Lapse

OF THE

COUNCIL

AND OFFICERS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

1920.

Patron.

His Masgesty Tue Kina.

COUNCIL.

His Grace Tor Duke or Beprorp, K.G., F.R.S., President.

'Tae Hon. Crecit Barine, M.A.

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

Lr.-Cot. 8. Monckton Cops-

MAN, M.D., F.B.S.

CHARLES DrumMonp,

Treasurer.

Huen §. GuApstone, Esq.,

M.A., F.R.S.E.

Siz Srpnry F. Harmer, K.B.E.,

M.A., Se.D., F.R.S., Vice-

President.

Pror. James P. Hit, D.Sc.,

FE.R.S., Vice-President.

WiiiiAmM Huntsman, Esa.

Pror. Ernest W. McBrips,

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

Ksq.,

Cont. Str Henry McManon, |

G.C.M.G., K.C.LE.

E. G. B. Mreapre-Watpo, Esq.,

Vice-President.

P,. CuauMers MircHe.u, Esgq.,

C.B.H., M.A., D.Se., LL.D.,

F.R.S., Secretary.

THE Karu or Onstow, O.B.E.

Magsor Arperr Pam, O.B.E.

Aprian D. W. Pottock,

Ksq.

His Grac—E Tur DUKE oF

Ruttanp, K.G.

THE Marquis oF Srico, F.S.A.,

Vice-President.

Masor RicHarp 8. Taytor.

A. Trevor-Barrys, Esq., M.A.

AntHony H. WINGFIELD, Esq.,

Vice-President.

PRINCIPAL OFFICERS.

P. Caatmers Mircuet., C.B.M., M.A., D.8c., LL.D., F.R.S.,

Secretary.

R. I. Pocock, F.R.S., F.L.8., Curator of Mammals and

Resident Superintendent of the Gardens.

D. Seru-Smiru, Curator of Birds and Inspector of Works.

Epwarb G. BouLencEr, Curator of Reptiles.

Miss L. E. Curusman, F.E.S., Curator of Insects.

Prof. R. T. Lerper, D.Se., M.D., Director of Prosectorium.

Dr. C. F. Sonnac, Ch.B., Anatomist.

Dr. N.S. Lucas, M.B., Ch.B., Pathologist.

Dr. G. M. Vuvers, M.R.C.S., L.R.C.P., Hon. Parasitologist.

F, Martin Duncan, F.R.M.S., Librarian.

F. W. Bonn, Accountant.

W.H Cote, Chief Clerk,

LIST OF CONTENTS.

1920, pp. 1-194.

EXHIBITIONS AND NOTICES.

Page

Mr. R. I. Pocock, F.R.S. Report on the Additions to the

Society's Menagerie during the months of November

AMC ECETMOET OAS foc. ..,. ares « CRP ates selene a wal 189

My. A. J. Etwes, F.R.S. Letter relative to the condition

of herds of Moose in Yellowstone Park., ............... 190

Mr. D. Sera-Suiru, F.Z.8. Remarks on an interesting

case of response of Sparrows to colour .................. 190

Mr. E. G. Boutencer, F.Z.S. Exhibition of, and remarks

upon, a remarkable new Land-Tortoise (Zestudo

OCCT UUGU) IN tacts eM, « 3-3. de SE CEE Oe 190

Mr. F. Martin Duncan, F.Z.8. Exhibition of, and remarks

upon, photomicrographs of Acari from the lungs of

AU ASRETRENESS TAGS SES ROR BENE A 05 oe crea Sg OAR 190

Wes Das 5 ie Burne, F.Z.8. Exhibition of, and remarks

upon, some skeletons of feetal Mammals prepared by

MACS ONES ede aes Cote oe sks os oe SME Selo e hehd cieeee aS cre a ISH

Mr. R. I. Pocock, F.R.S. Report on the Additions to the

Society’s Menagerie during the month of January,

JOG Mert ic es see eat rcisie nlc. < 2 sto oe SRR ce oe ie eee cnet. 191

Mr. F. Martin Duncan, F.Z.S. Exhibition of, and remarks

upon, photomicrographs of a female Mole-flea (//ystri-

CHO DSUUUG ACME DOAN ieick i ce cae RRR, oS Re at ranlciohe tact 1G),

Mr. R. I. Pocock, F.R.S. Report on the Additions to the

Society's Menagerie during the month of February,

1920

Page

Mr. E. G. Boutenaer, F.Z.S. Exhibition of, and remarks

upon, a Frog with a duplicate foot ...................5 192

Prof. J. P. Hint, F.R.S. Exhibition of, and remarks upon,

an Embryo obtained from a Kangaroo recently living

in, the Society's Menagerie ©) ...).8- 5.0.2.6. -2eeie tase tiee 192

Sir Frank Coryer, K.B.E., F.R.C.S. Exhibition of, and

remarks upon, a series of skulls of Macacus rhesus ... 192

Prof. H. Maxwett Lerroy, F.Z.S. Exhibition of, and

remarks upon, photographs of Egret Farms in Sind... 192

Mr. R. H. Burne, M.A., E:Z.S. Exhibition of, and

“remarks upon, a series of Pig’s mandibles from the

New. tHlelonidles cc abe eee cca gee eerie fearie eaiaie 193

Mr. R. I. Pocock, F.R.S. Exhibition of, and remarks

upon, two Fournier’s Hutia (Capromys pilorides)...... 193,

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

Menagerie during the month of March, 1920 ......... 193

Mr. Arruur Loveriper. Exhibition of, and remarks upon,

a collection of Snakes which he had obtained in Kast

Africa during the wears T9ho = 9 holy eee. ca. nace 193

Miss L. KE. Curesman, F.E.S. Exhibition of, and remarks

upon, a series of lantern-slides illustrating the life-

history and habits of the Ichneumon-fly, Rhyssa

DET SUMDSOTU: os wc athens os eae Wacies ested eee eee eee 194

Dr. P. Cuatmers Mircunii, C.B.E., M.A., LL.D., D.Sc.,

F.R.S. Exhibition of, and remarks upon, photo-

graphs of Abbot’s Duiker (Cephalophus spadix) ...... 194

Mr. D. Seru-Suiru, ¥.Z.S. Exhibition of, and remarks

upon, lantern-slides showing the display of a male

Monaul Pheasant (Lophophorus impeyanus) .........++. 194

Mi. L, Hocsen, M.A., B.Sc. Exhibition of, and remarks

upon, a specimen of Amblystoma ......-s..-..cs0+---- <0 194

10.

PAPERS.

A Revision of the Ichneumonid Genera ZLabium and

Pecilocryptus. By Rowtanp E. Turner and JAmeEs

Wearrrsrone 3 (Rext-foumresiel = 11.) Veice sees, eas Noe

. Description of the Larynx and (isophagus of a

Common Macaque (Jacacus fascicularis) exhibiting

several abnormal Characters. By C. F. Sonwnrac,

M.D., Ch.B., F.Z.S., Auatomist to the Society.

(INSTR AUC Te e5) GSTS) et Se Eo cic cio Gee a aor

Some Australian Opiliones. By H. R. Hoae, M.A.,

ID Ziosiy Cede hates WU 19 nn eco pee eet anes eee

. Revision of the English Species of Red Spider (Genera

Tetranychus and Oligonychus). By Sranuey Hirst.

(GRexsG =m are Oe cee. ls «ns image eer eeeriee era Seek

. On the Aortic Ligament in Indian Fishes. By

D. R. Buarracuarya, M.S8c., Department, Zoological

The Muir Central College, Allahabad, India, U.P.

(Plates VG Wi vand Next-figuresl os) see ceee. os. neces:

On some Lizards of the Genus Chaleides. By E.G.

Bouuencer, F.Z.8S. (Text-figures 1-4.) ...............

. Report on the Deaths in the Gardens in 1919. With

Notes on Avian Enteritis. By Naruantet 8S. Lucas,

M.B., F.Z.S., Pathologist to the Society. (With

AC NA BUSS) Pee ie) aetna 2 oo 5s Sell Meso ela

. An Apodous Ama calva. By Arnrnur Witievy, F.R.S.,

E.Z.S:, McGill University, Montreal .....0..........:.

. On the External Characters of the South-American

Monkeys. By R. I. Pococn, F.R.S. (Text-figures

Ee) Meee ei nce he SY. Ss. 5k pa MM ee Rn a i

The Comparative Anatomy of the Tongues of the

Mammalia.—l. General Description of the Tongue.

By OC. F. Sonnrac, M.D., Ch.B, F.Z.8., Anatomist

Lowther Societys. (Vext-tiguressO=07.)) .20.0.0cessse.- ees

Page

11. Notes on East African Lizards collected in 1915-1919,

with Descriptions of a new Genus and Species of

Skink and new Subspecies of Gecko. By ArrHuR

Lovertpen. (Text-figume wl S)het sored es aaa 131

12. On Abnormal Features in the Peritoneum of a Raccoon.

By C. F. Sonnrac, M.D., Ch.B., F.Z.S., Anatomist

to the Society. (Text-figures 18-19.) .................. 169

13. On Abnormalities of the Abdominal Arteries of a

young Panda. By C. F. Sonnrac, M.D., Ch.B.,

F.Z.S., Anatomist to the Society. (Text-figure 20.).. 175

14. On the External Characters of the Ratel (d/ellivora)

and the Wolverene (Galo). By R. I. Pocock, F.R.S.,

RO7AS. oo ( lext-figures 142185) tm ce mene reas Coc ee: 179

Alphabetical last ot Contributoncsnae teense rere Vil

iindexcot Witistratrons’) 230 peepee en eee eee ee eee eee x1

AV Iie dele lex KOs IL) | JUiJL iS) a

CONTRIBUTORS,

With References to the several Articles contributed by each.

(1920, pp. 1-194.)

Page

BuatracHarya, D., R., M.Sc.

On the Aortic Ligament in Indian Fishes. (Plates I.

wand, Next one ures T—))\\\). 0). Wald eeeebe Aen ce as -. 61

Boutencer, EH. G., F.Z.S., Curator of Reptiles.

On some Lizards of the Genus Chalcides. (Text-

AVOWOMEES) ela) ) CRRA oie ea ale as0 xs ota is earn ee eee ey wa 17

Exhibition of, and remarks upon, a remarkable new

Land-Tortoise (Testudo loveridgit) .......0.0.ecceseceee seen es 190

Exhibition of, and remarks upon, a Frog with a

Clio ONIN 22) NOVO) Maile cet RIES SS do QA cay 6 OC Re 122

Burne, R. H., M.A., F.Z.8.

Exhibition of, and remarks upon, some skeletons of

foetal Mammals prepared by Tadpoles ........................ 1g

Exhibition of, and remarks upon, a series of Pig’s

mandibles from the New Hebrides ........................05. 193

CuEgEsMAN, Miss L. E., F.E.S.

Exhibition of, and remarks upon, a series of lantern-

slides illustrating the life-history and habits of the

Ichneumon-fly, Rhyssa persudsortd ..,..ccecceeerene seers ees 194

vill

Conver, Sir Franx, K.B.E, F.R.C.S.

Exhibition of, and remarks upon, a series of skulls of

WIGCACUS TREGUS cn 3.2. ok cn ae

Duncan, F. Martin, F.Z.S.

Exhibition of, and remarks upon, photomicrographs of

Acari from the lungs of Macacus rhesus ............0.00.0.4-

Exhibition of, and remarks upon, photomicrographs of

a female Mole-flea (Hystrichopsylla talpe) ..................

Kuwes, A. J., F.R.S.

Letter relative to the condition of herds of Moose in

NWiellowstone Park: 25.2.2 een ey nae era

Hitt, Prof. J. P., F.R.S.

Exhibition of, and remarks upon, an Embryo obtained

from a Kangaroo recently living in the Society’s

Mienagerie *...... ci ii Ae ee ie En ester aie

Hirst, STANLEY.

Revision of the Engish Species of Red Spider (Genera

Tetranychus and Oligonychus). (Text-figures 1-5.) ......

Hoesen, L., M.A., B.Sc.

Exhibition of, and remarks upon, a specimen of

AGTOUYSEOMUG oe via vosintek see eee nsf Roa oe

Hoge, H. R., M.A., F.Z.S.

Some Australian Opiliones. (Plates I.-III.)............

Lerroy, Prof. H. Maxwett, F.Z.S.

Exhibition of, and remarks upon, photographs of

Egret Farms DTD SSLIYEL. oo sash etae One Ree eee

Page

190

191

190

192

LovERIDGE, ARTHUR.

Exhibition of, and remarks upon, a collection of

Snakes which he had obtained in East Africa during the

Bee tance) = LOH Otis ete ta. ss calc sists a ee RE eee ois Patches

Notes on East African Lizards collected in 1915-1919,

with Description of a new Genus and Species of Skink

and new Subspecies of Gecko. (Text-figure 1.) ............

Lucas, Naruantet 8., M.B., F.Z.S., Pathologist to the

Society.

Report on the Deaths in the Gardens, 1919. (With

AS CARESS) 2 bird ces'.k Soc ee See. OR eee: Lees

MirceHELL, P. CHatmers, C.B.E., M.A., D.Sc, LL.D.,

F.R.S., Secretary to the Society.

Report on the Additions to the Society’s Menagerie

during themonthvot March, 1920 ives.

Exhibition of, and remarks upon, photographs of

Abbot’s Duiker (Cephalophus spadix) ........,.+.sceceseeeeee

Pocock, Reecinaup I., F.R.S., F.L.S., F.Z.S., Curator of

Mammals and Resident Superintendent of the

Gardens.

On the External Characters of the South American

Monkeys (Rext-teures, L=3.)) (ie machete keenest.

On the External Characters of the Ratel (J/ellivora)

and the Wolverene (Gulo). (Text-figures 14-18.).........

Report on the Additions to the Society’s Menagerie

during the months of November and December, 1919 ...

Report on the Additions to the Society’s Menagerie

duemoyune month of January, UOA0Meindss.coc esses -c..-

Report on the Additions to the Society’s Menagerie

duriny) the month of February, [9209 3. .s-22:2-. 0:

Exhibition of, and remarks upon, two Fournier’s Hutia

( (OOTOUS JOUUCTOGHES)) Corcepnencen osonadceeas -bcaber be aaccoeoaeece

Proc. Zoou. Soc.—1920. b

193

131

193

149

179

189

191

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

Remarks on an interesting case of response of

Sparrows to colour

rr eC

Exhibition of, and remarks upon, lantern-slides show-

ing the display of a male Monaul Pheasant (Lophophorus

OPO TAO KCOCIS) NAEBPBPSPRE BARD AB ice 56 0%48 ch bohene begsbeumbonddtednaT ce

Sonnrac, Cuarues F., M.D., Ch.B., F.Z.8., Anatomist to

the Society.

Description of the Larynx and Csophagus of a

Common Macaque (Macacus fascicularis). (Text-figures

1-5.)

eee e ee Feet ere oo ree EOE HEE OE EY CELE EO Her Becesreseeseseseeeese

The Comparative Anatomy of the Tongues of the

Mammalia.—I. General Description of the Tongue.

(Text-figures 6-17.)

eee te tee ete we ee es eee essere eees see rtsees-ecsece

On Abnormal Features in the Peritoneum of a

Raccoon. (Text-figures 18 & 19.)

cere eee seer eres eet ees eee eee

On Abnormalities of the Abdominal Arteries of a

young Panda. (Text-figure 20.)

weet there eee essere esses sesees

Turner, Rowianp E., and WaArTERSTON, JAMES.

A Revision of the Ichneumonid Genera Zabiwm and

Pecilocryptus. (Text-figures 1-11.)

ee pest ese ereetsoce

WarteERSTON, JAMES. See TURNER, RowLAnp E.

Wittey, Arruur, F.R.S., F.Z.8.

An Apodous Amia calva

Page

190

194

169

175

INDEX OF

Algidia cuspidata, P\. IIT, p. 31.

Alouatta, pp. 93, 95, 105, 108.

villosa, pp. 102, 105.

Aotus, pp. 99, 99.

Ateles, pp. 92, 95, 111.

ater, p. 102.

paniscus, p. 103.

Cacajao rubicundus, pp. 92, 99, 105.

Callicebus moloch, pp. 92, 98.

wean - personatus, p. 110.

Callimico goeldii, pp. 92, 95, 98, 110.

Cebus, pp. 93, 95, 101, 105, 108, 111.

Chalcides delislii, p. 79.

—— guentheri, p.79.

lineatus, p. 79.

mauritanicus, p. 79.

— ocellatus, pp. 79, 81.

, var. botteg?, p. 83.

, var. polylepis, p. 83.

sepoides, p. 79.

—— thierryi, p. 79.

tridactylus, p. 79.

Clarias magur, Pl. I., p. 61.

Eutropiichthys vacha, Pl. L., p. 61,

p- 64.

Gulo, pp. 180, 183, 186.

Hapale gacchus, pp. 105, 106, 111.

Labium bivittatum, p. 9.

ILLUSTRATIONS,

Labiuin ferrugineum, pp. 14, 19.

hohartense, p. 13.

montivagum, pp. 2,9, 17, 18, 25.

—— occidentate, p. 13.

petitorius, pp. 8, 14.

pilosum, p. 10.

sculpturatum, p. 14.

—— subequale, p. 18.

—- vasseanum, p. 13.

walker, p. 5.

Lagothrie, pp. 93, 95, 108.

infumatus, p. 101.

Leontocehus rosalia, pp. 92, 105, 106,

111.

Macacus fascicularis, pp. 27, 29.

Macropsalis chiltoni, Pl. 1., p. 31.

Mammalia, Tongues of, pp. 116-125,

2, NPAs),

Mellivora, pp. 180, 182, 185, 186.

Monoxyommea henlei, P\. I11., p. 31.

trawli, Pl. WIL., p. 31.

-—— tuberculata, Pl. I1I., p. 31.

Nuncia smithi, P\. 11., p. 31,

Oligonychus quercinus, pp. 51, 59.

—- simplex, pp. 51, 59.

—— ulmi, pp. 51, 59.

ununguts, pp. 51, 59.

Panda, p. 176.

Pantopsalis gray, Pl. I., p. 21,

X1i INDEX OF ILLUSTRATIONS.

Pantopsalis halli, Pl. 1., p. 31.

pococki, Pl. 1., p. 31.

wattet, Pl. I., p. 3l.

Pecilocrypius nigromaculatus, pp. 17,

20.

Pseudeutropius garua, pp. 63, 65.

Raccoon, pp. 170, 172.

Rita buchanani, Pi. 1., p. 61.

Saimiris sciurea, pp. 93, 95, 98, 105,

106, 111.

» Scolecoseps boulengert, p. 159.

Tetranychus carpini, pp. 51, 52.

crategi, pp. 51, 53, 54.

—— lintearius, pp. 52, 53, 54.

populi, pp. 51, 58, 54.

—— schizopus, pp. 51, 52, 53.

talisee, pp. 51, 53, 54.

-— telarius, pp. 51, 52, 53, 54.

—— tiliarium, pp. 51, 52, 53, 54.

Trienobunus acuminatus, Pl. 11., p. 31.

Trienonyx cockayni, Pl, 11., p. 31.

variegata, Pl. I1., p. 31.

Wallago attu, Pl. I1., p. 61.

INDEX.

1920.—Pages 1-194,

[New names in clarendon type.

Systematic references in italics.

(z.8.1.) indicates additions to the Society’s Menagerie. |

Ablepharus boutont, var. peronit, 157.

wahlbergit, 158.

Acanthopterygti, 71.

Agama atricollis, 142.

colonorum, 140.

flavicauda, 141.

lionotus, 141.

vaillanti, 142.

Agamide, 140.

Ailie coila, 70.

Algidia, gen. n., 46.

cuspidata, sp. n., 47.

Alouatta, 93, 94, 96, 97, 101, 102,

104, 106, 107, 112, 113.

(Mycetes) villosa, 92.

Amia calva, 89.

Aniurus natalis, 89.

Ammotraqus lervia, 198.

Amphipnoide, 71.

Amphipnous cuchia, 71.

Amphisbenide, 145.

Anabantide, 71.

Anabas scandens, 71.

Anelytropide, 160.

Anguilliformes, 71.

Aotus, 93, 94, 96, 100, 104, 106,

118.

103,

ey,

—— (Nyctipithecus) trivirgatus, 92.

Proc. Zoou. Soc.— 1920.

| —— (Ouakaria), 93, 94, 96, 97,

Ateles, 91, 93, 94, 97, 100, 103,

109, 112, 118.

ater, 92.

Bagarius yarrellii, 70.

Bulistes erythrodon, 72.

stellaris, 72.

Balistide, 72.

Barbus sarana, 67, 70.

tor, (2.

Barilius bola, 70.

Belone cancile, 71.

strongylura, 71.

| Berycide, 71.

104,

Brachyteles, 91, 94, 103, 104, 112, 113.

Cacajao (Brachyurus, Ouakaria) rubi-

cundus, 92.

104, 112, 113.

Calamoichthys, 89.

103,

Callicebus, 93, 94, 96, 97, 100, 104, 106,

Te Ws}.

—~—- (Callithrix) moloch, 92.

—— personatus, 107, 109.

Callichrous macrophthalimus, 67, 70.

XIV

Callichrous pabda, 67, 70.

Cullimico, 94, 96, 97, 98, 100, 104, 109,

112, 113.

goeldit, 92.

Capromys pilorides, 193.

Carangide, 72.

Caranx atropus, 72.

gallus, 72.

Catarrhactes chrysolophus

192.

Catla buchanani, 65, 67, 70.

Cebus, 93, 94, 96, 97, 100, 103, 104,

106, 107, 112, 118.

-—— albifrons, 92.

apella, 92.

Cephalophus spadia, 194,

Chetodontide, 72.

Chalcides, (7.

——- bedriage, 78.

bottegi, 77, 80, 83.

delislii, 80.

—— guenthert, 80.

lineatus, 80.

— linose, 79, 80, 82.

—— mauritanicus, 80.

— occidentalis, 80.

ocellatus, 77.

polylepis, 80.

ragazztt, 80, 82.

(z. 8. L.);

sepoides, 80.

simonyi, 78.

—— tiliqugu, 80, 82.

— tridactylus, 80.

—— viridanus, 78.

vittatus, 80, 82.

Chameleon biteniatus, 168.

——- dilepis, 160.

tsabellinus, 168.

-—— gracilis, 160.

—— hehnelii, 163.

jacksoni vauerescece, 163.

melleri, 166.

Chameleontide, 160.

Channa orientalis, 90.

Chasmorhynchus nudicollis, 191.

Chatoessus chacunda, 71. *

manminna, 67, 71.

Chelys fimbriata, 190,

Chirocentride, 70.

INDEX.

Chirocentrus dorab, 70.

Cinnyris mariquensis (z. 8. L.), 189.

Cirrhina mrigala, 70.

reba, 70.

Clarias magur, 70.

Clupea alosa, 61.

chapra, 71.

— ilisha, 67.

sindensis, 67.

Clupeide, 71. .

Clupeiformes, 70.

Conepatus, 179, 184.

Coronella amabilis, 189.

zonata, 189,

Cricetulus migratorius (z. s. L.), 189.

Cuscus, 102.

Cynoglossus quinguelineatus, 72.

Cyprinide, 70.

Cypriniformes, 70.

Dipsosaurus dorsalis, 189.

Licheneis naucrates, 72.

Echeneidide, 72.

Elasmodactylus triedrus, 140.

Engraulis malabaricus, 71.

telara, 67, 70.

Ephippus orbis, 72.

Equus pryevalskii, 189.

Hremias spellti, 148.

Esociformes, 71.

Eurypyga hetias, 190.

Eutropiichthys vacha, 67, 70.

Felis caracal, 189.

leo, 189, 198.

—— tigris, 189.

Feylinia currori, 160.

Galeopithecus, 116.

Galera, 179.

Galietis, 179.

Gastropholis vittata, 147.

Gazella subgutturosa, 191.

INDEX. xV

Gazza equuleformis, 72.

Geckonide, 132.

Gerres filamentosus, 72.

Gerride, 72.

Gerrhosauride, 149.

Gerrhosaurus flavigularis, 150.

major, 149.

nigrolineatus, 150.

Gobiide, 72.

Gobioides tenius, 72.

Grisonia, 184.

Gulo, 179, 181, 182, 184, 186, 187,

193.

Hapale jacchus, 91, 97, 98, 104, 105,

106, 109, 112.

Hemidactglus brookii, 134.

citerntt, 132.

—— mabouia, 133.

——- ruspolti, 134.

—— squamulatus, 134.

Histiophorus, 89.

Holacanthus xanthurus, 72.

Flolaspis guenthert, 149.

Hystrichopsylla talpe, 191.

Ichnotropis capensis, 148.

Ictonyx, 179, 184, 187.

Julis lunaris, 72.

Labeo ceruleus, 70.

calbasu, 70.

diplostomus, 67, 70.

rohita, 65, 70.

Labium approximatum, sp. n.,

21.

—— associatum, sp. n., 20.

bicoler, 24.

—— bivittatum, sp. n., 8.

—— brevicorne, sp. n., 6.

— centrale, sp. n., 6.

clavicorne, 19.

Serrugineum, 19.

-—— fulvicorne, sp. n., 22.

—— hobartense, sp. n., 20.

| Labium longicorne, sp. n., 23.

| ——— montivagum, sp. n., 16.

| —— multiarticulatum, sp.r., 23.

—— occidentale, sp.n., 16.

petitorius Wrichs., 7.

, subsp. concolor, subsp. n.,

——- pilosum, sp. n., 10.

| ——sculpturatum, sp. n., 14.

—— spiniferum, sp.n., 12.

—— subzquale, sp. n. 15.

— subpilosulum, sp. n., 11.

—— vasseanum, sp. n., 13.

—— walkeri, sp. n., 4.

, Key to the Species of, 3.

Labride, 72.

Lacerta vawereselli, 147.

Lacertide, \47.

Lagothri«x, 94, 96, 97, 101, 102, 108,

104, 106, 107, 112, 113.

infumatus, 92, 109.

-— lagotricha (=humboldti), 92.

Laniatores, 36.

Latastia johnstoni, 148.

longicaudata, 147.

Lates calearifer, 72.

Leontocebus rosalia, 91, 105, 106, 109,

112.

Lethrinus miniatus, 71.

Lophophorus impeyanus, 194.

Lutra maculicollis (z. s. u.), 189.

Lygodactylus capensis (Smith),

subsp. n., 135.

Jischeri scheffleri, 136.

—— grote, 136.

— picturatus, 136.

Lygosoma ferrandii, 157.

sundevallii, 155.

Lyncodon, 179.

Mabuwia brevicollis, 152.

maculilabris, 152.

— megalura, 152.

—— quinqueteniata, 153.

striata, 153.

varia, 103.

Macacus fascicularis, 27.

rhesus, 190 192.

Xvi

Macrones aor, 67, 70.

~— cavasius, 70.

seenghala, 67, 70.

Macropus bennetti, 191.

Mamunalia, tongues of, 115.

Martes, 179, 181, 187.

Mastacembelde, 72.

Mastacembelus armatus, 72.

Meles, 179, 181.

Mellivora, 179, 181, 182, 184, 185, 187,

193.

Mephitis, 179, 184.

Monopeltis colobura, 145.

Monoxyomma heudei, sp. n., 44.

spinatwm, 44.

—— trailli, sp.n., 45,

tuberculatum, sp. n., 44.

Mugil corsula, 71,

Mugilide, 71.

Mugiliformes, 71.

Murena macrura, 71.

—— punctata, 71.

sathete, 71.

tessellata, 71.

Murenide, 71.

Mustela, 179, 181, 187.

putorius, 184.

Mydaus, 184.

Myocastor coypus, 192.

Myripristis murdjan, 71.

Mystax midas, 91.

mystax, 91, 96, 97, 98, 106, 109,

112.

—— ursulus, 91, 109.

Notopteride, 70,

Notopterus chitala, 67, 70.

kapirat, 70.

Nucras emini, 147.

Nuncia enderbe@i, 42.

smithi, sp. n., 42.

— sperata, 41.

sublevis, 41.

valdiviensis, 41,

(Edipomidas, 96.

edipus, 91, 94, 105, 106, 112.

Oligonychus quercinus, 59,

uli, d8.

INDEX.

Oligonychus simplex, 60.

—- ununguis, 59.

Ophichthys boro, 71.

Ophiocéphalide, 71.

Ophiocephalus marulius, 71.

punctatus, 71.

striatus, 71, 90.

Otolithus ruber, 72.

Palpatores, 32.

Panda, Abnormalities of the Abdominal

Arteries of a young, 175.

Pangasius buchanani, 70.

Pantopsalis, 32.

—— coronata, 33.

—— pgrayi, sp. n., 35.

— halli, sp. n., 34.

Jenningst, 39.

listeri, 33.

-—— nigripalpts, 3o.

—— pococki, sp.n., 34.

tasmanica, 33.

trippt, 33.

—— wattsi, sp. n., 33.

Pellona brachysoma, 71.

elongata, 71.

Percide, 72.

Phalangiide, 32.

Phascolarctos, 102.

Phelsuma laticauda, 139.

Pithecta, 91, 97, 112, 113.

Platycephahide, 72.

Platycephalus scaber, 72.

Platyphotis fasciata, 139.

Pleuronectide, 72.

Plotosus arab, 70.

Poecilocryptus nigripectus, sp. n.,

24.

-— nigromaculatus, 26.

Pecilogale, 179.

Polynemide, 71.

Polynemus plebeius, 71.

Polypterus, 89.

Procavia capensis, 189.

Psetiodes erwnet, 72.

Pseudeutropius garua, 61, 62, 63, 67,

68, 70.

Putorius, 179.

INDEX.

Raccoon, Abnormal features in the

Peritoneum of a, 169.

Report on the Deaths in the Gardens

in 1919, 85.

Rhanpholeon brevicaudatus, 166.

kerstenit, 167.

Rhyssa persuasoria, 194.

Rita buchanani, 67, 70.

Saccobranchus fossilis, 70.

Saimiris, 94, 96, 97, 100, 104, 106, 107,

112, 1138.

(Chrysothrix) sciureus, 92.

Scatophagus argus, 72.

Scelotes eggeli, 159.

Sciena maculata, 72.

Scienide, 72.

Scienoides pama, 72.

Scincide, 152.

Scolecoseps boulengeri, sp. n.,

159).

Scombresocide, 71.

Scombride, "72.

Serranide, 71.

Serranus angularis, 71.

Sillaginide, 72.

Sillago sihama, 72.

Silundia gangetica, 67, 68, 70.

Siluride, 70.

Simia, 104.

Sorensenella hicornis, 48.

— prehensor, 43.

Sparide, 71. —

Stromateide, 71.

Stromateus sinensis, 71.

Symbranchiformes, 71.

Synagris tolu, 72.

Tanqua tiara, 145.

Taurotragus oryx, 193.

Taxidea, 181.

Testudo loveridgii (z. s. u), 190, 191.

XVII

Tetranychus carpini, sp. n., 56.

— cratezgi, sp.n., dl.

—— populi, 52.

schizopus, 50.

—— talisiz, sp.n., 54.

——— telarius, 50.

tiliarium, O7.

-——, Key to the species (males), 49.

Therapon jarbua, 71.

Thynnus pelamys, 72.

Trachynotus insidiator, 72.

Triacanthide, 72.

Triacanthus brevirostris, 72.

Triznobunus acuminatus, sp. n.,

36.

Trienonyx aspera, 39.

cockayni, sp. n., 39.

coriacea, 39.

rapax, 39.

stewartius, 39.

testaceus, 39.

variegata, sp. n., 40.

Trichiuride, 72.

Trichiurus savala, 72.

Umbrina russellii, 72.

Ursus americanus, 191.

arctos, 191.

Varanide, 143.

Varanus albigularis, 143, 192.

— niloticus, 144.

Vormela, 187.

Wallago attu, 67, 70.

Xiphias, 89.

Zonuridx, 143.

Zonurus derbianus, 192.

--— tropidosternum, 145.

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1920, Parts I. & Il. (pp. 1-194).

EXHIBITIONS AND NOTICES.

Mr. R. I. Pocock, F.R.S8. Report on the Additions to the Society’s Menagerie during the

monthsrof November, and December, eu s arse reiersiellerelleyche eaieiciers = «1s =e = ale) eielceneaetle

Mr. A. J. Exwes, F.R.S. Letter relative to the condition of herds of Moose in Yellow-

WALONG Paves eck oe Shale osc ao whehenecalepe coer Menera Maman On ctar a eN omer ate Velatelaylatells ails le eveysters/ eloneeeemntets

Mr. D. Sern-Surrn, F.Z.S. Remarks on an interesting case of response of Sparrows to

Pi KN he a pa Se ROM m eines oon OO Te St Sloan bs anor meeP ear gars co”

_ Mr. E. G. Bounszyerr, F.Z.8. Exhibition of, and remarks upon, a remarkable new Land-

Tortoise (Testudo loveridgit) ..++.e.seeceee- Pel gen eia We cvateveNaiaieiar oro\ls evevansieeersteneseyeamte

Mr. . Martin Doncan, F.Z.8. Exhibition of, and remarks upon, photomicrographs of

Acari from the lungs of Macacus :hesuS s. 25.01. ccc fence cso een ee wounesiv ns

Mr. E. H. Burns, F.Z.S. Exhibition of, and remarks upon, some skeletons of foetal

Mammals prepared by. Cad poles) 5. <u teaers Gis lage’ ssanieieinjersin cia les cls pis leew eet anlar

Mr. R. I. Pococn, F.R.S. Report on the Additions to the Society’s Menagerie during the

iON Oud h eva SPO. neion anind do Ur ons sodn odGanmbDoatoOSOntuEsco nD eooOusuC

Mr. F, Martin Duncan, F.Z.8. Exhibition of, and remarks upon, photomicrographs of a

female Mole-Hea (Hystrichopsylla talp@) soo. canoe bese cees sewer sc ecsenecesnes

Mr. R. I. Pocock, F.R.S. Report on the Additions to the Society’s. Menagerie dune the

month of Bebraanys L920: is: alathcete australe tc cuelnpacee’s Ioicis ape (sporsaey ar atheie se ae

My. HE. G. Bovtenerr, F.Z.S. Exhibition of, aud remarks upon, a Frog with a duplicate

ROOtimnetslclavers erereiercrerarcts cielo s obatexeustettanenekeneyentte Cbs boca 0Un awe gae Sale ouster alene ouytiate

Prof. J. P. H1it, F.R.S. Exhibition of, and remarks upon, an Embryo obtained from a

Kangaroo recently living in the Society’s Menagerie ..... .

eC ee rare eco reese we eens

Sir Frank Coryer, K.B.E., F.R.C.S. Exhibition of, and remarks upon, a series of skulls

OP GA CUCIUSHESUS: ino ai) a care ss'e ate ss6 Se eee RTOs aon tented ora Fare evol tes i nara eee

Prof. H. Maxweut Lerroy, F.Z.8. Exhibition of, and remarks upon, photographs of

Heret Farms in Sind

Pe ee eC Ce rer)

Mr. R. H. Burne, M.A., F.Z.S. Exhibition of, and remarks upon, a series of Pig's

mandibles from the New Hebrides ....... dao eae 5

CC ee ee er ie rd

Mr. R. I. Pococs, F.R.8. Exhibition of, and remarks upon, two Fournier’s Hutia

(Capromys pilorides’ .eseversecccecerease bidee aroc Sharada sist ate ua) Nioveles sfataverse ctjoustestetteieys

The Szcrerary. Report on the Additions to the Society's Menagerie during the month of

Marelt; 1920)® iiagiercavereusie'a seis, a alaile a) Set aleparcretonae tetas couetcartwien a alcalePare: areieter anette: Cire fexeewer

Mr. Arruur Lovertper. Exhibition of, and remarks upon, a collection of Snakes which

he had obtained in East Africa during the years 1915-1919

Page

190

191

192

Contents continued on page 3 of Wrapper.

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PAPERS.

1. A Revision of the Ichneumonid Genera Labiwm and

Peeilocryptus. By RowLtanp E. Turner and Jamus

~ WATERSTON.

{Received November 14, 1919: Read February 10, 1920.]

(Vext-figures 1-11.)

Genus Lasium Brulleé.

Labium Brulle, Hist. Nat. Insect. Hymén. iv. p. 316 (1846).

The genus Labiwm, though the typical species was described

from New Guinea, is really characteristic of the southern half of

Australia. No specimens are in the British Museum from the

northern half of Australia, and we may certainly assume that the

genus does not occur in the north-eastern coast regions, unless at

high elevations where little collecting has been done. In the

extreme south of Australia and in Tasmania various species of

the genus are exceedingly common, swarming on the blossom of

Leptospermum. It is most remarkable that so little work should

have been done on a genus so conspicuous and so plentiful as this,

and that nothing should be known of its life-history. Before

Brullé detined the genus Erichson had already described a

Tasmanian species as /chnewmon petitorius, but the genus was not

again recognised till Cameron described L. ferruginewm in 1901,

Morley later (1915) adding another species, Z. clavicorne. The

genus is extremely isolated among the Ichneumonide, the most

striking characters being the long exserted labrum, the more or

less clavate antenne, the large ungues and empodium of the hind

Proc. Zoou. Soc.—1920, No. I. 1

2 MESSRS, R. E. TURNER AND J. WATERSTON ON THE

tarsi, and the strongly bent recurrent nervure. In one or two

aberrant species one or more of these characters may be only

feebly developed, but the long labrum is always present. The

best account of the genus is given by Cameron, but his description

of the generic characters is founded on a single species, and one

or two characters given are merely specific. Thus the petiolar

area and areola are usually separated, not confluent, and the

nervulus is often more or Jess postfurcal, as noticed by Morley.

Cameron is inclined to place the genus in the Jehnewmonine, as

a separate tribe; the great objection to this placing is in the

position of the spiracles on the petiole only a little behind the

middle, and much nearer to each other than to the apex of

the segment. It seems, however, on the whole, a more natural

position than either among the 7ryphonine, with which it was

placed by Brullé, or near Banchus, where Morley is inclined to

Text-figure 1.

AEN

SQW «

\\ (i

Ki,

Wi <

My : p TERZIVJ

= .

Labium monutivagum Turn. & Wtrst. Thorax, propodcon, and petiole

in profile.

place it. We give a full account of the sculpture of the median

segment (text-figs. 1, 3, and 10), as far as it is common to all the

Species: carina between the short basal area and areola usually

indicated by short lateral stumps ; areola and petiolar area some-

times confluent ; pleural and juxta-coxal are confluent anteriorly,

but separated up to 4 or beyond posteriorly. ‘The surface of the

basal area and areola is smooth, more or less rugulose round the

sides ; of the external area smooth or finely punctured: the an-

terior part of the spiracular area is also punctured, generally rather

strongly so. ‘The other are vary in sculpture, the juxta-coxal

always showing some strong ridges, indicated at least anteriorly.

Sculpture of the male usually coarser than in the female. The

sculpture of the arez is liable to considerable individual variation

in intensity.

Species of Thynnide of the genera Lirone and Tachynomyia

ICHNEUMONID GENERA LABIUM AND PCSCILOCRYPTUS. 3

bear a strong superficial resemblance to species of this genus.

1. abdominalis Guér. and Hirone celsissima Turn., both of which

look like Zabiwm, occur on Mt. Wellington at a considerable ele-

vation, where Labiwm montivagum is very plentiful. £. celsissima

even has the apical antennal joints yellow, a most abnormal

coloration for one of the Thynnide.

Key to the Species of Labium.

1. Antenne not more than 30-jointed

Antennz more than 30-jointed .....................

2. Second recurrent nervure only feebly bent in the

middle; nervellus straight, discoidella want-

ing; head, thorax, and median segment. black

with yellow markings Lc . DL. walkeri, sp.n., bd, p. 4

Second recurrent nervure sharply “angulate in

the middle, nervellus angulate, discoidella not

wholly wanting ;, head, thorax, and median

segment suo eM oeS with yellow mark-

TEI) cogon- 3.

3. Discoidella reduced toa short stump, ‘originating

halfway between the submediella and the

middle of the nervellus; median segment

yellow apically ............ . L. centrale, sp.n., p. 6.

