley, the largest example ex Natal Museum).

Distribution. Sumatra, on Belone schismatorhynchus.

Remarks. ‘The specimens described above show all the specific features of

belones, and are clearly conspecific. Van Kampen’s 13? mm. 9, however, had

the posterior lobes of the carapace meeting (‘sogar miteinander verwachsen

oder verklebt’) in the middle line, and extending backwards to cover half the

abdomen. This shows the danger of regarding the carapace length as a con-

stant specific character, and of utilizing it to separate species in a key.

Van Kampen mentions a pair of blunt spines between the maxillipeds, but

they are not traceable in the present material.

230 ANNALS OF THE SOUTH AFRICAN MUSEUM

This species is distinguished from alexandrensis Wilson, arcassonensis Cuenot,

and alosae Gould (cf. remarks of Wilson, 1923, p. 5, and Monod, 1928, p. 255)

by the lateral indentations of the carapace, and also the eyes and suckers, being

much farther forward; the relatively broader and more deeply incised

abdomen; the very robust proximal joints of the maxilliped with the two

abruptly narrower distal joints.

So far as I am aware, this species has not been recorded since its description.

Its occurrence in Natal waters, and on a fish of a different family, is therefore

interesting.

Fig. 3. Argulus multipocula n. sp. a, 2, respiratory area dotted; 6, portion of

margin of sucking disc; c, maxilliped, with edgewise view of 5th joint;

d, process of basal joint of 4th leg.

SOUTH AFRICAN PARASITIC COPEPODA 231

Argulus multipocula n. sp.

Fig. 3

@—Carapace broadly elliptical, with deep antero-lateral sinuses, posterior

lobes rounded, extending to middle of caudal lobes. Eyes distinct, separated

by a distance equal to that between eye and lateral margin of carapace.

Posterior respiratory area reniform, anterior area not clearly traceable, but

apparently subtriangular or oval. Abdomen ovate, cleft to a little beyond

middle, lobes apically pointed. Antennae 1 and 2 asin belones. Suckers partially

visible in dorsal view in the antero-lateral sinuses of the carapace. Rods

Fig. 4. Argulus capensis n. sp. a, 2, respiratory areas dotted; 5, antennae 1 and 2; ¢, por-

tion of margin of sucking disc; d, maxilliped, with one of the pair of accessory spines.

supporting the margin composed of very numerous extremely short cups

(multipocula) producing a transversely striated appearance. Accessory spines

between maxillipeds absent. Basal plate of maxilliped trapezoidal, entire,

without any lobes or spines, no scabrosities on any of the joints; 5th joint with

2 oval plates, basally fused, distally separate but adpressed, the dorsal one with

scarious, minutely crenulate anterior margin. Basal joint of 4th leg with

digitiform process. No flagella on any of the legs.

10 X 7mm. (anterior margin to tips of posterior lobes of carapace 9 mm.).

Locality. Richards Bay, Natal. One ovigerous 9, not attached to any host,

but found among the usual miscellaneous collection of littoral animals obtained

by shore collecting (University of Cape Town Ecological Survey, 1949).

232 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks. ‘The distinctive specific characters are the deep antero-lateral

sinuses, the entire basal plate and bifid nonunguiculate apex of the maxilliped,

and the striated appearance of the supporting rods of the suckers. The latter

feature invites comparison with A. melanostictus Wilson 1935 (see: Wilson, loc.

cit., 1944, p. 565, pl. 25, fig. 91) from California and Siam.

Argulus capensis n. sp.

Fig. 4

9—Carapace subcircular, slightly longer than broad (4 xX 3:75), front

broad, lateral indents distinct but shallow, posterior lobes broadly rounded,

reaching almost to base of abdomen. Eyes small. Anterior respiratory area

wholly in front of the posterior area. Abdomen ovate, cleft nearly to midway,

lobes ovate, bluntly pointed. Antenna 1 with small claw on anterior margin,

strong apical claw and small projection near hind margin. Antenna 2 4-jointed

sparsely setose, basal joint with rather strong spine. Supporting rods of sucker

composed of 10-12 cups, the basal one slightly the longest. Basal joint of

maxilliped with 3 strong spines, scabrous area pear-shaped. A pair of accessory

spines between the bases of the maxillipeds, but no additional pair. No

flagella on any of the legs.

6 mm. Whitish, the respiratory areas marked by dark interrupted line.

Locality. Zoetendals Vlei (Bredasdorp District), on the Cape Kurper

(Sandelia capensis). (Transvaal Mus. 1 @ 1941.)

Remarks. ‘This species is undoubtedly closely similar to A. personatus Cunn.

1913, from Lake Tanganyika. It differs in having no flagella on the legs, the

basal joint of 4th leg (2) not so strongly produced postero-laterally, and no

additional pairs of spines behind the pair between the bases of the maxillipeds.

Suborder CYCLOPOIDA

1913. Scott, T. & A., Brit. Parasit. Copepoda, i, p. 33.

1918. Sars, G. O., Crust. Norw., vi, p. 1.

1932. Wilson, Bull. U.S. Nat. Mus., no. 158, pp. 310 and 583 (key to

genera).

Thorax (cephalothorax) usually more robust than, and sharply demarcated

from, the narrow abdomen. Antenna 1 slender and elongate, or short and stout,

those of ¢ alike on both sides, sometimes prehensile. Antenna 2 uniramous

(some parasitic forms with rudimentary exopod), sometimes prehensile.

Mouth-parts varying according to habits. First 4 pairs of legs well developed,

but with exceptions in @ of some parasitic forms. Fifth pair of legs small,

simple, alike in both sexes. Two lateral or subdorsal egg-sacs.

The group includes free-swimming, commensal, and parasitic forms, both

marine and fresh water.

Sars includes Sabelliphilus in the family Lichomolgidae, and Cancerilla in the

Cancerillidae. Dogiel compares Entobius with Enterognathus and Myizlicola.

SOUTH AFRICAN PARASITIC COPEPODA 233

Key to South African genera

1. Cephalothorax ovate, abdomen segmented.

a. Each ramus of first 4 pairs of legs 3-jointed. Antenna 2 with 3 (2) apical

claws. On Polychaet worms. Sabelliphilus

b. Rami of legs unevenly jointed.

i. Rami of 2nd, grd, and 4th legs 3-jointed. Antenna 2 with 4 large

apical claws. On fishes. Ancistrotos

ii. First 3 pairs of legs more or less reduced; 4th absent. Antenna 2

with one stout apical claw. On Brittle-stars. Cancerilla

2. Cephalothorax narrow elongate, abdomen unsegmented. Antenna 2 not

prehensile, with apical spine-seta. On Polychaet worms. Entobius

Gen. Sabelliphilus M. Sars

1862. Sars, M., Forh. Vidensk. Selsk. Christiania, 1861, p. 139.

1918. Sars, G. O., loc. cit., p. 187.

1932. Wilson, loc. cit., p. 587 (in key).

Thorax in 9 (typically) narrow ovate, abdomen slender. Antenna 1

7-jointed. Antenna 2 prehensile, uniramous, 4-jointed, 2nd joint with 6 stout

teeth on inner surface, 3rd joint with curved claw, 4th joint subequal to or

shorter than 3rd, with 3 claws (normally). Each ramus of first 4 legs 3-jointed.

Fifth leg reduced to a single distinct joint with 2 setae.

Ectoparasitic on Sabellid Polychaet worms.

Sabelliphilus (?) bispirae M’ Intosh

Fig. 5 a-c

1904. M’Intosh, Mar. Invest. S. Afr., ui, p. 86, pl. 9.

Thorax ovate, not twice as long as wide, longer than abdomen, Ist segment

(2 g) and 2nd segment (Q) with angular, more or less projecting postero-

lateral corners. Abdomen narrow. Caudal rami narrow, 5 times as long as

wide. Antenna 2 with only 2 apical claws.

Ooi? mm..) G) O:7 mm.

Locality. Cape, on Bispira volutacornis (M’Intosh).

Remarks. M?Intosh notes that this species does not agree with Sabelliphilus in

having only 2 apical claws on antenna 2, and in other details; and quotes

Scott’s opinion that it cannot be satisfactorily ascribed to any described genus.

In Wilson’s key this species might be followed down to Ubperogcos, but

M’Intosh does not state the exact number of joints in antenna 2.

Gen. Ancistrotos Brian

1906. Brian, Copep. Parasit. Pesct. Ital., p. 33.

1911. Wilson, Proc. U.S. Nat. Mus., xxxix, pp. 384-6, and pp. 3091,

392 (with key to species).

Goals Gls, MonGle, Ibs. euce Typ jon (oy

1O 322 pide. loc. cit., pp. 364, and (in key) 505:

1935. Leigh-Sharpe, Parasitology, xxvii, p. 266.

TOGO. Ide lbiG:. xxx, py TOO!

1939. Yamaguti, Parasit. Copep. Japan, pt. 4, Cyclopoida 2 (Vol. Jub.

Prof. Yoshida II), pp. 410-13.

234 ANNALS OF THE SOUTH AFRICAN MUSEUM

Cephalothorax ovate, wider than the free segments. Abdomen 3 or 4

segmented. Antenna 1 6-jointed. Antenna 2 prehensile, with 4 apical claws.

Each ramus of ist leg consisting of a single widened joint; rami of gnd-4th

legs 3-jointed; 5th leg 1-jointed. Egg-sacs stout, moderately long.

Fig. 5. Sabelliphilus bispirae M’Int. a, 9; b, 3; c, antenna 2 g. Cancerilla durbanensis Steph.

d, ventral view of C. neozelanica Steph. 2; e, maxilliped; jf, 2nd leg (copies: a-c, after M’Intosh ;

d-f, after Stephensen).

Ancistrotos ostracionis (Richiardi)

1870. Richiardi in Bianconi, Spec. zool. Mosambicana, fasc. x1x/xx, p. 34.7.

(Bomolchus [sic.] 0, sine descr.).

1870. id., Arch. Zool. Anat., (2) ii, p. (? 53, quotation from this page in

Wilson, 1911). (Bomolochus 0.)

1911. Wilson, loc. cit., pp. 385, and (in key) 392.

Abdomen 3-segmented, last segment longest. Terminal claw of maxilliped

as long as basal joint, and without filaments. Both rami of 4th leg with plumose

setae (Wilson, in key, 1911).

Locality. Mozambique, on Ostracion cornutum (Richiardi).

Remarks. Bianconi includes among the Zoophyta a brief note on the

occurrence of Richiardi’s species.

SOUTH AFRICAN PARASITIC COPEPODA 235

Gen. Cancerilla Dalyell

1851. Dalyell, The Powers of the Creator, i, p. 223.

Hole. |) Sars, G. ©., loc, cit., p. 138.

1927. Stephensen, Vid. Medd. Dansk. Naturh. For., \xxxiil, p. 377.

1932. Wilson, loc. cit., p. 597 (in key).

1951. Heegaard, Vid. Medd. Dansk. Naturh. For., cxiii, p. 172.

Thorax in 9 very broadly ovoid, abdomen short, 3-segmented; in g ovate,

longer than broad, abdomen elongate, 5-segmented. Caudal rami smaller in

2 with stout apical claw. First 3 pairs of legs reduced in 9, better developed in

Gg; outer ramus of 3rd leg 1-jointed, inner ramus absent; 4th leg absent;

5th leg 1-jointed, with 2 setae in 9, 5 in J.

Cancerilla durbanensis Steph.

Fig. 5 ¢, f.

1933. Stephensen, Vid. Medd. Dansk. Naturh. For., xciii, p. 198, fig. 1.

©—Thorax broadly cordate, widest near anterior end, front margin very

slightly convex; genital segment very broad, nearly half as wide as thorax,

4 times as broad as long, lateral margin angulate. Second leg with both rami

long, narrow, outer ramus with 6 setae on both inner and outer margins, inner

ramus % length of outer, with 4 setae. Ovisacs globular. 1 mm.

Locality. Off Durban, 127-225 fathoms, on the cosmopolitan Brittle-star

Amphipholis squamata.

Remarks. C. tubulata Dalyell and C. neozelanica Steph. occur on the same

species of Ophiuran in Europe and New Zealand respectively. The South

African and the New Zealand species agree in the shape of the thorax, and

differ in this respect (as well as in minor details of the appendages) from the

European species.

Gen. Entobius Dogiel

1908. Dogiel, Zool. Anz., xxxili, p. 561.

1932. Wilson, loc. cit., p. 592 (in key).

g—Body narrow, with segmental constrictions but not truly segmented.

Five thoracic segments. Abdomen unsegmented, ending in a pair of semi-

articulated caudal styles. Antenna 1 3- or 4-jointed. Antenna 2 3-jointed,

with terminal spine-seta, but not prehensile. Mandibles present. One pair of

minute maxillae. Maxillipeds robust, prehensile, consisting of a basal joint

and apical curved claw. Four pairs of biramous legs, rami of a single joint,

setose on inner margin; 5th leg absent. Egg-sacs elongate, eggs multiseriate

proximally, uniseriate distally.

Entoparasites on marine Polychaets. The type species was found in the

intestine of a Terebellid (Lozmia).

Remarks. Dogiel regards the segment behind that which carries the last pair

of legs as the 5th thoracic segment, and the following segment as the 1st

abdominal segment. The latter, however, carries the ovisacs and is clearly the

genital segment of the thorax.

236 ANNALS OF THE SOUTH AFRICAN MUSEUM

Entobius euelpis Brnrd.

Fig. 6 a.

1948. Barnard, Ann. Mag. Nat. Hist., (xii), 1, p. 242, fig. 1.

9—In general agreeing with E. loimiae. Antenna 1 3-jointed (not 4).

Abdominal segment more flask-shaped, the caudal styles stouter and closer

together (resembling a bifurcation) ; each style with a minute apical spine-seta

but no subapical or marginal setae.

8 mm., breadth about 1 mm., ovisacs 6-7 mm.

Fig. 6. Entobius euelpis Brnrd. a, 2 (ovisacs not completely drawn). Notodelphys allmani Thorell.

b, 2; ¢, 3; d, 5th leg. Doropygus pulex Thorell. e, 2. Gunenotophorus globularis Costa. var. giganteus

Schell. jf, 2; g, and leg.

Locality. One ovigerous ? found in a bottle of slimy and much decomposed

Polychaets in the s.s. Preter Faure collection; uncertain whether from the Cape

area or Natal.

Suborder NOTODELPHYOIDA

1932. Wilson, Bull. U.S. Nat. Mus., no. 158, pp. 385 and 598 (key to

genera).

1948. Lang, Ark. Zool., xl, 3, no. 14, pp. 1-36.

Ento- or ecto-parasites, or commensals; the majority living within Ascidians.

Females usually much modified and not leaving host after maturity, males

retaining capacity for free-swimming.

SOUTH AFRICAN PARASITIC COPEPODA 237

Schellenberg (1922) adopts one family, but Sars (1921) and Wilson (1932)

more than one. In these circumstances sharply defined family diagnoses are not

advisable, and Schellenberg’s scheme is adopted.

Fam. NOTODELPHYIDAE

1921. Sars, Crust. Norw., vill, p. 29 (part, includes only 2 genera).

1921. Schellenberg, A. Norsk. Vid. Selsk. Skr., no. 3, p. 3.

1922. id., Mitt. Zool. Mus. Berlin, x, p. 219 (key to genera).

Key to South African genera 99

1. Ova contained in the swollen thoracic segments. Antenna 2 with apical

claw. 5 pairs of legs in normal ventral position (Notodelphyinae).

a. Rami of tst-4th legs setose.

i. 1st thoracic segment fused with head, 2nd and 3rd free. Brood

pouch ovoid, dorso-ventrally flattened. Caudal rami with plumose

apical setae. Notodelphys

ii. Head and ist three thoracic segments distinct. Brood pouch dor-

sally gibbous. Caudal rami with minute apical setules. Doropygus

b. Rami of 1st-4th legs non-setose. Head distinct, thoracic segments much

swollen. Caudal rami non-setose. Gunenotophorus

2. Ova contained in external ovisacs. Antenna 2 without apical claw (S.

African genera). 5th legs more or less dorso-lateral in position (Asczdicolinae)

a. 5th legs digitiform. Botryllophilus

6. 5th legs lamelliform.

i. Exopods of tst-4th legs spiniform. Enterocola

ii. Exopods of 1st-4th legs falciform Kanclopus

Gen. Notodelphys Allman

1847. Allman, Ann. Mag. Nat. Hist. (1), xx, p. 2.

1878. Brady, Brit. Copep. (Ray Soc.), i, p. 125.

1921. Sars, loc. cit., p. 30.

1922. Schellenberg, loc. cit., p. 225 (key to species).

1932. Wilson, loc. cit., p. 386.

1948. Lang, loc. cit., p. 4.

9—1st thoracic segment usually fused with head, 2nd and 3rd free, separated

by deep lateral incisions. Brood pouch arising from 4th segment, ovoid, dorso-

ventrally somewhat flattened. Caudal rami each with 4 plumose setae and a

spine on outer margin. Antenna 1 15-jointed. Antenna 2 3-jointed. Ist-4th

pairs of legs biramous, both rami 3-jointed; 5th pair very small, basal joint

produced externally into a digitiform process tipped with a long spine, ramus

ovoid, with a spine and a seta.

g—All segments, except 1st thoracic, free. 1st abdominal segment slightly

swollen, with (when mature) 2 spermatophores.

Antenna I I1- or 12-jointed, last 2 joints bent, feebly prehensile.

Remarks. Several species have been described, but they are not easily

separable. The position of the spine on outer margin of caudal rami has been

used, but Schellenberg thinks the 5th pair of legs may be a better differential

character (cf. figures of 7 species in Sars).

Key to South African species

I. Q 1st thoracic segment fused with head. 3 cephalic segment longer than wide,

postero-lateral corners not elongated. allmani

238 ANNALS OF THE SOUTH AFRICAN MUSEUM

2. Q 1st thoracic segment free. ¢ cephalic segment as wide as long, postero-lateral

corners elongated. weberi

Notodelphys allmant Thorell

Fig. 6 b—d.

1860. Thorell, A. Sv. Vet. Akad. Handl., ii, p. 31, pl. 1, play.

1869. Buchholz, <. wiss. ool., xix, p. 111, figs. (mediterranea).

To70. brady, loc: cit p. 120, pli 25. figs, 110:

TO2T.) Oats, LOGwClt.. p31, pls: Tae 10

1922. Schellenberg, loc. cit., p. 227 (in key), and p. 262 (locality record).

©—1st thoracic segment fused with head. Last abdominal segment quad-

rangular. Spine on outer margin of caudal ramus at about ? length of ramus.

5th leg see fig. 6d.

2475 mm., gd 1:9 mm. Pale yellowish, eggs greenish.

Localities. Simons Bay, in Ascidia sydneiensis (Schellenberg); Still Bay, in

Ascidians, coll. T. A. Stephenson. 4 ovig. 92 (S. Afr. Mus.); mouth of

Zwartkops River estuary, Port Elizabeth, in Ascidians (W. Macnae, Rhodes

University, ovig. 99, 33).

Distribution. West coast of Europe, Mediterranean.

Remarks. Some of the specimens I have seen resemble closely Stock’s figure

of the 2 weberi as regards the outline of the hind part of head with the 1st

thoracic segment; but the latter is certainly not a free segment.

Notodelphys webert Stock

1950. Stock, Amsterdam Nat., I, 2, pp. 37-42, figs. 1-3. (2 3).

See key above. 2 4-2 mm., g 2:2 mm. Yellowish-grey.

Locality. Knysna, in Ascidia canaliculata (Stock).

Gen. Doropygus Thorell

1859. Thorell, Ofvers. Vet. Akad. Forh., xvi, p. 339 (part).

reo: pad:, A. Suv. Vem Ak. Hondl., n-f, 11 sinow oun. slop

1O2T.)) OAalss OCH Clip 4a

1922. Schellenberg, loc. cit., p. 238 (key to species).

1932. Wilson, loc. cit., p. 387.

1940. lang, loc cit.194)6;

—Body somewhat laterally compressed and curved ventrally. Head and

first 3 thoracic segments distinct; 4th and 5th segments fused and overlaid by

the large gibbous brood pouch. Caudal rami elongate, sometimes coiled

distally, apical setae not plumose, usually minute. Antenna 1 usually 9-,

sometimes 10- or 13-jointed; first 2 joints enlarged. Antenna 2 2- or 3-jointed.

Ist-4th pairs of legs basally stout and muscular, biramous, rami 3-jointed, or

endopods of 2nd-4th pairs 2-jointed; 5th pair with a single one-jointed ramus

(sometimes biramous).

g§—Body cylindrical, tapering. All thoracic segments free.

Doropygus pulex 'Thorell

Fig. 6 e.

1860. Thorell, loc. cit., p. 46, pl. 6.

1878. Brady, loc. cit., p. 133, pl. 28, figs. 1-12.

SOUTH AFRICAN PARASITIC COPEPODA 239

noo (Sars! loc. icit... p. 42, pl. 20:

nee. ochellenbers, loc: cit, pp. 241 Gn Key), 240, 271, 272, figs: 26, 27.

aga24) VVilson, loc: cit.; p.. 389, fig. 239.

Last abdominal segment cleft (Q 3). Caudal rami not apically coiled,

apical setae minute. 5th leg uniramous. Apical claw of antenna 2 as long as

2nd joint.

2 3°38 mm., ¢ 1.5 mm.

Localities. Liideritzbucht, in Pyura stolonifera (‘Red Bait’) (Schellenberg) ; St.

James, False Bay, in Pyura stolonifera, coll. K.H.B. 1913 (S. Afr. Mus.); Klein-

mond, Cape, in Ascidian (Univ. Cape Town Ecolog. Surv. 1939); mouth of

Zwartkops River estuary, Port Elizabeth, in Ascidians (W. Macnae, Rhodes

University, ovig. 99); Durban, in Microcosmus oligophyllus var. wahlbergi

(Schellenberg).

Distribution. Europe, Mediterranean, east coast of N. America, Barbados,

West Africa, Australia, Japan.

Gen. Gunenotophorus Costa

1840. Costa, Fauna Regn. Nap. Cat. Crost., p. 7.

1864. Claus, <. wiss. Zool., xiv, p. 379 (Sphaeronotus, non Laporte 1832).

1869. Buchholz, ibid., xix, p. 144 (Gunentophorus).

1879. Kerschner, Denkschr. Ak. Wiss. Wien., xli, p. 156 (Gunentophorus).

1905. Norman, Mus. Norman., ed. 2, no. 3, p. 36 (Gunetotrophus).

1910. Stebbing, Gen. Cat. S. Afr. Crust., p. 550.

1921. Sars, loc. cit., p. 56 (Gunentophorus).

1922. Schellenberg, Mitt. Zool. Mus. Berlin, x, p. 257.

1932. Wilson, loc. cit., p. 602 (in key) (Gunentophorus).

—Thorax greatly inflated, segments more or less confluent, at least dor-

sally in ovigerous individuals. Head bent ventrally. Abdomen straight, only

3 segments distinct. Antenna 1 short, compressed, joints ill-defined. Antenna

2 3-jointed, with apical claw. Mandibles well developed. One pair of maxillae,

2 pairs of maxillipeds. tst-4th pairs of legs biramous, outer rami 3-jointed,

inner ramus of Ist pair 3-jointed, of 2nd and grd pairs 4-jointed, of 4th 2-jointed;

only the rst pair setose, closely applied to the mouth-parts, inner rami of 2nd-

4th pairs small and weak; 5th pair absent. Caudal rami curved outwards,

I-jointed, with minute apical setules. No external ovisacs. Male unknown.

In Ascidians.

Remarks. 1 have not seen Kerschner’s reasons for the change in spelling.

There is not a great difference in the meaning of the two compound words.

But if Stebbing’s statement (p. 550) is correct, viz. that Costa gave no definition

of the genus or the species, then maybe Costa’s name is a nomen nudem and

Kerschner’s name should be adopted. There is no justification for Norman’s

suggested name. Stebbing also points out that Costa’s specific name may

have to be replaced by thorelli Claus 1864.

One other species: spinipes Schell. 1922, in S. America.

Gunenotophorus globularis Costa

Fig. 6/5 g.

1840. Costa, loc. cit., p. 7.

1864. Claus, loc. cit., p. 379 (Sphaeronotus thorelli).

240 ANNALS OF THE SOUTH AFRICAN MUSEUM

1910. Stebbing, loc. cit., p. 550.

TOA (Sans, LOG: Clits, P-'5i75, pl. 2a, ales os

1922. Schellenberg, loc. cit., p. 258, fig. 38 (prp. 2), and var. giganteus

p- 258, fig. 39 (prp. 2), and pp. 268, 271.

Third joint of outer ramus of 2nd leg without spines.

Up to 6-5 mm., var. giganieus up to 8-4 mm.

Localities. Var. giganteus: False Bay, in Pyura stolonifera (‘Red Bait’) (Stebbing,

and S. Afr. Mus.); Ltideritzbucht, Simons Bay, and Plettenberg Bay (Schellen-

berg); Groen River, south of Hondeklip Bay (west coast), in Pyura stolonifera

(Univ. Cape Town Ecol. Surv. 1939).

Distribution. Mediterranean and coasts of France and Scandinavia.

Remarks. The variety differs from the typical form in having the outer ramus

of 2nd leg stout and very little longer than inner ramus, instead of slender and

nearly twice as long. All the South African specimens which I have seen,

including Stebbing’s material, belong to the variety.

Gen. Botryllophilus Hesse

1864. Hesse, Ann. Sct. Nat., ser. 5,1, Pp. 345.

Toei. Sars; locscit pez

1922. Schellenberg, loc. cit., pp. 281, 296.

1932. Wilson, loc. cit., p. 392.

1948. Lang, loc. cit.jipp.'8,,.44-

—Head distinct. The 5 thoracic segments more or less completely fused,

often swollen in adult. Abdomen 4-8-segmented. Caudal rami prehensile,

each with 4 stout curved spines. Antenna 1 short, basal joint broad, other

joints reduced. Antenna 2 with end joint short, 3rd with strong spines.

Maxilliped very stout, conical, 3-jointed, with apical claw. Ist-4th legs

biramous, rami I- or 2-jointed, outer rami sometimes symmetrical; 5th legs

unjointed, digitiform, supporting on either side the round ovisacs.

g—Cyclopiform. Abdomen 6-8-jointed. Genital segment broad.

Schellenberg (p. 285) has discussed the asymmetry of the legs in the 9.

Inhabits the gill-sac and cloaca of Ascidians.

Key to the South African species

1. Outer rami of 1st-4th legs asymmetrical. africanus

2. Outer rami approximately symmetrical. aspinosus

Schellenberg (p. 294) also records a species of this genus in Polycitor rentert

from Plettenberg Bay.

Botryllophilus africanus Schell.

1922. Schellenberg, loc. cit., p. 284, figs. 3 a-d.

Length 9 1-2 mm.

Locality. Angola, in Macroclinum angolanum (Schellenberg).

Botryllophilus aspinosus Schell.

Fig. 7 a.

1922. Schellenberg, loc, cit., p. 283, figs. 1, 2 a-c.

Length 9 2 mm.

SOUTH AFRICAN PARASITIC COPEPODA 241

Locality. Angola, in Styela hupfert (Schellenberg).

Distribution. Plymouth, England.

Gen. Fnterocola van Bened.

1860. Van Beneden, Bull. Ac. Belg., ser. 2, 1x, p. 154.

1909. Chatton & Brément, Bull. Soc. zool. Fr., xxxiv, p. 223 (references).

fg21. ‘soars, loc. cit.,'p. 76.

1922. Schellenberg, loc. cit., p. 287.

©—Head distinct. Thorax cylindrical, 4-5 segmented. Abdomen short,

conical, more or less distinctly segmented. Caudal rami lamellate or digiti-

form. Antenna 1 short. Antenna 2 1-2-jointed, lamellate. Mandibles absent.

Fig. 7. Botryllophilus aspinosus Schell. a, 9. Enterocola bilamellatus

Sars. b, 2; c, 2nd leg, with median lamina. Zanclopus cephalodisci

Calman. d, 9; e, 4th leg (copies: a, after Schellenberg; 6, c, after

Sars; d, e, after Calman).

1st-4th pairs of legs united by a median projecting lamina, flattened, biramous,

both rami 1-jointed, outer rami spiniform, inner rami with 2 spine-setae;

5th legs broadly lamellate, acting as ovitectrices.

§—Cyclopiform. Abdomen 5-segmented. Antenna 1 8-jointed. Antenna

2 3-jointed, with apical claw. tst-4th legs natatory, biramous, both rami

usually 3-jointed.

Only found in compound Ascidians (Krikobranchiata).

_Enterocola fulgens van Bened.

1860. Van Beneden, loc. cit.

1909. Chatton & Brément, loc. cit., p. 227.

1922. Schellenberg, loc. cit., p. 287.

©—Thorax without alate expansions (except the modified 5th pair of legs).

Caudal rami 14-2 times as long as wide. 1:5 mm.

Locality. ‘Table Bay, in Amaroucium erythraeum (Schellenberg).

Distribution. Europe, Mediterranean.

Remarks. In fig. 7 is given an illustration of an allied species, E. bilamellatus

Sars 1921, as an example of the genus.

242 ANNALS OF THE SOUTH AFRICAN MUSEUM

Gen. <anclopus Calman

1908. Calman, Mar. Invest. S. Afr., v (Tr. S. Afr. Phil. Soc., xvii), p. 178.

1913. Gravier, Deux. Exp. antarct. Frang., p. 68.

1922. Schellenberg, loc. cit., p. 295 (in list of genera, etc.).

1932. Wilson, loc. cit., p. 601 (in key to genera).

—Head distinct. Thorax cylindrical, 5-segmented. Abdomen obscurely

segmented. Caudal rami short, with 4 short spiniform points directed out-

wards. Antenna 1 short but conspicuous, I-jointed. Antenna 2 smaller than

antenna I. Ist-4th pairs of legs biramous, outer ramus ending in a sickle-

shaped claw, inner ramus subcircular with 4 marginal papillae; 5th legs large,

ovate.

g—Cyclopiform. Abdomen 5-segmented. Antenna 1 (in adult) 7-jointed.

Antenna 2 3-jointed, with 2 or 3 apical claws. tst-4th pairs of legs natatory,

biramous, rami 3-jointed; 5th leg consisting of 2 small papillae, each bearing

a seta.

In the stomachs of the zooids of Cephalodiscus (Pterobranchiata).

Kanclopus cephalodisc:t Calman

Bien 7d ae

1908. Calman, loc. cit., p. 178, pls. 18, 19.

1910. Stebbing, Ann. S. Afr. Mus., vi, p. 550.

Length 2 0-62 mm., $ 0-6 mm. ;

Locality. Agulhas Bank in Cephalodiscus gilchristt (Calman).

Suborder CALIGOIDA

1913. Scott, T. & A., Brit. Parasit. Copepoda, i, p. 43.

1932. Wilson, Bull. U.S. Nat. Mus., no. 158, pp. 397 and 604 (key to

genera).

Body segmented or not. Fourth thoracic segment usually movably articu-

lated with 3rd segment, but firmly attached to 5th. In some fixed parasites the

movable articulation is lost in the 9, the whole body becoming rigid. First, or

ist and 2nd, or Ist to 3rd thoracic segments often fused with head, forming a

shield-like carapace. Antenna 1 reduced to 1 or 2 joints. Antenna 2 pre-

hensile. A sternal fork (furca) sometimes present between bases of maxillipeds.

Usually 4 pairs of legs, but one or two pairs may be wanting. Fifth pair

rudimentary or obsolete. Two lateral ovisacs, usually filiform or cylindrical,

uniseriate.

Parasitic on aquatic mammals and fishes, rarely on Invertebrates; mostly

retaining the power of swimming and sometimes (especially males) captured

in plankton.

Remarks. Scott excludes the family Lernaeidae from the Caligoida, including it

with the Chondracanthidae and Lernaeopodidae in a suborder (tribe) Lernaeozda.

Wilson, however, includes the Lernaezdae in the Caligoida, and the Chondracan-

thidae and Lernaeopodidae in the suborder Lernaeopodoida, not recognizing

Lernaeoida.

The sternal fork (intercoxal plate: Lang, 1951, Ark. Zool., and ser., I, 6,

Pp. 499) is not found in all Caligorda, but it is not found in any other suborders.

SOUTH AFRICAN PARASITIC COPEPODA 243

Wilson’s families are here adopted, but without family diagnoses. The key

to the South African genera utilizes some of the characters in Wilson’s key,

but it is admittedly unsatisfactory.

Dysgamus St. & Liitk., 1861 occurs on the Whale Shark (Rhyneodon typicus)

(Wilson, 1944, Proc. U.S. Nat. Mus., xciv, pp. 531, 533), a shark which is

known to occur in South African waters. Heegaard (1943, Ark. Zool., xxxiv,

A, A. 18, p. 24, figs. 65-75) records specimens from a ‘Cape-line ship’ which he

would have considered to be the free-swimming stage of some Caligid, had not

Bassett-Smith (1899, Proc. Zool. Soc. Lond., p. 460) recorded ovigerous 99.

Wilson (1907, Proc. U.S. Nat. Mus., xxxi, p. 712) accepted the genus pro-

visionally pending the discovery of the ovigerous 9.

Key to the South African genera (99 only)

A shield-like cephalothorax. Body more or less depressed.

A. Head and first 3 thoracic segments fused, 4th segment free.

1. 4th segment without dorsal plates.

a. With frontal lunules. Caligus

b. Without frontal lunules.

i. 4th leg uniramous. Lepeophthetrus

ii. 4th leg biramous. [Dysgamus ]

2. 4th segment with a pair of dorsal plates Alebion

B. Head and first 2 thoracic segments fused, 3rd and 4th free, without

dorsal plates. Trebius

C. Head and first thoracic segment fused, 2nd-4th free (2nd and 3rd

sometimes fused); one or more segments with dorsal plates.

I. 3 pairs of dorsal plates (segments 2-4).

a. A 6th thoracic segment concealing the abdomen. Pandarus

b. No 6th segment. Perissopus

2. 2 pairs of dorsal plates (fused 2-3, and 4).

a. Ovisacs visible, elongate. Achtheinus

b. Ovisacs concealed, coiled. Cecrops

3. One pair of dorsal plates (segment 4).

a. Legs not foliaceous.

i. Ovisacs visible.

a. Plates of 4th segment and those of 5th with smooth

margins.

* end and grd segments not fused.

+ A 6th thoracic segment with a pair of

small dorsal plates. Dinemoura

tt No 6th thoracic segment. Echthrogaleus

** ond and 3rd segments fused, with lateral lobes Nesippus

B. Plates of 4th segment and those of 5th with serrated

margins. Philorthragoriscus

ii. Ovisacs concealed. Orthagoriscicola

b. Legs foliaceous, forming with the dorsal plates of 4th seg-

ment a skirt surrounding the genital segment and abdomen. Anthosoma

4. 2nd-4th segments fused, with a single dorsal plate covering

genital segment and abdomen. Lernanthropus

II. Body without shield-like carapace, more or less segmented and cylindri-

cal. Head without horns or processes for attachment.

A. Body distinctly segmented. 4 pairs of biramous legs. Nemesis

B. Body divided into head, neck, and unsegmented trunk. 2 pairs of

biramous legs. Hatschekia

244. ANNALS OF THE SOUTH AFRICAN MUSEUM

III. Body without carapace, cylindrical but not segmented. Head and neck

distinct; head with horns or processes for attachment.

A. Abdomen without processes.

1. Head with horn-like processes. Lernaea

2. Head with branched processes. Cardiodectes

B. Abdomen with plumose processes. Pennella

Fam. CALIGIDAE

1905. Wilson, Proc. U.S. Nat. Mus., xxviii, p. 532 (Caliginae).

1934. Gurney, Proc. Zool. Soc. Lond., 1934, 2, pp. 177 sqq., figs. (develop-

ment).

Gen. Caligus Miller

1785. Miiller, O. F., Entomostraca, p. 128.

1899. Bassett-Smith, Proc. Zool. Soc. Lond., p. 446 (list of species).

1905. Wilson, loc. cit., p. 555 (key to species).

1908. id., ibid., xxxill, pp. 594. sqq.

1913. Scott, T. & A., Brit. Parasit. Copep., p. 44.

1924. Brian, ‘Parasit. Mauritan.’ (Bull. Com. Etud. Hist. Sci. Afr. occid.

Fir), [email protected],) pp. 13 Sada:

1932. Wilson, Bull. U.S. Nat. Mus., no. 158, pp. 397, and 605 (in key).

1935. Brian, Ann. Mus. Civ. Genoa, lvii, p. 152 (Mediterranean species).

1936. Yamaguti, Parasit. Copep. Japan, pt. 2, Caligoida 1, pp. 2-9.

1937. Wilson, Hancock Pacific Exp., I1, 4, p. 24.

1939. Yamaguti, Parasit. Copep. Japan, pt. 5, Caligoida 3 (vol. Fubil. Prof.

Yoshida, I1), pp. 445-9.

1951. Pearse, Proc. U.S. Nat. Mus., ci, p. 344.

1954. Yamaguti, Publ. Seto Mar. Biol. Lab. Ill, 3, pp. 379 sqq.

Carapace large, oval or subcircular. Genital segment simple, without plates

or processes. Abdomen 1-4-segmented; caudal rami usually moderate or

small. Frontal lunules present. Maxilla 2 simple, spiniform. Sternal fork

present. 1st and 4th legs uniramous, 2nd and 3rd biramous; 4th leg 3-jointed,

rarely 4-jointed. Rudiments of 5th and 6th legs sometimes present. Ovisacs

elongate, uniseriate.

On a variety of hosts, mostly Teleost fishes.

Key to South African species

I. 4th leg 4-jointed.

A. 1st abdominal segment 4 times the length of and. pelamydis

B. 1st abdominal segment shorter than 2nd. [elongatus |

II. 4th leg 3-jointed.

A. Abdomen @ 4, 3 2-segmented. coryphaenae

B. Abdomen 1-segmented, or more or less distinctly 2-segmented.

1. Abdomen long, in 2 about as long as, in ¢ longer than, genital seg-

ment.

a. Caudal rami long, in 2 about half length of, in § longer than

abdomen. lalandei

b. Caudal rami short.

i. Caudal rami short. Abdomen distinctly 2-segmented, a little

shorter than genital segment. cossackit

ii. Caudal rami very short. Abdomen 1-segmented, a little

longer than genital segment. arit

SOUTH AFRICAN PARASITIC COPEPODA 245

2. Abdomen short, about half length, or less, of genital segment.

a. Caudal rami much longer than wide. engraulidis

b. Caudal rami about as broad as long. tetrodontis

C. elongatus Heegaard (1943, Ark. Zool., xxxiv, 4, A. 18, p. 11, figs. 21-31)

was described from specimens from a ‘Cape-line ship’, but not necessarily from

South African waters. No host was recorded.

eo =z

Fig. 8. Caligus coryphaenae St. & L. a, 2; b, abdomen 9; c, furca; d, 4th leg. Caligus cossackii

B-S. e, 2; f, furca (copies: a, from Brian, after St. & L.; 5b-d, after Brian; e-f, after Heegaard).

Caligus pelamydis Kroyer

1863. Kroyer, Naturh. Tidsskr., ser. 3, ii, p. 124 (fide Stebbing), p. 50

(fide Wilson), pl. 4, figs. 4 a-g.

1905. Wilson, loc. cit., p. 594, pl. 13, figs. 154-61, pl. 14, fig. 161 a.

1910. Brady, Deutsch Stidpol. Exp., xi (zool. iii), p. 589, fig. 69 (9).

1910. Stebbing, Ann. S. Afr. Mus., vi, p. 558.

1913. Scott, 1. & A., loc. cit., p. 57, pl. 7, figs. 2, 3 (@), pl. 9, figs. 1-5

(Q), pl. 71, fig. 14 (d).

1932. Wilson, loc. cit., p. 406, fig. 254.

1935. Brian, loc. cit., p. 183, figs. xii (after Wilson) and xiii.

—Carapace oval, less than half total length. 4th segment narrow, short.

Genital segment a little longer than wide, oblong, narrowing anteriorly,

truncate posteriorly. Abdomen as long as carapace, and slightly longer than

246 ANNALS OF THE SOUTH AFRICAN MUSEUM

genital segment, obscurely 2-segmented, Ist segment nearly 4 times length of

2nd; caudal rami shorter than 2nd segment, with plumose setae. Furca with

base much narrower than width across prongs, these short, broad, curved.

4th leg 4-jointed, last 3 joints with 1, 1, and 3 spines respectively, these 5 spines

set closely together like a comb. Ovisacs rather stout, as long as carapace plus

genital segment, about 30 eggs in each.

3°3 mm. Yellowish-white.

Locality. Simons Bay (Brady).

Distribution. Danish and British seas, Mediterranean, east coast of N.

America. On Pelamys sarda and Gymnosarda pelamys.

Caligus coryphaenae St. & L.

Fig. 8 a—d.

1861. Steenstrup & Liitken, A. Dansk. Videns. Sels. Skr. (5), v, p. 360,

pl. 4, fig. 7.

1894. Scott, Trans. Linn. Soc. Lond. (2), vi, p. 130, pl. 14, fig. 20 (ben-

goensis) (juv. 3).

1905. Wilson, loc. cit., p. 555 (bengoensis), and pp. 556, 559 (in key).

TOQS5 ide PAV 400) sever On Dana:

1995. Brian.) loc cit. pp. 202, dose xi KK

1936. Yamaguti, loc. cit., p. 5, pl. 4, figs. 40-54.

1949. Heegaard, Vid. Medd. Dansk. nat. For., cxi, p. 240, figs. 6-10.

@—Carapace oval, less than half total length. Genital segment longer than

wide, somewhat wider posteriorly (deeply lobed: Steenstrup & Liitken; not

so deeply: Brian’s and Yamaguti’s figures). Abdomen about as long as genital

segment, 4-segmented (junctions somewhat obscure: Brian’s figures), Ist

segment as long as the others together; caudal rami short, with plumose setae.

Furca narrow, with narrow prongs. 4th leg 3-jointed, 1 apical spine on and

joint, 4 spines on 3rd (Brian, Yamaguti). Ovisacs slender, elongate.

g—Carapace oval, more than half total length. Genital segment quadrate.

Abdomen a little shorter than genital segment, 2-segmented; caudal rami

short (Steenstrup & Liitken’s figures in Brian). Rudiments of 5th leg visible.

Length: 2 6-7 mm., ¢ 2-3 mm.

Locality. Delagoa Bay, on Squalus acanthias (Wilson).

Distribution. Atlantic and Mediterranean, on Coryphaena hippurus; Loanda;

Pacific.

Caligus lalande: Brnrd.

Fig. 9 a-c.

1948. Barnard, Ann. Mag. Nat. Hist. (xii), 1, p. 243, fig. 2.

—Carapace oval, less than half total length. 4th segment not laterally

prominent. Genital segment oval, longer than wide, posterior corners rounded,

slightly produced. Abdomen nearly as long as genital segment, obscurely

2-segmented, Ist segment slightly narrower than 2nd which is about 24 times

as long as Ist, straight-sided; caudal rami about 2 length of abdomen, elongate-

lanceolate, apical setae short, nonplumose. Furca widening distally, with oval

sinus between broad apically truncate prongs. 4th leg 3-jointed, one spine on

apex of 2nd joint, apex of 3rd joint with 1 long and 2 short spines each with

serrated scale at base. Ovisacs about 2 total length.

SOUTH AFRICAN PARASITIC COPEPODA 247

g§—Genital segment quadrate, longer than wide, hind corners quadrate.

Abdomen about twice as long as genital segment, obscurely 2-segmented, 2nd

segment about 2} times as long as Ist, sides gently converging; caudal rami

very elongate, about 3 total length, filiform, with 2 long and 1 short apical

spine-setae, nonplumose.

Fig. 9. Caligus lalandei Brnrd. a, 2; 6, 3; ¢, furca. Caligus tetrodontis Brnrd. d, genital segment

and abdomen 9; e, genital segment and abdomen @; /, furca.

@ 10°5 mm. (g mm. to end of abdomen); ¢ 16 mm. (g mm. to end of

abdomen), carapace 4 mm., abdomen 3 mm., caudal rami 7 mm.

Locality. Kalk Bay (Cape Peninsula), on Yellow-tail (Seriola lalandei), 4 99,

1.) C. Li. Biden.

Remarks. Noteworthy on account of the length of the caudal rami in both

sexes, but especially in the g. The @ differs from that of serzolae Yamaguti 1936

in the shape of the furca and the considerably longer caudal rami: the ¢ of

seriolae is unknown.

248 ANNALS OF THE SOUTH AFRICAN MUSEUM

Caligus cossackiu B-S.

Fig. 8 e, f.

1898. Bassett-Smith, Ann. Mag. Nat. Hist. (vii), 2, p. 85, pl. 4, figs. 3,

3 af (€).

1943. Heegaard, Ark. Zool., xxxiv, A. 18, p. 5, fig. 12 A-C (9).

@—Carapace subcircular, shorter than half total length, frontal plate con-

cave. 4th segment not laterally prominent. Genital segment longer than broad,

oblong, without ‘neck’, hind corners rounded, without processes or rudiments

of 5th legs. Abdomen distinctly 2-segmented, 2nd segment a little shorter

than Ist; caudal rami short, well developed. Lunules very prominent. Furca

with divergent prongs, slightly incurved apically. 4th leg 3-jointed. Ovisacs

moderately stout.

g6—More elongate. Genital segment oval. Caudal rami longer and more

profusely setose (Bassett-Smith).

295 mm., d 3 mm.

Locality. Madagascar, in gill cavity of Sparus berda (Heegaard).

Distribution. Persian Gulf and Ceylon, on Chrysophrys sarba (Bassett-Smith).

Remarks. As both Acanthopagrus berda and Austrosparus sarba are found in

South African waters, it is reasonable to expect that this parasite will eventually

be included in the South African fauna-list.

There seems to be a strong similarity between this species and affinis Heller

(see figure in Brian, Rev. Zool. Bot. Afric., xxxil, p. 178, fig. 1, 1939. Mouth of

Congo River, on Sphyraena).

Caligus aru B-S.

Fig. 10 a, b.

1898. Bassett-Smith, Ann. Mag. Nat. Hist. (vii), 2, p. 82, pl. 4, figs. 1,

1 a—d (9).

1948. Barnard, ibid. (xii), 1, p. 244.

—Carapace subcircular, considerably shorter than half total length. 4th

segment not laterally prominent. Genital segment slightly longer than broad,

oblong, with a ‘neck’ anteriorly of about the same width as 4th segment, hind

corners rounded, slightly produced, without processes or rudiments of 5th

legs. Abdomen t-segmented (Bassett-Smith: 2-segmented, 2nd segment

short, as long as wide), as long as genital segment (without the ‘neck’), slightly

tapering; caudal rami distinct, well developed, but very short. Lunules very

prominent. Furca with moderately divergent prongs, slightly incurved apically.

Ath leg 3-jointed. Ovisacs stout, not very much longer than abdomen.

4°5 mm.

Locality. Chinde, mouth of Zambezi River, on palate of Arius dussumieri,

3 OO 1K EB so 12).

Distribution. Ceylon, on Arius acutirostris.

Remarks. There is little doubt that these specimens should be regarded as —

conspecific with the Ceylon specimens in spite of certain differences. The

abdomen shows no trace of a division into two segments near the hind end,

and the caudal rami are quite well developed although small. The 4th leg is

composed (as usual) of 3 joints, there being no trace of a division near the apex

SOUTH AFRICAN PARASITIC COPEPODA 249

(as shown in Bassett-Smith’s figure). These differences may be due to the

hosts being different species, though of the same genus, but they scarcely

warrant the institution of a separate species.

Caligus engraulidis Brnrd.

Fig. 10 ¢, d.

1948. Barnard, Ann. Mag. Nat. Hist. (xii), 1, p. 244, fig. 3.

Q—Carapace oval, longer than rest of body. 4th segment small, not laterally

prominent. Genital segment wider than long, subquadrangular, hind corners

\ / sy \ ‘

rey ye

t Hh \ \

t i] 5 +

/ Cc

Fig. 10. Caligus arii B-S. a, genital segment and abdomen @; 3, furca.

Caligus engraulidis Brnrd. c, genital segment and abdomen Q, apical spines

of 4th leg further enlarged; d, furca.

peta — ae

rounded, only slightly produced, without processes or rudiments of 5th legs.

Abdomen a little less than half length of genital segment, oblong; caudal rami

a little shorter than abdomen, with 3 long plumose setae. Furca with short,

rather stout, and narrowly separated prongs. 4th leg with 1 long and 1 short

spine on apex of 3rd joint, both serrate on both margins. Ovisacs stout, about

as long as body.

3°5 mm.

Locality. Zwartkops River, Algoa Bay, on Engraulis (Anchoviella) holodon

250 ANNALS OF THE SOUTH AFRICAN MUSEUM

Caligus tetrodontis Brnrd.

Fig. 9 d-—f.

1948. Barnard, Ann. Mag. Nat. Hist. (xii), 1, p. 245, fig. 4.

@—Carapace about half total length. Genital segment cordate, a little wider

than long, hind corners rounded, not produced, 5th legs represented by 2-3

fine setules (better visible in immature 2). Abdomen a little less than half

length of genital segment, obscurely 2-segmented; caudal rami nearly as long

as broad, with plumose setae. Furca with narrow, divergent prongs. 4th leg

with 1 spine on apex of 2nd joint, and 3 spine-setae of varying lengths on apex

of 3rd joint, with 2 short biserrate spines (scales) at their base. Ovisacs stout.

g—Genital segment longer than broad, hind corners obliquely bevelled off,

showing tufts of setae (3 and 2 respectively) representing 5th and 6th legs.

Abdomen distinctly 2-segmented, 1st segment shorter but slightly wider than

2nd, with convex lateral margins, 2nd about as long as wide, with straight sides.

4-5 mm.

Locality. Port Elizabeth, on Tetrodon hypselogeneion. 2 ovig. 99, 2 juv. 929,

4 33d.

Remarks. Closely allied to parvus Bassett-Smith from Bombay, also parasitic

on a species of Tetrodon. ‘The present specimens, however, are larger (parvus:

2 3:4, 3 2:3 mm.), the abdomen and caudal rami are longer, and the prongs of

the furca longer. The abdomen of parvus is apparently only 1-segmented.

Gen. Lepeophtheirus Nordmann

1832. Nordmann, Mekr. Beitr. Wirbellos. Thiere, ii, p. 30.

1899. Bassett-Smith, Proc. Zool. Soc. Lond., p. 453 (list of species).

1905. Wilson, Proc. U.S. Nat. Mus., xxviii, p. 615 (key to species).

1908. id., ibid., xxxili, pp. 600 sqq.

1908. id., ibid., xxxv, pp. 439 sqq.

1913. Scott, T. & A., Brit. Parasit. Copep., p. 64.

1932. Wilson, Bull. U.S. Nat. Mus., no. 158, pp. 409, and 606 (in key).

1933. Gurney, Brit. Freshwater Copep., iii, p. 324 (development)

1936. Yamaguti, Parasit. Copep. fapan, pt. 2, Caligoida 1, pp. 10-15.

1939. id., pt. 5, Caligoida 3, pp. 449-53.

1944. Wilson, Proc. U.S. Nat. Mus., xciv, pp. 533 sqq-

Carapace large, broadly oval or subcircular. Fourth segment free, without

dorsal plates. Genital segment simple, without plates or processes. Abdomen

I-, 2- (or 4-) segmented; caudal ramimoderate. No frontal lunules. Maxilla

2 small, bifurcate. Sternal fork present. Ist and 4th legs uniramous, 2nd and

grd biramous; 4th leg 4-jointed, sometimes 3-jointed. Rudiments of 5th and

6th legs sometimes present. Ovisacs elongate, uniseriate.

Closely resembling Caligus, but without the frontal lunules.

Key to the South African species

1. Abdomen (2) well developed.

a. Furca with narrow acute prongs.

i. Carapace subcircular, considerably more than half total length. 4th leg

4-jointed. insignis

li. Carapace oval, very little more than half total length. 4th leg 3-jointed. sp.

6b. Furca with broad, blunt prongs. lichiae

SOUTH AFRICAN PARASITIC COPEPODA 251

2. Abdomen very short, concealed under genital segment.

a. Norudiments of 5th legs on genital segment (9). brachyurus

b. Rudiments of 5th legs on genital segment (). plotosi

Lepeophtheirus insignis Wilson

Big itr

1908. Wilson, loc. cit., p. 444, pls. 70, 71.

1948. Barnard, Ann. Mag. Nat. Hist. (xii), 1, p. 247.

Q—Carapace subcircular, about as wide as long, considerably more than

half total length. Eyes minute. 4th segment rather prominent laterally, not as

wide as genital segment, which is ovoid, longer than broad in adult. Abdomen

of one segment, oblong, shorter than genital segment in adult. Furca with

simple rather widely diverging prongs.

Fig. 11. Lepeophtheirus insignis Wilson. a, immature 9, 6th leg further

enlarged; 6, g; c. furca.

g—Similar to 9, but carapace relatively slightly larger. Genital segment

ovoid, with rudiments of 5th and 6th legs. Abdomen 2-segmented.

@ 11°75 mm., carapace 6-5 mm., ¢ 6-6 mm., carapace 4 mm. (Wilson).

Yellowish, chitinous ribs and thickenings on carapace purple, internal oviducts

orange, ovisacs yellow (Wilson). White marbled with reddish-grey (K.H.B.).

Locality. ‘Table Bay, on Sunfish (Mola mola). 1 immature 9, 1¢ (K.H.B.).

Distribution. Coast of Southern California, on Mola mola.

Remarks. The 3 (carapace 4 X 4 mm.) corresponds closely with Wilson’s

description. The 9 (carapace 5 X 5 mm.) clearly belongs to Wilson’s species

but is in a stage intermediate between the young 9 figured on pl. 71, fig. 51,

and the adult figured on pl. 70, fig. 37; the genital segment is more ovoid

than in fig. 51 but has not reached its full length as shown in fig. 37. The 6th

leg is narrower than in fig. 51; there is no trace of a rudimentary 5th leg.

(Comparison of pl. xxii, fig. 273, and pl. xxv, fig. 311, in Wilson’s 1905 paper

252 ANNALS OF THE SOUTH AFRICAN MUSEUM

with pl. 70, fig. 44, in the 1908 paper leaves a doubt whether these trispinose

appendages represent the 5th or the 6th legs; in the former paper they are

called the 5th, in the latter the 6th.)

Lepeophtheirus sp.

Two ovigerous 99 of a species very close to L. longispinosus Wilson (1908,

Proc. U.S. Nat. Mus., xxxiii, p. 604, pl. 52) were taken by Mr. C. L. Biden from

P| b A j . f 3

Fig. 12. Lepeophtheirus lichiae Brnrd. a,?; 6, furca. Lepeophtheirus brachyurus Heller. c, 2; d, furca.

Lepeophtheirus plotost Brnrd. e, genital segment and abdomen 9; ff, furca; g, 5th leg.

a Sand Shark (Rhinobatus) at Durban 1953. Wilson’s species was taken from a

Hammerhead Shark off the coast of North Carolina.

The present specimens agree with Wilson’s species in having the 4th leg

3-jointed, but differ in the shape of the furca which consists of 2 small acute

subparallel prongs.

5 mm., ovisacs 3 mm. Whitish.

Lepeophthetrus lichiae Brnrd.

Big. 1 21as 0:

1948. Barnard, Ann. Mag. Nat. Hist. (xii), 1, p. 247, fig. 5.

@—Carapace oval, about as broad as long, very slightly more than half

total length. Eyes minute. 4th segment laterally prominent, almost as wide as

SOUTH AFRICAN PARASITIC COPEPODA 253

genital segment, which is slightly broader than its median length; posterior

corners of genital segment lobately produced, with a strong straight spine

arising from ventral surface and projecting backwards, ? length of segment.

Abdomen a little longer than half median width of genital segment,

2-segmented. Furca with broad, apically rounded prongs.

7°5 mm.

Locality. Natal, on Lichia amia. Ovig. and nonovig, 99.

Remarks. ‘The spines on the genital segment are reminiscent of Gloiopotes.

Lepeophtheirus brachyurus Heller

Big. 12 ¢..4.

1865. Heller, ‘Novara’, Crustac., p. 185, pl. 16, fig. 4.

1906. Wilson in Herdman’s Ceylon Pearl Oyster Fish Rep., v., p. 190. pl. 1,

figs. I-10.

1948. Barnard, loc. cit., p. 248.

@—Carapace as broad as long, slightly longer than half total length, hind

margin strongly convex and extending farther backwards than lateral lobes.

4th segment less than half width of genital segment, which is about as long as

wide, subquadrangular with anterior angles bevelled off, hind margin very

slightly convex medianly; no rudiments of 5th or 6th legs. Abdomen extremely

short, inserted ventrally, scarcely visible dorsally. Furca with simple, diverging

prongs.

5 mm.

Locality. Durban, on Tetrodon hypselogeneion, 1 .

Distribution. Java, on Tetrodon calamariae; Ceylon, on T. séellatus.

Remarks. This specimen in so far as a comparison is possible (4th legs missing)

is very like Heller’s species. His figure shows the genital segment subcircular,

not subquadrate, and the 4th legs as 2-jointed.

Lepeophtheirus plotost Brnrd.

Fig. 12 e-g.

1948. Barnard, loc. cit., p. 248, fig. 6.

@—Carapace about as broad as long, slightly longer than half total length.

4th segment well developed. Genital segment oval, longer than wide, hind

corners rounded, not at all produced, with distinct rudimentary 5th legs.

Abdomen very short, broader than long, attached to dorsal surface of genital

segment; caudal rami shorter than abdomen, broader than long. Furca with

narrow prongs, sinus wide, U-shaped. Ovisacs stout, as long as genital segment.

3°5 mm.

Locality. East London, on gills of Plotosus anguillaris.

Remarks. ‘The three species listed by Bassett-Smith as parasitic on Bagrid

fishes: bagri Dana, quadratus Kr., and longipalpus B-S. all have a 2-segmented

abdomen. I have not seen descriptions of the first two, but the third has an

elongate abdomen.

254 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fam. EURYPHORIDAE

1932. Wilson, Bull. U.S. Nat. Mus., no. 158, p. 415.

Gen. Alebion Kroyer

1863. Kroyer, Waturh. Tidsskr. (3), 2, p. 239.

1907. Wilson, Proc. U.S. Nat. Mus., xxxi, p. 702 (key to species).

1932. id., loc. cit., pp. 418, 606.

1951. Gnanamuthu, Ann. Mag. Nat. Hist. (xii), 4, p. 1237.

1953. Capart, Bull. Inst. frang. Afr. Noire, xv, 2, p. 655.

Sexes nearly similar. Head and first 3 thoracic segments fused into a shield-

like carapace. Fourth segment free, with a pair of dorsal plates in 9, but these

much reduced or absent in g. Genital segment with hind corners usually

produced in 9, rounded in g. Abdomen 2-segmented, hind corners of basal

segment in 2 produced. Caudal rami lamelliform. First 3 pairs of legs

biramous, outer rami with large horny claws, rami of 1st leg 2-jointed, of 2nd

and 3rd 3-jointed, 4th legs rudimentary. No frontal lunules. No furca.

On sharks.

Alebion carchariae Kroyer

Fig. 13 a.

1863. Kréyer, loc. cit., p. 165, pl. 12 (page and plate quoted from Wilson,

1932). |

1907. Wilson, loc. cit., xxxi, p. 704 (in key).

T9392: 1d:, loc. cit., p. 422) 110+ 260:

1953- Capart, loc. cit., p. 655, fig. 3.

Q—Carapace |subcircular, about as wide as long, median lobe short,

truncate, with 2-4 denticles at each postero-lateral corner. Dorsal plates of

Ath segment subtriangular, separated by a triangular notch. Genital segment

widest behind middle, lateral margins distally fringed with spines; posterior

processes flattened, upper margins fringed with spines; centre of segment

gibbous, hind margin excavate, without spines or denticles.

g mm. (Wilson: 14 mm.).

Localities. Ifafa, Natal, on a shark (Dept. Zool. Rhodes Univ. Grahamstown,

1948); Durban, on Carcharinus (C. L. Biden 1951).

Distribution. N. Atlantic, Gape Verde Islands, Senegal.

Remarks. The present specimens agree with carchariae except the processes of

the genital segment are spinose along the upper margin as in crassus.

Fam. TREBIIDAE

1907. Wilson, Proc. U.S. Nat. Mus., xxxi, p. 669 (Trebinae [sic]).

1932. id., Bull. U.S. Nat. Mus., no. 158, p. 413 (Trebidae [sic]).

Gen. Trebius Kroyer

1838. Kroyer, Naturh. Tidsskr., ii, p. 30.

1899. Bassett-Smith, Proc. Zool. Soc. Lond., p. 461.

1907. Wilson, loc. cit., p. 672 (key to the 3 species known).

1932. id., loc. cit., p. 413, and 604 (in key).

1953. Capart, Bull. Inst. frang. Afr. Noire, xv, p. 665.

SOUTH AFRICAN PARASITIC COPEPODA 255

Sexes similar. Head and first 2 thoracic segments fused into a shield-like

carapace. 3rd and 4th segments free, without dorsal plates. Abdomen elon-

gate, 1-3-segmented. No frontal lunules. Furca present. All 4 pairs of legs

biramous, rami of rst leg 2-jointed, of the other legs 3-jointed (or inner ramus

of 4th leg 2-jointed). Rudiments of 5th and 6th legs on genital segment in J.

No adhesion pads. Ovisacs elongate, uniseriate.

On skates (rays) and Dogfish.

A SANS:

Fig. 13. Alebion carchariae Kroyer. a,Q. Trebius caudatus Kroyer. 6,9; ¢, maxilla1; d. maxilla 2;

e, furca; f, 1st leg.

Trebtus caudatus Kroyer

Fig. 13 b-f.

1838. Krdéyer, loc. cit., p. 30, pl. 1, fig. 4 (9).

1863. id., Naturh. Tidsskr. (3), ii, p. 149, pl. 10, fig. 1 (¢).

1907. Wilson, loc. cit., p. 681, pl. 15, figs. 11-13, pl 16, figs. 14-22.

1913. au T. & A., Brit. Paras. Copep., p. 81, pl. 22, figs. 1-2, pl. 54,

gs. I-II.

1934. Gurney, Proc. Zool. Soc. Lond., 1934, 2, p. 192, figs. 18-20 (develop-

ment).

256 ANNALS OF THE SOUTH AFRICAN MUSEUM

1941. Sproston & Hartley, 7. Mar. Biol. Assoc. Plymouth, xxv, p. 393

(bionomics).

1948. Barnard, Ann. Mag. Nat. Hist. (xii), 1, p. 249.

@—Carapace subcircular, well arched dorsally. Eyes obsolete in adult.

Genital segment oval, hind corners rounded, each with 3-4 denticles. Abdomen

subequal to carapace, longer than genital segment, fusiform, more or less

distinctly 3-segmented. Furca with narrow divergent prongs. Maxilla 2

apically bifurcate.

g—Genital segment barrel-shaped. Abdomen a little longer than genital

segment, 2-segmented.

Iomm.

Locality. ‘Table Bay, on skate (Raia marginaia). 13 992 in poor condition

(S. Afr. Mus.).

Distribution. British Seas, on skates and dogfishes.

Fam. PANDARIDAE

1907. Wilson, Proc. U.S. Nat. Mus., xxxiii, pp. 325, 345 (Pandarinae).

Gen. Pandarus Leach

1816. Leach, Encycl. Brit., ed. 5, Suppl., p. 405 (9).

1819. id., Dict. Sci. Nat., xiv, p. 535 (do) (Nogaus).

1840. Milne Edwards, Suite a Buffon (Crust.), iii, p. 460 (Nogagus pro

Nogaus).

1907. Wilson, loc. cit., pp. 346, 387 (key to species).

1913. Scott, T. & A., loc. cit., p. 94.

1932. Wilson, Bull. U.S. Nat. Mus., no. 158, pp. 432, and (in key) 607,

600.

Sexes dissimilar. Head and rst thoracic segment fused. Carapace smooth,

without grooves, postero-lateral lobes moderate, hind margin excavate, den-

tate. Free thoracic segments (3) each with a pair of plates, those on segment 2

dorso-lateral, the others dorsal. Genital segment large, posterior corners more

or less lobately prolonged. Segment 6 represented by a median plate con-

cealing the abdomen. Abdomen short, broad, 2-segmented, covered ventrally

by a short broad median plate. Caudal rami attached laterally to the ventral

median plate, flattened, subtriangular or more or less conical and acicular.

Four pairs of adhesion pads: at bases of 1st and 2nd antennae, between bases

of 1st maxillipeds and opposite 1st legs. Antenna 1 2-jointed; antenna 2 with

apical claw. Maxilliped 2 swollen, with a pair of knobs. Four pairs of

biramous legs, spinose and indistinctly jointed legs. Ovisacs elongate filiform,

uniseriate.

g—Carapace broadly oval, posterior lobes prominent, posterior margin

straight, usually with a pair of accessory lobes, lateral grooves distinct. Free

thoracic segments without dorsal plates, but the first with lateral lobes.

Genital and 6th segments fused, with 2 pairs of rudimentary legs. Abdomen

2-segmented; caudal rami laminate, with 4 plumose setae. Maxilliped 2 with

apical claw or corrugated knobs. All 4 pairs of legs biramous, both rami 2-

jointed, with long plumose setae.

On various sharks and dogfishes,

SOUTH AFRICAN PARASITIC COPEPODA P25) 7

Key to the South African species (29)

Dorso-lateral plates of 2nd segment not extending beyond ends of plates on 3rd

segment.

a. Caudal rami not very prominent. bicolor

6b. Caudal rami prominent. 3rd and 4th segments each with a pair of dorsal

spines. armatus

Dorso-lateral plates of 2nd segment extending beyond ends of plates on 3rd seg-

ment.

a. Dorsal plates of 3rd segment fused. cranchit

b. Dorsal plates of 3rd segment separate (in adult). smithit

Fig. 14. Pandarus bicolor Leach. a, 2; 6, 3, right corner of carapace cut away. Pandarus armatus

Heller. c, 2 (copy after Heller).

Pandarus bicolor Leach

Fig. 14 4, D.

Foto ieeach loc ycit...p. 405,:pl),20,, figs. 1, 2/2).

1900. Scott, T., Rep. Fish. Board Scotl., xviii, pt. 3, p. 157, pl. 6, figs. 33-8

(Nogaus 3).

1907. Wilson, loc. cit., p. 400, pl. 27 (9).

HOMO we OCOL leiec UAE OCs Cll.) O05.) pl. 21s) fig. 2) \ple22. figs. 5.) ©,

pl. 26, figs. 4-19, pl. 58, figs. 1-8 (2 @).

1O322)) VVilson, loc. cit-,p: 430i. 274 (2 3).

1948. Barnard, Ann. Mag. Nat. Hist. (xii), 1, p. 249.

©—Carapace widest posteriorly, hind margin dentate. Dorso-lateral plates

of and segment not reaching beyond plates of 3rd segment; the latter with

258 ANNALS OF THE SOUTH AFRICAN MUSEUM

narrow median sinus, more or less expanded anteriorly; plates of 4th segment

overlapping genital segment for about half its length, with angular sinus.

Genital segment with broadly rounded posterior lobes; 6th segment plate at

least half width of genital segment, subcircular. Caudal rami short, laminate,

triangular, thickened on anterior margin.

g—Carapace as wide as long, no accessory lobes on hind margin. 2nd

segment with lateral lobes (in adult), 3rd and 4th successively narrower;

genital segment wider than 4th, hind corners shortly lobed. Caudal rami

subtriangular.

@ g-IO mm., ovisacs 15-16 mm.; g 6-7 mm. Q yellowish, when adult with

dark chestnut brown patches on Ist joint of Ist antennae, carapace (except a

Y-shaped pale median mark and the hind lobes); dorsal plates of 3rd and 4th

segments brown; ¢ pale yellow.

Localities. Table Bay and False Bay, on Carcharius and Galeorhinus. 99 adult

and juv., gd (S. Afr. Mus.); Durban, from nostrils of Grey Shark (C. L.

Biden, 1953).

Distribution. British Seas, North Atlantic.

Remarks. The figure of a 3 given by Wilson 1932 after T. Scott is that of an

immature example. Wilson’s figures (1907 and 1932) of the caudal rami in

dorsal view scarcely convey the impression that they are really triangular;

Wilson’s figure (1907, pl. 27, fig. 123) of the ventral surface is more correct.

Pandarus armatus Heller

Fig. 14 ¢.

1865. Heller, Resse ‘Novara’, Crust., p. 202, pl. 19, fig. 4.

1907. Wilson, loc, cit., p. 395 (in key only).

Ig10. Stebbing, Ann. S. Afr. Mus., vi, p. 558.

1953. Capart, Bull. Inst. franc. Afr. Noire, xv, p. 659, fig. 7.

@—Carapace oblong, only slightly wider posteriorly. Dorso-lateral plates of

end segment not extending beyond plates of 3rd segment, 4 spines on hind

margin between the lobes. Plates of 3rd and 4th segments with moderate

median sinus, each with a spine near hind margin (i.e. a pair of spines on each

fused plate). Genital segment wider than 4th segment plates, lateral margins

convex, with deep notch anterior to the acute hind corners. 6th segment plate

rather bluntly rounded apically. Caudal rami prominent, extending con-

siderably beyond 6th segment plate.

8 mm.

Locality. Cape of Good Hope, on Scyllium africanum (Heller).

Distribution. Senegal, on Sphyrna tudes (Capart).

Pandarus cranchit Leach

1819. Leach, Dict. Scz. Nat., xiv, p. 535-

1907. Wilson, loc. cit., p. 403, pl. 28.

1912. Brian, Res. Sci. Camp. Monaco, fasc. 38, p. 14, pl. 3, fig. 1 (coloured).

1923. Wilson, Ark. Zool., xv, 3, p. 9.

1932. 1d. lOG (Cite, patio 27.

SOUTH AFRICAN PARASITIC COPEPODA 259

1935. id., Pap. Tortugas Lab. Carneg. Inst., xxix (452), p- 333, pl. 5, figs.

58-70, pl. 6, fig. 71.

1943. Heegaard, Ark. Zool., xxxiv, A 18, p. 27, figs. 76-8.

1948. Barnard, loc. cit., p. 249.

9—Carapace broad, widest posteriorly, length (incl. lateral lobes) nearly as

long as rest of body (excl. 6th segment). Dorso-lateral plates of 2nd segment

reaching beyond dorsal plates of 3rd segment to about half length of those of

4th segment; 3rd segment plates broad, sinus deep; 4th segment plates

covering at least half of genital segment, with broad but shallow sinus. Genital

segment with pointed posterior lobes, the outer margin of lobe more or less

concave; 6th segment plate subcircular but with distinct basal neck. Caudal

rami acicular.

g—TIafter Wilson] Accessory lobes on carapace present, longer than wide.

Genital segment with hind corners and a lateral lobe acute, rudiments of 5th

and 6th legs represented by spines.

8 mm., ovisacs 8 mm.; @ yellowish, more or less suffused dorsally with

dark chestnut brown, the median pale mark on carapace often more oblong

or cuneiform than Y-shaped.

Localities. Durban (Wilson); Natal coast, on Stegostoma fasciatum, 99, and on

Hammerhead Shark, 99; off Tugela River mouth, 9 (S. Afr. Mus.).

Distribution. North Atlantic, east coast of N. America, Cape Verde Islands,

Gulf of Guinea, Bay of Bengal, Pacific.

Remarks. In the present specimens the lobes of 2nd thoracic segment do not

extend so far back as in Wilson’s descriptions and figures; but that is not of

great importance.

Wilson (1907) makes zygaenae Brady a synonym of satyrus Dana. Perhaps

both might be combined with cranchi as there seems very little real difference.

Pandarus smithi Rathbun

fgaoe)) Ieathbun, Proc. U.S. Nat. Mus., 1x; p..315, pl. 5; fig: 3, pl. 7, fig.9:

rqo74)) Wilson, loc: cit:, p. 410, pls. 29, 30.

KOSI LOC. Clt.. |p. 434) le. 279.

1944. Brian, An. Mus. Argentino, xli, p. 202, pl. 5, fig. 40.

1948. Barnard, loc. cit., p. 249.

@—Carapace broad, widest posteriorly, length (incl. lateral lobes) almost as

long as rest of body. Dorso-lateral plates of 2nd segment extending beyond

ends of 3rd segment plates to about half length of those of 4th segment; the

dorsal area is demarcated by grooves, but, although described as a plate

(Wilson), has no free edges. Plates of 3rd segment completely separated (at

least in adult). Plates of 4th segment covering more than half genital segment,

with moderately deep notch. Genital segment with pointed posterior lobes,

their outer margin slightly concave; 6th segment plate slightly longer than

wide. Caudal rami prominent, trapezoidal, outer (thickened) and inner

margins nearly parallel, apical margin oblique, denticulate.

g—I[after Wilson] Accessory lobes on carapace present, broader than long.

Genital segment not wider than the preceding free segments, hind corners and

lateral lobe not very acute.

260 ANNALS OF THE SOUTH AFRICAN MUSEUM

299mm. Yellowish, greater part of carapace, except hind margin and a

transverse crescentic mark over the ocular area, and all the dorsal plates more

or less suffused with dark chocolate brown.

Localities. Table Bay, on Carcharias, 2 99 (S. Afr. Mus.); Durban, on

Carcharinus (C. L. Biden 1951, 99).

Distribution. East coast of N. America and Gulf of Mexico, Hawaiian and

Laysan Islands, Argentine.

Gen. Perissopus St. & L.

1861. Steenstrup & Liitken, K. Dansk. Videas. Sels. Skr. (5), v, Pp. 393:

1899. Bassett-Smith, Proc. Zool. Soc. Lond., p. 468.

1907. Wilson, Proc. U.S. Nat. Mus., xxxili, p. 352, and p. 340 (larval

stages).

1930. Leigh Sharpe, Mem. Mus. Roy. Hist. Nat. Belge (H.s.) III, 2, p. 7.

1932. Wilson, Bull. U.S. Nat. Mus., no. 158, pp. 424, and 608, 609 (in

key).

1945. Heegaard, Ark. Zool., xxxv, A 18, p. 18 (discussion on Nogagus).

1951. Gnanamuthu, Spolia <eylanica, xxvi, 1, p. 9.

—Body ovate, squarish posteriorly. Carapace smooth, without grooves.

Free thoracic segments (3) each with a pair of plates, those on 2nd segment

dorso-lateral but fused across middle line, those on 3rd and 4th dorsal, meeting

but not fused medianly. Genital segment large, postero-lateral corners quad-

rate, posterior margin with small excision through which abdomen and caudal

rami are more or less visible. Abdomen 1-segmented. Terminal joints of 2nd

maxilliped enlarged, fleshy, reniform, rough and scaly. Four pairs of biramous

legs, rami of 3rd and 4th pairs minute; 5th pair obsolete. Four pairs of

adhesion pads. Ovisacs straight, elongate, uniseriate.

g—Carapace longer than wide; end thoracic segment with small lateral

plates, 3rd and 4th segments without plates. Genital segment with 5th legs.

Abdomen 1-segmented, caudal rami large, with plumose setae. Maxilliped 2

with stout claw shutting against a pair of corrugated knobs. All 4 pairs of legs

biramous, 2-jointed, but joints of 4th pair more or less fused.

On sharks.

Perissopus dentatus St. & L.

Fig. 15 a.

1861. Steenstrup & Liitken, loc. cit., p. 393, pl. 12, fig. 25.

1887. Rathbun, Proc. U.S. Nat. Mus., x, p. 560, pls. 29, 30 (communis).

1899. Bassett-Smith, loc. cit., p. 468.

1907. Wilson, loc. cit., p. 354, pls. 17, 18 (communis).

1924. Brian, Parasitolog. Mauritanica, fasc. 1, p. 33 (communis).

1932. Wilson, loc. cit., p. 425, fig. 267 (communis).

1953. Capart, Bull. Inst. frang. Afr. Noire, xv, p. 662.

@—Carapace narrowed anteriorly, large, about 3~¢ length of genital seg-

ment. A large knob on ventral surface of carapace near the margin, external

to the large 2nd maxilliped. Plates on 2nd and 4th segments with denticulate

margins, those on 3rd segment with entire margins.

26mm. (Wilson: 3:75-4:25 mm.).

SOUTH AFRICAN PARASITIC COPEPODA 261

Locality. Durban, on Milk Shark (Scoliodon). 3 99, C. L. Biden, 1953.

Distribution. Eastern coast of N. America, on various sharks and dogfish

(Wilson); coast of Mauritania (Brian); Senegal (Capart).

Remarks. Wilson (1907) maintains communis as a valid species distinguished

from dentatus St. & L. by relative sizes of carapace and genital segment (1907,

P. 353 in key), the knob on ventral surface of carapace, and certain details

(1907, p. 361).

Fig. 15. Perissopus dentatus St. & L. a, 2; Nesippus alatus Wilson. b, 9. Echthrogaleus coleoptratus

(Guérm). ¢;9:

Gen. Dinemoura Latr.

1814. Rafinesque, Precis Som., p. 31 (Dinemurus).

1829. Latreille, Régne Anim., iv, Crust., p. 197.

1835. Burmeister, Nova Acta Ac. Leop. Carol., xvii, 1, p. 284 (Dinematura).

HOES e oCOtt, Dé Ax; Brit. Paras: Copep., p. 35.

1907. Wilson, Proc. U.S. Nat. Mus., xxxili, p. 374 (key to species ) (Dinema-

tura).

1932. Wilson, Bull. U.S. Nat. Mus., no. 158, pp. 430, and 608, 609 (in

key) (Dinematura).

1943. Dollfus, Bull. Inst. ocean. Monaco, 851, p. 7 (comparison Dine-

matura with Demoleus).

—Body elongate-oval, carapace with grooves. Dorsal plates on 4th seg-

ment moderate, divided by a deep sinus. Genital segment oblong. Segment 6

distinctly separated from genital segment, with a pair of dorsal plates, and

ventrally a rudimentary pair of legs. Abdomen small, one-segmented; caudal

rami laminate, with nonplumose setae. Two adhesion pads behind each

antenna 1. Maxilliped 2 with terminal claw closing between 2 knobs on basal

262 ANNALS OF THE SOUTH AFRICAN MUSEUM

joint. Four pairs of biramous legs, both rami of Ist 2-jointed, of 2nd and 3rd

3-jointed, of 4th laminate, 1-jointed; first 3 pairs with plumose setae, 4th

without setae or spines. Ovisacs straight, elongate, uniseriate.

g—Carapace broadly oval; 2nd segment with lateral processes, 3rd without

dorsal or lateral plates, 4th with or without lateral lobes slightly overlapping

genital segment. Genital and 6th segments fused. Abdomen 2-segmented;

caudal rami laminate, with 4 plumose setae. All 4 pairs of legs biramous,

rami of 1st and 4th 2-jointed, of 2nd and 3rd 3-jointed.

On various sharks.

Wilson (1907, p. 375) ascribes the establishment of the genus to Latreille in

1829, and makes no mention of Rafinesque 1814. Latreille seems to have

altered Rafinesque’s spelling. The original works are not accessible to me.

But Wilson’s combination ‘Dinematura Latreille’ is certainly not acceptable.

Key to the South African species

I. carapace + total length. Dorsal plates of 4th segment much longer than wide,

apically rounded. ¢ 4th segment without lateral lobes. producta

2. Q carapace $ total length. Dorsal plates of 4th segment wider than long, posterior

margin truncate. ¢ 4th segment with lateral lobes. latifolia

Dinemoura producta (Miiller)

Fig. 16 a, b.

1785. Muller, Entomosir., p. 132, pl. 21, figs. 3, 4.

1907. Wilson, loc. cit., p. 380, pl. 23 (references).

1913. Scott, T. & A., loc. cit., p. 86, pl. 22, fig. 9) Dl sain a

pl. 27, figs. 1-8.

1923. Fage, Bull. Soc. zool. Fr., xlviii, p. 281.

1923. Wilson, Ark: Zool., xv, 3, p. 8, pl. 2, figs. rI—reN( aye

rose. ad.; locscit., peagisaigs 270.

1944. Brian, An. Mus. Argent., xli, p. 202 (Dinematura p.).

1950. Matthews & Parker, Proc. Zool. Soc. Lond., cxx, p. 568, figs. 14, 15

(Dinematura p.).

—Carapace subcircular, about 4 total length. Dorsal plates of 4th segment

longer than wide, apically rounded. Genital segment with apices of dorsal

plates obliquely truncate. Abdomen quadrangular.

g—4th segment without lateral lobes, hind margin slightly sinuous. Genital

segment elliptical.

2 up to 20 mm., ovisacs 40-80 mm., ¢ 12:5 mm.

Locality. Durban (Wilson).

Distribution. Northern Atlantic. Usually on Lamna cornubica, also on Alopias,

Scymnus, Laemargus, and Cetorhinus.

Remarks. Matthews and Parker figure the ¢ and 9 2nd maxillipeds showing

differences in the specimens taken on Lamna and Cetorhinus and differences in

other appendages are noted. It is suggested that ‘the species has a number of

ecotypic variants associated with the different hosts’ (p. 568), the differences

being ‘determined by the nature of the host-skin on which the larvae settle’

(Pp. 573)-

SOUTH AFRICAN PARASITIC COPEPODA 263

Dinemoura latifolia St. & L.

Bis. wOuc, a,

1861. Steenstrup & Liitken, A. Dansk. Viden. Selsk. Skr. (5), v, p. 378,

(0) ONS nem Wop

F907. Wilson, loc. cit., p., 383, pls: 24, 25.

Bg22. id., loc. cit., p. 6, pl. 1, figs. 6—10, pl. 2, fig. 19 (a@mmat. 9).

Boao.) id., loc. cit., p:'492, fig. 271.

1936. Yamaguti, Paras. Copep. Japan, pt. 3, Caligoida 2, p. 9, pls. 5, 6.

1944. Brian, loc. cit., xl, p. 201 (Dznematura /.).

1954. Deboutteville & Nunes-Ruivo, Vie et Milieu, iv, p. 204, figs. 2, 3

(Dinematura l.).

ime

Fig. 16. Dinemoura producta (Miiller). a, 9; 6, 4th and genital segments g. Dinemoura latifolia

St. & L. c, 9; d, 4th and genital segments §. Achtheinus dentatus Wilson. e, 2; f, g. g, antenna

2 2 (copies: a, after Scott; b-d, after Wilson).

9—Carapace broadly ovate, about 4 total length. Dorsal plates of 4th seg-

ment wider than long, posterior margin truncate and slightly emarginate.

Genital segment with apices of dorsal plates bluntly rounded. Abdomen

kidney-shaped, twice as wide as long.

g—4th segment with lateral lobes. Genital segment somewhat wedge-

shaped.

2 up to 16 mm., ovisacs 30 mm., g 8:5 mm.

Locality. Durban (Wilson).

Distribution. Northern Atlantic and Mediterranean, Argentine, Pacific.

Gen. Echthrogaleus St. & L.

1861. Steenstrup & Liitken, K. Dansk. Videns. Selsk. Skr. (5), v, p. 380.

1907. Wilson, Proc. U.S. Nat. Mus., xxxill, p. 362 (key to species).

1932. id., Bull. U.S. Nat. Mus., no. 158, pp. 426, and 608, 609 (in key).

264 ANNALS OF THE SOUTH AFRICAN MUSEUM

°—Body ovate-elliptical. Carapace with grooves. Dorsal plates of 4th seg-

ment large, covering more than half genital segment. The latter as large as or

larger than carapace, with deep posterior sinus between rounded lobes.

Abdomen small, wholly or partly concealed beneath genital segment; caudal

rami large, with nonplumose setae. Four pairs of biramous legs, rami of Ist

pair 2-jointed, outer rami of and and grd’3-jointed, inner rami 2-jointed, rami

of 4th pair large, lamellate, indistinctly jointed. Ovisacs elongate, filiform,

uniseriate.

§—Carapace broader than long, with accessory lobes on hind margin.

and-4th thoracic segments free, each with a pair of small dorsal plates. Genital

segments much smaller than carapace. Abdomen visible in dorsal view,

2-segmented; caudal rami large, with plumose setae.

On sharks.

Echthrogaleus coleoptratus (Guérin)

Fig. 15 ¢.

1907. Wilson, loc. cit., p. 367, pl. 19.

1910. Stebbing, Ann. S. Afr. Mus., vi, p. 559 (not the specimen there

recorded, which = Philorthragoriscus, q.v.).

1923. Wilson, Ark. Zool., xv, 3, p. 13.

NCOOAN TNCs oes itty) oo VAT ally Bk

1936. Yamaguti, Paras. Copep. Japan, pt. 3, p. 7, pls. 4, 5.

1944. Brian, An. Mus. Argentino, xli, p. 202.

—Body more than twice as long as wide, carapace smaller than genital

segment. Margins of plates on 4th segment not serrate. Abdomen concealed.

g—Carapace subcircular. Lateral lobes of 2nd segment extending outwards,

tips concealed by lobes of carapace.

2 12-14 mm., ¢ 3:6 mm. Yellowish; carapace, especially the central portion,

and dorsal plates of 4th segment brown.

Localities. Durban (Wilson); Kalk Bay (False Bay), on Carcharinus glaucus.

1 9 (S. Afr. Mus.).

Distribution. Atlantic and Indian Oceans, Pacific, Argentine.

Remarks. In the General Catalogue of S.A. Crustacea Stebbing recorded a

specimen (‘No. 54’) from Orthagoriscus mola. In tube 54, as returned to the S.A.

Museum, with Stebbing’s autograph label, there were 3 specimens of Philor-

thragoriscus (not recorded by Stebbing) and no Echthrogaleus. The Sunfish

would be an unusual host for this latter genus. (cf. 1946, Dollfus, Ann. Soc. Sc.

Nat. Charante-Maritime, ii, 7, p. 71.)

Wilson’s record, and the present specimen, however, prove that the genus

does occur in South Africa, as would be expected.

Gen. Nesippus Heller

1865. Heller, Rezse ‘Novara’, zool., 11, Crust., p. 193.

1907. Wilson, Proc. U.S. Nat. Mus., xxxili, p. 424.

1932. id., Bull. U.S. Nat. Mus., no. 158, pp. 438, 607, 609 (in key).

1943. Heegaard, Ark. Zool., xxxiv, A 16, p. 6.

1944. Wilson, Proc. U.S. Nat. Mus., xciv, p. 538 (3 formerly ascribed to

Achtheinus).

SOUTH AFRICAN PARASITIC COPEPODA 265

©—Carapace broader than long, smooth, without grooves; 2nd and 3rd

segments fused, with lateral plates; 4th segment free, with a pair of fused

dorsal plates. Genital segment large, ovate, without posterior processes or

rudiments of legs. Abdomen small, concealed; caudal rami large, more or

less concealed. Antenna 2 uncinate. Maxilliped swollen, with terminal claw.

Four pairs of biramous legs, rami of first 3 pairs 2-jointed, of 4th 1-jointed.

Ovisacs elongate, filiform, uniseriate.

g—Carapace ovate. Free thoracic segments much narrower than carapace,

2nd segment with lateral lobes. Genital segment ovate, without rudiments of

legs (or these reduced to mere spines). Abdomen one-segmented; caudal rami

small, with plumose setae. Maxilliped with knobs or a terminal claw. Legs

resembling those of 9.

Nesippus alatus Wilson

Fig. 15 0b.

Hag) Valson, loc: cit., p. 426, pls.-34, 35 (2 3).

Hg22- 10+, loc. cit., p. 438, fis, 276: (9 3).

©—Dorso-lateral plates of 2nd segment large, alate, angular; dorsal plates

of 4th segment slightly overlapping base of genital segment.

6-7 mm., ovisacs 12-13 mm.

Locality. Durban, on hind part of tongue of Carcharodon carcharias (numerous

29, C. L. Biden, 1953).

Distribution. Eastern coast of N. America, on various sharks (Wilson).

Remarks. Usually found on the gill-arches, rarely on the floor or roof of the

mouth; Wilson (1932) says this is the only Copepod likely to be found inside

the throat of a shark.

Heller’s species orientalis and crypturus were taken at Java; neither of them

have the alate plates on 2nd segment characteristic of alatus, and the plates on

4th segment do not reach the genital segment.

Brian (1924. ‘Parasitol. Mauritanica,’ fasc. 1, p. 33) identifies Mauritanian

specimens with ortentalis. Capart (1953. Bull. Inst. frang. Afr. Noire, XV, 2,

pp. 658, 659, fig. 6) also identifies Senegal specimens with orientalis, and con-

siders that more abundant material will eventually show that alatus Wilson and

angustatus v. Ben. should be united with Heller’s species. Capart’s fig. 6

certainly suggests that the plates on 2nd segment are alate.

Gen. Achtheinus Wilson

1908. Wilson, Proc. U.S. Nat. Mus., xxxv, p. 450.

LOPiay 14.5 1b1d...xo0x, p. 630;

HED 102 ibides xhit,: ps 295.

1924. Kurtz, SB. Ak. Wiss. Wien. Abt., 1, cxxxill, p. 613 (Pholidopus).

1927. Wilson, Sczence, N.Y., Ixvi, p. 397 (validity of genus).

HO22es 1d, Arh.“ 00l., xiv eNO. TOs Dp. A.

1932. id., Bull. U.S. Nat. Mus., no. 158, pp. 608, 609 (in key).

1936. Yamaguti, Parasit. Copep. Japan, pt. 3, Caligoida 2, p. 11.

1944. Wilson, Proc. U.S. Nat. Mus., xciv, p. 539.

266 ANNALS OF THE SOUTH AFRICAN MUSEUM

9—Body ovate-elliptical; carapace (head + 1st thoracic segment) smooth,

without grooves; 2nd and 3rd segments fused, with a single pair of dorsal

plates; 4th segment free, with a similar pair of plates. Genital segment with

apical median sinus. Abdomen small, concealed beneath genital segment,

caudal rami large, partly or wholly concealed. Antenna 2 stout, with strong

claw. Maxilliped swollen, with slender apical claw. Four pairs of biramous

legs, rami of first 3 pairs 2-jointed, of 4th pair 1-jointed. Ovisacs straight,

elongate, uniseriate.

g—Carapace, broadly ovate. Free thoracic segments much narrower than

carapace, dorsal plates represented only by lateral lobes. Genital segment

with rudiments of 5th legs. Abdomen 1-segmented, caudal rami laminate,

with 4 setae. Antenna 2 without teeth. Maxilliped with strong terminal claw.

Legs resembling those of 9.

On sharks.

Pholidopus Wilson 1907 (pro Lepidopus Dana 1852 preocc.) and Achtheinus are

very closely related, but Wilson (1927) showed that Kurtz’s arguments were

ill-founded, and maintained the validity of the latter genus.

Key to the South African species (29)

1. Antenna 2 dentate. dentatus

2. Antenna 2 not dentate. pingutis

Achtheinus dentatus Wilson

Fig. 16 e-g.

Tori, \Wailson: loc eit., p.'630) pl. (67, ;hles.22—315( Oe

TO12) “ids locseitesip) 290:

1917. siren S. Afr. Mus., xvii, p. 41, pl. 8 (S. Afr. Crust., pl. 97)

(P&S

1921. Wilson, Proc. U.S. Nat. Mus., lix, p. 6, pl. 3, figs. 20-27 (¢ &

immature @).

1922. id., Nyt Mag. Naturv., |x, p. 107.

1923. id., Medd. Goteb. Mus. Zool. Avd., no. 19, p. 7, pl. 1, figs. 6-12 (9)

(parvidens).

1944. 1d., loc: cit., p. 537, pl. 33; figs. 196-200) (Cec troeaae

1944. Brian, An. Mus. Argentino, xli, p. 203, pl. 2, figs. 15-17, pl. 3,

figs. 18-29 (2 g).

—Carapace about as broad as long, posterior corners rounded (not acute

as might be thought from Stebbing’s figure). Antenna 2 falciform, with a

double row of stout teeth apically, 4 teeth on dorsal edge and 3 on ventral edge

of inner margin, and a pair of slightly larger teeth proximally to these two

rows. Dorsal plates separate from their bases. Abdomen with an accessory

lobe on each side at base.

g—Carapace slightly longer than broad (as broad as long if pressed flat),

hind margin as in 9. Lateral angles of 1st free segment subacute; on 2nd and

3rd segments rounded and overlapping the succeeding segment. Genital seg-

ment quadrate, hind corner with a minute spiniferous denticle (5th leg);

hind margin with a rounded exsert lobe on each side.

2 up to 8:5 mm., ovisacs 15-18 mm.; ¢ 3°75 mm. Yellowish.

Localities. Algoa Bay, on shark (Stebbing) ; Table Bay, on Acanthias (Wilson) ;

Kalk Bay, on Mustelus and Carcharias; ‘Table Bay, on Carcharodon (S. Afr. Mus.) ;

Natal, on dogfish (Natal Mus.).

SOUTH AFRICAN PARASITIC COPEPODA 267

Distribution. Coast of Peru and California, Argentine.

Remarks. Stebbing says Wilson’s figure shows the relative length and breadth

of the 2 carapace better than his; in fact Stebbing’s figure is an accurate

representation of the South African specimens. Wilson (1911) described and

figured the 2nd antenna as having only a single row of teeth, but corrected this

statement in 1921, and in 1944 (fig. 197).

Stebbing described what he thought was a 3, and appeared to have con-

sidered this the first description of the ¢ of this genus. Wilson, however, in

1912 had described a ¢ assigned to A. pinguis.

In 1921 Wilson, without reference to Stebbing’s 1917 paper, described a 3

assigned to dentatus, and also immature 99. Wilson’s immature 9, 3-5 mm. in

length, corresponds with the specimen described by Stebbing as a 3. Stebbing

found his specimen attached to the underside of the genital segment of a Q,

with the head pointing forwards. Among other specimens of the same lot there

are several such ‘pairs’, and from the position of the smaller specimens one

would naturally assume them to be males.

In 1944 Wilson, again without reference to Stebbing, retracted his 1912

description of the g of pinguis (assigning it to WNesippus*), and also his

description of the (supposed) ¢ of dentatus. ‘The supposed immature 99 are

now regarded as 3g.

Thus Stebbing was right in claiming the first description of the 3, not only

of dentatus, but of the genus Achtheinus.

Wilson’s parvidens is obviously a synonym.

Achtheinus pinguis Wilson

LO12: oo loc. cit., p. 235, pl. 31, figs. 8-14 (¢), pl. 32, figs. 15-21

Q).

HG Sem IGe Arc. KOOL. KV) Bnei he

IQ41. ae Mem. Mus. R. Hist. Nat. Belge, ser. 2, fasc. 21, p. 184,

gs. 5-7.

Not 1944. Wilson, loc. cit., p. 538 (¢ = Nesippus).

@—Similar to dentatus, but antenna 2 not dentate.

5°75 mm.

Localities. Cape of Good Hope, on Saw-shark (Pliotrema warreni) (Wilson) ;

Table Bay, on ‘dogfish shark’ (Wilson); off Dassen Island (West coast), on

Acanthias vulgaris (Capart).

Remarks. ‘The differences in plumpness and relative width of carapace

between this species and dentatus, mentioned by Wilson, would not seem to be of

great significance.

Unfortunately the only specimens (99) in the South African Museum from

Pliotrema have lost the 2nd antennae, and are therefore only presumed to be

pinguis. ‘They were collected in 1898 by Dr. Gilchrist, then Marine Biologist

to the Gape Government.

* See Wilson, Proc. U.S. Nat. Mus., xxxiii, p. 424.

268 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fam. CECROPIDAE

1907. Wilson, Proc. U.S. Nat. Mus., xxxili, p. 461 (Cecropinae).

1932. id., Bull. U.S. Nat. Mus., no. 158, p. 441.

The four genera, three of which occur in South Africa, are found most

commonly on Sunfishes (Molidae), but are found occasionally also on Diodon,

Thynnus, and other Teleosts, and on sharks.

More gregarious than the Pandaridae, and when thus congregated in bunches

cause sores and pits in the flesh of the host. Both sexes are incapable of

swimming.

Gen. Cecrops Leach

1816. Leach, Encycl. Brit. (suppl. to 4th-6th eds.), I, (2), p. 405.

1907. Wilson, loc. cit., p. 466.

1923. id., Amer. Mus. Nov., no. 80, p. 1.

1932. id., loc. cit., p. 441 (diagnosis of ¢ legs not quite accurate).

@—Carapace oval, deeply emarginate posteriorly, frontal plates fused with

carapace. 3rd segment with small fused dorsal plates; 4th segment with

larger fused plates. Genital segment with fused dorsal plates larger than

carapace. Abdomen with expanded ventral plates. Antenna 1 2-jointed.

Rami of tst-3rd pairs of legs 2-jointed, of 4th pair 1-jointed. Ovisacs numerous,

irregularly coiled, concealed, uniseriate.

Q—similar to 9 but 4th dorsal plates not completely covering abdomen.

4th pair of legs not much enlarged.

Margins of carapace and dorsal plates in both sexes smooth.

Two species, one of which is found chiefly on the Sunfish, usually attached

to the gills, but has also been found on Diodon, Thynnus, and Pleuronectes; the

other (exzguus Wilson 1923) on sharks.

Cecrops latreillit Leach

Fig. 17 a-c.

1899. Bassett-Smith, Proc. Zool. Soc. Lond., p. 465 (references).

1907. Walson;loc..eit:,)p: 460, pls. 36. 90.

1910. Stebbing, Ann. S. Afr. Mus., vi, p. 558.

1912. Brian, Res. Sci. Camp. Monaco, fasc. 38, p. 13 (references).

1932. Wilson, loc. cit., p. 442, fig. 278.

1936. Yamaguti, Paras. Copep. Japan, pt. 3, p. 10, pl. 6, figs. 62-66.

Ovig. 2 up to 30 mm., width 12-15 mm.; g¢ up to 17 mm., width 11 mm.

Uniform yellowish white.

Locality. From the Cape (Stebbing, and S. Afr. Mus.), on the Sunfish

(Orthagoriscus = Mola).

Distribution. Atlantic and Pacific Oceans.

Remarks. Heegaard (1943. Ark. Zool., xxxiv, 18, p. 26) records a specimen

from Thynnus, but suggests that this may be due to a mistake in labelling, or

accidental transference from the parasite’s normal host.

SOUTH AFRICAN PARASITIC COPEPODA 269

Gen. Orthagoriscicola Poche

1902. Poche, Zool. Anz., xxvi, p. 15.

1907. Wilson, loc. cit., p. 472.

nae2. id., loc.. cit... p: 443.

@—Carapace subcuneiform, shallowly emarginate posteriorly, lateral

margins serrate, frontal plates fused with carapace but their outlines indicated

by grooves. Dorsal surface spinulose. 2nd and 3rd segments free, without

dorsal plates; 4th segment with large fused plates, with serrate margins.

Genital segment with large overlapping dorsal plates, with serrate margins.

Abdomen with expanded lateral plates. Antenna 1 3-jointed. Rami of 1st and

and pairs of legs 2-jointed, of 3rd and 4th 1-jointed and laminately enlarged.

Ovisacs numerous, irregularly coiled, concealed, uniseriate.

Fig. 17. Cecrops latreillei Leach. a, ovigerous 2; 5, genital segment 9; left half of dorsal plate

removed to show ovisacs; ¢, g. Orthagoriscicola muricatus (Kroyer). d, 2; e, ventral view 2

genital segment with abdomen and 3rd and 4th legs; f, 3; g, ¢ spermatophores removed from

9. Philorthragoriscus serratus (Kroyer). h, 9; 2, 6 (copy after Wilson).

270 ANNALS OF THE SOUTH AFRICAN MUSEUM

g—Similar to 2 but carapace relatively wider. Genital segment much

swollen, its dorsal plates fused, with posterior sinus, margins serrate. Abdomen

and caudal rami relatively larger. Rami of Ist-3rd pairs of legs 2-jointed, of

4th pair as in 9.

A single species found on the Sunfish (Orthagoriscus = Mola), on the gills or

more frequently on the surface of the skin; has also been found on Selene vomer

(Carangidae).

Orthagoriscicola muricatus (Kr6éyer)

Fig. 17 d-g.

1899. Bassett-Smith, Proc. Zool. Soc. Lond., p. 467 (Laemargus m.).

1907. Wilson, loc. cit., p. 473, pls. 40, 41.

1910. Stebbing, Ann. S. Afr. Mus., vi, p. 559.

1912. Brian, Res. Scz. Camp. Monaco, fasc. 38, p. 14, pl. 2, figs. 1, 2, pl. 6,

figs. 5-8 (references).

1932. Wilson, loc. cit., p. 443, fig. 279.

Ovig. 2 up to 20 mm., width 7-9 mm.; ¢ up to 15 mm., width 8 mm. Pale

yellow.

Locality. Table Bay (Stebbing and S. Afr. Mus.), on Mola.

Distribution. Atlantic.

Remarks. Wilson states that the carapace of ¢ has the margins and dorsal

surface smooth; but that is not the case with the specimens I have examined,

in which the § carapace resembles that of the 2 though not so strongly serrate

and spinulose.

One ¢ in the S.A. Museum collection still has the two frontal filaments

attached though it is mature with fully developed spermatophores. The

filaments are nearly as long as the body.

Gen. Philorthragoriscus Horst

1897. Horst, Notes Leyden Mus., xix, p. 137.

1907. Wilson, Proc. U.S. Nat. Mus., xxxiii, p. 478.

1932. id., Bull. U.S. Nat. Mus., no. 158, p. 444.