Discoidella complete, originating much nearer to

the middle of the nervellus than to the sub-

mediella; median segment ae fulvo-fer-

ruginous ...... . LL brevicorne, sp. n., p. 6.

4. Face and clypeus fused, in the same » plane, the

dividing-line obsolete ; more or less pilose... 5.

Face and clypeus not entirely fused or in the

same plane; scarcely pilose....... 6.

5. Strongly pilose; hind femora ‘and median § seg-

ment black .. ; . LL. piloswm, sp. v., p. 10.

Feebly pilose ; hind femora and median ‘segment

fulvo-ferruginous BSNS Ae REA ors Sepa eA L, subpilosulum, sp. n., p. 11.

6. Hind empodia small, not more than one-third of

the length of the tarsal ungues; mesonotum

with a broad black fascia on cach side ......... EL. bivittatwm, sp.n., p. 8.

Hind empodia elongate, at least two-thirds of the

length of the tarsal ungues; mesonotum with-

out black lateral fascie ..... Oe

. Mesonotum opaque, closely and coarsely. pune-

tured ; dentiparal area bepstuted into a broad

well- developed tooth . 8.

Mesonotum shining, smooth or - finely punctur ed;

dentiparal area unarmed, or at most with an

“I

inconspicuous tooth ......... Forest i).

8. Scutellum and postscutellum yellow Fara ceeee LT. petitorium Urichs., p. 7.

Scutellum and postscutellam ferruginous .. . L. petitorium, subsp. con-

9. Flagellum of the antennze wholly black above, [eolor, subsp. n., p. 8

without pale apical joints......... 10.

Flagellum, if black above, then always Can

several of the apical joints conepiou easly fer-

ruginous or ochraceous .. Bence serios 11.

‘10. Mandibles very long and ‘straight, vertical ;

dentiparal area with a small, but distinct

tooth at the external apical angle ‘gatos EERE L. vasseanum, sp.n., 6, p.13

Mandibles normal, not elongate; dentiparal area

WINENE TT (Gl <ckcee sou lacuna oa. de Ae AMMAR BER Rne eco ic L. ferrugineum Cam., p. 19.

11. External area of the median segment more

coarsely sculptured than the deutiparal area ;

external subopaque, finely and closely punc-

tured, dentiparal smooth and shining ......... 12.

4 MESSRS. R. E. TURNER AND J. WATERSTON ON THE

Dentiparal area more coarsely sculptured than

the external or both smooth and shining ...... 13.

12. Antenne from 50- to 55-jointed ; hind tarsi fer-

ruginous, median segment with a transverse

yellow fascia oes litttttsses. i multiarticulatum, sp. n.,

Antenne 45- to 46- jointed; “hind tarsi black [p. 23.

from before the apex of the metatarsus, median

segment without a yellow fascia . bs . LL. fulvicorne, sp.u., p. 22.

13. Hind femora black, very narrowly yellow. at the

apex ...... . L. sculpturatum, sp.n., 3,

Hind femora ferr ‘uginous, at most narrowly black [p. 14.

SHE THOS) ORISA ENOL EYWERS food sovoasaonocsos boas2>oubongace 14,

14. Antennze more than 40-jomted ................0... 15.

Antennz less than 40-jointed ........................ 18.

16. Mandibles bidentate; basal tergites narrowly

margined with yellow ...................... L. hobartense, sp. n., p. 20.

Mandibles simple ....... 16.

16. Dentiparal area unarmed, ‘areola, external area,

and dentiparal areze smooth... tne ae eae L. longicorne, sp.n., 3, p. 23.

Dentiparal area armed, the carine strongly raised

forming small teeth at the outer angles........ 17.

17. Antenne 47-jointed; face yellow, closely ‘and

Cha yr FOUUVCWORECL asco asec ist nano sect asdane ano ca: LL. approximatum, sp. n., 6,

Antenne 43-jointed : face ferruginous, margined [p. 21.

with yellow laterally, centrally rugulose ...... L. spiniferum, sp. n., p. 12.

18. Third and fourth antennal joints subequal ...... i),

Third antennal jomt much longer than the

fowtthy eee. Ea Mae ene 20.

19. Mandibles str ‘ongly ‘bidentate ; ; “hind femora

shining, with Jar ee and rather sparse punctures. L. occidentale, sp. n., p. 16.

Mandibles very obscurely bidentate; hind femora

subopaque, very closely covered with minute

punctures......... L. subequale, sp.n., p. 15.

20. Distance between the recurrent and second trans- :

verse cubital nervures not equal to more than

one-third of the length of the latter nervure ;

nervulus very distinctly postfurcal ............... LL. clavicorne Morl., p. 19.

Distance between the recurrent nervure and the

second transverse cubital nervure equal to at

least half the length of the latter nervure ;

nervulus interstitial or very ey post-

WOKEN 5.6 oo Sb al 21.

21. Petiole black, ‘except ‘at the ‘apex ; “ mesopleure

extensively black superiorly....... LL. montivagum, sp. n., p. 16.

Petiole mainly dH Cus mesopleurze with

very little black .. Won iccbeebtenecnoudacasazong 4a CISRYIXGHAEO HI, Oe Ts, 1) 240)

Labium bicolor Brullé is not included in the Key, as we have not seen a

specimen. It is described from a male, and has the hind femora black as in

L. sculpturatum, to which, from the description, it appears to be related.

LABIUM WALKERI, sp.n. (Text-fig. 2.)

3d. Niger; mandibulis, apice brunneis, labro, clypeo, facie,

fronteque lateribus, genis, orbitis externis, scutello, basi anguste

nigro, postscutello, tegulis, mesopleuris maculis duabus, epimeris,

pedibus anticis inter medi sque, tarsis iIntermediis br unnescentibus,

coxis posticis apice, trochanteribus posticis, genubus, tibiisque

posticis dimidio basali, basi angustissime brunneo annulatis,

flavis; antennis subtus rufescentibus ; femoribus posticis medio

late rufescentibus, basi atque apicem versus nigrescentibus ;

ealcaribus pallidis ; alis subhyalinis, venis fuscis: petiolo apice

ICHNEUMONID GENERA LABIUM AND PGiCILOCRYPTUS. 5

in medio late testaceo, tergitis 2-5 brunneo-ferrugineis, apice

anguste flavo-limbatis; tergitis apicalibus fuscis; sternitis quarto

sequentibusque flavo-testaceis.

Long. 6 mm.

3. Mandibles simple; clypeus distinctly separated from the

face, convex; face smooth, subquadrate; vertex and temples

smooth and shining. Antenne 28-jointed, third and fourth

joints subequal, joints five to eight diminishing gradually, in the

funicle the first fifteen joints are cylindrical. “Thorax shining,

epimeral furrows crenulate throughout. Median segment with

the apical carina of the basal area complete, areola and external

areze smooth; petiolar and adjacent arex, the apical two-thirds

of the spiracular aree, and the dentiparal aree subrugulose ;

pleural ares smooth, anteriorly crenulate, with five or six strong

Text-figure 2.

Labium walkeri Turn. & Wtrst. Wings.

ruge in front of the coxa. Petiole without a subbasal tooth, but

the spiracular area prominent; spiracles well behind the middle ;

abdomen shining, smooth. Areolet (text-fig. 2) pentagonal, second

abscissa of the radius shorter than either of the transverse cubital

nervures, distance between the recurrent nervure and the second

transverse cubital nervure equal to nearly half the length of the

latter nervure; second recurrent nervure only feebly bent in the

middle, not abruptly angled as in other species of the genus.

Discoidella missing; nervellus straight, not angled.

Hab. Hobart, Tasmania; summer, 1891 (J. J. Walker).

Very distinct im the neuration from the larger species of the

genus. Probably this will eventually prove to be generically

distinct, but in the present state of our knowledge it is hardly

necessary to divide the genus.

6 MESSRS. R. E. TURNER AND J. WATERSTON ON THE

LABIUM CENTRALE, Sp. li.

2. Fulvo-ferruginea ; mandibulis, apice nigris, labro, clypeo,

facie, fronte, scapo, pedicello, propleuris antice, mesonoto lateri-

bus anguste, seutello, sulco basali lateribusque nigris, postscu-

tello, mesopleuris supra fasciaque infra, segmento mediano fascia

lata postica, coxis anticis intermediisque, tibiisque posticis

dimidio basali flavis; tarsis posticis articulo apicali, tibiisque

posticis dimidio apicali intus fusco-ferrugineis; alis hyalinis,

venis brunneis ; stigmate brunneo, macula basali flava.

Long. 6 mm.

?. Mandibles simple; line between the clypeus and face

distinct, less so in the middle than at the sides, the face and

clypeus sparsely, but not very finely punctured, face broader

than long; eyes rather strongly emarginate above the base of the

antenne; vertex and front smooth. Antenne 27-jointed, third

joint only a little longer than the fourth, twelfth funicular joimt

quadrate, thirteenth transverse. Notauli short; middle lobe of

mesonotum not prominent, rather coarsely punctured, the lateral

lobes very sparsely punctured ; the sharp anterior edge of the

mesopleure is rather short, extending about halfway towards the

spiracle and just extending a little beyond the first longitudinal

hollow ; epimeral sulcus crenulate throughout. Prepectus antero-

ventrally crenulate. Median segment with the apical carina of

the basal area only indistinctly indicated laterally ; areola very

broad, not clearly separated from the petiolar area, smooth,

punctured near the sides; external area subrugulose, dentiparal

and spiracular are a little more rugulose, the spiracular area tend-

ing to be punctured anteriorly ; pleural area distinctly separated

from the juxta-coxal on the posterior two-thirds, the carina

obsolete anteriorly. Petiole with the lateral teeth between the

spiracles and the base well developed. Hind coxe and femora

smooth, with a few scattered punctures. Nervulus postfureal ;

areolet pentagonal, distance between the recurrent nervure and

second transverse cubital nervure equal to half the length of that

nervure, second abscissa of the radius shorter than the secon

transverse cubital nervure; nervellus angled far below the middle

(at about three-fourths), discoidella represented only by a short

stump at that point, entirely obsolete beyond the stump, without

a vena spuria.

Hab. Killalpanima, 100 miles E. of Lake Eyre (H. J. Hillier).

LABIUM BREVICORNE, Sp. n.

2. Fulvo-ferruginea ; mandibulis, apice excepto, labro, clypeo,

facie, scapo, pedicello subtus, fronte lateribus, scutello, post-

scutello, mesopleuris maculis sub alis, tegulis, linea verticali supra

coxas anticas, pedibusque anticis intermediisque, illius tarsorum

articulo apicali excepto, flavis; antennis supra infuscatis usque

ad articulam vicesimum, subtus omnino articulisque septem

apicalibus ferrugineis; scutello fovea basali et Jlateribus, pro-

ICHNEUMONID GENERA LABIUM AND PCCILOCRYPTUS. 7

pleuris antice, segmento mediano area juxta-coxali, tiblis posticis

dimidio apicali, tarsisque posticis articulo apicali nigris.

Long. 7 mm.

Q. Mandibles simple; face and clypeus shining, very finely

and sparsely punctured, the line between the clypeus and face

distinct. Antenne 28- to 29-jointed, the third joint less than

half as long again as the fourth, which is slightly longer than the

fifth, the basal twelve joints of the funicle cylindrical. Vertex

and front smooth and shining. Notauli shallow and short,

middle lobe of the mesonotum not prominent anteriorly, smooth ;

mesopleure shining, epimeral sulcus crenulated throughout.

Median segment with the basal area very short, scarcely

extending beyond the posterior edge of the sulcus separating

the postscutellum and median segment, the apical carina only

indicated laterally ; areola broad and smooth, sparsely punctured

towards the sides, not distinctly separated from the petiolar

area, which with the adjacent aree is longitudinally rugulose ;

external area smeoth ; dentiparal area shining, but with a more

uneven surface; spiracular area anteriorly moderately and

posteriorly coarsely punctured; pleural area smooth, with a

few scattered punctures, juxta-coxal area rugose. Petiole with

a well-developed tooth between the spiracles and the base;

spiracles prominent, the petiole rather abruptly widened behind

the spiracles; abdomen shining. Hind coxe smooth, with at

most a few scattered punctures beneath. Second abscissa of the

vadius shorter than the second transverse cubital nervure,

distance between the recurrent nervure and the second trans-

verse cubital nervure equal to about three-eighths of the length

of the latter nervure ; nervulus slightly postfureal.

Hab. Swan River (Du Boulay); Yallingup, 8.W. Australia,

November, 1913 (Turner).

Lasium perirorium Hrichs. (Text-figs. 3 and 7 c.)

Ichnewmon petitorius Evichs. Arch, f. Naturges. p. 255 (1842).

Hab. KWaglehawk Neck, 8.H. Tasmania; February (Zurner).

Common.

This is distinguished from other species of the genus by the

opaque, strongly and very closely punctured mesonotum and meso-

pleure and the strongly developed teeth (text-fig. 3) of the denti-

paral avez. The division between the clypeus and face is well

marked, both clypeus and face closely punctured ; antenne of the

female 42-jointed ; median segment with the apical carina of the

basal area interrupted in the middle, external area smooth, denti-

paral area coarsely and irregularly longitudinally striate, spiracular

area rugose, juxta-coxal and pleural arez rugose-striate, postero-

intermedia] and postero-external arez transversely striate, areola

shining with a few large punctures, petiolar area indistinctly and

irregularly transversely striated. Second abscissa of the radius

and first transverse cubital nervure subequal, second transverse

8 MESSRS. R. E. TURNER AND J. WATERSTON ON THE

eubital nervure slightly longer, the distance (text-fig. 7, c) between

the recurrent nervure and the second transverse cubital nervure

equal to more than half of the length of the latter nervure.

Antenne 42-jointed.

Subspecies concolor, subsp. n.

@. Differs from the typical Tasmanian form in having the

scutellum and postscutellum ferruginous, only very slightly paler

than the rest of the thorax, not yellow as in the typical form.

The second abscissa of the radius is somewhat shorter than in

the typical form and the two transverse cubital nervures sub-

equal. Antenne with three or four joints less than in the typical

form.

Hab. Yallingup, 8.W. Australia ; October-December (Turner).

Text-figure 3.

\ i a

TERZI—~

Labium petitorium Erichs. Propodeon. (a) dorsal; (6) profile; (ex) coxa

of hind leg; (pé) petiole.

To show outlines of the aree. Sculpture of arew not detailed fully.

LAsiuM BIVITTATUM, sp.n. (Text-fig. 4, 6.)

2. Ochracea; labro, clypeo, facie lateribus, macula lineari

sub alis, scutello postseutelloque flavis; vertiee late, propleuris

dimidio inferiore, mesonoto fascia lata longitudinali utrinque,

tergito primo apice, coxis posticis macula magna dorsali apicali,

femoribus posticis basi et apice, tibiisque posticis dimidio apicali

nigris ; alis hyalinis, venis fuscis; antennis fuscis, infra ferru-

gineis, articulis 9 apicalibus fulvo-ochraceis,

$. Femine similis; antennis articulis 7 apicalibus fulvo-

ochraceis ; tibiis posticis parte basali minus late ochraceis quam

in femina.

Long., 9 10-11 mm., ¢ 8-10 mm.

2. Labrum and clypeus rather deeply but not very closely

punctured, the face much more closely and finely punctured,

face broader than long. Antenne 42-jointed, the third joint as

<p Slag go peer es a nL

ICHNEUMONID GENERA LABIUM AND PQG:CILOCRYPTUS. 9

long as the fourth and fifth combined, the fourth very little

longer than the fifth. Front and occiput microscopically punc-

tured, the space between the eyes and the ocelli almost smooth.

Thorax shining, mesonotum sparsely and finely. punctured on

the black fasciz, more closely anteriorly on the very prominent

median lobe ; notauli distinct, extending backwards beyond the

black lateral fasciz ; scutellum smooth and shining, with a few

small scattered punctures, postscutellum a little more closely

punctured. Median segment with the basal area smooth, short,

and separated from the areola only by lateral indications of a

carina; areola hexagonal, smooth, with a few scattered punctures,

narrowed posteriorly, the carina separating it from the dentiparal

area broadly obsolete behind the middle ; dentiparal area smooth

and shining; external area shining, microscopically punctured ;

spiracular area smooth, divided into three aree by two sharply

marked carine, both of which nearly touch the spiracle. Hind

Text-figure 4.

Hind tarsal ungues and empodia of Labiwm (a) montivagum Turn. & Wtrst.;

(b) bivittatwm Turn. & Wtrst.

coxe elongate, as long as the first tergite, nearly smooth dorsally,

but laterally and ventrally distinctly and rather closely punctured ;

hind femora closely and finely punctured. First transverse cubital

nervure strongly oblique, second abscissa of the radius very little

shorter than the second transverse cubital nervure, second

recurrent nervure separated from the second transverse cubital

nervure by a distance equal to about half the length of that ner-

vure. Ungues (text-fig. 4, >) of the hind tarsi much less robust

than is usual in the genus; hind empodium very small, not elon-

gate, only about one-third of the length of the unguis.

Hab. Yallingup, S.W. Australia; September 14—October 31,

1913 (Turner). A good series taken.

Very distinct from other species of the genus in the small

empodium, less robust hind tarsal ungues, prominent median lobe

of the mesonotum, and black markings. The antenne are also

less strongly thickened to the apex than in most species of the

genus. The groove between the face and clypeus is strongly

marked,

10 MESSRS. R. E. TURNER AND J. WATERSTON ON THE

LABIUM PILOsUM, sp. n. (Text-fig. 5.)

2. Ferruginea, fulvo-pilosa; antennis, scapo articuloque tertio

subtus flavis, occipite, macula circa ocello, propleuris, scutello

lateribus sulcoque basali, segmento mediano, femoribus posticis,

tibiisque posticis apice nigris; tarsis posticis infuscatis ; capite,

mesopleuris, scutello, postscutello, pedibusque flavo-ochraceis ; alis

sordide hyalinis, venis fuscis.

Long. 9 mm.

@. The whole insect, except the dorsal surface of the median

segment, sparsely clothed with pale fulvous hairs, which are

denser on the pleurz, legs, the sternites, and the vertex than else-

where. Labrum, clypeus, and face shining and sparsely punctured,

Text-figure 5.

Labium pilosum Turn. & Wtrst. In the front view the mandibles are partly

concealed by the labrum. In the profile the cheetotaxy of vertex and

antenna is not shown.

the clypeus quite smooth apically ; face (text-fig. 5) much broader

than long, the groove separating it from the clypeus obsolete

(2. e., clypeus and face in the same plane). Occiput more closely

and strongly punctured, front almost smooth. Antenne stout,

32-jointed, scarcely as long as the head, thorax, and median

segment combined, the third joint as long as the fourth and fifth

combined, the fourth distinctly longer than the fifth. Mesonotum

shining, finely punctured, rather closely on the median lobe, much

more sparsely posteriorly and laterally; the notauli very short, only

visible anteriorly, a very faint indication of a longitudinal carina

ICHNEUMONID GENERA LABIUM AND PCICILOCRYPTUS. 11

between the notauli. Scutellum and postscutellum smooth,

almost impunctate; mesopleure rather coarsely and not very

sparsely punctured on the anterior two-thirds, smoother pos-

teriorly ; vertical groove between the episternite and epimeron

crenulate on upper half, smooth ventrally. Basal area of median

segment broad and very short, the carina separating 1t from the

areola only indicated laterally, surface smooth; areola very

broad, not distinctly separated from the petiolar area, smooth,

with indications of transverse ruge posteriorly and with a few

scattered punctures; external area smooth, with a few indistinct

punctures ; dentiparal area coarsely, but rather sparsely, punc-

tured; spivacular area and juxta-cosal area with numerous

punctures, not quite as coarse as those on the dentiparal area ;

aree bounding the petiolar area rugose next to the bounding

carine. Petiole apically very broad, a little raised medially on

the basal half; all the tergites shining, with numerous minute

piliferous punctures ; second tergite fully twice as broad as long,

third tergite fully three times as broad as long. Areolet long,

second abscissa of the radius longer than the second transverse

cubital nervure; distance between the second recurrent and second

transverse cubital nervure equal to less than half the length of

the latter nervure.

Hab. 8. Australia.

This is very distinct in the pilosity of the whole insect, the

robust and broad abdomen, and the black and rather short

antenne.

LABIUM SUBPILOSULUM, Sp. n.

@. Fulvo-ferruginea ; antennis omnino, mandibulisque apice

nigris; capite, propleuris supra, tegulis, mesopleuris macula

magna sub alis anticis, scutello, postscutello, coxisque anticis

intermediisque flavis; tibiis posticis apice tarsisque posticis articulo

apicali infuscatis; alis hyalinis, iridescentibus, venis fuscis.

Long. 7 mm.

2. Mandibles simple; clypeus and face fused, without a

dividing-line, sparsely punctured, with fine hairs springing from

the punctures. Antenne 37-jointed, third joint distinctly shorter

than the fourth and fifth combined, the joints becoming trans-

verse about the twelfth. The emargination of the eyes is almost

obsolete; front and vertex smooth and shining. Mesonotum

shining, with a few sparse and very small punctures, the median

lobe rather more closely punctured, not prominent; notauli short

and shallow. Mesopleure anteriorly sparsely punctured on the

upper half, closely punctured rugulose on the lower half, smooth

posteriorly ; the epimeral sulcus crenulated on the upper half

only. Median segment with the apical carina of the basal

area broadly interrupted in the middle; areola and petiolar area

finely rugulose, not distinctly divided ; external area smooth and

shining; dentiparal are rugulose, without spines; spiracular

12 MESSRS. R. E. TURNER AND J. WATERSTON ON THE

and pleural arez finely punctured. Petiole with the basal teeth

blunt and not very prominent, spiracles only feebly prominent.

Hind cox and femora sparsely punctured. Apical tergites with

sparse hairs. Second abscissa of the radius very little shorter

than the second transverse cubital nervure, distance between the

recurrent and second cubital nervures not quite equal to half

the length of the latter nervure.

Hab. Victoria (C. French). 1 @.

In the fused condition of the face and clypeus this resembles

pilosum, but is much less pilose, very different in colour and in

the number of antennal joints. The antenne are very feebly

thickened to the apex, but are not as stout in this species as in

pilosum.

LABIUM SPINIFERUM, sp. n.

2. Fulvo-ferruginea; antennis 43-articulatis, articulis 20 basa-

libus fulvo-ferrugineis, 2i-34 infuscatis, 35-43 ochraceis ; labro

facieque marginibus, orbitis internis, genubus, mesonoto antice

lateribus angustissime, propleuris antice, coxis anticis, tegulis,

macula sub alis anticis, macula parva sub alis posticis, mesopleuris

macula magna mediana, macula magna ante coxas intermedias,

mesonoto angulis posticis anguste, scutello, postscutello, sulco

epimerali, segmentoque mediano fascia lata transversa apicali,

aveaque pleurali flavis; alis leviter infuscatis, venis fuscis.

Long. 10 mm.

Q. Clypeus and face rather strongly punctured, the face in

the middle rugulose longitudinally ; labrum very long, distinctly

longer than the clypeus, more finely punctured; mandibles

simple. yes distinctly, but shallowly emarginate a little above

the base of the antenne; third antennal joint fully as long as

the fourth and fifth combined, the eight basal joints of the funicle

cylindrical. Front and vertex finely punctured; mesonotum

shining, finely and evenly punctured, the median lobe promi-

nent ; notauli well developed, not very short; pleure shining,

almost smooth, the epimeral furrow crenulated on the upper

half; scutellum and postscutellum almost smooth. Median seg-

ment with the basal area not closed apically, the apical carina

being obsolete; areola almost smooth, distinctly divided from the

rugulose petiolar area; external and dentiparal ares finely and

sparsely punctured, each produced into a small tooth at the external

apical angle, that of the external area being stouter and less

acute than that of the dentiparal; spiracular area finely punctured-

rugulose; pleural area finely punctured, with larger punctures

intermingled. Spiracles of the petiole prominent, a very feebly

developed tooth on each side between the spiracles and the base.

Hind cox finely punctured above, closely punctured-rugulose on

the outer side and beneath. Second abscissa of the radius a little

shorter than the second transverse cubital nervure ; the distance

between the recurrent and second transverse cubital nervures

ICHNEUMONID GENERA LABIUM AND PQECILOCRYPTUS. 13

equal to slightly more than half the length of the latter

nervure.

Hab. Yallingup, 8.W. Australia ; October (Turner). 1 2.

Distinguished from other species by the minute, but distinct,

teeth on the median segment.

LABIUM VASSEANUM, sp. n. (Text-fig. 6, a, e.)

3. Fulvo-ferrugineus ; mandibulis, apice nigris, labro, clypeo

facie, scapo subtus, pronoto macula parva laterali, mesopleuris

macula magna, macula sub alis anticis, macula parva sub alis

posticis, macula ante coxas intermedias, scutello, postscutello,

segmento mediano macula magna apicali, areaque juxta-coxalli

flavis; antennis nigris, apice haud ferrugineis; alis subhyalinis,

venis fuscis.

Long. 9 mm.

Text-figure 6.

(a) Labium vasseanum Turn. & Wtrst. Head, profile.

(b) Labium hobartense ,, z Mandible.

(d) Labium subequale ,, x »

(e) Labiwn vasseanum ,, » »

3. Mandibles elongate, acute (text-fig.6, e) at the apex ; labrum

very long. Clypeus and face (text-fig. 6, w) shining, sparsely punc-

tured, ihe face longitudinally rugulose in the ane fe. Antenne

36- jointed, third joint more than half as long again as the fourth,

the fifth distinctly shorter than the fourth. eile conatiin shining,

very sparsely and finely punctured, the median lobe rather promi-

nent anteriorly and more closely punctured, notauli very short ;

mesopleure shining, with a few scattered punctures ; epimeral

sulcus crenulated on the upper half. Median segment with the

14 MESSRS. R. E. TURNER AND J. WATERSTON ON THE

apical carina of the basal area broadly interrupted in the middle ;

areola smooth, the carina dividing it from the petiolar area com-

plete; petiolar area smooth ; postero-intermedial area with a few

longitudinal striz ; external area with a few minute punctures ;

dentiparal area smooth in the middle, with a few punctures on

the sides and a minute tooth at the external apical angle; spira-

cular area finely punctured-rugulose; pleural and juxta- coxal

are not divided, smooth. Petiole narrow at the apex, without

teeth between the base and the spiracles. Hind empodia not very

long, not more than two-thirds of the length of the tarsal unguis.

Both transverse cubital nervures somewhat oblique and longer

than the second abscissa of the radius, the distance between the

recurrent nervure and the second transverse cubital nervure less

than half as great as the length of that nervure.

Hab. Yallingup, 8.W. Australia; October, 1913 (Turner).

Nearly allied to ZL. spiniferum, of which it may be the male,

but the colour of the antennz and the number of antennal joints

differ much, also the spines on the dentiparal and external aree

are much more distinct in spiniferum. ‘The elongate mandibles

are common to both.

LABIUM SCULPTURATUM, sp. n. (Text-fig. 7, a.)

3. Niger; mandibulis, labro, clypeo, facie, scapo subtus, orbitis,

pronoto lateribus anguste, linea supra tegulas, mesopleuris macula

magna mediana, fascia sub alis anticis, sulco epimerali, scutello,

postscutello, segmento mediano fascia apicali, petiolo fascia api-

cali, pedibusque anticis intermediisque flavis; tergitis secundo

sequentibusque, tibiisque trochanteribusque posticis rufo-ferru-

gineis ; antennis supra nigris, subtus rufescentibus, articulis

13 apicalibus ferrugineis ; alis subhyalinis, venis nigris.

Long. 11 mm.

Text-figure 7.

Aveolet (right wing):—(a) Labium sculpturatum Turn. & Wtrst. (b) Labium

Serrugineum Cam. (c) Labiwm petitorium Erichs.

3. Face much broader than long, face and clypeus sparsely

but not very finely punctured. Antenne 42-jointed, third joint

more than half as long again as the fourth, the fifth distinctly

shorter than the fourth, the joints as far as the sixteenth longer

than broad, those beyond transverse. Mesonotum with the middle

lobe prominent; notauli short, but deep and crenulate; the

median lobe densely punctured anteriorly, sparsely posteriorly ;

ICHNEUMONID GENERA LABIUM AND PG@CILOCRYPTUS. 15

lateral lobes and the scutellum sparsely punctured ; mesopleure

antero-ventrally closely and rather strongly punctured, smoother

elsewhere; the epimeral sulcus coarsely crenulate throughout.

Median segment with the basal area and areola confluent; areola

smooth, with a few punctures, posteriorly rugulose ; petiolar area

separated, transversely rugose, as also are the adjacent arew; ex-

ternal area smooth, very bluntly raised at the external apical

angle; dentiparal area coarsely rugulose, with a ‘small tooth at the

external apical angle; spiracular area coarsely punctured ante-

riorly, rugose posteriorly ; pleural and juxta-coxal aree strongly

striate, with a few punctures between the striae. Hind coxe

externally coarsely, hind femora much more closely and finely

punctured. Second abscissa (text-fig. 7, a) of the radius barely

longer than the second transverse cubital nervure; the distance

between the recurrent nervure and the second transverse cubital

nervure more than half of the length of the latter nervure.

Petiole without a tooth between the base and spiracles, the latter

very prominent; a carina beginning near the base and almost

reaching the apex, the central raised area strongly marked,

bearing throughout its length two parallel grooves.

Hab. Yallingup, S.W. Australia; November, 1913 (Zurner).

IG.

Distinct in the strong sculpture, also in the black colour of

the thorax, petiole, and hind femora,

LABIUM SUBEQUALE, sp. n. (Text-fig. 6, 2.)

2. Fulvo-ferruginea ; antennis nigris, articulis 10 apicalibus

ochraceis ; scapo subtus, mandibulis. labro, clypeo, facie, fronte

lateribus, orbitis externis, genis, scutello, macula sub alis anticis,

coxis anticis intermediisque, tibiis intermediis apice, tarsisque

anticis intermediisque, articulo apicali excepto, flavis; pronoto

antice, tegulis, area Juxta-coxall, tarsisque posticis articulo apicali

nigrescentibus; alis subhyalinis, venis fuscis.

Long. 9 mm.

2. Mandibles (text-fig.6, Z) obscurely bidentate, blunt, the second

tooth indicated by an external sulcus; elypeus and face sparsely

punctured, the line of division between them distinct, but not

strong. Antenne 36-jointed, the third and fourth joints sub-

equal, the third seen in profile slightly shorter than the fourth.

Mesonotum smooth, with scattered punctures, the notauli rather

long; propleure strongly crenulate in front ; mesopleure shining,

closely and very finely punctured on the lower half; epimeral

sulcus crenulate throughout, the lower crenulations more strongly

developed than usual. Median segment with the apical carina

of the basal area interrupted in the middle; areola smooth,

slightly rugulose at the sides; external area smooth ; dentiparal

area rugulose; spiracular area coarsely punctured anteriorly,

rugulose posteriorly ; pleural area striate-rugulose, separated from

the juxta-coxal area by a carina reaching to one-half, the latter

16 MESSRS. R. E, TURNER AND J. WATERSTON ON THE

area rugose. Petiole with the lateral teeth between the spiracles

and the base well developed ; spiracles prominent, obscurely striate

between the spiracles, the remainder of the segment smooth and

convex ; before the spiracles the raised area is distinct. Hind

coxe closely and rather finely punctured, femora very finely

punctured. Second abscissa of the radius about equal to the

second transverse cubital nervure, the latter fully twice as ‘long

as the distance between it and the recurrent nervure.

Hab. Yallingup, 8.W. Australia; October and November,

1913 (Turner).

LABIUM OCCIDENTAL, sp. n. (Text-fig. 6, ¢.)

Q. Fulvo-ferruginea; antennis infra obscure ferrugineis, supra

nigrescentibus, articulis 13 apicalibus rufo- _ferrugineis; scapo

subtus, mandibulis, apice nigris, labro, clypeo, facie, orbitis

externis, scutello, postscutello, macula ante tegulas, mesopleuris

maculis duabus, macula sub alis posticis, macula supra coxas

intermedias, segmento mediano fascia tramsversa obscura post

medium, coxisque trochanteribusque anticis intermediisque flavis;

fronte in medio, propleuris antice, mesopleuris antice et inter

maculas flavas, scutello postscutelloque Jateribus, petiole dimidio

basali, area juxta-coxali, tibiis posticis apice supra, tarsisque

posticis articulo apicali nigrescentibus; alis subhyalinis.

Long. 10 mm.

3g. Differt occipite, temporibus, mesopleuris, segmento mediano,

notaulis, mesoncto lateribus, petiolo, apice excepto, coxis posticis

basi et apice supra, femeribus tibiisque posticis apice, nigrescen-

tibus; tarsis posticis infuscatis; pedibus anticis intermediisque

omnino flavis.

2. Differs from swhequale in the strongly bidentate (text-

fig. 6 c) mandibles, the upper tooth much shorter than the lower,

in the much more strongly punctured face, in the generally

stronger puncturation, and in the sculpture of the hind coxe and

femora, which are Shimon with large punctures in occidentale,

whereas in subequale the hind femora are subopaque and very

closely covered with minute punctures.

Hab. Yallingup, S.W. Australia; October (Z'urner).

LABIUM MONTIVAGUM, sp. n. (Text-figs. 1, 4a, 8, 9, and 11a.)

3. Niger; mandibulis, apice excepto, labro, clypeo, facie,

orbitis anguste, callis humeralibus, linea sub alis anticis, meso-

pleuris macula parva mediana, macula parva sub alis posticis,

scutello, postscutello macula mediana, pedibusque anticis inter-

mediisque flavis; segmentis abdominalibus secundo Sequenti-

busque, femoribus posticis, apice nigris, tibiis posticis, apice

nigris, tarsisque posticis articulis quatuor basalibus rufo-ferrugi-

neis ; antennis subtus (articulis tribus basalibus exceptis) arti-

culisque 11 apicalibus supra, oehraceis; alis hyalinis, venis fuscis.

ICHNEUMONID GENERA LABIUM AND PG:CILOCRYPTUS. ii%

@. Differt scapo subtus flavo; thorace ferrugineo, flavo-

maculato, propleuris infra, mesopleuris antice, scutelloque sulco

basali nigris; segmento mediano, COXIS posticis, femoribusque

posticis ferrugineis, apice nigro-maculatis; pedibus anticis

intermediisque fulvo- ferrugineis.

Long. 2 ¢ 9-10 mm.

Text-figure 8.

Wings :—(a) Pecilocryptus nigromaculatus Cam. (b) Labiwm montivagum

Turn. & Wtrst.

@. Labrum, clypeus, and face rather sparsely, but not finely

punctured ; the dividing line between the face and clypeus rather

indistinct, the face broader than long; mandibles simple.

Antenne from 32- to 36-jointed, usually 35 or 36; third joint

equal to the two following. Front shining, sparsely and finely

punctured at the sides, rather more strongly below the ocelli,

occiput finely and closely punctured. Thorax shining; the meso-

notum sparsely punctured, more closely on the median lobe:

notauli very distinct, but short, more or less crenulate; scutellum

and postscutellum smooth and impunctate; propleur anteriorly

rugulose, posteriorly crenulate, centrally slightly raised and

rather sparsely punctured ; mesopleure ventrally rather closely

Proc. Zoou. Soc.—1920, No. II 2

18 MESSRS. R. E. TURNER AND J. WATERSTON ON THE

and not finely punctured, posteriorly smooth, the smooth area

narrowing ventrally ; epimeral suleus entirely crenulate, with

one or two large fover at its ventral extremity. Sternum shining,

moderately closely punctured, one or two large fovez outside the

intermediate coxe close to the extremity of the epimeral sulcus.

Median segment coarsely sculptured; apical carina of the basal

area broadly interrupted in the middle, the basal area smooth

and shining ; areola and petiolar area fully separated, the former

smooth, with one or two large punctures and rugulose towards

the sides, especially posteriorly, petiolar area transversely rugu-

lose; external areze nearly smooth, dentiparal and other dorsal

aree rugulose, except the anterior portion of the spiracular area,

which is coarsely punctured ; pleural (text-fig. 1) and juxta-coxal

are rugulose above, and towards the coxee with strongly marked

Text-figure 9.

TERZI 3 b

Labium montivagum Turn. & Wtrst. ¢ genitalia. (a) Stipes and in profile.

(6) Entire apparatus—right stipes removed—dorsal view. (ce) Apex of

volsella.

ridges, so that anteriorly this area is crenulate. Spiracles of the

petiole just behind the middle, much nearer to each other than

to the apex of the segment, a well-defined tooth (text-fig. 11, @) on

each side of the petiole, nearer to the base than to the spiracle ;

the ceutral dorsal portion of the petiole basally distinctly raised,

with a sulcus near each spiracle; gastrocceli distinct, finely punc-

tured, abdomen otherwise smooth. Second abscissa of the radius

a little shorter than the second transverse cubital nervure,

distance between the recurrent nervure and the second transverse

cubital nervure more than half as great as the length of the latter

nervure. Externally the hind coxe and femora are sparsely but

rather deeply punctured. Tarsal unguis etc., text-fig. 4, a.

Hab. Mt. Wellington, Tasmania; January and February, 1913

(Turner). A long series.

ICHNEUMONID GENERA LABIUM AND PCSCILOCRYPTUS. 19

The male has the face almost square, the sculpture is somewhat

coarser, especially on the median segment. The sculpture of the

areola in the male shows considerable variation; 1n some speci-

mens there are coarse transverse ruge, which are only rather

feebly indicated in others. The teeth on the sides of the petiole

are remarkable, also the sexual colour differences.

The female is the type.

LABIUM CLAVICORNE Morl.

Labiwm clavicorne Mor]. Revis. Ichneum. iv. p. 151 (1915).

This species is allied to montivagwm in having a distinct tooth

on each side of the petiole between the spiracles and the base

and also in the sculpture, but differs in the almost entirely

fulvous antenne, in the ferruginous colour of the front, vertex

and propleure, and in the position of the second recurrent ner-

vure which is received close to the apex of the areolet.

This is very doubtfully distinct from ferruginewm Cam., but

differs in the colour of the antenne. The other differences given

Text-figure 10.

Labium ferrugineum Cam. Propodeon ; dorsal view.

by Morley either fall within the range of individual variation or

ave taken from one or other of the three males marked by him

as co-types, one of which is certainly specifically distinct, not at

all nearly related to the other two, in which the thorax is mostly

black, having only the mesonotum and a patch on the meso-

pleure ferruginous. These are probably the true males of

clavicorne, which species probably takes the place of montivayum

on the mainland.

Hab. Victoria (Wrench).

LABIUM FERRUGINEUM Cam. (Text-figs. 7 6 and 10.)

Labium ferruginewm Cam. Ann. & Mag. Nat. Hist. (7) vii.

paos0i(l9Ol)\s 59.

This species seems to differ from elavicorne Morl. only in the

eolour of the antenne, which are black above and brownish on

the apical half beneath; the scape is yellow beneath in both

forms. Cameron’s type is in a dirty condition.

20 MESSRS. R. E. TURNER AND J. WATERSTON ON THE

The locality given is Australia,

The nervulus i is distinctly postfurcal, not differing appreciably

from clavicorne in this respect, in spite of Morley’s statement to

the contrary.

LABIUM HOBARTENSE, sp.n. (Text-fig. 6, b.)

2. Fulvo-ferruginea ; antennis supra nigro infuscatis, articulis

13 apicalibus ferrugineis; mandibulis dimidio basali, labro,

clypeo, facie, fronte lateribus, coxis trochanteribusque anticis

intermediisque, maculis duabus sub alis anticis, macula sub alis

posticis, macula supra coxas intermedias, sulco epimeral, scutello,

postscutello, segmento mediano fascia transversa post medium

coxam posticam attingente, tergitisque fascia apicali angustissima.

flavis; petiolo, area juxta-coxali, tarsisque posticis articulo

apicali nigrescentibus ; alis subhyalinis, venis nigris.

Long. 11 mm.

@. Mandibles bidentate (text-fig. 6, 6), rather short; face

shining, sparsely and finely punctured, broader than long. An-

tennee 42-jointed, third joint a little shorter than the fourth and

fifth combined.

Very similar to montivagum in other respects ; differing in the

yellow band on the median segment, in the yellow apical bands

of the tergites, and the less prominent basal teeth of the petiole.

Hab. Hobart, Tasmania; summer 1891 (/. J. Walker).

Might easily be mistaken for montivagum, but the structural

differences in the mandibles and antenne distinguish it at once.

LABIUM ASSOCIATUM, sp. n.

@. Fulvo-ferruginea ; mandibulis, apice nigris, labro, clypeo,

facie, scapo subtus, orbitis, scutello, macula sub alis anticis,

mesopleuris antice macula, coxisque trochanteribysque anticis

intermediisque flavis; scutello sulco basali, tibiis posticis apice

supra, tarsisque posticis, basi anguste ferrugineis, nigris; alis

hyalinis. iridescentibus, venis nigris.

Long. 8 mm.

2. Clypeus and face shining, very sparsely punctured, face

much broader than long ; inandibles simple. Antenne 36-jointed,

fulvous, somewhat infuscate above to beyond the middle, the

third jomt about ‘half as long again as the fourth, the fifth

distinctly shorter than the fourth. Eyes very feebly sinuate

opposite the base of the antenne; front and vertex shining,

almost smooth, witha few minute punctures, Mesonotum shining,

almost smooth, the median lobe sparsely punctured anteriorly ;

notauli, rather short ; .mesopleure smooth on the upper half,

sparsely and finely punctured on the lower half; epimeral sulcus

finely crenulate throughout. Median segment with the apical

carina of the basal area broadly inter rupted i in the middle; areola

not separated from the petiolar area, the dividing carina only

feebly indicated at the sides, areola georqoulh § in anes middle, finely

ICHNEUMONID QENERA LABIUM AND PGCILOCRYPTUS. 21

rugulose at the sides, petiolar area with the same sculpture as

the areola; the are adjacent to the petiolar area rugulose ;.

external area smooth, dentiparal area subrugulose, a minute

tooth at the external apical angle of both the external and

dentiparal ares; spiracular area strongly rugulose, anteriorly

punctured-rugulose ; pleural area superiorly rugulose with a few

deep punctures, ventrally with three or four large complete ruge,

which merge with thosé of the juxta-coxal area, the carina

dividing the two are well defined to about one-half. Petiole

with a well-developed tooth on each side between the base and

the spiracles; the petiole with a rather stronger sculpture than is

usual in the genus, behind the spiracles the surface in the middle

subrugulose, before the spiracles the raised median area is very

distinct. Hind coxe and femora smooth on the outside, with

large irregular punctures. Second abscissa of the radius a little

shorter than the distance between the recurrent nervure and the

second transverse cubital nervure, and only a little more than

half as long as the latter nervure.

Hab. Mundaring Weir, W. Australia; March 18, 1914

(Turner).

LABIUM APPROXIMATUM, Sp. 0.

3. Fulvo-ferrugineus ; mandibulis, apice nigris, labro, clypeo,

facie, froute lateribus, scapo subtus, scutello, postscutello, meso-

pleuris maculis duabus, macula sub alis posticis, macula supra

coxas intermedias, segmento mediano fascia. dorsali transversa

post medium, macula supra coxas posticas, coxisque trochanteri-

busque anticis intermediisque flavis; antennis ferrugineis, supra

nigris, articulis 15 apicalibus fulvo-ochraceis; coxis posticis apice

supra, mesopleuris macula post coxas anticas, area juxta-coxali,

petioloque ante spiracula infuseatis.

Long. 12 mm. :

3. Mandibles simple; clypeus and face closely, evenly, and

rather strongly punctured, the line between the face and clypeus

very distinct ; malar space short, only half as broad as the hase

of the mandible. Front smooth, with sparse punctures. Antenne

47-jointed, third joint almost as long as the fourth and fifth

combined, the joints becoming transverse at about the 22nd or

23rd. Mesonotum shining, finely and rather closely punctured,

the median lobe prominent aud more closely punctured; notauli

long. Scutellum and postscutellum shining, finely punctured ;

propleurz rather strongly punctured; mesopleure punctured on

the lower half, the epimeral sulcus strongly crenulate. Median

segment with the apical carina of the basal segment broadly

interrupted in the middle; areola shining, with a few lateral

punctures, its apical keel well defined ; external area finely and

closely punctured ; dentiparal area rugulose, both the external

and dentiparal arez with a small tooth at the external apical

angle, and a minute tooth also at the inner apical angle of the

dentiparal area; spiracular area anteriorly finely and closely

22 MESSRS. R. E. TURNER AND J. WATERSTON ON THE

punctured, posteriorly rugulose; pleural area rugulose-punctate ;

juxta-coxal area with six or seven strong striz. Basal tooth of

petiole blunt, not prominent, spiracles prominent; basal area

of petiole indistinct. Hind cox and femora shining, externally

closely and finely punctured. Second abscissa of the radius con-

siderably longer than the second transverse cubital nervure ;

distance between the recurrent nervure and the second transverse

cubital nervure equal to more than half the length of the latter

nervure.

Hab. Victoria (C. French). 1 3.

This is one of the males selected by Morley as a co-type of his

LL. clavicorne, to which it is not at all nearly allied.

LABIUM FULYICORNE, sp. n.

2. Fulvo-ferruginea; antennis articulis apicalibus fulvo-ochra-

ceis; mandibulis, apice excepto, labro, clypeo, facie lateribus,

mesonoto margine laterali anguste, scutello, postscutello, propleuris

infra, macula horizontali sub alis anticis ; suleoque epimerali flavis ;

tarsis posticis nigris, metatarso apice solum nigro; alis sordide

hyalinis, venis nigris, stigmate fusco-ferrugineo.

Long. 10-12 mm,

Q. Clypeus and labrum sparsely, face more closely punctured ;

clypeus not on the same plane with the face, divided from it by a

distinct groove, the face broader than long ; front smooth, occiput

with a few small punctures. Antenne 46-jointed, more than

three-quarters of the length of the whole insect ; third joint fully

as long as the fourth and fifth combined. Thorax shining and

almost smooth, the median lobe of the mesonotum alone distinctly

punctured ; notauli short, only distinct anteriorly. Hpimerat

groove very finely crenulated above, smooth below. Arez of the

median segment smooth and shining, the external, pleural, and

spiracular ares very finely punctured; petiolar area with a few

scattered punctures ; basal area very short, the carina separating

it from the areola narrowly broken in the middle, the areola

completely divided from the petiolar and dentiparal aree. Hind

coxe shining, very sparsely punctured, reaching as far as the apex

of the petiole. Second abscissa of the radius longer than the

second and nearly as long as the first transverse cubital nervure,

the distance between the second recurrent nervure and the second

transverse cubital nervure a little less than half as great as the

length of the latter nervure.

Hab. Yallingup, S.W. Australia; October and November, 1913

(Turner).

In this and some closely-allied species the external area of the

median segment is more distinctly sculptured than the dentiparal

area, the reverse being the usual condition in the genus. The

probable male of this species has the pleure and fore legs

almost entirely yellow, also a broad transverse band at the apex

of the median segment and the pleural aree. The number of

joints in the antennz is 41-44. . These males are much smaller

ICHNEUMONID GENERA LABIUM AND PGICILOCRYPTUS. 23

than the female, measuring from seyen to nine millimetres, and

the hind tarsi are not black.

LABIUM MULTIARTICULATUM, sp. n.

Q. Fulvo-ferruginea ; flagello supra fusco, articulis 13 api-

ealibus ochraceis, infra ochraceo ; scapo subtus flavo ; mandibulis,

labro, elypeo, facie, propleuris antice, fascia sub alis, mesopleuris

fascia lata maculaque arcuata supra coxas intermedias, scutello,

postscutello, mesonoto fascia apicali areaque pleurali, coxisque

anticis intermediisque flavis; alis subhyalinis, venis fuscis, stig-

mate fusco-ferrugineo.

¢. Femine similis ; flagello articulis 10 apicalibus solum supra

ochraceis ; mesonoto lateribus flavo-marginatis, mesopleuris fere

omnino flavis, trochanteribusque anticis intermediisque flavis.

Long. 9° 3 10-11 mm.

@. Labrum long, rather narrowly rounded at the apex; face

distinctly broader iiten long, face and clypeus shining, sparsely,

but rather deeply punctured. Antenne usually from 51-. to

53-jointed (51 in the type 2 ), sometimes with one or two joints

more or less; third joint a little shorter than the fourth and fifth

combined ; front and vertex shining, almost smooth. Mesonotum

shining, finely punctured, rather closely on the median, very

sparsely on the lateral lobes, the median lobe not prominent,

notauli short ; mesopleure shining, sparsely and finely punctured.

more closely on the lower than on the upper half; epimeral sulcus

crenulate on the upper half only, the striz oblique. Median

segment with the apical carina of the basal area obsolete, except

at the sides; areola shining, sparsely and finely punctured, the

basal carina of the petiolar area well defined ; external arew finely

punctured, dentiparal areze smooth and shining ; ; Spiracular area

punctured ; pleural area very sparsely punctured, not separated

from the smooth juxta-coxal area; petiolar and adjacent arez

punctured. Petiole without any clearly-defined lateral teeth

between the spiracles and the base, spiracles not prominent.

Second abscissa of the radius very slightly longer than the second

transverse cubital nervure ; the distance between the recurrent

and the second transverse cubital nervure equal to about half the

length of the latter nervure.

Hab. Yallingup, 8.W. Australia; October and November, 1913

(Turner).

Near jfulvicorne, but may be distinguished by the greater

number of antennal joints. The third antennal joint is distinctly

shorter than in fulvicorne and the antenne less clavate. The

male of the present species is generally larger and more robust

than that of fulvicorne.

LABIUM LONGICORNE, sp. 0.

36. Fulvo-ferrugineus ; mandibulis, apice nigris, labro, clypeo,

facie, scapo subtus, scutello, postscutello, propleuris antice, pro-

sterno, mesopleuris maculis duabus sub alis anticis, suleo epimerali,

24 MESSRS, R. E. TURNER AND J. WATERSTON ON THE

coxisque trochanteribusque anticis flavis; antennis ferrugineis,

seapo pedicelloque supra leniter infuscatis; alis hyalinis, venis

fuscis.

Long. 9 mm.; antennarum long. 7 mm,

3. Very near LZ. multiarticulatwm, but differs in the colour of

the antenne, in the absence of a yellow apical band on the median

segment, in the distinctly shorter third antennal joint (in muléi-

articulatum this joint is twice as long as broad, in the present

species rather less than half as long again as broad), and in the

smooth external area of the median segment. The antennse

are 49-jointed.

Hab. Swan River, W. Australia (Du Boulay).

Lasium sicotor Brulleé.

_ Labium bicolor Brullé, Hist. Nat. Insect. Hymén. iv. p. 316

(1846). o.

Hab. New Guinea.

This species, the type of the genus, is unknown to us.

Genus Pacrtocryprus Cam.

Pecilocryptus Cam. Ann. & Mag. Nat. Hist. (7) vii. p.527 (1901).

Cameron’s description of this genus is good, and he is probably

correct in connecting the genus with the Phygadeuonini, though,

as he remarks, it has no near relationship with any other genus.

Cameron only gives Australia as the locality for his species,

which, however, we can now record from Tasmania, together

with a new species described below.

PacrLocryPrus NIGRIPECTUS, Sp. nN.

2. Flava; mandibulis apice extremo, antennis, articulis sex

basalibus subtus flavis, fronte supra antennas, area circa ocellos,

occipite, mesonoto fascia longitudinali lata antice, usque ad

medium attingente, lateribus et margine postico late, mesosterno,

scutello basi, segmento mediano basi, lateribus late, supra augus-

tissime, tergito tertio, apice extremo excepto, tergito sexto,

valvulis terebre, femoribus posticis macula magna extus intusque,

tibiisque posticis apice nigris ; alis hyalinis, iridescentibus, venis

fuscis.

Long. 5°5 mm.; terebre long. 2°5 mm.

@. Antennz 22-jointed, the basal joints very slender, the

apical joints strongly thickened, the last seven broader than

long, the five basal joints of the flagellum long and slender, joints

6 to 9 slightly thickened, from the tenth more abruptly

thickened, attaining the greatest breadth at the fifteenth. Face

longer than broad by at least one-third, with a short narrow

suleus, which is medially enlarged and extends downwards to the

middle of the face. Clypeus and face smooth and shining. Eyes

ee ee ee eee

ICHNEUMONID GENERA LABIUM AND PC@CILOCRYPTUS.

Text-figure 11.

(a) Labiwm montivagum Turn. & Wtrst. (b) Pecilocryptus nigro-

maculatus Cam.

26 THE ICHNEUMONID GENERA LABIUM AND PC@CILOCPYPTUS.

separated from the base of the mandibles by less than the breadth

of the mandible. Front concave, the concave area‘extending as

far back as the anterior ocellus, vertex smooth and shining.

Thorax smooth and shining, notauli indistinct. Median segment

very feebly punctured, rather dull, spiracles very small, oval ;

basal area very small, transverse; areola quadrate, a_ little

longer than broad, longer than the petiolar area; dentiparal area

subquadrate and about as large as the areola: external area

transverse; spiracular area divided by a keel arising from the

side of the basal area just before the apex, and continued out-

ward, touching the spiracle posteriorly ; of the two arez so formed

the anterior one (containing the spiracle) is pentagonal, the

posterior one is quadrate, about a third longer than broad ; both

the postero-intermedial and the postero-external are appear to

be defined ; the confluent pleural and juxta-coxal ares form a

hexagon, Spiracles of the petiole before the middle; the base

of the petiole with a raised median space bounded by definite

edges, and extending backwards to the highest part of the petiole ;

second tergite with the gastroceeli distinct, the segment rather

longer than wide; sutures 2-5 rather deepand distinct. Surface

of the abdomen dull to the middle of the second tergite, shining

apically. Hind cox and femora distinctly intumescent. Areolet

pentagonal, longer than high, the radial side equal to the inner

and almost equal to the outer, but shorter than the two lower

sides; second recurrent nervure strongly oblique.

Hab. Mt. Wellington, Tasmania, 2300 ft., March 12-21, 1913

(Turner).

The areolet is much shorter on the radius than in mgro-

maculatus Cam., the face 1s much narrower and differently sculp-

tured; there is no tooth at the apex of the dentiparal area, the

basal area is much shorter, and in-nigromaculatus the carina

between the spiracular and pleural are is anteriorly obsolete ;

in nigripectus the spiracles of the petiole are nearer to the base

than in nigromaculatus.

P@cILOCRYPTUS NIGRoMACULATUS Cam. (Text-figs. 8@ and

115.)

Pecilocryptus nigromaculatus Cam. Ann. & Mag. Nat. Hist.

(7) vii. p. 528 (1901). @.

The locality given for the type is merely Australia. Two

females in the British Museum are from 8. Tasmania, Mt. Wel-

lington, January 15—February 6, 1913 (Zurner), and Haglehawk

Neck, February 12—March 3, 1913 (Turner).

Unless indication to the contrary is given the types of the

species described in this paper are females, and all are deposited

in the British Museum.

ON THE LARYNX AND G&SOPHAGUS OF A COMMON MACAQUE. 27

2. Deseription of the Larynx and Cisophagus of a Common

Macaque (Macacus fascicularis) exhibiting several Ab-

normal Characters. By C. F. Sonntac, M.D., Ch.B.,

F.Z.S., Anatomist to the Society.

"Received January 10, 1920; Read February 10, 1920. |

(Text-figures 1-5.)

The specimen which forms the subject of the present com-

munication was preserved among a series of tongues in the

Society’s Prosectorium. When a disseetion was made, the rare

conditions shown in text-fig. 2 were revealed, and the normal

state is illustrated in text-fig. 1 for comparison. One of the

Text-figure 1. Text-figure 2.

Text-fig. 1—The normal tongue, larynx, and cwsophagus. A. Ridge of mucous

membrane. C. Posterior surface of larynx clothed with mucous membrane.

E. Recess.

Text-fig. 2.—The abnormal tongue, larynx, and esophagus. B. Posterior car-

tilaginous plate. A, C, Eas in text-fig. 1.

28 DR. C. F. SONNTAG ON THE LARYNX AND

conditions is unique, and raises problems for solution by the

embryologist and physiologist. The soft palate and the posterior

wall of the pharynx were divided in a vertical direction, but the

posterior wall of the cesophagus was only divided from behind

forwards and above downwards till the mucous membrane

appeared. I did not divide any more, as I considered that it

was better to preserve the abnormality of the cesophagus entire.

In the normal state, however, the posterior wall of the cesophagus

was divided completely and the two halves thrown outwards.

By that means an uninterrupted view of the tongue, larynx,

pharynx, and cesophagus is obtained.

In the normal state the epiglottis has a notched border and a

median fissure running vertically down the inner surface, so that

the cartilage appears to be composed of two symmetrical halves.

It slants upwards and forwards, and the upper aperture of the

larynx is small. The aryteno-epiglottidean folds, with their

contained cartilages of Wrisberg, are small, and appear globular.

The posterior surface of the larynx is smooth.

In the abnormal specimen the epiglottic border is devoid of a

notch, and the cartilage stands up vertically, so that very little

of its imner surface is visible. There is no vertical median

fissure on that surface, so that it appears like an arc of a circle.

The aperture of the larynx is capacious, and the aryteno-epiglottic

folds with their cartilages are oval; they are more prominent

than in the normal condition. Indeed, the whole larynx appears

larger. Its postero-lateral aspects exhibit a number of ridges

and sulei running horizontally round the upper part and ter-

minating on a strong vertical median ridge.

The cesophagus of the normal specimen appears as a wide space

behind the larynx. Its walls are ridged vertically, and an unin-

terrupted view of the back of the larynx, invested by transparent

mucous membrane, is obtained.

In the abnormal specimen a broad cartilaginous plate covers

up the whole of the larynx below the pitcher-beak produced

by the arytenoid cartilages. It in turn is, with the exception of

its upper end, concealed | by a vertical partition of mucous mem-

brane. It measures 1°5 em. long, 1 mm. thick, and *75 em. in

vertical height at its mid point, but it tapers towards its extre-

ities. It is convex backwards on its outer surface, and concave

backwards on its inner one. Its left extremity runs into the

end of a plate on the posterior surface of the larynx, but its right

end fuses with the posterior surface of the larynx beyond the

right extremity of the latter plate. As the anterior plate

is convex backwards, the two plates fit into one another, and the

orifice of the cesophagus is curved (text-fig. 3); it is tightly

closed. The anterior plate is not so high as the posterior one,

but extends further down the ceesophagus, and is firmly adherent

to the cricoid cartilage.

In text-fig. 5 a sagittal section has been made of the normal

neck, and the simplicity of the structures can be seen. It

oe ae ee

(ESOPHAGUS OF A COMMON MACAQUE. 248)

contrasts greatly with the abnormal state which is depicted in

text-fig. 4. In the latter one sees the mucous membrane (A)

reflected on to the posterior cartilaginous plate (B). Then comes

the cesophageal opening. Before it are the anterior cartila-

ginous plate (D), a small groove, and the posterior surface of the

larynx (C).

The folds of mucous membrane (A) merit attention. In the

normal condition a vertical ridge runs down the lateral walls of

the pharynx and cesophagus, beginning at the posterior faucial

pillar at the level of the upper border of the tonsillar pit, and

ending at the level of the middle of the larynx. At the level of

the most expanded part of the larynx, several horizontal folds

run round the entire circumference of the pharynx and ceso-

phagus. The vertical ridge, the posterior pillars of the iauces,

and the side of the larynx enclose a small triangular recess.

Text-figure 3. Text-figure 4. Text-figure 5.

2. 2 (oF

OPENING

OESOPHAGEAL

OE£SOPHAGEAL

OPENING

Text-fig. 3—The cesophageal opening. Text-fig. 4.—Sagittal section of

abnormal specimen. ‘Text-fig. 5.—Sagittal section of normal specimen.

In these D is the anterior cartilaginous plate, and A, B, C are the same

as in text-figs. 1 and 2.

In the abnormal specimen a ridge runs horizontally round the

cesophagus, beginning at the angles of the epiglottis, and it

bounds a deep recess on each side of the larynx. The mucous

membrane is reflected on to the posterior cartilaginous plate

(text-fig. 4), thereby forming a small pocket. From the ridge

horizontal folds turn out and run round the cesophagus.

The special points which arise from a consideration of the

above abnormality are :—

1. The origin of the cartilaginous ring.

2. The manner in which it affects the normal mechanism of

deglutition.

The great rarity of this condition, and the absence of specimens

showing gradations between the normal and the cartilaginous

ring described above, make all opinions as to its nature purely

hypothetical. My own opinion is that the development of the

cricoid cartilage has gone wrong, and the halves, or one of them,

which fuse to close in the posterior part of the laryngeal cavity,

have or has sent processes backwards; these have joined behind

to form the catilaginous plate (text-fig. 2, B), or one has grown

round and rejoined the cartilage again. The anterior plate

text-fig. 2, D) is immovably fixed to the back of the cricoid

30 ON THE LARYNX AND GSOPHAGUS OF A COMMON MACAQUE.

cartilage, and the entire ring is internal to the mucous lining of

the csophagus. The structure round the cesophagus seems to

have broken into it from the cricoid cartilage.

If the tongue and larynx are moved by the observer as they

do naturally when the animal swallows, the posterior part of the

larynx comes up against the posterior plate (text-fig. 2, B) and

obstructs the cesophagus. Of course, that probably does not occur

in life. Again, the semilunar mouth of the esophagus is firmly

closed by the cartilaginous plates, and one gets the appearance

of a sphincter. ‘The specimen had been preserved too long to

permit of a dissection of muscles which might move the posterior

plate, so I must leave the problem of how this animal swallowed

unsolved.

Both the specimens described in this communication are now

- preserved in the Museum of the Royal College of Surgeons,

London.

Py oZ SO ZO eh OGGra reese

ZEA

AUSTRALIAN OPILIONES.

Pl. Ul.

HOGG,

1920,

PZ. 8:

VV AAS

UV7-Y

Bale & Danielsson, Ltd.

AUSTRALIAN OPILIONES.

“SaNOMIdO NVINvaLsny

‘P}] ‘uossjaiueg g ajeg

Hi id SOO OCGleS. 27nd

ON SOME AUSTRALIAN OPILIONES. 31

3. Some Australian Opiliones.

By H: R. Hoec, M.A. EZ.

[ Received December 27, 1919: Read February 10, 1920. |

(Plates I.-IIL.*)

I am indebted to Dr. Charles Chilton of Canterbury College,

Christchurch, New Zealand, for a large collection of Spiders and

their allied orders, gathered over a seriés of years from most

parts of New Zealand. Amongst them are a number of Opiliones,

and these with a few others I am now recording.

Of the three suborders Palpatores, Laniatores, and Cypho-

phthalmi, the two former only are represented—Palpatores by the

genera Macropsalis Sor. and Pantopsalis Sim. of the family

Phalangiide Thorell, and Laniatores by the families Trisno-

nychidee Sor. and Trienobunide Pocock.

Only three species of the genus Macropsalis have been recorded—

the type J. serritarsis Sor.r from Sydney, N.S. Wales; one

collected by myself at Macedon, Victoria, described under my

name by Mr. R. I. Pocockt; and Jf. chiltoni$ from Stewart

Island, N.Z., described by myself.

The New Zealand specimens of this genus sent to me at

various times have all come from Stewart Island, the southern-

most remnant of the now broken land, and none from the Main

Islands, whereas Pantopsalis is widely distributed over both the

North and South Islands. d/acropsalis was generally supposed to

be an Australian form, and as Stewart Island lies in the belt of

westerly winds which blow all the year round, and the genus has

not been recorded from any other part of New Zealand, it may

have been originally introduced from Australia after the sepa-

ration of the islands. In the Stewart Island examples of this

species (JZ. chaltont) the well-known apophysis at the distal end

of the palpal patella varies in size individually, but the banding of

the palp in yellow and white is always the same, and I look on

the specimens as constituting but one species.

Of Pantopsalis probably ten species may be taken as established.

The curious difference in the length and shape of the man-

dibles among the specimens of this genus is remarkable, and it

has been a matter of uncertainty whether this difference is

sexual, dimorphic in one sex, or specific. In the whole family

the sexes are not easy to distinguish accurately without spoiling

the specimens, and many of the species have been described from

single examples. Judging from the specimens in this collection,

added to the evidence hitherto available, it would appear that

* For explanation of the Plates see p. 48.

+ “Opiliones,” W. Sérensen, in Koch & Keyserling, Arachn. Austr., Suppl. p. 55.

£ “Some new Harvest Spiders,” R. I. Pocock, Proc. Zool. Soc. Lond. 1902, vol. ii.

p. 398. , :

§ “Some New Zealand and Tasmanian Arachnide,” H. R. Hogg, Trans. N.Z.

Inst. vol. xlii. (1910), p. 277.

oe MR. H. R. HOGG ON SOME

the males have the long mandibles and the females the short,

but discrepancies have to be cleared up before this can be

accepted for a certainty. It may be regarded as quite certain

that the difference is not specific, both forms being found in

members of the same species.

White’s short description* of the type species P. listerz, and

Simon’s note on the original types in the Paris Museum t, show

that the specimens originally described had the long mandibles,

but the sex was not determined. Mr. Pocock, judging from

specimens, said to be females, in the British Museum, records

P. listeri among those species with the shorter mandibles,

thereby distinguishing it from his P. albipalpis, which had them

long. Therefore, if his determination is correct, there are both

long and short mandibles in that species.

P. trippi Poe. with long mandibles, is a male.

P. tasmanica Hoge Bae eo 3 a dried specimen, sex undeterminable.

P. nigripalpis Poc. Pras san sf is a male.

P. wattsi, sp. u. Q (from the shape of the genital oper-

P. pococki, sp. n. § 2 ae 28 2 cula I take to be males.

P. coronata Poe. with short mandibles, sex not recorded.

P. halli, sp. n. seas 53 operculum broader and straighter in

specimens with short mandibles.

Mandibles long in specimens cer-

tainly males.

P. grayi, sp. n. sagas a operculum broader, female.

On the other hand, Mr. Pocock first described his P. jenningst

as having short mandibles and as a female, but later found it to

bea male. Here, however, the basal segment of the mandible is

twice as long as the palp, and the second segment six times as

long as wide, which is hardly a short mandible when compared

with that of the females recorded above.

Mr. Pocock gives a synopsis of the species described (Ann. &

Mag. Nat. Hist. May 1903, p. 438), and this I supplement

below :—

Suborder PALPATORES.

Family PHALANGIID.

Genus PANTOPSALIS Simon.

a. Patella, tibia, and distal joint of palpi white or yellowish

white.

@, Mandibles red-brown; carapace black; eye-tubercle

SMMOOPM + sth.40 eeeb bee ssaece tec) UA areteepR eee ea each eens MILESCEMOMN abies

a. A group of small spicules on front border ............ ? albipalpis Poe.

8. About half a dozen strongish spicules in front ...... trippi Poc.

61, Mandibles pale yellow-brown; carapace cream-colour,

with dark patch in front of eye-tubercle spiculed

in front; eye-tubercles smooth........................... watési, sp. n.

* Adam White, Proc. Zool. Soc. Lond. vol. xvii. (1849), p. 6.

+ HE. Simon, C.R. Soc. Ent. de Belg. May 2, 1879, p. 16.

AUSTRALIAN OPILIONES. 33

6. Patella, tibia, and distal joint of palpi dark brown, yellow-

brown or deep yellow.

a*, Kye-tubercle denticulated ; carapace dark brown.

a3, Palpi black brown .............ccc00cceceeeeeceseereesenneee nigripalpis Pec. and

subsp. spiculosa Poc.

68, Palpi pale yellow-brown ...................00eeeeeseeeess pococki, sp. n.

6°, Eye-tubercle smooth.

a‘, Spiniform tubercles on frontal area; palpi uni-

HorlAlly IDEVES, 6. ccdenanee noboncoe cbabhaesececeaDocacesoopane . GEOG SO A HOG:

6+. Carapace wholly smooth.

a. A short spine on trochanter i.; palpi yellow,

banded with brown; carapace tortoise-shell ... gray, sp. n.

6°. Trochanter 1. smooth.

a®, Palpi yellow, blotched with grey; carapace

dark yellow-brown .......... seccesees eee. tasmanica Hoge.

66, Palpi reddish brown; red and: grey stripes on

rear segments of abdomen Seema coronata Poc.

c®, Palpi orange; carapace callomeiarer. Ww ith

brown patches; legs bright yellow-brown ... halli, sp. n.

PANTOPSALIS LISTERI White.

Proc. Zool. Soc. 1849, p. 6.

Ty pe species.

PANTOPSALIS NIGRIPALPIS Poc.

Pocock, “Some new Harvest Spiders,’ Proc. Zool. Soc.

Lond. 1902, vol. 11. p. 399; 2d., “Some new Tropical and

Southern Opiliones,” Ann. & Mag. Nat. Hist. ser. 7, voi. xi.

1903, p. 438.

A specimen with the eye-tubercle and surrounding portion of

the carapace more strongly denticulated, Mr. Pocock has named

as a subspecies of the above, P. spiculosa.

PANTOPSALIS CORONATA Poc.

Pocock, Ann. & Mag. Nat. Hist. supra, p. 436.

PANTOPSALIS TRIPPI Poc.

Pocock, Ann. & Mag. Nat. Hist. supra, p. 437.

PANTOPSALIS JENNINGSI Poc.

Pocock, Ann. & Mag. Nat. Hist. supra, p. 437.

PANTOPSALIS TASMANICA Hogg

Hogg, “Some New Zealand and Tasmanian Arachnide,”

Trans. N.Z. Inst. vol. xlii. 1910, p. 279.

PANTOPSALIS WATTSI, sp.n. (PI. I. fig. 3.)

The cephalic area is dark yellow-brown in front over a median

space as far as and including the eye-tubercle. This is bounded

by a wide horseshoe belt of bright cream-colour, beyond which

Proc. Zoot, Soc.—1920, No. II, 3

34 MR, H. R. HOGG ON SOME

again as far as the side margin it is yellow-brown, but intruding

at intervals into the lighter colour.

The eye-tubercle is smooth, brown and yellow, with the eyes

yellow.

The abdomen is unfortunately partially broken but apparently

brown.

The mandibles are long, pale yellow-brown, strongly bespiculed

in rather regular bands. ‘The palpi, remarkably long, are brown

on the basal half of the femur, yellow on the distal half, and

pale yellow on the remaining joints. The legs are banded

alternately with brown and yellow, and are also very long.

On the under side the cox, operculum, and mouth-parts are

yellow-brown, dark in the two former and lighter in the latter.

The space between the eye-tubercle and the front of the carapace

is bespiculed.

The measurements (in millimetres) are as follows :—

Ceph. lg. 3, lat. 25 in front, 4 behind; mand. 20; palpus 82 ;

Ist leg 374, 2nd 65, 3rd 373, 4th 453.

This specimen, apparently a male, was collected at Hawera, in

the North Island, N.Z., by Mr. G. 8S. Watts, and I have named

it after him.

PANTOPSALIS POCOCKI, sp. n. (Pi. I. figs. 3 a—d.)

The cephalic part of the carapace is dark brown, the abdo-

minal portion the same in the middle with pale yellow-grey

at the sides, into which the darker colour runs in places. The

posterior segments are grey, with transverse marginal streaks of

black-brown:

The mandibles are dark red-brown, long, profusely covered

with rather large spicules, and are moderately thin until the

thickening at the distal end.

The femoral joint of the legs black-brown, the remaining joints

red-brown. The palpi are uniformly yellow-brown till quite at

their distal end, which is yellowish grey.

The carapace, including the eye-tubercle, is almost wholly

covered with small spicules. ‘The eyes are large and black.

The under side, including the coxal and sternal area, is black-

brown.

The measurements (in millimetres) are as follows :—

Ceph. lg. 27, lat. 33; abd. lg. 27, lat. 32; mand. 233; palpus 47;

Ist leg 30, 2nd 56, 3rd 26, 4th 37.

This specimen, collected on Mt. Dick, N.Z., by Mr. T. Hall,

IT have named after Mr. R. I. Pocock, to whom the students of

the Opiliones are much indebted for his valuable papers.

PANTOPSALIS HALLI, sp. n. (PI. I. figs. 4 ad.)

The sides of the cephalic part are mottled yellow and dark

brown, a broad median area being paler yellow and light brown.

The eye-tubercle is pale yellow-brown, rather darker in the

median longitudinal streak ; eyes quite black.

AUSTRALIAN OPILIONES. 35

The dorsal abdominal part has a still paler median area, light

yellowish grey, narrower in front and broadening posteriorly to

a scolloped pattern near the rear end. This is bordered by a

dark brown streak on each side, separating the median area from

the bright yellow-brown sides.

The mandibles and palpi are bright yellow, and the legs pale

yellow-brown, except the distal end of femora 3 and 4, which is

darker.

The carapace and abdomen are both quite smooth and destitute

of granulation, as are also the mandibles, which are of the quite

short type, less than the length of the body.

The coxee and under side of the abdomen are pale yellow-

brown, quite smooth, and the segmental divisions almost obliter-

ated, being just indicated by darker transverse streaks.

The patellar joint of the palpi is slightly longer than the tibial,

both joints broadening anteriorly and being profusely covered

with short bristles.

The genital operculum is thick and broad.

The measurements (in millimetr ey) are as follows :—

Ceph. Ig. 2, lat. 14 in front, 24 behind; abd. lg. 25, lat. 3.;

mand. 32; palpus 6; Ist leg 18, and 423, 3rd 194, dth 29.

Four specimens, much shriveled and without label, but from

New Zealand. Three are males with long mandibles and the male-

shaped operculum, and one is a fariale with seemingly broader

and straighter operculum and short mandibles. Also one female,

collected on Mt. Algidus, Rakaia Gorge, South Island, by

Mr. T. Hall, to whom the species is dedicated. It is to his

efforts that this large and valuable collection is mainly due.

PANTOPSALIS GRAYI, sp. n. (PI. I. figs. 5 a-c.)

The cephalic part of the carapace is bright yellow-brown. The

abdominal part is rather darker brown in the median area, pale

and dark mingled at the sides. The under side is pale yellow from

the front to as far as the base of the genital operculum, brown on

the abdominal segments. ‘he coxe are pale yellow, with a dark

brown band on each at the end nearest to the trochanter.

The legs are yellow, banded with brown. The mandibles pale

yellow, blotched with brown. The palpi yellow, banded with

brown on the femur and patella, more faintly on.the tibial and

distal joints.

The mandibles are of the short type, not so long as the body,

and, though covered with short bristles, are otherwise smooth.

The carapace is wholly smooth, but there is a short spine on

trochanter i. The eye-tubercle is nearly in the centre of the

cephalic region. The genital operculum is broad in front.

The measurements (in millimetres, but legs mostly broken)

are as follows :—

Ceph. lg. 24; abd. lg. 34; mand. 327; palpus 5; Ist leg 31.

A single specimen, apparently a Peinalel collected at Hukanui,

Waikaremoana, North Island of N.Z., by Mr. W. R. Gray, after

36 MR. H. R. HOGG ON SOME

whom I have named the species, was found in close proximity to

a male specimen, which agrees closely with M. Simon’s description

of the Paris Museum types of P. listeri White; but the two are

very different in coloration.

Suborder LANIATORES.

Family TRI Z#NOBUNID® Poe.

Proc. Zool. Soc. Lond. 1902, vol. ii. p. 400.

Genus TRIZNOBUNUS Nor.

W. Sorensen in Koch & Keys. Die Arachn. Ausitr.,

Suppl. p. 59 (1886).

Dr. Sdrensen described the type species of this genus 7’. bicari-

natus from Sydney, and Mr, R. I. Pocock 7’. pectinatus from

Tasmania. To these I have now to add a New Zealand form

from the collection of Mr. T. Hall. In this the curious structure

of the eye-tubercle is still more elaborated than in the other two,

since it has four pairs of lateral branches. ‘There is, further;

an apical spine, and the horns on each side of the base are

more powerful. The whole creature is profusely bespined and

granulated.

Dr. Sdrensen (loc. cit.) merely regarded Triawnobunus as a

second genus in the family Trienonychide, but Mr. Pocock very

rightiy writes it down as a separate family. The quite different

type of sternum, together with the smaller and weaker palpi,

apart from the very remarkable development of the eye-tubercle,

show that it has proceeded along considerably different lines

from the above-named family. The form of the eye-tubercle is a

specialisation of that seen in Hridanus Thor. and Phistobunus Poc.

of M. Simon’s family Phalangodide.

Synopsis of S'pecies.

a. ye-tubercle having two pairs of lateral branches in front of

the eyes.

a}. Two longitudinal rows of spinous tubercles reaching

from the base of the eye-tubercle to the anterior rear

SECMMEM Ge erie eicicteveatepetahe onal mcciecetoamle arden eanen Meer

b}. A single median longitudinal row of spinous tubercles

reaching from the base of the eye-tubercles to the

anterior rear segment .........ceeeeeeeeeeeeeersceeteeteeseeese pectinatus Poe.

b. Hye-tubercle having three pairs of lateral branches in front of

PIAGEWES: razubedsucecuateest ec Atte ace aac ae CESSES EES eR ET ECTRa eS

bicarinatus Sor.

acuminatus, sp. n.

TRIZNOBUNUS ACUMINATUS, Sp. n. (PI. II. figs. 6 ad.)

Colour. Upper side greyish yellow all over; the small tubercles

forming a pattern on the back are rather yellow. The under side

is more yellow all over, with two dark brown patches on each side

of the anal segment. Both joints of the mandibles are so thickly

covered with black network on yellow ground as to appear black.

AUSTRALIAN OPILIONES. Bil

The palpi are palish yellow, banded with black; the legs have the

femur, patella, and tibia darker and greyer, the tarsal joint being

quite ight yellow.

One specimen, which I take to bea female, is much lighter and

greyer above than the others, and on the under side pale yellow,

without the black bands on the palpi. The genital operculum is

more thickly covered with bristles and the sternum is corrugated,

whereas in the others it is not. The spinous tubercles on the

segments are more in number (about 14). Otherwise there is

not much difference.

As in Mr. Pocock’s 7’. pectinatus from Tasmania, which this

seems to resemble more than Dr. Sérensen’s 7’. bicarinatus from

Sydney, the dorsal carapace is decorated with longitudinal and

transverse rows of small round tubercles, forming ringed areas

smooth in the inside. There are four longitudinal and about five

transverse rows of granules, but no median longitudinal row of

larger granules.

The very remarkable eye-bearing tubercle, rising and pointing

‘upwards and forwards from the front margin, is broadest at the

base, and tapers to a point at the anterior end. ‘The eyes are

situated one on each side, about half-way from base to summit.

Just behind the eyes a pair of long, pointed tubercles jut out one

from each side. ‘These are followed by three more similar pairs,

and the whole is topped by one median tubercle ending in a sharp

point. The intermediate space between these is filled up with

small granular and pointed tubercles. On the front edge of

the carapace on each side of the eye-tubercle are five spinous

tubercles.

Between the last row of small tubercles and the first of the

rear segments is a pair of moderate-sized spinous tubercles rather

wide apart.

On each of the rear segments is a transverse row of long

spinous tubercles, about nine in number in the males, but more

numerous (about 12 or 14) and not quite so large in the female.

On the under side the coxe of the palpi and first and second

pairs of legs are profusely covered with granular tubercles, but

the 8rd and 4th coxe are granular only along their margins.

Each segment has a row of round tubercles rather larger than

the above.

The soft mouth-parts of the inner end of the palpal coxa form

a very distinct upper jaw, divided longitudinally into two parts.

The same parts of the coxa of the first pair of legs similarly form

the lower jaw, each part lying on one side of a soft median lip.

Coxa ii., on the other hand, ends in a flat chitinous piece, which,

turning forwards, clasps the mouth-parts, and would appear to

support and hold them together.