Q9—Carapace rounded, deeply emarginate posteriorly, lateral margins

serrate, frontal plates fused. 4th segment with a pair of large plates fused at

base, but deeply cleft, margins serrate. Genital segment not quite as large as

carapace, with fused dorsal plates, apically cleft, margins finely serrate, antero-

lateral angle acute. Abdomen concealed, without lateral expansions; caudal

rami large. Antenna I 2-jointed. Rami of ist-3rd pairs of legs 2-jointed, with

spines and setae; peduncles of 3rd pair large, concealing the small 4th pair

of legs, the rami of which are 1-jointed, with spines only. Ovisacs elongate, not

concealed.

g—Carapace much larger than rest of body, 2nd and 3rd segments fused,

with small lateral plate. 4th segment with rudimentary dorsal plates. Genital

segment quadrangular, with short fused dorsal plates at its base, postero-

lateral angles acute.

A single species parasitic on the Sunfish; originally placed in Dinemoura

(Pandaridae).

SOUTH AFRICAN PARASITIC COPEPODA PAG ia

Philorthragoriscus serratus (Kroyer)

Bie 7s a:

1863. Kroyer, Naturh. Tidsskr., p. 176, pl. 8, figs. 4 a-1 (Dinematura s.).

Hea) blorst, loc. cit, p. 137; pl7.

w9o7. | VWilson, loc: cit., p- 470,'pls..42, 43:

1912. Brian, Res. Sci. Camp. Monaco, fasc. 38, p. 12, pl. 2, fig. 3, pl. 6,

fig. g.

1922. Wilson, Nyt Mag. Naturv., |x, p. 108.

feg2.) tid:, loc: cit., p-445, fis. 280.

1948. Barnard, Ann. Mag. Nat. Hist. (xii), 1, p. 249.

27-5 mm., width 4:5 mm., ovisacs (Wilson) 15 mm.; g 5 mm., width 4 mm.

Yellowish, clouded with grey (Wilson).

Locality. Table Bay, on Mola (S. Afr. Mus.).

Distribution. Atlantic.

Remarks. A single non-ovigerous 2 was found among the specimens of

Orthagoriscicola identified by Stebbing, and overlooked by him; three 99

returned by Stebbing labelled as ‘No. 54. Echthrogaleus coleoptratus’.

Wilson (1922) records this species from Squalus acanthias.

Fam. ANTHOSOMIDAE

1922. Wilson, Proc. U.S. Nat. Mus., lx, 5, pp. 1, 19, 20 (Dichelesthiidae

part, Anthosominae).

1932. id., Bull. U.S. Nat. Mus., no. 158, p. 446 (Anthosomidae).

Gen. Anthosoma Leach

1816. Leach, Encycl. Brit. (suppl. to 4th-6th eds.), I, 2, p. 406.

1922. Wilson, loc. cit., p. 23.

Hosen) ides loc. cit., p. 446.

9—Body short, stout; head and 1st thoracic segment fused to form a large

oval carapace. 4th segment dorsally with 2 large overlapping plates con-

cealing the genital segment and most or all of the abdomen. Abdomen

I-segmented, caudal rami narrow, unarmed. Antenna 1 6-jointed. Antenna 2

3-jointed, stout, porrect, with apical claw. Maxillipeds stout, with strong

apical claw. First 3 pairs of legs large foliaceous, inner margin of 1st and 2nd

pair notched but without rami; 4th pair of legs absent. Ovisacs slender,

elongate, eggs uniseriate.

g—similar but smaller; 4th segment without dorsal plates. Inner margin

of 1st and 2nd legs notched, with 2 small rami, those on 1st leg ovate, the inner

one on 2nd leg 2-jointed, with 3 apical curved spines, moved by powerful

muscles.

Parasitic on sharks.

Anthosoma crassum (Abildgaard)

Bice 1G:

1794. Abildgaard, Skr. nat. Selsk. Copenhagen, ii, p. 46, pl. 5, figs. 1-3.

(Caligus c.)

1816. Leach, loc, cit., p. 406, pl. 20, figs. 1-6 (smithit).

272 ANNALS OF THE SOUTH AFRICAN MUSEUM

1899. Bassett-Smith, Proc. Zool. Soc. Lond., p. 468.

1915. Stebbing, Ann. S. Afr. Mus., xv, p. 58 (record only).

1922. Wilson, loc. cit., p. 23, pl. 1, figs. 1-8.

Omer, ich, Alle, K@olkey ivy By Os RS

1924. ids,eProc: (U.S. Natwius.. xiv ps 1S:

1932, id., loc: cit., p) 446, fig. (28i:

1936. Yamaguti, Paras. Copep. Japan, pt. 3, Caligoida 2, p. 12.

1944. Brian, An. Mus. Argentino, xli, p. 208, pl. 5, figs. 43, 44.

1948. Debouteville, Bull. Mus. Hist. Nat. Paris, xx, 5, p. 446.

1952. Birkett & Burd, Ann. Mag. Nat. Hist. (xii), v, p. 391. fig.

Fig. 18. Anthosoma crassum (Abildg.). a, 9, with 2nd antenna further enlarged; 6, g; ¢, 1st leg

6; d, 2nd leg ¢.

215 mm., d 9 mm., ovisacs 25 mm. Carapace yellowish-brown or amber,

dorsal plates and foliaceous legs greyish-white, abdomen brown.

Localities. Durban (Stebbing, Wilson); Table Bay, False Bay, and Durban,

on Porbeagle and species of Carcharias (S. Afr. Mus.).

Distribution. Europe, east and west coasts of N. America, New Zealand,

Japan, Argentine. On various sharks: Lamna, Carcharias, Isurus, Oxyrhyncha,

Cetorhinus, Selache.

Remarks. Wilson says this parasite is usually found in the throat attached

to the gill-arches. Birkett & Burd record a heavy infestation in the skin of the

underside of the shark, Selache, extending around the [axils of the] pectoral

fins and into the mouth and gill-slits.

The South African Museum specimens were collected by Mr. C. L. Biden

(1928 and 1953) from among the teeth of Porbeagles and species of Carcharias.

The parasites were present in enormous numbers, closely packed together;

the flesh of the jaws was raw and suppurating, and in some cases most of the

teeth had fallen out. As Birkett & Burd remark, such an infestation must

seriously endanger the health and even the life of the host.

SOUTH AFRICAN PARASITIC COPEPODA 273

Gen. Lernanthropus Blainv.

1822. Blainville, 7. de Physique, xcv, p. 443.

1832. Nordmann, Mikrosc. Beitr., ii, p. 45 (Epachthes).

1864. id., Bull. Soc. Nat. Moscou, xxxvii, p. 510 (Stalagmus).

1922. Wilson, loc. cit., p. 30 (key to species).

1932. id., loc. cit., p. 447, and 609 (in key).

1935. id., Pap. Tortugas Lab. Carneg. Inst., xxix (452), p. 338.

1936. Yamaguti, Parasit. Copep. Japan, pt. 3, pp. 13-18.

1940. Gnanamuthu, Rec. Ind. Mus., xlv, p. 291.

1950. id., Parasitology, xl, p. 277.

1954. Deboutteville & Nunes-Ruivo, Bull. Inst. frang. Afr. Notre, xvi,

pp. 141-58.

1954. Yamaguti, Publ. Seto Mar. Biol. Lab. II, 3, pp. 387 sqq.

Q—Head and ist thoracic segment fused into a carapace with lateral margins

curved down ventrally. 2nd-4th segments fused, covered by a single dorsal

plate prolonged backwards over genital segment and abdomen; latter 1- (or

2-) segmented. Antenna 1 filiform, joints more or less fused. Antenna 2 and

maxillipeds prehensile, uncinate. 1st and 2nd pairs of legs biramous but rudi-

mentary; rami of 3rd and 4th pairs modified: each leg of the 3rd pair formed

of fused inner and outer ramus, folded along middle (cross-section semi-

circular), fleshy; rami of 4th legs elongate, extending backwards, fleshy.

Ovisacs elongate, filiform, eggs uniseriate.

g—Carapace with flat margins. 2nd-4th segments fused with genital seg-

ment, no dorsal plate. Abdomen 1-segmented, visible dorsally. tst-3rd legs

as in 9, rami of each 4th leg fused, elongate, projecting backwards.

Key to the South African species

1. Ventral plate present. petersi

2. Ventral plate absent. paradoxus

Lernanthropus petersi v. Ben.

Fig. 19.

1857. Wan Beneden, Bull. Ac. Roy. Belgique, xxiv, n.s. I, pp. 51-63, pl.

1864. Nordmann, loc. cit., p. 510 (Stalagmus p.).

1899. Bassett-Smith, Proc. <ool. Soc. Lond., p. 472 (reference to van

Beneden omitted).

1922. Wilson, loc. cit., p. 35 (in key).

1939. Brian, Rev. ool. Bot. Afr., xxxii, p. 183, fig. vi.

—A dorsal plate covering entire body, only rami of 3rd and 4th legs

visible dorsally; posterior margin of plate pointed. A ventral plate similar to

the dorsal plate, covering bases of 3rd and 4th legs but leaving genital segment

and abdomen uncovered. (Characters from Wilson’s 1922 key.)

10 mm.

Localities. Mozambique, on gills of Serranus goliath (= Epinephelus fuscogut-

tatus) (van Beneden); Belgian Congo, host unknown (Brian).

Lernanthropus paradoxus (Nordm.)

1832. Nordmann, Mikrogr. Betr., uu, p. 45 (Epachthes p.).

1833. Burmeister, Act. Ac. Leop. Car. Nat. Cur., xvii, p. 307, pl. 14, fig. 12.

274 ANNALS OF THE SOUTH AFRICAN MUSEUM

1840. Milne Edwards, Hist. Nat. Crust, 111, p. 499

1899. Bassett-Smith, Proc. Zool. Soc. Lond., p. 460.

1922. Wilson, loc. cit., pp. 22, 36, and 35 (in key).

—A dorsal plate covering entire body, only rami of 3rd and 4th legs

visible, but no ventral plate. 3rd and 4th legs, genital segment, and abdomen

entirely visible in ventral view. 3rd legs biramous. (Characters from Wilson’s

1922 key.)

8-7 mm.

(

nase man ~ =

aur ae.

—, tn

Fig. 19. Lernanthropus petersi van Ben. Dorsal and ventral views 2 (copies from

van Beneden; the ventral view shows the 2nd antennae turned downwards as in

Brian’s figure).

Locality. Gape of Good Hope, on Mugil sp.

Remarks. Wilson (1922, p. 36) says that Nordmann in his text referred to

pl. 12, figs. 12-14, that no such plate was published, and that the only illustration

of this species is that published by Burmeister.

Fam. EUDACTYLINIDAE

1922. Wilson, Proc. U.S. Nat. Mus., 1x, 5, pp. 1, 19, 20

part, Kudactylinae).

id., Bull. U.S. Nat. Mus., no. 158, p. 452 (Hudactylinidae).

(Dichelesthiidae

1932.

SOUTH AFRICAN PARASITIC COPEPODA 275

Gen. Nemesis Risso

e204 Risso, esi. Vai. Hur, v, p. 135.

1913. Wilson, Proc. U.S. Nat. Mus., xliv, p. 236.

1922. id., loc. cit., p. 58 (references and key to 5 species) (characters of

¢ incorrectly stated).

1923. Fage, Bull. Soc. Zool. Fr., xlviii, p. 282.

1932. Wilson, loc. cit., p. 460 (key to 3 species).

9—Head and ist thoracic segment fused. Carapace elliptical, longer than

wide. ist-3rd free segments wider than long, each covered with a dorsal plate

whose sides curve round on to ventral surface; 4th free (= 5th) segment

narrower than preceding segments; genital segment narrower than 4th.

Abdomen narrow, 2-3-segmented; caudal rami tipped with spine-setae.

Antenna I 10-15-jointed. Antenna 2 uncinate. Maxillipeds large, uncinate.

4 pairs of biramous legs, rami of ist pair differing from those of and-4th; 5th

pair rudimentary, uniramous. Ovisacs elongate, filiform, uniseriate. Sperma-

tophores spherical, attached to genital segment.

g—Similar, but narrower. Dorsal plates of 1st-3rd free segments not curved

over ventrally to such an extent as in 9; 4th free segment much shorter than

3rd; genital segment large. Abdomen 3- or 4-segmented; caudal rami larger

than in 9. tst-4th pairs of legs larger, more strongly setose.

Gill-parasites on sharks.

Nemesis pallida Wilson

Fig. 2074) b.

1932. Wilson, loc. cit., p. 464, pl. 30 b-p.

1935. id., Pap. Tortugas Lab. Carn. Inst., xxix (452), p. 340.

1948. Barnard, Ann. Mag. Nat. Hist. (xii), 1, p. 250.

@—Carapace regularly oval, greatest width in centre. 1st free segment

without notch on lateral margin; 4th free segment wider than long, but

narrower than 3rd, hind margin evenly curved, sternal plate just visible

laterally. Genital segment wider than long. Abdomen 3-segmented. Antenna 1

13-14-jointed. No trace of 5th pair of legs on the 4th free segment.

g—Abdomen 4-segmented.

2 4°5 mm., ovisacs 7-8 mm., § 4 mm. As preserved dirty yellowish-grey,

ovisacs yellow, spermatophores dark maroon.

Locality. ‘Table Bay, on gills of Thresher shark (Alopias) (S. Afr. Mus.).

Distribution. East coast of N. America.

Remarks. In general features these specimens correspond with Wilson’s

description, but the hind margin of the 5th (4th free) segment in 9 cannot

be described as 3-lobed.

Fam. DicHELESTHIIDAE

1922. Wilson, Proc. U.S. Nat. Mus., |x, 5, pp. 1, 19-21. (Dichelesthidae,

part, Dichelesthiunae.)

1932. id., Bull. U.S. Nat. Mus., no. 158, p. 476. (Dichelesthiidae restricted.)

Lamproglena Nordmann has the abdomen nearly as long as the trunk,

obscurely 3-segmented. L. monodi Capart (1944. Bull. Mus. Hist. Nat. Belg.,

276

ANNALS OF THE SOUTH AFRICAN MUSEUM

xx, 24, p. 8, fig. 2) occurs in the Belgian Congo on the fresh-water fishes

Serranochromis thumbergi and various species of Haplochromis.

Fig. 20. Nemesis pallida Wilson. a, 9, spermatophore shown on left, base of

ovisac on right side; b, g§. Hatschekia acuta Brnrd. c, 2, with ventral view

of abdomen; d, antenna 2; e, maxilliped.

Gen. Hatschekia Poche

1902. Poche, ool. Anz., xxvi, p. 16.

1913. Wilson, Proc. U.S. Nat. Mus., xliv, p. 239.

1922. id., loc. cit., p. 81 (key to species).

1932) jude loc ycit. o-04:70:

1939. Yamaguti, Paras. Copep. Japan, pt. 5, Caligoida 3, vol. Fubil. Prof.

Yoshida, 11, pp. 458-69.

1951. Pearse, Proc. U.S. Nat. Mus., ci, p. 357.

1953. Yamaguti, Publ. Seto. Mar. Biol. Lab., III, 2, p. 225.

1954. id., ibid., III. 3 pp. 392 sqq.

9—Head rounded or transversely oval. First 2 thoracic segments more or

less free,

but often fused, each with a pair of biramous legs. Remaining

thoracic segments and genital segment forming a trunk, subcylindrical, more

or less elongate, but sometimes elongate-ovate. Abdomen short, 1-segmented,

or absent

. Antenna 1 3-6-jointed, often indistinctly jointed. Antenna 2 stout,

with apical claw. Maxilliped slender, uncinate. Two pairs of biramous legs,

sometimes rudiments of 3rd and 4th pairs. Ovisacs cylindrical, short or

moderately long, eggs uniseriate.

SOUTH AFRICAN PARASITIC COPEPODA 249/57

6—Head rounded, separated from the fused thoracic segments. Abdomen

more or less distinct; caudal rami larger than in 9. Antenna 1 longer and

stouter than in 9. Maxilliped very long, slender, projecting far beyond margin

of head. ‘Two pairs of biramous legs, rami 1-jointed.

Parasitic on gills of various Teleost fishes.

Hatschekia acuta Brurd.

Fig. 20 c=.

1948. Barnard, Ann. Mag. Nat. Hist. (xii), 1, p. 250, fig. 7.

°—Head transversely oval. First 2 thoracic segments completely fused, as

wide as (or nearly) head; trunk cylindrical, about 6 times as long as wide,

smooth, without indents or constrictions, postero-lateral corners acute. Antenna

1 obscurely 3- or 4-jointed. Ovisacs elongate.

4mm., ovisacs 7-8 mm.

Locality. Fish Hoek, False Bay, on gills of Brama rai (coll. K.H.B. 1935).

Remarks. Appears to be distinguished from other species by the acute

postero-lateral corners of the trunk, but in this respect resembles the Japanese

conifera Yamaguti 1939, from the gills of Stromateoides.

Fam. LERNAEIDAE

1905. Stebbing, Mar. Invest. S. Afr., iv. p. 116.

1917. Wilson, Proc. U.S. Nat. Mus., liii, pp. 1-150, pls. 1-21 (revision of

family).

1932. id., Bull. U.S. Nat. Mus., no. 158, p. 479.

1933. Gurney, Brit. FW. Copep., iii, p. 334 (restricted).

In 1917 Wilson admitted four subfamilies: Lernaeinae, Lernaeenicinae [sic],

Lernaeocerinae, and Pennellinae; in 1932 the Pennellinae are given full family

rank. Gurney included only the first subfamily in the family Lernaezdae.

Life-history. The Copepodid §¢ and 99 infest the gills or fins of fishes, but

the gg do not develop beyond the fourth Copepodid stage when they become

sexually mature. The 99, after fertilization, seek a second fish host, a species

different from the first host, which is necessary for the maturing of the eggs.

The 99° burrow into the second host until the head reaches the dorsal aorta

or one of the main blood-vessels. Horns and variously shaped processes grow

out from the head and fix the parasite in the tissues of the host; and a hard

cyst forms around the head of the parasite.

Parasitic on various fishes, marine and fresh water.

Key to the South African genera (29)

1. 2-4 soft horns symmetrically arranged on head. Ovisacs sack-like, eggs multi-

seriate. Fresh water. Lernaea

2. Head with branched processes but no horns. Ovisacs filiform, eggs uniseriate.

Marine. Cardiodectes

Gen. Lernaea Linn.

1758. Linnaeus, Syst. Nat., roth ed., p. 655.

1914. Cunnington, Proc. Zool. Soc. Lond., ii, p. 819 (Lernaeocera).

1917. Wilson, loc. cit., p. 36.

278 ANNALS OF THE SOUTH AFRICAN MUSEUM

1918. id., Bull. Bureau Fish., xxxv, 1915-16, p. 163, pls. 6-15.

1920. id., Bull. Amer. Mus. Nat. Hist., xliii, p. 5.

1925. lLeigh-Sharpe, Parasitol., xvii, p. 245.

1928. Wilson, Swed. zool. Exp., White Nile, pt. 5, no. 3, p. 13.

1932. Monod, Ann. Parasitol., x, p. 345, and pp. 359, 378 (list of species

and bibliography).

1933. Gurney, loc. cit., p. 336 (development).

1944. Capart, Bull. Mus. Hist. Nat. Belg., xx, no. 24, p. 2.

1950. Harding, Bull. Br. Mus. (N.H.), I, 1, pp. 3-27, figs. (key to species).

@ (adult)—Head a rounded knob, behind which one or two pairs of horns,

simple or forked, soft, conical. Neck soft, thin, gradually enlarging into trunk,

which has a pregenital prominence in front of vulva. Abdomen short, blunt.

A pair of maxillipeds as well as 2 pairs of maxillae. 4 pairs of biramous legs,

and a 5th pair of 1-jointed stumps just in front of vulva. Ovisacs cylindrical

or ovoid, eggs multiseriate.

Parasitic on fresh-water fishes. In Africa 8 species have been found on

Polypterus, Labeo, Barbus, Clarias, Distichodus. In North America the Black-bass

and Blue-gills are attacked.

Only one record actually from South Africa.

Lernaea barbicola Leigh-Sharpe

Fig. 21 a:

1930. Leigh-Sharpe, Parasitol., xxii, p. 334, figs. 1-6.

1950. Harding, loc. cit., p. 23, fig. 89.

Locality. Transvaal, on the tail of a young Barbus. Length 7-2 mm.

The following species occur in Lake Tanganyika or Lake Nyasa, on Polypterus,

Bagrus, Clarias, Tilapia, Haplochromis, etc., and may be found in Rhodesia or the

Transvaal:

haplocephala (Cunn)., diceracephala (Cunn.), bagri Harding, tilapiae Harding,

barilii Harding, palati Harding, lopharia Harding, bistricornis Harding, tuberosa

Harding.

Gen. Cardiodectes Wilson

1917. Wilson, loc. cit., p. 50.

1928. Brian, Boll. Mus. Zool. Unww. Genova, viii, no. 26, p. I.

1934. Leigh-Sharpe, Szboga Exp. monogr., 29 b, p. 143.

1936. Markevitsch, Treubia, xv, 4, p. 407.

1937. Stekhoven, Mem. Mus. Hist. Nat. Belge, ser. 2, fasc. 9, p. 13.

1953. Capart, Bull. Inst. franc. Afr. Notre, xv, p. 665.

@ (adult)—Whole anterior surface of head covered with dichotomously

branched ampulliform processes forming a more or less spherical mass,

radiating chiefly from short horns which are continued along the lateral

margins. Neck curved, trunk straight, abdomen hemispherical. 4 pairs of

legs, the first two close together and biramous, 3rd pair uniramous, 4th pair

without rami. Ovisacs straight, elongate, eggs uniseriate.

oy)

SOUTH AFRICAN PARASITIC COPEPODA 279

The parasite penetrates the isthmus and buries its head in the heart of the

fish; but is sometimes attached to other parts of the body. Hosts: various

species of Scopelus (Myctophidae), Stolephorus (Engraulidae), and Apogon

(Apogonidae).

C. frondosus Stekhoven 1937 appears to be a synonym of bellotti (Richiardi)

and hardenbergi Markev. 1936 a synonym of rubosus L.-S. 1934. But see:

Capart, 1953.

Fig. 21. Lernaea barbicola L-S. a, 2 (copy after Leigh-Sharpe). Cardiodectes

medusaeus (Wilson); 0, 9.

Cardiodectes medusaeus (Wilson)

Fig. 21 b.

1908. Wilson, Proc. U.S. Nat. Mus., xxxv, p. 458, pl. 76, figs. 99, 100

(Lernaeenicus m.).

1912. Brian, Res. Sct. Camp, Monaco, fasc. 38, p. 27, pl. 10, figs. 1-5

(Lernaeenicus m.).

1917. (May). Stebbing, Ann. S. Afr. Mus., xvii, p. 42 (Lerneaenicus m.).*

1917. (June). Wilson, loc. cit., p. 52, pl. 3, figs. 15-23.

1933. Pesta, <ool. Anz., civ, p. 278, fig. (Lernaeenicus m.).

1950. Kirtisinghe, Parasitology, xl, p. 84, figs. 36-9.

—Two pairs of horns, the anterior flattened, the posterior pair more or less

lobate, covered with nodular processes forming a semiglobular mass resembling

a mulberry. Ovisacs about twice as long as body.

Body (excluding curved portion within host) 5-8 mm., ovisacs 13-16 mm.

Localities. Off Saldanha Bay, on Scopelus argenteus Gilch. (= hectorts Gnthr.)

(Stebbing); off Cape Peninsula and Cape Point, on Myctophum hectoris and

coccot (S. Afr. Mus.).

Distribution. Pacific coast of N. America, and off Japan; near Mauritius;

Ceylon (on Anchoviella).

* Lerne ae nicus is correct.

280 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fam. PENNELLIDAE

1917. Wilson, Proc. U.S. Nat. Mus., liii, p. 103 (Lernaeidae part, Pennel-

linae).

1932. id., Bull. U.S. Nat. Mus., no. 158, p. 489 (Pennellidae).

Gen. Pennella Oken

1815. Oken, Lehrb. Nat. (3), 1, p- 358.

1877. Wierzejski, Z. wiss. Xool., xxix, p. 562, pls. 32, 33 (larval stages).

1905. Stebbing, Mar. Invest. S. Afr., iv, p. 117 (references).

1913. Quidor, Deux. Exp. Antarct. Franc. Copep. Parasit., p. 197. (Penella, —

S2¢)\.

1917. Wilson, loc. cit., p. 105 (key to species).

1928. Leigh-Sharpe, Parasttol., xx, p. 79.

TOU. ads \ibid:) xxii1, 9. 10g:

1992.) Wilson} loc: jert., p. 460:

1932. Kirtisinghe, Parasttol., xxiv, pp. 137, 548.

1938. Monod, Bull. Trav. St. d’Aquic. Péche Castiglione, p. 3 (reprint).

1943. Heegaard, Ark. Zool., xxxiv, A 18, p. 28.

1952. Rose & Hamon, Bull: Soc. Hist. Nat. Afr. Nord, xliv, p. 172.

(Pennella varians, larval stages on Cephalopods)

Q (adult)—Head more or less globular, truncated, with 2 or 3 horns, usually

chitinous and usually unbranched. Neck long, cylindrical, passing insensibly

into the trunk which is wider, straight, and transversely rugulose. Abdomen

shorter than trunk, with a row of plumose appendages along each side, and a

pair of minute caudal rami with long setae. Antennae dorsal; antenna 1 with

few joints, antenna 2 chelate. Mouth-parts obsolete. First 2 pairs of legs

biramous, close together, 3rd and 4th pairs uniramous, all rami 2-jointed,

setose. Ovisacs filiform, several times the length of the body, eggs uniseriate.

Copepodid g—Head fused with ist segment; 2nd, 3rd and 4th segments

free and diminishing in width; 5th and genital segments fused. Abdomen

one-segmented, caudal rami short, wide, setose. Antenna 1 indistinctly jointed.

Antenna 2 2-jointed, with stout chela. Mandibles, 2 pairs of maxillae, and a

pair of maxillipeds well developed. Swimming legs like those of 9; 5th pair

absent.

Adult 99 are parasitic on Fishes (Mola, Diodon, Lophius, Exocoetus, Thunnus,

Germo, Naucrates, Remora, Xiphius, Histiophorus, and Tetrapturus) and on Whales.

Pennella is the only Copopod parasitic on Mammals.

Life-history. The larva is presumed to hatch as a typical Nauplius. At the

first Copepodid stage (fig. 22 g, h) the larva attaches itself to the gills of

Cephalopods. In subsequent stages the gg and 99 mature and fertilization

occurs. The @ then leaves the Cephalopod and seeks the second host, in which

it buries its head, seeking one of the larger blood-vessels. A large cyst forms

round the head of the parasite, and this becomes very hard and persists some

time after the death of the parasite.

Young 99 in the final stage have been described under the names Baculus

Lubbock 1860 (Tr. Linn. Soc. Lond., xxiii, p. 190, pl. 29, fig. 40) and Hessella

SOUTH AFRICAN PARASITIC COPEPODA 281

Brady 1883 (see Challenger Rep., viii, p. 136, pl. 55, figs. 9-13; also Brian, 1912,

loc. cit., infra pl. 6, fig. 10). These three figures represent successive stages

leading to the adult 2 form.

1758.

1905.

IQI2.

1913.

IQI7.

1928.

1932.

1954-

Pennella filosa (Linn.)

Big 220050.

Linnaeus, Syst. Nat., p. 819 (Pennatula f.).

Stebbing, loc. cit., p. 119 (as orthagorisct).

Brian, Res. Sci. Camp. Monaco, fasc. xxxviii, p. 16, pl. 3, figs. 2-4,

ply Oj ie ro:

Quidor, loc. cit., pl. 1, figs. 5-8, pl. 4, figs. 35, 36.

Wilson, loc. cit., p. 119, pl. 15, figs. 125-7, pl. 16, figs. 128-33,

pl. 17, figs. 134-9.

Leigh-Sharpe, loc. cit., p. 82, figs. 3, 7.

Wilson, loc. cit., p. 490, fig. 295 a, b.

Deboutteville & Nunes-Ruivo, Vie e¢ Milieu, iv, p. 215, figs. 7, 8.

Fig. 22. Pennella filosa (Linn.). a, 2; 6, dorsal view of head 9. Pennella crassicornis St. & L.

c, d, two views, dorsal and slightly oblique, of head 9; e, ventral (buccal) surface of head 9.

Pennella balaenopterae K. & D. f, ventral surface of head 9. Pennella varians St. & L. g, h, Gope-

podid stage of 9 and ¢ respectively (copy after Wilson, from Wierzejski).

—Two short stout horns projecting nearly at right angles to head, some-

times (more or less developed) a third horn medio-dorsal between the bases of

the other two. Neck varying in length; trunk about twice as wide as neck.

282 ANNALS OF THE SOUTH AFRICAN MUSEUM

Abdomen about half length of trunk, plumes branched. Ovisacs very slender,

at least twice length of body.

One of the specimens in S. Afr. Mus.: trunk 50 mm., abdomen 23 mm.,

ovisac 280 mm. Neck pale yellowish, trunk and abdomen dark slaty brown.

Locality. ‘Table Bay (Stebbing, and S. Afr. Mus.), on Mola mola.

Distribution. Atlantic, Mediterranean, east coast of N. America. Recorded

also as parasitic on the Swordfish and Tunny.

Pennella exocoett (Holten)

1802. Holten, Skr. Naturh. Selsk, v, p. 136, pl. 3, fig. 3.

1913. Quidor, loc. cit., p. 205 (in key), pl. 1, fig. 3, pl. 4, fig. 39.

1913. id:, ibid., p. 209, pl. 1, fig..1, pl. j2,. fig. 26, plaiiey fee aeeplena.

fig. 37 (louveller).

1915. Calman, West Indian Bull., xv, p. 120, pl.

1917. Wilson, loc. cit., p. 112 (in key), 115.

1928. Leigh-Sharpe, loc. cit., p. 83, fig. 4.

1954. Deboutteville & Nunes-Ruivo, Bull. Inst. france. Afr. Notre, xvi, — |

p. 163, fig. 16 a-g.

A small species, as one would expect from its host: Flying-fishes (Exocoetus).

Quidor’s specimen described as Jzouvillei measured about 37 mm. Not yet

recorded from South African waters.

Pennella crassicornis St. & L.

Fig. 22 c-e.

1861. Steenstrup & Litken, K. Dansk. Videns. Selsk. Skr. (5), v, p. 416,

pl. 14, fig. 34 (on Hyperoodon).

1912. Brian, Res. Sct. Camp. Monaco, fasc. 38, p. 18, pl. 3, figs. 5-9 (on

Mola).

1917. Wilson, loc. cit., p. 113 (in key to species).

1938. Monod, loc. cit., p. 3, fig. 1 (on Xzphius).

1954. Deboutteville & Nunes-Ruivo, Vie et Milieu, iv, p. 217, fig. 9.

In the South African Museum there are three specimens, consisting of head

and neck only, taken from the belly of a Minke (Lesser Rorqual or Piked

Whale) (Balaenoptera acutorostrata) stranded on the beach in Table Bay 1914.

The neck of the longest specimen is 90 mm. in length.

They do not correspond with the figures given by Brian for the specimens

which he assigned with a ? to crassicornis. But they are very like the figures

given by Monod for two Algerian specimens. The medio-dorsal horn varies in

length; in the specimen where it is longest it is somewhat bulbously enlarged

in its distal half. Around the buccal surface of the head are several hard, blunt

knobs varying in number and size in the 3 specimens; the central portion is

filled with spongy tissue without such definite granules or nodules as seem to be

indicated in Monod’s figure. P. louville: Quidor (1913. loc. cit.) from a

Flying-fish, as shown in pl. 2, fig. 26, appears to have a similar but more effusive

growth of knobs on the buccal surface,

SOUTH AFRICAN PARASITIC COPEPODA 283

I have not seen the original description of crassicornis, and merely record these

specimens as corresponding with Monod’s specimens.

Pennella orthagorisct Wright

1917. Wilson, loc. cit., p. 124.

Beads) id., Proc. U.S. Nat. Mus., lxiv, p. 12.

1928. Leigh-Sharpe, loc. cit., p. 81, fig. 2.

1932. Wilson, loc. cit., p. 492, fig. 295 d.

Stebbing’s 1905 description of the horns of his specimen corresponds with

those of specimens from a Mola caught in Table Bay and received by the South

African Museum in 1936. Wilson diagnoses Wright’s species as having 2 (or 3)

slender horns, longer than head and directed obliquely backwards; and

abdomen 4-2 length of trunk. Whether Wright’s species can be maintained as

distinct from filosa seems rather doubtful.

Hosts: Mola mola, Germo alalonga.

Pennella balaenopterae K. & D.

Fig, 22 7:

1877. Koren & Danielssen, Fauna Littor. Norveg, pt. 3, p. 157, pl. 16,

figs. I-Q.

1905. Turner, Tr. Roy. Soc. Edinb., xli, 2, p. 409, pls. 1-4 (anatomy).

1910. Quidor, Bull. Mus. Paris, p. 97.

Fg13id., loc.cit., p. 205 (in key), pl. 1, fig. 14, pl. 4, fig. 32.

1913. id., ibid., p. 206, pl. 1, figs. 15-18, pl. 4, figs. 29, 34 (antarctica).

1917. Wilson, loc. cit., p. 116, pl. 15, figs. 119-24 (antarctica), and pp. 112,

113 (in key).

1928. Leigh-Sharpe, loc. cit., p. 86, fig. 6.

1939. Legendre, Bull. Soc. zool. Fr., lxiv, p. 312 (comparison with

germonia).

1944. Brian, An. Mus. Argentino, xli, p. 215.

In the South African Musuem are two specimens from whales taken at

Saldanha Bay, and one from a Fin Whale stranded in False Bay. The latter

has no head, and a specimen of the barnacle Conchoderma virgatum is firmly fixed

over the vulva. The thorax measures 65 mm. and the abdomen 40 mm.

The measurements of the former two specimens are: neck 170 and 130 mm.,

thorax 42 and 40 mm., abdomen 30 and 25 mm., respectively. The ovisacs

are 180 mm. in the one, and the single sac in the other 350 mm. long.

Although Wilson keeps antarctica as a separate species, there seems every

likelihood that with further study all Quidor’s species, about which he himself

(loc. cit., p. 204) was somewhat diffident, viz.: antarctica, charcoti, anthony and

cette1, are forms of balaenopterae.

Mathews (1938. Discovery Rep., xvii, p. 126) records Penella [sic] on a

Sperm Whale caught at Durban, and (ibid., p. 238) on South African examples

of Sei-whales; but the parasite is rare.

284 ANNALS OF THE SOUTH AFRICAN MUSEUM

LERNEOPODOIDA

1913. Scott, T. & A., Brit. Paras. Copep., p. 141 (Lernaeoida part).

1932. Wilson, Bull. U.S. Nat. Mus., no. 158, pp. 492, and 613 (key to

genera).

1945. Gurney, Ann. Mag. Nat. Hist. (xi), 12, p. 121 (remarks on classifi-

cation).

1947. id., 7. Mar. Biol. Assoc., xxvil, 1, p. 133 (remarks on classification).

Body usually without movable articulations, often without any trace of

segmentation. Sexes very dissimilar, the male a pygmy clinging to the female,

which is immovably attached to the host. Antenna 2 small, but sometimes

prehensile. Maxillae 2 in 9 often modified into ‘arms’, separate or fused at

their tips, or completely fused, with a ‘bulla’ for attachment. During develop-

ment the maxillipeds sometimes migrate from their normal position until they

lie between or even in advance of the bases of maxillae 2. Eggs in the two

ovisacs multiseriate (extracted from Wilson).

Key to the South African genera (99 only)

I. Parasitic on fishes.

A. Body more or less elongate. Parasite attached to host by antennae 2,

maxillae 2, or maxillipeds, or by combination of two or more of these

appendages.

1. Antenna 2 prehensile. Maxilla 2 non-prehensile.

a. Head and trunk with paired lateral processes.

b. No lateral processes on head; only one pair at hind corners of

genital segment.

2. Antenna 2 non-prehensile. 2nd maxillae usually fused, at least at

their tips, with apical bulla for attachment.

a. Maxillipeds between bases of 2nd maxillae, both close to

mouth; 2nd maxillae longer than head, which is more or less

in line with trunk or bent forwards.

i. Trunk without processes. Abdomen distinct. Fresh

water.

ii. Trunk with 2 posterior processes ventral to the ovisacs.

No abdomen.

b. Mazxillipeds close to mouth, 2nd maxillae far removed.

Head more or less in line with trunk or bent backwards.

i. 2nd maxillae short, fused.

a. Antenna I 3-jointed; antenna 2 uniramous.

B. Antenna 1 4-jointed; antenna 2 biramous.

ii, 2nd maxillae short, separate, fused only (if at all) at

tips, with bulla.

a. Tips of and maxillae not lobate. Antenna 1

3-jointed.

B. ‘Tips of 2nd maxillae lobate. Antenna 1 4-jointed.

iii. 2nd maxillae long, separate, but fused at their tips, or

tips enlarged.

a. Bulla small.

B. Bulla large (or tips of 2nd maxillae enlarged or

branched).

B. Head and (wholly or in part) neck buried in host.

1. Genital segment with 2 short posterior processes.

2. Genital segment with long, digitate posterior processes.

3. Genital segment with 2 profusely branched posterior processes.

IJ. Parasitic on Crustacea, Body spherical, attached by 2nd maxillae and

maxillipeds,

Chondracanthus

Acanthochondria

Achtheres

Lerneopoda

Clavella

Clavellopsis

Eubrachiella

Parabrachiella

Brachiella

Charopinus

Medesicaste

Strabax

Sphyrion

Sphaeronella

SOUTH AFRICAN PARASITIC COPEPODA 285

Fam. CHONDRACANTHIDAE

1899. Bassett-Smith, Proc. Zool. Soc. Lond., p. 488 (part).

1930. Oakley, Parasitology, xxii, pp. 182 sqq.

1932. Wilson, Bull. U.S. Nat. Mus., no. 158, p. 493 (no definition).

1951. Deboutteville, Arch. Kool. exp. gen., |xxxvil, notes no. 4, p. 139

(classification).

Remarks. The Zoological Record (\xvii, 1930, Crust., p. 41) lists Chondracan-

thoides [sic = odes| and Chondracanthopsis as new genera ‘ex Wilson MS’.

Oakley (loc. cit., p. 182) says: “The descriptions of Chondracanthodes and

Chondracanthopsis were communicated to me in manuscript by Wilson.’ He

proceeds, however (p. 196), to give as genotype of the latter C. nodosus Miller

1779. As this is a well-known species the Rules of Nomenclature are satisfied,

and the genus must be credited to Oakley, although a full generic description

was not given until 1932 by Wilson.

For the other ‘n.g.’ on the other hand, Oakley gave as genotype an

undescribed species ‘deflexus’. Both genus and species were first described by

Wilson in 1932. ‘The result is thus:

Chondracanthopsis Oakley 1930, genotype C. nodosus (Miller) 1779.

Chondracanthodes Wilson 1932, genotype C. deflexus Wilson 1932.

Gen. Chondracanthus De la Roche

1811. Dela Roche, NV. Bull. Sct. Soc. Philom. Paris, 2 (44), p. 270.

1899. Bassett-Smith, loc. cit., p. 491 (part).

1922. Wilson, Ark. Zool., xiv, 10, pp. 8-12.

1930. Oakley, loc. cit., p. 186 (genotype: zez de la Roche).

1932. Wilson, loc. cit., p. 497.

1939. Yamaguti, Paras. Copep. Japan, pt. 6 (vol. Jub. Prof. Yoshida, 11),

PP- 531-3:

1943. Heegaard, Ark. Xool., xxxiv, A 18, p. 30.

—Head separated from thorax by a more or less constricted neck, with

barb-like process at each hind corner. First two thoracic segments more or less

free; other segments fused with genital segment, with paired processes.

Abdomen very small, without caudal rami. Antenna 2 prehensile. Two pairs of

fleshy biramous legs. Ovisacs multiseriate.

g—Head large, more or less fused with thorax. Caudal rami present.

Antenna 2 strongly prehensile. Mandibles and maxillipeds falcate. Two pairs

of uniramous legs.

Key to the South African species

1. Without dorsal processes. merluccit

2. With medio-dorsal crest-like processes.

a. Head longer than wide. lophi

b. Head circular. congiopodi

286

1802.

1899.

1913.

21923.

1923.

1930.

1932.

1933-

1948.

ANNALS OF THE SOUTH AFRICAN MUSEUM

Chondracanthus merluccit (Holten)

Bie. 23a Nb.

Holten, Skr. Natur. Selsk., v, 2, p. 135, pl. 3, fig. 2.

Bassett-Smith, loc. cit., p. 494.

Scott, T. & A., Brit. Paras. Copep., p. 180, pl. 20, fig. 10 (9), pl. 47,

fig. 8 (Q), pl. 53, figs. 16-19.

Wilson, Ark. Zool., xv, 3, p. 14. (Chondracanthus sp. nova 1 9.)

id., Medd. Goteb. Mus. zool. Avd., no. 19 (Goteb. Vet. Handl. (4)

xxv, 6), p. 10, pl. 1, fig. 13 (9), pl. 2, figs. 14, 15 (2) (stramineus).

Oakley, loc. cit., p. 188, fig. 2.

Wilson, loc. cit., p. 498, fig. 298 and pl. 1, fig. a.

Saby, Proc. Zool. Soc. Lond., 1933, 4, p. 865, pl. 2, figs. 7-9

(anatomy).

Barnard, Ann. Mag. Nat. Hist. (xii), 1, p. 251. |

@—Head wider behind than in front, and gibbous dorsally, a small pro-

jection at postero-lateral corner. Thoracic segment with antero-lateral

Fig. 23. Chondracanthus merluccii (Holten). a, dorsal view 9; 6, 3. Chondracanthus lophi Johnston.

c, dorsal view 2; d, dorsal profile 2; e, ventral view of abdomen 9. Inc and d the dotted lines

indicate the differences in a Plymouth specimen and the Cape specimens. Acanthochondria

lepidionis n. sp. f, ventral view 9, left antenna 1 and right antenna 2 removed; g, dorsal view of

head 9; h, 3.

SOUTH AFRICAN PARASITIC COPEPODA 287

corners slightly prominent. Genital segment with a pair of long processes

anteriorly, a ventral median knob, and hind corners produced. Abdomen

small. Basal joint of antenna 1 large.

g—Head much inflated dorsally. Attached to 2 near vulva.

2 up to 12 mm., ovisacs 14 mm., g 0-6 mm.

Localities. Agulhas Bank, on Merluccius capensis, 1 2 with 3 (Wilson); Table

Bay, on gills and floor of mouth of the Stockfish, M. capensis (S. Afr. Mus.,

22, 3d).

Distribution. Europe, east coast of N. America.

Remarks. In Ark. Zool. 1923 Wilson records a single 9 taken in Table Bay on

Gadus [sic.] capensis by Dr. Holub in 1894, and says: ‘. . . apparently the same

species as one obtained... from. . . Merluccius bilinearis, on the New England

coast. It is a new species... and will soon be published.’ I have not been able

to trace this advertised description; but as Wilson has recorded C. merluccii

from M. bilinearis in 1932, it would appear that he revised his opinion and

decided it was not a new species.

Wilson’s second 1923 record refers to a single 9 collected by Skoog in 1912

on the Agulhas Bank. There seems no reason for regarding it as distinct from

merluccit.

Chondracanthus lophit Johnston

Fig. 23 c—e.

1836. Johnston, Loud. Mag. Nat. Hist., p. 181, fig. 16.

1862. ‘Turner & Wilson, Trans. Roy. Soc. Edinb., xxiii, p. 67, pl. 3.

1899. Bassett-Smith, loc. cit., p. 494 (references).

1913. Scott, T. & A., loc. cit., p. 179, pl. 52, fig. 4 (2), pl. 56, figs. 16-18.

1927. Goggio, Publ. Staz. zool. Napoli, vill, pp. 427 sqq., figs.

1930. Oakley, loc. cit., p. 187, fig. 1, B.C.D.

1948. Barnard, loc. cit., p. 251.

9—Head longer than wide, with a lateral projection near hind corner.

Thoracic segment indistinctly bisegmented, with 2 lateral processes, the

hinder one (in Cape specimens) rounded, scarcely developed; a pair of bifid

ventral processes (legs). Genital segment constricted in middle, anterior

portion with an obscurely bifid lateral process, posterior portion with two

(dorsal and ventral) postero-lateral processes. Abdomen small, dorsally con-

cealed by a more or less elongate process. A mid-dorsal crest with more or less

developed processes on thoracic and genital segments. Antenna 1 obscurely

jointed, strongly expanded at base. Ovisacs curved or twisted.

10-12 mm.

Locality. Agulhas Bank, on Lophius piscatorius (99 S. Afr. Mus.).

Distribution. British Seas.

Remarks. In comparison with a Plymouth specimen certain differences in the

development of the lateral processes and dorsal crest were found, which

are indicated in fig. 23 c, d.

The ovisacs break up, and the nauplii hatch within a few hours (Sproston,

1942. 7. Mar. Biol. Assoc. Plymouth, xxv, p. 442).

288 ANNALS OF THE SOUTH AFRICAN MUSEUM

Chondracanthus congiopodi n. sp.

Fig. 24.

—Cephalothorax subcircular, gibbous posteriorly on dorsal surface, ‘barbs’

large, lobate. Trunk unsegmented, but with 4 lateral processes separated by

deep indents, 4 medio-dorsal knobs, a pair of posterior processes extending

beyond abdomen. Antenna 1 fusiform, 2-jointed. Two pairs of uniramous

fleshy legs, ovate or more or less boot-shaped, the ‘toe’ pointing medianwards.

Ovisacs shorter than body, stout.

95 mm., ovisac 3 mm., ¢ 0-6 mm.

Fig. 24. Chondracanthus congiopodi n. sp. a, dorsal view 9, with antenna 1 further enlarged;

b, ventral view 9; c, lateral view 2; d, maxilla, maxilliped and mandible 9; e, variations in

shape of leg 2; f, d.

Locality. ‘Table Bay, on gills of Horse-fish (Congiopodus torvus). (S. Afr. Mus.

22; $S-)

Remarks. Although the legs are not definitely bifid, there seems no reason

why this species should not be included in Chondracanthus.

Gen. Acanthochondria Oakley

1927. Oakley, Parasitology, xix, p. 466 (genotype: cornuta O. F. Miller).

TOO; ide wibid.. cai pao:

1932. Yu & Wu, Bull. Fan. Mem. Inst., 111, 4, pp. 57-71 (Chondracanthus

spp.)

1933. Saby, Proc. Zool. Soc. Lond., 1933, 4, p- 861, pl. 1 (anatomy).

1939. Yamaguti, Paras. Copep. Japan, pt. 6, vol. Fub. Prof. Yoshida, 11,

PP- 533-41.

1940. Heegaard, Vid. Medd. Dansk nat. For., civ, p. 87.

1944: ids ibid, eval.) page

SOUTH AFRICAN PARASITIC COPEPODA 289

Q9—Head separated from thorax, without barb-like processes. First 2

thoracic segments free; other segments fused with genital segment. No dorsal

or ventral processes except a pair at posterior corners of genital segment.

Abdomen very small, no caudal rami. Antenna 2 prehensile. Two pairs of

uniramous or feebly biramous legs.

Acanthochondria lepidionis n. sp.

Fig. 23 f-h.

—Cephalothorax subcylindrical, very slightly widening posteriorly, where

there is a slight constriction between it and the following thoracic segment.

Head dorsally indicated by a slightly more strongly chitinized brown area,

with a darker brown median line. A constriction between 2nd and 3rd

thoracic segments, the latter completely fused with genital segment, its identity

marked only by the position of the 2nd pair of legs. Genital segment longer

than rest of body, and wider; posterior corners rounded, shortly produced.

Abdomen very small, wider than long, with 2 small acute processes (? degene-

rate caudal rami). Antenna 1 expanded basally. Two pairs of small rudimen-

tary bifid legs.

9 4-4°5 mm., g 0°5 mm.

Locality. Off Cape Point, 300 fathoms, on gills of Lepzdion capense.

Remarks. Resembling longicephalus Yu & Wu, and pings Yu & Wu in the

long cephalothorax; the latter species has the posterior corners of the genital

segment conically produced.

Fam. LERNEOPODIDAE

1915. Wilson, Proc. U.S. Nat. Mus., xlvii, pp. 565 sqq.

1932. id., Bull. U.S. Nat. Mus., no. 158, pp. 509 and (in key) 613 sqq.

The importance of the male in delimiting the genera has long been recognized

and Wilson has very usefully brought together illustrations of 17 generic types

of male on plates 25-28 of his 1915 revision of the family.

Gen. Achtheres Nordmann

1832. Nordmann, Mikr. Beitr. wirbell. Th., 1, p. 63.

1915. Wilson, Proc. U.S. Nat. Mus., xlvii, p. 617.

1933. Gurney, Brit. FW. Copep., ili, p. 359.

@—Trunk without posterior processes. Abdomen distinct.

g—Cephalothorax in line with body-axis, smaller than the segmented trunk.

Remarks. This genus contains species parasitic on the gills of fresh-water

fishes, mostly North American. As one species is found on the Small-mouth

Black Bass Micropterus dolomieu, which has been imported into South Africa, the

genus is included here; it is possible that the parasite has been imported with

the fishes.

Achtheres microptert Wright

Fig. 25

1882. Wright, Proc. Canad. Inst., n.s., 1, p. 249, pl. 2, figs. 1-11.

1915. Wilson, loc. cit., p. 620, pl. 34, figs. 64—7, pl. 35, figs. 68, 69.

¢ 4-4'5 mm.

290

1822.

IQI5.

1918.

1932.

1939-

ANNALS OF THE SOUTH AFRICAN MUSEUM

Gen. Lerneopoda Blainv.

Blainville, 7. de Phys., xcv, p. 442.

Wilson, Proc. U.S. Nat. Mus., xlvii, p. 631 (Lernaeopoda).

Leigh-Sharpe, Parasitology, xi, p. 256.

Wilson, Bull. U.S. Nat. Mus., no. 158, pp. 617, 619 (in key)

(Lernaeopoda).

Yamaguti, loc. cit., p. 549 (Lernacopoda).

pace; one or two thoracic segments distinct. ‘Trunk ovoid with a cylindrical

Fig. 25. Achtheres micropteri Wright. a, b, dorsal and lateral views 9;

c, 3 (copies after Wilson).

or foliaceous posterior process on either side of the minute genital process

ventral to the ovisacs; no anal lamellae. Antenna 1 4-jointed. Maxillipeds

inside the 2nd maxillae, both close to mouth; 2nd maxillae long, separate, tips

joined and enlarged into a disc or with a bulla.

g—Unusually large (2-3-5 mm.). Cephalothorax in line with trunk, with a

dorsal carapace, and separated by a groove from the unsegmented trunk;

anal lamellae enlarged, turned forwards dorsally. Antenna 1 4-jointed.

Antenna 2 apically chelate.

Remarks. ‘The ventral position of the posterior processes separates the genus

from Lerneopodina Wilson 1915.

1837.

1913.

1Q15.

1923.

Lerneopoda gale: Kroyer

Fig. 26 a, b.

Kroyer, Naturh. Tidsskr., i, p. 272, pl. 3, fig. 5 af.