Coxa lili. is similarly hard but black, and the pair seems to

form “‘areuli genitales” on each side of the genital operculum

and sternum. The end of coxa iv. is a narrow ridge of hard

black substance, a continuation of the sides of the triangular

38 MR. H. R. HOGG ON SOME

sternum. The latter is like the lower end of the sternum in

Trienonyx, but without the narrow stem; consequently the

operculum is brought much closer to the mouth-parts, as in

genera so far removed as Stylocellus Westw. The head of the

penis is bluntly triangular, and its upper and lower sheaths, as

far as can be seen when not extended, are terminated in a

transverse line.

The mandibles ave shorter and weaker thanin 7riwnonya; the

first joint does not reach as far as the end of the eye-tubercle.

The palpi also are short and weak, and are not longer than the

dorsal carapace,

The legs are fringed the whole distance along the femora,

patelle, and tibize with long spines ranged on each side of the

respective joints.

The measurements (in rol ipacilies) are as follows :—

Ceph. ls. 1j, lat. 13; abd. lg. 22, lat.21; mand. 1; palpus 22;

Ist leg 5, 2nd 8, 3rd 64, 4th 83.

This species differs from 7. seniors Poe. in having a larger

number of projections along the sides of the eye-tubercle, in’

having a larger number of spinous tubercles on the rear segments,

and in being without the median row of larger tubercles on the

dorsal carapace.

I have four specimens, of which three are certainly males.

They were collected by Mr. T. Hall near the Holliford River,

L. Wakatipu district in the province of Otago.

Family TRI#ZNONYCHID.

Genus TRI#NonyxX Sor.

W. Sorensen in Koch & Keys. Die Arachn. Austr., Suppl. p. 58.

This genus, being now restricted to those species in which the

eye-tubercle is marginal, includes, with new ones that 1 am now

recording, only seven in number. These may be distinguished as

follows :—

a. ay coarsely granulated.

. Transverse row of granules on carapace, and a pair of

spinous tubercles at posterior end of same ............ 7. rapax Sor.

b'. No spinous tubercles on carapace.

a*, Carapace chocolate-brown ; rear segments orna-

mented with round flat orange spots, with a

short bristle on each .............0.....000ee000ee 2, cockayni, sp. n.

6. Carapace deep brown; a series of low tubercles,

hut no bright spots on rear segments.

a. Distal end of tarsus elongated..................... T. coriacea Poc.*

(3. Distal end of tarsus nodular .................... ZZ. aspera Poe.

* Mr. Pocock (Ann. & Mag. Nat. Hist. May 1903, p. 445) refers to a species in

this section under the name of 7. verrucosa. This I aim unable to trace the origin

of, and he allows me to state that he was, he believes, intending to refer to the

species named 7. coriacea.

AUSTRALIAN OPILIONES. 39

6. Carapace smooth or finely granulated.

a’, Carapace yellow-brown, with a dark brown rectan-

gular pattern round the margins and a similar |

pattern behind the eye-tubercle; yellow spots in

the posterior area between the two patterns ......... T. variegata, sp. n.

63. Carapace orange, two black patches on the yellow eye-

tubercle ; legs very pale yellow ; a row of fine spines

on upper surface of each segment of abdomen ...... T. testaceus Hoge.

e®. Carapace dark yellow-brown, a row of tubercles along

its posterior border in front of first rear segment ;

a longitudinal row of short bristles on the median

Ihave 8 a ae ee eevee emery ce y's | Roopa a opty,

TRLENONYX RAPAX Sor.

W. Sorensen in Koch & Keys. Die Arachn. Austr., Suppl.

1886, p. 58.

Locality. Fiji.

TRLENONYX CORIACEA Poc.

R. I. Poeock, “Some new Harvest Spiders,” Proc. Zool. Soc.

Lond. 1902, vol. 11. p. 403.

Locality. Auckland, N.Z.

TRIZNONYX ASPERA Poc.

Loe. cit. supra, p. 404.

Mr. Pocock states that this species, only located as Austrahan,

differs from other species in the formation of the end of the

tarsal joint, which, instead of being elongated, has the last portion

spherical or nodular.

TRIENONYX TESTACEUS Hogg.

‘H.R. Hogs, “Some New Zealand and Tasmanian Arach-

nide,” ‘Frans. N.Z. Inst. vol. xli. 1910, p. 280.

Locality. New Zealand only.

TRIBNONYX STEWARTIUS Hogg.

H. R. Hogg, loc. cit. supra, p. 281.

Locality. Stewart Island, N.Z.

TRIHNONYX COCKAYNI, sp. n. (PI. II. figs. 7 a—d.)

Colour, Male: orange, shaded in patches at the side, and in

the middle line and on the eye-tubercle with brown; females

browner all over. Transverse lines of round orange spots on the

rear dorsal segments, the ventral segments yellow- -brown and

smooth. The mandibles, and more faintly the upper sides of the

palpi, have a brown network pattern; the legs are yellow, banded

with brown.

The dorsal surface is coarsely granulated. No raised tubercles

on the segments, but a short bristle on each round spot and no

spines on anterior margin of carapace. The eye-tubercle is

marginal, straight in front, but sloping posteriorly. On the

40 MR. H. R. HOGG ON SOME

ventral surface the oral segment of coxa ii. is well developed,

pointing prominently forward, but is flat and does not stand up

from the surface; the mouth-parts, on the other hand, of coxa i.

and the palpal coxa stand up well above the surface-level.

The mandibles are smooth, the basal segment 13 times as long

as wide, with two small spines on the inner side of the anterior

end. The palpi robust, the trochanter with three spines under-

neath, in front, on the outer side; three powerful spines under

the femur; two small spines under the patella; three long spines

on each side underneath the tibia; three small spines on each

side and a large terminal spine on the distal joint.

The extended sheath of the male penis is bilobed, the penis

itself terminated by fine bristles.

The coxe of the anterior three legs are corrugated and

tubercled, that of the 4th pair smooth.

In the female the sternum between the third pair of coxee is |

corrugated and narrow.

The measurements (in millimetres) are as follows :—

Ceph. lg. 2, lat. 3; abd. lg. 4, lat. 4; palpus 64; Ist leg 11,

2nd 14, 3rd 94, 4th 13.

One male and four females were collected by Dr. Cockayne

from Kapiti Isiand, off the S.W. coast of the North Island of New

Zealand.

TRIENONYX VARIEGATA, Sp.n. (PI. II. figs. 8 a-c.)

Colour. Female: pale greyish yellow, with dark brown rect-

angular pattern reaching from the eye-tubercle to the first rear

segment, also a smaller and lighter coloured similar rectangle

reaching from behind the eye-tubercle to the posterior end of the

cephalic part, and transverse rows of yellow spots round on the

carapace, oval on the segments; the under side is pale all over.

The legs are pale yellow, banded with dark grey; the mandibles,

and femoral and tibial joints of palpi are deep black, speckled

with small yellow spots. The eye-tubercle yellow-grey in the

middle, dark grey at the sides; eyes orange.

The dorsal surface 1s smooth but coriaceous, the spots thereon

flat, with short bristles, but the segmental spots slightly raised.

The marginal eye-tubercle, rather high and conical, has a wide

circular base, narrowing to a blunt point anteriorly, but there is

nothing either spinous or tubercular about it. The eyes are at

the side. There are no spines on any part of the carapace.

The mandibles are smooth, without tubercles, the first segment

about twice as long as wide.

The palpi are rather slight, the usual spines on the under side

little more than low tubercles, and none on the upper side.

The sternwm between the 3rd pair of coxe is wide and corrugated,

in the male, with an upper and lower division, and above this a

rectangular lip between the prominent oral parts of coxe i. The

oral portions of coxe ii. are hardly formed at all and in no wise

raised,

AUSTRALIAN OPILIONES. Al

The measurements (in millimetres) are as follows :—

Q. Ceph. Ig. 13, lat. 13; abd. Ig. 22, lat. 23; mand. 21;

palpus 34; 1st leg 5, 2nd 8, 3rd 53, 4th 72.

S. Ceph. Ig. 17, lat. 13 ; abd. lg. 23, lat. 24 ; mand. 12 ;

palpus 32; Ist leg 5, 2nd 8}, 3rd 43, 4th 84.

There are three females from the Holliford River and one

young from Mt. Remarkables, L. Wakatipu. One from Mt.

Oakden ; also six specimens from Paradise, L. Wakatipu, of which

four are apparently males, with more prominent spines on the

palpi and without the clear pattern on the back. They were all

gathered by Mr. T. Hall.

The males are, as mentioned by Mr. Pocock, generally more

constricted in the cephalic part of the carapace than the females.

The wide sternum and well-formed lip above it are features

which might almest be held to be of generic importance, but I

have not made them so in consideration of other resemblances

the species shows to the genus 7’ricenonyx.

Genus Nunera Loman.

Nuneia J. C. C. Loman, Zool. Jahrb. Syst. xvi. 1902, p. 214.

Nuncia R. I. Pocock, Ann. & Mag. Nat. Hist. ser. 7, vol. xi.

May 1903, p. 440.

* Following the limitations adopted by Loman and Pocock, we

may assume that the species to be attributed to this genus have

the ocular tubercle rising at a short distance from the front

margin of the carapace, that it is smooth and rather low and

without any prominence thereon ; the carapace itself smooth, the

sternum in both sexes narrow, and the tarsal claws of the third

and fourth pairs of legs with short side claws springing from

near the base of the median claw. On this basis we have the

following :—

NUNCcIA SPERATA Loman.

Dr. J. C. C. Loman, loc. cié. supra.

From Stephen’s Island on the N. coast of the South Island of

New Zealand.

NUNCIA VALDIVIENSIS Sor.

Trienonyx valdiviensis W.Sérensen in Koch & Keys. Die

- Arachn., Austr.

Nuncia valdiviensis R. I. Pocock, Ann. & Mag. Nat. Hist.

supra.

From Chili.

NUNCIA SUBL&VIS Poc.

Trienonyx sublevis R. I. Pocock, Proe. Zool. Soc. Lond.

1902, vol. ii. p. 404.

Nuneia sublevis R. 1. Pocock, Ann. & Mag. Nat. Hist, supra.

Locality. Otago, 8. Island, New Zealand.

49 MR. H. R. HOGG ON SOME

NUNCIA ENDERBHI Hoge.

Trienonyx enderbei WH. R. Hogg, in Dr. Chilton, “ Sub-

antarctic Islands of New Zealanc , Wellington, N.Z.,

1909.

This species, from the Enderby and Auckland Islands, clearly

comes into the genus Vuncia.

NUNCIA SMITHI, sp.n. (PI. II. figs. 9 a-c.) .

Colour. Male: carapace rather deep red-brown, mingled with

black patches and streaks, but hardly suggesting much of a

pattern. The edges of the segments are marked in some cases

with bright white transverse “lines. The mandibles are dark

brown and the palpi bright yellow, with black network pattern

on the lower portions of the femoral, patellar, and tibial joints.

The under side is paler yellow-brown.

The female is pale yellow with brown markings, and in both

cases the legs are yellow with brown bands.

The eye- ee rele 18 low (but rather higher in the male than in

the female), slightly removed from the front margin of the cara-

pace. The latter is finely granulated, and the dorsal surface is

without warts or spines. The first joint of the mandibles is about

twice as long as broad, with a bunch of short spines near the base;

the fingers long and fine. On the basal part of the femur of the

palp are one long spine and two short ones, and there is another

half-way up. On the inner side is a row of short blunt tubercles

with a bristle on each, and on the outer side a row of bristles ;

on the tibial joint are two long spines on the inner side and two

warty knobs.

The true maxillary process of coxa ii, is flat, and clasps on each

side the soft white mouth-parts of coxa i., but behind this is a

large prominence standing up from the surface and pointing

rather backwards. At the outer end of coxa iv., Just above the

spiracle, are three short tubercles followed by a rather long club-

shaped knob. The tarsal claws of legs iii. and iv. have each a

pair of moderately large side claws springing from the base. The

sternum is narrow, corrugated, with a triangular base and a

spear-headed distal end.

The measurements (in millimetres) are as follows :—

Ceph. lg. 17, lat. 22; abd. lg. 33, lat. 32; mand. 37; palpus 1};

Ist leg 7, ond 102, 3rd 7, Ath 10.

I have one male and one female collected by Mr. W. W. Smith

at Hawkes Bay, North Island of N. Zealand, and two males by

Dr. Chilton from Picton at the north of the South Island.

Genus SORENSENELLA Poe. ;

R. I. Pocock, “Some new Harvest Spiders,” Proc. Zool. Soc.

Lond. 1902, vol. 11. p. 409.

The two species of this genus deseribed by Mr. Pocock have

the eye-tubercle placed shortly behind the front margin of the

AUSTRALIAN OPILIONES. 43

carapace and bearing on its summit a short dentiform tubercle.

The claws of the third and fourth pairs of legs have the side

branches longer than the median claw in both sexes. I have a

male and female which I am placing in the genus following

(Monoxyomma), in which this type of claw appears in the male

only, the female having the side claws shorter than the median.

SoRENSENELLA PREHENSOR Poc.

R. I. Pocock, loc. cit. supra.

Locality. New Zealand (only),

SORENSENELLA BICORNIS Poc.

R. I. Pocock, Ann. & Mag. Nat. Hist. ser. 7, vol. xi. May

1903, p. 439.

Male and female from Christchurch, N.Z.

Genus MonoxyomMa Poc.

R. I. Pocock, Ann. & Mag. Nat. Hist. ser. 7, vol. x1.

May 1903, p. 444.

Mr. Pocock formed this genus for those members of the family

Triznonychide in which the ocular tubercle, rising distinctly

behind the anterior margin of the carapace, is protected by a

‘Jong suberect spine. He also gives as generic characters a pair

of long spines on or about the third abdominal segment, and the

dorsal valve or sheath of penis trilabiate.

The pair of dorsal spines is far from being peculiar to this genus.

It is present in a median bifurcated form in one case, and wanting

in another, among the species which I am compelled to consider

as belonging to this genus since they otherwise conform to the

ty pe.

The third character suffers under the disability of being a

sexual one; and it is a matter of considerable difficulty, in the

majority of cases, to ascertain whether it appertains to a par-

ticular species or not.

The following synopsis may help to distinguish the species

below recorded :—

a. Dorsal carapace roughly granular all over ......... M. tuberculatum, sp. n.

6. Dorsal carapace smooth or small granules only.

a}, A pair of spinous tubercles near rear end of

(CEAPROEXOSY 34) chocadeaduindeaad eee Bene Santen Eerisen 2005006

61. No pair of spinous tubercles near rear end of

carapace.

a*, A bifurcated spinous tubercle on median

line; no spines on anterior margin ; palpi

15 times as long as body ...............-.-..-... ML. hendei, sp. n.

62, No spinous tubercles on carapace; a pair 0

short spines, one on each corner of anterior

margin; palpi nearly twice as long as body. JM. trailli, sp. n.

M. spinatwm Poe.

44 MR. H. R. HOGG ON SOME

MonoxyomMaA sPInatum Poe.

Ann. & Mag. Nat. Hist. loc. cit. p. 445.

The type species, sent by Dr. Brown from Hill Grove,

N.S. Wales. ;

MoNoOXYOMMA HENDEI, sp.n. (PI. III. figs. 10 a-d.)

Colour. The cephalic part is yellow-brown smirched with black-

brown, chiefly at the sides. The thoracic part and segments

almost wholly black-brown. The mandibles yellow-brown with a

slight network of black, and the palpi the same but the network

heavier. The under side of the carapace is very similar, yellow-

brown and black being mixed up without much semblance of

pattern. On the segments, however, the two colours are in

transverse bands. On the legs the same mixture occurs, the pale

being more in the joints. On the femur and tibia the two colours

appear in longitudinal bands.

The median area of the carapace is slightly convex, this part

being circumscribed by a ridge in front, and at the sides with a

flat streak between it and the margin. The surface is coriaceous,

without spines or tubercles, except as below, either thereon or

projecting from the margin. The eye-tubercle is situated on the

inner side of the ridge above mentioned ; it is conical and rather

tall, with the horn-shaped projection from its highest point bend-

ing slightly forward. This is about the same height as the lower

tubercle, from which it springs. Near the posterior end of the

carapace there is a large bifid tubercle in the middle line, with

two transverse rows of small single ones, each with a bristle in

the middle, between it and the first of the segments.

On each of the three abdomiual segments is a row of similar

warty tubercles, rather larger than the above, each with a short

bristle.

The mandibles and palpi are fairly long and strong. On the

femoral joint of the latter are two longitudinal rows of quite

small spines; on the tibial and distal joints the usual longer

spines. On the trochanter, femur, patella, and tibia of all legs are

rows of small warty tubercles with short spines, and similar but

smaller on the coxe.

The measurements (in millimetres) are as follows :—

Total length 22, breadth 2; mand. 2; palpus 4; Ist leg 63,

2nd 92, 3rd 7, 4th 93.

One male from Hende’s Ferry, Central Westland, N.Z., which

1 have named after Mr. J. W. Hende, the collector.

MonoxyoMMA TUBERCULATUM, sp.n. (Pl. III. figs. 11 a-c.)

Colour. Males: a uniform dark yellow-brown over whole of

carapace. The posterior edge of each rear segment is distinctly

marked out by a pale transverse line; the mandibles and palpi

AUSIRALIAN OPILIONES. 45,

are rather bright yellow. The legs ashy grey. The under side

is yellow from the front as far as the rear coxe; dark brown

behind this on the sterna, which are bordered with paler edging.

Two specimens, which I take to be females, are ashy grey above

with pale grey spines and almost wholly yellow underneath, being

there grey only at the extreme end; on the upper side there is a

distinct rhomboidal pattern in pale grey on a dark ground.

The cephalic part of the carapace is thickly covered with coarse

granulations ; it is bounded posteriorly by a distinct depression

separating it from the abdominal portion. The front border has

one long median spine, flanked on each side by five smaller

spines. The eye-tubercle takes its rise at a point well behind

the front margin; it is roughly hemispherical, with the eyes

looking upwards, and a median horn about equal to the height

of the basal part on which it stands. Behind the suleus above

mentioned the granulations form themselves more into longi-

tudinal and transverse rows, but without achieving any very

definite pattern. About midway between the dividing sulcus

and the rear end are a pair of rather large spinous tubercles, and

behind these four more, of which the middle two are the largest.

The rear segments have regular rows of spinous tubercles,

about ten on front row to four on the back row.

The basal joint of the mandibles is thick and bulges upwards.

On the basal part of the second joint are several spinous

tubercles.

The palpi are thick and powerfully bespined on all joints. In

length they just equal the body. ‘hose of the supposed female

are “slighter and less strongly armed.

The “legs are covered with spinous granules « as far as the tibial

joint, and the spaces between the trochanters of ii. and ili., and

uli. and iv. are armed with three or four spinous tubercles. On

the claws of tarsi 11. and iv. are two quite small side wings

inserted about the middle of the claw.

I have little doubt about the sexes, but on removing the oper-

culum of one supposed male, there was nothing at. the bottom of

a deep hollow but a granular mass.

The measurements (in millimetres) are as follows :—

Ceph. lg. 14, lat. 235 abd. lg. 33, lat. 32; mand. 23; palpus 5;

Ist leg 77, 2nd 142, 3rd 11, Ath 1g,

These are three males ‘and two females from Mt. AJgidus,

Rakaia Gorge, South Island, N.Z.

MoNoxYOMMA TRAILLI, sp. n. (PI. III. figs. 12 a-f.)

Colour. Male: a black-brown border along the front edge of

the carapace, along the sides, and the same colour over the rear

seements; in the median area there is a large yellowish patch.

On the segments large, round faint yellow spots in transverse

VOWS. The mandibles are yellow, with a black network pattern

AG MR. H. R. HOGG ON SOME

on both joints. The palpi are orange, with dark blotches on the

femur and inside of tibia. The legs are dingy yellow, ringed with

brown. On the under side the whole of the cephalic part is

orange, and the segments black-brown. In the female the pale

area of the carapace is paler and more cream-colour than in the

male, and the femur and patella of the palp are more continuously

black-brown, otherwise the coloration is much the same.

The carapace in both sexes is of a dull smoothness without

granulations, and the only spinous tubercles are one at each

front corner at the side of the trochanter of the first leg. The

spots on the segments are slightly raised, with a bristle on each.

The eye-tubercle is clearly removed from the front margin, and

the portion between the eyes is produced forward into a peak

about as high as the portion of the tubercle below the eyes.

The mandibles are remarkably short, but the first joint bulges

upwards above the base. On the other hand, the palpi are

nearly twice as long as the whole body, the femoral joint bowed

like the first joint of the mandible, and powerfully bespined both

above and below. The other joints are similarly spined.

The tarsal joint of legs 111. and iv. in the male has the modifi-

cation of the claws which Mr. Pocock considered one of the

points of his genus Sorensenella—viz., the side claws longer than

the median; but in the female the claws are normal, the side

claws being only half the length of the median, but springing

from the base, thus showing that this character is only sexual in

some cases. .

The sternum in the male is of the narrow type, triangular at

the base and spear-headed at the distal end, with the oral part

of coxe ii. meeting above it and no visible lip; but in the female

it is broad, as I have above described it in 7ricenonyx variegaia,

with a well-defined lip in front. It would appear, therefore, as

if this broad sternum, where it occurs, is a sexual character. I

have re-examined the specimens I believe rightly taken to be

males of 7’. variegata (indeed, in one of them the penis is exposed ) ;

and, although the sternal depression is still wide, a slightly-

formed narrow sternum is to be seen in the median line, and the

lip is above it.

The measurements (in millimetres) are as follows :—

Male. Ceph. 1g. 22, lat. 31; abd. lg. 22, lat. 4; mand. 2; palpus 9;

Ist leg 83, 2nd 14}, 3rd 9, 4th 13.

Mr. W. Traill, after whom I have named them, sent the

specimens, one male and one female, from Stewart Island.

ALGIDIA, gen. nov.

This genus differs from the others in the family in having the

genital operculum, in both sexes, furnished with denticular

tubercles, each with a short bristle at its apex, extending along

the front margin and toa greater or shorter distance down the

AUSTRALIAN OPILIONES. A7

sides. The eye-tubercle, which is situated the length of its

diameter behind the front margin of the carapace, bears wart-

like protuberances in its median line, varying from one only to a

row of three or four. The mandibles are short and weak. The

palpi rather slight, but tuberculated and strongly bespined. The

rear segments and trochanters are strongly bespined and the

carapace profusely granulated.

ALGIDIA CUSPIDATA, sp.n. (PI. IIT. figs. 13 a-e.)

Colour. Male: carapace dingy yellow, with two dark, broad,

longitudinal stripes, beginning one each side of the eye- tubercle

and reaching to the level off the fourth trochanter, where it

turns outwards to the margin. The mandibles and palpi are

darker, the latter covered with pale wart-like protuberances and

the former with black network pattern. The legs are yellow

with dark grey rings; the under side yellow-grey. The female

is paler on the carapace; the dark stripes not so conspicuous, but

the warty pattern more regular. The legs, mandibles, and palpi

about the same as in the male.

The eye-tubercle is hemispherical ; the median row of warts

three or four in the males, fewer in the females.

The carapace in the male is strongly constricted behind the

cephalic part, the sides straighter in the female. On the front

margin in the male there are seven long spines in front of the

eye-tubercle and three smaller behind each corner. In the female

are three formidable spines in the centre, flanked by two small

ones, and none at the side. The median area and sides of the

carapace are thickly covered with warty pustules, but there are

none on the darker stripes.

On the abdominal portion in the male are four transverse rows

of powerful spines, while in the female there are only a few large

ones near the centre line, and a fewer number of smaller ones at

wide intervals reaching to the sides.

The trochanters of all the legs and the intervals between are

strongly bespined. The legs themselves have small denticu-

lations, with a short bristle on each as far as the distal end of the

tibial joint. The metatarsal joints smooth, the tarsi with short

hairs. The tarsal claws are weak; those of iii. and iv. with short

side wings springing from about the middle. On the under side

the coxe are all bordered with rows of warty prominences, the

front margin of coxe i. having a row of sharp pointed black

spines.

The margin of the genital operculum is divided into about

ten or eleven distinct scopulations or pustules, each terminated

with a bristle.

The mandibles are short and weak, and covered with short

black beady pustules.

The palpi are about the length of the carapace in the male,

48 ON SOME AUSTRALIAN OPILIONES.

strongly bespined with sharp bristly points; those of the

female are longer and rather slimmer, but equally bespined.

The measurements (in millimetres) are as follows :—

Male. Ceph. lg. 14, lat. 13; abd. Ig. 2, lat. 22; mand. 22;

palpus 34; 1st leg 43, 2nd 8, 3rd 67, 4th 83.

Female. Ceph. lg. 13, lat. 14; abd. lg. 24, lat. 2; mand, 12;

palpus 43; 1st leg 54, 2nd 92, 3rd 83, 4th 103.

The males are three from Mt. Algidus, Rakaia Gorge, and one

from Mt. Remarkables, near L. Wakatipu.

The females are one from Mt. Starve-all, near Nelson, and one

from Canterbury. All these localities are in the South Island of

New Zealand.

There can be little doubt that they are all of the same species

from the similar coloration of their parts.

EXPLANATION OF THE PLATES.

Puate I.

Fig. 1. Macropsalis chiltont Hogg. § with short mandibles. Underside showing

mouth-parts, genital operculum, and coxe.

. Macropsalis chiltoni Hogg. Profile of male.

3. Pantopsalis pococki, sp.n. (a) Dorsal view; (6) profile; (¢) Pantopsalis

wattsi, sp. n., cepbalic part of carapace, dorsal view.

4. (a) Pantopsalis halli, sp.n., 6; (6) 2; (c) genital operculum of g; (d)

genital operculum of g turned back showing organ.

. Pantopsalis grayi, sp.n. 2. (a) Ventral view of carapace; (6) mandible ;

Prats II.

Fig. 6. Trienobunus acuminatus, sp.n., 6. (a) Dorsal view of carapace; (b) ven-

tral view of carapace; (c) eye-tubercle enlarged; (d) genital operculum

turned back, showing dorsal and ventral sheaths and organ—distal end

only.

7. Trienonyx cockayni, sp.n. (a) Under side of cephalic part of 9; () under

side of cephalic part of 6; (c) genital organ of g protruded from

operculum—tront view ; (@) ditto—side view.

8. Trienonyx variegata, sp.n. (a) Dorsal view of g ; (6) ventral view of 3 ;

9. Nuncia smithi, sp.n. (a) Dorsal view of carapace of 2 ; (6) ventral view,

anterior portion of carapace; (c) tarsal claw of iv.

Prarez III.

Fig. 10. Monoxyomma hendei,sp.n. . (a) Dorsal view of carapace; (6) ventral

view of carapace ; (c) profile; (d) tarsal claw of iv.

11. Monoxyomma tuberculatum, sp. n. (a) Dorsal view of carapace ; (0) ventral

view of carapace ; (c) eye-tubercle from side.

12. Monowyomma trailli, sp.n. (a) Dorsal view of g3; (6) ventral view of

cephalic part of ¢; (c) ventral view of cephalic part of 2; (d) eye-tubercle ;

(e) iv. tarsal joint showing claws of 6; (jf) iv. tarsal jomt showing claws

of 2.

13. Algidia cuspidata, gen. et sp.n. (a) Dorsal aspect of g ; (b) dorsal aspect

of ¢ ; (c) under side of cephalic part of ¢ showing genital operculum ;

(d) under side of cephalic part of 9 showing genital operculum ; (e) tarsal

joints of iv. leg.

ON THE ENGLISH SPECIES OF RED SPIDER. 49

4, Revision of the English Species of Red Spider (Genera

Tetranychus and Oligonychus). By Stanury Hirst.

(Published by permission of the Trustees of the British Museum.)

[Received December 8, 1919: Read February 24, 1920.]

(Text-figures 1-5.)

The following note deals with the English species of mites

commonly called Red Spiders, and is almost entirely based on

material collected by the author, the coloration being described

from living specimens. <A good deal of confusion still exists in

the classification of this group—for instance, the name Tetra-

nychus telarius is not always used for the same species by recent

authors, and several distinct species (7'. tiliariuwm, T. lintearius,

T. populi, and Oligonychus quercinus) ave wrongly considered as

synonyms of that species. Three new species are described in

the present note, but one of them (7" talisiew) is undoubtedly an

intreduced form.

My best thanks are due to Lieut.-Col. Sir David Prain (Kew

Gardens), Prof. W. Bateson (John Innes Horticultural Station,

Merton), and to the authorities of the Royal Horticultural

Society's Gardens, Regent’s Park, for kindly allowing me to

collect material in the gardens under their charge.

The drawings illustrating this note have been made by Mr. Percy

Highley and Mr. A. J. Engel Terzi.

Order PROSTIGMATA.

Superfamily Trombidioidea.

Family TETRANYCHIDA!.

The two principal genera Vetranychus and Oligonychus can

be readily distinguished from one another by the following

characteristics :—

Claw of legs always simple and unmodified (never bifurcated nor

furnished with teeth); it has a slight protuberance near the base

bearing minute hairs ......... esccceeeeee, Genus Oligonychus Berlese.

Kirst claw of male either atti: in nine heetl or bifurcated. Claw

of the other legs in both sexes usually either ending in fine hairs or

foifinn Gabed Grete ete ceen certain atin snc mahaea ee Genus Tetranychus Dufour.

Genus Terranycuus Dufour, 1832.

Key to the species of Yetranychus (males) occurring in

England :—

(Terminal finger of palp minute and inconspicuous or

EOCENE eri de oA Rern econ e aaa ear ee RRM che I LA gt a eal

(erminal tingeyiwellideveloped | -:..:.... cee. -enesesee eee 4,

Terminal finger of palp absent.

All the claws of the legs bifurcated ........................ Detranychus schizopus

IL, (Zacher.

, Terminal finger present, but minute. ‘

Claw of first leg alone bifurcated

Proc. Zoou. Soc.—1920, No. Vin ae © 4

iw)

50 MR. SLANLEY HIRST ON THE

Penis strongly curved and furnished with a spine and

9. tubereletatt the end! 0 7.8ste.ce cesses eee eee eae T. crategi, sp. 0,

| Penis straight or only slightly curved. the end unarmed ......... 3.

[ Penis fairly long, the end blunt .............0..0.5..seneee eee T. populi C. L. Koch.

3.5 Penis shorter and pointed at the end....................... 1. talisie, sp. n.

L (introduced species).

(Psa sopuelal SBHHOYS Aly WO) Biavel benguaonacoacs acvece coosvavssnavaosacacad Be

‘UlCPantstwithowt baths aU, Ae Me CPR ies: kre 4G:

g Claw of first lee armed with short teeth or spines ...... T. telarius, L.

"i (Claws of both first and second leg with teeth ............ T. lintearius Dufour.

(Penis strongly sinuous. Offshoots of claw of first leg

Te ae Oe Cred: PME ES obclaceabtindosgasanadee: T. carpini Oudemans.

| Penisstraighter. Offshoots of claw of first leg stronger,

(UY spimitormy Baie carta misscasesrs a erence teen emmee nce: T. tiliarium Hermann.

1. TerraANnycHus scuizopus Zacher, 1913.

Tetranychus schizopus Zacher, Berlin, Mitt. biol. Anst. 1913,

heft 14, pp. 38 & 40, text-fig. 4.

27. salicis C. L. Koch, Deutsch. Crust., Myr., Arachn. 1838,

heft 17, no. 18.

Schizotetranychus schizopus Trigardh, Stockholm Medd. Cen-

tralanst. Forséksv. Jordbruksomr. 1915, vol. 109, p. 19.

3. Penis of the same type as in the species found on the

hawthorn (7. cratwgi), being curved when viewed laterally, and

with one of the terminal barbs greatly elongated so as to form a

spine, whilst the other is in the form of a minute inconspicuous

tubercle.

Palp. Terminal finger or cone of palp apparently absent in the

male of this species. Dorsal sensory finger well developed (see

text-fig. 3a, 1 & 2, for the explanation of these terms).

Claws of all the legs bifurcated, the two branches into which

the first leg is divided being in this sex split again for a short

distance at the tip.

@. Palp. Terminal finger of palp well developed and fairly

stout, being about as long as or longer than the neighbouring

rod-like setz. Dorsal sensory finger normal in appearance.

The claws of the legs do not end as fine hairs as in the female

of 7. telariws (and all other species of the genus seen by the

author), being divided to form two prongs or claws as in the

male, but the prongs of the first legs are entire instead of being

split again at the extreme tip as in the male sex.

Colour rather variable, greenish, pale yellow, reddish, and

orange-red specimens being met with. Small dark patches are

sometimes present on the dorsum.

Eggs sometimes pale amber in colour, sometimes reddish.

Length of male (including mandibular plate) 340 », of female

350-535 p.

Host plants. Various species of willows.

ENGLISH SPECIES OF RED SPIDER. 51

Norz.—Trigirdh has created a new genus (Schizotetranychus)

for this species, which certainly is very peculiar in having all the

claws bifid in both sexes. The penis is, however, very like that

of 7. crategi. Moreover, the male of 7. populi has the first leg

shaped like that of 7. schizopus. On the whole, I think it is not

necessary to place this species in a separate genus.

Text-figure 1.

a. Lateral view of penis of Tetranychus schizopus. b. Ditto of T. populi. & j.

Ditto of T. tiliarium. d, e, f. Ditto of 7. talisie. g. Ditto of 7. crategi.

h. Ditto of T. carpini. i. Ditto of 7. telarius. k. Ditto of Oligonychus

ununguis. 1. Ditto of O. quercinus. m. Ditto ot O. ulmi. n. Ditto of

O. simplex.

2. TETRANYCHUS CRATAGI, Sp. n.

g. Penis very like that of 7’. schizopus in shape, being strongly

curved and with a long spiniform process.at the end, also a very

minute and inconspicuous tubercle which no doubt represents

the barb of the other side.

Palp. Terminal finger in the form of a very minute and in-

conspicuous cone or tooth. Dorsal sensory finger apparently

variable in length. ,

Claw of first leg in this sex distinctly bifid, being divided

into two strong prongs; one or two frail hairs are present on

these prongs, and no doubt correspond to the spines present in

T. tiliarium.

2. Palp. Terminal finger somewhat short but very wide, being

rather conical in shape; 1t seems to be a little shorter than the

two rod-like setz near it. Dorsal sensory finger moderately

developed.

Olaws of legs ending in long hairs as in 7. telarius.

Measurements. 3, length 340; 2, 335-460 p.

Host plant. Hawthorn hedges in the outskirts of Salisbury,

Wilts.

52 MR, SUANLEY HIRST ON THE

3. TerraAnycuus Poputt C. L. Koch, 1838. |

Tetranychus popult C. L. Koch, Deutsch. Crust., Myr., Arachn.

1838, heft 17, no. 14.

T. telarius (ad part.) Canestrini, Prospett. Acarof. Ital. 1890,

p. 434; Berlese, Acari, Myr., etc. in Ital. reperta, 1889, fase. lvi.

no. 5.

Text-figure 2

a. Claws (empodia) of first leg of Tetranychus schizopus 6, lateral view of same.

b, Ditto. Ventral view. c. Claws of posterior leg of 7. schizopus. d. Ditto of

first leg of 7’. telarius 6. e. Ditto of second leg of 7. lintearius g. jf. Ditto

of first leg of T. carpini 3. g. Witto of TZ. tiliarium S. h. Ditto of

T. tiliarium §.

3. Penis rather like that of 7. tiliariwm, being only slightly

curved ; it is much longer than in 7’. talisiw, sp. n., and also

differs in having the end blunt.

Palp. Terminal finger or cone very minute as in 7’. talisie ;

dorsal sensory finger long.

First claw of fnnle ifareated exactly as in 7’. schizopus, Paice

ENGLISH SPECIES OF RED SPIDER. 53

prong being again split for a short distance at the extreme tip.

Claws of posterior legs, as in 7’. telarius, ending in hairs.

2. Palp. Terminal cone rather short but wide.

Claws of legs as in 7’. telarius.

Eggs whitish.

Measurements. $, length 3204; 2,380. Strongly chitinised

part of penis about 43, in length, but, if the more weakly

chitinised part is included. it measures altogether about 65 p.

Host plant. Lombardy poplars, Regent’s Park, London.

Text-figure 3.

a. Terminal segments of palp of T.crategi g. b. Ditto of T.schizopus 8. c. Ditto

of JT. talisie 8. d. Ditto of T. populi g. e. Ditto of T. lintearius g.

f. Ditto of T. telarius 3. g. Ditto of TP. tiliariwm g. 1. Terminal finger

or cone. 2. Dorsal sensory finger. :

Norre.—C. L. Koch described his 7. populi from Lombardy

poplars (‘‘italienischen Pappel”), so that there can be little doubt

as to the correctness of my identification of this species. The

Common Red Spider (7. telariws) is sometimes also present on

Lombardy poplars and also on ordinary poplars, but the males of

the. two species of mites are very unlike one another.

54 MR. STANLEY HIRST ON THE

4, TerrANYCHUS TALISIA, Sp. n.

3. Penis short and straight, the terminal half slender and

ending in a sharp point ; there is no trace of barb at the end; it

is very much shorter than that of 7. tiliarium or 7. popult.

Palp. Terminal finger or cone very minute, but usually longer -

than wide, being conical and pointed. Dorsal sensory finger short

and slender, but the width seems to vary in different specimens.

Text-figure 4.

a. Terminal segments of palp of 7. crategi 9. 6. Ditto of T. populi 2. c. Ditto

of 7. talisie 2. d. Ditto of 7. lintearius P. e. Ditto of T. tiliariwm 2»

f. Ditto of 7. telarius 2.

Claw of first leg bifid when seen from above; when viewed

laterally, each fork bears two hairs (one dorsal, the other ventral) : a

as in 7’. carpini, but they are still more inconspicuous and difficult

to see, being very fine. The claws of the three posterior legs end

in hairs as in 7’. telarius.

@. Palp. Terminal finger or cone fairly well developed and

rather stout, being about equal to the length of the neighbouring

rod-like sete. Dorsal sensory finger slightly club-shaped.

ENGLISH SPECIES OF RED SPIDER, 5D

Colour. Body pale yellowish green; sometimes minute dark

specks, arranged in four small clusters, are present at the sides

and posterior end of the dorsum.

Measurements. 3, length 220-250; 2, 340-380 p.

Host plant. South American tree (Zalisia princeps), growing in

Tropical Greenhouse, Kew Gardens.

5. TeErRANYCHUS TELARIUS L.

Acarus telarius (ad part.) Linneus, Syst. Nat. 1758, Ed. 10,

p- 616.

Tetranychus telarius C. L. Koch, Deutsch. Crust., Myr., Arachn.

1838, heft 17, no. 12.

T. russeolus C. L. Koch, op. cit. heft 17, no. 15.

T’. urtice C. L. Koch, op. cit. 1835, heft 1, no. 10.

Acarus telarius Boisduval, Ent. Hort. 1867, pp. 82-84, text-

fig. 4.

A. cucumeris Boisduval, op. cit. p. 84.

A. vitis Boisduval, op. cit. pp. 92-93.

Tetranychus althee Von Hanstein, Zeitschr. wiss. Zool. 1901,

Ixx. p. 74.

1’. telariws Ewing, Ann. Soc. Ent. America, 1913, vi. pp. 455—

457, text-fig. 1.

7’. althee Trigirdh, Stockholm Medd. Centralanst. Forsoksv.

Jordbruksomr. 1915, vol. 109, pp. 86-40 & 57.

3. Penis strongly curved near the end, and the tip furnished

with two minute but distinct barbs.

Palp. Terminal finger of moderate length (about as long as the

two rod-like sete situated near it). Dorsal sensory finger varying

slightly, but usually short and slender (sometimes, however, it is

somewhat club-shaped).

Claw of first leg in this sex with a little dorsal offshoot, the

rest of the claw being divided into six short slender forks or teeth

(three on each side).

Q. Palp. Terminal finger fairly well developed, but not very

long. Dorsal sensory finger also fairly well developed.

Colour usually pale, being either whitish, greenish, or yellowish

in tint, with dark lateral spots or patches varying in extent,

sometimes quite extensive. More rarely pink or reddish indi-

viduals are met with.

This species occurs on many plants, and was confused by

Linneus, and later by Hermann, with the form occurring on the

lime-tree (the latter creates the name tiliarvwm, but says that the

species occurring on the lime-tree is also found on Althea,

a typical plant host for 7’. telarius proper). This mistake is

followed by many other authors, including recent authors of great

repute, and several other quite distinct species—viz., 7’. lintearius

Dufour (from gorse), 7’. populi C. L. Koch (from poplar trees),

and the species living on the oak (Oligonychus quercinus)—have

also wrongly been regarded as synonyms of 7’. telarius.

56 MR. STANLEY HIRST ON THE

T. telarius is our Common Red Spider, being found on many

plants, both in the open and in greenhouses, It is especially fond

of hollyhocks (Althea), but attacks numerous other cultivated

flowers and also weeds. It often infests strawberries, beans, mint,

peaches, cucumbers, and melons (in hothouses), also grape-vines.

The hop-gardens in Kent and elsewhere are often greatly

damaged by this pest. It sometimes attacks bushes and trees—

for instance, elders, rose-trees, laburnum, poplars, ete.

6. TETRANYCHUS LINTEARIUS Duf., 1832.

Tetranychus lintearius L. Dufour, 1832, Ann. Sci. Nat. (1)

Xxv. pp. 276-283, pl. ix. figs. 4 & 5.

1’. lintearius Lucas, Ann. Soc. Ent. France, 1868, (4) vii.

pp. 741-743.

7’. telarius (ad part.) Canestrini, Prospett. Acarof. Ital. 1890,

iv. p. 434.

T. telarius (ad part.) Berlese, Acari etc. in Ital. reperta, 1889,

fase. lvi. no. 5.

T. telarius Berlese, Gli Insetti, 1912, 11. p. 95.

6. Penis very like that of 7’. telariws, but wider and shorter.

Palp. Terminal peg-shaped, sensory bristle comparatively short

and of moderate width. Dorsal sensory finger short and club-

shaped.

Legs. Both the first and second Jegs of the male have the claw

modified so as to form a kind of comb of minute teeth, whereas

in 7’. telarius the claw of the second leg ends in the usual six

fine hairs.

@. Palp. Terminal finger rather short and wide, being shorter

than in 7’. telarius. Dorsal sensory finger rather short but

not slender.

Measurements. 2 450-530 mp.

Host plant. Gorse (Ulex ewropeus}; many hundreds of speci-

mens collected by the author from infected bushes on Malvern

Hills in 1917; unfortunately only one male example was present

in this material. The masses of white web formed by this species

are very obvious, and at once attract attention. Considerable

damage is done to the gorse by this mite in certain areas on the

Malvern Hills, and it is sometimes necessary to fire the bushes to

get rid of it.

7. TETRANYCHUS GARPINI Oudemans.

3. Penis very like that of 7’. tiliariwm, but always strongly

sinuous, instead of being practically straight as in that species.

Palp. Terminal finger very like that of 7. taliariwm, being long

and slerder. Dorsal sensory finger very slender.

Claw of first leg bifid; each of the two prongs consists of a

fairly strong middle tooth or spine, from which spring a dorsal

and a ventral hair or seta (in 7. tiliariwm the dorsal and ventral

offshoots are much stronger, being spiniform like the middle one).

ENGLISH SPECIES OF RED SPIDER. 57

@. Palp. Terminal finger very like that of 7’. taiarium, being

well developed and of moderate stoutness. Dorsal sensory finger

quite slender.

Colour pale greenish or yellowish, sometimes with dark lateral

spots.

Measurements. ¢, length 215; 2, 340 p.

Host plant. Hornbeam (Carpinus betwluws) ; a number of speci-

mens from a tree of this species growing in London.

8. TETRANYCHUS TILIARIUM (Herm.), Koch 1838.

Trombidium tiliariuwm (ad part.) Hermann, Mem. Apt. 1804,

pp. 42, 43, pl. i. fig. 12.

“3 telarius (ad part.) Hermann, op. cit. pp. 40, 41.

wa socium Hermann, op. cit. p. 43, pl. ii. fig. 13.

Tetranychus tiliariwm Koch, Deutsch. Crust., Myr., Arachn.

1838, heft 17, no. 12.

5 socius Koch, op. cit. heft 17, no. 16.

as telarius (ad part.) Canestrini, Prospett. Acarof.

Ital. 1890, iv. p. 434.

- telarius (ad part.) Berlese, Acari etc. in Ital.

reperta, 1889, fase. lvi. no. 5.

ts telarius Von Hanstein, Zeitschr. wiss. Zool. 1901,

Ixoxe Wand 4.

3. Penis very long and slender, being in the form of an almost

straight (only slightly sinuous) style.

Palp. Terminal finger long and slender, being more slender

than in the female. Dorsal sensory finger short but slender.

Claw of first leg bifurcated, each fork bearing three minute but

sharp denticles ; claws of the other legs ending, as in 7’ telarius,

in six fine hairs.

2. Palp. Terminal finger well developed, being fairly long and

of moderate width. Dorsal sensory finger short and slender.

Colour. Pale green, yellowish, or whitish, with minute dark

markings, chiefly lateral in position, but occasionally there are

one or two minute dark specks in the centre of the dorsum

as well.

Egg whitish.

Measurements. 3, length 260-305 w; 2, length 420-490 p:

penis about 57 pw in length.

Host plant. Lime-tree (Tilia ewropea). I have also found

specimens of this mite on a hawthorn and on a willow, also in

abundance on hazels near Exeter.

Nove.—Hermann was the first author to use the name tiliariwm

as a specific name, but he mentions 1t as occurring also on Alcea

[Althea] rosea, and his specimens from this plant are probably

referable to 7’. telarius. Fortunately, C. L. Koch limits the

species to examples from lime-trees only, and there is a good deal

of reason in this, for it occurs on very few other plants.

58 MR. STANLEY HIRST ON THE

Genus Oxticonycuus* Berlese, 1886.

It is difficult to separate the species of Oligonychus by means

ofakey. The principal differences between the species of this

genus are in the structure of the penis and terminal finger of the

palp (see accompanying figures).

1. OtigonycHus utmi C. L. Koch, 1835.

Tetranychus ulmi OC. L. Koch, Deutsch. Crust., Myr., Arachn.

18385, heft 1, no. 11.

T. pilosus Canestrini & Fanzago, Atti Soe. Ven. Trent. 1876,

v. pp. 133-134.

1’. ulmi Berlese, Acari dann. piant. coltiv. 1886, p. 22.

T. pilosws Berlese, Acari ete. in Ital. reperta, 1889, fase. lvi,

no. 6.

Paratetranychus pilosus Zacher, Berlin Mitt. biol. Anst. 1913,

heft 14, pp. 38-39, text-fig. 1.

P. pilosus Trigirdh, Stockholm Medd. Centralanst. Forséksv.

Jordbruksomr. 1915, vol. 109, no. 20, pp. 21-29 ete., text-figs.

3. Penis strongly curved, being shaped as figured.

Palp. Terminal finger short, being less than half the length of

the two neighbouring rod-like sete near it. Dorsal sensory finger

also short.

2. Palp. Terminal finger quite short but very stout, being

much shorter than the rod-like sete near it. Dorsal sensory

finger short and slender.

Another characteristic point is that the hairs of the body in

this species spring from slight tubercles.

Oolour. Dorsum dark reddish at the sides, and:usually with a

paler reddish central band, the anterior end of the body also pale

red. Very minute w hitish specks arranged in longitudinal

series (one at the base of each hair) are also present on the

dorsum, the hairs themselves being pale; appendages flavous.

Some specimens are dark (almost blackish), brown, and the central

pale reddish band may be absent.

Eggs reddish.

Measurements. 3, length 260-310 w; 9, 380-470 p.

Host plants, etc. This species is frequently met with on elms

in this country, causing a characteristic discoloration of the

leaves. It is obviously the species described by C. L. Koch

under the name wlmi, as being not uncommon on that tree in

Regensburg. His figure is quite good and recognisable, and it

is a great pity that the later name pilosws has been used instead

of his name. The species sometimes attacks roses, especially

rambler roses, discolouring the leaves greatly and obviously

weakening the plants. Various fruit trees—such as pear-trees,

apple-trees, plum-trees, etc.—are also affected by this species, but

the damage is usually very slight. It also occurs on the service

* The name Paratetranychus has been used by recent authors for this genus,

but, as Banks points out (Ent. News Philad. 1917, vol. xxviii. p. 197), it should

be replaced by the earlier,.name Oligonychus.

ENGLISH SPECIES OF RED SPIDER. 59

tree, horse~chestnut, copper-beech, Scoteb Jaburnum (Kew

Gardens), and American flowering currant. Although usually

found in the open, in one case I discovered acacias (Acacia lon-

gifolia etc.) and a Sapindus saponaria under glass infested with

numerous specimens of this mite.

”

Text-figure 5.

a. Terminal part of palp of Oligonychus quercinus 2. b. Ditto of O. ulmi g.

ce. Ditto of O. simplex g. d. Ditto of O.ununguis . e. Ditto of O. quer-

cinus $. f- Ditto of O. ulmi 9. g. Ditto of O. simplex 2. h. Ditto of

O. ununguis &.

2. OLIGONYCHUS UNUNGUTS Jacobi, 1905.

3. Penis very strongly curved, being very like that of O. ulmi

in shape.

Palp. Terminal finger small, but much longer than wide; it

is shorter than the dorsal sensory finger.

2. Palp. Terminal finger very like that of O. ulmi, being

short but not so wide and almost truncate when viewed laterally,

whereas in O. ulmi it is somewhat narrowed or compressed near

the base, the distal half being wider, so that it is knob-shaped.

Colour. This species is deeply pigmented, the abdomen being

blackish, but there is usually a pale brownish central streak

posteriorly ; cephalothorax ochraceous or red; legs ochraceous.

Measurements. 3, length 260 4; 9, 370 pw.

Host plant. Pinus sylvestris ; Oxshott, Surrey.

3. OLIGONYCHUS QUERCINUS Berlese 2, 1886.

Tetranychus quercinus Berlese, Acari dann. piant. coltiv.,

Padua, 1886, p. 23.

T. telarius Berlese, Acari etc. in Ital. reperta, Ordo Prostig-

mata, p. 58.

60 ON THE ENGLISH SPECIES OF RED SPIDER.

2 Tetranychus yothersi McGregor, Ann, Soc. Ent. Amer. 1914,

vii. pp. 855-357, pl. xliii. figs. 1-8.

dg. Penis very like that of O. simplex Banks (from cate-palms),

being small and curved at the end.

Palp. Terminal finger absent or very minute and inconspicuous.

Dorsal sensory finger short and very siender.

2. Palp. Terminal finger not nearly so wide as in O. ulmi,

being short but only of moderate width ; sometimes it is slightly

clavate. Dorsal sensory finger quite short and slender.

Colour whitish or pale brownish with black markings.

Eggs variable in colour, being either whitish, pale yellow, or

reddish.

Measurements. 3, length 220 uw; 2, 270-310 pm.

Host plants. Oak-trees.

Nore.—Targioni Tozzetti seems to have been the first to observe

Tetranychus occurring on oak-trees, but he does not give the

species a name.

In his ‘ Acari dannosi alle piante coltivate’ Berlese copies

Tozzetti’s brief descriptions, and gives the name quercinus to this

mite, and mentions also the ‘* Tetranico verde del leccio ” (Z. vires-

cens), Which is probably the same species. Oligonychus brevipodus.

also from the evergreen-oak, seems to be another synonym of this

species. Both Canestrini and Berlese in later publications

regards this species as a synonym of Jetranychus telarius. The

specimens found on oaks in this country (at Wimbledon Commen

and Kew) belong, however, to the very distinct species of Parete-

tranychus (= Oligonychus) described above, which is easily recog-

nised by the structure of the palp.

I take the opportunity to include the diagnosis of a species of

Oligonychus from date-palms in Mesopotamia :—

4, OLIGONYCHUS SIMPLEX Banks @, 1914.

2 Tetranychus simplex Banks, J. Ent. Zool. Claremont, Cal.

GME sale 70s DUG

3. Penis. Chitinsed part of penis very short, and it terminates

as a very minute curved hook.

Palp. Terminal finger very like that present in O. ulmi, being

rather short, but it is slightly blunter. Dorsal sensory finger

spindle-shaped, being rather short and slender, Spine on upper

surface of palp borne on a distinet angular projection.

@. Palp. Terminal finger more slender than in O. ulmi, but

still it is fairly stout. Dorsal sensory finger fairly long.

Measurements. 3, length 275-290 w; 2, 280-385 p.

Host plant. Date-palm, Basra, Mesopotamia ; numerous speci-

mens collected by Capt. P. A. Buxton.

P.Z.S.1920. BHATTACHARYA, Pl. 1.

Bale & Danielsson, Ltd

AORTIC LIGAMENT IN INDIAN FISHES.

P.Z.S.1920, BHATTACHARYA, PI. II.

Bale & Danielsson, td

AORTIC LIGAMENT IN INDIAN FISHES.

ON THE AORTIC LIGAMENT IN INDIAN FISHES. 61

5. On the Aortie Ligament in Indian Fishes. By D. R.

Buarracuarya, M.Sc., Zoological Department, The

Muir Central College, Allahabad, India, U.P.*

{Received December 9, 1919: Read February 24, 1920.]

(Plates I. & I1.t and Text-figures 1—5.)

ConTENTS. Page

Tere SHH OLOGY SL ONIY Tlaveea cane heute Se ReeMeBecoMeatk Moat ae wa nose dododeede ta Ol!

Il. The Aortic Ligament in Pseudeutropius garua taken as

a Type .. s3 soe | (OH

the igoasition ond i iectmnent: Re the earner Sin (GH

2. The Gross Structure and Histology of the Ligament... 64

JOD Wine Ipeenanerniy mA OYA INISMVES Son cee ash ane con coooue ace ceaneeossese 67

IV. The Occurrence and Absence of the Ligament in various

V. Some Suggestions concerning Ane Role aah Relationships

of the Aortic Ligament ........... eee ste OO)

Appendix: Methods of Preparation of Maia A ae ei 70

1. Introduction.

The existence of a longitudinal ligament, evidently closely con-

nected with the aorta, and extending over its entire length, was

first noted by me in the Siluroid fish, Psewdewtropius garua,

the year 1916. It was not, however, until some months later,

when Dr. W. N. F. Woodland, who had independently observed the

same ligament in the same fish, and at the same time chanced to

alight upon Mr. Burne’s Note ‘‘On Elastic Mechanisms in Fishes

and a Snake” (Proc. Zool. Soc. London, 1909), suggested to me

that the presence of a ligament actually inside the aorta was

sufficiently remarkable to warrant further enquiry, that I seriously

took up the work, and the present paper records my observations

of the dorsal aorta in over 80 species of fishes, both marine and

fresh-water. The short paper by Burnet constitutes the whole

of the literature on the subject known to me, and I have been

unable to discover if any other exists. Burne found that an

elastic ligament exists in the dorsal aorta of Clupea alosa, and he

termed this ligament the ‘“ hgamentum longitudinale ventrale.”

He did not, in his Note, go into any details on the subject, and

only casually mentions its structure and, according to him, its

probable function. His conclusions, however, regarding the

function of the ligament seem to me to be doubtful. I may also

emphasize in this place that this aortic ligament (as I prefer to

call it) is not to be confused with the better known, because more

* Communicated by W. N. F. Woopuanp, F.ZS.

+ For explanation of the Plates see p. 74.

£ Burne, R. H., ‘* On Elastic Mechanisms in Fishes and a Snake,” Proc. Zool. Soc.

London, 1909, pp. 201-203.

62 MR. D. R. BHATTACHARYA ON THE

conspicuous, dorsal elastic ligament (“ligamentum longitudinale

superius ”), which runs longitudinally in the vertebral column at

the bases of the neural spines and above the spinal canal.

Before proceeding further, I wish to offer my sincere thanks to

Dr. Woodland for suggestions offered during the progress of my

work, and for assistance in the writing up of this paper for

publication, especially in connection with Section V.

II. The Aortic Ligament in Pseudeutropius garua taken

as a Type.

I have selected the aortic ligament of Psewdeutropius garua as a

type for detailed description because this is well developed in this

species, because this fish is plentiful in the local bazaar, and

because the ossification of the vertebral column in this fish is not

so pronounced as in many other species, and so there is less

hindrance to proper decalcification of the vertebral column for

section-cutting purposes.

1. The Disposition and Attachments of the Ligament.

The aortic ligament in this fish is a compact band of highly

elastic tissue running longitudinally izside the dorsal aorta. In

its fully formed condition, it appears to hang down into the

cavity of the aorta, suspended by a fold from the median dorsal

wall of the aorta (Plate II. fig. 7)—in fact, the suspending fold

surrounds the ligament in much the same way as the mesentery

surrounds the gut (text-fig. 4). Anteriorly, the major portion of

the ligament ends beneath the basioccipital bone. ‘The ligament

pierces through the dorsal wall of the aorta, just at the point where

the most posterior efferent branchial vessels meet, and breaks up

into bundles of elastic fibres which become severally attached to

the basioccipital bone and give the whole structure a fan-shaped

appearance (text-fig. 1, a,f's.). At this place there is generally

a pad of cartilaginous tissue developed. A small portion of the

ligament, however, after perforating the dorsal wall of the aorta,

runs anteriorly close beneath the basioccipital in the median line,

and, after dividing into two, runs beneath the parasphenoid bone

and becomes attached to the vomer by a series of branches. The

dissection here had to be done under the binocular microscope as

the branches were exceedingly fine and delicate.

At the posterior end the ligament is attached to the hypural

bone, or rather the bone which represents the fused urostyle and

the hypural, close to the point of junction of the latter with the

last vertebra. There is a definite concavity in the bone here, in

which cartilaginous tissue 1s developed. The ligament here also

spreads out and breaks up into smaller bundles of fibres, which

acquire a strong attachment with the bone (text-fig. 1, pfis.). A

small portion of the ligament runs posteriorly for a short distance

and becomes attached to the middle region of the hypural bone.

tow eimai

ie opiate Stee +

netlist

AORTIC LIGAMENT IN INDIAN FISHES. 63

Text-figure 1.

t--------l gor,

Peudeutropius garua (nat. size). The dorsal aorta has been opened from the

ventral side to show the disposition of the ligament.

Vom., vomer ; lig.a.’, the branches of the anterior portion of the ligament which

become attached to the vomer; par., parasphenoid ; lig.a.’, the two main

branches of the anterior portion of the ligament; bas., basioccipital ; Zig.a.,

the anterior portion of the ligament lying outside the dorsal aorta; lig.per.,

the place where the ligament perforates through the dorsal wall of the aorta 3

eff.a., efferent branchial vessels; a,f-s., anterior fan-shaped structure; s..c.,

round masses of connective tissue or sub-vertebral masses; i.v.c., sub-inter~

vertebral masses of connective tissue ; lig., aortic ligament ; aor., aorta; ver.,

vertebra ; p.f.s., posterior fan-shaped structure ; hy., hypural bone.

64 MR. D. R. BHATTACHARYA ON THE

Before it acquires an attachment with the bone, the ligament

pierces through the wall of the aorta close to the junction of the

last vertebra with the urostyle.

The aortic ligament, as already mentioned, has no connection

whatever with the well-known ‘ligamentum longitudinale supe-

rius” (Hertwig*) which lies above the spinal cord, is attached

anteriorly to the exoccipital bones and posteriorly to the urostyle,

and is flattened dorso-ventrally (text-fig. 2, D.lig.).

Text-figure 2.

D. fig.

Transverse section through the caudal region of Hutropiichthys vacha

(X 14).

D.lig., dorsal ligament; sp.c., spinal cord; e¢.¢., connective tissue; lig., aortic

ligament ; cav.a., cavity of aorta.

2. The Gross Structure and Histology of the Ligament.

Text-figures 3 and 4 will show that in transverse section the

ligament looks more or less oval in shape, being thinner dorso-

ventrally than from side to side. On opening the aorta from the

ventral side and soaking out the blood with pieces of blotting-

paper, the ligamert looks like a white flat sheet of elastic tissue,

* Dr. O. Hertwig, ‘ Handbuch der Entwickelungslehre der Wirbeltiere,’ p. 450.

AORTIC LIGAMENT IN INDIAN FISHES. 65

resting on—or, more correctly, supported by—regularly arranged

white round masses of connective tissue (text-fig. 1, lig.). At

first sight these round masses look like adipose tissue, but on

closer inspection they are found to be aggregations of adventitia

tissue which form the outermost layer of the wall of the aorta.

Text-figure 3.

Text-figure 4.

Text-fig. 3.— Pseudeutropius garua (X 15). Transverse section of the aorta through

the sub-vertebral region (A—B) of text-fig, 1.

d.w., dorsal wall of aorta; con., sub-vertebral mass of connective tissue; Zig., aortic

ligament; cav., cavity of aorta; iné., intima.

Text-fig. 4.— Pseudeutropius garua (X15). Transverse section of the aorta through

the sub-intervertebral region (C-D) of text-fig. 1.

d.w., dorsal wall of the aorta; con.’, sub-intervertebral mass of connective tissue ;

n., neck or the thin suspensory fold of the ligament; Jig., ligament.

Kach of these round masses of connective tissue occurs beneath

the body of each vertebra and fits into a depression in the

centrum on its ventral side. In certain fishes, as in Labeo rohita

and Catla buchanani, these depressions in the centrum acquire

the form of deep notches; and in such cases the masses of con-

nective tissue have to be carefully scooped out from the cavities

Proc. Zoou. Soc.—1920, No. V. Ip

66 MR. D. R. BHATTACHARYA ON THE

in order to take out the aorta intact for histological purposes.

These masses acquire a strong attachment with the connective

tissue of the vertebral column. Each of the big masses of con-

nective tissue lies beneath the body of a vertebra, and I have,

therefore, called it ‘“sub-vertebral mass,” in contradistinction to

the thinner layer of adventitia which connects the adjoining big

masses and which I have referred to as ‘‘ sub-intervertebral

mass”? because it lies beneath the intervertebral region of the

adjoining vertebre (text-figs. 8 and 4, con.,con.). These ‘‘ ver-

tebral masses” of connective tissue are segmentally arranged in

relation to the vertebre, for they are found beneath each vertebra

from the first to the last. These masses seem to serve as a

cushion or pad for the strong and tough elastic ligament. During

the lateral flexions of the body, in which the aorta also takes

part, the ligament though itself stationary is displaced from its

median position inside the aorta. But it is never allowed to go

beyond the range of these masses, and probably it never actually

touches the wall of the aorta. Thus the ligament is protected

from undue strain which would result from direct contact with

the vertebral column, since the aorta lies in close contact with

its ventral surface. In the ‘“sub-intervertebral” region the

adventitia layer of the ligament and the dorsal wall of the aorta

are much thinner (Plate I. fig. 2). But the aorta in this region

is more strongly attached to the connective tissue of the vertebral

column than in the “ sub-vertebral” region.

Microscopie examination of the transverse sections (text-fig. 4)

shows that the ligament lies in the middle coat or media of the

dorsal wall of the aorta, the intima being reflected over the prc-

jection of the ligament into the aorta lumen. Muscle-fibres are

practically altogether absent from the dorsal region of the aorta

wall (though smooth muscle-fibres are present in the lateral and

ventral regions), and it is noteworthy that it is in this position

that the longitudinally-elastic ligament is situated. Outside the

ligamentous layer is the thick and stout adventitia layer, though

in between the two a thin layer of muscular fibres, elastic fibres,

and connective-tissue cells may generally be found (Plate II.

ise).

On a more minute examination of thin longitudinal and trans-

verse sections, under high power, the ligament seems to consist of

bundles or groups of elastic fibres (Plate II. fig. 11, bun.), both of

the finer and coarser varieties. The fibres branch and unite and

run longitudinally throughout the length of the aorta. Owing

to their extreme elasticity and to the fact that the fibres are so

thickly clustered as to give the impression that they are running

in definite bundles, they acquire a wavy shape towards the exposed

sides and especially near the cut ends of the ligament (Plate I.

fig. 2, lig.). In a transverse section the bundles of the elastic

fibres of the ligament may generally be seen to lie transversely in

the lower or ventral portion of the ligament and more or less

vertically in the upper or dorsal portion of the ligament (Plate II.

par Saga ti

AORTIC LIGAMENT IN INDIAN FISHES. 67

fig. 11, lig.). This arrangement seems to be fairly constant

throughout the length of the ligament.

Ill. The Ligament in other Fishes.

The ligament in a transverse section looks either circular (e. g.

in Notopterus chitala, Labeo diplostomus, and Macrones aor) or

oval (e. g. in Hutropuchthys vacha, Rita buchanani, and Engraulis

telara) or crescent-shaped (e. gy. in Chatoessus manminna, Clupea

sindensis Clupea ilisha). The ‘suspensory fold of the ligament

consists of a double layer of intima with a mass of connective

tissue in between. This mass of connective tissue may be very

thick, so as to give rise to round or oval masses—the ‘sub-

vertebral” masses (e. g. in Pseudeutropius garua, Hutropiichthys

vacha), or it may be thin (¢. g. in Callichrows macrophthalmus,

Callichrous pabda, Barbus sarana). Sometimes the suspending

fold is so thin and elongated that in a transverse section it looks

like the “neck” of the ligament (text-fig. 4, 2.). In some cases,

the suspending fold is so deep in the trunk- -region that the

ligament comes to lie quite close to the ventral wall of the aorta.

Ta such cases (text-fig. 3) in the ‘“sub-vertebral” region the

ligament with its suspensory fold and connective tissue fills up

more than half the cavity of the aorta. The “neck” of the

ligament is thinner in the ‘‘ sub-intervertebral” region than in

the ‘‘sub-vertebral ” region (text-fig. 4). In some cases there is

no suspending fold in the “sub-intervertebral” region and the

ligament is developed in the wall of the aorta, and in a transverse

section looks like a small protuberance (e. g. in Rita buchanani,

Plate I. fig. 4, lag.). Dorsally to the suspensory fold or the neck

of the ligament there is always either a thinner or more generally

a thicker layer of connective tissue or adventitia. In some cases

(e.g. in Chatoessus manminna and Rita buchanani) a cartilaginous

pad is developed in this layer in the “ sub-vertebral” region

(Plate I. fig. 3).

In the majority of cases the ligament ends in the basioccipital

bone. In some the greater portion of the ligament becomes

attached to the basioccipital bone, but a small branch pierces

through the dorsal wall of the most anterior region of the aorta

and runs forwards to end in the vomer or parasphenoid bone

(e. g. in Hutropuchthys vacha, Pseudeutropius garua, Clupea ilisha,

Silundia gangetica). In others (e.g. in Macrones seenghala and

Catla buchanant) the ligament pierces through the dorsal wall of

the aorta a short distance behind its origin, and is attached to

the ventral side of the fused mass of anterior vertebre. The

aorta here runs in close contact with and inside a bony groove

formed on the ventral side of the fused mass of anterior

vertebrae

In Wallago attu, in which the ligament is very well developed,

I have worked out in detail both by dissection under the binocular

microscope and by microtome sections the anterior termination of

68 MR, D. R. BHATTACHARYA ON THE

the ligament. Fig. 7 (Plate IJ.) represents diagrammatically the

exact disposition of the ligament in the anterior trunk-region and

beneath the basioceipital bone. Tracing the ligament in the

trunk-region from the posterior to the amber ior end, we find that

behind the region of the third vertebra the ligament is well deve-

ae and hangs deep down into the cavity of “the aorta (Plate II.

HVA) coment “the region of the third vertebra the lhgament

tes not hang so deep down, but gradually ascends towards the

dorsal wall of the aorta, and consequently the suspensory fold

becomes narrower dorso ventrally, though thicker from side to

side (Plate I. fig. 13). Just beneath the region where the second

vertebra ends, the ligament enters the wall of the aorta (Plate IT.

fig. 12). Here, in the outer dorsal wall of the aorta (in the

adventitia layer), a thick mass of cartilaginous tissue is developed

(Plate IT. fig. 11, ¢.), which, as I have already said, acts like a pad

or cushion, and is always situated close behind where the hgament

is attached to the bone. Figs. 9, 10, and 11 (Plate IT.) show the

ligament running in the wall of the aorta and gradually ascending

till we find that the ligament actually perforates the dorsal wall

of the aorta and becomes attached to the bone (Plate II. figs. 7

and 8, lig per). A branch, however, runs through the wall of the

aorta and pierces through its most anterior region (fig. 7, lig.a.')

close to the place where the posterior efferent br. ae vessels meet.

Tt then runs close beneath the basioccipital and is finally attached

to the vomer as in Pseudeutropius garua. In Silundia gangetica

(Plate I. fig. 5) the ligament pierces through the dorsal wall of

the aorta a short distance behind its origin. The major portion

of the ligament becomes attached to the basioccipital bone, and

a branch runs forwards beneath the basioccipital and the para-

sphenoid. Towards its anterior end the ligament breaks up in a

fan-like manner and forms a thin sheet of elastic and connective

tissue (fig. 5, @. ). Posteriorly the ligament in all cases ends

either in the last vertebra or in the fused uvostyle and hypural

bone.

TV. The Occurrence and Absence of the Ligament in various

Fishes.

The aortic ligament is by no means a structure of universal

occurrence. It is confined to some of the Teleost fishes alone,

being totally absent from the Cyclostomes, Hlasmobranchs,

Amphibians, Reptiles, Birds, and Mammals. In the Teleostomes

it seems to be more of a generic feature than of a family one. The

tabular statement (pp. 70- 72) shows that, whereas it is present

in the various species of a genus, it Is absent from other genera of

the same family. I have adopted here Goodrich's classification

chiefly, as given in Lankester’s ‘A Treatise on Zoology,’ but I

have also followed in some places Day’s system of classification

(‘ Fauna of British India, Fishes,’ vols. 1, and il.).

The ligament is present in all the Clupeidee and Cyprinide that

I have examined, but in the large family of Siluride it seems to

AORTIC LIGAMENT IN INDIAN FISHES, 69

be absent from a few genera, though present and well-developed

in the majority.

V. Some Suggestions concerning the Role and Relationships

of the Aortic Ligament.

If a fish possessing an aortic ligament be completely stripped

of its muscles, the vertebral column (with its dorsal elastic

ligament and the attached aorta) becomes thrown into a series of

serpentine curves. If now the aortic ligament be removed, the

curves disappear. This is proof (1) that the lateral muscles

keep the aortic ligament on the stretch; (2) that the aortic

ligament, which is not closely adherent along its course to the

vertebral column, is antagonistic in action to the dorsal elastic

ligament, the latter tending to keep straight the vertebral

column, the former tending to curve it; (3) that the aortic

ligament is more powerful than the dorsal elastic ligament ,

(4) that when the vertebral column becomes flexed by muscular

action, the dorsal elastic ligament becomes stretched and the

aortic ligament slackened. These obvious conclusions, however,

do not assist us much in comprehending the function of the

aortic ligament. What this function can be, it is at present

only possible to guess at. It would seem that it must be different

to that of Reissner’s fibre in the nerve-cord and of the dorsal

elastic ligament, if only because of its different situation and

attachments. It would also appear that, in view of the fact that

the aorta in these, as in other fishes, assumes the functions of

a heart (the actual heart being separated from the aorta by the

gill capillary system), the aortic ligament must either act as an

auriculo-ventricular valve or actively assist in the propulsion of

blood along the aorta. Since it is difficult to conceive in what

way any alteration of position or tension of the vertical curtain

of tissue formed by the ligament and its suspensory fold in the

aorta can enforce the contractions of the aortic wall, the only

feasible explanation of the aortic ligament is to suppose that it

acts as a longitudinal valve preventing forward regurgitation of

blood. According to Burne, Stewart suggested that since the

aortic ligament, owing to its stoutness and tension, remains

straight during the lateral flexions of the body, it, with its

suspensory fold, must act as a diagonal curtain which sweeps the

blood posteriorly in the aorta during the swimming movements

of the fish. The flexions of the body originating anteriorly and

passing posteriorly, successive portions of the aortic ligament

curtain will assume a diagonal position in the aorta as the flexion

proceeds posteriorly, and this process, it was suggested, may aid

in the propulsion of the blood posteriorly. Since, however, the

fold. hangs loosely in the aorta and never completely closes it,

it seems more likely that this diagonal curtain will be of more

use as a valve which to some extent prevents regurgitation of the

blood forwards, than as a mechanism which would have to be of

MR. D. R. BHATTACHARYA ON THE

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AORTIC LIGAMENT IN INDIAN FISHES. 73

powerful construction to be at all effective. For the suspended

ligament to be of any use as a diagonal curtain, it is evident that

the aorta, as part of the body, will have to undergo flexion,

the curtain remaining taut and straight, but, curiously enough,

comparison of the statements in the tables (pp. 70-72) shows that

it is just in those fishes in which, owing to a deep thick body

and envelopment by deeply grooved vertebral and stout hemal

arches, the aorta cannot experience much lateral flexion, that the

ligament exists,

Text-figure 5.

r--\ -“g.

dor.

A diagrammatic representation of the position of the ligament inside

the aorta during the lateral flexions of the body.

aor., aorta; lig., ligament.

On the other hand, the aortic ligament is absent or but feebly

developed in all, or most of, those fishes in which the body is

slender, and the median (especially the median dorsal) fins

elongated in form, or the caudal region very much elongated

narrow and tapering), 7. e., in just those fishes in which flexion

of the body and therefore of the aorta must be most marked.

This correlation of facts, founded on my examination of over

80 species of fishes, certainly does not appear to favour the

hypothesis as to the mode of action of the aortic ligament

suggested by Professor Stewart.

Since I have no theory of my own to offer concerning the

raison @étre of the aortic ligament, I will merely add that it is

74 MR. D. R. BHATTACHARYA ON THE

evident that though the ligament is well developed in the Siluridze

(a primitive group), yet it cannot be regarded as a primitive

structure, seeing that it is not developed in many other primitive

groups of fishes.

As regards the development of the ligament, [ have not been

able to study this for lack of material. It is, however, evident

that the ligament must arise as a special development of the

inner dorsal wall of the aorta (Plate IT. figs. 12, 13, 14), the elastic

fibres of the middle and inner coats of the aorta becoming

aggregated to form the continuous elastic ligament. This view

is borne out by the fact that in the “sub-intervertebral ” regions

of the aorta in Pita buchanani (Plate I. fig. 4), and also at the

anterior and posterior ends of the aorta, the ligament pierces

through its dorsal wall and runs through the middle coat of the

aorta for some distance.

Appendix: Methods of Preparation of Material.

The marine specimens were fixed in 4 per cent. formalin, and

the fresh-water specimens, which were available locally, were

fixed in Potassium bichromate solution. The smaller specimens

were decalcified in a solution of 3 per cent. nitric acid in 70 per

cent. aleohol, which was changed every alternate day for from 3-5

weeks. Portions of trunk and tail region were imbedded in hard

wax and sections 8 w thick were cut. The sections were stained

on the slide, mostly in Delafield’s Hematoxylin, though I have

also, at times, used Borax Carmine and Picro-indigo-carmine for

differential staining with remarkably good results.

EXPLANATION OF THE PLATES.

Prats I.

Fig. 1. Dorsal aorta of Olarias magur (X 25). Note the absence of any trace of

a ligamentous structure in the dorsal wall of the aorta. c.aoz., cavity

of aorta; d.w.a., dorsal wall of aorta.

. Longitudinal section through the vertebral column and dorsal aorta of

Eutropiichthys vacha in the caudal region, showing the relative positions

of the ,dorsal ligament and the aortic ligament (x 5). The aortic

ligament in the caudal region is more closely attached to the dorsal wall

of the aorta than in the trunk-region.’ d.lig., dorsal ligament; sp.c.,

spinal cord; d.w.a., dorsal wall of aorta; lig., aortic ligament ; v.2.a.,

ventral wall of aorta ; cav.a., cavity of aorta; s.v.m., sub-vertebral mass

of connective tissue.

. Transverse section through the sub-vertebral region of the dorsal aorta in

Rita buchanani (X16). car., cartilaginous tissue; lig., aortic ligament.

. Transverse section through the sub-intervertebral region of the dorsal aorta

in Rita buchanani (X 16). lig., aortic ligament.

. Anterior termination of the aortic ligament in Silwndia gangetica (X » nat.

size). aor., dorsal aorta; lig., aortic ligament; a., anterior position of

aortic ligament which, passing through the dorsal wall of the aorta,

becomes attached to the basioccipital bone ; c., the point of origin of the

anterior branch of the ligament after it perforates the aorta; 0., basi-

occipital; par., parasphenoid; a.lig., anterior branch of the ligament ;

a., the tibres of the ligament which spread out to form a thin sheet.

Fig. 6. Ventral view of the skull and anterior vertebree in Wallago attw (nat.

size). par. parasphenoid; bas.o., basioccipital; 1s¢ v., first vertebra ;.

p.t., post-temporal ; 2nd v. second vertebra.

eat ee

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AORTIC LIGAMENT IN INDIAN FISHES. 15

Puate IT.

Fig. 7. A diagrammatic longitudinal representation of the position of the ligament

inside the dorsal aorta in Wallago attu. The lateral wall of the aorta

has been removed to show the position of the ligament. Jig., aortic

ligament; sws.f., suspensory fold; 3rd v., third vertebra; 2nd v., second

vertebra; 1st v., first vertebra; ba.o., basioccipital; d.w.a., dorsal wall

of aorta; v.2.a., ventral wall of aorta: lig.per., the place where the

ligment perforates through the dorsal wall of the aorta and becomes

attached to the bone; cav.a., cavity of aorta; lig.a.’, anterior branch of

the ligament running through the wall of the aorta; lig.a.’’, the place

where the anterior branch of the ligament perforates through the most

anterior dorsal wall of the aorta; lig.a.’/’’, the anterior branch of the

ligament after it comes out of the aorta and runs beneath the basioccipital

bone; eff:a., efferent arteries.

Fig. 8. Transverse section through the aorta of Wallago attu in the region where

the ligament perforates through the dorsal wall of the aorta. This section

is supposed to have passed through A-B region of fig. 7 (X18). c¢.aor.,

cavity of aorta; d.2v.a., dorsal wall of aorta; lig.per., the place where the

ligament perforates through the dorsal wall of the aorta and becomes

attached to the bone.

Vig. 9. Transverse section through C-D region of fig. 7. The ligament (lig.) here

runs through the upper region of the dorsal wall of the aorta (X 18).

Lettering as in fig. 8.

Fig. 10. Transverse section through E-F region of fig. 7. The ligament here

runs through the lower region of the dorsal wall of the aorta (xX 18).

Lettering as in fig. 8.

Fig. 11. Transverse section through G—H region of fig. 7 (x 85). d., adventitia ;

e., cartilaginous tissue; 6., media; int., intima; q@., elastic fibres; Jig.,

ligament ; ba2., bundles of elastic fibres.

Fig. 12. Transverse section through I-J region of fig. 7. A slight proliferation

takes place inside the cavity of the aorta in the region where the ligament

is situated (X16). car.c., cartilage-cells; lig., ligament; d.w.s., dorsal

wall of aorta.

Fig. 13. Transverse section through K-L region of fig. 7. The proliferation of the

dorsal wall of the aorta containing the ligament grows deeper (X 18).

Lettering as in fig. 8.

Fig. 14. Transverse section through M—N region of fig. 7. The proliferation reaches

nearly its maximum and the ligament acquires its characteristic shape.

Posteriorly the ligament flattens out, being narrower dorso-ventrally than

from side to side (X 18). Lettering as m He. 8.

ON SOME LIZARDS OF 'THE GENUS CHALCIDES. lan

6. On some Lizards of the Genus Chaleides.

By E. G. Boutencer, F.Z.8

[Received February 4, 1920: Read February 24, 1920. |

(Text-figures 1-4.)

At a recent meeting of the Society a paper was read by Major

Stevenson-Hamilton in which the subject of the geographical

distribution of the varieties of various African mammals was

touched upon, and it was pointed out that one would be justified

in treating some of the varieties as distinct species were it not

for the existence of intermediate forms. This paper brought to

my mind some notes I had made about ten years ago on the

classification and distribution of the Skink Chalcides ceellatus! a

species inhabiting Southern Hurope, Northern and N.-Eastern

Africa, and $.W. Asia, which presents an extraordinary amount

of variation: in fact, the structural difference between the two

extreme forms is so great that, were it not for the wonderfully

complete manner in which they are connected, they could not

possibly be denied specific rank. I have recently gone over

again the material in the British Museum, and completed my

notes on this subject, which I now have the honour to bring

before the Society.