Scott, T. & A., Brit. Paras. Copep., p. 197, pl. 60, figs. 4-6, pl. 68,

figs. 9-15, pl. 63, fig. I (2 3).

Wilson, loc. cit., p. 635.

Monod, Bull. Inst. ocean. Monaco, no. 427, p. 5; figs 1a (2 3), 2.

Se da eee

SOUTH AFRICAN PARASITIC COPEPODA 291

®—Trunk obovate, dorso-ventrally flattened; posterior processes cylindrical.

and maxillae nearly or quite as long as trunk, with apical bulla. Ovisacs

typically rather slender and about as long as total length.

Q (trunk) 7-10 mm., 2nd maxillae 4-5 mm., ¢ 2°15 mm. (Wilson).

Locality. Kalk Bay (False Bay), in cloaca of Musielus laevis (S. Afr. Mus.

BD):

Distribution. British Seas.

oom

s pe ECR

1026 af ae 7 @

{ > VO

aese “oun

——— MALO SS

fesreens

eeee:

are

©,

SECS

Was eS

Fig. 26. Lerneopoda galei Kroyer. a, 9; b, 3 of L. bidiscalis (redrawn

from Wilson after Kane). Clavella pagelli (Kroyer). c, 2; d, 3.

Remarks. In the present specimen the ovisacs are rather stout and not quite

as long as trunk, but may not be fully developed.

The 3 here figured belongs to another species, but shows the generic

characters.

Gen. Clavella Oken

1815. Oken, Lehrb. Nat. (3), 1, p. 358.

1822. Blainville, 7. de Phys., xcv, p. 438 (Lernaeomyzon).

1832. Nordmann, Mikr. Beitr. wirbell. Th., 2, p. 53. (Anchorella, non

Cuvier).

1910. Stebbing, Ann. S. Afr. Mus., vi, p. 561 (Lerneomyzon).

1915. Wilson, Proc. U.S. Nat. Mus., xlvii, p. 666 (key to species).

1918. Leigh-Sharpe, Parasitology, xi, p. 118.

1920.,, id., 7. Mar. Biol. Assoc. Plymouth, n-s., xu, p. 332 (excretory

system).

292 ANNALS OF THE SOUTH AFRICAN MUSEUM

1925. id., Parasitology, xvii, p. 194 (revision of British species).

1932. Wilson, Bull. U.S. Nat. Mus., no. 158, p. 513.

1933. Saby, Proc. Zool. Soc. Lond., 1933, 4, p. 857 (uncinata, anatomy).

1934. Gurney, ibid., 1934, 2, pp. 196, sqq. (uncinata, development).

1939. Yamaguti, loc. cit., pp. 553-6.

1939. Poulsen, Vid. Medd. Dansk naturf. Foren., cii, p. 223 (uncinata,

biology).

—Cephalothorax long, bent backwards at an angle with axis of trunk,

with or without dorsal carapace. Trunk ovoid, without posterior processes,

abdomen, or anal lamellae; an unpaired genital process often present.

Antenna I 3-jointed. Antenna 2 uniramous. 2nd maxillae fused, usually short,

sometimes absent.

g—Cephalothorax and trunk folded together, forming an unsegmented

ovoid mass, hind end rounded, not projecting below bases of 2nd maxillae.

Antenna 1 2-jointed. Antenna 2 uniramous, 3-jointed. Maxilla 1 bipartite.

No anal lamellae.

Key to the South African species

Cephalothorax @ distinctly longer than trunk. pagelli

2. Cephalothorax 2 about same length as trunk. denticis

Clavella pagelli (Kroyer)

eo Olena.

1863. Kroéyer, Naturh. Tidsskr. (3), ii, p. 295, pl. 16, fig. 3 (Anchorella p.).

1865. Heller, ‘Novara Crust., p. 242, pl. 24, fig. 6 (A. canthart).

1910. Stebbing, loc. cit., p. 526 (Lerneomyzon p.).

1913. Scott, T. & A., Brit. Parasit. Copep., p. 224, pl. 69, figs. 1-3 (can-

thart).

1915. Wilson, loc. cit., pp. 668, 669, and 671, 672 (in key) (canthart and

pagellt).

1924. Brian, Parasitol. Maurit., p. 52 (Clavellodes c.).

—Cephalothorax distinctly longer than trunk, curved backwards, with

dorsal carapace, expanded into 2 lobes where it joins the trunk. Trunk ovoid,

usually longer than wide and width usually greater than dorso-ventral depth,

but sometimes subspherical, a small knob-like genital process. 2nd maxillae

very short, fused, forming a lobe between the lobes of the cephalothorax, with

a clavate bulla ending in a brush of minute bristles. Ovisacs about as long as

cephalothorax, cylindrical, slightly tapering distally.

g—See figure 26 d. Dorso-ventral depth greater than ‘head to tail’ length.

9 cephalothorax 3-4 mm., trunk 2 mm., ovisacs 3 mm., g (major diameter)

0-5 mm.

Locality. Cape: on Cantharus bleekeri (sic., now Pachymetopon blochi) (Heller) ;

on gills of White Steenbras (Pagellus lithognathus) (S. Afr. Mus. several 99

and g<).

Distribution. European seas, on Cantharus and Pagellus.

Remarks. T. & A. Scott include pagelli Kréy. with a ?, and adopt Heller’s

specific name. Wilson keeps the two separate in his rather unsatisfactory key.

SOUTH AFRICAN PARASITIC COPEPODA 293

There would seem to be little doubt that canthart is a synonym of pagelli.

Among the present lot those specimens with subspherical trunk have the

ovisacs shorter and more distinctly tapering than the others.

Clavella denticis (Kroyer)

1863. Kroyer, loc. cit., p. 296, pl. 16, fig. 4 (Anchorella d.).

1665. Heller, loc. cit., p. 243.

1910. Stebbing, loc. cit., p. 562 (Lerneomyzon d.).

1915. Wilson, loc. cit., pp. 668, and (in key) 672.

—(Characters from Wilson’s key.) Cephalothorax slender, linear, about

same length as trunk; latter ovoid, considerably longer than wide, narrowed

anteriorly, posteriorly truncate. Genital process present. Ovisacs thick,

ellipsoidal. 2nd maxillae?

Locality. Cape, on Dentex rupestris (Heller).

Distribution. European seas.

Gen. Clavellopsis Wilson

1915. Wilson Proc. U.S. Nat. Mus., xlvii, p. 686 (key to species).

1939. Yamaguti, loc. cit., pp. 556-62.

1950. Kirtisinghe, Parasitology, xl, p. 84.

9—Cephalothorax usually shorter and thicker than in Clavella. Trunk

squat, often wider than long, sometimes with posterior processes, but no

abdomen or anal lamellae; an unpaired genital process present. Antenna I

4-jointed. Antenna 2 biramous, endopod 1-jointed, exopod 2-jointed. 2nd

maxillae short, broad, fused, with folds of skin or swellings, with apical bulla.

g—Cephalothorax and trunk at right angles, latter strongly arched dorsally,

unsegmented, hind end projecting below bases of 2nd maxillae. Antenna 1

3-jointed. Antenna 2 biramous, each ramus I-jointed. 1st maxillae tripartite.

Remarks. Differs from Clavella in general body form, 1st and 2nd antennae,

2nd maxillae in 9, and 1st maxillae in J.

Wilson (p. 688) gives a reference to his own original description of Clavella

robusta; but the paper is not quoted in his bibliography (p. 721), and seems to

have been omitted from the Zoological Record.

Key to the South African species

1. No posterior processes. fallax

2. Two dorsal and 2 ventral posterior processes. hostilis

Clavellopsis fallax (Heller)

Fig. 27 a-c.

1865. Heller, ‘Novara’, Crust., p. 241, pl. 24, figs. 4, 5.

1924. Brian, ‘Parasit. Maurit’., p. 53.

1943. Heegaard, Ark. Zool., xxxiv, A 18, p. 32, figs. 88-94.

—Cephalothorax longer than trunk, with dorsal carapace, and a lobe on

either side at junction with trunk. Trunk dorso-ventrally flattened, broader

than long, front and hind angles rounded, genital process about }-4 length of

trunk. 2nd maxillae short, fused, lobulate, bulla mushroom-shaped. Ovisacs

stout, cylindrical, apically rounded.

294. ANNALS OF THE SOUTH AFRICAN MUSEUM

g—Dorso-ventral depth about equal to ‘head to tail’ length.

—Cephalothorax 2°5, trunk 1:5 X 2, ovisacs 2°5-3°5 mm.; ¢ I mm.

Localities. On gills of White Steenbras (Pagellus lithognathus) (loc.

unrecorded) ; East London, in mouth of Pagrus nasutus. (S. Afr. Mus, 99, 33.)

Distribution. Mediterranean, on Dentex vulgaris.

Remarks. Kurz (1877. ertschr. Wiss. Zool., xxix, p. 393, pl. 25, figs. 5, 6,

29, 51, 52) has described C. sargi which, according to Wilson’s key (loc. cit.,

p. 687), differs in having a cylindrical genital process 4 length of trunk instead

)

Weave

Fig. 27. Clavellopsis fallax (Heller). a, 2 from Pagellus; 6, 2 from Pagrus, with ¢ attached to head;

c, 6. Clavellopsis hostilis (Heller), d, e, dorsal and ventral views 9.

of a minute one, and smooth instead of wrinkled 2nd maxillae. Yamaguti

(loc. cit., 1939, p. 556, pl. 48, figs. 144-148) records sarg: from Japanese Sparid

fishes. :

In the South African specimens from Pagellus the thoracic and maxillary

lobes vary in distinctness; and the genital process in the specimens from Pagrus

is larger and more pointed than in those from Pagellus.

Clavellopsis hostilis (Heller)

Big 2700 e.

1865. Heller, ‘Novara’ Crust., p. 243, pl. 24, figs. 7, 7 a.

1915. Wilson, loc. cit., pp. 669 and (in key) 702 (? Brachiella).

1924. Brian, ‘Parasitol. Maurit’., p. 51 (Clavella h.).

—Cephalothorax curved ventrally, about as long as trunk. Head without

carapace. Trunk broader than long, tumid, with dorsal median concavity;

2 dorsal and 2 ventral posterior processes, of about equal length, the 2 dorsal

ones far apart, the 2 ventral ones approximate; a small genital process.

Antenna I 4-jointed. Antenna 2 biramous. 2nd maxillae short, entirely fused,

wrinkled, with small bulla. Ovisacs stout, apically rounded.

SOUTH AFRICAN PARASITIC GCOPEPODA 295

Cephalothorax and trunk each 1:25 mm., width of trunk 2 mm., ovisac

2-5 mm.

Locality. Natal, on gill-rakers of Umbrina robinson (S. Afr. Mus. 1 9).

Distribution. Mediterranean, on Umbrina cirrhosa (Heller).

Remarks. The present specimen appears to belong to Clavellopsis (1st and

2nd antennae), and is certainly very similar to Heller’s species. No ¢ is

present, and Heller had no g; and as Wilson remarks the 3 is necessary for

an exact generic determination. Wilson’s suggestion to transfer Heller’s species

to Brachiella seems rather a strange one.

C. appendiculata Kirtisinghe 1950, on Chirocentrus, has similar posterior

processes.

Gen. Charopinus Kroyer

1864. Kroyer, Naturh. Tidsskr., (3), ti, p. 361. (Nomencl. Preuss. Ak.,

gives date 1863; Marschall, and Neave give 1864.)

1913. Scott, T. & A., Brit. Paras. Copep., p. 188.

1915. Wilson, Proc. U.S. Nat. Mus., xlvii, p. 652 (key to species).

1928. Leigh-Sharpe, Parasitology, xx, p. 276.

1932. Wilson, Bull. U.S. Nat. Mus., no. 158, pp. 511 and (in key) 617,

610.

1939. Yamaguti, loc. cit., p. 552.

1946. Capart, Bull. Mus. Hist. Nat. Belg., xxii, no. 10, pp. 1-6, figs.

—Cephalothorax elongate, often flexed backwards, head usually without

carapace. Trunk swollen, pear-shaped, flattened dorso-ventrally, with a pair of

posterior processes dorsal to the ovisacs; no genital process, abdomen or anal

lamellae. Ovisacs long, cylindrical. Antenna 1 indistinctly 4-jointed. Antenna

2 biramous. 2nd maxillae elongate, either fused at their tips with a bulla of

varying shape, or each one apically enlarged into a chitinous bar, or knob, or

processes.

g—Cephalothorax separated by a constriction from the trunk, which is

segmented, with anal lamellae. Antenna 1 indistinctly 4-jointed. Antenna 2

biramous.

Remarks. Distinguished from Brachiella by the (usually) more elongate 2nd

maxillae in 9, and the segmented J.

T. & A. Scott (loc. cit., pp. 190, 191) describe the posterior processes as

situated ‘ventrally’, and on pl. 55, figs. 1 and 5 label ventral view as ‘dorsal’.

Parasitic in the nasal passages or spiracles, or on the gills, of sharks and

skates.

Charopinus dubius 'T. Scott

Fig. 28

1900. Scott, T., 18th Ann. Rep. Fish. Board Scotl., p. 130, pl. 7, fig. 15.

EGUg) COLL. & As loc! Cit., p:, 190, pl. 55, He. 5:

1915. Wilson, loc. cit., p. 654 (in key).

9—Cephalothorax slightly shorter than trunk, which is dorso-ventrally

flattened, pear-shaped in dorsal view, concave dorsally; posterior processes

slightly shorter than trunk. Maxilliped with very small apical spine instead of a

296 ANNALS OF THE SOUTH AFRICAN MUSEUM

claw (cf. Wilson, 1915. loc. cit., pl. 41, fig. 113. Brianella corniger). and

maxillae longer than trunk, firmly joined at their tips to a large boat-shaped

chitinous plate.

Cephalothorax 5 mm., trunk 6 mm., width 5 mm., posterior process 5 mm.,

2nd maxillae 7 mm., ovisacs 9-10 mm.

Locality. Port Elizabeth, on a skate (1 Q sent by Dr. van Hille, Rhodes

University, 1948).

Distribution. Scotland.

Remarks. The ¢ of this species is unknown.

Fig. 28. Charopinus dubius T. Scott. a, 9; 6, end view of bulla; c, antenna 1; d, antenna 2;

e, maxilliped.

Gen Brachiella Cuvier

1830. Cuvier, Régne Anim., ed. 2, iil, p. 257.

1905. Miculicich, Zool. Anz., xxviii, pp. 599 sqq-

1913. Scott, T. & A., Brit. Paras. Copep., p. 203.

1915. Wilson, Proc. U.S. Nat. Mus., xlvii, p. 698 (key to species).

1928. Leigh-Sharpe, Parasitology, xx, p. 25.

1932. Wilson, Bull. U.S. Nat. Mus., no. 158, pp. 520 and (in key) 618,

619.

1933. Saby, Proc. Kool. Soc. Lond., 1933, 4, p. 873, figs. (merluccu and

obesa, anatomy).

1939. Yamaguti, loc. cit., p. 566.

1951. Gnanamuthu, Spolia <eylan, xxvi, 1, p. 13.

@—Cephalothorax elongate, more or less flexed backwards, head with

dorsal carapace. Trunk swollen, flattened dorso-ventrally, with rows of pits,

grooves, or knobs on dorsal and ventral surfaces in some species; 2 or 4 posterior

processes and an unpaired genital process, no abdomen or anal lamellae.

Ovisacs long, slender. Antenna 1 2-4-jointed (usually 3). Antenna 2 biramous.

2nd maxillae long, usually separate and joined at tips by a bulla, but sometimes

fused.

SOUTH AFRICAN PARASITIC COPEPODA 297

g6—Cephalothorax separated by a constriction from trunk. Trunk fusiform,

narrower than cephalothorax, unsegmented, with small anal lamellae.

Antenna I 3-jointed. Antenna 2 biramous, both rami 1-jointed.

Remarks. Wilson (1915. p. 700). refers to B. lophii M. Edw. 1840, and says

it had not been seen by other investigators. Evidently he had overlooked the

Fig. 29. Brachiella sp. a, lateral view of 2 with dorsal view of

posterior processes further enlarged. Parabrachiella australis

Wilson. 6, 2; c, g.

record of T. & A. Scott (1913) who placed the species in Clavella. Moreover

Wilson in his key (1915. p. 702) characterizes lophii as having pear-shaped

‘posterior processes’; reference to Milne Edwards’s description and figure,

however, indicates that these are the ovisacs.

Key to the South African species

1. Two posterior processes; genital process short. B. sp.?

2. Four posterior processes; genital process long. macrura

298 ANNALS OF THE SOUTH AFRICAN MUSEUM

Brachiella sp.

Fig. 29 a.

Two 98, without $4, from the axil of the pectoral fin of Mugil capito, Table

Bay, are not in good condition.

Head with carapace. Trunk longer than wide, with 2 short digitiform

posterior processes and a small rounded genital process. 2nd maxillae separate,

joined at tips, with minute bulla.

Cephalothorax 1:5 mm., trunk 1-5 mm.

Brachiella macrura Wilson

1920. Wilson, Bull. Amer. Mus. N.H., xiii, p. 7, pl. 3, figs. 23-8.

1939. Brian, Rev. Zool. Bot. Afr., xxxii, p. 196.

1953. Capart, Bull. Inst. frang. Afr. Noire, xv, p. 669.

@—Trunk longer than wide; 2 dorsal and 2 ventral posterior processes of

about equal length; genital process long, about half length of posterior pro-

cesses. 2nd maxillae fused at base, then separate to the bulla. Ovisacs slender,

elongate.

Cephalothorax, trunk, and posterior processes each about 4 mm., trunk

width 2 mm., ovisacs 9 mm.

Localities. Mouth of Congo R., on gills of Snapper (NMeomaenis fulgens)

(Wilson); mouth of Congo R. (Brian): Senegal and Togo, on Ofolithus

(Capart). :

Remarks. Brian considers that this should be a synonym of chevreuxit van

Beneden 1891 (on an Elasmobranch); Capart, with some hesitation, main-

tains the two species.

Gen. Parabrachiella Wilson

1915. Wilson, Proc. U.S. Nat. Mus., xlvu, p. 713.

1932. id., Bull. U.S. Nat. Mus., no. 158, pp. 519 and (in key) 618, 6109.

1933. Saby, Proc. Zool. Soc. Lond., 1933, 4, p. 867, figs. (insidiosa, anatomy).

1945. Ringuelet, Notas Mus. la Plata, zool., x, no. 86, p. 129.

Q—Large size. Cephalothorax separated from trunk by a distinct groove. |

Trunk cylindrical, without pits or grooves, with 1 or 2 pairs of posterior pro-

cesses, and a minute genital process; no abdomen or anal lamellae. Antenna 1

4-jointed. Antenna 2 biramous, exopod 2-jointed. 1st maxillae bi- or tri-

partite. 2nd maxillae short, united only at their tips which are lobate, with

small bulla.

§—Body without trace of segmentation, no carapace, trunk swollen dorsally.

A pair of small caudal rami. Antenna 1 3-jointed.

Remarks. Distinguished from Brachiella by the groove separating cephalo-

thorax and trunk in 9, and the unsegmented and unconstricted body in 9.

Parabrachiella australis Wilson

Fig) 200), c:

1923. Wilson, Medd. Goteb. Mus. zool. Avd., no. 19 (Goteb. Vet. Handl.

(4), xxv, 6), p. 8, pl. 2, figs. 16-23 (9 Q).

9—Trunk with 2 pairs of posterior processes, and a minute genital process.

end maxillae short, tips expanded like a hand, with 5 digitate or lobate pro-

cesses (more or less separate), bulla attached to the ‘palms’.

SOUTH AFRICAN PARASITIC COPEPODA 2G9

9 trunk 6 mm., including processes 9g mm., J 3 mm.

Localities. Cape Barracouda, on gills of Stockfish (Merluccius capensis)

(Wilson); no locality, on gills of Stockfish (S. Afr. Mus. 1 9, 1 Q).

Remarks. Closely allied to znsidiosa (Heller 1865. ‘Novara’, Crust., p. 2309,

pl. 24, fig. 1), which is parasitic on Merluccius vulgaris in Europe.

Fig. 30. Eubrachiella sublobulata n. sp. a, b, lateral and ventral views adult 9; c, another specimen

2 to show knobs at base of 2nd maxillae; d, juvenile 2, with ¢ attached, to show subcuticular

lobes; e, antenna 2; f, mandible; g, maxilliped; h, g with caudal ramus further enlarged.

Gen. Eubrachiella Wilson

1915. Wilson, Proc. U.S. Nat. Mus., xlvii, p. 716.

1932. id., Bull. U.S. Nat. Mus., no. 158, pp. 618, 619 (in key).

—Cephalothorax about as long as trunk, more or less flexed ventrally, no

(distinct) dorsal carapace. Trunk stout, short, separated from cephalothorax

by a groove, no (distinct external) processes. Ovisacs stout. Antenna I

3-jointed. Antenna 2 biramous. Ist maxillae tripartite. 2nd maxillae short,

stout, separate throughout their length, or united at tips by a bulla.

g—Cephalothorax separated from trunk by a constriction, trunk posteriorly

bent forwards. Antenna 1 3-jointed. Antenna 2 uniramous.

Remarks. ‘This genus was established by Wilson to contain the two Antarctic

species antarctica Quidor 1906, and gainii Quidor 1912. The present specimens,

found on the subantarctic genus Congiopodus, appear to be congeneric.

300 ANNALS OF THE SOUTH AFRICAN MUSEUM

Eubrachiella sublobulata n. sp.

Fig. 30.

°—Cephalothorax thick, straight in juvenile, curved ventrally in ovigerous

specimens, carapace indicated anteriorly, a lateral rounded protuberance (not

very prominent) in front of 2nd maxillae. Trunk broad, but longer than wide,

dorso-ventrally flattened; a medio-dorsal bifid knob at junction with cephalo-

thorax and a medio-dorsal bifid knob (or two processes adnate at their base)

on hind margin; both these bifid projections arise from the internal tissues, and

each pair is united into a single knob by the external cuticle; similar lateral

knobs covered by the cuticle along the sides, usually in groups of three: antero-

laterally, laterally, dorso-laterally and ventro-laterally at the hind corners; a

medio-ventral trifid knob on hind margin; no two specimens are exactly alike.

Antenna I 3-jointed. Antenna 2 biramous. Ist maxillae tripartite. 2nd maxil-

lae short, stout, wrinkled, separate at base but united at tips, with a bulla; a

knob on each ventrally, and another knob-like process at their bases projecting

laterally; the latter appear to belong to the cephalothorax, but may belong to

the trunk region (see fig. d, juvenile). Ovisacs short, stout.

6—Body gibbous, with distinct constriction between cephalothorax and

trunk.

2 cephalothorax (if straightened) about 1-5 mm., trunk 1-5-2°5, gj 1 mm.

Locality. Table Bay, on gills of Horse Fish (Congtopodus torvus) (S. Afr. Mus.

29, 5d).

INCERTAE SEDIS

Gen. Medesicaste Kroyer

1863. Kroyer, Naturh. Tidsskr. (3), 2, p. 386.

1899. Bassett-Smith, Proc. Zool. Soc. Lond., p. 489.

1927. Oakley in Leigh-Sharpe & Oakley, Parasitology, xix, p. 464 (new

definition).

1932. Wilson, Bull. U.S. Nat. Mus., no. 158, pp. 495 and (in key) 614,

616.

@—Head and neck cruciform, bulla on anterior part of head. Neck long and

thin, enlarged at base. Body dorso-ventrally flattened, longer than broad,

marked with faint transverse and longitudinal grooves. Thoracic appendages

absent. Posterior processes convergent, enclosing a small 1-segmented abdomen.

Ovisacs cylindrical.

g—Very similar to ¢¢ of Chondracanthus. Antennal area marked off by a

definite constriction.

Remarks. The above diagnosis is after Oakley (1927). He says that the

general resemblance of the 9 to that of the Sphyridae is striking, and ‘but for

the shape of the posterior part of the genital segment of 9, and the general

characteristics of the 3, it would be extremely tempting to remove it to that

family’. Earlier in the same paper (pp. 457, 458) Leigh-Sharpe remarked

about Heller’s species Medesicaste capense [sic]: ‘. . . from the figure bears a

striking resemblance to a Rebelula which has lost its posterior processes. . . .

In particular the Copepod bears a strong resemblance to Rebelula bouviert

(Quidor) as described and figured by Wilson 1919 [Proc. U.S. Nat. Mus., lv,

p. 579, pl. 53, figs. 34-40, pl. 54, figs. 41-44]. One of Heller’s figures of the 9,

however, indicates articulated appendages as well as knobs, and his figure of

SOUTH AFRICAN PARASITIC COPEPODA 301

the 3, though little like those of the Chondracanthidae, is even less like those of the

Sphyritdae.’ He proposes (p. 459) that M. penetrans should be provisionally

regarded as a synonym of Rebelula bouviert.

The resemblance to R. bouviert seems to be exaggerated; the 3 and incom-

plete $ specimens described below confirm the validity of Heller’s species and

show that it cannot possibly be regarded as a Rebelula.

Fig. 31. Medesicaste penetrans Heller. a, whole animal 2; 6, ventral view of abdomen; c¢, d, dorsal

and ventral views of genital segment 2 (S. Afr. Mus. specimen) ; e, $ (a and 6 copies from Heller),

Medesicaste penetrans Heller

Riga or

1605. Heller, ‘Novara’, Crust., p. 235, pl. 25, figs. 1, 2.

1910. Stebbing, Ann. S. Afr. Mus., vi, p. 560.

Q—(description after Heller). Head transversely ovate, separated by a

constriction from the elongate cylindrical thorax which is bilobately expanded

posteriorly; followed by a more slender elongate neck. Genital segment

quadrangular, bilobed anteriorly and posteriorly; abdomen small.

(S. Afr. Mus. specimen.) The neck is broken off shortly before the genital

segment. The latter approximately as broad as long; and more rounded-

quadrate than Heller’s figure; 2 knob-like anterior processes dorsally, con-

tinued ventrally into a collar-like thickening around base of neck. The hind

margin is trilobate, the larger broadly rounded lateral lobes each with a

narrower process ventrally. The short and slender 1-segmented abdomen is

302 ANNALS OF THE SOUTH AFRICAN MUSEUM

concealed in dorsal view. ‘he dorsal and ventral surfaces of the genital seg-

ment show a number of pits between the smooth swollen margins and the

central area.

The g, attached ventrally at base of abdomen, closely resembles Heller’s

figure.

Heller: 2 28 mm., g 0-5 mm. S. Afr. Mus. specimen: 2 genital segment

4°5 xX 4°5 mm., ovisacs 6-5 mm., 9 0-75 mm.

Locality. Cape, on Trigla capensis (Heller): Kalk Bay, in mouth of Gurnard

(Trigla) 1903. (S. Afr. Mus. 1 mutilated 2 with attached J.)

Remarks. ‘There would seem to be no doubt that this specimen is referable to

Heller’s species; both were found at the Cape on gurnards.

The only other species is M. triglarum Kroéyer, which was regarded as a

synonym of asellinum (Linn.) by Scott (1913). Oakley, however, (loc. cit.)

resurrected Lernentoma Baird for the latter species, keeping triglarum in Medesi-

caste, and defined the differences between these two genera and Oralien Bassett-

Smith.

The shape of the cephalothorax of the present species can scarcely be called

cruciform; and the genital segment shows no lateral indentations as are

shown in Oakley’s figure of M. triglarum (loc. cit., fig. 7 C). These differences,

however, are not important enough to require the removal of M. fenetrans to

another genus.

Gen. Strabax Nordm.

1864. Nordmann, Bull. Soc. Imp. Nat. Moscow, xxxvil, p. 477.

1899. Bassett-Smith, Proc. Zool. Soc. Lond., p. 490.

1912. Brian, Res. Scz. Camp. Monaco, fasc. 38, p. 33.

1932. Wilson, Bull. U.S. Nat. Mus., no. 158, pp. 615, 616 (in key to

genera).

1948. Barnard, Ann. Mag. Nat. Hist. (12), 1, p. 251.

9—Body divided into 3 distinct regions. Cephalothorax enlarged, more or

less transverse to the neck-like trunk, with 6 fleshy knobs, 2 anterior, 2 ventral

and 2 posterior, head small. Trunk cylindrical; genital segment transverse

with large digitiform processes. Abdomen obsolete (unless the median dorsal

process be regarded as the abdomen). Antennae and mouth-parts reduced in

adult. Legs absent. Ovisacs cylindrical, multiseriate.

g—Pygmy. Cephalothorax inflated, hinder part of body segmented.

Caudal rami setiform. Antenna I 4-5-jointed; antenna 2 prehensile. Mouth-

parts see infra. ‘Two (or three) pairs of minute 2-jointed, uniramose legs.

Remarks. The hammer-like cephalothorax, neck, and expanded genital

segment with pits on dorsal surface (as in Rebelula and Periplexus) afford points

of similarity with the Sphyritdae. In fact it is rather surprising to find that

Strabax cannot be included in this latter family.

Oakley (1930. Parasitology, xxii, p. 185) considered that Wilson (1917, p. 34)

intended to transfer this genus to the Sphyrizdae but was deterred by the Chondra-

canthine 3. Asso few undamaged specimens were known, Oakley thought best

to leave it in the Chondracanthidae, while recognizing the extreme doubtfulness

of its position.

SOUTH AFRICAN PARASITIC COPEPODA 303

Wilson’s key (1932) is very misleading. Of the four alternative methods of

fastening to the host given in sect. 1 (loc. cit., p. 613), the present genus falls

under the second (head and neck buried in tissues of host), which leads to

sect. 60 and the Sphyriid genera.

As regards the processes of the genital segment, Bassett-Smith and Wilson

(neither of whom presumably examined an actual specimen) state that there

are eight; Brian (loc. cit., pl. 8, fig. 8) shows 9, the two ventral ones being

bifurcate, and calls them all ‘faisceaux abdominaux’. Bassett-Smith says

‘abdomen pyriform 4 as long as processes’. The present specimen, however,

leads to the conclusion that all these processes arise from the genital segment

and that the abdomen is so reduced as to be obsolete. If this be so, there are

5 dorsal processes, and 2 ventral bifurcate ones, i.e. counting the tips g in all

in the present specimen, but 11 in Brian’s figure. This raises the question

whether Brian’s specimen represents a different species, or whether the artist

has perhaps shown too many processes.

Brian gives figures of the ¢ and its appendages. The present specimen does

not fully correspond with his figures. Both antennae and the appendage

labelled by Brian ‘pmx 1’ correspond in general, but that labelled ‘pmx 2’ is

different, and I find no trace of the jointed appendage labelled ‘p 1’ (loc. cit.,

pl. 5, figs. 4 and 5). I have not seen Nordmann’s paper, but Brian says the

original description is incomplete, especially as regards the antennae and

mouth-parts. Herein lies another possibility that the present specimen may

correspond with the genotype, and Brian’s specimen represent a different

species.

A further point concerns the nomenclature of the g appendages. Brian gives

figures labelled mx, pmx 1, pmx 2, p I, p 2, p 3. Omitting the jointed appen-

dage p 1, which I do not find in my specimen, the stout appendage with bifid

unguis (pmx 1) is presumably the 2nd maxilla, and the 2- or 3-jointed appen-

dage (pmx 2) with falcate unguis in my specimen is the maxilliped.

One may also refer to the discrepancy in Wilson (loc. cit.). His fig. 298 e,

pl. 38, fig. f, and pl. 30, fig. e all represent appendages similar in structure

(2nd joint with a patch of spinules below unguis); the first is labelled ‘and

maxilla’, the second ‘maxilla’, and the third ‘maxilliped’; and the descriptions

of the first two do not correspond with their figures.

Very few specimens have been recorded, and a search for more material

would be useful. All the recorded specimens are from Scorpaenid fishes.

Strabax monstrosus Nordm.

Fig. 32.

1864. Nordmann, loc. cit., p. 18 (fide Brian), pl. 5, figs. 1-10.

1948. Barnard, loc. cit., p. 253, fig. 8 (2 3).

1954. Nunes-Ruivo. Vie et Milieu, suppl. 3, p. 118, figs. 2 (2), 3 (4).

@—Cephalothorax obliquely transverse to neck. ‘The latter shows 4 pairs of

pale longitudinal lines due to internal muscle strands; the posterior portion is

304. ANNALS OF THE SOUTH AFRICAN MUSEUM

bent and shows 2 or 3 transverse grooves, none of which completely encircle

the neck and do not represent segments. Genital segment dorsally with 3 pairs

of pits. Digitiform processes slightly constricted near the bluntly rounded tips.

¢ attached ventrally near the vulva between the ventral bifurcate processes.

4 chitinous dorsal patches on posterior segments. Antenna 1 5-jointed;

antenna 2 stout, prehensile. Maxilla 2 (?) stout, unguis with tooth on outer

margin. Maxilliped 2-jointed, slender, with apical curved unguis. Ist and

and legs very small, 2-jointed, tipped with 2 setae; 3rd leg minute.

Fig. 32. Strabax monstrosus Nordm. a, b, dorsal and ventral views 9, the latter showing attached

6; c, 6; d, 1st and 2nd antennae J.

@ ‘hammer’ about 5 mm., neck 4 mm., processes 3-4 mm., Ovisacs 5 mm.;

g 0-9 mm. Hammer and neck dark grey, the knobs pale, processes dirty buff,

ovisacs yellow.

Locality. Port St. Johns, hammer embedded in the upper corner of the gill

chamber of Scorpaenodes guamensis.

Distribution. Mediterranean and off north-west coast of Africa. On gills

and palate of Scorpaena porcus and scrofa, and Sebastes.

Fam. SPHYRIIDAE

1919. Wilson, Proc. U.S. Nat. Mus., 1v, pp. 549 sqq. (revision).

1932. id., Bull. U.S. Nat. Mus., no. 158, p. 524.

—Adult divided into 3 regions: expanded head, narrow neck, and dorso-

ventrally flattened trunk. Abdomen minute, anal lamellae present. A pair of

SOUTH AFRICAN PARASITIC COPEPODA 305

posterior processes. Ovisacs long, cylindrical, multiseriate. Legs degenerate.

In juvenile 2 pairs of antennae, the 2nd pair chelate; 2 pairs of maxillae, the

2nd pair uncinate, one pair uncinate maxillipeds. |

g—Curved or straight, unsegmented or feebly segmented. Two pairs of

antennae, the 2nd pair chelate; 1st maxillae biramous, 2nd pair uncinate;

maxilliped uncinate.

Exclusively on marine fishes. The females are usually attached near the

dorsal fin, the vent, or in the gill chamber. The female in burrowing into the

flesh of the host endeavours to reach one of the blood-vessels, usually the dorsal

aorta. The males, having found the females, become more or less permanently

attached to them.

Gen. Sphyrion Cuvier

1840. Milne Edwards, Hist. Nat. Crust., 111, p. 525.

1900. Stebbing, Mar. Invest. S. Afr., i, p. 59.

1912. Quidor, Arch. Zool. Paris, ser. 5, vol. 10, notes: pp. xxxix-xlu, figs.

1919. Wilson, loc. cit., p. 566.

1928. Leigh-Sharpe, Parasitology, xx, p. 179.

1932. Wilson, loc. cit., p. 530.

1914. Tyvold, Bergens Mus. Aarb., 1914-15, pp. 1 sqq., figs.

@—Head (cephalothorax) greatly expanded laterally, more or less hammer-

shaped, but variable. Trunk much expanded, dorso-ventrally flattened, often

pitted. Abdomen obsolete, anal lamellae knob-like. Posterior processes

profusely branched.

6—Body folded upon itself, unsegmented.

Remarks. Wilson admits only two species: Jaevigatum Guérin-Mén. and

lumpi (Kroyer).

Both the Preuss. Akademie Nomenclator and Neave give Milne Edwards as the

author of the genus. Stebbing and Wilson credit the genus to Cuvier; Stebbing

quoting the French form ‘Les Sphyrions’, Wilson the Latin form. If Cuvier

used the Latin form as well as the French, the case seems clear; but if not, the

authorship of the genus should go to Guérin-Méneville whose work is referred

to by Milne Edwards and therefore antedates Milne Edwards (viz. 1839, vide

Wilson, 1919. loc. cit., p. 568) (see Stebbing, loc. cit., p. 62).

Key to the species

1. ‘Hammer’ knobbly, neck short, trunk broader than long. laevigatum

2. ‘Hammer’ smooth, neck long, trunk longer than broad. lumpi

Sphyrion laevigatum Guérin-Mén.

Fig. 33 a.

1871. Cunningham, Tr. Linn. Soc. Lond., xxvii, p. 501, pl. 59, fig. 12

(Kingz).

1900. Stebbing, loc. cit., p. 60, pl. 4 (9).

1917. Brian, Bull. Inst. ocean. Monaco, no. 324, p. 3, text-figs. (9).

1919. Wilson, loc. cit., p. 575.

1922. Philipps, New Zeal. Ff. Sci. Tech., iv, p. 315, fig.

1928. Leigh-Sharpe, loc. cit., p. 182, fig. 3 (Kingz).

1944. Brian, An. Mus. Argent., xli, p. 199, pl. 9, figs. 76-81 (kingi).

306 ANNALS OF THE SOUTH AFRICAN MUSEUM

—-Hammer variously knobbed, no two specimens exactly alike, but knobs

more or less symmetrically arranged. Neck relatively short, shorter than width

of hammer in fully grown specimens; in one non-ovigerous specimen (S. Afr.

Mus.), however, neck twice as long as width of hammer, and even if the hammer

had grown wider the neck would have been relatively much longer than in

normal specimens. ‘Trunk in fully grown specimens broader than long.

Fig. 33. Sphyrion laevigatum Guérin-Men. a, 9. Sphyrion lumpi (Kroyer). 5, 2 from Antimora;

¢, anterior portion 2 from Cottunculoides; d, g (redrawn from Wilson).

Largest specimen: width of hammer 28 mm., length of neck 10 mm.,

width of trunk 18 mm., ovisacs 40-50 mm. In the exceptional specimen

mentioned above: width of hammer 12 mm., length of neck 25 mm., width

of trunk 15 mm.

Localities. Off Table Bay and Cape Point, near the dorsal fin or vent of

Macrurid fishes and King Klip (Genypterus) (S. Afr. Mus.); Agulhas Bank, on

‘Cape Salmon’ (probably Afractoscion) (Natal Museum); Cape Town, host

unknown (Brian).

Distribution. North Atlantic, Antarctic, New Zealand.

SOUTH AFRICAN PARASITIC COPEPODA 307

Remarks. The exceptionally long-necked specimen is interesting. It was

embedded near the vent of a Macrurid fish, but there is no obvious reason for

the length of the neck, other specimens similarly situated having typical short

necks.

Leigh-Sharpe maintains king: distinct from Jlaevigatum on account of the

nodulose (4 pairs of processes) head, the swollen neck where it joins the trunk,

and the posterior processes not being arranged in three ‘series’. I do not think

any of these characters will be found to be constant and distinctive enough for

specific differentiation. Quidor’s (1912) species should probably also become

synonyms.

Sphyrion lumpi (Kroyer)

Fig. 33 b-d.

1845. Kroyer, Danmarks Fiske, 1, p. 517 (Lestes 1.).

Ergun cc NA. scott, Brit. Parasit. Copep., p: 164,.pl. 51, figs. 3, 4:

ropa Lyvold, loc. cit., p. 12.

1919. Wilson, loc. cit., p. 570, pls. 50-2.

RO2e.) Wcigh-Ssharpe, loc: cit., p. 179, fig. 1-

HOo2 a Wilson, loc: ‘cit:, p- 530, fig. 315.

1939. Nigrelli & Firth, oologica., xxiv, pl. 1, figs. 1-3, and pls. 1-4.

1948. Barnard, Ann. Mag. Nat. Hist. (xii), 1, p. 251.

Q—(large nearly complete specimen). Hammer smooth, head prominent,

the ends of the hammer somewhat truncate and bilobed. Neck relatively

very long. Trunk pear-shaped, longer than broad.

Width of hammer 25 mm., length of neck 35 mm., width of trunk 18 mm.,

length 22 mm.

(Anterior portion only.) Two lateral processes projecting forwards enclosing

the head, which has 2 knobs on ventral surface but no indications of mouth-

parts (cf. Leigh-Sharpe, fig. 1). Length 4 mm., width 3 mm.

Localities. Off Cape Point, on Antimora australis (Gadidae) ; (anterior portion)

off Cape Point, 310-560 fathoms, behind pectoral fin of Cottunculotdes inermis.

(S. Afr. Mus.)

Distribution. Iceland (on Cyclopterus); Atlantic coast of North America (on

Sebastes).

Remarks. ‘The large specimen in the South African Museum had been named

laevigatum by Stebbing, but not recorded by him. The specimen has been

desiccated, and is not perfect, lacking the branched posterior processes. The

smooth hammer and long neck seem to indicate its reference to this species.

The specimen consisting of only the anterior portion resembles fig. 3, in

T. & A. Scott, but the horns of the hammer are relatively larger than the head.

Fam. SPHAERONELLIDAE

1897. Hansen, Choniostomatidae, pp. 1-206, 13 pls.

1910. Stebbing, Ann. S. Afr. Mus., vi, p. 562.

1930. Monod, Senckenbergiana, xii, 6, p. 336.

Minute parasites on Malacostracan Crustacea (Caridea, Mysidacea,

Cumacea, Isopoda, and Amphipoda).

308 ANNALS OF THE SOUTH AFRICAN MUSEUM

Gen. Sphaeronella Salensky

1868. Salensky, Arch. Naturg., xxxiv, p. 301.

1897. Hansen, loc. cit., pp. 4, 87 (in key), 98.

Q—Head small, more or less defined from the oval or globular trunk.

Affixed to host by uncinate maxillae and maxillipeds. Abdomen absent. Legs

small or wanting.

Parasitic in the brood-pouch of Gumacea, Isopoda, and Amphipoda.

Sphaeronella capensis Hansen

1897. Hansen, loc. cit., pp. 100 (in key), 131, pl. 5, figs. 4 a-c, pl. 6,

figs. 1 a—d.

1910. Stebbing, loc. cit., p. 562.

Size about half a millimeter.

Locality. Cape of Good Hope, in marsupium of the Amphipod Lemboides afer

(Stebb.).

List of South African hosts with the parasites recorded from them in South

African waters. In some cases the parasites recorded from other parts of

Africa are included in [_ ], but the list is not intended to be a complete list of

all the parasites recorded from each host.

MARINE

Polychaet Worms

Bispira volutacornis Sabelliphilus bispirae

Host ? Entobius euelpis

Echinoderms

Amphipholis squamata Cancerilla durbanensis

Crustacea (Amphipoda)

Lemboides Sphaeronella capensis

Ascidians

Amaroucium erythraeum Enterocola fulgens

Ascidia sydneiensis Notodelphys allmant

», canaliculata » webert

Macroclinum angolarum Botryllophilus africanus

Microcosmus oligophyllus Doropygus pulex

Polycitor reniert Botryllophilus sp.

Pyura stolonifera (“Red Bait’) Doropygus pulex

Gunenotophorus globularis

Styela hupferi Botryllophilus aspinosus

Pterobranchiata

Cephalodiscus gilchristi Kanclopus cephalodrscr

Fishes

Acanthias (see also Squalus) Achtheinus dentatus

» pinguis

SOUTH AFRICAN

Acanthopagrus berda

Alopias vulpes

Anchoviella holodon

Antimora australis

Arius dussumeiri

Atractoscion

Austrosparus sarba

Brama rai

Cantharus see Pachymetopon

Carcharinus (Carcharias)

see also Eulamia and

Sharks unspecified

Carcharodon

Cetorhinus maximus*

Chrysophrys see Austrosparus

Conger vulgaris

Congiopodus torvus

Coryphaena hippurus

Cottunculoides inermis

Cymatoceps nasutus

Dentex see Petrus

Diplodus cervinus (trifasciatus)

», sargus (rondeleti)

Echeneis naucrates

Engraulis see Anchoviella

Epinephelus fuscoguttatus

Eulamia acuta

Exocoetus species

Galeorhinus canis

Genypterus capensis

Germo alalonga

Gymnosarda pelamys

Tsurus glaucus

Lepidion capense

Lichia amia

Lophius piscatorius

* 1948. Deboutteville, Bull. Mus. Paris., xx, no

PARASITIC GCOPEPODA

399

[Caligus cossackit]

[Dinemoura producta]

Nemesis pallida

[Pie lamnal

Caligus engraulidis

Sphyrion lumpi

Caligus ari

Sphyrion laevigatum

[Caligus cossackit |

Hatschekia acuta

Pandarus bicolor

5 smith

Echthrogaleus coleoptratus

Achtheinus dentatus

Anthosoma crassum

Alebion carchariae

Nesippus alatus

Achtheinus dentatus

[Dinemoura producta]

[ Anthosoma crassum|]

[ Nemesis lamna]

[Congericola pallida]

Chondracanthus congiopodt

Eubrachiella sublobulata

Caligus coryphaenae

Sphyrion lumpr

Clavellopsis fallax

[Caligus ligusticus|

[ Clavellodes macrotrachelus|

| Lepeophtheirus longipes|

Lernanthropus peterst

Pandarus smithit

[Pennella exocoett |

Pandarus bicolor

[Pseudocaligus apodus|

Sphyrion laevigatum

| Pennella orthagorisct |

Caligus pelamydis

Anthosoma crassum

Acanthochondria lepidionis

[Caligus mauritanicus|

Lepeophtheirus lichiae

[ Nemesis lamna]

Chondracanthus lophit

- 5, Pp. 446, 447.

ANNALS OF THE

Macrurus species

Merluccius capensis

Mola mola and lanceolata*

Muzgil species

Mustelus species

Myctophum species

[ Neomaenis (Congo)

Orthagoriscus see Mola

Ostracion cornutus

Otolithus

Pachymetopon blochir

Pagellus lithognathus

5) mormyrus

Pagrus see Cymatoceps

Pelamys sarda

Petrus rupestris

Pliotrema warrem

Plotosus anguillaris

Polynemus

Pomatomus saltator

[ Pseudotolithus

Raia marginata

Rhinobatus

Sargus see Diplodus

Sciaena aquila

Scoltodon

Scopelus see Myctophum

Scorpaenodes guamensis

Scylliorhinus (Scyllium)

* 1946. Dollfus, Essai de Catalogue des Parasites Poisson-lune Mola mola (L. 1758) et autres

Molidae. Ann. Soc. Sci. Nat. Charante-maritime, n.s. III, fasc. 7. (Copepodes, pp. 70-2.)

SOUTH AFRICAN MUSEUM

Sphyrion laevigatum

> lumpr

Chondracanthus merluccti

Parabrachiella australis

Lepeophtheirus insignis

Cecrops latreillea

Orthagoriscicola muricatus

Philorthragoriscus serratus

Pennella filosa

3 orthagorisct

Achtheinus dentatus

Lernanthropus paradoxus

Brachiella sp.

Achtheinus dentatus

Lerneopoda galei

Cardtodectes medusaeus

Brachiella macrura]

Ancistrotos ostracionis

Brachella macrura

Clavella pagelli

Clavella pagellt

Clavellopsis fallax

[Caligus ligusticus]

[Clavellopsis fallax]

Caligus pelamydis

Clavella denticis

Achtheinus pinguts

Lepeophtheirus plotose

[Argulus dartevellet |

[Caligus mauritanicus |

Argulus alexandrensis|

Trebius caudatus

Lepeophtheirus sp.

[Caligus mauritanicus |

[ Lepeophtherrus longipes |

[Sceaenophilus tenuis]

| Polyrhynchus sciaenae|

[ Lernanthropus gislert |

[ Brachiella sciaenophila]

Perissopus dentatus

Strabax monstrosus

Pandarus armatus

Achtheinus dentatus

>» pingurs

SOUTH AFRICAN PARASITIC COPEPODA

Seriola lalandet Caligus lalandet

Serranus goliath see Epinephelus

Sharks unspecified Alebion carchariae

Perissopus dentatus

Dinemoura producta

ms latifolia

Achtheinus dentatus

Skate unspecified Charopinus dubius

Sparus see Acanthopagrus

Sphyraena commersont Argulus belones

[Caligus affinis]

Pandarus cranchu

Sphyrna species Pandarus armatus

5 cranchi

[ Nemesis robusta]

Squalus acanthias Caligus coryphaenae

Achtheinus dentatus

Stegostoma fasciatum Pandarus cranchit

Temnodon see Pomatomus

Tetrodon hypselogeneion Caligus tetrodontis

Lepeophtheirus brachyurus

Thunnus thynnus [Pennella filosa|

Trigla species Medesicaste penetrans

Umbrina robinson Clavellopsis hostilis

Aiphias gladius [Pennella filosa|

9 -« Crassicornis |

Keus [Argulus zei|

Kygaena see Sphyrna

Mammals

Balaenoptera acutorostrata Pennella crassicornis

» Species 5, balaenopterae

FREsH WATER

Amphibians

Tadpoles Dolops ranarum

Fishes

Barbus sp. Lernaea barbicola

Barbus gunningt and swierstrae Dolops ranarum

Cichlidae [Chonopeltis inermis]

[ Lernaea]

[ Lamproglena]

Eutropius Dolops ranarum

Gnathonemus Chonopeltis inermis

Heterobranchus [Dolops ranarum]

Huro salmonoides (imported) Dolops ranarum

Hydrocyon [ Argulus]

[ Lates Dolops ranarum|

Marcusenius Chonopeltis inermis

312 ANNALS OF THE SOUTH AFRICAN MUSEUM

[ Micropterus dolomieu (imported)

[ Polypterus

Protopterus

Sandelia capensis

Serranochromis thumbergi

Synodontis melanostictus

Tilapia mossambica and other species

Achtheres microptert |

Lernaea haplocephala|

| Dolops ranarum|

Argulus capensis

[ Lamproglena|

Chonopeltis inermis

Dolops ranarum

[ Lernaea]

11. The Breeding and Growth of Hymenosoma orbiculare Desm. (Crustacea,

Brachyura). By G. J. BRoEKHUYSEN, PH.D., Department of Zoology,

University of Cape Town. (With 13 text figures.)

INTRODUCTION

The growth and reproduction of the South African shore crab Cyclograpsus

punctatus was described earlier (Broekhuysen, 1941). The present paper deals

with the Crown crab Hymenosoma orbiculare.. Whereas Cyclograpsus lives on the

upper part of the shore and is common among broken rocks, Hymenosoma lives

on the lower part of the shore and is restricted to quiet areas where fine sands

and mud accumulate. It is thus common in lagoons and estuaries. Cyclograpsus

can withstand exposure to air for considerable periods and prefers this, but

Hymenosoma either buries itself in damp sand covered by a shallow layer of

Fig. 1. Hymenosoma orbiculare Desm.

Large specimens from the Sand Vlei Estuary, natural size. Left: male; right: female.

Byes)

314 ANNALS OF THE SOUTH AFRICAN MUSEUM

water or moves down the shore with the ebbing tide and often extends below

tide marks. Thus Barnard (1950) has recorded it from 45 fathoms. Neverthe-

less there are certain similarities between the two crabs: both tolerate a

very wide range of temperature and salinity.