In papers written nearly 30 years ago my father, dealing with

the matter, came to the conclusion that this species could be

divided into five distinct varieties or subspecies, characterized

mainly by the coloration and by the number of scales round the

body, which was found to vary between 24 and 40—a range of

variation far greater than is to be found in any other lizard *.

The five forms then described were the forma typica, and the

varieties ragazzi, tiligugu, vittatus, and polylepis. To these must

be added the var. occidentalis (Ch. simonyi Stdr.).

The position of the nostril has been used as a specific character

in the lizards of the genus Chalcides, the species viridanus, of the

Canary Islands, and bottegi, of Somaliland, being regarded as

specifically different from C. ocellatus, mainly from the fact that

the opening is pierced in advance of the suture between the

rostral and the first labial instead of exactly above it, as is

normally the case in the typical C. ocellatus. C. bottegi was

described from a single specimen preserved in the Genoa Museum,

and was stated to be closely related to C. ocellatus, but differed,

apart from having the nostril pierced in advance of the rostral

and first labial, in the body being much more slender and the

scales of the vertebral rows being more than twice as broad as

* Boulenger. Ann. & Mag. N. H. (6) v. 1890, p. 144.

Tr. Zool. Soc. xi. 1891, p. 138, ea Xvi.

53 Ann. Mus. Genova (2) xii. 1891, p. 12.

a » Xvi. 1896, p. 581.

Anderson. Zool. Egypt, Rept. p. 210 (1898).

78 MR. E. G. BOULENGER ON SOME

long *. On examination of a large material since received at the

British Museum, I find that this form cannot be accepted as a

distinct species, the nostril being almost as often pierced above

the suture in question as in advance of it; while in a number of

specimens of the typical (. ocellwtus the nostril is pierced in

advance of the rostral and the first labial. ‘The body of the form

bottegi is, I find, not always more slender than in the typical

C. ocellatus, in which there is considerable variation in this respect.

The number of scales, however, is less than in the typical

C. ocellatus, being as a rule 24, as in the var. ragazzii, but

dropping sometimes to 22. The degree of enlargement of the

two median rows of dorsal scales varies considerably both in the

form in question and in the typical C. ocellatus. If the position of

the nostril caunot in this genus be regarded as of specific value,

the question arises whether C. viidanus, which apart from this

character agrees so closely with C. ocellatus, must also be only

allowed the rank of a variety to be added to the numerous other

forms which are embraced in the specific conception of C. ocellatus.

I find, however, that the head has a different shape, the snout

being less convex—a difference which finds expression in the

proportions of the upper labials, all or most of which are not

deeper than long.

There are two forms of (. viridanus—-the typical, from Tenerife,

Gomera, and Hierro, with the sides and belly black and 26-32

(usually 28) series of scales; and the var. simonyi, from Gran

Canaria, with the belly yellow, greenish white or grey, the head

somewhat larger and better defined than in the preceding, and

28-34 series of scales.

As pointed out by Steindachner, the Chaleides of Fuertaventura

must be regarded as a variety (var. occidentalis) of C. ocellatus T.

Great individual variation in form is to be found in the

structure of these lizards, especially in the proportions of the

limbs and body. In the var. bottegi the latter may vary to a very

great extent, namely from 18 to 28 per cent. In the distance

between the axilla and groin the variation is also often great.

The variations show that little importance can be attached to the

proportions of the body and limbs, there being an overlap, for

instance, in the length of the limbs between the two species

CO. ocellatws and OQ. bedriage. The latter lizard was described as

differing specifically from C. ocellatus in the proportions of the

limbs, and in the nostril being pierced in advance of the suture

between the rostral and first labial. It has been shown that

neither of these characters can be regarded as absolute. I have

ascertained, however, that in C. bedriage the fourth labial

normally takes the place of the subocular, and not the fifth, and

* Boulenger. Ann. Mus. Genova (2) xviii. 1898, p. 719, pl. x. fig. 1, and (3) v.

1912, p. 330.

+ Lanzarote and Fuertaventura, waterless and treeless and nearer the African coast,

differ greatly from the other Canary Islands in their fauna, which is nearly identical

with that of the neighbouring Sahara.—Tristram, Brit. Assoc. 1893.

LIZARDS OF THE GENUS CHALCIDES. 79

that therefore it may, provisionally at least, retain its specific

rank.

In the small island of Linosa, between Tunisia and Malta,

lizards similar to, but easily distinguishable from, the typical

C. ocellatws are found, and have been regarded as the young of

the var. tiligugu, which occurs in Tunisia and Malta. They differ

from the typical form in the small size (the largest specimen

measuring only 80 mm. without the tail), in having the gular

Text-figure 1.

Yi SERIES A.

a. Ch. ocellatus.

b. Ch. sepoides.

c. Ch. delislii.

d. Ch. mauritanicus.

SERIES ]3.

a. Ch. thierryi.

y b. Ch. lineatus.

c. Ch. tridactylus.

" d. Ch. guentheri.

b c d

Reduction in the hind limb.

region spotted, and in the under surface being slate-colour. They

are dorsally brown, spotted all over with small black and white

ocelli. An indistinct paler dorso-lateral band is sometimes pre-

sent. The number of scales round the body is 30 in all speci-

mens, the two median rows being enlarged. These lizards are

undoubtedly distinct from all the other forms of the species

ocellatus, and for them I propose the varietal name of linose.

C. thierryi was originally described as a var. of C. bottegi: it is,

80 “MR. E. G. BOULENGER ON SOME

however, a very distinct species, quite different from the nume-

rous forms of C. ocellatus *. In its shorter not so unequal toes, in

its large ear-opening, and in its long, thick tail, it approaches the

groups including C. lineatus, tridactylus, guenthert; and my

father has given it as his opinion, that, although derived from the

same stock as C. ocellatus, it represents one of the pentadactyle

forms from which the more degenerate types referred to above

have been evolved ; whilst 2 continuous degeneration can be traced

from C. ocellatus through C. sepoides to C. delisliti and C. mauri-

tanicus.

In all, therefore, we now have, apart from the typical form,

seven varieties of the lizard C. ocellatws, and it is interesting

from the evolutionary point of view that they are geographically

connected, it being possible to trace every link in the chain from

the short and stout variety with as many as 40 scales from

Morocco, which must be regarded as the most generalized form,

to the long and slender type with only 22 scales round the body

from Abyssinia and Somaliland.

The general reduction in the number of scales takes place

as follows :—

1. var. polylepis (84-40 scales). Morocco.

,», occidentalis (80-32 ,, ). I. of Fuertaventura.

», vittatus (80-34 ,, ). Tangier.

- », chguern (28-34 ,, ). Sardinia, Sicily, Malta,

S. Italy, Algeria and

Tunisia, N.of the Sahara.

5. forma typica (26-32 ,, ). Arabia, Persia, Kgypt to

Algerian Sahara, Syria,

Cyprus, Greece, Eritrea.

> oo bo

6. var. linose (30 5) ke dlhvor Ibreaesey.

(ee ragazeru (2 a ae scall:

8. ,, bottegi (22-24 ,, ). Abyssinia, Somaliland.

The following are definitions of the 8 forms into which

C. ocellatus may be divided :—

1. Var. polylepis Blgry. 34-40 seales round the body, the two

median dorsal rows not enlarged; light brown above, with-

out ocelli, but with a round yellowish spot on each scale,

forming regular longitudinal series sometimes separated by

dark lines: young with vertical black-and-white bars on the

sides of the neck. Maximum length from snout to vent

155 mm.

2. War. occidentalis Stdr. 30-32 scales round the body, the two

median dorsal rows not enlarged; coloration as in the

preceding, but the yellow spots less numerous. Maximum

length from snout to vent 100 mm.

* Ch. bottegi var. thierryi Tornier, Arch. f. Nat. 1901, p. 87.

Ch. thierryi O. Neumann, Zool. Jahrb., Syst. xxi. 1908, p. 401.

Ch. pulchellus Mocquard, Bull. Mus. 1906, p. 466.

a en en en

LIZARDS OF THE GENUS CHALCIDES.

*$N7Y]]200 saproynyy JO NOINGLystp [eompdvadoar

°Z OINSY-4X9q,

“HIZZWOWe

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Proc. Zoot. Soc.—1920, No. VI.

82 MR. E. G, BOULENGER ON SOME

Text-figure 3.

POLYLEP|S OGELLATUS

lOCCIDENTALIS

i RAGAZZII

BOTTEG}

Relatienships and distribution.

3. War. vittatus Blgr. 30-34 scales round the body, the two

median dorsal rows not enlarged; brown above, without

spots or ocelli, with a light dorso-lateral and a dark brown

or black lateral band, both sharply defined, Maximum

length from snout to vent 115 mm.

4, Var. tiligugu Gmel. 28-34 scales round the body, the two

median dorsal rows not or but feebly enlarged ; olive or

brown above, with black and white ocelli and a well-defined

yellowish dorso-lateral band edged with black below. Maxi-

mum Jength from snout to vent 150 mm.

5. Forma typica Blgr. 26-32 scales round the body, the two

median rows not or but feebly enlarged ; yellowish or brown

above, with black and white ocelli, sometimes confluent

into irregular transverse bands; a light dorso-iateral band

sometimes present. Maximum length from snout to vent

140 mm,

6. Var. linose,n. 30 scales round the body, the two median dorsal

rows not enlarged; dark brown above, ocellated all over,

with or without a more or less distinct pale dorso-lateral

band ; belly grey; gular region spotted with black. Maxi-

mum length from snout to vent 80 mm.

7. Var. ragazzit Bler. 24 scales round the body, the two median

dorsal rows feebly enlarged ; pale greyish brown above, with

an ill-defined paler dorso-lateral band; no ocellar spots

except on the posterior part of the body, the hind limbs, and

the tail; crowded black spots form a lateral band from

nostril to above axil, passing through the eye and above the

ear-opening. Maximum length from snout to vent 75 mm.

LIZARDS OF THE GENUS CHALCIDES. 83

Text-figure Ay

Ch. ocellatus, var. polylepis. Ch. ocellatus, var. bottegi.

8. Var. bottegi Bler. 22-24 scales round the body. the two median

dorsal rows more or less strongly enlarged; yellowish or

greyish brown with black and white ocelli, with a dark, often

black-edged dorsal band along the median 1ows of scales and

a dark brown or black lateral band, the two separated by a

sharply-defined pale area. Maximum length from snout to

vent 150 mm.

The two extreme forms are represented on text-figure 4.

ON DEATHS IN THE GARDENS IN 1919. 85

7. Report on the Deaths in the Gardens in 1919. With

Notes on Avian Enteritis. By Naraanien S. Lucas,

M.B., ¥.Z.8., Pathologist to the Society.

[Received February 6, 1920: Read February 24, 1920.]

(With 4 Charts.)

The total deaths in the Gardens for the year 1919 amount

to 926.

The total is composed as follows :—

MViteianiialsi 4. ce ssnaceee 299

IBHIRC Gy: TRE ERENT) ae 368

Reptiles, etc. .......... te 2408)

JE) 117 22, 208 oe eee 50

In the following table are shown :—

In column I. animals in Gardens at beginning of year ;

_ I. “a added during the year ;

vi III. total of animals in Gardens ;

% IV. total of deaths ;

% V. percentage of deaths.

I. ROMMEL SUMAN) NR,

IWIEWODNOTANIS on ccccsccsnccaseaece OUD 279 954 299 31%

IE WINGIS assesses escneddospocsesen | AUZNG) 801 1947 368 19°75

Rep tilesys cat eeectucdete) 209) 658 937 209 22/°/,

The large percentage of deaths among the mammals is to be

accounted for by the high mortality among the macaques.

The usual table giving the deaths from various diseases is not

given, as no figures are available this year.

The subject of Enteritis deserves special mention.

The high rate of mortality from this disease is shown by the

charts for 1919. These emphasize the importance of an attempt

being made to deal with the disease.

Enteritis is the name given to inflammation of the intestine.

It begins as congestion and a consequent catarrh. The mucous

membrane inside is pink, deepening to red, and the contents are

liquid due to an excess of mucus and usually milky.

The congestion deepens and hemorrhages occur, so that the

whole gut becomes a deep red, and contents become blood-stained

and eventually black from altered blood. .

The final stage shows sloughing of the mucous membrane

lining the intestines, so that the walls are thin and the contents

dark and thickened by the destroyed cells.

It can be acute or chronic. The final stage of sloughing is

usually seen in this acute type. Often owing to the weakened

86 DR. N. S. LUCAS ON

Chart No. I.

Other

Causes.

Lnteritic,

Chart No. IT.

Enteritic.

Other

Causes.

Enteritic.

Other 0

Causes,

Other

Causes.

Enteritic

fo)

In these charts the deaths caused by enteritis are shown in a continuous line, those

due to all other causes in an interrupted line.

The birds are grouped according to the food eaten, though in the last group the chief

point is that these birds live mostly in the open and their food is scattered on the

ground and not placed in a receptacle. Enteritis is less prominent in this group,

on the whole, and most prominent in Group II., where the food is of the sort

which forms a good culture medium for bacteria and is almost always put into

receptacles,

DEATHS IN THE GARDENS IN 1919. 87

state of the bird bronchitis or pneumonia slips in and finishes

the illness.

~The cause of the inflammation is irritation, and this may be

either mechanical or toxic.

The mechanical source would be foreign bodies in the intestine,

e.g. grit. This appears a less likely cause.

The toxic cause may be either bacterial in origin or brought

about by poisons from unsuitable or decomposed food.

Which of these causes is the true one or which the prepond-

erating one is the subject of the investigation which has now

to be made.

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’

MR. A. WILLEY ON AN APODOUS AMIA CALVA. 89

8. An Apodous Amia calva. By Anraur WILLEY, F.R.S.,

F.Z.S., MeGill University, Montreal.

{Received March 13, 1920: Read April 13, 1920.]

It is known that a good many interrelated genera of fishes

differ from each other by the presence in one and absence in the

other of ventral fins. Perhaps the classic and primitive example

of this contrasting condition is that of the Crossopterygian fishes,

Polypterus and Calamoichthys, upon the theoretical interest of

which Gegenbaur (1895) laid some stress. Calamoichthys is a

Crossopterygian eel, the Mastacembelide are Actinopterygian

eels (Giinther), the Murenoids are Malacopterygian eels—all

lacking ventral fins. A far-removed contrast of the same kind is

found in the Swordfishes: Histiophorus with ventral fins, Xiphias

without ; and this may serve as a sample of the rest.

Only in a few species has the absence of ventral fins been noted

as a rare mutation. Brindley (1891) recorded the capture, in the

River Cam, of a White Bream without ventral fins. Ten years

later, EKigenmann and Cox (1901) described a specimen of the

Yellow Catfish (Amiurus natalis) from Turkey Lake, Indiana,

showing absence of all trace of ventral fins. Some further

references will be found in Gemmill (1912).

Last November (1919) a male Ama calva, which had been

caught in the Richelieu River on the south side of the St. Lawrence

in the province of Quebec, was purchased from the market in

Montreal. Its length was twenty inches and it was in perfect

condition except for one strange defect, the utter absence of the

ventral fins. The specimen is preserved in the Peter Redpath

Museum, McGill University. The addition of Amia to the

meagre list of occasional apodous mutants should contribute

towards the ultimate evaluation of the phenomenon. Its rarity

and incidence show that the absence of ventral fins from fishes

which normally possess them is no ordinary malformation, though

there is at present no means of testing its behaviour as a unit

character experimentally.

Gegenbaur gave reasons pointing to the ventral fins of recent

ganoids and teleosts having lost at least part of their original

function and being consequently in a state of flux and retro-

gression. In most teleostomes they seem to persist because they

have been inherited, rather than for any particular use they may

be to the animal. Accordingly their loss would not react in-

juriously upon the organism, but might be an advautage to it.

Bateson (1894) made no attempt to deal with this remarkable

variation, doubtless through lack of corroborative data. With

the increase of instances it seems likely that it will take its

place as a standard illustration of natural mutation amongst fishes,

99 MR. A. WILLEY ON AN APODOUS AMIA CALYA.

especially since it falls into line with ascertained anatomical

relations.

According to Giinther (1880), fishes living in limited localities

or concealing themselves in mud are apt sometimes to lose their

ventral fins. One of the local names for Amia is Mudfish, another

is Beaver-fish (poisson castor), a third is Bowfin. The last of these

may have reference to the rounded arcuate shape of the caudal

fin, like a stretched bow. Wherever it occurs it frequents

marshy places, and its habits resemble in many ways those of. the

oriental Tankfish (Ophiocephalus striatus), known in Southern

India as the “ murrel” and in Ceylon as the ‘“Jula.” This species

has ventral fins, but the closely-related Paddy-field fish (Channa

orientalis) is without them. The Swordfishes and some other

pelagic and deep-sea fishes show that. the presence or absence of

ventral fins does not depend on one class of habits alone.

There are certain other wavering characters in Teleostean

fishes, which, taken in conjunction with the admitted decadence

of the ventral fins, suggest the hypothesis that the presence or

absence of such deep-seated characters is linked up with their

use or disuse, and that they do not necessarily dwindle away

to vanishing point, but may simply drop out of the factorial

system.

REFERENCES.

. A. C. L. Gunrusr, 1880.—The Study of Fishes, p. 615.

. H. H. Brinptey, 1891.—“ On a Specimen of the White Bream

(Abrams blicca Bloch) without Pelvic Fins.” Proc. Zool.

Soc. pp. 108-9, pl. x.

. W. Bareson, 1894.—Materials for the Study of Variation.

. C. GecenBAur, 1895.—“ Das Flossenskelet der Crossopterygier.”

Morph. Jahrb. xxii. pp. 119-160, 5 figures.

5. C. H. Ergrnmann and U. O. Cox, 1901.—‘‘ Some Cases of

Saltatory Variation.” Amer. Nat. xxxv. p. 33.

6. J. F. Gemmit, 1912.—The Teratology of Fishes, p. 55. Glas-

cow, Ato.

H> OO

a a ree

_—

EXTERNAL CHARACTERS OF THE SOUTH AMERICAN MONKEYS. 91

. On the External Characters of the South American

Monkeys. By R. I. Pococx, F.RB.S.

[Received February 23, 1920: Read March 16, 1920.]

(Text-figures 1-13.)

ContTENTs.

Page

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“Me INfostiilse 08. cjekce eco sassuid ovecssnitesnh cece reek eee eee

i Mheshan! Gee RCAC HOM ARES heer serait) Quen

iat deena Feet. 1 KA DO ee een pera 47

The Tongue ..... ophl gtinas Sasa OAs

The External Genigalea ae eS Male Peer sence ee ote a3 UNS)

The External Genitalia of the Hemalen ee a iscokon 105

ET ayy Cee WRN 5 aloes Sen ee ee RS Oman ee eee JI)

Introduction.

The observations recorded in this paper are based mainly upon

the Platyrhine Monkeys that have died in the Zoological Gardens

during the past ten years* ; and the subject-matter is treated on

the lines adopted in my paper-on the Lemurs and Varsius (P.Z.S.

1918, pp. 19-53). Since I described the hands and feet and the

ears of the Hapalide in 1917 (Ann. Mag. Nat. Hist. (8) xx.

pp. 247-258), my notes, so far as that family is concerned, are

in the present case restricted for the most part to the external

genitalia, the species examined being Hapale jacchus, Mystax

ursulus, midas and mystav, Edipomidas aedipus, and Leontocebus

rosalia.

Of the Cebide, I have seen examples of all the admitted genera,

except Pithecia and Brachyteles ; but 1 have not seen both sexes

in all cases, and in many instances immature specimens only have

been available. These defects are regrettable, since the external

genitalia promise to yield valuable diagnostic characters for the

genera.

The immaturity of specimens also makes their specific identity

doubtful. Very little indeed appears to be known of the range

of variation in colour and structure within specific limits. It

is not an uncommon event, for instanee, to receive in our

Zoological Gardens immature examples of <Ateles exhibiting

characters of at least two admitted species and assuming perhaps

those of a third with growth.

I have examined examples of the following genera. Where the

species are doubtful they are marked with an interrogation,

* T am indebted, however, to Prof. J. P. Hill, F.R.S., for the opportunity to

examine adult examples of Callicebus personatus nigrifrons collected by himselt

at Minas Geraes.

92 MR. R. I. POCOCK ON THE EXTERNAL

and the better-known text-book names for the genera are put in

brackets :—

Callimico goeldti Thos.; ¢ ad., from the Ma River, Bolivia.

Callicebus (Callithrix) moloch Hoftm. (?); two immature 9°.

n personatus ; several adults of both sexes (Prof. J. P.

Hill).

Saimiris (Chrysothrix) sciureus Linn.; ¢ and 2 ad.

Cacajao (Brachyurus, Ouakaria) rubicundus Geoftr.; 2 ad.

Aotus (Nyctipithecus) trivirgatus (?); 9 ad.

Cebus albifrons (?) and apella(?); 3, 2 immature.

Lagothri« lagotricha (=humboldti); 2 ad.: and L. infumata ;

¢ immature.

Ateles ater Linn.; ¢ immature: and an immature ¢ partaking

of the characters of two or three alleged species.

Alouatta (Mycetes) villosa Gray (?); Q@ immature, so named

from Elliot because the immature specimen was all black.

Text-figure 1.

Mas

ae take ie D

——————

B &

. Nostrils of Leontocebus rosalia.

3 Callimico goeldii.

55 Callicebus moloch.

33 Cacajao rubicundus.

. Side view of muzzle of the same.

. Nostrils of Ateles.

. Side view of muzzle of the same.

Qa Om b>

The Nostrils.

Contrary to the current belief, all the South American Monkeys

are not, strictly speaking, Platyrhine. There is great variation

in the shape and situation of the nostrils. Typically both in the

CHARACTERS OF THE SOUTH AMERICAN MONKEYS. 93

Hapalide and the Cebide the internarial septum is wide, greatly

surpassing In width the longest diameter of either nostril, and

the nostrils look outwards and shightly forwards, but so slightly

in some cases as to be only just visible when the face is viewed

from the front, as in Cebus, Ateles, Cacajao (Owakaria), and Oalli-

cebus (text-figs. 1 & 2, A-C). But in two of the genera—dAotus,

Text-figure 2.

= ee

A. Nostrils of Cebus.

B és Lagothr Be

Cc Saimiris sciurea.

TD. Side - y lew of muzzle of Aofus.

E. Half protile view of the same, enlarged.

©. Nostrils of the same.

G. Nostrils of Alouatta.

H. The same from above.

I. Side view of muzzle of the same.

x He

as recorded by H. O. Forbes, and Alcwatta—the nostrils *are less

ieee are visible to a areat extent from the front, ‘and are

éparated by a septum ahieh hardly exceeds the long diameter

94 MR. R. I. POCOCK ON THE EXTERNAL

of either. According to H. O. Forbes’s account, the nostrils of

Brachyteles seem to resemble those of Alowatta. (Text-fig. 2,

D-I.)

The nostrils also vary in shape in a very interesting manner.

In Callicebus they are practically civeular, and nearly so in

Cacajao. In Cebus they are longitudinally ovate. In Ateles the

upper edge has an *S?-like curve, the posterior portion of the

orifice being a narrow slit, owing to the presence of the down-

wardly projecting lobe which constricts the nostril behind. This

Icbe is also present in Alowatta, but it is relatively larger in

Aotus than in any member of the Cebide. In that genus, indeed,

the nostrils with their comparatively narrow septum and well-

developed posterior lobe are of a more primitive type, and more

resemble the nostrils of the Strepsirhine Primates (Lemurs) than

do those of any genus of the Haplorhine Primates, including

even Tarsus.

To sum up—the nostrils are typically platyrhine in Hapalide,

Callimico, Callicebus, Saimiris, Cebus, Cacajao, Lagothria, Ateles ;

and stenorhine in dotus, Alowatia, and, it is alleged, in Brachy-

teles.

The Kars.

In my paper on the genera of Hapalide, the ears of Hapale,

Mystax*, Leontocebus, and Gidiponidas were described, and it

was pointed out that Gidipomidas may be distinguished by the

suppression of the free edge of the pinna from a point just below

the level of the upper portion of the antitragal thickening.

In the Cebide the pinna of the ear generally resembles that

of the Hapalide, showing variations in suppression analogous to

those of that family. In the majority of cases (Aotus, Callicebus,

Alouatta, Cacajao, Cebus, and Callimico) the pinna is provided

with a freely projecting laminate margin, which terminates

inferiorly just beneath the antitragal thickening as in all the

Hapalide except Gdipomidas.

In the ear of Cebus, which will serve as well as another as

typical of the group, the intertragal notch is bordered in front

by a comparatively small tragus and behind by an enlarged,

prominent antitragus, which has a well-developed ridge on its

inner surface and is defined behind by a notch from the strong

ridge of -the antihelix, which curves upwards and forwards,

dividing above into two branches—one, less well-defined, passing

forwards and upwards towards the upper edge of the pinna in

front, the other, which runs horizontally forwards, constituting

the well-defined and shelf-like supratragus (plica principalis).

Che anterior end of the supratragus is overlapped and concealed

by the backwardly folded edge of the antero-superior portion of

the pinna; but this backwardly folded edge is carried only a

* The only example of Hapalide examined by me since that paper was written

was a specimen of Mystax mystax. In this the ear resembles that of IZ. midas.

a ~~

CHARACTE&RS OF THE SOUTH AMERICAN MONKEYS, 95

very short distance below the supratragus, which is set high

above the middle of the ear, there being a long space between

the inferior termination of the back wardly folded edge and the

Text-figure 3.

Hana M4

WON

yy aes

NSN

We Ly a ae

EARS.

A. Cebus. B. Aotus. C. Saimiris. WD. Callicebus, E. Callimico. F. Alouatta.

G. Ateles. H. Ateles (with rim overfolded). I. Lagothrix.

‘ a. Point where free edge of rim ceases,

tragus. Between the curved elevation, formed by the antihelix,

and the edge of the pinna there is a semicircular fossa, extending

from the antitragus, where it is deep, up to the fossa above the

96 : MR. R. I. POCOCK ON THE EXTERNAL

supratragus. The upper edge of the pinna is always folded ;

but the posterior edge may be unfolded, partly folded, or folded

throughout its extent. In the latter case the fossa behind the

antihelix is especially well defined. Sometimes the upper edge

of the pinna shows an angular projection, sometimes it 1s evenly

rounded; but I have not worked at the variation of the ears

with a view to their possible systematic value in the determination

of species. (Text-fig. 3, A.)

The ear of a specimen of Alowatta resembles that of Cebus,

except that the antitragus is less well developed and the semi-

circular fossa behind the ridge of the antihelix does not extend

so far beneath the antitragus. The entire edge of the pinna is

folded, (Text-fig. 3, F.)

In a specimen of dotus the shape of the pinna recalls that of

Mystax. The antero-superior portion of the edge is the only

part that is folded. There is no definite fossa behind the ridge

of the antihelix except inferiorly behind the massive antitragal

thickening. The posterior edge is convex below, concave above.

At the summit of the concavity it runs out into a short, obtusely

angular point, above which the edge extends obliquely upwards

and backwards to the rounded top of the ear. (‘l’ext-fig. 3, B.)

In Cacajao the ear is very like that of Cebus, but the inferior

portion of the submarginal fossa does not extend so far forwards

below the antitragus. The posterior and inferior edges of the

pinna are unfolded.

There is nothing in the ear of Callimico that calls for particular

notice. The tragus is hardly developed, but the antitragus is

massive; the superior edge of the pinna is folded to a slight

extent, the posterior edge being flat, and the postero-inferior

portion is not so prominently rounded as in typical members of

the Cebide. (Text-fig. 3, HE.)

In Oallicebus and Saimiris the postero-inferior portion of the

pinna is somewhat reduced, its free margin ceasing at a point

below the notch defining the posterior margin of the antitragus,

and the fossa behind the antihelix, which is very shallow in

Saimiris, falls short of the antitragus inferiorly in both genera.

This partial suppression of the free margin of the pinna inferiorly

foreshadows, in a measure, the condition seen in the two genera

to be considered next. (Text-fig. 3, C, D.)

The above-mentioned genera have normally formed and nor-

mally developed pithecoid ears; but in Ateles and Lagothrix

the ear is modified in a manner similar to that of @dipomidas

in the Hapalide. In Ateles the upper portion of the pinna is as

well developed asin Cebus, with the anterior edge folded and the

upper and posterior edge folded or flat; but the inferior portion

of the pinna has no free margin below a point approximately on

a level with a line half-way between the supratragus and the

antitragus, there being no postero-inferior laminate lobe and no

fossa impressing the antitragus or the area just behind it. For

the vest, the ear is normal, the tragus, antitragus, supratragus,

Pa. *

CHARACTERS OF THE SOUTH AMERICAN MONKEYS. 97

and the ridge of the antihelix being well developed. (Text-fig. 3,

G, H.)

In Lagothrix the ear shows similar suppression of the lower

half of the margin of the pinna; but the upper portion is also

reduced and stands away from the head to a comparatively small

extent, being fleshy and thickly covered with hairs. (Text-

fig. 3, I.)

By the structure of the ear, therefore, the genera of Cebide

fall into two groups :—(1) comprising Ateles and Lagothrix, in

which the inferior portion of the pinna has no freely projecting

laminate margin; (2) comprising Alowatta, Cucajao, Saimiris,

Callicebus, Cebus, and Callimico, in which the free laminate

margin extends right round the pinna inferiorly to a point below

the intertragal notch.

Dr. Boas (Ohrknorpel etc. der Siug. 1912, pp. 199-206, pl. 23)

describes and figures the ears of several species of Cebus, of

Alouatta, Ateles, Saimiris, Mystax, and Hapale. So far as my

observations on these genera extend, they agree completely with

those of Dr. Boas.

The Hands and Feet.

In a paper upon the genera of Hapalide or Marmozets (Ann.

Mag. Nat. Hist. (8) xx. p. 249) I pointed out that the hands

and feet of Leontdcebus differ from those of Hapale, Mystax, and

(dipomidas in the elongation of the palm and sole, and that the

hand further differs in the presence of webs tying the second

digit to the third, and the third to the fourth, the latter web

being of considerable depth, so that the two digits in question are

only separable from a distance a little on the proximal side of the

joint between the first and second phalanges.

I have since found that this character does not always hold, for

in an example of Mystax mystax, subsequently examined, I found

the third and fourth fingers of the right hand webbed almost to

the same extent as in Leontocebus ; while the left hand hardly

differed in that respect from the hands of examples of other

species of Mystax, and of all specimens of Hapale and of Gdipo-

midas examined. In this connection it is interesting to recall

that W. A. Forbes recorded in the case of Pithecia satanas a

similar abnormal instance of interdigital webbing, the third and

fourth digits of both hands being tightly tied together to the

bases of the claws (P. Z. 8. 1882, p. 442).

The hands and feet of the Hapalidee differ from those of all the

other South American Monkeys, except Callimico, in two partic-

ulars :—(1) The hallux is extremely reduced in size, so that when

it is turned forwards its apex falls short of the distal margin of

the plantar pad ; (2) the nails of all the digits, except the hallux,

are converted into strongly compressed, curved, pointed claws like

those of a Squirrel. The capacity of these claws for maintaining

a secure hold on the rough bark of trees compensates for the loss

of grasping power in the foot, due to the feeble development of

Proc. Zoou. Soc.—1920, No. VII. 7

98 MR. R. I. POCOCK ON THE EXTERNAL

the hallux. I believe the Marmozets to be specialised Cebide,

derivable from them by reduction in bodily size, by the loss of

the third molar above and below, and by adaptation of the hands

and feet for holding to the roughnesses of bark in the way

above described.

Text-figure 4.

. Right hand of Callimico goeldii, adult, approx. nat. size.

. Right foot of the same.

. Right hand of Saimiris sciwrea, adult. X 3.

. Right foot of the same. X 3. ‘

. Right hand of Callicebus moloch, young. X

. Right foot of the same.

The hands and feet of Callimico need no detailed description,

since they resemble in all important points those of Hapale or

Mystax. (Text-fig. 4, A, B.)

CHARACTERS OF THE SOUTH AMERICAN MONKEYS.

Text-figure 5.

“a,

A. Right hand of Cacajao rubicundus, from below.

ght foot of the same.

B. Right foot of the same.

C. Right hand of Aotus, from below.

D. Ri

100 MR. R. I. POCOCK ON THE EXTERNAL

The hands and feet of the typical Cebidw while exhibiting an

interesting range in structural variation—e. g., in the develop-

ment of the pads, the relative lengths and spacing of the digits as

described below under the different genera—have certain features

in common, which may be briefly referred to by way of

introduction.

The talons are always narrow and compressed and not infre-

quently acuminate, but are never so strongly compressed, curved,

and pointed at the tip as in the Hapalide and Callimico. As in

the Hapalide, the pollex, when present, is a short edition of the

other fingers, and is never truly opposable to them even to the

extent that it is in the Old World Monkeys, being set much

closer than in the latter to the base of the second digit, although

the space between them varies to a certain extent according to

the genera. The hallux is typically well developed, although

somewhat reduced in Aéeles. It can be extended at right angles

to the long axis of the foot, and it projects approximately from

the middle of the side of the latter, nearly half-way, that is to

say, between the second digit and the tip of the heel.

The digits of both hands and feet, apart from exceptional cases,

are free from webbing—that is to say, they are separated almost

down to the plantar pad when viewed from the lower side.

Digits 3 and 4 both on the hands and feet are frequently sub-

equal; and since they are occasionally subequal in Lemurs and

in the Catarhine Monkeys, there is in this respect a complete

gradation between the Lemurs, in which digit 4 typically sur-

passes 3, and the Catarhine Primates, in which digit 3 typically

surpasses 4.

The plantar and digital pads are as a rule not well defined, but

in Aotus they are especially well developed, and they surpass the

average in Saimuris.

In Cebus the palm of the hand is tolerably long and digits 2,

3, 4, and 5 are subequally spaced; digit 1 (the pollex) is com-

paratively long, and a little further removed from digit 2 than

the latter is from digit 38. The foot is much longer than the

hand. (Text-fig. 6, A, B.)

The hands and feet of Callicebus do not appear to differ

materially from those of Cebus. (Text-fig. 4, H, F.)

Saimiris has the pads better defined than in either of the

preceding, and the palm of the hand is relatively broader.

(Text-fig. 4, C, D.)

In Aotus the hand is also relatively broader than in Cebus and

Oallicebus, and the pads are exceedingly well developed and

coarsely striate. It seems probable that the exceptional develop-

ment of the pads and sensory striz in this genus is an adaptation

to the nocturnal habits of this Monkey, the specialised tactile

sense compensating for imperfect nocturnal vision*, (Text-fig. 5,

CD»)

* W. Kidd (‘The Sense of Touch,’ pp. 84-38, 1907) has figured and described the

hands and feet of Hapate, Saimiris, and Cebus from the point of view of the sensory

ridges.

CHARACTERS OF THE SOUTH AMERICAN MONKEYS. 101

In Cacajao the length of the hand as compared with the foot

is about the same as in Cebus; but in the former genus the

pollex is shorter and the interval between digits 2 and 3 is a

Text-figure 6.

- Right hand of Cebus, from below.

. Right foot of the same.

. Right hand of Lagothri« infwnatus, from below.

. Right foot of the same, with abnormal basal webbing between digits 2

and 3.

x 2 approx.

little greater than between 3 and 4 or 4 and 5, thus foreshadow-

ing the more marked inequality in spacing that obtains in

Alowatta and Lagothrix. (Text-fig. 5, A, B.)

102 MR. R. I. POCCCK ON THE EXTERNAL

In Alouatia and Lagothri« the hand is relatively larger and its

digits are longer as compared with the foot than in the genera

previously mentioned, and the space in the hand between digits

2 and 3 is markedly greater than between digits 1 (pollex) and 2

or 3and 4. The hands are thus in a sense zygodactylous, like

Text-figure 7.

A. Right hand of Alouatta villosa, young.

B, Right foot of the same.

C. Extremity of tail of Ateles ater.

the paws of Phascolarctos and of Cuscus amongst the Marsupials,

there being a grasping interval between digits 2 and 3 instead of

between digits 1 and 2 as in Lemurs and Catarhine Monkeys *.

* From a photograph of a living Pithecia pithecia I judge that there is a wide

space between digits 2 and 3 in that genus also.

CHARACTERS OF THE SOUTH AMERICAN MONKEYS. 103

Both in Alouatta and Lagothria the pollex is normally developed

for the family, being perhaps a little shorter relatively than in

Cebus but longer than in Cacajao. The hallux is also of normal

length and strength*. (Text-fig. 6, C, D, and text-fig. 7, A, B.)

In Afeles, as is well known, the hand differs from that of other

genera of Cebide, except Brachyteles, in having the pollex func-

tionless and at most forming a small excrescence on the side of

the palm ; and it is noticeable that digits 2, 3, 4, and 5 are evenly

spaced as in Cebus, Callicebus, etc., and that there is no grasping

space between digits 2 and 3 as there is in Alouatta and Lago-

thric. The hand of Ateles is therefore not derivable from the

type of hand seen in Lagothrix, but from the more primitive type

of hand seen in Cebus for example. (Text-fig. 8, A, B.)

Text-figure 8.

A. Right hand of Ateles paniscus, from below.

B. Right foot of the same.

The foot of Ateles is characterised by the reduction in the size

of the hallux, which is both shorter and weaker than in other

genera, the big muscular lobe at the base, representing the

““ball” of the human thumb, being scareely visibly developed.

* In a specimen of D. infumatus the second and third digits of the foot were

anited basally by a short web. Possibly this was an abnormality, since no such

variation occurred in an example of L. lagotricha.

+ The only genus of Monkeys which resembles Ateles [and Brachyteles| in the

suppression of the pollex is the Catarhine Colobus. In the latter, however, the

suppression of the pollex is clearly foreshadowed in Pithecus (= Presbytes = Semno-

pithecus), where it is greatly reduced.

104 MR, R. I. POCOCK ON THE EXTERNAL

Setting aside Callimico, which in the structure of the hands and

feet falls in with the Hapalide, the genera of Cebidee, so far as

known to me, may be arranged as follows by their extremities :-——

a. Pollex suppressed and functionless; hallux short, with basal

muscular lobe scarcely developed ... Ateles [and probably Brachyteles |.

a’. Pollex and hallux normally developed.

b. Feet relatively short as Soares with the hands; a wide

erasping space between digits 2 and 3 of the hand.

Alouatta, Lagothrix.

. Feet long as compared with the hands; digits 2 to 5 of the

hand generally approximately evenly spaced, space between

2 and 3 a little wider in Cacajao.

ec. Pads exceedingly well developed and coarsely striated ...... Aotus.

ce’. Pads not noticeably strongly developed and comparatively

weakly striated .................. Cebus, Callicebus, Saimiris, Cacajao.

The hands and feet of Ateles show an interesting resemblance

to those of the Orang Utan (Simia) amongst the Anthropoid Apes.

In the latter the pollex and hallux are both small, the Ape

depending mainly for its hold upon the clutch of digits 2 to 5,

which form powerful suspensory hooks. The same applies to

Ateles, which, like the Ape, is no jumper, but climbs by reaching

from bough to bough, commonly in an inverted position, aided by

its prehensile tail.

Lagothrix and Alowatta with comparatively short feet and

highly prehensile tails also climb by reaching instead of by

jumping from branch to branch. The rest of the genera with

relatively longer feet jump, so far as my experience goes, to a

certain extent. The species of Cebws indeed, despite their pre-

hensile tails, are tolerably good jumpers, but the only South

American Monkeys which appear to rival in arboreal activity the

more agile Catarhine Monkeys are the Marmozets, which can

cover a surprising distance with a leap.

The Tongue.

As in Lemurs, and some other Mammals, the lower surface of

the tongue in Platyrhine Monkeys is provided with a lamella—the

Jrenal lamella—in close connection with the orifices of the sub-

lingual salivary gland. In all cases observed this lamella ends

distally in a pair of slender juxtaposed processes, and its free

lateral margin runs backwards on each side towards the base of

the tongue*. (Text-fig. 9.)