The ecological niche occupied by H. orbiculare shows much similarity to that

occupied by Carcinus maenas of the Northern hemisphere (Broekhuysen, 1936).

Carcinus, however, seems to prefer a more solid bottom than HAymenosoma,

although the species was

very common in the Wadden

Sea, N.W. of Holland,

where the bottom is sandy.

Hymenosoma orbiculare

Desm. has been described

by Barnard (1950). The

adult ¢ and 9 are illustrated

in fig. 1; and in fig. 2 an

example of the first crab

(post-larval) stage. During

the course of the estuary

survey carried out by the

University of Cape Town,

the species has been

recorded from the following

localities: Lambert’s Bay,

Steenberg’s Cove, Berg

Fig. 2. Hymenosoma orbiculare Desm. River Mouth, Saldanha Bay,

First crab (post-larval) stage. A Megalopa stage Langebaan Lagoon, Mil-

has not been observed. nerton Estuary, Hout Bay,

Sand Vlei Estuary, False

Bay, Klein River Lagoon, Breede River, Great Brak River Mouth, Knysna

Lagoon, Sundays River Mouth, Bushman River Mouth, Keiskamma River

Mouth, The Haven, Port St. Johns, Durban Bay, St. Lucia Bay, Kosi Bay,

estuaries near Inhambane (Portuguese East Africa). Balss recorded it from

Liideritzbucht and Barnard from Olifants River Mouth, so that its total dis-

tribution is from South West Africa around the Cape to Portuguese East Africa

(see fig. 3). Also recorded from Zanzibar (Lenz, 1905).

Hymenosoma geometricum Stimpson was described from False Bay. Barnard

(1950) considers this to be merely a deep-water variety of H. orbiculare. ‘The

identity of the two species will be discussed further in the present paper.

MATERIAL AND METHODS

The bulk of the material was collected in Sand Vlei Estuary at Muizenberg

between the Vlei Bridge and the Foot Bridge (see fig. 4). During the course

of the investigation the estuary was subjected to wide variations in salinity

BUD)

THE BREEDING AND GROWTH OF HYMENOSOMA ORBICULARE DESM.

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ANNALS OF THE SOUTH AFRICAN MUSEUM

316

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THE BREEDING AND GROWTH OF HYMENOSOMA ORBICULARE DESM. 317

indicated by a limited number of salinity determinations which have been

tabulated in Table I.

Table I

Salinities determined in the Sand Vlei Estuary during the course of the

investigation (see for stations fig. 4). The figures with an asterisk were bottom

sample readings.

Station || March| April May June July | August | September | December | January

I = |8'31-9'0!) 7°4-7°6 | 3°3-31°8 | 075-58 = 33°7-34°3 = rae

2 ay 8-4. | F2-T5) | 28-2570 104-577 | 0-5-0'6* |.8:4-32°3 | 24-2 20°1

3 8-9 79 | 7°2-7°5 | 0°8-25:2 | 0-4-5°3 0:65) 0:0-21-2 21°4

4 g°2 3°3-8°2 | 2:0-23'1 1°5—-28°9 21°5

4°8*

5 2°8 3°3

The salinites given in the above table are in 9/9.

Random samples of crabs were collected at regular intervals from February

1947 to February 1948. The total was 1,494 females and 867 males. After

sexing, the maximum width of the carapace was measured. The degree of

maturity was judged from the shape of the abdomen, and the adults were

dissected to determine the developmental stage of the gonads. Berried females

were recorded and the developmental stage of the developing embryos noted.

During the course of the investigation a considerable number of Hymenosoma

were kept in the laboratory. They were in glass dishes with a little water.

Sand and shell fragments were provided as a substratum. These crabs were

under constant observation.

THE REPRODUCTIVE STAGES OF THE MALE

A total of 752 males collected from Sand Vlei had a carapace breadth

exceeding 14 mm. By dissecting, three stages could be distinguished in the

reproductive system: (a) gonads undeveloped or small; (5) gonads developing;

The monthly changes in the percentage of crabs which fell into these three

categories is shown in Table II and text-figure 5.

Table IT

Developmental Stages of the Gonads of Males from Sand Vlei during the

period February 1947 to February 1948 (figures given are percentages).

318 ANNALS OF THE SOUTH

AFRICAN MUSEUM

Date Gonads undeveloped | Gonads developing | Gonads well developed | Total number of crabs

February I 38 61 94

March 5 26 69 175

April 4 13 83 78

May 10 24 65 78

June 4 26 70 23

July ) 28 72 25

August 2 37 61 41

September - - - I

October ~ - - 3

November 67 2 31 52

December 21 54 26 39

January 8 4! 51 143

No records are available for September and October as the water-level in

the estuary was so high that collecting was difficult. It will be seen that most

of the males had fully developed gonads between February and August. In

November (and possibly October) they had small or resting gonads and in

December to January they were growing.

It therefore appears that during spring and early summer most males were

unable to copulate as their gonads were not mature. The breeding-season was

in late summer and in winter, and covered a period of eight months. It must

be stressed that these conclusions refer only to crabs in Sand Vlei Estuary and

UNDEVELOPED

Ej DEVELOPING

UU] WELL DEVELOPER

PERCEN TAGE

Re ad 8 SS = a, OE a a ees

Ge Gy ne ey = 2 ae ie eg OS ae Fa)

bE SS Sea ee Osan

Fig. 5

The development of the gonads in males. During

the months September and October too few males

were obtained to make observations possible.

differences may well occur

in crabs inhabiting other

localities. Thus’ ‘of 25

mature males caught at

Lambert’s Bay in October

1947, 4 per cent had un-

developed gonads, 12 per

cent developing gonads,

and 84 per cent well-

developed gonads. As will

be seen later, the develop-

mental cycle of the male fits

in rather well with that of

the female.

THE REPRODUCTIVE STAGES

OF THE FEMALE

A total of 1,314 females

with a carapace width

exceeding 14 mm., were

examined and of those

which were not in berry,

THE BREEDING AND GROWTH OF HYMENOSOMA ORBICULARE DESM. 319

687 were dissected and the state of the gonads recorded in the same three

categories as was used for the males, namely: (a) ovary undeveloped or

small; (bd) ovary developing; (c) ovary well developed.

A total of 327 females were in berry, so that this represents a fourth phase

in the reproductive cycle. In order to determine whether these berried females

had resting or active gonads, 245 of them were dissected and their condition

recorded under the same categories as before.

The results are summarized in Tables III and V and text-figures 6 and 7.

The upper part of text-figure 6 shows the percentages of berried females, while

the lower part gives the percentages of dissected females not in berry with

gonads in one of the three different developmental stages.

Table II

The activity of the reproductive organs of female crabs at Sand Vlei during

the period February 1947 to February 1948.

| Total number

Date Females not in berry | Females in of crabs

] berry examined

scree | eee) f

Number of

Undeveloped | Developing Well- females larger || Percentage of

ovaries ovaries developed than 14.mm. | females in

ovaries dissected | berry

% % % | %

12°2. 747 71 28 I 93 I 136

ee 96 4 oO 112 oO 152

253 100 O O 47 ) 67

15°4 100 O ) 37 O 40

6°5 68 30 I 47 O 50

27°5 3 26 71 35 52 67

6-6 II 56 33 9 33 27

19°20.6 44 22 BB 9 64 25

18-7 17 ) 83 6 85 39

30°7 ia 33 66 12 78 54

12°8 13 25 63 8 79 39

26°8 8 25 67 12 76 51

23°9 33 33 oe 3 94 54

13°10 10 20 70 10 55 22

13°11 25 33 42 12 20 25

25711 54 31 15 52 30 7

16°12 64 29 7 41 10 49

g't. 48 93 6 I go 4 243

23°1 96 4 Oo 82 O 97

Table III and text-figure 6 show that there is a definite periodicity in the

reproductive cycle of the female. The breeding-season is in the winter and the

spring. During summer the ovaries are in a resting stage, but by May

development starts to take place.

320

ANNALS OF THE SOUTH AFRICAN MUSEUM

80 ee

aaa (r

FEMALE S

IN BERRY

3 °F ia Hl

20}

Te! |

e . e s

e = e ° Ld s }

| Le ©

*.*.1 DEVELOPMEN-

:."+).1 TAL STAGES

-"*:.1 OF OVARIES

Pore a Ue

epee iinnin

PERCENTAGES

Faults

| UNDEVEL. OVARIES

c===| DEVELOPING OVARIES

(TT) WELL DEVEL. OVARIES

Fig. 6

The development of the gonads in females. The upper half gives the percentages of females

in berry, while the lower half represents the percentages of dissected, not in berry, females

with gonads in different developmental stages.

THE BREEDING AND GROWTH OF HYMENOSOMA ORBICULARE DESM. 321

While the investigation was carried out plankton obtained in plankton

netting in the Sand Vlei Estuary was also examined for the presence of larval

stages. The results are given in Table IV.

Table IV

The occurrence of Hymenosoma zoeae in Sand Vlei Estuary.

| |

Date June July | August September | October | November | December

Quantity some |some-many; some many many few-some few

As could be expected the presence of zoeae coincided with the months of the

year that females were in berry.

As the breeding-season falls mainly in the winter, it coincides with the

months of heavy rain and consequently Sand Viei fills up with rain water.

This eventually causes a strong current which flows into the sea so that large

numbers of zoea larvae are lost to the estuary. As will be mentioned later the

outflowing water will have a salinity too low for the zoeae to remain alive.

Table V

Analysis of samples of mature females from other localities.

Total number

of females Not in berry In berry

Lambert’s Bay 226 = 100% yh = Bue 142)— 6307,

October 1946 of these:

18% with undeveloped ovaries.

35% with developing ovaries.

48% with well-developed ovaries.

Lambert’s Bay 246 = 100% TO2— OG

December 1946 Ofir7-

72% with undeveloped ovaries. 54. = 22%

15% with developing ovaries.

14% with well-developed ovaries.

Hermanus 160 = 100% 142 = 88% LOne—s iy,

September 1947 of137:

2% with undeveloped ovaries.

37% with developing ovaries.

61% with well-developed ovaries.

Berg River Mouth || 94 = 100% 36 = 38% 58 = 62%

September 1949 of 24:

54% with well-developed ovaries.

* An additional 74 (46%) had still traces of hatched eggs.

322 ANNALS OF THE SOUTH AFRICAN MUSEUM

Although there is a possibility that the tide sometimes may carry zoeae

larvae into the estuary, those swept out will outnumber those swept in. The

maintenance of the Hymenosoma population in the estuary must be in great part

due to the limited number of zoeae which hatch when the mouth of the estuary

is blocked by a sandbank, or when the mouth is still open while very little water

flows out and the tide enters at high water. The possibility of crabs invading

the estuary from False Bay cannot be completely ruled out.

As has been mentioned previously in addition to the crab material which

was collected at Sand Vlei some was collected from other localities. Hymeno-

soma was extremely common in a small blind lagoon at Lambert’s Bay, and in

October and December 1946 material was collected and examined. In

September 1947 material was collected from the large Hermanus lagoon and in

September 1949 crabs were collected and examined from the Berg River

Mouth.

The results are given in Table V. (See previous page.)

Many crabs were also kept in the laboratory. The salinity of the water

in which these were kept varied from 2°7°/9) to 30:9°/o9. This variation

is not abnormal because under natural conditions the species occurs in

places where large variations in salinity do take place. The temperature

varied between 13° C. and 15° C. In seventy-one cases these crabs extruded

ege-batches. In Table VI these have been tabulated according to the months

in which they occurred.

Table VI

Egg-batches extruded by 34 crabs kept in the laboratory.

Month Number of | Percentage of total Month Number of | Percentage of total

egg-batches | number of egg-batches egg-batches | number of egg-batches

January 2 2°8 July II 15'5

February 4 5:6 August 15 21°3

March I 1-4 September 7. 9°9

April I 1°4 October 13 18°3

May I 1-4 November 8 II3

June 4 56 December 4 56

The information given in Table V indicates that the breeding-season for

Hymenosoma females from Lambert’s Bay, Hermanus, and the Berg River

Mouth is similar to that from the Sand Vlei Estuary crabs. Table VI shows

that under the stated laboratory conditions, egg-laying could take place in any

month of the year, but mainly occurred from July to December. This also

agrees with what was found at the Sand Vlei Estuary (see text-figure 6).

THe RELATIONSHIP BETWEEN THE GONAD ACTIVITY OF THE Two SEXES

If text-figures 5 and 6 are compared, it is clear that the short resting-period

of the gonads of the males (from October to December) coincides with the

THE BREEDING AND GROWTH OF HYMENOSOMA ORBICULARE DESM. 323

last half of the breeding-period of the females. This may be expected, because

copulation during the egg-carrying period must be down to a minimum. A

similar state of affairs was found to exist in Cyclograpsus punctatus (Broekhuysen,

1941).

NUMBER OF EGG-BATCHES EXTRUDED BY EACH FEMALE DURING A SINGLE

BREEDING-SEASON

In order to ascertain whether [===] UNDEVELOPED

one female produces more than E=j DEVELOPING

A : (00 WELL DEVELOPED

one batch of eggs in a breeding-

A total of 245 females in berry

were dissected and the results

100

season, berried females were

dissected and the state of the

gonads recorded. If only one ied

egg-batch was produced, the

gonads of berried females could 60 :

be expected to be in a resting e aes

stage; if, however, more than one AG O me

<q BOO

batch of eggs was extruded, the = ee

gonads could be expected to be 9 a8

in various stages of development. eo ee

° ° : >

is graphically expressed in text- eS POLS OI? :

figure 7. Fig. 7

Developmental stages of the gonads in females

Table VII in berry.

State of development of ovaries of berried females.

% females with % females with % females with Total number of

Date undeveloped ovaries | developing ovaries | well-developed ovaries crabs

May 04 6 fe) 16

June 24 77 O 13

July 39 41 20 54.

August 39 42 19 62

September 53 31 16 51

October 83 17 Oo 12

November 54 43 3 37

Table VII and fig. 7 show that in May most of the berried females had

undeveloped or resting gonads. During the rest of the breeding-season (June

to September) the ovaries were in various stages of development while the

crabs were carrying developing embryos. This is strong evidence that more

324 ANNALS OF THE SOUTH AFRICAN MUSEUM

than one batch of eggs is produced per breeding-season. As shown below

(Table VIII) these results were confirmed by observations on crabs kept in the

laboratory.

Table VIII

The interval between the hatching of one batch of eggs and the laying of

the next in between consecutive moultings in laboratory cultures.

Interval in days \ Number of cases || Interval in days | Number of cases || Interval in days | Number of cases

9 2 17 3 36 I

10 I 18 2 39 I

II 2 19 I 42 I

12 2 21 2 43 2

14 2 32 2 48 I

15 2 33 2

Average interval 23 days

From Table X the average incubation time of eggs kept under the stated

laboratory conditions appears to be 43 days. The average interval between

batches was 23 days (see Table VIII). ‘The total time from the attachment of

one egg-batch until the attachment of the next during the breeding-season

was therefore about 66 days. The breeding-season during the period of the

investigation lasted from May until October covering approximately 150 days.

Although information obtained on crabs kept in the laboratory under controlled

conditions may differ to some extent from what is the case for crabs under

natural conditions, the possibility of two to three batches of eggs in one breeding-

season may be provisionally accepted.

THE DEVELOPMENT OF THE EGGs IN THE EGG-BATCHES

In the eggs of the marine Brachyura the following ten developmental stages

can be distinguished by external examination of the living eggs:

Stage I. No segmentation has occurred and no external cleavage can be

detected.

Stage II. The first cleavages have taken place.

Stage III. A considerable number of cleavages have taken place.

Stage IV. The first indication of invagination has become visible.

Stage V. A distinct division between a yolk-free and a yolk-containing

part can clearly be seen. This stage covers the whole period

between the first indication of the forming of the germ-layers and

the formation of the eye-pigment.

Stage VI. The first indication of the eye-pigment is externally visible.

Stage VII. The first indication of the chromatophores which will form

pigment bands has become visible.

THE BREEDING AND GROWTH OF HYMENOSOMA ORBICULARE DESM. 325

Stage VIII. The larvae are clearly pigmented, a fair amount of yolk remains,

but is becoming reduced in quantity.

Stage IX. The yolk has nearly disappeared, the egg-shells are rupturing,

and the larvae are emerging.

Stage X. Only dead eggs and empty egg-shells remain.

Fig. 8. Hymenosoma orbiculare Desm.

Left: first zoea stage; right: last zoea stage.

As in Cyclograpsus punctatus (Broekhuysen, 1941) it is the first zoea stage

which escapes from the female abdomen (fig. 8, left).

During the period May 1947 to the first week of January 1948 a total of

323 ege-batches of berried females were examined and the stage of development

of the embryos recorded. The results are given in Table IX. Individual

variation in the development of one batch was slight and this was also found

to be the case in other marine crabs (Broekhuysen, 1936, 1941). The duration

of each stage was not the same and the laboratory experiments showed that

the duration from stage I to beginning of stage [IV was approximately the

same as from stage IV to beginning of VII and stage VII to IX. In text-

figure g stages I to III have been lumped together and the same has been done

for IV to VII and VII to IX.

326 ANNALS OF THE SOUTH AFRICAN MUSEUM

Table 1X

The development of attached embryos during the 1947 breeding-season

expressed in percentages.

Dead | Total number

Date®|| Stage ds| “Al |) HT | Va OVE SEE | VTE eggs of crabs

Yo |, Yo yor il feral azo: |e: |) ou ||) ven ||) ol eran a

27°5 3 97) S25 4) 19 Wag 6 fo) 6 oO oO 32

6-6 co) 1G al eie2e| arora! 4d 6 fo) 6 ) fo) 17

20°6 Oo CONmm Tf ot Qe aoe Gotha Oo Oo o | 56 9

18-7 O o | 16 B25 S20 25 fo) O 32

30°7 O Oe i A | eI sg Oy HO as fo) Oo 2 42

12°8 13 3 13 30) tO 3 ste) oO oO 30

26:8 5 5 | 21 5 24 A 5) 5, || 24 Por aes 3 38

23°9 4 Anil aLA: 4S 6 6 Oo | 49 2 (0) 2 51

13°10 17 17 25 8 8 O 8 8 oO (o) 8 12

13°11 22 DE 50. Wy 21 fo) O ) ) fo) ) 9

25°11 Oo 3 6 fo) 8 6 6 | 69 ) 3 36

16-12 O 20 13207732017 40 O Oo Oo Oo oO 5

gl Oo 0) Oo fo) fo) Odi 10.490 (0) O 10

Fig. g indicates that the majority of females produced two and perhaps three

batches of eggs during the breeding-season of 1947 at Sand Vlei. This, there-

fore, confirms what has been said on page 324. Fig. 9 also indicates a con-

siderable individual variation.

INCUBATION TIMES OF EGGS KEPT UNDER LABORATORY CONDITIONS

As has been mentioned previously crabs were kept in the laboratory while

the field-work was in progress. The salinity of the water in which the crabs

were kept was known at the beginning and the water-level was marked on the

glass. At intervals distilled water was added to compensate for evaporation.

At the end of the experiment (in some cases lasting over a year) the water in

some of the dishes was titrated and the salinity determined. In this way a

rough estimate of the salinity conditions during the experiment was obtained.

As will be shown further on, the developing eggs of Hymenosoma orbiculare are

rather euryhaline and the limited salinity changes of the water in the dishes will

probably not be very important. The glass dishes containing the crabs were

kept in a controlled temperature room. Most of the time the temperature was

13° C. but sometimes the temperature went down to 12° and up to 15° for a

short time. The development of 43 batches of eggs extruded by crabs kept

under these conditions was studied. The incubation time of all 43 batches was

determined with an accuracy of two days. The results are given in Table X.

THE BREEDING AND GROWTH OF HYMENOSOMA ORBICULARE DESM. 32°77

27 MAY IS NOVEMBER

6 JUNE

28 NOVEMBER

r: JUNE is

Ii6 DECEM

18 JULY BER

12 AUGUST : 9 JANUARY

26 AUG.

a . a a ae

: OCTOBER

il Ww Wik

DEVELOPMENT STAGES

SEAT aane

fe)

PERCENTAGES

Fig. 9

The development of the attached embryos during the 1947 breeding-season expressed in

percentages and three groups of development stages each of which takes about the same time.

(The observations for 30 July are not graphed as they are practically the same as for 18 July.)

328 ANNALS OF THE SOUTH AFRICAN MUSEUM

Table X

Incubation time of eggs under laboratory conditions.

Maximum Maximum

temperature salinity range Incubation time in days Average

range

(Op ss

12—13 1529 49 49

12—I4 14——29 43, 44, 47 45

15—29 42, 43, 44, 44, 44, 44, 44, 45, 45, 47, 48, 48 45

16—29 44 44

I2—I5 14—29 40 40

15—29 43, 45 44

13—14 14——29 43 43

I15—29 39, 49, 41, 41, 42, 42, 42, 44, 44 42

16—29 38, 39 39

13—I5 I4——I9 41, 42 42

15 41 41

I5—29 37, 38, 40, 41, 42, 43, 45 41

14 15—29 38 38

Average of all observations 43

The figures in the above table show that 43 days was the average incubation

time of the eggs of H. orbiculare at a temperature between 12° C. and 15° C.

The figures also suggest that the more constant the temperature, the shorter

the incubation time. A greater amount of variation of the temperature seems

to increase the incubation time, as was also found to be the case for Cyclograpsus

punctatus (Broekhuysen, 1941, p. 344).

In July 1947 a female with a carapace width of 18-7 mm., kept in a glass

dish without any sand or shell fragments, extruded eggs which got attached to

the hairs of the pleopods in the normal way. The presence of a soft sub-

stratum, therefore, seems not to be essential for a normal attachment of the

eggs in this species. In this respect H. orbiculare seems to be different from

C. punctatus and C. maenas (Broekhuysen, 1936, 1941) in both of which the

extruded eggs did not get attached if sand and shell fragments were not

supplied.

In laboratory cultures there were twelve cases of two batches of eggs being

produced which developed normally although copulation had taken place

only once, i.e. before the first batch of eggs. In two cases, three batches were

produced after one copulation; the eggs of the first two batches developing

normally while those of the third batch developed abnormally and never

produced zoeae. There were four cases where three batches were extruded

after one initial copulation and the eggs of all three batches developed normally

into zoeae.

This shows that a female H. orbiculare can produce as many as three normal

batches of eggs in succession although fertilized only by one initial copulation.

THE BREEDING AND GROWTH OF HYMENOSOMA ORBICULARE DESM. 329

In some cases, however, the eggs of the third batch were not properly fertilized.

The fact that one copulation suffices for the fertilization of more than one

batch of eggs has also been found to hold good in other Brachyura (Gosse,

1852; Williamson, 1900; Churchill, 1917-18; Duncker, 1934; Broekhuysen,

1936, 1941).

In all the cases where small immature female crabs were put into the glass

dishes and reared apart from males, the eggs developed only into an irregular

cleavage stage and then died. In one instance four egg-batches were produced

by such a female during a period of thirteen months. In this case none of the

eges developed further than into an irregular cleavage stage. In some cases

females which had just moulted when caught were put into a glass dish. After

some weeks these females moulted again, but as no male was in the dish,

refertilization could not take place. Nevertheless the egg-batches produced

later contained fertilized eggs, showing that the moulting process had not

affected the spermatozoa stored in the female.

These cases also show that copulation is not essential for extrusion of eggs.

The time between copulation and egg-extrusion varies considerably as can

be seen from Table XI. ;

Table XI

The interval between moulting and the next egg-extrusion in the crabs kept

in the laboratory.

Copulation took place after the moult No copulation took place

Date of Date of Date of Interval in Date of Date of

moulting copulation egg-laying days moult ing egg-laying Interval

14°3 | 17°3 315 77 25°3 3°6 59

30°4. 30°4. about 54

24:6 1°4 6:8 126

22°5, 22°5 57 43 6-4 1°7 85

255 ois 78 73 13°4 6-7 83

19°6 21-6 29°7 39 27°4. 19°6 52

21°6 21°6 27°83 66 27°4. 10°8 104

3°38 a 7°10 64 28-4. 25°7 87

3°38 ? 22°10 79 6:5 about 57

2:7

11°8 11°8 8-10 57 12°5 10°8 89

11°8 11°8 26-11 106 155 4°7 49

22°8 22°8 about 84 155 18-7 63

1511

Q°Il II‘ll 28°1 78 19°5 18°8 90

Q'Il II‘1l 12°2 94. 26°5 BNioy| 65

I5‘11 nigh 5:2 81 26°5 23°9 118

I-12 OG) Q°2 69 46 48 60

| 10°6 10°8 60

Average 71 13°6 20°8 67

17°6 7c 142

4°7 11°9 68

Average 80

330 ANNALS OF THE SOUTH AFRICAN MUSEUM

As shown later, copulation coincides with moulting of the female. If the

two parts of ‘Table XI are compared it will be seen that there is a difference of

nine days between the average intervals, and copulation therefore seems to

have a slight stimulating effect as regards egg-extrusion.

During the laboratory experiments several observations of the method

whereby females with hatching eggs aided the zoea’ larvae to escape from

underneath the abdomen were made. On such occasions females stood on their

pereiopods and bent their abdomens backwards. The zoeae then poured from

beneath the abdomen into the surrounding water.

COPULATION

In Hymenosoma orbiculare copulation takes place between a hard male and a

soft, newly moulted female. In this respect this species behaves in the same way

as C. maenas and many other crabs. Copulation is preceded by an embrace of

the female by the male, some time before she actually moults. The male holds

the female underneath him with his pereiopods, but the female is in the normal

position (i.e. dorsal side up). Actual copulation was observed several times

in the laboratory. The following notes were taken on one of the occasions:

I1°11. °47 Female recently moulted, leathery. Add male from dish 12. After a few seconds the

male mounts the female. ‘This male removed and substituted by male from dish 15. The latter,

after a few seconds, approaches the female which is busy digging into the sand. Male is first

on top of female, but then gets underneath the female. He then turns over on to his legs and in

doing so keeps female underneath him and turns her over on to her back and copulation

commences.

In the limited number of copulations observed, the process lasted more than

half an hour and probably longer. After copulation had occurred, the female

was kept embraced by the male for a considerable time, sometimes more than

a day.

No actual experiments were carried out with a view to establishing the

presence or absence of sex recognition. The general impression, however,

was that sex recognition was poorly developed if present at all. The procedure

in H. orbiculare seemed very similar to that suggested by Broekhuysen (1937)

for C. maenas, i.e. the seasonal periodicity in copulation is only caused by the

seasonal moulting act in sexually mature females, when the female due to its

soft condition cannot evade or resist a male trying to copulate. These remarks,

however, are tentative and more experimental work is necessary.

GROWTH

In order to obtain information on the rate of growth, the number of moults,

the time required for hardening after moulting, the existence of sexual dimor-

phism and the average maximum age, all the crabs collected in Sand Vlei

estuary were sexed and measured. The index of size used was the width of the

carapace in millimetres. Crabs which were just about to moult, or had just

moulted were noted. As has been mentioned a fairly large number of crabs

THE BREEDING AND GROWTH OF HYMENOSOMA ORBICULARE DESM. 331

were kept in captivity in some cases for periods of over a year. The growth of

those which were caught while still small was recorded in order to supplement

the data obtained in the field.

MOuLTING

As shown earlier, moulting not only marks a stage of growth, but in the

present species it also controls the possibility of successful copulation. Moulting

is thus essential to the animal but is also a dangerous period, for the crab is

practically defenceless and open to attack by predators, including its own

species. It has been noted that males copulate with soft, just moulted females

and this must decrease the danger of soft females being attacked by males,

considerably to the advantage of the females. In some cases females which

moulted in the laboratory were killed by males in the same dish. If this was

not an abnormal occurrence due to captivity, it means that the moulting

female is not altogether immune to attacks by males. Hymenosoma orbiculare and

also the other species of Brachyura where males copulate only with soft females

seem to have some advantage over species in which copulation occurs between

two hard crabs.

The process of moulting in H. orbiculare is identical to what takes place in

other Brachyura and has been described for Carcinus maenas and Cyclograpsus

punctatus by Broekhuysen (1936 and 1941). One interesting difference between

the hardening of the new shell of C. punctatus and that of H. orbiculare was found.

While H. orbiculare always remained submerged during the process, it was

found that in C. punctatus it is essential that the crab should only be submerged

part of the time for the hardening process to proceed normally.

The duration of the hardening process was observed in the laboratory. For

52 crabs between 8 and 22 mm. carapace-width at a temperature between

12 and 15° C., the average time was four days and the extremes two and nine

days. Unfortunately the records do not permit one to determine the effect of

temperature or the size of the crab on the duration of the hardening process.

The increase in size after moulting was recorded for 99 crabs kept in

captivity under fairly constant temperature. These records include a certain

number of observations on crabs kept in very low salinities and some which

were kept in water with a salinity of over 3 5/00:

As the increase in size of crabs kept in abnormal salinities was roughly the

Same as the increase for crabs in normal salinities, the records were combined.

Also, no significant difference was found between males and females. A

summary of all the records gave the following results:

Size range Number of crabs | Percentage increase in carapace

width

5—8 mm. 15 22%

9—12 mm. 18 24.%

13—16 mm. 48 15%

17—22 mm. 18 11%

332 ANNALS OF THE SOUTH AFRICAN MUSEUM

Thus also in H. orbiculare the rate of increase decreases as the crab gets

bigger and older.

PERIODICITY IN MOULTING

As females only copulate after moulting, it was thought possible that mature

females would show a distinct periodicity in moulting, correlated with the

breeding-season.

In Table XII percentages of moulting females and males, and females in

berry have been combined. Only crabs larger than 14 mm. have been

considered. ‘The information is expressed graphically in fig. to.

Table XII

% of moulied | % of females | Total number % of moulted Total number

Date Females in berry of females males of males

February 1947 3 ; I 136 5 108

March 37 oO 119 14 184

April 35 a) 40 5 78

May 21 27 117 5 ZoMe

June 19 52 52 13 31

July 8 81 93 13 30

August 4. 78 go 2 43

September 13 04. 54 (0) I

October 5 55 22 oO 3

November 13 27 99 6 53

December 4 10 49 5 39

January 1948 4 2 338 4 159

From Table XII and fig. 10 it can be seen that mature females can and do

moult during the whole of the year, but that there is a definite maximum in

moulting activities during the months March, April and May. Fig. 10 also

shows that this increase in moulting precedes the breeding-season. The

observations available for the males seem to indicate that there is no clear

maximum for the moulting in this sex.

In fig. 11 the comparable figures for crabs kept in the laboratory have been

graphed. If the two figs. 10 and 11 are compared it will be noticed that they

differ little for the females but quite distinctly for the males. The females kept

in captivity also show a sudden increase in the number of moulting crabs in

March, April and May, the same as was found in the crabs at Sand Vlei. For

the males the number of observations of captive crabs is limited but they do

show a clear maximum of moulting in March. This was not found in the crabs

at Sand Vlei.

It should be stressed that fig. 11 is based on crabs kept under rather uniform

conditions and may therefore be more accurate in a comparison of the

behaviour of females to males. In any case it is interesting to note that in these

THE BREEDING AND GROWTH OF HYMENOSOMA ORBICULARE DESM. 333

laboratory crabs, the

moulting activity of the

males precedes. the

maximum moulting

activity of the females,

in other words the males

have already moulted

and become hard when

the females moult and

are ready for copula-

tion. In the field where

conditions are not

uniform this adaptation

apparently becomes less

striking.

SEXUAL DIMORPHISM

In Hymenosoma orbi-

culare there is relatively

little external difference

between the two sexes

except for the shape of

the abdomen, and the

2ABS

NUMBER OF MOULTING CRA

MONTHS

Big Tt

The number of moultings taking place in the

100 --~MOULTED FEMALES

904 ——MOULTED MALES 74

! *+t++ FEMALES IN vA

804 BERRY jr,

(e)

PERCENTAGES

ev bh

o Oo

Fig. 10

The percentages of (a) moulted or moulting females,

(6) moulted or moulting males and (c) females in berry,

plotted against the months of the year. The crabs

were collected at random at the mouth of Sand Vlei.

development of the pleopods,

which are adapted to carry-

ing eggs in the female. The

chelae, however, are some-

what bigger and broader in the

case of the male (fig. 1).

In order to determine

whether there was a _ con-

sistent difference in size be-

tween males and females of

the Sand Vlei population, a

total of 1,417 females, of

which 316 were females in

berry, and 856 males were

measured. The results are

given in Table XIII and

different months in crabs kept in captivity. graphed in fig. 12.

334 ANNALS OF THE SOUTH AFRICAN MUSEUM

—---— ALL FEMALES

MALES

pH FEMALES WITH

OVA

NUMBER OF CRABS

6 ree RRM Saraue Rll Sm ~

(5,5.6.7.8. 10.11.12.13.14,15,6.17.18.19.20.22. 24. 26.

CARAPACE -WIDTH IN MILLIMETRES

Fig. 12

The distribution of all the crabs measured over classes differing one millimetre. The size is

expressed in carapace-width in mm. (females with ova means females in berry).

THE BREEDING AND GROWTH OF HYMENOSOMA ORBICULARE DESM. 335

Table XIII

| The carapace width in millimetres of crabs from Sand Vlei

Carapace Females (those in | Females in berry | Males| Mean carapace Mean carapace

width berry included ) only width for females width for males

5 mm. ~ - I

5 I - 2

6 - - I

7 I - I

8 I - I

9 ~ ~ I

10 I ~ I

II I - 2

12 19 a 5

13 66 12 28

14 254 12 44, 16 18

15 381 21 105

16 241 24 132

17 126 35 118

18 108 ais 94.

19 IOI 64 84

20 103 52 86

21 43 22 60

22 20 12 39

23 2 I 36

24. 2 I 10

25 = < 3

26 - = 2

Total 1,471 316 856

The mean for females is 16 mm. and for males 18 mm. The difference is

very small and was tested statistically. The t-test of significance showed that

at the 1 per cent level of significance the estimated ¢ was 2-819 and the 5 per

cent level 2-074. The calculated value for t was 0-136. There was therefore no

significant difference in size between the two sexes in the population of

H. orbiculare at Sand Vlei at the time of the investigation.

The shape of the curves in fig. 12 may suggest the presence of two year-classes

in both females and males.

THe RATE oF GROWTH

Since crabs can only increase in size when they moult the rate of growth is

dependent on the increase at a moult and the frequency of the moults.

As has been mentioned before the experiments in the laboratory did not

reveal any significant difference between males and females in the increase in

size after a moult. The increase after moulting is mentioned on page 331. As

regards the number of moults, or rather the duration of the interval between

two successive moults, it is difficult, if not impossible, to obtain information on

crabs living under natural conditions. The only information available, there-

336 ANNALS OF THE SOUTH AFRICAN MUSEUM

fore, comes from crabs reared in the laboratory where conditions were not

quite natural. This may or may not have affected the duration of the intervals

between successive moults.

In Table XIV the available information has been tabulated. The females

not in berry and those in berry as well as the sexes have been kept separate.

The crabs have been divided into five size-classes.

Table XIV

Interval between successive moults in days, in crabs reared in the laboratory.

(Extremes are given in brackets.)

FEMALES MALES

Not in berry In berry

SLC GR ee

mm. Average Number Average Number Average Number

duration of cases duration of cases duration of cases

5 40(18—58) 5 30(18-40) 5

5-8 33(25-45) 11 35(22-57) 4

Q-12 52(30-go) 12 37(38-96) By.

13-16 89(59-139) II 219(216—221) 2 32(81-214) 4

17-22 128(42-183) 5 174(109-221) II 66(51-79) 4

This table reveals some interesting points:

(a) From the table it appears that the interval between successive moults, even

in crabs of less than 5 mm., was considerable. For the smallest sizes this may be

partly due to the young crabs taking some time to settle down to laboratory

conditions after their capture.

(b) The interval between two successive moults even for crabs of the same size-

class varied a lot.

considerably longer than those for females not in berry and males. The

number of observations on males, however, may be too small to enable any

positive conclusions, but they suggest that where the female growth can be

affected adversely by breeding, this may not be the case for the males. This

point will be mentioned again later. .

(d) The interval gets longer as the crab gets older and therefore bigger and in

this respect H. orbiculare behaves in the same way as other Brachyura.

In order to determine the development of the crabs at Sand Vlei, size/

frequency graphs were plotted for each month of the year and for both sexes.

These are given in fig. 13. While every attempt was made to obtain random

samples, there is no doubt that a high proportion of the smallest size groups,

particularly those under 7 mm., evaded capture. To this extent the samples

and the graphs constructed from them are biased.

THE BREEDING AND GROWTH OF HYMENOSOMA ORBICULARE DESM.

Ly TOTAL

aA NUMBER

OF CRABS

és 4

99 138

FEB: 1947

OF CRABS

NUMBER

(Mil) MALES

| SEX?

E41) FEMALES i

86 178

99 392

JAN. 1948

(5 6 8 10 12 14 16 I& 20 22 24 26 28

CARAPACE WIDTH IN MILLIMETRES

Fig. 13

Size/frequency of males and females for the different months. The crabs

were all collected at the Sand Vlei mouth.

337

338 ANNALS OF THE SOUTH AFRICAN MUSEUM

Even so the following important points are shown:

(a) In 1947 a considerable number of very small H. orbiculare made their

appearance in the catches in September. These were the new generation of

the year. In addition there were a considerable number of female crabs of

16 to 22 mm. carapace-width which were probably over 1 year old. Hardly

any males were caught. In October the position had changed very little. In

November a distinct group of female and male crabs of 12 to 18 mm. had

appeared. It is unlikely that these consisted of crabs of the same season as they

appeared too big for that. It is impossible to decide where these crabs came

from, although the possibility of an invasion into the estuary from the shallow

sea along the shore of False Bay cannot be ruled out. A certain number of

crabs hatched in 1946 were still present, but by December most of these

seemed to have vanished.

(b) If we examine fig. 13 we shall see that in November and December the

curves for the males and the females more or less coincided. In January,

February and March the males gained more and more so that by April there

was a considerable difference in size between the males and the females, the

former being the largest. In May, however, this difference suddenly disappeared

due to a sudden increase in the size of the females. It should be remembered

that March, April and May are months in which the females showed an

increase in their moulting activity (see figs. 10 and 11), prior to the beginning

of the breeding-season. During June, July and August the males again gradually

gained on the females. This was the time when most females had egg-batches

and therefore were not in a condition to moult. This did not affect the males

(see Table XIV) which apparently continued to increase in size.

males and in September nearly all the males from the older generation had

disappeared.

During the course of the investigation a number of very small Hymenosoma

crabs were collected and reared in captivity. Some of the results of this part

of the investigation have been combined in Table XV.

Table XV

Some small Hymenosoma crabs reared in captivity at 13° to 15° C.

Increase in size in Number of Salinity in parts

Sex millimetres moults Time in days* per thousand

3 Rae 2 55 35

3 A BONG eas 3:5 5 230 35 — 36

3 4 to 5 — 16 5 329 36 — 36

y 4 to5— 13 4 169 31 — 35

Q 6-8 — 14:6 4 230 | 5— 14

2 4 to5 — 13 4 252 Ops

* The number of days covers the period from the date of moult at which the smallest size

mentioned was reached until the moult at which largest size mentioned was reached.

THE BREEDING AND GROWTH OF HYMENOSOMA ORBICULARE DESM. 339

The above table indicates that crabs of about 14.mm. are more than 8 months

old. It should be remembered, however, that these crabs were kept under

unnatural conditions and fig. 13 seems to indicate that the crabs grow faster

under natural conditions.

The size at which the crabs of the Sand Vlet population became adult

In males it was not easy to decide macroscopically when the crab had become

mature. In females, however, the change in shape of the abdomen is an obvious

indication of maturity. Moreover the development of the ovaries can usually

be seen without the aid of a microscope. Extrusion of eggs is the surest sign of

maturity having been reached. During the investigation the smallest females

with ‘adult’-shaped abdomen and the largest females with ‘juvenile’-shaped

abdomen were recorded and it soon became evident that a lot of variation

eccurred. Some females with a carapace-width as small as 12:2 mm. had

‘adult’ abdomens, while others as large as 17 mm. still had somewhat ‘juvenile’

abdomens. The smallest female carrying a batch of eggs was 12:6 mm. in the

Sand Vlei population. Dissections showed that in one instance a female as

small as 12-2 mm. had well-developed ovaries, indicating that she was mature.

Dissections also revealed that some males of 12:3 mm. and 12:5 mm. had well-

developed genital tubes, indicating that they were mature. In one instance a

female of 15-5 mm. was collected that had a rather immature abdomen but

dissection revealed well-developed ovaries. The shape of the abdomen,

therefore, is not always decisive.

As will be seen later different environments may affect the minimum size at

which maturity is reached.

Although the above data shows that H. orbiculare females from Sand Vlei

could become mature at 12-13 mm., they were usually mature at 13-14 mm.

carapace-width. This is confirmed in fig. 12 in which among others the size/

frequency of berried females has been plotted.

SEx-RATIO

When all the sexable crab samples from Sand Vlei were added together it

was found that 1,471 or 63-2 per cent were females and 856 or 36:8 per cent

were males. It is interesting to note from fig. 13 that this predominance of

females over males was specially marked in the January population. From

then on the difference between males and females decreased gradually until in

April the situation was such that there were more males than females. From

April onwards the females again became predominant. No satisfactory

explanation can be given for the large predominance of females over males. It

is very unlikely that the males were overlooked during collecting. The increased

predominance of females after May may be partly due to the fact, noticed

during collecting of samples, that the females usually do not expose themselves

so readily as the males and that, therefore, the chances of being swept away

to the sea by the current are greater for the males,

340 ANNALS OF THE SOUTH AFRICAN MUSEUM

DiIscussION

Although the fact that H. orbiculare is often found in estuaries indicates that

the species is euryhaline, there seems to be a certain minimum salinity tolerance,

as these crabs do not occur in waters with a constant very low salinity. In order

to estimate this minimum salinity limit, crabs were kept in water of a low

salinity and under controlled temperature conditions. These experiments

showed that within a temperature range of 12°-15° C., a salinity of 5 per

thousand interferes with the normal development of the eggs. Eggs extruded

under these conditions did not develop beyond the first cleavage stages.

However, when females carrying embryos in an advanced stage of development

were kept under the same conditions, the embryos did hatch sometimes but

the resulting zoea larvae died immediately. A salinity of 0-2-1-4 per

thousand and a temperature of 12°—14° C. had the same effect on development.

Although salinities of 1-5 per thousand at a temperature of 12°-15° C.

seriously affect the early development of this species if the crabs are exposed

to these conditions long enough, it apparently has little or no effect if the crabs

are only exposed to it for a short time. This is shown by the figures in Table I.

From this table it can be seen that during part of June, July and August 1947

the salinity at the mouth of Sand Vlei fell well below 4 per thousand but this

apparently did not affect the development of the larval stages in the egg,

although it may have affected hatching zoeae and eggs freshly extruded. As

the salinity in estuarine sands changes slowly, even when the water above is

almost fresh, the habit of the females to dig into the surface layer of the sandy

substratum may have saved the eggs from any adverse influence due to too low

a salinity.

During the low-salinity experiments in the laboratory, several females

extruded eggs. Low salinities, therefore, apparently do not prevent females

from extruding eggs. Nothing definite can be said regarding the effect of

prolonged exposure to high salinities, but there are strong indications that

although the species may survive and even develop under these conditions, it

has some effect.

Barnard (1950) in his Monograph on the South African Decapoda mentions a

variety of H. orbiculare from deeper water in False Bay, which at first sight seems

to be specifically distinct from the normal orbiculare. Although Barnard

stresses the difference he does not consider it necessarily another species.

During the course of this investigation the present author had an opportunity

of examining fairly large numbers of these deep-water Hymenosoma from False

Bay. In addition to the extensive granulation, mentioned by Barnard, there

was a striking difference in the size when compared with H. orbiculare from the

mouth of Sand Vlei. In Table XVI the sizes of these deep-water crabs are

compared with those of the Sand Vlei population.

THE BREEDING AND GROWTH OF HYMENOSOMA ORBICULARE DESM. 341

Table XVI

Comparison of crabs dredged below 12 fathoms in False Bay with those from

Sand Vlei estuary:

False Bay (dredged) Sand Vler

Females

Maximum carapace-width 10 mm. 24mm.

Average carapace-width 7-9 mm. 18 mm.

Smallest 9 with mature abdomen 6—7 mm. 13-14 mm.

Smallest berried female 6 mm. 12°6 mm.

Males

Maximum carapace-width 12 mm. 26 mm.

Average carapace-width 7-6 mm. 18 mm.

The total number of deep-water crabs available was 96 99 and 85 gg. The

differences in size between the two populations are very striking. If the deep-

water crab is the same species as H. orbiculare which populates the mouth of

Sand Vlei, the difference must have been caused by external conditions. The

factors which may be concerned are:

(a) constant high salinity in the case of the False Bay crabs, and a lower

average salinity, but varying tremendously, in the case of the Sand Vlei

population ;

(b) considerable depth in the case of the False Bay crabs, and relatively

shallow water in the case of the Sand Vlei crabs;

Without more detailed field-work and experiments, it is not possible to be

certain which of these factors is or are the more important. There is some

evidence, however, that salinity may be important. In 1948 during an investi-

gation of the ecology of St. Lucia Estuary on the Zululand coast (Day, Millard

and Broekhuysen, 1954) it was found that salinities in this estuary were very

high (34-53 per thousand). Specimens of H. orbiculare were collected and

measured, and it was found that females were becoming mature at 5 and 6 mm.

carapace-width and one female of 4 mm. had a ‘mature’ abdomen. Several

berried females were only 5 and 6 mm. These crabs were therefore comparable,

at least as far as size, with those from False Bay. Although few measurements

are available the Hymenosoma crabs inhabiting Langebaan Lagoon also seem

to be of small size. The smallest mature female measured was 10 mm., while

a female of 7 mm. carrying an egg-batch was collected in Saldanha Bay into

which Langebaan Lagoon opens. The salinity in this lagoon is near the

salinity of normal sea-water and therefore fairly high.

It seems, therefore, that high salinity is at least one factor which decreases

the size of the mature crabs. It also increases the size of the chelae in the

males and causes excessive granulation. There are also indications that there

342 ANNALS OF THE SOUTH AFRICAN MUSEUM

is a difference in breeding habits between the deep-water Hymenosoma from

False Bay and those from Sand Vlei.

When the Hymenosoma results are compared with those for Cyclograpsus

punctatus (Broekhuysen, 1941) it is evident that there is a great deal of similarity

between the two species. Both are winter-breeders. In C. punctatus breeding

covers the period May to November and in H. orbiculare the period is from June

to November. In both cases the females produce several egg-batches in the one

breeding-season. Whereas C. punctatus has a second, minor breeding-season in

the summer this is not evident in H. orbiculare. In both species the gonads of

the male show a periodicity in their activity which is adapted to the rhythm in

the females. There is also a great similarity as regards the incubation period

of the eggs of the two species. The incubation time of the eggs of C. punctatus,

at a constant temperature of 16°5° C., is little over a month, while at a tem-

perature of 12°-15° C. it took the eggs of H. orbiculare 38 to 48 days to develop

and hatch.

In Cyclograpsus copulation takes place between two hard crabs while in

Hymenosoma it only takes place between a hard male and a freshly moulted

female. In C’. punctatus no seasonal difference between the moulting periods of

the two sexes was observed, while in Hymenosoma orbiculare such a difference did

seem to occur. This is probably related to the fact that in the latter copulation

only takes place when the female has recently moulted. In neither species is

there a significant difference in size between the two sexes. In C. punctatus

from the shore of False Bay the majority attained an age of two to three years.

The present investigation indicates that the majority of H. orbiculare die in

their second year. In both species there were more females than males, although

this appeared to be much more pronounced in the case of Hymenosoma.

SUMMARY

(1) Hymenosoma orbiculare is a crab which occurs in the shallow waters of

estuaries as well as in deeper water along the shore of South Africa and Portu-

guese East Africa. Most of the material on which this paper is based was

collected in the False Bay area between February 1947 and February 1948.

(2) The population, showed a definite periodicity in the activity of the

gonads and the breeding-season appeared to be the winter months.

(3) Hymenosoma females extruded two or three egg-batches during the

breeding-season..

(4) The development of the eggs was followed in the field and in the

laboratory.

(5) Females will only allow males to copulate after the female has moulted

and is still soft.

(6) Although moulting may occur at any time of the year, the females

showed a sudden increase in moulting a few months before egg-extrusion.

THE BREEDING AND GROWTH OF HYMENOSOMA ORBICULARE DESM. 343,

Under laboratory conditions males also showed a periodicity, which seemed

to be adjusted to that of the females. Under natural conditions this was not so

evident.

(7) Evidence is given that during the breeding-season males increase more

rapidly in size than the females. This may be due to the fact that the majority

of females are carrying eggs and therefore do not moult during that period.

(8) The percentage increase in size of the crab after each moult shows a

eradual decrease as the crab grows larger.

(9) The size at which the female crabs became mature was established for

the Sand Vlei population for the period of the investigation. In crabs from

other localities it was found that the size varied according to environment.

(10) There was evidence that the majority of crabs, in the population

studied, reached an age of somewhat over one year, after which a heavy

mortality occurred.

(11) There are the usual Brachyuran sexual differences between the male

and the female Hymenosoma orbiculare; but there is no significant difference

between the sizes of the two sexes.

(12) Of all the crabs collected and sexed 63-2 per cent were females and

36:8 per cent males.

(13) The influences of low and high salinities are discussed and the life

history of H. orbiculare is compared with that of C. punctatus.

ACKNOWLEDGEMENT

I have pleasure in thanking Professor J. H. Day for his valuable criticism.

REFERENCES

Barnard, K. H., “Descriptive Catalogue of South African Decapoda Crustacea (Crabs and

Shrimps).’ Ann. S. Afr. Mus., XX XVIII, 1-864, 1950.

Broekhuysen, G. J.,“On Development, Growth and Distribution of Carcinides maenas (L.).’

Arch. Neerl. Zool., 11, 257-399, 1936.

——, ‘Some notes on Sex Recognition in Carcinides maenas (L.).’ Arch. Neerl. Zool., 111, 156-64,

1937-

——., ‘The Life-History of Cyclograpsus punctatus, M. Edwards: Breeding and Growth.’ Trans.

Roy. Soc. S. Af., XXVIII, 4, 331-6, 1941.

Churchill, E. P., ‘Life History of the Blue Crab’. Bull. U.S. Bur. Fisher., 36, 1917-18.

Day, J. H., Millard, N. A., Broekhuysen, G. J. B., “The Ecology of South African Estuaries,

Part IV, The St. Lucia System.’ Trans. Roy. Soc. S. Af, XXXIV, 1, 129-56, 1954.

Duncker, H., ‘Gefangenschaftsbeobachtungen an Sesarma cinerea Milne Edwards.’ Zool. Jahrb.

Syst., 66, 285-90, 1934.

Gosse, P. H., ‘On the Sloughing of the Spider-Crab (Maia squinado).’ Ann. Nat. Hist., X,

210-12, 1852.

Williamson, H. C., ‘II. Contribution to the Life-History of the Edible Crab (Cancer pagurus

Linn.), 18th.’ Rep. Fisher. Board Scotl., 3, Sci. Invest., 77-143, 1900.

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PART VI, containing :—

12. Neue Lamiinae (Cerambycidae) aus der South African Museum. Von Dr.

STEPHAN BREUNING.

13. Beitrag zur Kenntnis der Malacodermata Afrikas. Von W. Wirrmer (mit

einer Abbildung im text).

14. Some new Caddis flies from the Western Cape Province. By K. M. F. Scott,

Ph.D., FLR.E.S. (with 6 text-figures).

15. [wo new species of Boriomyia (Hemerobiidae) from South Africa. By Bo

TJEDER (with 15 text-figures).

16. Four new species of Gregarines from mountain planes of the Cape oe

By A. D. Harrison (with 11 text-figures). :

17. Fiinf Dactylispa-Typen Péringuey’s. Von Erich UHMANN (mit einer

Abbildung im text).

Title-page and Index to volume.

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BY THE RUSTICA PRESS (PTY.) LIMITED, COURT ROAD, WYNBERG, CAPE

12. Neue Lamiinae aus dem South African Museum (Coleoptera Cerambycidae).

Von Dr. STEPHAN BREUNING.

Dank dem freundlichen Entgegenkommen der Herren Dr. A. J. Hesse und

Dr. H. Andreae konnte ich die Mehrzahl der unbestimmten Lamiinae des

South African Museums untersuchen. Unter diesem interessanten Material

fand ich eine Anzahl neuer Formen, deren Beschreibungen hier nachfolgen.

Alle Typen befinden sich in dem oben angefiihrten Museum.

Prosopocera (Dalterus) hessei n. sp.

Sehr langgestreckt. Fihler um ein Drittel langer als der Kérper (2), das

dritte Glied kaum langer als das vierte. Untere Augenloben etwas langer als

die Wangen. Stirn etwas hoher als breit, sehr dicht und sehr fein punktiert.

Halsschild ziemlich dicht und fein punktiert und jederseits mit zwei wenig

ausgesprochenen runden postmedianen Beulen bewehrt, je eine auf den

Seiten der Scheibe und eine am Seitenrand. Schildchen trapezformig.

Fliigeldecken im basalen Viertel dicht und ziemlich fein gekérnt, danach sehr

dicht und fein punktiert.

Dunkelrotbraun, graulila tomentiert. Die zwei riickwartigen Drittel der

Fligeldecken mit sehr kleinen wenig deutlichen weisslichen Flecken iibersaht.

Lange: 19 mm.; Breite: 6 mm.

Typ: 1 2 von Rhodesia: Sebakwe, leg. D. Dodd.

Diese Art unterscheidet sich leicht von allen anderen Arten des Subgenus

Dalierus Fairm. durch das Fehlen eines Halsschildseitendornes sowie die

gleichzeitige Anwesenheit einer dichten Granulierung auf dem basalen Viertel

der Fliigeldecken.

Prosopocera (Dalterus) fossulata Breuning.

Das Mannchen dieser Art war noch nicht bekannt (1936, Nov. Ent., 3.

Suppl., fasc. 24, p. 188). Nun liegt mir ein solches von Damaraland: Otji-

_kondo, vor. Die Fithler sind um zwei Drittel langer als der Kérper, die

Glieder drei und vier in ganzer Lange verdickt. Die Stirn ist mit einem langen,

schmalen, zweispitzigen Horn bewehrt. Sonst wie das 9.

Falsotragiscus n. gen.

Sehr langgestreckt. Fihler fein, kaum kiirzer als der Kérper, unterseits

nicht gefranst; das erste Glied diinn und ziemlich lang, das dritte merklich

345

VOL. XLI. PART 6.

JUL 2 8 1958

346 ANNALS OF THE SOUTH AFRICAN MUSEUM

langer als das vierte oder das erste. Fiihlerhécker weit auseinanderstehend,

kaum vortretend. Augen fein facettiert, sehr stark ausgeschnitten. Stirn

breiter als hoch. Halsschild quer, vor der Basis eingezogen und quer vertieft,

mit zwei feinen Querfurchen—eine am Vorderrand, die andere am Hinter-

rand—und einem kleinen konischen Seitendorn bewehrt. Fliigeldecken sehr

langgestreckt, parallel, kaum breiter als der Halsschild, apikal verrundet.

Kopf nicht riickziehbar. Prosternalfortsatz schmal, an seinem Vorderrand

abgestutzt. Mesosternalfortsatz vorn senkrecht abgestutzt. Metasternum

von normaler Lange. Mittelhiifthéhlen offen. Beine wenig lang, die Schenkel

leicht gekeult, die Mittelschienen mit Dorsalfurche, die Klauen divergent.

Typ: peritragoides Breuning.

Diese Gattung gehort in den Tribus Tragocephalini, neben Pseudotragiscus

Breun. Sie unterscheidet sich von letzterer Gattung durch ein nicht gekeultes

erstes Fihlerglied, kaum vortretende Fihlerhocker, nicht vortretende

Schultern u.s.w.

Falsotragiscus peritragoides n. sp.

Untere Augenloben so lang als die Wangen. Die ganze Oberseite sehr fein

und wenig dicht punktiert.

Dunkelbraun. Kopf und Halsschild ockergelb tomentiert. Eine dunkel-

braune Querbinde verbindet die Fiihlerhécker. Eine ziemlich breite schwarze

Langsbinde reicht vom Vorderrand des Scheitels bis zur Halsschildbasis.

Jederseits des Scheitels eine schwarze Makel. Auf den Seiten der Halsschild-

scheibe jederseits drei hintereinander liegende runde schwarze Makeln und

auf den umgeschlagenen Seitenteilen jederseits zwei hintereinander liegende

runde schwarze Makeln. Schildchen schwarz tomentiert. Fliigeldecken rot,

fein gelb tomentiert; auf jeder Decke vier kleine schwarze Makeln: eine

runde nahe dem Seitenrand hinter der Schulter, zwei preapikale, die eine

nahe der Naht, schief gelagert, die andere rund, auf der Mitte der Scheibe

gelegen, und eine apikale. Unterseite des K6rpers graugelb tomentiert, jedes

Abdominalsegment mit zwei seitlichen, nahe dem Vorderrand gelegenen,

kleinen schwarzen Makeln. Beine fein gelb tomentiert. Fiihler braunschwarz

tomentiert.

Lange: 11 mm.; Breite: 34 mm.

Typ von Damaraland: Karibib, leg. A. Ullmann, 1917.

Pseudochariesthes transversevitticollis n. sp.

Sehr langestreckt. Untere Augenloben dreimal so lang als die Wangen.

Fliigeldecken parallel.

Rot. Oberseite gelb tomentiert. Halsschild mit zwei grauen Querbinden,

die eine am Vorder-, die andere am Hinterrand, einer kurzen und schmalen

Mittellangsbinde, die die gelb tomentierte Zone nicht iberschreitet, und

jederseit in der Mitte mit zwei kleinen, runden dunkelbraunen Makeln: die

eine auf den Seiten der Scheibe, die andere an der Grenze der Scheibe und der

NEUE LAMIINAE AUS DEM S.A. MUSEUM (COLEOPTERA CERAMBYCIDAE) 347

umgeschlagenen Seitenteile. Auf jeder Fliigeldecke eine kleine dunkelbraune

Makel am Hinterrand des Schildchens, eine schmale graue Nahtbinde, die die

beiden vorderen Drittel durchlauft, und neun runde grau tomentierte und

schmal dunkelbraun umrandete Makeln: eine auf der Schulter, eine am

Seitenrand hinter der Schulter, eine an der Naht am Ende des ersten Drittels,

eine premediane auf den Seiten der Scheibe, eine mediane am Seitenrand—

letztere beiden seitlich miteinander verbunden—eine an der Naht am Beginne

des apikalen Drittels, eine am Seitenrand am Beginne des apikalen Viertels,

eine preapikale an der Naht, und eine apikale—letztere beiden miteinander

vereinigt. Unterseite des Korpers, Beine und Fihler fein graugelb tomentiert.

Lange: 114 mm.; Breite: 34 mm.

Typ ohne Fundortangabe.

Diese Art steht besonders der ficta Breun. nahe, unterscheidet sich aber

von ihr wie auch von allen anderen Arten dieser Gattung durch das Vor-

handensein zweier grauer Querbinden auf dem Halsschild.

Ceroplesis thunbergi Fahr. m. supplementaria nov.

Wie die Stammform, aber mit einigen kleinen roten Flecken zwischen der

postmedianen Querbinder und der Apikalbinde.

Typ.: 1 ¢ von Maputaland, Nord-Zululand, leg. H. W. Bell-Marley, 1908.

1 Allotyp, 9, ohne Fundortangabe.

Von hamilton Auriv. unterscheidet sich diese Form durch die breitere und

regelmassigere postmediane rote Querbinde der Fliigeldecken.

Ocax subaequalis n. sp.

Dem inaequalis Thoms. nahe stehend, aber die ganze Oberseite feiner

punktiert, die postbasale Langscrista der Decken aus sechs kleinen Zahnchen

gebildet.

Die Tomentierung ist recht 4hnlich, aber der Hinterkopf ohne schwarze

Pubescenz, die beiden schwarzen seitlichen Halsschildlangsbinden nach vorn

bis zum Hinterrand der Augen verlangert, die dreieckige postmediane schwarze

Seitenmakel der Decke im Nahtdrittel als schmale Querbinde bis zur Naht

verlangert, die kurze schwarze Querbinde im apikalen Deckenviertel durch

eine sehr kleine, schief gelagerte schwarze Makel nahe der Naht ersetzt, die

preapikale schwarze Seitenrandmakel langer. Die ganze basale Halfte der

Fihlerglieder vom dritten ab, weiss tomentiert.

Lange: 12 mm.; Breite: 5 mm.

Typ von Natal: Durban, leg. H. W. Bell-Marley.

Idactus plurifasciculatus n. sp.

Sehr langgestreckt. Fihler merklich kiirzer als der K6rper; das erste

Glied massig lang, das dritte viel langer als das vierte. Untere Augenloben

348 ANNALS OF THE SOUTH AFRICAN MUSEUM

etwas kurzer als die Wangen. Fiihlerhécker mdssig hoch. Stirn so hoch als

breit. Halsschild mit vier feinen Querfurchen, zwei nahe dem Vorderrand,

zwei nahe dem Hinterrand, fiinf kleinen Scheibenhéckern: ein kleiner

konischer in der Mitte, die anderen vier kaum angedeutet, und einem ziem-

lich kleinen stumpf konischen Seitenhécker. Schildchen halbrund. Fligel-

decken sehr langgestreckt, gewdlbt, apikal verrundet, wenig dicht und sehr

fein punktiert. Jede Decke mit fiinf kleinen Haarfaszikeln: zwei postbasale,

eines auf der Scheibenmitte, das zweite seitlich davon, zwei postmediane auf

der Scheibe, eines neben dem anderen, und ein preapikales, hinter dem

inneren postmedianen gelegen.

Dunkelbraun, weisslich tomentiert und dicht braun marmoriert, besonders

dicht auf den Schlafen und den Halsschildseiten. Auf jeder Decke zuweilen

zwei hellbraune wenig scharf ausgepragte Scheibenmakeln: eine postbasale

hinter den beiden vorderen Haarfaszikeln gelegen und eine postmediane

hinter den breiden postmedianen Haarfaszikeln gelegen.

Lange: 16 mm.; Breite: 54 mm.

Typ von Prince Albert, leg. C. West, 1 Paratyp von Namaqualand und

1 Paratyp von O’okiep, Namaqualand, 20. XII. 1885.

Diese Art steht dem multifasciculatus Breun. nahe, unterscheidet sich aber

insbesondere durch das Fehlen von Kanten auf den Decken und die geringere

Zahl von Haarfaszikeln.

Olenecamptus tesselatus Dist. m. postobliteratus nov.

Wie m. vittatus Breun., aber ohne weisse Preapikalmakel, so dass die

weisse Scheibenbinde am Beginn des apikalen Viertels endigt; die Basalmakel

bleibt von der Binde getrennt.

Typ.: 1 von Siid Rhodesia: Sebakwe, leg. D. Dodd, 1 Paratyp ditto.

Tambusoides n. gen.

Langoval, gedrungen. Fihler fein, etwas mehr als zweimal so lang (¢)

oder etwas kiirzer (2) als der K6rper, die drei ersten Glieder nicht gefranst, -

die weiteren deutlich gefranst, dichter beim ¢ als beim Q, die letzten Glieder

beim ¢ noch dichter gefranst; das erste Glied kurz und dick, das dritte so

lang als das vierte, viel langer als das erste, merklich langer als eines der

weiteren Glieder. Fithlerhécker einander ziemlich genahert und hoch. Augen

fein facettiert, sehr stark ausgeschnitten, die unteren Augenloben so hoch als

breit. Halsschild quer, gewélbt, mit zwei hohen konischen Scheibenhéckern

und einer nur schwach angedeuteten stumpfen postmedianen Seitenbeule.

Fligeldecken lang, gewélbt, an den Schultern merklich breiter als der Hals-

schild, apikal verrundet. Jede Decke mit einem kurzen und hohen basalen

Kamm auf der Scheibe und zahlreichen grossen Haarfaszikeln. Schultern

eckig vortretend. Kopf riickziehbar. Prosternalfortsatz wenig breit, niedriger

als die Hiiften, regelmdssig gerundet. Mesosternalfortsatz breit, an seinem

NEUE LAMIINAE AUS DEM S.A. MUSEUM (COLEOPTERA CERAMBYCIDAE) 349

Vorderrand mit zwei kleinen vorstehenden nebeneinander gelegenen Hoéckern

besetzt. Metasternum leicht verkiirzt. Beine massig lang, kraftig, die Schenkel

gekeult, die Mittelschienen ohne Dorsalfurche, die Klauen gesperrt.

Typ: multifasciculatus Breun.

Diese Gattung gehort in den Tribus Crossotini. In meiner Bestimmungs-

tabelle der Gattungen dieses Tribusses (1942, Nov. Ent., 3. Suppl., Fasc. 72,

p. 8) reiht sie sich bei der Nummer to ein und unterscheidet sich von Tambusa

Dist. durch die hohen Fihlerhécker, das Fehlen eines Halsschildseitendornes,

apikal verrundete Fltigeldecken, u.s.w.

Tambusoides mulitfasciculatus n. sp.

Untere Augenloben viel kiirzer als die Wangen. Kopf und Halsschild

ziemlich dicht und grob punktiert. Schildchen quer trapezformig. Fliigel-

decken dicht und sehr grob punktiert, die Punkte apikalwarts etwas feiner

werdend. Der basale Kamm an seinem Vorderrand in einen vortretenden

Hécker auslaufend und in seiner riickwartigen Halfte von einem grossen

gelben Haarfaszikel tberlagert. Auf jeder Decke ausserdem sieben grosse gelbe

Haarfaszikel: eines hinter der Schulter, seitlich des den Kamm iiberragenden

Faszikels, drei im mittleren Teil und drei im apikalen Drittel, die in Form

zweier gewellter Querbinden angeordnet sind. Metasternum und Abdomen

dicht und grob punktiert.

Dunkelbraun, Kopf braun tomentiert, mit rosa untermischt. Halsschild

rosa tomentiert, hinter den Scheibenhéckern und auf den Seitenteilen mit

braun untermischt. Schildchen braun tomentiert mit breiter ockergelber

Mittellangsbinde. Fligeldecken und Unterseite des Kérpers strohgelb tomen-

tiert, die Punkte kahl bleibend. Beine feiner strohgelb tomentiert und mit

braun untermischt, fast einfarbig braun in der apikalen Halfte der Schienen.

Fuhler dunkelbraun tomentiert, die basale Halfte der Glieder vom fiinften ab

und der apikale Teil des elften Gliedes weisslich tomentiert.

Lange: 16-20 mm.; Breite: 7-84 mm.

Typ: 1 g vom St. Johns River. Allotyp: 1 2 von Port St. Johns, VII.

1894.

Tetraulax rhodesianus n. sp.

Untere Augenloben kaum zweimal langer als die Wangen. Halsschild

ausser den beiden kreisf6rmig angeordneten Furchen auf der Scheibe mit

einer in Form eines Dreieckes angeordneten Furche. Fliigeldecken sehr fein

punktiert. Tomentierung anders.

Dunkelbraun, hellbraun tomentiert. Auf jeder Decke zwei breite weisse

Querbinden: eine postbasale und eine mediane, die im Nahtviertel und im

Seitenrandviertel in der Langsrichtung miteinander verbunden sind. Einige

gekriimmte weisse Linien im apikalen Drittel. Metasternum weiss tomentiert,

350 ANNALS OF THE SOUTH AFRICAN MUSEUM

Abdomen teilweise weisslich tomentiert. Basales Drittel der Fiithlerglieder vom

dritten ab weiss tomentiert.

Lange: 8 mm.; Breite: 2? mm.

Typ von Siid Rhodesien: Umtali.

Cloniocerus aureovittatus n. sp.

Dem bohemant White nahestehend, aber die Fligeldecken in ganzer

Ausdehnung dicht punktiert, die Punkte sehr grob in den drei vorderen

Vierteln, sowie die Tomentierung abweichend:

Schwarz, schwarz tomentiert. Halsschild mit zwei breiten goldfarbenen

Langsbinden, deren je eine den Seitendorn deckt. Fliigeldecken mit zwei

goldfarbenen Querbinden: eine schmale premediane und eine breitere post-

mediane; das apikale Viertal mehr oder weniger dicht goldfarben marmo-

riert. Unterseite grésstenteils goldfarben tomentiert. Schenkel und Schienen

im basalen Drittel golden geringt. Die basale Halfte der beiden ersten Tarsen-

glieder sowie des vierten ‘Tarsengliedes und die Basis des dritten Gliedes, rot

und weisslich tomentiert. Die Fihlerglieder vom dritten (oder vom vierten)

bis zum siebenten Glied nach der Basis gelbrot geringt, diese Ringe fein

weisslichgelb tomentiert.

Lange: 10 mm.; Breite: 3 mm.

Typ von Gt. Winterhoek, Tulbagh, Cape Province, 1,500 m., XI. 1916,

leg. Lightfoot. Ein Paratyp ditto.

Stathmodera reticulata n. sp.

Fiihler etwas langer als der K6rper, das erste Glied sehr dicht und fein

punktiert, im dorso-apikalen Teil verdickt, das dritte Glied etwas kiirzer als

das vierte, etwas langer als das erste. Fiihlerhocker massig vortretend. Untere

Augenloben etwas langer als die Wangen. Halsschild mit einer kraftigen

Mittellangsrinne und sehr zahlreichen (iiber 30) sehr feinen aber scharf

eingeschnittenen Langsfurchen, die nach riickwarts gegen die Mittellinie zu

konvergieren. Fliigeldecken sehr dicht und tief punktiert, die Intervalle ein

dichtes Netzwerk bildend. Auf jeder Decke ein feinen Langskamm, der den

Seitenrand begleitet, und ein kurzer preapikaler Langskamm auf der Scheiben-

mitte. Die apikale Randecke in einen Zahn ausgezogen. Stirn ohne abstehende

Haare.

Schwarz, sehr fein weisslichgrau tomentiert. Die Stirn, eine breite ~

Langsbinde auf dem Scheitel, die umgeschlagenen Seitenteile des Hals-

schildes, das Schildchen, sowie drei Makeln auf jeder Decke (eine schief

gelagerte hinter der Schulter, eine runde postmediane auf der Scheibenmitte

und eine runde preapikale nahe der Naht), gelb tomentiert. Unterseite und

Beine fein grau tomentiert. Fiihler schwarzbraun tomentiert, der apikale

Teil der Glieder drei bis zehn weisslich tomentiert.

NEUE LAMIINAE AUS DEM S.A. MUSEUM (COLEOPTERA CERAMBYCIDAE) 351

Lange: 7-8 mm.; Breite: 2 mm.

Typ von St. Lucia Lake, X. 1934, leg. H. W. Bell-Marley. Ein Paratyp

ditto.

Eumdia vagefasciata n. sp.

Fihler fast um die Halfte langer als der K6rper, das dritte Glied apikal

in eine Spitze ausgezogen, das vierte so lang als das fiinfte oder erste. Untere

Augenloben weniger als zweimal so hoch als breit, viermal so lang als die

Wangen. Halsschild fast zweimal so breit als lang, nicht punktiert. Fliigel-

decken apikal breit aber sehr schwach abgestutzt, durchwegs dicht und sehr

fein punktiert.

Dunkelbraun. Kopf, Halsschild und Schildchen strohgelb tomentiert.

Fliigeldecken weisslichgrau tomentiert. Jede Decke mit zwei unscharf aus-

gebildeten braunen Querbinden: eine in der Mitte und eine gewellte am

Beginn des apikalen Drittels, sowie mit drei unscharf ausgebildeten gelben

Querbinden: eine basale, eine gewellte postbasale und eine apikale. Unter-

seite weisslichgrau tomentiert, diese Tomentierung gegen die Seiten der

Sterna und der Abdominalsegmente in gelb iibergehend. Beine graugelb

tomentiert. Das erste Fihlerglied strohgelb tomentiert, die weiteren Glieder

weisslichgrau tomentiert.

Lange: 11 mm.; Breite: 3 mm.

Typ von Transvaal: Leydsdorp, 1905, leg. J. Naughton.

Eunidia bifuscofasciata n. sp.

Fiihler um ein Drittel langer als der Koper, das dritte Glied apikal in

eine Spitze ausgezogen, das vierte so lang als das fiinfte oder erste. Untere

Augenloben zweimal so hoch als breit, sechsmal so lang als die Wangen.

Halsschild quer, kaum punktiert. Fliigeldecken apikal verrundet, durchwegs

sehr dicht und sehr fein punktiert.

Dunkelrotbraun, weisslichgrau tomentiert. Zwei breite Querbinden auf

den Fliigeldecken: eine in der Mitte und eine preapikale ohne Pubeszenz,

Schenkel sowie die drei ersten Fiihlerglieder ziemlich dunkel braun tomentiert,

die weiteren Fiihlerglieder fein rotbraun tomentiert.

Lange: 6 mm.; Breite: 13 mm.

Typ von Siid Rhodesien: Sebakwe, 1902, leg. D. Dodd.

Eunidia subnigra n. sp.

Fihler um ein Drittel langer als der Korper, das dritte Glied apikal in

eine Spitze ausgezogen, das vierte so lang als das fiinfte, etwas kiirzer als das

erste. Untere Augenloben zweimal so hoch als breit, fiinfmal so lang als die

Wangen. Halsschild quer, nicht punktiert, vor der Basis und hinter dem

Vorderrand stark eingezogen. Fliigeldecken apikal verrundet, durchwegs

dicht und sehr fein punktiert.

352 ANNALS OF THE SOUTH AFRICAN MUSEUM

Schwarz, durchwegs sehr fein dunkelbraun seidenglanzend tomentiert.

Lange: 10 mm.; Breite: 2? mm.

Typ von Nord Rhodesien: Pemba, 1918, leg. Father Casset.

Mimogmodera n. gen.

Sehr langgestreckt. Fihler fein, unterseits vom vierten oder funften

Glied ab wenig dicht aber lang gefranst, um ein Drittel langer als der K6rper;

das erste Glied lang und diinn (aber immerhin merklich dicker als die

weiteren Glieder), oberseitig konvex, das dritte Glied so lang als das vierte,

etwas kiirzer als das erste, das vierte etwas langer als das fiinfte, die weiteren

langsam an Lange abnehmend. Fihlerhécker sehr klein, einander genahert

und vorstehend. Augen grob facettiert, stark ausgeschnitten, die unteren Loben

klein und quer. Stirn héher als breit. Halsschild gewolbt, merklich langer als

breit, ziemlich weit vom Vorderrand sowie ziemlich weit vom Hinterrand

merklich eingezogen, der mittlere Teil seitlich kraftig gerundet. Fliigeldecken

gewolbt, sehr lang, parallel, etwas breiter als der Halsschild, apikal ziemlich

schmal verrundet. Kopf nicht riickziehbar. Prosternalfortsatz schmal, nied-

riger als die Hiiften, gleichmdssig verrundet. Mesosternalfortsatz schmal, zum

Vorderrand allmahlich abfallend. Metasternum von normaler Lange.

Mittelhiifthohlen offen. Beine kurz, ziemlich kraftig, die Schenkel gekeult,

die Mittelschienen ausgeschnitten, die Klauen gesperrt.

Typ: rufula Breun. Diese Gattung ist der Gattung Stenidea Muls. ver-

wandt.

Mimogmodera rufula n. sp.

Erstes Fiithlerglied sehr dicht und sehr fein punktiert. Untere Augenloben

etwas kiirzer als die Wangen. Schildchen schmal, apikal verrundet. Die ganze

Oberseite des Korpers fein und sehr dicht, die Unterseite sehr fein und dicht

punktiert.

Rot, sehr fein weisslichgelb tomentiert. Fliigeldecken mit einigen unscharf

ausgebildeten dunkelbraunen Scheibenflecken.

Lange: 44-5 mm.; Breite: 1-1} mm.

Typ. von Natal: Durban, B.B.-Bush, XI. 1908, leg. H. W. Bell-Marley.

Ein Paratyp ditto. Weitere Paratypen im British Museum von Pondoland:

Port St. Johns, und Natal.

Ayllisia albifrons n. sp.

Sehr langgestreckt. Fithler fast zweimal so lang als der K6rper, das

dritte Glied merklich langer als das vierte, kaum langer als das erste, die sechs

ersten Glieder unterseits schiitter und kurz gefranst. Untere Augenloben etwas

kirzer als die Wangen. Kopf und Halsschild sehr dicht und sehr fein punk-

tiert. Halsschild so lang als breit, seitlich leicht verrundet. Schildchen halb-

rund. Fliigeldecken sehr lang, parallel, apikal verrundet, durchwegs sehr

dicht und fein punktiert.

NEUE LAMIINAE AUS DEM S.A. MUSEUM (COLEOPTERA CERAMBYCIDAE) 353

Dunkelbraun, weisslich tomentiert. Scheitel mit drei ockergelben Langs-

binden. Halsschild mit sieben schmalen ockergelben Langsbinden. Fligel-

decken rot, fein grau tomentiert, jede mit weisslichen Langsbinden: eine

Nahtbinde, eine Seitenrandbinde und drei Scheibenbinden, und in der

rickwartigen Halfte mit einer weiteren weisslichen Langsbinde zwischen der

zweiten und der dritten (von der Naht aus gezahlt) Scheibenbinde. Fihler

dunkelbraun tomentiert, die drei basalen Viertel des dritten Gliedes, die

basale Halfte der Glieder vier bis zehn, sowie das basale Viertel und die apikale

Halftes des elften Gliedes rotweisslich tomentiert.

Lange: 10 mm.; Breite: 2 mm.

Typ von Rust en Vrede, Oudtshoorn District, Cape Province.

. Anauxesis andreaei n. sp.

Fihler ungefahr zweieinhalbmal so lang als der KGrper, das dritte Glied

viel kiirzer als das vierte, viel langer als das erste. Untere Augenloben um die

Halfte langer als die Wangen. Der ganze Korper sehr dicht und sehr fein

punktiert. Halsschild zweimal so lang als breit. Schildchen fiinfeckig. Fliigel-

decken apikal ausgeschnitten, die Nahtecke spitzig, die Randecke in einen

ziemlich langen spitz-dreieckigen Lappen ausgezogen.

Dunkelbraun, sehr fein gelbbraun seidenglanzend tomentiert. Stirn,

Wangen, Schildchen und jederseits eine Langsbinde, die die umgeschlagenen

Seitenteile des Halsschildes deckt und dann auf den Seiten der Unterseite bis

zum Ende des KG6rpers reicht, ziemlich dicht weisslich tomentiert. Die

ausserste Basis des fiinften Fithlergliedes, sowie die apikale Halfte des siebenten

Gliedes (mit Ausnahme des Aussersten Endes dieses Gliedes) weiss und weiss

tomentiert.

Lange: 18-20 mm.; Breite: 2}-24 mm.

Typ von Zululand: N’kandhla Forest, I. 1937, leg. R. F. Lawrence.

Zwei Paratypen: Zululand: Mfongosi, leg. W. E. Jones.

Anauxesis nigroantennalis n. sp.

Fuhler mehr als zweimal so lang als der KGrper, die Glieder drei bis fiinf

unterseits sehr schiitter und kurz gefranst, das dritte Glied viel kiirzer als das

vierte, viel langer als das erste. Untere Augenloben um die Halfte langer als

die Wangen. Der ganze Kérper Ausserst dicht und sehr fein punktiert. Hals-

schild zweimal so lang als breit. Schildchen fiinfeckig. Fliigeldecken apikal

schwach, schief, ausgeschnitten, die beiden Ecken spitzig.

Schwarzbraun, schiitter weisslichgrau tomentiert, die Tomentierung

dichter auf den Wangen sowie auf je einer schmalen Langsbinde, die die

umgeschlagenen Seitenteile des Halsschildes und die Seiten der Sterna bis zum

Hinterrand der Metepisternen durchlauft.

354 ANNALS OF THE SOUTH AFRICAN MUSEUM

Lange: 104-11 mm.; Breite: 1-14 mm.

Typ ein 2 von Portugiesisch Ost-Afrika: Masiene, leg. R. F. Lawrence.

Ein Paratyp ditto.

Obereopsis meridionalis n. sp.

Langgestreckt. Fiihler etwas kiirzer als der K6rper, das dritte Glied so

lang als das vierte, etwas langer als das erste. Untere Augenloben merklich

langer als die Wangen. Stirn um die Halfte breiter als einer dieser Loben.

Kopf und Halsschild dicht und fein punktiert. Halsschild so lang als breit, mit

zwei breiten aber seichten Querdepressionen, eine premediane und eine post-

mediane. Schildchen dreieckig. Fliigeldecken langgestreckt, apikal ziemlich

breit verrundet, grob punktiert, die Punkte gereiht. Hinterschenkel das zweite

Hinterleibssegment kaum tiberragend.

Gelbrot, sehr fein gelblich tomentiert. Die umgeschlagen Seitenteile des

Halsschildes grésstenteils schwarz. Auf jeder Fliigeldecke eine breite schwarz-

liche Langsbinde am Seitenrand, die sich im premedianen Teil nahtwarts

stark verbreitert (sich hierbei aber etwas aufhellt) und die dann, sich ebenfalls

aufhellend, fast das ganze apikale Viertel deckt. Drei schwarze Flecken auf

dem Mesosternum: ein mittlerer und je ein seitlicher. Zwei kleine schwarze

Flecken auf dem Hinterrand des Metasternums. Hinterschienen, ausser basal,

ziemlich dunkelbraun. Tarsen braun. Fihler dunkelbraun.

Lange: 8mm.; Breite: 14 mm.

Typ von Cape Town, Rondebosch, X. 1883.

Obereopsis pseudocapensis n. sp.

Sehr langestreckt. Fiihler ziemlich fein, um ein Viertel langer als der

Korper, das dritte Glied so lang als das vierte, merklich langer als das erste.

Untere Augenloben zweimal so lang als die Wangen. Stirn merklich breiter

als einer dieser Loben. Kopf und Halsschild sehr dicht und sehr fein punk-

tiert. Halsschild merklich langer als breit, seitlich fast gerade. Schildchen

schmal dreieckig. Fliigeldecken sehr lang, in der Mitte sehr leicht eingezogen,

apikal schief abgestutzt (beide Ecken deutlich aber stumpf), sehr dicht und

massig fein punktiert, die Punkte mehr ober weniger regelmassig gereiht.

Sterna sehr dicht, Abdomen dicht und sehr fein punktiert. Hinterschenkel das

zweite Abdominalsegment kaum wuberragend.

Gelbrot, fein gelblich tomentiert. Auf dem Halsschild eine massig breite

schwarze Mittellangsbinde, die sich basalwarts leicht verbreitert; die umge-

schlagenen Seitenteile schwarz tomentiert. Auf jeder Fliigeldecke eine schwarze

Langsbinde, die von der Schulterbeule abgeht, am Ende des basalen Viertels

den Seitenrand erreicht um sich apikalwarts langsam zu verlieren. Sterna

schwarz. Eine grosse schwarze Quermakel auf der Mittel des ersten, zweiten

und dritten Abdominalsegmentes. Die zwei ersten Fihlerglieder dunkelbraun,

die weiteren Glieder rot.

NEUE LAMIINAE AUS DEM S.A. MUSEUM (COLEOPTERA CERAMBYCIDAE) 355

Lange: 9-10 mm.; Breite: 1}-2 mm.

Typ von Cape Town, leg. Péringuey.

Diese Art hatte ich, irregefiihrt durch eine falsche Determination hierher-

gehoriger Exemplare im British Museum, mit Nitocris capensis Péringuey identi-

fiziert, weshalb ich von letzterer Art sagte (1951, Ent. Arb. Mus., Frey, p. 3), sie

gehére in die Gattung Obereopsis. In Wirklichkeit weicht Nitocris capensis Per.

stark von der vorliegenden Art ab und reiht sich in die Gattung Phytoecta Muls.

ein. Ich gebe von letzterer Art hier nachfolgend die Beschreibung auf Grund

eines Paratyps von Péringuey in der Sammlung des Cap-Museums.

Phytoecia (Blepisanis) capensis (Péring.)

Nitocris capensis Péringuey, 1888, Trans. S. Afr. Philos. Soc., 1V, p. 184.

Sehr langgestreckt. Fiihler um ein Viertel langer als der Kérper, mAssig

dick, das dritte Glied etwas langer als das vierte, merklich langer als das erste.

Untere Augenloben mehr als dreimal so lang als die Wangen. Stirn merklich

breiter als einer dieser Loben. Kopf und Halsschild sehr dicht und ziemlich

fein punktiert. Halsschild so lang als breit, cylindrisch. Schildchen dreieckig.

Fliigeldecken sehr lang, apikal verrundet, dicht und grob punktiert, die Punkte

gereiht. Hinterschenkel das zweite Abdominalsegment wenig iiberragend.

Gelbrot, fein gelblich tomentiert. Die umgeschlagenen Seitenteile sowie

eine massig breite Mittellangsbinde auf dem Halsschilde sind schwarz. Auf

jeder Fliigeldecke eine schwarze Langsbinde am Seitenrand, die sich vor der

Mitte nahtwArts merklich verbreitert und dann das ganze apikale Viertel deckt.

Sterna (ausser entlang der Mittellinie) sowie der mittlere Teil der beiden

ersten Abdominalsegmente schwarz. Der apikale Teil der Tibien braun.

Tarsen dunkelbraun. Die drei ersten Fiihlerglieder dunkelbraun, die weiteren

rot.

Lange: 7-10 mm.; Breite: 14-1? mm.

Cape Town, Rondebosch, X. 1883, leg. Péringuey.

In der Bestimmungstabelle der Untergattung Blepisanis (1951, Ent. Arb.

Mus., Frey, Il, p. 30) schiebt sich diese Art bei der Nummer 69 ein, wobei sie

von disconotaticollis Breun. durch ganz rotgelb gefarbten Kopf, schwarze

umgeschlagene Seitenteile des Halsschildes etc. abweicht.

Phytoecia (Blepisanis) holonigra n. sp.

Sehr langgestreckt. Fithler wenig stark, merklich langer als der K6rper,

das dritte Glied kaum langer als das vierte, merklich langer als das erste, das

vierte etwas langer als das fiinfte. Untere Augenloben um die Halfte langer

als die Wangen. Stirn fast um die Halfte breiter als einer dieser Loben. Kopf

dicht und sehr fein punktiert. Halsschild so lang als breit, wenig dicht und

sehr fein punktiert, mit vier kleinen Scheibenbeulen besetzt: zwei vor der

Mitte der Mittellinie genahert und zwei hinter der Mitte mehr seitlich gelagert.

Schildchen gross, apikal verrundet. Fliigeldecken sehr lang, in der Mitte

356 ANNALS OF THE SOUTH AFRICAN MUSEUM

leicht eingezogen, apikal verrundet, durchwegs sehr dicht und fein punktiert.

Hinterschenkel den Hinterrand des zweiten Abdomunalsegmentag kaum

erreichend.

Schwarz, fein hellgrau tomentiert. Die Wangen, das Schildchen sowie

eine unscharf ausgebildete Langsbinde auf dem Scheitel und dem Halsschild,

dichter weiss tomentiert. Unterseite und Beine fein dunkelgrau, die Fihler

schwarz tomentiert. Die abstehenden Haare weisslichgrau gefarbt.

Lange: 9 mm.; Breite: 12 mm.

Typ: 1 ¢ ohne Fundort.

In meiner Bestimmungstabelle der Untergattung Blepisanis (1951, Ent.

Arb. Mus., Frey, II, p. 27) schiebt sich diese Art bei der Nummer 17 ein,

weicht aber von moreana Breun. und czlicrae Breun. durch viel schlankere

Gestalt, viel weniger dichte Punktierung des Halsschildes etc., ab.

Phytoecia (Blepisanis) pseudolateralis n. sp.

Langgestreckt. Fithler etwas langer als der Korper, wenig stark, das

dritte Glied so lang als das vierte oder fiinfte, merklich langer als das erste.

Untere Augenloben zweimal so lang als die Wangen. Stirn fast zweimal so

breit als einer dieser Loben. Kopf und Halsschild sehr dicht und sehr fein

punktiert. Halsschild so lang als breit, fast cylindrisch. Schildchen halbrund.

Fliigeldecken sehr lang, in der Mitte kaum eingezogen, apikal verrundet, dicht

und ziemlich grob punktiert, die Punkte gereiht. Hinterschenkel den Hinter-

rand des zweiten Abdominalsegmentes kaum erreichend.

Gelbrot, fein gelblich tomentiert. Jede Fliigeldecke mit einer dunkel-

-braunen Langsbinde am Seitenrand, die auf der Schulterbeule beginnt und

sich nach hinten allmahlich verbreitert um das ganze apikale Fiinftel zu

decken. Das erste Fihlerglied dunkelbraun, die weiteren Glieder rot,

dunkelrotbraun tomentiert.

Lange: 84 mm.; Breite: 14 mm.

Typ: ein 2 von Siid West Afrika, Kaokoveld: Kaross.

In meiner Bestimmungstabelle der Untergattung Blepisanis (1951, Ent.

Arb. Mus., Frey, II, p. 29) schiebt sich diese Art be1 der Nummer 51 ein, wobei

sie von foade Pasc. m. rufinitibialis Breun. durch feine Fliigeldeckenpunktierung,

rotgelbe Farbung der umgeschlagenen Halsschildseitenteile und der Sterna

etc., abweicht.

Phytoecia (Blepisanis) argenteosuturalis n. sp.

Der latesuturalis Breun. sehr nahe stehend, aber die Fiihler etwas dicker,

die unteren Augenloben zweimal so lang (g) oder etwas langer (9) als die

Wangen, die Stirn um die Haifte breiter (g) oder mehr als zweimal so breit

(2) als einer dieser Loben, sowie die Tomentierung abweichend.

Stirn ganz oder wenigstens grdésstenteils schwarz. Halsschild ohne vordere

schwarze Makel; die riickwartige Halsschildmakel, das Schildchen, sowie

NEUE LAMIINAE AUS DEM S.A. MUSEUM (COLEOPTERA CERAMBYCIDAE) 357

eine schmale Nahtbinde auf den Fliigeldecken silbrig tomentiert. Das vierte

Abdominalsegment sowie die Beine einfarbig rotgelb. Die Fihlerglieder vier

bis elf rot.

Typ: 1 g von Upper Sources of Olifants River, Ceres Division, Cape

Province, XII. 1949. Ein Allotyp (@) ditto.

In meiner Bestimmungstabelle der Untergattung Blepisanis (1951, Ent.

Arb. Mus., Frey, II, p. 29) schiebt sich diese Art bei der Nummer 58 ein,

wobei sie von Jatesuturalis Breun. und von neaver Aur. m. aurosternalis Breun.

durch grésstenteils schwarze Stirn, silbrig tomentierte Fliigeldecken-Nahtbinde

etc., abweicht.

Scapogoephanes n. gen.

Langgestreckt. Fiihler fein, um ein Drittel langer als der K6rper, unter-

seits ziemlich kurz und schiitter gefranst; das erste Glied kurz und ziemlich

dick, das dritte so lang als das vierte, etwas langer als das erste, das vierte viel

langer als eines der folgenden Glieder. Fiihlerhécker weit auseinanderstehend,

nicht erhaben. Augen ziemlich grob facettiert, sehr stark ausgeschnitten.

Halsschild schwach quer, gewdlbt, seitlich verrundet. Fliigeldecken lang,

etwas breiter als der Halsschild, in den vorderen drei Vierteln parallel, vor der

Mitte leich niedergedriickt, apikal verrundet; jede Decke mit zwei kaum

angedeuteten Beulen auf der Scheibe: eine postbasale und eine postmediane.

Kopf nicht riickziehbar. Prosternalfortsatz schmal, niedriger als die Hiiften,

verrundet. Mesosternalfortsatz nach vorn allmahlich geneigt. Metasternum

von normaler Lange. Mittelhiifthéhlen geschlossen. Beine miassig lang;

Schenkel gekeult, Mittelschienen dorsal ausgeschnitten, Klauen gesperrt. Der

eanze Korper und die Beine mit abstehenden Haaren besetzt.

Typ: pusillus. Breun. Die Gattung kommt neben Acartus Fahr. zu stehen.

Scapogoephanes pusillus n. sp.

Untere Augenloben etwas kiirzer als die Wangen. Schildchen halbrund.

Fliigeldecken sehr dicht und sehr fein punktiert.

Dunkelrot, schiitter strohgelb tomentiert. Die Fliigeldecken zum Teil,

die basale Halfte der Schienen, das zweite Fiihlerglied und die basale Halfte

des dritten und vierten Gliedes, sowie der basale Teil der weiteren Glieder

hellrot. Die abstehenden Haare der Oberseite schwarz, die auf dem Fliigel-

deckenseitenrand und den Beinen fahlgelb.

Lange: 34 mm. Breite: 14 mm.

Typ von Siidafrika: Blauberg (20 km. nérdlich von Kapstadt).

13. Beitrag zur Kenntnis der Malacodermata (Col.) Afrika’s. Von W. WitTTMER,

Buenos Aires. (Mit einer Abbildung im Text.)

Die in der vorliegenden Arbeit beschriebenen Arten entstammen Bestim-

mungssendungen, die ich vom South African Museum, Cape Town und vom

Transvaal Museum, Pretoria erhalten habe. Den Herren Dr. A. J. Hesse,

Cape Town and C. Koch, Pretoria danke ich verbindlichst fiir die freundliche

Uberlassung von Dubletten fiir meine Sammlung.

MALACHIIDAE

Dinometopus humeropictus nov. spec.

6 Schwarz, Kopf gelb, Hinterrand der Stirn bis zum Hinterrand der

Augen schmal schwarz gesdumt; Fihler gelb, vom 5. oder 6. Gliede an

gebraunt; Basalrand des Halsschildes in der Mitte kurz und Ausserst schmal

weisslichgelb; Seiten der Fliigeldecken mit einer langen, weisslichgelben

Makel, an der Basis beginnend, ca. 3/4 der Lange einnehmend, (Naht schwarz),

auf den Schultern eine kleine, langliche, isolierte, schwarze Makel. Vorder-

beine gelb, nur die Schenkel auf der Oberseite dunkel, Mittelschienen leicht

aufgehellt.

Kopf (fig. 1) mit den Augen breiter als der Halsschild, Interokular-

aushohlung breit, von Auge zu Auge reichend, Vorderstirn eine fast dreieckige

Platte, welche zahnartig nach oben, zwischen die Augen gerichtet ist, Spitze

breit abgerundet, davor deutlich eingeschniirt, in der Mitte gegen den Clypeus

kaum eingedriickt, jederseits iiber den Fiihlerwurzeln mit einer kleinen,

langlichen, schwach erhabenen Beule. Fihler kraftig, fast so lang wie der

ganze Korper, 2. Glied knétchenf6rmig, 3. so lang wie das 4. Halsschild nur

wenig breiter als lang, Seiten vorne, fast bis zur Mitte, fast parallel, dann

verengt und vor der Basis etwas eingeschniirt, Scheibe in der Mitte leicht

aufgewolbt, Basis leicht quer eingedriickt und Basalrand etwas aufgeworfen,

Oberflache mikroskulptiert, fein staubartig behaart. Fliigeldecken nach

hinten kaum verbreitert, Oberflache fast glatt mit feinen, zerstreuten Haar-

punkten.

Lange: 2 mm.

Fundort: South West Africa, Abachaus, Otjivarongo 12. 1949, leg. G. Hobohm.

Holotypus in der Sammlung des Transvaal Museum, Pretoria.

309

360 ANNALS OF THE SOUTH AFRICAN MUSEUM

Neben D. cavifrons Boh. und andreaet m. zu stellen, die eine ahnlich

geformte Vorderstirn haben; durch die eigentiimliche Zeichnung der Fligel-

decken ist die neue Art leicht von den beiden anderen Arten zu unterscheiden.

Dinometopus swellendamensis nov. spec.

3S Schwarz, ausserste Kante des Seitenwalles der Interokularaushéhlung

schmal braun gesdumt, der mediane Zahn gegen die Spitze verschwommen,

schwach aufgehellt, ebenso die beiden Seitenplatten an der Basis des medianen

Zahnes uber den Fiihlerwurzeln. Fihlerglied 1 auf der Unterseite, 2 und 3

fast ganz, 4 etwas aufgehellt. Fliigeldecken mit einem ziemlich breiten,

gelblichbraunen, durchgehenden Querbande, an den Seiten etwas breiter als

an der Naht. Spitzen der Vorderschienen kaum aufgehellt.

Kopf (fig. 2) mit den Augen so breit wie der Halsschild, Interokularaus-

hohlung nicht sehr breit, die Augen nicht erreichend, Zwischenraum zwischen

Auge und Interokularaushoéhlung mit einem langlichen Eindruck, medianer

Zahn an der Basis sehr breit, fast dreieckig, fast in einer Flache mit der Vorder-

stirn legend, Spitze mit wenigen, langeren Haaren besetzt, Seitenplatten tiber

den Fithlerwurzeln wenig stark erhoht, flach, leicht eingedriickt, durch einen

tiefen, punktformigen Eindruck voneinander getrennt. Fuhler ziemlich lang,

fast von Ko6rperlange, 2. Glied so lang wie das 3. Halsschild breiter als lang,

Seiten stark gerundet verengt, gegen die Basis starker als nach vorne, Basalrand

bis zur Mitte der Seiten deutlich abgesetzt, Scheibe leicht gewolbt, Punk-

tierung kaum wahrnehmbar. Fliigeldecken nach hinten kaum erweitert,

erloschen punktiert.

Lange: 2 mm.

Fundort: South Africa, Tradouw Pass 4,000 ft., Swellendam District,

Oct. 1925, leg. K. H. Barnard. Holotypus in der Sammlung des South African

Museum, Cape Town.

Durch die gelbbraune, durchgehende Quermakel auf den Fliigeldecken,

welche an der Naht nicht unterbrochen ist, und den vollstandig schwarzen

Halsschild von den anderen Arten leicht zu unterscheiden.

Colotes hesset nov. spec.

3S Kopf gelborange, Augen und eine langliche, quere Makel, vom Hals-

schildvorderrand teils verdeckt, schwarzlich. Fihler gelb, ausserste Basis des 1.

Gliedes leicht angedunkelt, grésster Teil des 5., fast die ganze Oberflache des

6., 7. und 8. mit einem immer kleiner werdenden Flecken, schwarz. Hals-

schild, Vorder- und Mittelbeine gelb; Hinterbeine und Abdomen schwarz.

Fligeldecken tiefblau, leicht griinlich schimmernd, Seiten in der Mitte ziem-

lich breit, weisslichgelb gesaumt, unter den Schultern beginnend bis vor den

Spitzen.

Kopf (fig. 3) mit den Augen etwas breiter als der Halsschild, Stirne stark

verdickt, in der Mitte zwischen den Augen mit einem deutlichen Langs-

BEITRAG ZUR KENNTNIS DER MALACODERMATA (COL.) AFRIKA’S 361

eindruck, Stirne neben und unter den Augen ausgerandet, wodurch dieselben

stark vom Kopf abstehen, Oberflache fein gewirkt, matt. Fihler nicht sehr

lang, 1. Glied langlich, gegen die Spitze allmahlich verdickt, ungefahr so lang

wie das 3. und 4. zusammengenommen, fast langer als breit, 3. etwas langer

als das 4., 5. eine Spur langer und etwas breiter als das 4., 6. und folgende unter

sich gleich lang und gleich breit. Halsschild breiter als lang, Seiten vorne

zuerst fast parallel, dann gegen die Basis fast geradlinig verengt, Oberflache

fast noch feiner als der Kopf chagriniert, matt. Flitigeldecken nach hinten

leicht erweitert, Oberflache fein chagriniert, matt, dazwischen fein, zerstreut

punktiert.

Lange: 2,2 mm.

Fundort: South Africa, Mfongosi, Zululand. Holotypus in der Samm-

lung des South African Museum, Cape Town. Herrn Dr. A. J. Hesse

gewidmet.

Die Bildung des Kopfes erinnert etwas an C. bigibbosus m., doch ist die Stirne

bei der neuen Art noch viel starker angeschwollen und neben den Augen

ausgerandet. Die Farbung ist vollstandig verschieden.

Colotes nasifrons nov. spec.

6 Kopf schwarz, vordere Halfte des Langswulstes auf der Stirne und

Wangen um die Fihlergruben herum, weisslichgelb. Fihler gelb, vom 5.

oder 6. Gliede an leicht gebraunt. Halsschild und Beine gelborange, ersterer

mit einer schwarzbraunen, kaum langeren als breiten Makel, den Vorderrand

bertihrend, die Basis nicht erreichend. Fliigeldecken blaugriin metallisch,

Seiten in der basalen Halfte schmal, weisslichgelb gesaumt.

Kopf (fig. 4) mit den Augen etwas schmdler als der Halsschild, kurz nach

der Basis bis zum Clypeus mit einem ziemlich breiten, stark erhaben Langs-

wulst in der Mitte, der vor den Augen am starksten erhaben ist, in der Mitte

auch etwas breiter als an den Enden, Oberflache des Kopfes fein mikro-

chagriniert, matt. Fihler etwas langer als der halbe Korper, 1. Glied

langer als das 2. und 3. zusammengenommen, an der Basis sehr schmal, gegen

die Spitze knétchenartig verdickt, 2. etwas langer als breit, viel schmaler als

das 1. an der Spitze, 3. ungefahr so lang wie das 4. und folgende, unter sich

gleich dick. Halsschild gut um die Halfte breiter als lang, Seiten stark gerundet,

Basalecken etwas starker verrundet als die vorderen, Rand in den Basalecken

etwas deutlicher, Oberflache fein mikrochagriniert, matt. Fliigeldecken fein,

ziemlich dicht punktiert.