I have not seen a sufticient number of specimens to establish

generic differences in the structure of this frenal lamella. Its

apparent narrowness in some vases and breadth in others may be

due to contraction or expansion in accordance with the mode of

preservation. The greatest variation was noticed in a common

Marmozet (Hapale jacchus), in which the edge of the lamella was

denticulated; and this condition was correlated with somewhat

coarse beading of the edge of the tongue. In this particular

* This lamella corresponds, I believe, to Wharton’s papillze of human anatomists,

and not to the sublingua of the Lemurs.

CHARACTERS OF THE SOUTH AMERICAN MONKHKYS. 105

specimen the lamella recalled in appearance the sublingua of the

Lemurs. (Text-fig. 9, A, B.)

Text-figure 9.

A. Lateral view of tongue of Hapale jacchus.

B. Lower view of the same.

C. Lateral view of tongue of Leontocebus rosalia.

D. Lower view of tongue of the same.

: 33 a Alouatta.

FE. +5 ss Cacajao.

G. Saimiris, with frenal lamella turned aside.

H. Tongue of Cebus, with the tip raised to show the relations of the frenal

lamella.

The Anal and Genital Areas of the Female. »

In the Marmozets Hapale jacchus, Cidipomidas edipus, and

Leontocebus rosalia the orifice of the vulva lies between a pair of

simple labia, and there is no visible external pendulous clitoris.

The most noticeable difference between these species in the

106 MR. R. I. POCOCK ON THE EXTERNAL

Text-figure 10.

\ Be

S tee?

= ZES

BEY | YA

aS NSS U WE.

SS ZZ)

SS . MEA

SS ZS

st: WES)

—— <P,

SS tS

a N{ pase

a —ss

4 5 {ra

a ‘SS A

Zt 7

LR iS

2 Ley 06 of

: CODES

oe “Sys Mis

G ANS

ZA, &

SSS

FO G

External Generative Organs of female.

A. Leontocebus rosalia, from behind.

B. Hapale jacchus, from behind.

C. The same, from the front.

D. Aotus, from behind.

Ki. Saimiris sciwrea, from behind.

F. Tip of clitoris of the same, from the front.

G. Side view of clitoris of the same.

a., anus; 7., ischial prominence; el., clitoris.

CHARACTERS OF THE SOUTH AMERICAN MONKEYS. 107

naked ano-genital area is the presence in Z. jacchus of a number

of whitish pustules round the anus, on the perineum and on the

ischial prominences. These exude under pressure a sebaceous

substance, which may be odoriferous. There is no trace of them

either in LZ. rosalia, in Mystax, or in Gdipomidas edipus. (Text-

fig. 10, A-C.) |

In the Cebide the ano-genital area varies considerably accord-

ing to the genus. In immature and unpaired females the urino-

genital orifice is a longer or shorter slit, above which, but

concealed from view by membrane (the Aymen), lie the apertures

of the urethra and of the vagina. (Text-fig. 11, B, D.)

In nearly all cases—-that is to say, in Cullincbue Saimiris, Aotus,

Alouatta, and Cebus—the ischial prominences and the area between

the anal and genital orifices and above the anus are naked or

scantily hairy; but in Lagothriz the hair grows tolerably thickly

and closely round the anus and genital area, covering the ischial

prominences and perinzeum.

In an adult, but unpaired, female Aotus the urino-genital

orifice is a very short and narrow slit bounded by small labia, but

there is no visible clitoris, the perineum and the area to the right

and left of the labia being somewhat swollen. The condition

observed resembles tolerably closely that of the Hapalide. (Text-

fig. 10, D.)

ein an adult Callicebus personatus (preserved in alcohol) the

perineum is comparatively short and the generative area is piri-

form, broad above, and narrowed below, The orifice being flanked

by thickened labia inferiorly converging upon a small glans-

shaped, grooved clitoris, which is hardly, if at all, pendulous. A

young ©. moloch resembles the last described, but the orifice

is merely a narrow slit, the thickened labia being undeveloped.

(Text-fig. 12, A, B.)

In an immature Alowatta the rima is an elongated slit, wider

above than below and flanked by swollen labia. The clitoris

is short and subconical, with a slightly expanded apex. (Text-

fis.)

In an adult female Lagothrix the elongated rima is flanked by

a pair of much swollen labia, forming together an oval area nar-

rower below, where it passes into a short subglobular, ungrooved

clitoris, defined proximally by a constriction*. JLagothria and

Alouatta resemble one another tolerably closely in the structure

of the female external genitalia. (Text-fig. 11, A.)

In an adult but unpaired female Saimiris the clitoris is long,

slender, pendulous, and grooved practically to the tip, which,

although tapering and not incrassate, has a shield of thickened

integument on its anterior surface. (Text-fig. 10, E-G.)

In Cebus the clitoris is elongated, slender, and grooved, prac-

tically to the apex as in Saimiris, but it is more penis-like, the

* This specimen of Lagothrix had been injected for anatomical purposes, and

possibly the genitalia were abnormally turgid, owing to the injected preservative.

108 MR. R. I. POCOCK ON THE EXTERNAL

tip being incrassate and enveloped in a definite preputial fold of

integument. (‘lext-fig. 11, C, D.)

Text-figure 11.

External Generative Organs of female.

A. Lagothrizx. B. Alouatta. C. Cebus. D. Cebus.

a., anus; 4., ischial prominence ; cZ., tip of clitoris; w., orifice of urethra; and

v., orifice of vagina exposed by cutting away the hymen in D and shown

diagrammatically with the hymen intact in B.

x6 30

CHARACTERS OF THE SOUTH AMERICAN MONKEYS. 109

I have had no opportunity of examining the dead body of a

female Ateles, but in living examples the clitoris attains a quite

unusual size. It is pink in colour and subcylindrical in shape,

and so greatly exceeds in size and prominence the penis of the

male, that people unacquainted with the facts invariably confuse

the sexes of this genus. This clitoris is apparently ungrooved,

and its apex is truncated and not incrassate.

The Anal and Genital Areas of the Male.

In the Marmozet Hapale jacchus the naked circum-anal in-

tegument, the ischial prominences, the scrotum, and the area

above the scrotum are studded with white glandular pustules,

like those of the female. The penis, which emerges from the

centre of the naked area of pustular skin, the lower half of which

is formed by the anterior surface of the scrotum, is subacuminate,

ending distally in a subcylindrical prominence, which lies above

and overhangs the transversely crescentic urino-genital orifice.

(Text-fig. 13, A-D.)

In Leontocebus rosalia the pustules, as in the female, are

absent; and the penis differs from that of H. jacchus in ending in

a definite hemispherical glans, carrying on the lower half of its

distal surface the transversely crescentic urino-genital orifice.

(Text-fig. 13, HE, F.)

In Mystax ursulus the subcaudal integument is without glan-

dularv pustules, and the penis ends in a small hemispherical glans,

which is described in my notes as bilabiate. That epithet might

also be applied to the glans of Leontocebus; but whereas in the

latter the labia are apparently dorsal and ventral, in Mystax

they appear to be lateral and to close over the small subcentral

orifice of the urethra.

Fresh material, however, is required to substantiate these

apparent differences in the glans in the genera of Hapalide.

In the Cebide the penis differs to a greater or less extent in

the genera examined. In Saimiris it terminates in a somewhat

cordate glans with longitudinally elongated orifice on its under-

side (text-fig. 13, G, H). In Callicebus personatus the generative

area of the male closely resembles that of the female, the scrotum

recalling the labia of the generative orifice and the penis re-

sembling an enlarged clitoris. The penis, which is short, ends in

an ovate glans, with an elongated orifice on its inferior or pos-

terior side as in Saimiris (text-tig. 12, C-F). In Callimico (text-

fig. 12, G), the penis has a hemispherical glans with terminal

orifice.

In Ateles the penis is subcylindrical from the base to the tip,

which is a little expanded. Its distal extremity is a nearly flat,

subcircular area, the orifice piercing the middle of its lower half

(text-fig. 13, K-M). In an immature example of Lagothrix infu-

matus the penis apparently differs from that of Ateles in having

the orifice in the middle of the terminal field. In neither of

110 MR, R. I. POCOCK ON THE EXTERNAL

Text-figure 12.

> G3 A

SNVipasatid

f a G

VE Oe 2

External Generative Organs.

. Callicebus personatus, ° ad., from behind.

. The same, from the side.

. Callicebus personatus, § ad., from behind.

. The same, from the side.

. Tip of penis of the same, from the front.

¥. The same, from behind.

G. Callimico goeldii, $ ; penis and scrotum from behind.

Hoan eS

a., anus; 7., ischial prominence; cl., clitoris; /., labium of yulva; p., penis

s., scrotum.

CHARACTERS OF THE SOUTH AMERICAN MONKEYS.

ABH Rune OOW

Text-figure 13.

External Generative Organs of males.

Rear end, with tail raised, of Hapale jacchus.

. Tip of penis of the same.

Naked area round penis of the same.

. Side view of extended penis of the same.

. Tip of penis of Leontocebus rosalia.

. Side view of extended penis of the same.

Lower view of extended penis of Saimiris.

. Side view of same with bristle passed through urethra.

Lower view of scrotum and extended penis of Cebus.

. Side view of extended penis of Afeles.

. Side view of tip of penis of A¢eles (another specimen).

. Tip of the same.

. Tip of penis of Cebus.

a@., anus ; 7., ischial prominence ; s., scrotum ; 0., orifice of penis.

stil

IW) MR. R. I. POCOCK ON THE EXTERNAL

these genera is the organ provided with a baculum (os penis). In

Cebus, however, the terminal portion of the penis is strengthened

with a baculum as in the Catarhine Monkeys, and the tip of the

baculum forms a small prominence in the middle of the subcir-

cular terminal area of the penis, thrusting the orifice to the right

of the middle line. For the rest the penis of Cebus differs from

that of Ateles in the gradual expansion of its distal portion up to

the truncated tip. (Text-fig. 13, J, N.)

The Tait.

I have no new facts to add to those already recorded about

the tails of South American Monkeys. In the four genera of

Hapalide (Hapale, Mystax, Cidipomidas, and Leontocebus) and

in six of the genera of Cebide (Callimico, Callicebus, Saimiris,

Pithecia, Cacajao, and Aotus) the tail is evenly bushy or hairy

throughout and not prehensile. Only in Cebide of the genera

Cebus, Lagothrix, Brachyteles, Ateles, and Alowatta is it prehensile.

Since, therefore, the tail is prehensile in only fifty per cent. of the

genera of South American Monkeys and in a minority even of

the Cebide, the prevalent idea, derived from the text-books, that

caudal prehensility is characteristic of Platyrhine Primates is

indefensible.

In nearly all the species of Cebus the tail, although prehensile,

is evenly hairy to the tip above and below. Only in the Central

American form (C. capucinus) is the end naked below. In this

genus practically the only modification of the tail is the develop-

ment of the flexor muscles at the expense of the extensors, so

that the organ is normally curled in its distal half. So far, there-

fore, as the tail is concerned, Cebus bridges the interval between

such genera as Saimiris or Aotus, for example, and the genera

Lagothrix, Brachyteles, Ateles, and Alowatta, in which the prehen-

sile power and tactile sensibility of the tail reach a maximum.

In the latter genera the distal portion of the tail is highly mobile

and sensitive, with the skin of its lower side naked and trans-

versely ridged, like the underside of an Elephant’s trunk (text-

fig. 7,C). It serves, indeed, the purpose of an additional hand

not only for climbing, but for grasping objects beyond the reach

of the arms*. Iam unable to satisfy myself, however, that this

special modification of the tail is proof or even strong evidence of

attinity between the genera that exhibit it. Ateles and Brachy-

teles are probably closely allied; but Lagothrix and <Alowatta

differ from both very markedly in the structure of the hand ;

and Alouatta differs at all events from Lagothric and Ateles in its

stenorhine nostrils, its ears and cranial characters.

* T have seen a Spider-Monkey (A¢eles) in the Gardens thrust its tail through the

bars of its cage and take food from visitors which it could not reach with its hand.

Capuchins (Cebus) apparently never attempt this; but I have seen them use their

tails for carrying paper bags and other things, very much as the Rat Kangaroo

(Bettongia) carries tufts of grass,

CHARACTERS OF THE SOUTH AMERICAN MONKEYS. WMS

The discovery of Callimico, with the dentition of the Cebide

and the hands and feet of the Hapalide, raises the question as

to the status to be accorded to these two groups. Callimico,

hitherto, has been included in the Cebide on the strength of the

retention of the last molar. Personally, I should attach more

importance to the extremities, and regard Callimico as a primitive

Marmozet. In that case the Callimiconine will be a subfamily of

Hapalide.

The genera of Cebide have been classified in various sub-

families, Elliot admitting :—

1. Cebine (Ateles, Brachyteles, Lagothrix, and Cebus),

2. Alouattine (Alowatia).

3. Pitheciine (Pithecia, Cacajao, Saimiris).

4, Aotinee (Aotus).

_ Without in any particular endorsing this classification, I do not

feel prepared to interfere with it. There appear to me to be no

particular reasons for associating Saimiris with Cacajao or Cebus

with <Ateles. Elhot extricated himself from the difficulty of

Callicebus by placing it, quite indefensibly of course, with the

Hapalide.

Proc. Zoou. Soc.—1920, No. VIII. 8

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ON THE ANATOMY OF THE TONGUES OF THE MAMMALIA, WLS

10. The Comparative Anatomy of the Tongues of the

Mammalia.—I. General Description of the Tongue.

By C. F. Sonnrac, M.D., Ch.B., F.Z.S., Anatomist to

the Society.

[Received January 22, 1920: Read March 16, 1920.]

(Text-figures 6-17.)

A considerable body of literature dealing with the development

and histology of the tongue has appeared in recent years, and a

number of individual tongues and groups have been described.

No one has, however, taken the various structures and shown

how they vary in different animals in exactly the same manner as

I have described here. It is the object of my series of papers to

fill this gap, and show how the tongue is of value for purposes

of classification. In this paper I have indicated the different

directions in which the various structures can be modified, and I

have defined the terms used in the sueceeding ones.

The tongue is a muscular organ enveloped in mucous membrane

except at the base and the posterior part of the inferior surface

where the muscles, nerves, vessels, and lymphatics enter and

leave it. On the dorsum and inferior surface the mucosa differs,

being firmly adherent in the former situation and_ loosely

attached to the subjacent tissues in the latter. On the dorsum,

jateral border, and, iff most cases, on a bownding zone of the

inferior surface it is covered with innumerable papille which

make these parts rough to the touch. The roughness reaches its

greatest degree of development in the Felids, in which the

individual papille stick imto the finger hke so many pins. In

the non-papillary part of the inferior surface, the mucosa is

smooth, but it may have ridges and fissures. These do not,

however, affect the smoothness.

Shape.

Mammalian tongues differ greatly in shape, being oval, conical,

spatulate, triangular, or vermiform, but these forms are not of

any value for comparative purposes.

Size.

Two sets of measurements must be made—the greatest width,

and the length—and the latter is divided into two—the lengths

of the oral and pharyngeal parts. In most cases the greatest,

width is situated at the attachments of the anterior faucial

pillars to the dorsum, but in spatulate tongues it may lie farther

forwards.

The length is measured from the apex to the mid point of the

glosso-epiglottidean fold. The oral part extends from the apex

to the median circumvallate papilla, when there is one, or to

116 DR. C. F. SONNTAG ON THE ANATOMY

the point where the lateral rows of papilla would meet the mid

line if produced backwards and inwards; the pharyngeal or

glandular part stretches from that point to the glosso-epiglottic

fold. Sometimes the smooth mucous fold from the tongue to

the epiglottis is regarded as a third zone (Owen) *.

The relative proportions which these two divisions bear to one

another differs, but the former is always the larger. Of all the

tongues which I examined, the greatest disparity between them

existed in the Indian Fruit Bat, in which the oral part measured

4-7 cm., whereas the pharyngeal part was only -4cm. long.

The Apex.

The apex may be round, flat, or pointed, and it may be entire

or notched, the latter being small, narrow and deep, or wide and

shallow. It always bears papille, but these are not always

visible to the naked eye. In the Red-fronted Lemur, the

papillae are conical and connected to a brush-like set of fine

ridges on the inferior surface. In the Californian Sea-Lion, the

long conical papille lining the broad apical notch are very |

noticeable. In the Tantalus Cercopitheque, the apex is closely

set with prominent fungiform papillee.

Text-figure 6.

eR SERN

INDIAN ABYSSINIAN

FRUIT BAT CERCOPITHEQUE

ae pas a oe ”

MLIFORMIAN BROWN TANTALUS GALEOPITHECUS

524 LION LEMUR CERCOFITHEQUE

The different forms exhibited by the apex of the tongue. Note also the median fissure

taking the form of a line of separation between two lateral masses of conical

papillee in the Californian Sea- Lion.

In Galeopithecus, and some of the Cetacea, the apex is lobu-

lated, and the lobules are fine in the former, but large and

prominent in the latter. From the notches between the lobules

in Galeopithecus, fine fissures pass backwards and inwards across

the inferior surface of the tongue. Such an arrangement is of

value for cleaning the posterior surface of the incisor teeth.

It is difficult to discover the function of the large lobules on the

tongues of the Cetacea, however.

In some tongues—e. g., the Capromys—the apex bears numerous

follicular apertures.

When the tongue is protruded in most animals the apex is

seen, but the Elephant’s apex is bound down to the floor of the

* Sir Richarl Owen, ‘The Comparative Anatomy of the Vertebrata,’ vol. tit.

p. 201.

OF THE TONGUES OF THE MAMMALIA. 117

mouth, and a piece of the anterior part of the dorsum appears to

take its place.

Median Fissures and Ridges.

A median fissure may extend backwards over the dorsum. for

a variable. distance from the apex, and it may be continuous or

interrupted. It may be merely a line of separation between two

lateral masses of conical papille, as in the Californian Sea-Lion,

or it may indent the mucous membrane to a variable depth. In

the Negro Tamarin and the Stoat, the tongue is small, and the

median fissure, which is deep, especially in the former, is a marked

feature.

The median fissure may be restricted to the front, middle,

or back of the tongue, and it may be narrow throughout,

or widen from before backwards.

In some of the Lemurs, Marmosets, and Carnivora, there may

be a median ridge instead of a sulcus.

Text-figure 7.

LAA

MACAQUES ‘SPOTTED (AVY HANGAROO AYE -AYE

Different forms of mesial furrows and ridges on the inferior surface of the tongue.

Fissures are shown in the Macaques and Spotted Cavy, and ridges im the

Kangaroo and Aye-Aye.

The inferior surface, like the dorsum, may possess a median

fissure or ridge, but either is restricted to the free part. The

muscles prevent it going any farther back. It is narrow and deep,

or broad and shallow, and it may end abruptly, or open into a

triangular depression into which the frenum passes (text-fig. 7).

In Galeopithecus, and the Spotted Cavy, small transverse

fissures pass horizontally out for a variable distance from the

posterior extremity of the median furrow. Again, the tongue of

Galeopithecus has small lateral fissures passing inwards from the

marginal lobulations.

In the Kangaroo, as in other Marsupialia, there is a firm, hard

median ridge passing back from the apex to the frenum, and, in

the Aye-Aye, a median ridge runs along the surface of the

sublingua, but it is not the same as that described above. It

represents the lytta.

Transverse Ridges and Fissures.

Transverse ridges and fissures are of two kinds—artificial

and real,—the former appearing in preserved spccimens as the

118 DR. C. F. SONNTAG ON THE ANATOMY

result of contracture of the muscles. It is necessar , therefore,

to examine fresh material if one desires to study these characters

properly. Speaking generally, the real fissures are regular and

symmetrical, or nearly so. They are complete or incomplete, the

former extending right across the dorsum, and the latter

stretching only from the median fissure to the edges of the

tongue. ‘They may be straight or curved, with their convexities

forwards.

In the Lemurs and Marmosets, they are shallow, broad, and

curved ; there are one or more complete ones behind the apex,

and several incomplete ones behind these on both halves of the

dorsum. In the Stoat, the fissures are narrow and deep, and

they pass in straight lines obliquely forwards and outwards from

the median fissure to the lateral borders of the organ.

The arrangement of the papille on these fissures and ridges

varies. In the Stoat they are only on the ridges, but in the

Lemurs and Marmosets they are on either.

Text-figure 8.

LEMURS STOAT

Different forms of transverse fissures and ridges on the dorsum of the tongue.

Intermolar Hlevation.

In many animals the posterior part of the oral division of the

dorsum is elevated, thereby producing the posterior lobule or

intermolar elevation, so-called from its situation. In the Ornitho-

rhynchusit appears as a broad and expanded part of the dorsum,

and it bears two triangular processes, or lingual teeth, on its

anterior border. These are of value to the animal when it

catches its food while swimming in the water, for they direct the

insects into the cheek-pouches, whence they can be removed for

eating when the animal is at leisure.

The Rodentia have well-developed elevations, which offer a

marked contrast to the low-lying anterior part of the dorsum.

Hach of these divisions has a different function to perform.

The papillose anterior part is gustatory, whereas the posterior

lobule raises the food up to the molar teeth.

The elevations differ in the abruptness in which they rise up

from the posterior end of the anterior division.

The Lingual Papille.

The lingual papillee are of three kinds—circumvallate, fungiform,

and conical. At the sides there are a number of fissures and

OF THE TONGUES OF THE MAMMALIA. 119°

intervening lamine termed folate papille, or, more properly,

lateral organs. In this series of papers they will be described as

such.

The Circumvallate Papille.

The circumvallate papille, also known as the fossulate or wall

papille, or, briefly, vallate papille, vary in number from one to

twelve, and the numbers may vary in different individuals of the

same species. They are the largest and, in most cases, the fewest

in number of all the papille. They are usually disposed in a

single row on each side, but there are two in the Binturong. In

the Capybara they appear as slits and ridges.

Text-figure 9.

ROO

FELINE DOUROUCOULI BLACK HOWLER CERCOPITHEQUES TREE KANGA

MONKEY

6 7

WHITE-NOSED COATI DESERT FOX RUFFED LEMUR

COMMON BADGER CHIMPANZEE

BINTURONG CAPYBARA

The different types of arrangement of the circumvallate papille.

Details in text.

Arrangement.—When there is only one papilla, it lies in the

mid line; when there are two, there is one on each side, and

they are described as right and left lateral, the tongue being

held with the apex away from the observer. If there are three,

they are grouped in the form of a triangle, and the individuals

are described as the posterior and right and left lateral papille.

In most cases the apex of the triangle is posterior, but in the

120 DR. CG. F. SONNTAG ON THE ANATOMY

Tree Kangaroos it is anterior. When there are more than three

papille, they are arranged in chains, giving the appearance of the

letter V, Y, T, or U.

Structure.—Each papilla has a cylindrical or club-shaped body

rising from the bottom of a cylindrical depression, and the space

between the body of the papilla and the wall of the depression is

termed the fossa, the bottom of which is termed the fundus.

The papilla may stand up higher than, be flush with, or

recessed below the surrounding dorsum, and the protruding

Text-figure 10.

17 18

The different macroscopic appearances presented by the circumvallate papille.

Details in text.

form is the commonest. When it is recessed, as in the Mono-

tremata and Marsupialia, the object attained is protection.

Poulton* showed how the vallate papilla of the Ornithorhynchus

consists of a ridge of delicate cells deeply recessed, and the vallum

ean contract over them to shelter them from noxious influences,

These variations are shown in text-fig. 10, nos. 1, 2, 3.

The fossa may appear as a mere slit round the papilla, or it

may be patulous, especially when the tongue is fresh. In the

Rhesus and Common Macaque Monkeys, the fosse of the two

lateral vallate papille exhibit recesses at the anterior and

posterior papillary poles (text-fig. 10, no. 4). When the tongue

has been preserved these recesses close up, however.

* Poulton, EH. B., Proceedings of the Zoological Society of Londen, 1883, p. 599.

OF THE TONGUES OF THE MAMMALIA. 121

Each fossa may contain one or more papille (text-fig. 10,

nos. 5-9). When there are two, they appear as separate

cylinders, or as two halves of the same oval. Moreover, the

several cylinders may be the same or of different sizes.

The papilla may be round, oval, or keeled, and its surface may

be smooth, polished, and glistening, or granular. When it is

granular, it may appear finely dotted, or covered with coarse

tubercles. Sometimes there is a central depression or umbilicus

(text-fig. 10, nos. 9-12). The granules may represent secondary

papillee.

The vallum may slope and appear as a mound on which the

papille are set prominently, but that is not common; it occurred

in the Indian Muntjac. It usually takes the form of a zone

round the papilla and fossa, and it may be smooth, furrowed,

lobulated, or covered with conical papillae. When it is lobulated,

the divisions may be round, oval, rhomboidal. There may be

two rows—an inner one of round, and an outer one of reniform

elements.

Text-figure 11.

LL.CP. OQ yy BLOB

The blood-vessels of the vallate papillary region of Macacus rhesus.

i.L.C.P. and R.L.C.P., right and left lateral papille; P.C.P., posterior vallate

papillee.

The tongue of the Common Badger exhibits an interesting

form of vallate papillary region. It is coloured brown, and the

papillae appear as if they are beneath the surface, and shine

through the mucous membrane, which looks as if it is composed of

transparent mosaics.

The different forms of vallum are shown in text-fig. 10,

nes. 13-20.

If the tongue is removed soon after death, the blood-vessels

supplying the vallate papillary region may be seen injected with

blood. In the Rhesus and Common Macaque Monkeys, two

arteries pass forwards in the middle line, and, when they reach the

papillary region, they divide into two branches-—a mesial and a

lateral one. The mesial branches pass between the posterior

papilla, and the lateral ones pass to the outer sides of their

corresponding papille, whenee they can be traced running out to

122 DR. C. F. SONNTAG ON THE ANATOMY

the lateral vallate papillae. When veins are visible, they pursue

the same course as the arteries (text-fig. 11).

The fresh papillee look like delicately tinted glass balls, but the

preserved ones are dead white in colour.

The Fungiform Papille.

The fungiform papille, which have a bright red colour in the

fresh tongue, are more numerous than the circumvallate and

fewer than the conical papille, and they do not exhibit the same

diversity of types as the other forms. ‘They vary in appearance,

number, and arrangement.

They are small globular bodies lying in depressions of the

mucous membrane, and are sessile or pedunculated. They project

for a variable distance above the surface of the dorsum. In most

eases the projecting part is hemispherical, but the complete

pedunculated papilla is seen in Man, the White-collared Man-

gabey, the Chimpanzee, the White-nosed Coati, and others.

Text-figure 12.

’ The different macroscopic appearances presented by the fungiform papille.

Details in text.

In Man and the Mangabey, the pedunculated papillz lie close to

the attachments of the faucial pillars to the dorsum; in the

Chimpanzee they are at the back of the tongue, and in the Coati

they take the place of the apical vallate papilla.

The papilla has a smooth, granular, or umbilicated surface, and

the umbilicus may be present in the sessileand pedunculated type

(text-fig. 12.). The different forms are not, however, charac-

teristic of any order of the Mammalia, and transitional types are

to be seen between them and the conical papille.

Fungiform papille are present on the dorsum, lateral borders,

and, in many cases, on the bounding papillary zone of the

inferior surface. They are absent in Galeopithecus and the

Arctic Fox, and difficult to see in the Dog.

As a rule they are situated at the apex, lateral parts, and

posterior division of the dorsum, and scanty or absent on the

central part. They are clustered behind the apex ; in the middle

third of the dorsum they are in transverse rows, and, at the

OF THE TONGUES OF THE MAMMALIA, 123

posterior part of the lateral part, they are in oblique chains

passing forwards and outwards parallel to the rows of vallate

papille. Those in the middle of the dorsum, in front of the

vallate papillary zone, are arranged in clusters as in Man, the

Macque Monkeys, and some of the Lemurs and Marmosets. In

the White-collared Mangabey, a V-shaped band of fungiform

papille meets with the lateral organs on each side, and partitions

off the oral division of the dorsum from the pharyngeal part

(text-fig. 13).

On the lateral borders of the tongue, the papille are arranged

in a chain, and they may be prominent or insignificant. On the

inferior surface they may either have no definite arrangement,

or they may be disposed in rows, of which the inner one is of

large, and the outer one of small elements.

Text-figure 13.

MACAQUES MANGABEY

The arrangement of the fungiform papille.

The fungiform papille may be covered with secondary ones

and possess taste buds. ‘The histology will be described in the

future papers of this series.

The Conical Papille.

The conical papille are the most numerous, and they vary

more than any other group in distribution, size, and form.

In some of the Primates they are restricted to the oral part

of the dorsum, but in most animals they are on both oral and

pharyngeal parts. When they are present on the pharyngeal

part, they are more discrete than those on the oral division.

Moreover, the character of the papillae on the two parts may be

so different that the tongue can at once be referred to the family

to which the animal belongs. In the Felide, for example, the

large recurved conical papille on the oral part of the dorsum are

pathognomonic, but the Canide have small and medium-sized

papillae on the oral part, and long and shaggy ones on the

pharyngeal part.

Arrangement.—In most animals the papille are aggregated

into clusters behind the apex; they are disposed in transverse

rows in the middle third of the dorsum and in oblique rows in

the posterior third. They are, therefore, disposed in the same

manner as the fungiform papille, but they are dotted over the

whole of the dorsum.

124 DR. C. F. SONNTAG ON THE ANATOMY

There are exceptions to the above arrangement, however: in

the Indian Fruit Bat the arrangement in transverse rows is

maintained right up to the apex, and in the Stoat there are no

transverse rows at all: they are oblique almost to the apex, and

there is a small cluster of papille behind it. In the Abyssinian

Cercopitheque the papille are closely set and not arranged in

rows at all. These various forms are shown in text-fig. 14.

Size.—The papille behind the apex are small in size, and

their true character can only be made out after magnification.

As a general rule they increase in size from before backwards,

and the central members of the rows are larger than the lateral

ones. In some of the Felide, and in the Indian Fruit Bat, there

is a central area with large papille, and a bounding zone con-

taining small ones. In some animals there is a marginal fringe

of long hair-like conical papille.

Text-figure 14.

COMMON TYPE STOAT

The arrangement of the conical papille.

The papille are cornified to a variable extent and confer a

rough feeling on the dorsum. In the Felide, the roughness is so

marked that the sharp cornified papille stick into the examining

finger like so many pins; this arrangement must be of the

greatest value for removing flesh from bones. The roughness

may vary in different species of the same genus. No contrast

could be greater, for example, than that which exists between the

tongues of the Abyssinian and Preuss’s Cercopitheque. In the

latter the tongue is comparatively smooth, but in the former it is

very rough, and, when it is examined with the hand lens, the

conical papille look like a field of long grass.

The shape and size of the papille, and the number of points,

exhibit many variations, but very few types are characteristic of

any order. The different kinds can be arranged in groups, but

there are transitional forms linking them together.

In the first, or filiform group (text-fig. 15, nos. 1-4), the

papilla is long and coarse, or fine, and a cluster may arise from a

single point. Sometimes the papilla gives off a bundle of hairs

from its trunk, or, as in some of the Marsupialia, they come off

in the form of a ring. These were called coronate papille by

Poulton. The filiform group is linked to the cylindrical (text-

OF THE TONGUES OF THE MAMMALIA. 125

fig, 15, nos. 6-9) and fusiform types (text-fig. 15, no. 10) by the

papilla shown in text-fig. 10, no. 5. Cylindrical papille may be

long or short, and bear one or more points (text-fig. 15, nos. 9,

25, 26).

Piekeasme may be recurved, and have an entire or divided —

apex (text-fig. 15, nos. 11-13), and the base from which it springs

may be smooth or granular; also the point may be rounded and

umbilicated (text-fig. 15, no. 14).

Text-figure 15.

29 30 5

The different macroscopic appearances presented by the conieal papille.

Details in text.

The triangular group may have straight, serrated, or curved

sides, and the apex may have one, two, or many points (text-fig.

15, nos. 15-18). |

The members of the oval group may have plain or pointed

sides, and the most complex form resembles the calyx and corolla

of some flowers (text-fig. 15, nos. 19-23).

In most animals the points of the papille in the centre of the

dorsum are directed backwards, and those at the sides look

126 DR. C. F. SONNTAG ON THE ANATOMY

backwards and inwards. In many of the Ruminantia, however,

there is an area at the back of the oral part of the dorsum on

which the papille, which are large, point in all directions.

The papille on the edges of the tongue give these a serrated

appearance, the points being directed backwards, and the dis-

position is in vertical chains.

On the inferior surface of the tongue the papille are small, but

the Banded Mongoose has large discrete ones arranged in trans-

verse rows with the points directed backwards and outwards.

The functions of the papille are numerous, but taste is not

one of them. They are tactile, retentive, or mechanical, and the

nature of the mechanical action differs in different animals.

In the Felide it is rasping, and in the Bats it may be suctorial

in virtue of the action of muscle-fibres near tne apex. They are

assisted in their work by the shape of the mouth, the nature and

strength of the lingual muscles, and the co-existence of ridges

and tubercles on the palate. Hard insects can, for example, “be

crushed between the cornified papilla and the palatal tubercles.

The conical papille may overlap the fungiform and vallate

papill, or there may be a non-papillary zone between them (text-

ise, IPA, sao / teh; 8)

The Lateral Organs.

The lateral organs, or foliate papille, consist of a number of

fissures and elevations at the posterior extremity of the free part

of each lateral border of the tongue. In most cases they extend

for a variable distance on to the pharyngeal part of the dorsum

each side. Both organs may be identical, or one may be more

pronounced and exhibit a greater degree of development.

The fissures may be long or short, and they may be curved

with the convexity forwards, sinuous, or straight. They may

be described as primary or complete, when they separate two

adjacent ridges, or secondary or incomplete, when they lie in the

middle of a ridge and partially divide it into two. Many tongues

have both forms. They vary greatly in number, the greatest

which I detected being fifteen in the Spotted Cavy. Both organs

may have the same number of fissures, or there may be more in

one of the two in any animal.

The ridges may not protrude above the surface of the dorsum,

as in the “Spotted Cavy, or they may appear as a series of ov al

bodies arranged in a straight line or a curve. The linear

arrangement is sgen in the Macaque and Cercopitheque Monkeys,

and the curve in the Brown Lemur and some other animals.

Again, the bodies may be grouped in pairs, as in the White-

collared Mangabey and the Bald-headed Ouakari. These modifi-

cations are shown in text-fig. 16.

Sometimes the lateral organs appear as distinct bodies which

are oval, lozenge-shaped, vermiform, or wedge-shaped (text-

fig. 16). Their margins may be serrated or plain, and their

OF THE TONGUES OF THE MAMMALIA. 127

upper limit may be bordered with conical papille or lymphoid

nodules. Both organs may be identical in size, but there may be

disparities in size, distinctness, and number of ridges and fissures.

Text-figure 16.

Te 717 7 RN 789)

SPOTTED CAVY RHESUS and CommoN BLACK HOWLER

MACAQUE MACAQUE MONKEY

Joo \o\ (Go) aallalo)

BALD-HEADED ; WH/TE-COLLARED WH/TE -FRONTED MOLOCH

QUAKARI MANGABEY CAPUCHIN TEETEE

144 ;

Cini = “=< (dave

RED-FRONTED RING-TAILEO NEGRO TANTALUS

LEMUR LEMUR TAMARIN CERCOPSTHEQUE

The different forms of lateral organ. Details in text.

To sum up, the lateral organs may consist of :—

A. Fissures and flat laminze, as in the Spotted Cavy.

B. Fissures and protruding lamine, as in the Macaques and

Mangabeys.

C. Definite organs, as in the Ring-tailed Lemur, Kabbit, etc.

These organs are richly supplied with taste-buds.

The Lymphoid Tissue and Glands.

Several papers dealing with the glands of the tongue have been

written by J. B. Haller * and others. I will not, therefore, enter

into a description here, but I will refer to it in the systematic

papers. I will only point out here that the orifices of glands are

not visible in all tongues, even when the tongue is examined with

the hand lens.

The Inferior Surface.

The inferior surface presents for examination a papillary

border, a mesial furrow or ridge, a frenum, the openings of the

sublingual glands, and the sublingual plate or its remnant the

plice fimbriatee.

The papillary border of the tongue may run completely round

it, or it may be broken at the apex. Its antero-posterior length

* Haller. J. B., Archiy fiir Mik. Anat. 1909, p. 368.

128 DR. C. F. SONNTAG ON THE ANATOMY

and breadth vary, and the papille on it are conical alone, or both

conical and fungiform; but it never bears fungiform papille

alone. The various arrangements and degrees of aggregation

have already been described, and attention has been paid to the

ridges and furrows.

Wharton’s Ducts open at the root of the frenum. In Man and

some of the Anthropoids, they open on the sublingual papille, or

caruncule sublinguales, which are overlapped by folds of mucosa

or sublingual plicee.

In many monkeys, the ducts open on the apex of a triangular

body (text-fig. 17) which has plain or serrated sides and a bifid

or entire apex. Also, the inferior surface may have small pointed

processes, as in Preuss’s Cercopitheque. In the Saki the apex

is rounded and entire, and the body bears lateral lobes ; in the

Squirrel Monkey there is a round entire apex, but the body is

not lobulated; in the Moloch Teetee the whole body is lobulated ;

and in the Black Howler Monkey the apex is bifid and the

halves are rounded. The sublingual-duct body is set astride the

frenum.

Text-figure 17.

ol ome A

MAN

PREUSS 5S

MANGA REI CERCOPITHEQUE aL

BLACK SQUIRREL MOLOCH

HOWLER MONKEY TEETEE

The different forms exhibited by the openings for Wharton’s Ducts.

Details in text.

The above body is not to be confounded with the sublingua

which is characteristic of the Lemuroidea, and represented by

the plice fimbriate of the Anthropoidea. These folds extend

backwards for a variable distance along the lateral aspect of the

inferior surface from the apex. The sublingua has already been

fully described by Poeock *.

The frenum extends from the posterior end of the free part of

the inferior surface to the floor of the mouth. It originates from

the plane surface of the tongue, or from the bottom of a

triangular depression. It varies greatly in length.

Structures for cleaning the Teeth.

Some animals have structures for cleaning the teeth, and

these may be apical, as in Galeophithecus, or lateral as in the

* Pocock, R. I., Proceedings of the Zoological Society of London, 1918, p. 19.

OF THE TONGUES OF THE MAMMALIA. 129

Ruminantia. In the Red-fronted Lemur it takes the form of a

number of long apical conical papille, but in Galeopithecws ib

appears as a number of apical lobules. In the Slow Loris it is

on the apex of the sublingua.

Apparatus for Purposes of Suction.

In some of the Bats, theapical conical papille can be arranged

in the form of a suctorial ring by means of muscular contraction.

Pigmentation of the Tongue.

The fresh tongue has a pink colour with fine dark red dots in

most animals, but some animals have peculiar pigmentation, which

may even be so characteristic that the species to which they

belong can be told at once. The Moloch Teetee has a black

tongue; the Grizzled Spider Monkey has a brown tongue with a

white cross on the dorsum; the Common Badger has a brown

vallate papillary area; and the Giraffe and Aurochs have

tongues of a leaden hue.

The disposition of the pigment varies in different members of

the same species, but the differences are not of great value for

comparative work; in the Moloch Teetee, for example, I have

seen one specimen with a completely black tongue, but another

had a wedge-shaped area devoid of pigment at the base.

The Litta.

On the under surface of the tongue, and sometimes within its

substance, there is a remnant of the glosso-hyal cartilage,

known as the litta, which varies in situation and character.

It lies below the tip of the organ in the Dasyure, but it is

absent in the Phalangers, in which it is replaced by a thickening

of the frenum. It is salle apical in the Rhinoceros.

In the Carnivora, it is developed, especially in the Dog, in

which it is termed the worm. It is elastic and assists in lapping.

In the Kinkajou, it is large, ligamentous, and ensheathed, and

in the Aye-Aye it takes the form of a nodulated and hooked

crest on the ventral aspect of the sublingua.