Lange: 2 mm.

Fundort: South Africa, Algoa Bay, Capland, leg. Dr. Brauns. Holotypus

in de Sammlung des South African Museum, Cape Town.

Neben C. frontalis Champ. zu stellen, von dem die Art durch die

verschiedene Farbung der Fithler und Bildung des Kopfes zu unterscheiden

ist.

362 ANNALS OF THE SOUTH AFRICAN MUSEUM

Aitalus kocht nov. spec.

§ Einfarbig schwarz mit schwachem griinlichem Schimmer, erste 2 bis 3

Fiihlerglieder, manchmal auch die Basis des 4. und 5, Vorderschienen, oft

auch alle Tarsen, Mittel- und Hinterschienen seltener, rétlich oder gelb-

lichrot.

Kopf mit den Augen kaum breiter als der Halsschild, Stirne fast flach,

zwischen den Augen, nach vorne, zwei erloschene Eindriicke, Oberflache fein

mikroskulptiert. Fihler die Schulterbeulen iiberragend, Glieder vom 3. an

stumpf gezahnt, vom 4. an etwas breiter als lang, Halsschild breiter als lang,

Seiten gegen die Basis stark verrundet verengt, Basalecken mit der Basis

vollstandig verrundet, Scheibe leicht gewodlbt, mikroskulptiert, fein, ziemlich

lang, sparlich, weisslich behaart. Fligeldecken 1 bis 2 Abdominalsegmente

unbedeckt lassend, nach hinten nur wenig verbreitert, erloschen punktiert,

fast glatt, weisslich, leicht abstehend, wenig dicht behaart.

Lange: 2,2 mm.

Fundort: South West Africa, Otjiwarongo 4.1950 leg. C. Koch (Holo-,

Allo- und Paratypen) ; Abachaus (Damaraland) 12.1951 leg. G. Hobohm (Para-

typen) in der Sammlung des Transvaal Museum, Pretoria. 7

Neben A. sulcicollis Champ. zu stellen, die Fihler der neuen Art sind

starker gezahnt, die Fliigeldecken weniger stark metallisch glanzend, die

Gestalt kleiner, weiss behaart, bei sulczcollis schwarz.

Hedybius (s.str.) hobohmi nov. spec.

6 Schwarz, Kopf gelb, dusserste Basis des Scheitels, meist vom Hals-

schildvorderrand verdeckt, seitlich bis zum Hinterrand der Augen vorstossend,

schwarz; Fihler ebenfalls schwarz, erste 4 Glieder gelb, 1., 3. und 4. mit

einem mehr oder weniger grossen, dunkeln Flecken auf der Oberseite.

Kopf (fig. 5) mit den Augen so breit wie der Halsschild an seiner breitesten

Stelle, zwischen den Augen tief und breit ausgehohlt, in der Mitte eine lange,

schmale, ziemlich spitz auslaufende, schrag aufstehende Lamelle, Vorderstirn

fast herzformig, stark abstehend, in der Mitte tief langsgefurcht, jederseits in

einen diinnen, etwas kiirzeren Fortsatz ausgezogen als die M§ttellamelle.

Fihler langer als der halbe K6rper, Glieder vom 4. an schwach, stumpf

gezahnt, 3. etwas langer als das 4. Halsschild breiter als lang, etwas vor der

Mitte am breitesten, Seiten an dieser Stelle schwach stumpfwinklig vor-

springend, Oberflache leicht gewolbt, fast glatt, fein greis behaart. Fligel-

decken nach hinten kaum verbreitert, Punktierung zerstreut, wenig tief,

Behaarung ziemlich dicht, grob, schrag abstehend. Vordertibien leicht

gekriimmt und vor der Basis innen, schwach ausgerandet.

3 Fuhlerglieder gelblich. Kopf einfach, zwischen den Augen erloschen,

BEITRAG ZUR KENNTNIS DER MALACODERMATA (COL.) AFRIKA’S 363

Erklarungen zu den Abbildungen:

Fig. 1 , Kopf von Dinometopus humeropictus n. sp. 3. Fig. 2, Kopf von Dinometopus swellendamensis

n. sp. g. Fig. 3, Kopf von Colotes hessei n. sp. 3. Fig. 4, Kopf von Colotes nasifrons n. sp. 3.

Fig. 5, Kopf von Hedybius (s. str.) hobohmin. sp. g. Fig. 6, Kopf von Philhedonus minutedentatus

Nn. sp. d.

364. ANNALS OF THE SOUTH AFRICAN MUSEUM

hufeisenformig eingedriickt. Halsschildseiten weniger stark vorstehend in der

Mitte als beim g. Vorderschienen einfach.

Lange: 2,7—3 mm.

Fundort: South West Africa, Abachaus, Damaraland 12.1951 leg. G.

Hobohm, Holo- Allo- und Paratypen; Abachaus, Otjivarongo 12.1949 leg.

G. Hobohm. in der Sammlung des Transvaal Museum, Pretoria. Dem

Entdecker gewidmet.

Durch die einfarbig schwarze Farbung von Halsschild und Fliigeldecken

von allen anderen bisher beschriebenen Arten der Gattung verschieden.

Hedybius (Subg. Hedybiinus) simpliciceps nov. spec.

6 Kopf schwarz, vom Vorderrande der Augen an, in gezackter Linie,

orangerot. Fiuhler schwarzbraun, erste 2 Glieder vollstandig orange, 3. bis 5.

auf der U'nterseite orange, oben und seitlich mit zunehmender, schwarz-

brauner Farbung, dunkle Farbung nicht scharf abgetrennt von der hellen,

allmahlich ineinander tbergehend. Halsschild orangerot mit breitem, durch-

gehendem, schwarzem Langsband oder mit schwarzer Makel, die Basis nicht

bertthrend. Schildchen schwarz. Fliigeldecken mit dunkelgriinem oder

blaulichem Metallschimmer. Beine gelb, nur der Kamm am 2. Gliede der

Vordertarsen schwarz, manchmal die Hintertarsen angedunkelt.

Kopf mit den Augen kaum so breit wie der Halsschild, Stirne ziemlich

flach und glatt, nur die Seiten der Stirn am Augenoberrand ein wenig erhoht,

zwischen den Augen ein schwacher Quereindruck, der in der Mitte durch

einen kleinen dreieckigen Eindruck unterbrochen ist; der quere Eindruck

zwischen den Augen trennt gleichzeitig den glatten oberen Teil der Stirne

vom leicht kérniggewirkten unteren, gegen den Clypeus wird die Vorderstirn

wieder ganz glatt. Fihler die Koxen der Hinterbeine knapp erreichend,

Glieder stumpf gezahnt, 3. so lang wie das 4. Halsschild breiter als lang,

Seiten gerundet verengt, Basalecken mit der Basis vollstandig verrundet,

Scheibe leicht gewélbt, glatt, mit einzelnen aufrechtstehenden, langen Haaren

besetzt. Fliigeldecken dicht, nicht sehr grob punktiert, Behaarung dicht,

schwarz, aufstehend.

Lange: 4 mm.

Fundort: South Africa, Richmond District, Cape Province 3. 1931 leg.

Museum Staff. Holo-, Allo- und Paratypus in der Sammlung des South

African Museum, Cape Town, 2 Paratypen in meiner Sammlung.

Die Art gleicht in der Farbung etwas dem H. aulicus F., nur dass die Beine

bei der neuen Art gelb sind. Sie kann mit keiner anderen Art verwechselt

werden, weil der Kopf fast keine Geschlechtsmerkmale aufweist.

Philhedonus minutedentatus nov. spec.

d Kopf, Beine und Fihler schwarz, nur bei letzteren ist die Spitze des

1. Gliedes und die Unterseite des 2. und 3. mehr oder weniger aufgehellt.

BEITRAG ZUR KENNTNIS DER MALACODERMATA (COL.) AFRIKA’S 365

Halsschild orangerot mit einer schwarzen Makel in der Mitte, den Vorderrand

beriihrend, basal ein Drittel freilassend. Fliigeldecken dunkelgriin mit

schwachem bronzefarbenem Schimmer.

Kopf (fig. 6) mit den Augen schmaler als der Halsschild, Stirnbasis in der

Mitte unter und etwas vor dem Halsschildvorderrand ausgehdhlt und mit

einem kleinen Zahn versehen, der in der Mitte in einer Ausbuchtung des

Vorderrandes des Halsschildes steht, davor jederseits ein kleiner Zahn (nicht

immer gut zu erkennen). Seitlich zieht sich die mediane Aushohlung gegen die

Schlafen und verbreitert sich iiber den Augen stark, den Oberrand derselben

bertthrend. Oberflache fein mikroskulptiert, matt, mit vereinzelten kurzen,

dicken, silberweissen Haaren besetzt. Fiihler kurz, stark gezahnt, 3. Glied

langer als breit, 4. und 5. eher breiter als lang, folgende ungefahr so lang wie

breit. Halsschild breiter als lang, Seiten gegen die Basis gerundet verengt,

Basalecken vollstandig mit den Seiten verrundet, Vorderrand in der Mitte

ausgerandet und etwas eingedriickt, Oberflache fein mikroskulptiert, matt, mit

vereinzelten, kurzen, silbrigweissen Haaren. Fliigeldecken schwach erloschen

gerunzelt skulptiert, starker als der Halsschild behaart.

Lange: 3,5 mm.

Fundort: South Africa, Bushmanland, leg. Alston. Holotypus in der

Sammlung des South African Museum, Cape Town, Allotypus in meiner

Sammlung.

DASYTIDAE

Dasytes crenicollis nov. spec.

Schwarz, Halsschild und Fligeldecken oft mit sehr schwachem, griin-

lichem oder blaulichem Schimmer, Spitze des 1. Fihlergliedes, 2. bis 4.,

Spitze der Schenkel, Tibien und Tarsen rotgelb.

Kopf mit den Augen eher etwas schmaler als der Halsschild, Stirne mit

zwei seichten Langseindriicken zwischen den Augen, Oberflache dicht mit

kleinen, runden, narbenartig erhéhten Punkten bedeckt, die in der Mitte

eingedriickt sind. Behaarung sparlich, kurz, silberig. Fihler die Schulter-

beulen nicht ganz erreichend, 1. Glied ziemlich stark verdickt, um die Halfte

langer als breit, 2. kaum kiirzer als das 1., aber schm4ler, 3. und 4. lang und

schmal, 3. langer als das 4., vom 5. an gegen die Spitze etwas verdickt, 6. und

foleende etwas starker verdickt als das 5., 10. nur wenig langer als breit, 11.

fast doppelt so lang wie das 10. Halsschild schmaler als die Fliigeldecken an

den Schultern, etwas breiter als lang, Seiten gerundet, gegen die Basis etwas

starker verengt als nach vorne, Seitenrand fein gekerbt, Scheibe gewélbt,

Oberflache wie der Kopf, jedoch weniger stark skulptiert, Haare kurz, wie auf

dem Kopfe, etwas nach dem Zentrum des Halsschildes gerichtet. Fliigel-

decken nach hinten kaum verbreitert, ziemlich tief, dicht, verworren punk-

tiert, Behaarung kurz, nicht sehr dicht, gelblichweiss. Tarsen lang und diinn,

fast so lang wie die Schienen.

Lange: 2-3 mm.

366 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fundort: South Africa, Seven Weeks Poort, Ladismith, Cape Province,

Nov. 1935. Holo- und Allotypus in der Sammlung des South African Museum,

Cape Town, 2 Paratypen in meiner Sammlung.

Die Art stelle ich provisorisch in die Gattung Dasytes, da sie wie die meisten

bisher aus Siidafrika beschriebenen Formen dieser Gattung einer Revision

bediirfen, sobald mehr Material vorliegt. Der kleine Halsschild, dessen Seiten

gekerbt sind, erinnert etwas an Danacaea.

14. Some new Caddis Flies (Trichoptera) from the Western Cape Province—I. By

K. M. F. Scott, Ph.D., F.R.E.S., Department of Zoology, University of

Cape Town. (With six text-figures.)

SUMMARY

A new species, Petroplax curvicosta (family Sericostomatidae, genus

Petroplax Barnard), is described, together with its larva and pupa. The young

stages of P. prion Barnard are also described, and a fuller account given of

the larva and pupa of P. caricis Barnard (briefly described by Barnard, 1934,

p- 319). The generic diagnoses of the larvae and pupae are revised, and the

wing venation of the imagos discussed. A key to the five known species of

Petroplax (based on ¢ imagos) is appended, also a series of sketches to facilitate

the identification of the females (of which only four are known).

INTRODUCTION

During the past three years a survey of the Great Berg River was carried

out from the Zoology Department of the University of Cape Town by Mr.

A. D. Harrison of the Council for Scientific and Industrial Research. The

author was associated with some of this work, and has pleasure in acknow-

ledging grants from the C.S.I.R. (first a Research Assistantship and recently

a Senior Bursary) which made this possible and which have enabled her to

continue work on some of the animals collected, particularly the Trichoptera

and Chironomidae. Grateful thanks are also due to Mr. Harrison for much

assistance; to Professor Day of this Department for criticism and advice, and

also to Dr. K. H. Barnard for his kindly interest in this work, as well as for the

loan of specimens and literature in the South African Museum.

In the course of the survey a considerable amount of caddis material

accumulated; this includes many larvae and pupae as well as imagos caught

in the vicinity of the river. Several of the caddis appear to be new species; in

other cases known species have been correlated by breeding them out from

larvae or pupae in the laboratory. The new forms will be described in this and

subsequent papers.

Two papers by Dr. K. H. Barnard (1934 and 1940) provide a firm founda-

tion for further studies of South African caddises, and have been used as a

basis for the present work. Reference should be made to these papers for a

complete bibliography of earlier and contemporary work; also for descriptions

367

368 ANNALS OF THE SOUTH AFRICAN MUSEUM

of families, genera and known species and keys, none of which will be repeated

here unless they have been altered to cover new species. Barnard’s techniques

have been used for the denudation of wings and clearing of genitalia, and the

same terminology has been used, though in the case of the male genitalia the

terms used by Mosely and Kimmins (1953) have been added in parentheses.

The wing notation used is also the same: that of Tillyard.

The holotypes of new species will be added to the South African Museum

collection, and paratypes will be sent to the British Museum (Natural History)

wherever possible.

Genus Petroplax Barnard

1934. Petroplax Brnrd, Trans. Roy. Soc. S. Afr., vol. XXI, Part 4, p. 316.

Remarks. The new species described, P. curvicosta, agrees with the generic

characters except in certain aspects of the wing venation. In the generic

diagnosis the venation is described as follows: ‘In forewing apical forks, 1,2,5,

in 9 also fork 3, i.e. M simple in 4, forked in 9. Thyridial cell very long.

1A and 2A in @ joining distally near hind margin, in 2 proximally. No cross

vein between Cuz and 1A in 9. In hind-wings forks 1,2,5. Discoidal cell open.’

In P. curvicosta (fig. 1A, B, C) forks 1 and 2 are present in the 3 fore-wing,

fork 5 is, however, absent as Cui does not reach the margin but joins an

anastomosis connecting the arculus to Mg + 4. In the @ fore-wing forks 1, 2

and 3 are present, and a false 5 between Cur and Cuz. Thus Mi and M2 are

separate in the 2 but not in the g, and Mg + 4 is present in both sexes. Cur

is not branched in either sex. The thyridial cell is long, and 1A and 2A join

proximally in the 9, as in the generic diagnosis; in the ¢ however 2A runs

into the hind margin midway to the arculus. There is no cross-vein between

Cu2 and 1A in the 9; there are, however, indications of a cross-vein between

Cur and Cu2 in both g and 9. In the hind-wing forks 1, 2 and 5 are present

in both sexes as in the generic diagnosis, but there are definite indications of

the presence of a cross-vein closing the discoidal cell, also of one joining

Mg + 4 to Cuia.

In consequence of these differences a careful examination was made of

specimens of P. prionit Brnrd and P. caricis Brnrd from our collection, as well as

specimens kindly loaned by the South African Museum of both these species

and of P. phleophila Brnrd. It appears from this examination that in the ¢ fore-

wing in these species Cui and Cua do not normally reach the hind margin but,

together with 1A, join an anastomosis running from the arculus to M3 + 4;

thus only forks 1 and 2 are present. The only other known species of Petroplax

is P. anomala Brnrd, whose venation (Barnard 1940, fig. 12a) is quite different;

in it forks 1, 2 and 4 are present, and the base of 3 (M1 and M2 are separate

but Me joins Mg and fails to reach the margin); Cui does reach the margin

but remains unbranched, so that fork 5 is absent in this species too. In the fore-

wing of the known females forks 1,2 and 3 are always present, also a false 5

between Cur and Cuz. There is no cross-vein between Cu2 and 1A. In the

pans

a ae ge

SOME NEW CADDIS FLIES FROM WESTERN CAPE PROVINCE 369

Fic. 1. Petroplax curvicosta sp. n., 3 and ? imagos.

A,B, Fore and hind-wings of g. (, fore-wing of 9. D,E, dorsal and lateral views of ¢

genitalia. F, ventral view of gth sternite of J. G,H,J, dorsal, lateral and ventral views of 2

genitalia.

370 ANNALS OF THE SOUTH AFRICAN MUSEUM

hind-wing forks 1,2 and 5 are always present, and the discoidal cell is usually

closed, either faintly or distinctly; it is, however, open in g4 of P. caricis and

P. priont. A cross-vein is usually present between Cura and M3 + 4.

It is therefore suggested that in the generic diagnosis the description of the

wing venation should be amended as follows: ‘In fore-wing apical forks 1 and 2

always present, in 9 also fork 3 and a false 5. Thyridial cell very long. 1A and

2A in 8 join proximally, in ¢ 2A may join 1A distally or may run into margin

before reaching arculus. No cross-vein between Cuz and 1A in 9. In hind-

wing forks 1,2,5. Discoidal cell may be open or closed in ¢ but is closed in 9.’

In examining the specimens it was noticed that a pair of eversible mem-

branous processes occurs on the face of the g, between the maxillary palps, in

P. prion, P. phleophila and P. caricis. ‘These were not seen in P. curvicosta, but

may possibly be present though not everted in the three males available.

Key to species of Petroplax (33)

1. Bend in fore-wing; fur-like sensory patch on basal joint of antenna;

androconia absent: P. curvicosta sp. n.

No bend in fore-wing; no fur-like sensory patch on antenna;

androconia present or absent: BR ens 6%,

2. Cug2 and 1A suppressed in fore-wing; Cur reaches margin; large

patch of androconia. P. anomala Barnard.

Cuz and 1A present in fore-wing (though Cur and Cue do not

usually reach the margin) Sicsuist eee

3. Upper penis cover short, not deeply cleft; 2 recurved spines at

sides as well as pair on penis; no androconia. P. phleophila Barnard.

Upper penis cover long and deeply cleft; no recurved spines at

sides (pair present on penis as usual); small patch of androconia

4. Strong brush of setae on claspers; spines on penis rather short

(less than one-third length of upper penis cover). P. prionit Barnard.

No brush of setae on claspers; spines on penis very long (as long

aS upper penis cover). P. caricis Barnard.

LARVAE AND PUPAE OF Petroplax SPECIES

Barnard (1934, pp. 318 and 3109, fig. 13 j-q) gave brief descriptions of the

larva and pupa of P. caricis Brnrd. These were accompanied by drawings of

some of the parts, but the whole larva was not drawn. The greater part of the

larval and pupal descriptions was given in the generic diagnosis, as this was the

only species whose young stages were known. Thus no full description of any

larva or pupa of the genus has so far been available. Since the identification

of larvae and pupae without the necessity of breeding out adults is of great

importance for the work of the stream survey, it seemed desirable to include a

full description of the young stages of both P. curvicosta and P. caricis in this

paper. In the case of P. prionii the description of the larva is necessarily brief

as it is based on larval remains extracted from the pupal cases. Whole larvae

of P. curvicosta and P. caricis were, however, available, and pupae or pupal

SOME NEW CADDIS FLIES FROM WESTERN CAPE PROVINCE 371

pelts and cases of all three species. The young stages of P. phleophila and

P. anomala are as yet unknown.

The study of these additional larvae and pupae makes it desirable to

amend the generic diagnosis slightly as follows:

Generic diagnosis of larva of Petroplax Barnard (1934, p. 316, emend.)

Mandibles with internal tufts, may be feeble in left mandible. No prosternal

spine. Mesonotum less strongly chitinized than pronotum, metanotum mem-

branous. Hind leg longest. Dorsal tubercle on abdomen segment 1 obsolete.

Fic. 2. Semi-diagrammatic sketches of 9 genitalia of Petroplax spp.

(Dorsal views drawn to same scale with the aid of a micrometer

eye-piece. )

A, P. caricis Brnrd. _B, P. phleophila Brnrd. __C, P. curvicosta sp. n.

D, P. prionit Brnrd.

Simple filiform gills on segments 2-8 (dorsal, lateral and ventro-lateral).

Lateral line represented only by minute sclerotized tubercles on segments 3-8.

Anal holdfast small. Case tubular, widening slightly towards mouth, composed

of fine sandgrains with a few larger grains at one or both ends.

Generic diagnosis of pupa of Petroplax Barnard (1934, p. 316, emend.)

Labrum nearly as long as broad, rounded. Mandible strong, serrulate on

inner margin. Mid-tarsus fringed. First abdominal segment with transverse

thickening, sometimes followed by a band of minute scabrosities. Simple

presegmental gills present on segments 2-8 (dorsal, lateral and ventro-

lateral). Lateral line present on segments 6,7,8 and posterior part of 5.

Presegmental dorsal plates on segments 3-6, postsegmental also on segment 5,

the latter smaller than the former. Anal appendages lanceolate, apices acute

and upturned. Case somewhat similar to that of the larva, but apparently

usually added to at the front and cut down at the hind end; anchored fore and

aft to stones by silken threads. Membrane closing posterior end partly covered

372 ANNALS OF THE SOUTH AFRICAN MUSEUM

with sand-grains and perforated by a narrow slit (the pupa lies with its ventral

surface towards the concave side of the case, and the slit at the hind end is

vertical, i.e. dorso-ventral). Front end of case closed by a convex lid more or

less covered with sand-grains and perforated by a narrow, curved slit which

lies at right angles to the one at the base.

Petroplax curvicosta sp. n.

Fig. 1 A-J.

A golden-brown species with a bend in the costa of the fore-wing and a

fur-like patch of setae on the basal joint of the antenna in the male imago.

Imago (in alcohol). Characters as given in the generic diagnosis (Barnard

1934, p. 316) except as indicated above in the remarks. Head: face yellowish,

vertex dark brown, head bearing warts from which spring long thickened

yellow or brownish setae; a few of the setae are black and spine-like. Antennae

annulate, brown and yellow; basal joint in g broad, almost triangular in lateral

view, densely clothed on three sides with a fur-like mass of stout sepia-coloured

setae. Maxillary palp 3 2-jointed, setose, short and upturned; basal joint

bulbous, brownish; 2nd joint very small, pale, attached to the dorsal side of

the first. Maxillary palp 2 5-jointed, pale yellowish-brown. Labial palps pale

yellowish-brown. Thorax: prothorax fuscous, bearing hairy warts; meso- and

metathorax dark chestnut brown, shining, nearly hairless except for two pairs

of raised patches in the centre of the mesothorax; membranous parts cream.

Legs: tibial spurs 2,2,4; femora and tibiae brownish-yellow; tarsi annulate

yellow and brown; tibiae and tarsi bear a number of short dark spines.

Abdomen: tergites light sepia, sternites paler, pleura cream.

Wings: 3 5:5-6°5 mm.; 2 6-0-6-7 mm. (fig. 1 A-C). Fore-wing: mem-

brane pale brown with darker neuration, except for costa which is white in 9;

hyaline areas indicated by dotted lines in figure (not as clear as in the other

species as the membrane is paler). Wings densely pubescent. In the 3 short

thickened pale gold setae form two broad streaks, one over the cubitals, the

other along the anterior border of the wing; the rest of the wing is covered with

longer brown and gold hairs. Fringe brown and gold. 9: whole fore-wing

mingled brown and gold; many erect hairs on the proximal part of the wing.

Pterostigma present in both sexes, also two faint white patches of hair over the

hyaline areas. Hind-wings: iridescent, fuscous, thinly pubescent, with long

hairs along Cur as far as fork and along proximal part of 1A, also a tuft at

base of 1A. Fringe long. An oval bare patch at the base of M, proximal to the

junction with Cu. Venation: ¢ fore-wing (fig. 1A) with costa thickened and

bent near the middle; forks 1 and 2 only. 9 fore-wing (fig. 1C) with forks 1,2,3

and (5). Hind-wings (fig. 1B) show a faint vein closing the discoidal cell, and

one joining M3 + 4 and Cutia; forks 1,2,5, present; g and 2 hind-wings

similar,

SOME NEW CADDIS FLIES FROM WESTERN CAPE PROVINCE 373

why

Yyy

Fic. 3. Larva of P. curvicosta sp. n.

A, Case of larva. B, lateral view of whole larva. CC, head capsule from behind, showing gular

sclerite. D, clypeus and anteclypeus. E, dorsal view of anterior end of larva. F, posterior

end of larva (left side). G, head and pronotum from front, with antenna enlarged. H, anal

claw, showing variations. J,K, left and right mandibles. L, labrum. M, labium and

maxilla. N, prothoracic leg, with plaque d’appui. P, metathoracic leg.

374 ANNALS OF THE SOUTH AFRICAN MUSEUM

Genitalia: 3 (fig. 1 D-F). Pre-anal (superior) appendages club-shaped,

dark, setose; dorsal plate triangular, the apex bifid to form two short points.

Upper penis cover apically slightly cleft, apices rounded; a pair of downwardly

directed, recurved, strongly chitinized processes at the sides and slightly

proximal to the cleft; proximal to each of these a tuft of setae. Penis with 2 long

stout sub-terminal chitinized processes; tip of penis slender. Claspers (inferior

appendages) with 2 branches, the upper lobate, the lower narrower, both

incurved and setose. goth sternite (ventral plate) with two long narrow

processes. A spatulate ventral process on the 7th sternite, almost semicircular

in shape. @ (fig. 1 G,H,J). Supra-anal plate cleft, lobes apically rounded

and with a slight dorsal keel; cleft narrow. Vaginal structure strongly

chitinized.

Locality: Great Berg River, Groot Drakenstein (A.D.H. and K.M.F.S.)

November 1953, 2 gd and 4 99, bred out in laboratory from pupae collected

under stones near margin out of the main current, found in the deeper pools

and not in very shallow water. Also 1 ¢ imago caught in the vicinity. Larvae

were collected at Groot Drakenstein and at Driefontein, higher up the Berg

River; of these one larva was bred out to a 9 imago. Imagos emerge on the

surface of the water.

Remarks: In general features this species belongs to the genus Petroplax

Barnard, but is quite easily distinguished from the other four known species.

The ¢ may be identified at once by the bend in the wing and by the fur-like

patch of setae on the basal joint of the antenna; the genitalia are very similar

to those of P. phleophila but are more heavily chitinized. The 9 genitalia show a

general resemblance to those of the other species, but the four known species

are easily separable by comparing the proportions of the supra-anal lobes and

the width of the cleft between them. (See fig. 2 A—D.)

Larva (fig. 3 A-P): Described from entire larvae and compared with

larval remains extracted from pupal cases. Larva eruciform; head hypogna-

thous; body slightly arched; legs and thorax very hairy. Length up to 64 mm.

(pupae up to 8 mm. were however found, so that 64 mm. is probably not the

maximum length); widest at metanotum and first abdominal segment. Head

rounded, chestnut brown in colour with a pattern of paler marks, surface of

chitin pitted in places; anterior margin yellowish, ridged, the two lateral

ridges each bearing a small upright antenna. Clypeus with one large and two

smaller pairs of lateral indentations and 13 bristles. Eyes prominent, sur-

rounded by pale areas. Gular sclerite triangular, fused with the genae except

at the oral margin, sutures visible. Mouthparts: labrum with a shallow central

excision and a transverse row of 6 long pale-coloured bristles; 3 pairs of blade-

like bristles along the anterior margin. Mandibles each with a broad cutting-

edge with 3 blunt teeth at one end; a brush of hairs on the inner side and a pair

of bristles on the ridged outer side; the left mandible bears an additional central

brush of short hairs. Maxilla bristly, maxillary palp 4-segmented and tipped

with papillae; maxillary lobe armed with knobs and sword-shaped spines and

SOME NEW CADDIS FLIES FROM WESTERN CAPE PROVINCE 375

Fic. 4. Larva of P. caricis Brnrd.

A, Case of larva (actual size slightly longer than larva). B, lateral view of whole larva.

C, head and pronotum from front, with antenna enlarged. D, clypeus and anteclypeus.

E, dorsal view of anterior end of larva. F,labrum. G, anal claw. H, posterior end of larva

(left side). J,K, left and right mandibles. L, labium and maxilla. M, metathoracic leg.

N, prothoracic leg and plaque d’appui. P, head capsule from behind showing gular sclerite.

376 ANNALS OF THE SOUTH AFRICAN MUSEUM

bristles. Labium conical, bearing 2 short palps. Thorax: pronotum hairy,

anterior part sclerotized, chestnut-coloured, posterior part yellowish with darker

chitinized spots; mesonotum hairy, only slightly chitinized, yellowish with

brown spots; metanotum membranous, with a yellow transverse bar on each

side of which is a row of long hairs. Legs: prothoracic legs short, heavily built,

hairy (except tarsus), edged ventrally with strong spines; plaque d’appui

oblong, truncate, with an upturned point. Meso- and metathoracic legs long,

slender and very hairy, metathoracic legs longest. Abdomen: white; flattened

lateral tubercles on segment 1. Simple filiform gills present as shown in

fig. 3 B (dorsal and lateral gills on segments 2 and 3; ventro-lateral on 2-8;

there also appear to be rudimentary lateral gills as shown—presegmental on

segments 4—7 and postsegmental on 3-7). Abdomen ends in a pair of raised

‘foot-like’ oval lobes, each bearing a single long strong black spine flanked by

2 or 3 smaller ones, the ‘foot’ edged along the ventral and inner borders with

short black spines. Anal claws with 3 barbs, the two smaller ones being

variable in size.

Case: A wide curved tube neatly made of sand-grains; posterior end a

truncated cone with a circular central aperture.

Pupa (fig. 6 D, G, K, N): Labrum rounded; mandibles with outer edge

angular, a pair of setae near the angle; articular condyle on outer corner of

base. Antennae as long as body. Each dorsal plate may bear either two or

three hooks; minute scabrosities are scattered over the dorsal surface but not

concentrated in any particular part. Simple pre-

segmental gills present on segments 2-8 as in table TABLE I

I. Anal appendages fairly stout, hairy, apex short, .———————

slender and upturned, armed with several long black De ae

spines. —]----] ——

Case tubular, up to 8 mm. in length, neatly | I

formed of small sand-grains, with a number of larger | II XO eS

grains attached round each end; apertures in basal | III Oo ee

membrane and lid have serrated edges. The size and | IV XIX

number of stones on the lid and round the ends of the | V Xa

case seems very variable, and several large grains | VI xX | X

may be attached round the anterior end as in P. | VII XE

caricus (fig. 6 M); the type of sand-grain used for the | VIII X

walls, and the shape of the slits, seem however to be

constant.

Petroplax caricis Barnard

Figs. 4 A-P and 6 A,C,F,J,M.

1934. Barnard (larva and pupa: pp. 316 and 319, fig. 13 j-q). Larvae and

pupae collected by Dr. K. H. Barnard from Platteklip stream, 500 ft., Table

Mountain, Cape Peninsula.

ee ee

SOME NEW CADDIS FLIES FROM WESTERN CAPE PROVINCE 377

Larva: Eruciform, head hypognathous, body strongly arched; legs and

thorax very hairy. Length up to 8 mm.; body widest at metanotum and Ist

abdominal segment. Head rounded, chestnut brown in colour, patterned with

paler marks, surface of chitin pitted, anterior margins dark, heavily chitinized,

ridges, a small antenna on each lateral ridge. Clypeus with one pair of lateral

indentations and indications of a second pair; 13 (in one case 14) bristles. Eyes

prominent, surrounded by pale areas. Gular sclerite small, triangular, largely

fused with the genae. Mouthparts: labrum with a shallow central excision, a

transverse row of 6 long yellowish bristles, and 2 or 3 pairs of short peg-like

B Cc D

Fic. 5. Larva of P. prionit Brnrd.

A, clypeus and anteclypeus. B, labrum. C,D, left and right mandibles. E, antenna.

F, anal claw, showing variations. G, maxilla and labium. H, plaque d’appui.

bristles along the anterior margin. Mandibles each with a broad cutting-edge

rising to a single apical tooth, a short brush of hairs on the inner side (also a

smaller brush on the left mandible), and a pair of bristles on the ridged outer

side. Maxilla and labium similar to those of P. curvicosta. Thoracic nota also

much as in P. curvicosta, but pattern of chitinized spots slightly different.

Legs and plaque d’appui also very like those of P. curvicosta. Abdomen

whitish, lateral protuberances on 1st abdominal segment flattened. Simple

filiform gills present as shown in fig. 4 B: dorsal and lateral on segments 2 and

3; ventro-lateral on 2-8; also rudimentary lateral gills as indicated—pre-

segmental on segments 4-8 and postsegmental on 3-7. Abdomen ends in a

pair of oval lobes, each bearing a tuft of 3-4 long strong black spines, and

ringed with smaller spines and hairs. Anal claws with 3 barbs (the third may

be minute).

Case: A wide curved tube made of small sand-grains, with a few larger

grains attached to the membrane closing the posterior end and forming a ring

round a circular central aperture.

378 ANNALS OF THE SOUTH AFRICAN MUSEUM

Pupa: Labrum rounded; mandibles with outer margin rounded and

sinuous, ending in condyle; one pair of lateral setae. Transverse thickening

on 1st abdominal segment followed by a patch of small rounded scabrosities

(smaller scabrosities occur scattered over most of the dorsal surface; these are

more prominent than in P. curvicosta). Simple presegmental gills as in table II.

Dorsal plates each bear 2 hooks. Anal appendages

not as stout as in P. curvicosta, apices short, acute and TABLE II

upturned, bearing a few long black spines, also eee ene

smaller spines and hairs. Di} heey

Case tubular, made of sand-grains of two sizes,

smaller ones in the basal half and larger ones in the | I

apical half; a number of large sand-grains attached | II xX

round the posterior end partly covering the basal | III Xi: [ea

membrane. Lid convex and partly covered with | IV DG i. €

sand-grains; apertures in lid and bottom of case have | V eX

smooth edges. A few very large grains may be | VI XS

attached round the anterior end of the case. VII DSi eG

VIIl xX

Petroplax prion Barnard

Figs. 5 A-H and 6 B,E,H,L,P.

1934. Barnard (imago: p. 319 and figs. 14 h-l).

No identifiable larvae were included in the Berg River collection, but

several pupae were collected from the Berg River at Groot Drakenstein,

together with those of P. curvicosta, and bred out in the laboratory; the

imagos which emerged (2 gd, 1 2) were identified as P. prion Barnard.

Larval remains were extracted from the pupal cases and drawings. (fig. 5 A-H)

and the following brief description made from them.

Larva: Head sclerites chestnut brown with paler markings, area round

eye pale, anterior margins yellowish; the sclerites are very similar to those of

P. curvicosta. Small vertical antennae on the lateral ridges. Clypeus with 2

pairs of lateral indentations and 13 bristles. Mouthparts: labrum with a shallow

central excision flanked by 3 pairs of strong blade-like setae, on the dorsal side

a transverse row of 6 setae. Mandibles rather rounded, with a broad cutting

edge rising to a single tooth; a slight lateral ridge flanked by a pair of setae;

the left mandible with two small brushes of setae, the right mandible with one

large brush. Maxilla very bristly, setae thickened and with serrated tips;

maxillary palp 4-segmented and tipped with papillae; maxillary lobe with

strongly developed spines of several types; labrum conical, bearing 2 short

palps. Legs, thoracic nota and plaque d’appui very similar to those of the other

two species. Abdomen appears to end in lobes studded with heavy spines and

setae much as in the other species. Anal claws with 3 or 4 barbs; the smallest

barb may be closely applied to the others and difficult to see.

SOME NEW CADDIS FLIES FROM WESTERN CAPE PROVINCE 379

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Fic. 6. Pupae of Petroplax spp. (Mandibles, labra and anal processes all drawn to same scale

with the aid of a camera lucida attachment.)

A, lateral view of pupa of P. caricis Brnrd. B, pre- and post-segmental dorsal plates from 5th

abdominal segment of P. prionti. C,D,E, mandibles of P. caricis, P. curvicosta and P. prionii.

F,G,H, labra of P. caricis, P. curvicosta and P. prionit. J,K,L, anal processes of P. caricis, P. curvi-

costa and P. prionii. M,N,P, pupal cases of P. caricis, P. curvicosta and P. prionit showing also

membranes from hind ends of cases, lids, and slits in lids enlarged.

380 ANNALS OF THE SOUTH AFRICAN MUSEUM

Pupa: Labrum rounded; mandibles with outer margin rounded, ending in

condyle, one pair of setae. Antennae as long as body. Transverse thickening

on ist abdominal segment followed by a patch of small scabrosities, smaller

scabrosities scattered over the dorsal surface (these are smaller than in P.

caricis, but larger than in P. curvicosta). Simple presegmental gills on segments

2-8 as in table III. Each dorsal plate normally bears 2 hooks, but occasionally

one hook is double. Anal appendages stout, bearing

long black spines, shorter spines and hairs. Apices TABLE III

acute and upturned, longer than in the other 2 = ————————

species. De) ey

Case: Tubular, leneth up to 8 mm., neatly formed! | nem ieee ee

of small sand-grains, usually with a band of larger I

grains round the upper end; a number of large sand- i zi x

grains attached to the basal membrane closing the It xX |X |x

posterior end; slit straight with crenate edges. Lid Ny x |X

convex, made of sand-grains neatly fitted together Mi x | Xx

with silk, central slit curved and with crenate edges. va x | Xx

As in the other species several very large sand-grains a x .

may be attached round the anterior end of the case.

x is rudimentary.

REFERENCES

Barnard, K. H., 1934. ‘South African Caddis Flies (Trichoptera).’ Trans. Roy. Soc. S. Afr.,

XXI, Part 4, pp. 291-392, 52 text-figs.

Barnard, K. H., 1940. ‘Additional Records, and Descriptions of New Species, of South African

Alder-flies (Megaloptera); Mayflies (Ephemeroptera); Caddis flies (Trichoptera); Stone

flies (Perlaria), and Dragonflies (Odonata).’ Ann. S. Afr. Mus., XXXII, Part 6,

pp. 609-61, 19 text-figs.

Hickin, N. E., 1946. ‘Larvae of the British Trichoptera.’ Trans. R. Ent. Soc. Lond., vol. 97,

Part 8, pp. 187-212, 15 text-figs.

Hickin, N. E., 1949. ‘Pupae of the British Trichoptera.” Trans. R. Ent. Soc. Lond., vol. 100,

Part 11, pp. 275-89, 10 text-figs.

Hickin, N. E., 1952. Caddis, a short account of the biology of British caddis flies with special reference to

the immature stages. London, Methuen & Co. Ltd., pp. 1-50, 47 text-figs, 4 plates.

Mosely, M. E., 1939. The British Caddis Flies (Trichoptera). London, G. Routledge & Sons.

Ltd., pp. 1-320, 631 text-figs.

Mosely, M. E., and Kimmins, D. E., 1953. The Trichoptera (Caddis Flies) of Australia and New

Kealand. British Museum, London, pp. 1-550, 364 text-figs.

15. Two new species of Boriomyia from South Africa (Neuroptera, Hemerobiidae).

By Bo TyeprErR, Falun, Sweden. (With fifteen text-figures.)

This paper forms the first report of a study of material of the order

Neuroptera, belonging to the South African Museum, Cape Town. My thanks

and acknowledgements are tendered to the Director and Dr. A. J. Hesse for

their confidence in entrusting to me the study of the rich collections of their

Museum.

Fic. 1. Boriomyia barnardin. sp. Holotype 3.

Boriomyia barnardi n. sp.

Figs. 1-9

Holotype 3. A dried and pinned specimen in good condition.

The frons is shining black. The vertex is yellowish brown. The mouth-

parts and the antennae are pale yellowish brown.

The pro- and mesonotum have a broad, yellowish brown median longi-

tudinal stripe, bordered laterally with blackish-brown. The metanotum is

blackish-brown; its scutellum a little paler, brownish. The legs are testaceous.

The tibiae of the anterior legs have three elongate, dark spots on their anterior

surface: one below the knee, one near the middle and one before the apex.

The fore-wings are elongate-oval. Their membrane is pale, with pale

fuscous sagittate markings and with large fuscous patches as indicated in fig. 1.

381

382 ANNALS OF THE SOUTH AFRICAN MUSEUM

The anal region is heavily dark coloured. The veins are blackish with pale

interruptions. ;

The hind-wings are pale with pale venation.

The abdomen has brownish tergites and pale testaceous sternites.

The ¢ genitalia (figs. 2-9). ‘The oth sternite (IX) is large and prominent,

rounded. The parameres (pa) are narrow with very acute apices, without

Fics. 2-9. Boriomyia barnardi n. sp. Holotype ¢.

2, apex of abdomen, lateral; 3, apex of right ectoproct, inside, in obliquely lateral aspect;

4, parameres, lateral; 5, ditto, dorsal; 6, gonarcus, lateral; 7, ditto, dorsal; 8. hypandrium

internum, lateral; 9g, ditto ventral.

superprocessus. ‘Their apophysis proximus (app) appears very broad (high)

when viewed laterally. The hypandrium internum (hyi) appears in dorsal

view nearly parallel-sided. The gonarcus (gs) has a very long mediuncus (mu),

the dorsal margin of which has two prominent teeth. Its apex is downwards

directed and acute. The entoprocessus (ent) are large and end each in a down-

wards and a little inwards directed very acute apex. The ectoprocts (epr) are

band-like, rather short and very broad in their proximal portion. Their apex

1 The terminology used here is that which I have recently brought forward in a paper

‘Genital structures and terminology in the order Neuroptera’ (Ent. Meddelelser, Copenhagen,

Pp. 22, 1954).

TWO NEW SPECIES OF BORIOMYIA FROM SOUTH AFRICA 383

is produced downwards-inwards and bears an internal, straight row of short

but strong teeth. The callus cerci is small and bears 18 trichobothria.

Size: length of body about 7 mm., of fore-wing 7,5 mm., of hind-wing

6,6 mm.

Female unknown.

Habitat: Cape Province, Kleinmond, February 1927; holotype 4, leg.

Dr. K. H. Barnard. In the South African Museum.

I have much pleasure in naming this species in honour of its discoverer.

This species belongs to the nervosa-group, represented in South Africa

p. 127, f. 2, 3, 1929). B. barnardi n. sp. is similar to B. nubila Kimm. but easily

distinguished by the ¢ genitalia.

Boriomyia fumosa n. sp.

Figs. 10-15

Holotype 9. A dried and pinned specimen in less good condition (the

head, the intermediate and the hind-legs are lacking).

The pro- and mesonotum have a broad, yellowish median longitudinal

stripe, which is blackish punctured. Their lateral borders are black. The

Fic. 10. Boriomyia fumosan. sp. Holotype 9.

metanotum is brownish black. The anterior legs are yellowish; their femora

have a distinct reddish-brown longitudinal stripe along the anterior surface;

their tibiae have a brownish ring below the knee and an ante-apical reddish-

brown elongate spot on the anterior surface.

The fore-wings are oval. Their membrane is smoky, the sagittate markings

being to a great extent confluent, especially in the distal portion and along the

384. ANNALS OF THE SOUTH AFRICAN MUSEUM

hind margin. There are dark brown shadings and spots as indicated in fig. 10.

The anal region is heavily dark-coloured. The venation is brownish black with

pale interruptions.

tip

ane

' =2

Fics. 11-15. Bortomyia fumosan.sp. Holotype 9.

II, apex of abdomen, lateral; 12, praegenitale and subgenitale with gonapophyses posteriores,

lateral; 13, ditto, dorsal; 14, ditto, ventral; 15, spermatheca.

The hind-wings are pale with pale venation.

The abdomen is uniformly brownish.

The @ genitalia (figs. 11-15). The oth tergite—viewed laterally—is in

its upper portion band-like; on level with the spiracle of the 8th segment it

suddenly broadens into a large and projecting lower portion. The subgenitale

(spg) is parallel-sided and has a small, triangular apex. The gonapophyses

posteriores (gp) are broad and pale, each ending in a rather acute apex.

Proximally they are united into a plate, lying over their bases. A small prae-

genitale (prg) is present, the shape of which is indicated in the figs. 12-193.

TWO NEW SPECIES OF BORIOMYIA FROM SOUTH AFRICA 385

The shape of the spermatheca (spm) is given in fig. 15. It should be noted

that its duct has three tooth-shaped extensions. The gonapophyses laterales

(gl) are almost triangular, their apex being rather narrow and subacute. The

ectoprocts (epr) are elongate with rather smoothly rounded apex. Their

cercal callus bears only 12 trichobothria.

Size. Length of body approximately 6 mm., of fore-wing 8 mm., of hind-

wing 7 mm.

The gis unknown.

Habitat: Cape Province, Grahamstown, February 1933; holotype 9, leg.

Dr. R. F. Lawrence. In the South African Museum.

Boriomyia fumosa n. sp. is a species of considerable interest because the

presence of a praegenitale has never been noted before in this genus. In other

respects the 2 genitalia resemble those of the subnebulosa-group. B. fumosa n. sp.

cannot, I am sure, be the female of B. barnardi n. sp. or B. nubila Kimm., which

both belong to the nervosa-group, in which group the subgenitale and the

gonapophyses posteriores of the females are differently shaped. It is to be

hoped that South African entomologists will succeed in finding the ¢ of

B. fumosa. Only when the ¢ has been studied can the definite position of the

species be stated.

16. Four new species of Gregarines from Mountain Cockroaches of the Cape Peninsula.

By A. D. Harrison. (With eleven text-figures.)

The Gregarines described in this paper were taken from the following

cockroaches indigenous to the mountains of the Cape Peninsula:

Aptera cingulata (Burm.) “Table Mountain Cockroach’, Temnopteryx

phalerata (Sauss.) and Melanosilpha capensis (Stal.).

Method of Study

The alimentary canal was removed from freshly killed insects, slit open

and extracted on a slide in 0°75 per cent solution of NaCl, in which all but one

of the species remained viable long enough for examination and measuring.

Only mature individuals were measured and, in cases where the gregarines

were associated in syzygy, only the anterior partner or primite was measured

as the protomerite of the posterior partner, or satellite, is flattened.

Permanent whole mounts of the species examined could not be prepared

using absolute alcohol, xylol and balsam as the specimens shrank badly during

the dehydrating process; glycerine-jelly mounts were fairly satisfactory but not

permanent. The best aqueous mounting medium was found to be ‘Gum

Chloral’. In this medium specimens were completely cleared without shrinkage;

in fact, specimens which had already shrunk during staining, resumed the

original shape when the slide was gently warmed. This medium sets hard.

Specimens were fixed in either Schaudin’s or Bouin’s fixative and stained

in Heidenhain’s iron haematoxylin which did not wash out in the aqueous

medium used.

It was found necessary to cut sections of gregarines zn sztu in the gut of the

host so that the epimerite and other features could be studied. The mid-gut

was removed from a freshly killed host and put immediately into fixative.

After fixing it was treated in hydrofluoric acid, to remove sand-grains, and

finally embedded in wax using the methyl benzoate-celloidin method. Sections

were stained in Delafield’s or Heidenhain’s iron haematoxylin and counter-

stained.

When sections were stained by the Feulgen method the nuclei of the

gregarines did not react though those of the host’s gut cells reacted strongly

and stained a dark purple. This supported the findings of previous workers.

As cysts of three of the species were discovered it was possible to study

their dehiscence. Previous workers have been very definite in stating that cysts

387

388 ANNALS OF THE SOUTH AFRICAN MUSEUM

do not dehisce under normal atmospheric conditions but only in air saturated

with moisture; some even keep them in a drop of water in a sealed cell while

others keep them in a moist chamber. Acting on the advice of Sprague, the

cysts were kept in sealed petrie dishes containing just enough water to keep

the air saturated; the cysts themselves were not in contact with the water.

Under these conditions the cysts dehisced satisfactorily. In the case of two of

the species they also dehisced in the open air of the laboratory which was fairly

humid, these investigations being carried out during winter and spring when

rain was frequent.

Classification

M. E. Watson (1916) gives a very full synopsis of the then-known Families

and Genera of the Tribe Cephalina, Delage (syn. Legion Septata, Lankester), of

the Sub-order Eugregarinae, Léger, based on the classification of Minchin and

Poche. In this synopsis she gives all the essential diagnostic points of each

genus. This table was used for classifying these gregarines.

All the species fell into the family Gregarinidae Labbé, for which Watson

gives the following characteristics: associative or solitary, satellite with

septum, epimerite symmetrical and simple, cysts with or without spore ducts.

Three definitely, and one almost certainly, fell into the genus Gregarina Dufour,

for which Watson gives the following characteristics: biassociative, epimerites

small and globular or cylindrical, spores dolioform to cylindrical, cysts

dehisced by spore ducts.

For differentiation of species Watson gives the following characteristics:

size, both medium and average; ratio of length of protomerite to total length;

ratio of width of protomerite to width of deutomerite; general shape of the

body; shape of the protomerite and of the deutomerite; character of the inter-

locking device between the sporonts in syzygy; size and shape of the nucleus;

colour and character of the protoplasm, and the shape of the cysts and their

method of dehiscence.

A species can only be fixed by the above characters when a large number

of individuals are considered as there is a great deal of variation between

individuals. For instance, mature individuals taken from a heavily infected or

from a starved host are not only smaller but also narrower than those taken

from lightly infected or from well-fed hosts. (See discussion on the dimensions

of Gregarina gibbsi from T. phalerata.) Also, the protoplasm of starved gregarines

is far less dense and less granular than that of well-fed ones.

A far more constant feature that does not vary with nutrition, etc., and

which can be used for the differentiation of species, is the structure of the

nucleus as it appears when stained in Heidenhain’s iron haematoxylin or in

other haematoxylin strains. As far as is known, this character has not been

used before in the case of this genus.

When stained the nucleus appears to be vesicular and contains a number

of clearly visible, darkly staining nucleoli or ‘karyosomes’. In the gregarines

FOUR NEW SPECIES OF GREGARINES FROM CAPE PENINSULA 389

described here the number of these varies from one to three in one species, to

almost twenty in another; however, in any actual species the number was

fairly constant or lay between well-defined limits when mature individuals

were considered. Moreover, it appears that these nucleoli vary in staining

properties in different species; in three of the species they stained very darkly,

almost black, but in the fourth, though clearly visible, they were hardly darker

than the rest of the nuclear substance, though the same technique was used.

These differences in the distribution of nucleoprotein appeared to be very

characteristic, and it would seem probable that other variations would be

discovered if more species were to be studied in a similar manner.

The different appearances of the nuclei of the four species are illustrated

in figure 11 and further discussed in the descriptions. The use of the nuclear

appearance was found most useful in relating starved and well-fed individuals

of the same species and also in distinguishing between individuals of the two

different species which were found in Melanosilpha capensis, especially in serial

sections.

Watson, in her general description of the Gregarinidae, states that, as the

eregarines increase in size, the nucleoli increase in number and decrease in

size until they are scattered irregularly throughout the nucleus and cannot be

counted. In these species, however, it was found that, though the nucleus of

a very small cephalont starts with only one nucleolus which divides up as it

grows, it ceases to do so at a fairly early age and each species develops its own

characteristic number.

Watson goes on to make the suggestion that this supposed breaking-up of

the nucleoli into large numbers before gamete formation, hastens this process

and reduces the time that cysts take to develop. She also states that the time

taken for cysts to develop and dehisce in the Genus Gregarina is two days. In

the case of one of the present species the cysts took twelve days to develop and

this was the species with over fifteen nucleoli, whereas in the species with one to

three nucleoli, the cysts took five days to develop. The time taken for cysts to

develop was found to be very constant for each species.

Gregarina fastidiosa n. sp.

Host: Aptera cingulata (Burm.)

Figs. 1-3, I1a.

Specimens of Aptera cingulata were collected on the lower slopes of the

mountain at St. James, Cape Peninsula; most of the specimens found were

females and only three males were captured during the whole period (March

to June 1943). Nymphs of most instars, except the very earliest, were captured

and examined.

These cockroaches live under dense bushes, in crevices in rocks or in any

sheltered place into which they can crawl. They are usually found singly but

occasionally two or three are found together; the males discovered were with

390 ANNALS OF THE SOUTH AFRICAN MUSEUM

females. Very young nymphs are sometimes found in groups of four or five,

but even these usually occur singly.

All the mature females examined were heavily infected to a greater or

lesser degree. In all cases there must have been well over a hundred parasites

in the gut; one specimen in particular must have had many hundreds in the

mesenteron and hepatic caeca. This specimen died in captivity after a few

hours and, when examined 24 hours after death, the gregarines were all alive

and in a very active condition. Another interesting case was that of a female

which was examined after it had been dead for some days at least as the gut

and organs were in an advanced state of decay and were quite unrecognizable;

FIG. I.

Gregarina fastidiosa n. sp., outlines from life, nuclei inserted from sections.

x 46.

here many dead gregarines were found but in moist places there were many

still alive.

Cysts were only taken from females and older nymphs.

Of the males examined two were heavily infected but all the parasites

were very immature and only cephalonts (i.e. individuals with epimerites)

were found.

The guts of the males seemed very empty and it would appear that they

do not feed when mature.

The third male examined had very large testes filled with actively motile

spermatozoa; the gut in this case seemed to have degenerated and was filled

with air bubbles. No parasites were discovered.

All nymphs examined were infected. The earlier instars were lightly

infected with young cephalonts while the later instars also contained mature

sporonts in syzygy and even had gametocysts in the posterior gut.

FOUR NEW SPECIES OF GREGARINES FROM CAPE PENINSULA 391

Gregarines were found in all parts of the gut except the crop and the

gizzard. The hepatic caeca and anterior mesenteron contained mostly

cephalonts; the caeca were frequently packed tight with them. Mature

sporants, mostly in syzygy, occurred in the middle and posterior mesenteron,

a few occurred in the hind gut and some passed out in the faeces.

Cysts were occasionally found in the anterior parts of the mesenteron but

there were more in the middle and posterior parts; most cysts, however, were

found in the hind gut and in the faeces.

Description

This gregarine is fairly large, quite white in colour, the body is divided

into protomerite and deutomerite by means of a well-formed septum and

there is a well-marked constriction at the septum. The sporonts are

biassociative when in syzygy. ‘The cephalont bears a small, knob-shaped

epimerite on a short stalk.

Mature Sporonts (fig. 1a—d). ‘These lie free in the mesenteron, often in syzygy.

Dimensions were taken from live specimens but this involved certain diffi-

culties, the chief one being that very few of the specimens on a slide were in a

suitable position for measuring as they were either undergoing bending move-

ments, were lumped together, or were adhering to débris. Also specimens did

not remain long in the 0-75 per cent NaCl but soon died and became dis-

tended; this difficulty was overcome by keeping them in the fluid contents of

the gut and by adding as little saline as possible. The forward movement of

the gregarines was not very troublesome as this species is very sluggish.

The following figures were obtained from thirty specimens taken from

different hosts.

Average Medium Range

Total length ch .. 594 microns 582 microns 743-443 microns

Length of protomerite .. 120 2 119 aM 143-87 ns

Width of protomerite .. 218 sig 187 M3 229-214 ad

Length of deutomerite .. 474 3 463 600-400 Pe

Width of deutomerite .. 352 bb 314. i 357-300 My

Ratios required for species determination, based on average figures:

Length of Protomerite

Total Length

Width of Protomerite

Width of Deutomerite

The greatest variation is in the length of the deutomerite (200 microns).

The most constant feature is the width of the protomerite—a variation of only

15 microns.

I: 4.9

Yo 1:6

General Shape. The deutomerite is more or less oval with the anterior end

flattened where it joins on to the protomerite. The greatest width is usually

towards the anterior end but not always. The deutomerite often has a very

slight waist in the middle and the posterior end is rounded.

392 ANNALS OF THE SOUTH AFRICAN MUSEUM

The protomerite is over twice as broad as it is long.

Cross-sections are quite circular showing that the gregarine is not flattened.

There is a rather unusual variation in the structure of the septum between

the protomerite and the deutomerite: thin strands of the septal cytoplasm run

forward from the septum into the protomerite and they tend to converge at

Fic. 2.

Gregarina fastidiosa n. sp. a. epimerite surrounded by remains of host

cell, arrow indicates point of breakage. x 4930. b. section showing

strands running forward from septum. c. stained developing spore

from smear, showing eight dumbell-shaped nuclei. x 2000.

the anterior end. (Fig. 2b.) These are only obvious in sections and usually

only one appears but occasionally two or three.

The Epicyte. ‘This is rather thick, and thickest in the position of the septum

where there is a well-marked constriction. The usual longitudinal ridges

appear as well as the myonemes just underneath it.

FOUR NEW SPECIES OF GREGARINES FROM CAPE PENINSULA 393

The Nucleus. ‘This occurs in any position in the deutomerite. It is fairly large

and the diameter averages about 60 microns. It contains from one to three

nucleoli which stain very darkly in haematoxylin. (Fig. 11a.) Most indivi-

duals have two nucleoli; when three are present, one is larger than the other

two.

The Interlocking Device. ‘This consists of a raised ridge round the top of the

protomerite of the satellite which fits on to the posterior part of the deutomerite

of the primite, rather like a sucker. The deutomerite is slightly squeezed out of

its normal shape to take it. A characteristic feature of this species is the

weakness of this interlocking device; even the gentlest handling separates the

sporonts.

The Epimerite. ‘This is knob-like with a short stalk and the entire structure,

both knob and stalk, is inside one of the cells of the gut wall. The effect on

the host cell is as follows: the cell, normally oblong, becomes round, its striated

border disappears and the contents degenerate entirely, neighbouring cells

also become shortened, the extent of this depending on their nearness to the

epimerite which thus fits into the base of a small pit in the gut wall. (Fig. 3.)

In sections the shape of the epimerite is usually obscured by darkly staining

degenerated cell contents.

When cephalonts are freed from the gut wall the epimerite is usually left

behind but those taken from a host which had been dead for 24 hours came

away easily with the epimerite intact surrounded by the gut wall cell which

had assumed a spherical shape. The epimerite could be seen inside and

appeared to have a slight ridge around the top (fig. 2a); specimens were also

found with the epimerite half broken away and the point of breakage is shown

in the same figure. The fact that the rounded cell was not the epimerite was

not obvious unless specimens were examined very carefully; this could easily

lead to erroneous descriptions.

Cysts. These are found in the faeces, hind-gut and a few in the mesenteron.

They are oval in shape and shining white in colour; a few spherical cysts were

discovered but these did not develop and, when artificially ruptured, were

found to contain no spores.

The average size of 20 cysts selected at random from various hosts was

730 X 438 microns (major and minor diameters). The measurements ranged

from 910 X 450 microns to 500 X 440 microns. The thinnest cyst was

640 X 400 microns.

The cysts dehisce by means of from 7 to 12 spore ducts which are from

120 to 140 microns long; the majority of the spore ducts are functional, the

spores push through the ducts and form chains often 2 cm. long. In a cyst

ruptured artificially after 24 hours, fully formed ducts were formed which

stained very darkly in haematoxylin.

It was noticed that when a drop of water collected around a cyst, the

cyst burst by means of a simple rupture and released the spores in a cloud

which dried into a solid mass.

394 ANNALS OF THE SOUTH AFRICAN MUSEUM

Cysts dehisced in five days after extraction from the host. They dehisced

in the sealed petrie dish and seven out of fourteen cysts exposed to the atmo-

sphere also dehisced. Cysts placed in a sealed petrie dish containing anhydrous

calcium chloride failed to dehisce.

Spores. ‘These are barrel-shaped and very uniform in size, the dimensions being

7°1 X 4:2 microns. The barrel shape is given by an exterior coat of mucus, and

when this was removed in glacial acetic acid, the spores appeared oval. The

spore wall could not be penetrated by ordinary fixatives and stains so that

sporozoites could not be seen; however, very young spores were obtained by

rupturing a cyst artificially after 48 hours; these had no resistant wall and,

Fic. 3.

Gregarina fastidiosa n. sp., epimerite in situ showing effect on host tissues. cyt. undegenerated

cytoplasm. e, epimerite. m. nucleus. vac. vacuole with degenerated cytoplasm.

FOUR NEW SPECIES OF GREGARINES FROM CAPE PENINSULA 395

when stained, were found to contain eight bilobed nuclei arranged in two rows

of four, one row at each pole. (Fig. 2c.)

Gregarina gibbsi n. sp.

Host: Temnopteryx phalerata (Sauss.)

Figs. 4-6, 115.

Specimens of T. phalerata were collected on the slopes and summit of St.

James mountain during the late autumn and winter. These cockroaches were

found under loose stones, pieces of wood and other objects and usually occurred

in small groups of three or four and sometimes more.

There appeared to be no striking differences between males and females,

as in A. cingulata, and, as the degree of infection in both sexes appeared to be

much the same they were not dealt with separately as regards statistics.

Fic. 4.

Gregarina gibbsi n. sp., outlines from life, nuclei inserted from

sections. x 46.

Although these cockroaches were found together in groups, the infection

rate was not heavy; 32 per cent of all hosts were infected and only to per cent

heavily.

The gregarines were found in the anterior mesenteron but none in the

hepatic caecae. All cysts were discovered in the hind-gut or rectum.

Description

G. gibbsi is of medium size and is white in colour; it embodies all the

general features of the genus. There is a slight constriction at the position of

the septum.

Sporonts are biassociative when in syzygy but one unique case was seen

where three had associated in line (fig. 4c).

Mature Sporonts. These lie free in the mesenteron and are nearly always in

syzygy. Dimensions were taken from live individuals and no difficulty was

396 ANNALS OF THE SOUTH AFRICAN MUSEUM

experienced in keeping specimens alive in 0:75 per cent saline. Only the

primite was measured.

When this species was first studied and the dimensions and ratios examined,

it was found that those from heavily infected hosts were much thinner than

those from lightly infected hosts. Before this relationship between proportions

and infection rate was noticed it was thought that two species, at least, were

involved as the ratios of width of protomerite/width of deutomerite varied

from 1 : I'1 to 1 : 2:6. However, a complete range, linking these two values,

was later obtained and the nuclear structure was found to be the same in all

individuals. Thus it was concluded that they were all conspecific.

The following figures were obtained from specimens from both heavily

and lightly infected hosts.

Average Medium Range

Total length fe .. 477 microns 442 microns 614-371 microns

Lenethiof protomentte 2 177 4 qi i 100-57 Be

Width of protomerite .. 105 ne 86 - 114-86 M

Length of deutomerite .. 400 ss oom A 514-314 es

Width of deutomerite ) .2 171 3 114. ss 227-114, "

Ratios required for species determination, based on average dimensions:

Length of Protomerite

Total Length

Width of Protomerite

Width of Deutomerite

1:6

T 2 16

The greatest variation is in the length of the deutomerite (200 microns)

and the least variation is in the width of the protomerite (28 microns).

General Shape. The shape of the deutomerite may vary greatly. In the larger

specimens it may be more or less oval and twice or one and one half times as

long as broad; in the smaller specimens it may be more elongated and three

times as long as broad. The posterior end may be rounded or slightly pointed

(fig. 4a—e).

The protomerite is dome-shaped and may bear a slight papilla at its anterior

end at the position where the epimerite was attached.

The Epicyte. This is moderately thick and tough enough to make handling of

live specimens easy. The usual longitudinal ridges are present and myonemes

are visible below.

The Nucleus. This occurs anywhere in the deutomerite. The nucleoli stain

darkly and are often clumped together so that they are difficult to count.

Mature sporonts have from 12 to 20, but the majority have 15. Young cepha-

lonts usually have fewer, 8 or 6 or less (fig. 11).

The Interlocking Device. ‘This is similar to that described for G. fastidiosa. How-

ever, here the syzygy is very firm and individuals do not come apart with

normal handling and can be easily mounted together.

FOUR NEW SPECIES OF GREGARINES FROM CAPE PENINSULA 397

The epimerite is knob-shaped and

is borne on a very short stalk.

When studied in whole mounts it

appears to be slightly flattened at

the top around which there

appears to be a very distinct

ridge (fig. 5). Serial sections

showed that in this case not only

one host cell was destroyed, as in

the previous species, but also

several surrounding cells were

completely broken down with the

apparent disappearance of cell

walls (fig. 6). on eae,

: : Gregarina gibbsi n. sp., epimerite (drawn from

Cysts were found in the hind-gut whole mount). x 360.

and faeces. They are oval, white

and glistening. They were not very numerous and few were discovered and only

five measured. ‘These gave average dimensions of 428 x 311 microns and

ranged from 457 X 392 to 400 X 300 microns.

A characteristic feature is that there is a very thick gelatinous layer out-

side the true cyst wall; this layer is present in the other species also but is much

thinner and hardly noticeable.

The cysts dehisced in 12 days. Dehiscence took place normally both under

normal atmospheric conditions and also in the damp chamber. Six cysts were

placed in drops of water in a damp chamber and, after twelve days, only one

had dehisced; the rest were removed and allowed to dry, these dehisced after

two days (fourteen days in all). The inhibiting effect of the water is probably

linked with the presence of the thick, gelatinous coat.

Cysts dehisce by means of from 8 to 12 operational spore ducts which are

quite normal in shape. However, when they dehisce there appear two or three

extraordinary ducts of unusual length. These are thinner than usual and

appear flattened and no spores are released through them, although the drop

of oil that invariably passes up the normal spore ducts just ahead of the spores,

also passes up these and hangs on the end. The normal ducts are approxi-

mately 250 microns long, while the abnormal ducts are 3,600 microns long.

The spores are exuded in chains, although a few cases were observed

where they collected in a large clump at the end of a duct.

Fic. 5.

The spores are barrel-shaped and very uniform in size. They were 8-5 xX 4:2

microns.

Species of Gregarina from host: Melanosilpha capensis (Stal.)

Specimens of M. capensis were collected from the top of St. James mountain

during the winter months of 1943. This small, black cockroach was found only

398 ANNALS OF THE SOUTH AFRICAN MUSEUM

on the top of the mountain in dry situations, usually under loose stones or

small rocks perched on top of boulders. They were found usually in small

groups of about half a dozen adults, mostly females, with large numbers of

small nymphs. The females are apterous but the males, which are few in

number, are winged. (Only 7 out of 100 individuals examined were males.)

Two distinct species were found in this host which have been named

Gregarina sandont and Gregarina impetuosa.

Fic. 6.

Gregarina gibbsi n. sp., section of epimerite in situ showing effect on host

tissues. d. c. degenerated cells. e. epimerite. n. nucleus. x 360.

Infection. These statistics were obtained from 100 individual hosts:

Gregarina sandoni Gregarina impetuosa

Adult of both sexes infected .. 38 per cent 6 per cent

Males infected Bi at 29 per cent 14. per cent

Females infected a - 39 per cent 5 per cent

As will be seen G. sandoni was found more often than G. zmpetuosa. Only

one host was at all heavily infected with G. impetuosa and that was a male. Only

13 per cent of the hosts infected with G. sandoni were heavily infected.

FOUR NEW SPECIES OF GREGARINES FROM CAPE PENINSULA 399

Gregarina sandoni sp. nov.

OS 7 Te

This was found in the anterior and middle parts of the mesenteron and

very young cephalonts in the hepatic caeca.

It is a fairly large gregarine, quite white in colour, with a clearly marked

septum; there is no marked constriction of the epicyte at the septum but

merely a slight dent. The species is extremely sluggish and hardly moves at all.

The sporonts are biassociative, and very characteristic of this species is

the very early age at which they form syzygies: young sporonts, only half or

even less than half the size of mature sporonts, are commonly to be found in

syzygy. These are liable to be mistaken for specimens of G. impetuosa but can

easily be distinguished by their sluggishness, the structure of their nuclei and

the presence of extra-nuclear bodies (which will be discussed later).

Mature Sporonts (fig. 7a-c) were found in the anterior or middle parts of the

mesenteron. Dimensions were taken from live specimens and no difficulty was

experienced in keeping them alive in 0-75 per cent saline. Only primites were

measured.

The following were the dimensions:

Average Medium Range

Total length ah .. 498 microns 451 microns 642-357 microns

Length of protomerite .. 80 a 86 . 100-57 RS

Width of protomerite .. 134 5 114 > 157-114, 3

Length of deutomerite .. 418 is 443 54.2-300 -

Width of deutomerite .. 274 # 257 i 357-214. ed

Ratios for species determination based on average figures:

Length of Protomerite

Total Length

Width of Protomerite

Width of Deutomerite

i 8 (Oye

lf @ 2

The greatest variation is in the length of the deutomerite (242 microns)

and the most constant feature is the width of the protomerite (43 microns

variation).

General Shape. The deutomerite is more or less oval in shape but it is slightly

pointed at the posterior end. The greatest width is usually towards the posterior

end in adults but in younger individuals the anterior is usually the widest part.

The protomerite is usually two-thirds as long as it is wide.

The Epicyte. ‘This is very thin and mature sporonts are very easily ruptured

even when handled very gently. This feature is not so noticeable in immature

specimens. The usual longitudinal ridges and circular myonemes are present.

The nucleus occurs anywhere in the deutomerite. It is fairly large and averages

about 50 microns in diameter. It contains 4 or 5 nucleoli which are rather

characteristic as they stain only slightly darker than the rest of the nucleus in

400 ANNALS OF THE SOUTH AFRICAN MUSEUM

Heidenhain’s iron haematoxylin and are often difficult to see. Nevertheless,

when viewed in section under an oil immersion lens, they appear as very

definite structures with definite outlines. They are often clumped together at

one side of the nucleus. (Fig. 11.)

Apart from the nucleus there is, in the deutomerite, a most characteristic

extra-nuclear body. ‘This is usually about a quarter the size of the nucleus and is

spherical with a very smooth surface. Occasionally it is broken into two, three

Fic. 7.

Gregarina sandoni n. sp., outlines from life. a-c. from normal host. e. from starved host. d. the

only individual in a well-fed host. x 76.

or more smaller bodies and may be elongated instead of spherical. The body

stains fairly darkly in haematoxylin but not at all by the Feulgen method. No

similar structure was found in the other species nor was any reference found to

it, or any similar structure, in the literature. (Typical examples have been

inserted in the outline drawings, fig. 7a—-e.)

The interlocking device is similar to that described for G. fastidiosa. In this

species the syzygy is fairly firm and the primite and satellite are not easily

separated with normal handling.

The epimerite is of the normal type found in this genus. The whole structure is

inside a cell of the gut-wall against which the protomerite is applied. The con-

FOUR NEW SPECIES OF GREGARINES FROM CAPE PENINSULA 401

tents of the cell, including the nucleus, break down and the effect on the

surrounding cells is the same as that described for G. fastediosa.

Few specimens were obtained in whole mounts with the epimerite still

intact; in fact, this structure seems to be exceptionally easily shed and this may

explain why so many immature sporonts are found free in the gut with the

early formation of syzygy.

Cysts are oval and are white or light buff in colour. They are found in the

faeces and hind-gut. The average dimensions for the shortest and longest

diameters are 330 and 500 microns respectively. The cysts range from 282 x

420 to 420 X 670 microns. The largest cyst recorded here was very much

bigger than the average size but most of the cysts did not vary very much from

this average.

Cysts dehisced in three days after they were removed from the gut. They

did not dehisce when exposed to the atmosphere but only in the damp chamber.

Dehiscence was by 7 or 8 spore ducts most of which seemed to be operational.

However, when a cyst was artificially ruptured just prior to dehiscence it was

found to contain 20 ducts. It would seem that all ducts are not extruded

simultaneously and, after the first 7 or 8 ducts are extruded the pressure inside

is too low to force the others out.

Spores are dolioform and are extruded in long chains. They are very regular in

size and are 7-1 microns long and 3°55 microns wide (measured under an oil

immersion lens), exactly twice as long as broad. No success was obtained in

attempts to make them exsporulate by putting them in fluid from the

mesenteron and hepatic caeca.

An interesting example of the dependence of the gregarines upon the

nutrition of their host was discovered when a specimen of M. capensis was

starved for a month and then opened. The mature sporonts were well below

average in size and much thinner, especially the deutomerites. Their average

dimensions were:

Total length ve se .. 464 microns

Length of protomerite eA: Ss

Width of protomerite .. aa TOA an

Length of deutomerite .. a 400 z

Width of deutomerite .. 55 ES 5

One of these starved individuals is illustrated in figure 7e. The endo-

plasm of these starved gregarines was less dense than normal. Cysts of normal

size and shape were discovered but their contents appeared to be coagulated

into dense lumps leaving the rest of the cyst transparent. These cysts did not

dehisce and no spores were formed in them.

Figure 7d was the only parasite in an apparently well-fed host and provides

an interesting contrast to 7e.

402 ANNALS OF THE SOUTH AFRICAN MUSEUM

Gregarina impetuosa n. sp.

Figs. 8-10, 11d.

All specimens of this species were found in the anterior mesenteron of the

host.

It is a small gregarine with a well-marked septum, at the position of which

there is a slight constriction of the epicyte.

The mature sporonts are _ bi-

associative and the cephalonts bear

a knob-shaped epimerite on a short

stalk.

Most characteristic of this species is

its activity; the sporonts glide forwards

across a slide covered in gut debris at

about 500 microns per minute, pushing

the débris aside as they move. This

activity made measuring and drawing

difficult but it was possible on rare

occasions when the gregarines were

slowed down by some exceptionally dense

obstruction.

Mature Sporonts (fig. 8). These lie free in

the mesenteron and are usually associated

in pairs. They are unaffected by the

0°75 per cent saline and remain alive in

Fic. 8. it for long periods.

Gregarina impetuosa n. sp., outlines from life, The following dimensions were

nuclei inserted from sections. x 100. taken from the primite only:

Average Medium _ Range

Total length ai .. 2QI microns 300 microns 357-228 microns

Length of protomerite .. 57 ss 57 53 72-A2 x

Width of protomerite .. 66 ie 71 ys 86-43 re,

Length of deutomerite .. 234 a 243 fs 286-185 es

Width of deutomerite .. 126 Hs 114, Bi 171-71 KS

Ratios based on averages:

Length of Protomerite

Total Length

Width of Protomerite

Width of Deutomerite

Eo? 1.9

The feature that shows the least variation is the length of the protomerite

(28:7 microns) while the greatest variation is in the length of the deutomerite

(101 microns).

FOUR NEW SPECIES OF GREGARINES FROM CAPE PENINSULA 403

Fic. 9.

Gregarina impetuosa n. sp., showing early development of karyosomes and

effect on parasitized cells. This shows the youngest parasite discovered.

d. deutomerite. e. epimerite. k. karyosomes or nucleoli. n. nucleus.

p. protomerite. x 800.

Fic. 10.

Gregarina impetuosa n. sp., a later stage. Lettering as in fig.9. X 570.

4.04. ANNALS OF THE SOUTH AFRICAN MUSEUM

General Shape. ‘The deutomerite is approximately oval in shape and the greatest

width is usually towards the posterior end, though not always. The deutomerite

has a slight waist in the middle.

The protomerite is only slightly wider than it is long. There is usually a

papilla on the anterior end where the epimerite was attached.

The epicyte is fairly thick and the gregarines are seldom, if ever, ruptured with

normal handling. The usual longitudinal ridges and circular myonemes are

present.

The nucleus (fig. 11d) occurs in any position in the deutomerite and is, on the

average, 34 microns in diameter. In the mature sporont there are from 6 to 8

nucleoli which stain darkly with haematoxylin though a few cases were

Fic. 11.

Nuclei stained in Heidenhain’s iron haematoxylin showing ‘karyosomes’ or nucleoli.

a. G. fastidiosa. b. G. gibbsi. c. G.sandoni. d. G. impetuosa.

FOUR NEW SPECIES OF GREGARINES FROM CAPE PENINSULA 405

observed where some of the nucleoli in a single nucleus stained a much lighter

colour than the others. In the youngest cephalont (fig. 9) there was only one

nucleolus; this specimen was the smallest cephalont encountered in any of

these four species. The nucleolus must divide up at a very early stage as in a

somewhat larger but still very young cephalont it had already divided up

into five (fig. 10).

The epimerite was only seen in sections and all cephalonts examined were very

young. It was knob-shaped and borne on a very short stalk and the whole

structure was inside a cell of the gut-wall.

The interlocking device is similar to that described for Gregarina fastidiosa. In this

species the syzygy is very firm and the pairs are not easily separated by normal

handling.

Cysts. Although well over 100 specimens of M. capensis were examined no cysts

of this species, and, consequently, no spores were discovered.

Although the mode of dehiscence of the cysts and the shape of the spores

of this species are not yet known, the known features, such as the knob-shaped

epimerite and the biassociation of the sporonts well before cyst formation, have

led to this species being placed in the genus Gregarina Dufour.

Diagnostic differences between Gregarina sandont and Gregarina impetuosa

Gregarina sandom Gregarina impetuosa

(a) Large gregarine. Small gregarine (see average

dimensions).

(6) Very sluggish. Very active.

(d) Nucleoli stain only slightly darker Nucleoli stain much darker than

than rest of nucleus. the rest of the nucleus.

(e) Extra-nuclear body present. No extra-nuclear body.

This paper was originally written in 1943 and presented in part fulfilment

of the degree of M.Sc. at the University of Gape Town. Thanks are due to

Dr. H. Sandon, then Senior Lecturer in Zoology, for help and guidance, and

to Dr. A. J. Hesse of the South African Museum for his help in identifying the

cockroaches. A set of slides has been deposited in the South African Museum.

REFERENCES

Bhatia, B. L., 1938. Fauna of British India, Protozoa (Sporozoa).

Bhatia, B. L., and Setna, S., 1924. ‘On some new cephaline gregarines.’ Parasitology, XVI,

pp- 279-88.

Bush, S. F., 1928. ‘A study of the gregarines of the grasshoppers of Pietermaritzburg, Natal.’

Ann. Nat. Mus., V1, pp. 97-169.

Gibbs, A. J., 1946. ‘Stylocephalus ingeri, sp. nov., a cephaline gregarine found in the gut of

Gonocephalum arenarium (Coleoptera).’ Trans. Roy. Soc. S. Afr., XXXI (2), pp. 169-80.

Sprauge, V., 1941. “Studies on Gregarina blattarum with particular reference to the chromosome

cycle.’ Illinois Biol. Monogr., XVIII, No. 2.

Watson, M. E., 1916. ‘Studies in gregarines.’ Illinois Biol. Monogr., II, No. 3.

Watson-Kamm, M., 1922. ‘Studies in gregarines.’ Jllinois Biol. Monogr., VII, No. 1.

17. Finf Dactylispa-Typen Péringuey’s.

165. Beitrag zur Kenntnis der Hispinae (Coleopt. Chrysomelidae).

Mit einer Abbildung.

Von ErtcH UnMann, Stollberg-Sachsen.

Wenn man die siidafrikanischen Dactylispa-Arten nach den Beschrei-

bungen Peringuey’s bestimmen will, so steht man vor einer schwer lésbaren

Aufgabe. Seine Beschreibungen reichen bei weitem nicht mehr aus, weil,

wegen der Fille der Hispinae-Arten des siidafrikanischen Raumes, eine Betrach-

tung der Objekte nach Gesichtspunkten erfolgen muss, von denen Péringuey

nichts wissen konnte. Es war damals, 1898 und 1908, nicht vorauszusehen,

wie gross die Artenzahl sein kénnte. Auch heute wissen wir noch nicht, wie

viele Arten eine genaue Durchforschung Siidafrikas liefern wird.

Herrn Dr. H. Andreae vom Siidafrikanischen Museum in Kapstadt ist es

zu. danken, dass wir heute tiber die Dactylispa-Arten Péringuey’s gut unter-

richtet sind. Sein ausserordentlich grosses Entgegenkommen erméglichte mir

das Studium von Typen, und wo das nicht anging, hat Herr Kollege Andreae

durch genauestes eigenes Studium zur Klarung der gestellten Fragen beige-

tragen. Seine Studien haben es erméglicht, dass die Arten Péringuey’s in

meinem Schliissel der Dactylispa-Arten Afrikas, der in einer belgischen Zeit-

schrift erscheinen soll, mit ziemlicher Sicherheit untergebracht werden konnten.

Ihm und dem Siidafrikanischen Museum in Kapstadt sage ich hiermit

nochmals meinen herzlichsten Dank.

Dactylispa capicola (Péring.)

1898. Ann. S. Afr. Mus., i, p. 121. (Hispa c.)

Die Untersuchung des Materials ergab folgendes: ‘Bei capicola sind ausser

dem eigentlichen Typ (mit rotem Zettel) zwei solche Stiicke vorhanden.’

Eins davon habe ich gesehen aus Natal: Durban. ‘Der Typus ist fein weisslich

behaart. Auf den Decken stehen die Haare zwischen den Punkten; oft ver-

schmutzt und schwer sichtbar.’ Péringuey: ‘nearly glabrous’. Die Decken

sind aufrechtstehend behaart. ‘Der Halsschild ist mit Ausnahme der Flachen

anliegend behaart, wenig dicht, die Haare am Seiten- und Hinterrand nach

der Seite, am Vorderrande nach vorn gelagert, nur zwischen den Vorderrand-

dornen halb aufgerichtet. Die Stirn ist niedergedriickt und langsrunzlig. Die

407

408 ANNALS OF THE SOUTH AFRICAN MUSEUM

Halsschildflachen sind gross, sie nehmen etwa ein Viertel bis ein Fiinftel der

Oberflache ein.’ Seitenbewehrung deutlich 2,1. ‘Fithlerglied 1 (von der Seite

gesehen) vorn leicht abgeschragt, die untere Kante etwas vorgezogen, die

Aussenecke (von oben gesehen) leicht vorspringend. Der Unterschied zwischen

den Seitenranddornen und den Spitzenranddornen der Decken ist deutlich,

aber viel schwacher als bei gracilis.”. Die Seitenbewehrung ist halb so lang wie

der Halsschild breit. ‘Der Abstand der Punktreihen voneinander ist grésser

als der Abstand der Punkte in den Reihen, deshalb treten die Zwischenraume

in der Langsrichtung mehr hervor. Rippen kann man das eigentlich nicht

nennen.’ Péringuey: ‘elytra costulate’.

Bei dem Stiick von Durban ist die Oberflache der Decken so stark skulp-

tiert, dass zwischen den einzelnen Dornen in der Langsrichtung tatsachlich

erhabene Rippenteile erscheinen.

Dactylispa congrua (Péring.)

IGOG:> loo. cits 123: (Espa)

Typus: Natal, Durban.

Der Typus ist gut erhalten, es fehlt ihm aber die rechte Decke. Farbung:

Kopf, Halsschild, Schildchen und Unterseite schwarz, Fihler gelbbraun, 1.

und 2. Fihlerglied schwarz, 3. schw4rzlich-braun, Beine gelbbraun, Decken

dunkelbraun mit schwachem Bronzenschimmer, mit schwarzen Dornen.

Behaarung: Innenseite der Augen mit hellem Haarsaum, Wangen deutlich

behaart, Halsschild auf der Scheibe und auf dem Abfall neben den Vorder-

dornen goldgelb behaart, die beiden kleinen glatten Flachen kahl; Decken

deutlich behaart, jeder Punkt mit einem goldgelben Harchen vom Durch-

messer eines Punktes. Stirn zwischen den Fihlern mit kurzem Kiel, in der

Mitte etwas eingedriickt, hinten mit kurzer, schwacher Furche, etwas schmaler

als eine Augenbreite, vom glatten Hals fein abgesetzt, nur wenig hoher als

dieser. Fihler bis hinter die Schultern reichend, schlank, etwas schmaler als

die halbe Stirn, zur Spitze nur schwach verdickt. Glied 1 etwas kraftiger als

die anderen, 2 schmiéler, ellipsoidisch, 3 so lang wie 1, schmaler als 2, zylin-

drisch, 4-5 zylindrisch, jedes kiirzer als 3, 6 kurz, so lang wie 2, 7-11 eine

schwache Keule bildend, 7 konisch, etwas so lang wie 5, 8-10 schwach quer,

jedes so lang wie 6, 11 eif6rmig, um die Spitze langer als 10. Alle Glieder fein

behaart, die Keule pubeszent. Halsschild auf der Scheibe dicht punktiert, mit

sehr feinem Mittelkiel, mit je einer kleinen, runden, kahlen Flache beiderseits,

dahinter mit seichtem Quereindruck. Neben den Vorderranddornen mit recht

kleinem Borstenkegel, Hinterecken mit vorgezogenem Borstenkegel. Beweh-

rung schlank, Vorderranddornen mit fast gleichen Aesten, diese kirzer als

eine Halsschildbreite, Seitendornen 2,1, das vordere Paar so lang wie die

Vorderdornen, mit kurzem, breiten Stiel, der freie Dorn entfernt, etwa halb

so lang wie einer der Seitendornen. Antebasalrand fein. Schildchen dreieckig,

schagriniert. Decken glanzend, hinter den Schultern wenig eingezogen, ohne

BEITRAG ZUR KENNTNIS DER HISPINAE 409

betonte Zwischenraume II, IV, VI, VIII. Punktreihen gut zu verfolgen, 7.

hinter dem Schulterdorn 3 beginnend, g. und 10. in der Mitte vereinigt.

Eindruck schwach. Dornen ziemlich lang, schlank, ohne verdickte Basis.

Naht hinter den schlanken Schliessdérnchen fein bedornt. Raum I mit einer

Reihe von Zusatzdornen; Raum II mit II 1-5, I1 5 diinn; IV mit IV 1,3,5 und

einigen diinnen Zusatzdornen; VI auf der Schulter mit den Achseldornen

1,2,3, dann mit VI 2,3,4; VIII mit VIII 4 und 5. Letzterer mit verstarkter

Basis, vorn und hinten mit einem Zusatzdorn. Randdornen der linken Decke

17, die langsten kaum langer als die langen Scheibendornen, nach dem Hinter-

winkel zu und zur Naht allmahlich kiirzer. Jede Klaue mit einem Basalzahn-

chen. 4. mm.

Zur Beschreibung Péringuey’s: ‘thorax pubescent, narrowly grooved

longitudinally in the centre. ...’ Die vermeintliche Langsgrube ist der Mittel-

kiel. ‘elytra . . . glabrous’ stimmt nicht. ‘(elytra) having on each side three

dorsal rows of moderately long spines’ ist recht summarisch, vielleicht meint

er mit der 1. Reihe die Dornen des II. Raumes, mit der 2. Reihe die des IV.

Raumes und mit der 3. Reihe die Dornen des VI. Raumes mit VIII 5. ‘a few

short spines along the suture’ ist richtig beobachtet.

Dactylispa gratula (Péring.)

1898. loc. cit., p. 123. (Hispa g.)

Herr Kollege Dr. Andreae, der den Typus untersucht hat, stellt das

Folgende fest (von mir etwas geadndert). Die Art ist durch die Bildung der

Stirn besonders ausgezeichnet. Die Stirn hat zwei weitlaufig punktierte und

behaarte Hocker, die in ihrer ganzen Lange durch eine glatte, dreieckige,

vorn zu beiden Seiten des Stirnkieles flach auslaufende Grube getrennt sind.

Die deutlich, aber nicht hoch abgesetzten Hinterrander der Hocker schliessen

ein stumpfwinkliges Dreieck ein, das ebenso glatt und glanzend ist wie der

Hals und mit ihm in einer Ebene liegt, aber durch eine feine Linie abgetrennt

ist. Fiihler ebenso lang und schlank wie bei D. pubicollis Chap. Alle Glieder

langer als breit, das 1. noch langer als bei D. pubicollis, etwas vor der Mitte am

breitesten, zur Spitze schwach verjiingt, am Vorderrande weniger scharf

abgestutzt, etwas langer als das 3.

Glied 3-6 an Lange allmahlich abnehmend, 7 deutlich langer als 6,

7-10 an Lange langsam abnehmend, 11 um zwei Drittel langer als 10, aber

etwas schmaler, etwa zweimal so lang wie breit. Hlalsschildscheibe matt,

dunkel, Bewehrung und Flachen dunkel-gelbbraun, kurz pubeszent, nach

meiner Lesart (Uh.) in der Mitte mit sehr feinem Langskiel (‘very faintly

grooved longitudinally in the middle’ bei Péringuey). Seitenflachen des

Halsschildes klein und schmal, deutlich gewolbt, glatt und glanzend. Sie heben

sich von der sonst punktierten, matten, dunklen Oberflache scharf ab. Sie

umschliessen eine fast halbkreisf6rmige Flache. Schildchen mit ovaler, tiefer,

an der Basis scharf begrenzter, zur Spitze hin flach auslaufender Grube, die den

410 ANNALS OF THE SOUTH AFRICAN MUSEUM

gréssten Teil des Schildchens einnimmt. In den Punkten der Decken stehen

feine, farblose Haare, etwa so lang wie der Durchmesser eines Punktes. Bei

schlechter Beleuchtung kaum erkennbar. Naht im basalen Drittel mit kurzen

Dornen, in der Spitzenhalfte mit feinen, nach hinten geneigten Dérnchen von

etwa 0,05 mm. Lange. Raum I mit kurzen Dornen im mittleren und Spitzen-

drittel; Raum II mit starken Dornen, die zwei ersten basalen und der letzte

apikale kirzer; Raum IV kurz hinter der Mitte und am Absturz mit einem

Dorn, der ebenso stark ist wie die Dornen des II. Raumes; and der Basis, vor

und zwischen den starken Dornen mit je einem kurzen Dorn; Raum VI mit

starken Dornen, die in der Basalhalfte langer als die des IJ. Raumes sind;

Raum VIII links mit einem kleinen Dorn kurz hinter der Mitte, einem starken

am Absturz (VIII 5), rechts mit je einem starken hinter der Mitte und am

Absturz, je einem kleinen vor dem ersten starken, unmittelbar vor und hinter

dem zweiten (VIII 5), und vor der Spitze. Rand links mit 13 Dornen, so

lang wie die von VI, aber dinner, rechts mit 17. In der Basalhalfte sind die

Dornen langer, die Spitzenranddornen nach der Naht zu stark verkirzt.

Hinter der Schulter messen sie 0,4 mm., am Nahtwinkel 0,1-0,15 mm.

4 X 2mm. Natal (Malvern), G. A. K. Marshall.

Uber die Scheibendornen schreibt Péringuey: ‘having two dorsal series

of moderately long dark spines, set at some distance from one another [nach

meiner Deutung sind das die Dornen von II und VI], with two shorter spines

on the posterior part,. between the first and second rows [bezieht sich auf die

Bedornung von IV]; the suture is also spinulose [wie oben].’

Dactylispa inanis (Péring.)

1898. loc. cit., p. 124. (Hispa 1.)

Typus: Natal, Malvern, VI, 1897 (7893).

Dem Typus fehlt die rechte Decke, sonst ist er gut erhalten. Gelbbraun,

schwarz: Augen, auf dem Hals ein Fleck hinter jedem Auge, ein Saum um jede

Halsschildflache, ein kleines Fleckchen in jeder Ecke der Schildchenbasis, das

scheinbar auch ein wenig auf den Hinterrand des Halsschildes tibergreift

(vielleicht dort nur durchscheinend), die Dornen der Deckenscheibe mit

Ausnahme von Achseldorn 1, ein Streifen auf der Schulterflache innen, ein

Fleck auf dem II. Zwischenraum an der Basis, am Hinterwinkel einige Rand-

dornen (angedunkelt) und die Seiten der Hinterbrust. Behaarung: diinn,

wenig auffallend, hellgelb, ein Saum am Innenrande der Augen, die Mitte der

Scheibe des Halsschildes, aber undicht, jeder Punkt der Decken mit einem

diinnen Harchen. Fiihler ziemlich dicht behaart. Stirn in der Mitte ein-

gedriickt, mit kurzen Kiel zwischen den Augen, vom glanzenden Hals

abgeschniirt, allmahlich zu ihm hinabgewolbt. Filer bis hinter die Schultern

reichend, kraftig, zur Spitze nur ganz allmahlich verdickt. Glied 1 gross,

stark, ellipsoidisch, Glied 2 fast kuglig, etwa ein Drittel so lang wie I, Glied 3

so lang wie 1, aber schmiler, zylindrisch, wenig langer als 4, Glied 4-6 einander

BEITRAG ZUR KENNTNIS DER HISPINAE 411

ahnlich, von abnehmender Lange, 7 schwach konisch, ein wenig verdickt, so

lang wie 4, 8-10 unter sich fast gleich, etwas langer als breit, 11 wenig langer

als 10. Halsschild in der Mitte der Scheibe gewolbt, punktiert, beiderseits

stark niedergedriickt, mit feinem Mittelkiel, beiderseits mit gewélbter, glatter

Querflache, der niedergedriickte Teil unpunktiert, unbehaart, Ante-

basalrand fein, davor flach quergefurcht. Vordere Borstenkegel klein, auf

dem Vorderrande neben den Vorderdornen, hintere Borstenkegel an der

Spitze der rechtwinkeligen Hinterecken. Bewehrung stark, kraftig. Vorderast

der Vorderdornen kiirzer als der Hinterast. Seitenbewehrung 2,1, der Seiten-

rand dort konvex erweitert, der mittlere Dorn etwas langer als der vordere,

der freie Dorn etwa halb so lang wie der mittlere. Schildchen dreieckig, mit

abgerundeter Spitze. Decken parallel, Raum II, IV, VI teilweise schwach

rippenformig. Punktreihen regelmdssig, die intrahumeralen an der Spitze

nach aussen gebogen. Dornen krdaftig, mit verdickter Basis. Naht mit sehr

kleinen Zahnchen, die nach dem Hinterrande zu etwas grésser werden,

Schliessd6rnchen etwas grdsser. Raum I vor der Mitte mit einem einzelnen

Dérnchen; Raum II mit IT 1-5 (II 2 grésster Scheibendorn, II 5 klein); IV mit

IV 1,3,5; VI mit 3 ziemlich gleichen Achseldornen, mit VI 2,3,4; VIII mit

VIII 2,3 (beide klein), 4 und 5 (Doppeldorn). 3 Basaldérnchen und ein

Schildchen-Zusatzdérnchen. 17 Randdornen, die vom Hinterwinkel bis zur

Naht an Lange abnehmen, die Spitzenranddornen recht kurz. Klauen-

zahnchen undeutlich.

Zur Beschreibung Péringuey’s: ‘faintly grooved longitudinally in the

middle’, muss ‘carinate’ heissen. Nach der Beschreibung der Dornenverteilung

auf den Decken ist die Art nicht zu erkennen. Péringuey meint, es seien 2

Reihen Dornen da, das waren die auf Raum II und IV, ‘with two isolated ones

in the second interval’ bezieht sich auf IV 3 und IV 5; ‘two set close to one

another, etc.’ bezieht sich auf den Doppeldorn VIII 5.

Dactylispa pretiosula Péring.

1908. Ann. S. Afr. Mus., 5, p. 337-

2 Typen. Malvern, Oct. 22, 1907. Beide Typen sind sehr gut erhalten.

Sie sind auf ein Stiick Karton geklebt. Ich wahle das grdéssere, rechte Stiick

als Lectotypus (Abb. 1). Farbung: Glanzend, gelbbraun, schwarz: Augen,

Umegebung der drei glatten Halsschildflachen, Randdornen des Aussen-

winkels und die Dornen der Deckenscheibe mit jeder Basis, auf der Scheibe

selbst die Schulter auf der Innenflache, die Schulterkehle, Aussenwinkel um

VIII 5 herum und einige Langsstreifen auf den Zwischenstreifen, z.B. ein

Fleckchen hinterm Schildchen auf der Schildchenreihe, die Naht (ange-

dunkelt), der III. Zwischenraum im 2. Zwischenstreifen, auf dem 3. Zwischen-

streifen der V. Raum im Anschluss an die Schulterschwarzung, eine Querbinde

zwischen den Dornen II 5 bis VIII 5 (das Schwarz zuweilen stellenweise mit

schwachem Metallschein). Angedunkelt ist der glatte Hals und die beiden

412 ANNALS OF THE SOUTH AFRICAN MUSEUM

Ire

a) rar

= PAA i

wie SNES ers

Oey a = ee,

oer is) x oR D <) PORE Siete

Ay reat Onin 3 ;

< S = a ee . ~ "Mee. Deo

as EMS sera Bare Ries

= OO igs Ae ean Soe eae

as = 5 CN ge, =

—¢ Aes ox ; ee

ee “ioe 7D Ay

o a) y

foe ~ 3 Sees Oy ey

c 4S ?

(se)

ESS

WEBB be Ta

(PI J " ee

A a in BS

lis,

se,

Dactylispa pretiosula Péring. Lectotypus.

B = Basaldérnchen. S = Schliessd6rnchen an der Naht. Z = Schildchen-Zusatzd6rnchen,

1,2,3, = Achseldornen. 1,2,3, VI 2 = Schulterdornen. II, IV, VI, VIII = gerade (Zwischen)

rdume. Von den Dornen sind auf der rechten Decke in ihrer wahren Grésse gezeichnet: II

224-5 Vite Sens OW bee OWN leo ora a Vie Aas

BEITRAG ZUR KENNTNIS DER HISPINAE 413

Basalglieder der Fiihler und noch einige Stellen auf den Decken. Die helle

und die dunkle Farbung sind nicht scharf abgegrenzt.

Unterscheide: Zwischenraume oder Raume liegen zwischen den einzelnen

Punktreihen; Zwischenstreifen oder Streifen liegen zwischen den Rippen,

zwischen Naht und I. Rippe (II. Raum), oder zwischen I. und II. Rippe (das

ist zwischen II. und IV. Raum) und so weiter.

Behaarung. Innenrand der Augen mit hellem Haarsaum, Scheibe des

Halsschildes mit einzelnen Harchen, jeder Punkt der Decken mit einem langen,

gekriimmten, hellen Harchen. Auf dem Halsschild sind die Harchen sicher

zum Teil abgerieben. Fihler fein behaart. Stirn mit den gewdlbten Augen

in einer Flucht verrundet, erstere zwischen den Fihlern mit kurzem Kielchen.

Kopf vom glatten Hals abgeschniirt, Stirn zum Hals hinabgewoélbt. Hals mit

mattem Fleck in der Mitte (Stridulationsapparat?). Fiihler die Schultern

erreichend, diinn, Glied 7-11 nur unmerklich verdickt. Glied 1 von der

iiblichen Starke, 2 ellipsoidisch, langer als breit, 3-5 zylindrisch, unter sich

fast gleich, 3 so lang wie 1, 6 wie die vorhergehenden, etwas kiirzer als 5, 7 so

lang wie 5, 8-10 doppelt so lang wie breit, 11 um die Spitze langer als to.

Halsschild auf der Scheibe niedergedriickt und dicht gerunzelt-punktiert, mit

glattem Mittelstreifen (Kielchen) und einer schragen, ovalen, gewdélbten

Flache beiderseits. Antebasalrand fein. Vordere Borstenkegel klein, unweit

der Vorderdornen, hintere Borstenkegel an den spitz vorspringenden Hinter-

ecken. Bewehrung kurz und krAaftig, der freie Dorn etwas kiirzer als die anderen,

unter sich fast gleich langen Dornen. Schildchen breit, dreieckig, mit

schwachem Eindruck. Decken hinter den Schultern nicht eingezogen, zur

Spitze etwas verbreitert, mit regelmassigen Reihen und Raumen. 9g. und Io.

Reihe in der Mitte kurz vereinigt. Schildchenreihe und gerade Raume

rippenartig, der VIII aber nur schwach konvex. Naht mit einem Paar senk-

rechter Schliessdérnchen und einer Reihe winziger Zahnchen; Raum I mit

einem einzelnen Doérnchen hinter der Mitte; II mit II 1,2,4,5; IV mit IV

1,3,5; VI auf der Schulter mit den Achseldornen 1,2,3, dann VI 2,3,4,; VIII

mit VIII 4 und dem Doppeldorn VIII 5 auf erhabener Basis. Randdornen

beiderseits 14, die des Seitenrandes so lang wie die Dornen der Scheibe, am

Hinterwinkel und bis zum Nahtwinkel von abnehmender Lange, die nahe am

Nahtwinkel recht kurz. Eindruck deutlich. Klauenglied mit einem Zahnchen

an jeder Klaue. Lectotypus 3,5 x 2 mm. (vor der Spitze), Paratypoid etwas

kirzer und schmdler, 34 x 14 mm.

Es ist nicht ausgeschlossen, dass D. pretiosula Péring. und D. hirsuta Gest.

identisch sind. Die ausfiihrliche Beschreibung Gestro’s passt ganz auf unsere

Art, vielleicht haben unsere beiden Typen auf den Decken etwas mehr

schwarze Streifen. Eine Synonymierung muss aber bis zum Vergleich der

Typen unterbleiden. Bei D. hirsuta gibt es gewisse Abweichungen, auf die ich

schon in friiheren Arbeiten hingewiesen habe. Man weiss nicht, ob sie

individuell oder artbedingend sind. Vielleicht handelt es sich hier um einen

Formenkreis einander sehr ahnlicher Arten.

414 ANNALS OF THE SOUTH AFRICAN MUSEUM

Zur Beschreibung Péringuey’s: (1) ‘Pallide straminea’, die Typen sind

nachgedunkelt. (2) ‘fere glabra’, nur auf den Vorderk6érper zu beziehen. (3)

‘Caput punctulatum’, die Stirn ist nicht punktuliert, aber schagriniert. (4)

‘striga’, bezieht sich auf den kurzen Mittelkiel des Halsschildes. (5) ‘inter-

stitiis vix distincte albido setulosis’, bei x 22 ist die Behaarung deutlich.

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