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ON EAST AFRICAN LIZARDS. 131

11. Notes on Hast African Lizards collected 1915-1919,

with Description of a new Genus and Species of

Skink and new Subspecies of Gecko. By Arraur

LOVERIDGE.

[Received March 5, 1920: Read March 30, 1920. ]

(Text-figure 1.)

The following notes are based on a collection of 900 lizards and

chameleons collected by the writer during the campaign in Kast

Africa. Over a theusand specimeus were collected, but one box

containing lizards frem Kerogwe, Handeni, etc., and handed to

the 8. & ‘I’. for transmission to the base, never reached its des-

tination. Another bex containing half-a-dozen jars of lizards

and chameleons was lest en route for England. Besides these

there are a good many specimens in the Nairobi Museum collected

by the writer which were net breught home, and therefore were

not available for study for the present paper.

Representatives of all the families of Kast African lizards were

taken, more than half the genera and about one-third of the

recorded species. The numbers are distributed as follows :-—

Number of

Family. Genera. Species. specimens.

Geckonidaes.0.9./.0.). we 12 255

PAvGarTIGeDs mera ee eet sth 1 5 55

Pommansiders Wy. ite eke: ] ] 12

NWiamreiatilach (5. Jeene Comer Ss 1 2 13

Amphisbeenide ............ ] 1 i

bacertidesr: AU Ay. eee: 5 6 69

Gerrhosauride ............ i 2 46

Scimeidcen twee. saat 12 330

Anelytropidz ............ 1 1 1

Chameleontide............ 2 9 112

ot lee 900

The most representative collection was made in Ex-German Hast

Africa, where the writer spent two and a half years. Six months

were spent in Portuguese Hast and one and a half years in British

East Africa. The field-notes are not so extensive as would have

been the case had cenditions not been so adverse. For nearly

twelve months all specimens had to be carried on the saddle until

camp was made, and an opportunity arose to send them back to

the base on the returning supply lorries.

The principal localities mentioned in the foliowing pages

are :—

British Hast Africa: West Mt. Kenia, Mt. Siswa, Mt. Margaret,

Kedong Valley, Thika, Nairobi, Kabete, Kagiado, Bissel, Voi.

Mbunyi, Mombasa.

2 MR, A. LOVERIDGE ON ;

Ke-German Hast Africe: West Mt. Longido, Aruscha, Moschi,

Kahe, Palms, Tsame, Gonya, Mikomasi, Mombo, Kerogwe, Han-

deni, Lukigura, Makindu (Msiha River), Ngeri-Ngeri, Mikesse,

Mkuyuni, Matombo, Duthumi, Tabora, Dodomo, Kongwa, Moro-

govo, Dar-es-Salaam.

Zanzibar.

Portuguese East Africa: Lambo (on mainland 3 miles from

Mozambique), Delagoa Bay.

The identification was eirried out at the British Museum at

South Kensington, where I had the advantage of examining the

types of a oveat many of the species referred to in the following

pages, as well as large series for comparison from many localities,

‘The notes were afterwards worked up at home.

I should like to take this opportunity of thanking Mr. G. A.

Boulenger for the great kindness which he showed me. Not

merely by according me free access to his papers and the collec-

tions in his care, but at all times so readily giving advice,

examining specimens, or making lengthy translations from the

German text. Without his kindly oversight 1 should never have

completed these notes, or, if completed, should without doubt

have made many blunders.

It is with the object of showing my appreciation for the help

so freely given that I have associated Mr. Boulenger’s name with

the only new lizard found in the collection, an interesting limb-

less hurrowing Skink described in the following pages.

Only one local race has been given a subspecific name, though

on colour grounds the Mombasa (Frere Town) form of Lygo-

dactylus pictur atus, the Dodoma specimens of Agama lionotus,

and the Longido specimens of Mabuia brevicollis might be con-

sidered by some as meriting formal names.

Measurements are given to the nearest eighth of an inch,

followed by the exact measurements in millimetres of the length

of head and body followed by the tail length when intact.

IT am indebted to Mr. H. A. Baylis for identifying the parasitic

worms found in many species of lizards. Some of these are pos-

sibly new, and as the descriptions have not yet been published,

only the generic names are given in the following pages.

GECKONID!.

HEMIDACTYLUS CrtERNII (Blgr.).

Blgr. Ann. Mus. Gen. (3) v. 1912, p. 329.

A single male was collected at Nairobi on 3.iv.15. It was

found beneath a stone on the hillside. Total length 27 inches

(38°35 mm.).

The type locality of this recently described species is in Somali-

land. The type is in the British Museum, and the above speci-

men was identified for me by Mr. Boulenger. This new record

shows the species to be widely spread.

BAST AFRICAN LIZARDS. 13333

HEMIDACTYLUS MABOUIA (Gray).

Blgr. Cat. Liz. i. 1885, p. 122.

This is undoubtedly the commonest of the East African Geckos.

Seventy-one specimens were collected. In British East at Voi ;

in German East at T'same, Mkomasi, Kerogwe, Lukiguva,

Makindu, Amani, Kongwa, Morogoro, Dar-es-Salaam, and Du-

thumi; in Portuguese Hast at Lumbo and Delagoa Bay. It is

commonly found in houses and out-buildings, but is almost as

common on trees.

The coloration is very variable. Those taken in the burnt-out

interior of trees were practically black, others hiding in crevices

of Paupau-trees were a pale straw-colour; some of the very

largest were found on trees at Lukigura and Makindu, where

their darkly barred and mottled skins by accentuating the small

tubercles led me to suppose them to be a distinct species. ‘Those

found in houses, particularly where the walls were whitewashed,

were pale flesh-colovr or almost transparent. The change of

colour may also be influenced by their feelings apparently, for on

one occasion I witnessed two of them fighting—the victor was

pale grey and the pursued brown-black.

The largest specimens were taken at Morogoro. Both male

and female measured 73 inches (86°102 mm.). It was curious

that both these large specimens should be in the same propor-

tions of head and body to tail, for the tail of the male was a

reproduced one, while that of the female was intact.

An extraordinary percentage of the Morogoro specimens had

regenerated tails, no fewer than twenty out of the thirty-one

collected. It occurred more frequently in males than in females,

and I attributed this to the combats that take place, which are

presumably amongst the males. In an outhouse I saw a speci-

men with bifid tail, but did not succeed in catching it. At

Kerogwe I caught seventeen specimens without one dropping its

tail; eleven of these had their original tails, five males and one

female had secondary growths. I put one of these Geckos into a

vivarium with an Underlined Sand-Snake (P. subteniatus); the

rapidity with which the snake gave chase was almost incredible ;

the weather being hot, the snake darted and doubled about the

case. The Gecko dropped its tail, but the snake, undeceived by

the wriggling of the severed tail, shortly after seized the Gecko,

and when the latter became limp from the poison—swallowed it.

The eggs of this species are almost globular except for a

flattening at the point of attachment; they are soft and sticky

when laid, and thus adhere to the bark, in whose crevices or

beneath which they are deposited. Two are produced at a time ;

in diameter they are from 10 to 12 mm.; the shell soon hardens

and becomes very brittle, so that it is difficult to detach them

from the bark without breaking them ; the surface of the shells

is finely granulate and the colour is opaque or bluish-white.

Eggs were taken on July 29th at Makindu, on March 9th and

134 MR. A. LOVERIDGE ON

December 20th at Morogoro, on October 16th at Lumbo. The

last-mentioned hatched out the same day; the emerged young

one measured just under 22 inches (31°31 mm.), which is extra-

ordinary when one considers the size of the egg. Two eggs

collected upon some books at Mombasa on November 17th, 1919,

also hatched out within a few days, so that the species probably

breeds at any time of the year.

Flies and spiders are their usual food, but I have also taken

small beetles, and one particularly gorged specimen had a big

brown cockroach 40 mm. long in ifs stomach.

This species is particularly liable to small red acarine parasites,

which generally are found scattered about the ventral surface.

HEMIDACTYLUS SQUAMULATUS (Torn.).

Tornier, Thierw. Ost-Afr. Rept. 1896, p. 10.

Two males only, collected at Morogoro, the first under an ox-

hide on 8.v.17 measured just under 22 inches (33°26 mm.), but

the tail was reproduced, the second on 9.iv.18was 3% inches

(42:42 mm.).

Compared with specimens in the Biitish Museum from Voi,

Tsavo, Samburu, and Kitui.

HEMIDACTYLUS BROOKU (Gray).

Blgr. Cat. Liz. i. 1885, p. 128.

Thirty specimens were collected at Morogoro, Duthumi, and

Lumbo; at the former place they were mostly to be found in

grass-huts and among rubbish heaps, but at the other localities

they were all found on trees.

The largest male measured 3} inches (40°40 mm.) and was from

Lumbo; the largest female measured 34 inches (54°33 mm.),

though the tail was reproduced: it was taken at Morogoro.

Female with eggs taken 14.11.17.

‘The types of this species from Borneo and Australia in the

British Museum were examined, as well as the fine series from

African localities. A. brookii has a wonderfully wide distribu-

tion in Kast Africa from Somaliland to Portuguese Hast ; it also

occurs on the West Coast.

HEMIDACTYLUS RUSPOLIL (Bigr.).

Bler. Ann. Mus. Gen. (2) xvii. 1896, p. 6.

Seventeen specimens, of which three were from Mbunyi, B.E.A.;

one from Longido West; twelve from Morogoro; and one from

Duthumi, G.E.A. The Mbunyi and Longido specimens were

taken from fissures and crevices of thorn-trees, the Morogoro

ones beneath stones and dug out of an earth-bank, where they

lived in holes.

Six of these were males, of which the largest measured 5 inches

(67-60 mm. regenerated), from Mbunyi. By far the largest of

ee neghiee

EAST AFRICAN LIZARDS, 135

the eleven females was from Duthumi; this specimen measured

63 inches (77°82 mm.). In both of these the tail was repro-

duced and carrot-shaped ; the female was strikingly so, measuring

17 mm. across at the base and tapering to a point; the body only

measured 23 mm. across. In life these Geckos have an orange-

colour, much brighter on the tail; this was so as regards the

Morogoro specimens, but I did not notice it at Mbunyi.

The following is a note from my diary of the colour of a living

Morogoro specimen :—‘‘ The head and back are of a very dull

orange-colour; a black stripe passes through the eye; three

pairs of rather indefinite ocelli on the sides are connected by

black saddle-like markings bordered with whitish granules; the

tail is brilliant orange ringed with black, the rings becoming less

distinct on the lower surface; throat, belly, and under-surface of

legs transparent white.”

LYGODACTYLUS CAPENSIS MOSSAMBICA, Subsp. nov.

Blgr. Cat. Liz. 1885, p. 160.

Fifty specimens of a Gecko were collected at Lumbo, P.E.A.,

which agreed with the description of L. capensis (Smith) in all

particulars, with the exception of the scaling on the underside of

the tail. In ZL. capensis the underside of the tail is covered with

large imbricate scales except in regenerated tails, where occa-

sionally one finds broad transverse scales in a median series. In

all the specimens collected at Lumbo this transversely dilated

median series was a constant feature both in the original and

renewed tails.

There is an exceptionally fine series of LZ. capensis in the

British Museum, and specimens were examined from the following

localities :—South Africa; ‘Transvaal (Zoutpansburg, Rusten-

burg, De Kaap Goldfields); Natal (Lower Unkomaas River) ;

Rhodesia (Mazoe); Zululand (Indukuduku, Ngoye Hills) ;

Portuguese Gazaland (Jiku, Kurumadzi River); Portuguese

Kast Africa (Delagoa Bay, Shire Highlands); Mashonaland (Mt.

Chirinda) ; Tanganyika: French Congo( Benito River): Nyassa-

land (Fort Johnson, Zomba); Angola (Chiyaka District); Ben-

guella (Interior); South Somaliland (Lush). All these agree

with the original description.

Specimens from Beira and Cogano, P.K.A., however, were

intermediate between the typical L. capensis and the Lumbo race,

one or two of the Beira specimens being practically indistinguish-

able from those collected at Lumbo. In view of the large series

of specimens collected and the constancy of the character, I

propose to designate this local race as a subspecies under the

name of mossanbica.

The largest male measured 22 inches (35:31 mm.), and the

largest female just over 23 inches (35°22 mm. regenerated).

These specimens I regard as the types. The average length of

nineteen males was 61 mm, (30°31 mm.), and of thirty-one

136 MR. A. LOVERIDGE ON

females 57 mm. (29°28 mm.); a few specimens with regenerated

tails are included in these averages.

Coloration from notes made from the living Gecko, as follows :—

‘“Grey or olive-brown above, darker or lighter according to.

habitat. Black lateral lines spotted with cream commence at

nostrils and disappear at, or on hind legs. A pair of dorso-

lateral lines bordered on their inner edge by fawn-coloured lines

commence on frontal region and vanish on tail. In some speci-

mens these lines were broken into a series of dots, in others they

were very indistinct. Whole dorsal surface much mottled with

darker and lighter spots. Throat pure white, bespeckled in

males; rest of under surface yellowish-white. Regenerated tails

plumbeous.”

Eggs two in number, pure white, more bird-shaped than most

Gecko eggs. Measured7 mm.x6mm. Laid in crevices of bark.

Eggs collected on 27. vii. 18 hatched on 16.x.18. Newly emerged

young measured 24 mm. (13°11 mm.).

Blue-bottle fly, large beetle larva, and small brown beetles were

found in stomach. On one occasion I saw a young Gecko seize a

small staphylinid beetle and drop it quickly, shaking its head

vigorously as does a person after taking a nauseous draught.

The type specimens and others have “heen given to the British

Museum. Co-types have been donated to the National Museum

of Wales, Manchester Museum, American Museum of Natural

History, Smithsonian Institute, Prof. Barbour at Harvard

University, and Nairobi Museum, British Hast Africa.

LYGODACTYLUS FISCHERI SCHEFFLERI (Sternf.).

Sternfeld, Ergebn. Deutsch. Zentr. Afr. Exp. 1918, iv

p- 206.

Two males taken on thorn-trees at Mbunyi (15.v.16). The

larger measures 27 inches (26°30 mm.).

LYGODACTYLUS GROTEI (Sternf.).

Sternfeld, §.B. Ges. Naturf. Berlin, 1911, iv. p. 245.

Twenty-one specimens in all were collected—nine males, nine

females, and three immature young. Localities—Morogoro,

Msiha, Duthumi. The favourite haunts of this species were the

banana-palms and paupau-trees, on whose stems they disported

themselves in the sunshine; occasionally they were found on

shrubs, stumps, or low bushes.

The largest male measured 2? inches (32°38 mm.), the largest

female 22 inches (33°37 mm.). On February 24th, 1917, a pair

were seen ti coitu on a spray of mimosa thorn. It was about

8 a.m., and the sun was causing the heavy dew-drops to glisten.

LycopactyLus PicturAtus (Peters).

Blgr. Cat, Liz. i. 1885, p. 161.

Forty specimens available for present paper; large series from

EAST AFRICAN LIZARDS. ey

Kerogwe and Handeni were lost. in transit. Seen at Mombasa

Island, Zanzibar, Mombo, Palms, Ngeri Negeri. Collected at

Kerogwe, Handeni, Morogoro, and Dar-es-Salaam.

A purely arboreal Gecko living on tree-trunks; the following

notes were made at various localities :—

Zanzibar (30. xi1.14). Was disappointed in not securing a

Gecko, whose body was bark-colour but whose head was like a

patch of yellow lichen.

Handeni (26. vi. 16). Caught sight of a small grey Gecko with

a yellow head similar to those seen at Palms a few days ago.

They frequent large trees and come down within a couple of feet

of the ground ; should anyone approach, they glide round to the

opposite side of the trunk and then hasten towards the top. On

two trees the yellow heads of these Geckos rendered them con-

spicuous, but on a third, which was overgrown with tufts of grey

and yellow lichen, the yellow head broke up their outline and

rendered them inconspicuous; this seems to be the probable

explanation of their somewhat peculiar coloration. By far the

most interesting thing about them was the tip of the tail, upon

which were transverse lamelle similar to those on digits of most

Geckos. This arrangement serves them as a fifth foot, thus pro-

viding additional grip.

Handeni (27. vi.16). Hunted most of the likely trees for

Geckos similar to the specimen. taken yesterday; most of them

were too quick for me, save those on one stump not more than

ten feet high. In passing this, I fancied seeing something glide

round, so dodged to and fro until I caught sight of a retreating

tail. Having broken off all the smaller twigs, I put my arms

around the stump near the ground, then ran them up to a height

of five feet, where I tied a white handkerchief loosely around to

keep them from coming down. Then swarming up, I broke off

all the remaining branches, raised the kerchief within a foot of

the top, and then captured one by one the male and two female

Geckos which were there. The male was very handsomely

coloured, with a velvety-black throat merging into a bright

yellow stripe along the centre of the belly. The grey and yellow

of the back and head (upper surface) were also much more vivid

than in the specimen taken yesterday. The females possess the

power of changing colour with great rapidity, bemg quite brown

when first sighted, but changing to a dull imitation of the male

when pursued.

Kerogwe (5.vii.16). The Yellow-headed Geckos are very

abundant here, almost every third tree is inhabited by them.

Several pairs of their hard-shelled eggs were found under the

bark of the trees.

Morogoro (2.11.17). Yellow-headed Geckos very abundant

hereon the trunks of the Acacia-trees, which border many of the

roads ; these Geckos are always to be found on the sunny side of

the tree-trunks during the mornings. The male is handsome

blue-grey with a brilliant yellow head, which fades in spirit, so

138 MR, A. LOVERIDGE ON

that only some longitudinal dark striations or mottlings are to

be seen in the preserved specimen. The chin and throat in adult

males only are velvety black, extending back as far as the fore

legs ; immature specimens have unmarked throats or like those

of females. ‘The rest of the under-surface as far back as the vent

is orange, whilst small yellow patches mark the underside of the

legs. The tail is grey beneath. The throat of the female is

white mottled with a network of black lines; the upper surface

of the head in adults is pale yellow, barred or mottled with

yellow; the back is brown or greyish, with indistinct darker

markings. When struck sharply on the back with a cane so as

to be instantly killed, this Gecko frequently falls over backwards,

and remains attached to the tree by the sucker-apparatus of the

tail tip.

Lang, in his field-notes on Lygodactylus picturatus gutteralis *,

speaking of this tail, writes: ‘but the really unique feature

distinguishing it from all. other Geckos is the adhesive pad on

the tip of the tail”; this is evidently a slip, as this is a generic

character +.

The largest males taken measured 33 inches (43°43 mm.) and

largest female 33 inches (3841 mm.). It is very unusual for this

species to drop its tail, and most of the specimens taken had

intact tails.

At Morogoro (1.1.17) a pair were seen courting. ‘ After

recent heavy rains the sun came out bright and fresh this

morning, and almost every tree in the avenue had a pair of

Yellow-headed Geckos in brilliant colour on its sunny side. A

pair were courting, the female chasing off the male every time he

approached. He arched his neck in an unusual manner and ex-

posed his throat, presumably to exhibit the velvety-black patch

peculiar to the male.”

Eggs were found at Morogoro on 1.11.18, at Dar-es-Salaam on

11.iv.18 (in one group of three it is just possible that two Geckos

had laid together; two eggs being the usual number), and at

Kerogwe on 5. vii. 16.

In the ‘Catalogue of Lizards’ the colour of L. picturatus is

given as “head and anterior part of body bright yellow, with

dark brown or black lines and spots.......” At Frere Town,

which is situated on the mainland opposite Mombasa Island,

from which it is only separated by a channel not more than

500 yards across, is a very distinct form whieh has no yellow

head, nor yellow on any part of the body. When first seen I

thought it was a distinct species, but after a careful examination

T can detect no structural difference between it and picturatus,

which I have seen on Mombasa Is. though never collected. In

the British Museum is a specimen of the Frere Town form,

labelled ‘‘ Mombasa,” and collected by A. Blayney Percival, Esq.,

* Schmidt, Bull. Amer. Mus. of Nat. Hist. xxxix. 1919, p. 465.

+ Mr. Schmidt points out to me that the lamelle on the tail are not even a

generic distinction, as they are also found in Diplodactylus palmatus Mocq.

EAST AFRICAN LIZARDS. 139

which conceivably may have been taken on the adjoining main-

land. In the Nairobi Museum are two specimens collected at

Jilore, Giriama, and presented by Mr. T. B. Nair, which are also

the Frere Town form. Unfortunately the colouring of the head

does not remain in preserved specimens.

The coloration of the Frere ‘Town form in life is as follows :

Male: Head white, transverse black band across supra-ocular

region, a second in parietal region; three longitudinal black

lines unite these two to form a gridiron pattern; a third cross-

band on nape. A double row of large black spots, sometimes

united to form transverse bands, on back from nape to base of

tail ; there may be as many as eleven pairs of spots. A black

stripe commencing at nostril passes through eye and over fore-

limb, where it ends above axilla. Very narrow black line

borders tips of upper and lower labials, continues below ear-

opening and on to the fore-limb, where it disappears before

reaching elbow. Nine to twelve vertical black stripes of

irregular shape along each side. Throat black, not extending

beyond an imaginary line drawn from ear to ear, except for two

narrow line-like continuations, only a couple of scales in width,

which extend back to fore-limbs. Belly white, projecting for-

ward into black throat like the arms of the letter ‘‘U.”

Female as in male, except for white throat, which has an inner

and outer chevron-shaped black marking.

Fourteen specimens were collected. of which the largest male

measured 2% inches (36°36 mm.) and female 3 inches (36°38 mm.).

PLATYPHOLIS FascraTA (Blegr.).

Blgr. P. Z. 8. 1890, p. 80.

Two males and a female taken on thorn-trees at Mbunyi.

Largest male 4 inches (64°37 mm.),and female 5 inches(80°45 mm.).

Both specimens had reproduced tails. Two ovules 2 inch diameter

in females.

The type specimen on which the genus was founded as well as

the species was collected at Mombasa; another specimen in the

British Museum which was examined came from Maziwi, which

the label states is between Mombasa and Kagiado, therefore not

very far distant from Mbunyi. Yet another specimen has been

recorded by Boulenger from Upper Ganali, Juba River.

PHELSUMA LATICAUDA (Boettg.).

Blgr. Cat. Liz. i. 1885, p. 215.

Two males taken at Dar-es-Salaam (10.xi.18) measure

43 inches (59-53 mm.) and 42 inches (59:60 mm.). I should not

have got these specimens, which dwell in the tops of the lofty

palms, had I not been passing as some natives were cutting

branches for thatching ; three specimens were seen but only two

captured. These differ from one another in several respects :

the first has a median cleft on rostral, absent in the second ; it

140 MR. A. LOVERIDGE ON

has 9 upper and 8 lower labials. It has three scales: between

naso-rostrals, and the dorsal scales are distinctly keeled; in both

of which points it differs from the other specimen, which has two

scales between the naso-rostrals, and the scales unkeeled on the

dorsal surface of tail.

Colour-notes in life are: ‘‘ Dark green colour, finely freckled

with red on back and base of tail.”

It has been previously reported from Nossi Be, Johanna,

Farquhar Is., Zanzibar, Comoro Is., Madagascar, but not from

the mainland of East Africa.

ELASMODACTYLUS TRIEDRUS (Bler.).

Blgr. Rev. Zool. Afr. ii. 1913, p. 104, pl. v. fig. 2.

Two specimens of this rare Gecko were taken—a male from

Kongwa (21.iv.17), measures 53 inches (70°67 mm.); and a

female whose locality is somewhat uncertain, though I believe it

was taken in the neighbourhood of Morogoro, measures 44 inches

(57°57 mm.).

AGAMID4.

AGAMA COLONORUM (Gray).

Blgr. Cat. Liz. i. 1885, p. 356.

Thirty-four specimens were obtained at Thika, Gonya, Kongwa,

Morogoro, Mkuyuni, Duthumi, and Lumbo. The largest Morogoro

male measured 9% inches (121-111 mm., tip of tail missing). largest

Morogoro female 11 inches (105°175 mm.), seventeen specimens were

collected at Morogoro. Of fourteen specimens coilected at Lumbo

the largest male measured 13 inches (116 213 mm.), and female

12 inches (104°204 mm.).

Coloration was very variable; the following note was made on

a Lumbo male :—‘* Dirty cream or pale grey, with dark brown

vermiculations. Dorsal crest pale blue, particularly bright on

neck. Head dark brown above, vermiculated with same on sides ;

indistinct red stripe from eye to eye: irregular line of a fine

brick-red colour originates behind ear and is lost in a large patch

of same colour just above fore-leg. Chin vermiculated with rich

blue, converging to a blue patch on throat. Belly and under-

surface of tail dirty white.”

A female was killed at Morogoro on 14 i1.17 with 12 eggs in

ovary measuring 15x10 mm. Another was killed on 1.iv.18

with 10 eggs 18x10 mm.

Ants of several species, including the fierce little cock-tail

species, beetles, large cricket, and millipedes were found in

stomachs.

Physaloptera was taken in the stomach of this species.

This species is very arboreal; if found upon the ground it is

usually not far from a tree, to which it flies for refuge.

A very large specimen was found partly digested in the

stomach of a Hissing Sand-Snake (Psammophis sibilens) at Lumbo

(Get S)INS)))5

EAST AFRICAN LIZARDS. 141

Many specimens are infested with a small acarid (Pterygosoma

agame) beneath the ventral scales. A new nematode (Oochoristica

agame) * was found in several.

AGAMA FLAVICAUDA (Werner).

Wern. Zool. Anz. xx. 1897, p. 264.

Agama caudospina Meek, Field Mus. Nat. Hist. vii. 1910,

No. 11, p. 407.

Several specimens in the Nairobi Museum were collected by the

writer at West Kenia (23. xi.15). A large male measures 83? inches

(100°120 mm.). These specimens were taken in the thatch of

a pig-sty, and in an isolated pinnacle of earth where they had

taken up their abode in the holes excavated by Sand-Martins,

as well as in natural crevices. There are also specimens in the

Museum collected by Mr. Allen Turner at Kegamaia, near

Mt. Elgon.

There can be no doubt that Agama caudospina recently

deseribed by Meek from Elementeita, B. E. A., should be relegated

to the synonymy of this species. Werner’s s description was based

on a single male of unknown locality. Both names are very -

descriptive of this species, which is quite unlike any other Agama

collected. :

AGAMA Lionotus (Bler.).

Bler. P. Z.8. 1896, p. 214, pl. viii.

Thirteen specimens from Voi, Mbunyi, Longido West, and

Dodoma. Was also seen at Kahe and Kongwa; there are

specimens in Nairobi Museum collected by the writer near the

Kedong Valley.

The type specimen from 8.E. of Lake Rudolph, as well as

specimens from Mt. Kenia and Ngaya (south of Lake Victoria),

B.E.A., which are in the British Museum, were examined.

Arranging the specimens geographically from the type locality

southwards one finds a marked increase in the scale-rows.

Rudolph. M. Type. 65 scale-rows round mid-body.

Voi. NevwiGamelG: 03 .

Mibmnyis Mor ye. 75

ioncado: wee Mey eel 16.’ 80

iDedomany Mia 7 Sqxa18.). 79

9 F. be) 78 33 39 99

” M. ” 80 ” ” 5

9 Bp ” 82. ” ” ”?

i M. : 87

I draw attention to this for a purpose. These Dodoma speci-

mens are strikingly different in life to those collected elsewhere.

The throat of the males from other localities was invariably

# Baylis, Parasitology, x1. 1919, p. 409.

+ 3 males examined.

142 MR. A. LOVERIDGE ON

scarlet in the breeding adults. J was at once struck by the hand-

some throats of the Dodoma specimens as they bobbed their heads

up and down upon the rocks where they basked in the sun. The

throat of these males has a pear-shaped scarlet patch 14 scales

wide and 30 to 32 scales long; it is surrounded by a rich navy-

blue border 9 scales in width; outside the posterior part of this

is a semicircle of scarlet commencing narrowly at base of jaws

and widening on the throat. ‘These males are tar larger than the

type or any collected elsewhere. Both the largest males measured

114 inches (140150 mm., tails missing), the largest female

measured 112 inches (112:°173 mm.).

There appear to be no scale characters whereby the Dodoma

specimens can be distinguished ; they have a far larger number

of scale-rows round mid-body than the type, but as the cross-

country series show an intergradation in this character, it appears

to be a pity to multiply local races by giving them names.

A couple of notes made on Longido specimens are interesting,

as they show something of the chameleon-like possibilities of this

Agama :— 28. i. 16. ‘‘ Was successful in shooting one of the scarlet-

headed, blue-bodied Agamas. As soon as it was put in formalin

allthe bright colouring departed, and it became a study in browns.”

2.11.16. “ A brown Agama was basking on a rock, and I killed it

very suddenly with a smart blow across the back, causing it to

fall over with a little quiver, its back evidently broken. I placed

it in a black bag, and on my return to camp was surprised to find

its head of a brilliant scarlet and the body bright blue, exactly

the reverse of what occurred with a shot specimen a few days ago.

It appears probable that all the Agamas hereabouts are only

colour variations of the one species.” Of a male shot in the

Kedong Valley I wrote, 19. vii. 15: “Shot a brilliantly coloured

Agama, head rich brick-red, and the body, more particularly

underneath, a vivid ultramarine blue.”

All the specimens were found upon rocks and the same type of

sandy thorn-bush country.

The two Dodoma females contained eggs, the larger having 8

and the smaller 7, measuring 20 x 12 mm. (8. xii. 18).

AGAMA VAILLANT! (Blgr.).

Blgr. Ann. Mus. Civ. ser. 2, vol. xv. 1895, p. 12.

Five specimens in all were obtained, two being from Mbunyi

and three from Voi. One of the Mbunyi specimens measured

85 mm. in body, but the tail was mutilated. The largest Voi

specimen measured 93% inches (80:170 mm.). Found on reddish

sandy soil, their colouring rendering them inconspicuous,

AGAMA ATRICOLLIS (Smith).

Bler. Cat. Liz. i, 1885, p. 358.

Specimens in the Nairobi Museum were collected in the neigh-_

bourhood, where it 1s very common on trees. Two specimens

EAST AFRICAN LIZARDS. 143

collected at Dodomo on thorn-trees appear to belong to this

species. They are very small if so, the largest measuring

9 inches (90°140 mm.). Both are males with vermiculations on

the throat, which has a blue patch on the basal portion.

A female taken at Nairobi (3.iv.15) had 10 eggs in ovary,

20 x 10 mm.

ZONURIDS.

ZONURUS TROPIDOSTERNUM (Cope).

Blgr. Cat. Liz. ii. 1885, p. 252.

A single specimen was taken at Makindu (Msiha River) and

eleven at Morogoro. ‘The largest male (Morogoro) measured

7s inches (90°90 mm., tip of tail missing), the largest female

(Makindu) measured 6? inches (95°75 mm.).

These specimens entirely bear out the remarks made by

Nieden* after an examination of nineteen specimens from many

localities in G.H.A. All the specimens agree with Z. tropido-

sternum in the roughness of the head-scales, sand the granular

interstices between the flank-seales. All except two agree with

tropidosternum in that the fronto-nasal scale is in contaet with

the rostral. In these two the nasals separate the fronto-nasal

from the rostral, which is the key character for Z. cordylus. No

reliance can be placed on the arrangement of the head-scales in

this genus as a guide to specific character under these circum-

stances. It is rather interesting to note that one of these

specimens in which fronto-nasal and rostral are not in contact

was taken on the same day with a normal male and female, and

not only so, but within a few yards of them, two of the three

were seen to emerge from a hole at the base of a wall; the third

was killed at the same wall only a short distance from the hole.

Two specimens were found in a half-drowned condition in road-

side gutters, having evidently been washed out of some retreat

by the heavy rains. The favourite haunt appeared to be hollow

trees, into whose interiors they would retreat and from which it

was difficult to get them. The Makindu female was brought into

camp in a hollow log which had been cut for fuel; she had

remained while it was chopped down. She had four large eggs in

the ovary.

Termites were taken from the stomachs of four of the

specimens.

Parasitic worm (Oocharistica zonuri) proved to be new +.

VARANID4.

VARANUS XANTHEMATICUS ALBIGULARIS (Daud.).

Blgr. Cat. Liz. 11. 1885, p. 307.

The only specimen of the White-throated Monitor met with

* Nieden, Mitt. Zool. Mus. Berlin, 1913, vii. p. 71.

+ Baylis, Parasitology, xi. 1919, p. 406.

144 MR. A. LOVERIDGE ON

was found preserved in a German house at Morogoro. I do

not believe it was collected in the neighbourhood. ‘Total length

28 inches (300°410 mm.).

VARANUS NILoTICUS (Linn.).

Blgr. Cat. Liz. ii. 1885, p. 317.

Met with at Gonya, Msiha, Morogoro, Dar-es-Salaam, Duthum1,

and Lumbo. At the last-mentioned place I did not see it myself,

but heard of it several times, and the probability is that 1t was

this species and not the white-throated.

The following are the measurements of those obtained :—

M. Head and body 554mm. Tail 850mm. Morogoro. 19.1.18.

» » 460 », 850 8.1.17.

M. ‘ » 407 pH OV Dar-es-S’m. 15. vi. 18.

M. at ey watt », 490 Morogoro. 1. iv. 18.

F. 3 » 3870 » Oe ee Liv. 18.

M. 3 » 230 > oO 35 13.11.18.

No} HS “, 1653 » 235 a Q7. iii. 18.

It, \ » 142 » 240 Gonya. 29.v. 16.

M. a Pe 5 lee 5, 190 Morogoro. 10. iv. 17.

M. 5 pp, dlPA0) py LL 7/) 55 liv. 18.

2 . ao ,» 170 i 1. xi. 16.

Skin. ,, » 435 » Gal Msiha. 14. yi. 16.

In some of the foregoing the tail was missing at the tip.

Along the river-bank at Morogoro they were very common,

though more often heard than seen. The first intimation that

one was in the vicinity would be a rush through the undergrowth

followed by a splash. If you were fortunate you might be in

time to see the creature emerge on the opposite bank and crawl

into its hole. At other points along the river where the banks

were high and cliff-like they might be seen basking on some ledge

or drawing themselves up with the aid of their powerful claws.

A good many were captured alive. The usual procedure was for

me to wade down the river, with a native walking through the

undergrowth along either bank a little im advance. On the

Monitor taking to the water, I remained quiet and watching where

it emerged and marked down its hole. We could then dig out

the hole until the creature was located at the terminus. The hole

would then be closed with the shovel till only a small opening

remained, large enough for its head to come through, but not to

let the body pass if it madearush. As soon as the head made

an appearance, a widely-forked stick would be placed on it by a

person standing above and the animal held to the ground. The

spade would be pulled away, and a few exciting moments followed

as the creature struggled, scratched, and lashed about with its

tail. As soon as a favourable moment occurred the tail would be

seized with one hand, and with the other (wrapped in a cloth for

preference) the Monitor would be securely grasped by the neck

and transferred to a sack,

EAST AFRICAN LIZARDS. 145

It is a most awkward creature to handle; not only can it give

a severe bite, but a ash from the tail of even a small specimen is

severe, owing to it turning the dorsal keel over sideways as it

strikes. It makes good use of its claws to scratch when seized,

and on this account alone I have dropped Monitors which I had

securely by the neck,

Hearing that a couple of very large Monitors paid a daily visit

to the cook-~house, which was situated at the edge of a bank or

slide which sloped steeply down to the river nearly 200 ft. below,

I walked along the opposite bank of the river one day. A bell

was rung at 12 P.M. each day, and from 12.30 to 1.30 P.M. all is

quiet. It is then that the reptiles came up for scraps. Whether

the ringing of the bell had any significance for them it would

be difficult to say. Whilst walking along the opposite bank I

descried one of the lizards lying just below “the top near the cook-

house, but hidden from view from anyone on that bank. After

crossing the river I scrambled up thirty foot or so of the slide,

and found that the Monitor had disappeared. Even as I looked,

however, its head appeared over the top of the bank, and I fired

at it with a*22 Winchester. The bullet caused it to bound over

the bank where it lay quiet, for as it heard the bullet whistle

past, if imagined the danger came from above.

In its new position it exposed its whole length to me, and I put

three bullets into it as fast as I could load; after each it gavea

jump, but kept under the bank. Someone, hearing the firing,

came to the edge of the bank and looked over, thus disturbing

the Monitor, which fled down the bank like a great dog, disappear-

ing into some bushes on the brink of a cliff that rose shear from

the river forty feet below. I feared that it had gone over this,

but my boy retrieved it from the very edge. It was not in the

least spoilt by the three bullet-holes, and I had to give it a

tremendous dose of chloroform to kill it. The creature was a

male and measured 55 inches over all. Its stomach contained

meat from the cook-house and crabs. Crabs’ claws on the partly

submerged rocks in the river are generally a sign that, Monitors

are in the neighbourhood. In the stomach of another specimen I

have found the remains of a toad. As is well known, they often

come to fowl-houses for the eggs, which seems to be one of their

favourite articles of diet.

Ticks are commonly found about the anal region of Monitors.

A worm, Tanqua tiara (v. Linst), was found in one specimen

at Morogoro (6. iv. 18).

AMPHISBHZNIDA.

Monore.tis cotopura { Bler.).

Blgr. Ann. 8. Afr. Mus. v. 1910, p. 495.

Of the ten species of Amphisbeenide found in East Africa this

was the only one met with by the writer. his is its first record

Proc. Zoot. Soc.—1920, No. X. 10

146 MR. A. LOVERIDGE ON

from the East African coast I believe, as it was described from

three specimens collected in Barotseland by the Rev. L. Jalla.

Type in the British Museum.

A male and half-a-dozen females were taken at Lumbo,

P.E.A., between August 20th and October 31st, 1918. The

male measured 151 inches (346°44 mm.) and the largest female

192 inehes (440-50 mm.).

By Europeans and natives alike this strange creature was

called a snake. It is decidedly more like a flesh-coloured worm

than a lizard. In life the skin is loose and moves freely over the

body ; it is so transparent that one may see the pulsation of blood

in the blood-vessels. The scales, which are almost square in out-

line, are united in rings around the body. The eye is barely

distinguishable as a small black speck. ‘The little white tongue

is continually extruded from the mouth after the manner of

snakes. he mouth is situated on the lower surface as in bur-

rowing snakes (Lyphlops), but not so pronouncedly as in sharks.

The rostral shield is enormously developed and spade-like. The

tail is abruptly truncated, ending in a bone-like knob or shield,

doubtless developed for the same purpose as the terminal shield

characteristic of the Indian burrowing snakes of the family

Uropeltide. ;

Two specimens taken at the end of August contained 4 eggs

each; these measured 35x10 mm. and 35x 9 mm. respectively.

Another specimen laid 4 eggs either during the night or in the

early morning of September 20th. No two of these eggs were

of the same size; their measurements in millimetres were as

follows :—35 x 8, 32x 8, 309, 26x 9.

No trace of food was found in the stomachs of any of the

specimens.

At 2.15 in the afternoon of September Ist I was called to catch

a “snake”; the sun was beating fiercely upon the sand at the

time. The “snake” proved to be one of these lizards, which had

come to the surface and was wriggling about on the scorching

sand. On drawing out the last few inches of the creature which

still remained in the sand, the cause of its appearance upon the

surface at such an uncongenial hour was apparent. Its vent and

tail was smothered in ants of a subterranean species, which I have

previously noted will eat a dead body from beneath, but appears

to hate the light.

At 3 p.m. on September 20th I obtained another specimen

under precisely similar conditions, though in this case the lizard

was wriggling along the surface of the ground, leaving a trail of

ants behind it, while only a few were still clinging tenaciously to

its tail.

The following morning yet another was brought to me. It had

severe hemorrhage in the intestinal region, and died during the

day. As it was found above ground, I have no doubt that it was

also a victim of the voracious ants.

EAST AFRICAN LIZARDS. 147

LACERTID &.

GASTROPHOLIS VITTATA (Fischer).

Blgr, Cat. Liz. 11. 1887, p. 7.

Two specimens of this scarce lizard were obtained ; both were

females, and neither specimen had the long tail intact. The larger

was found bottled without data in a German house at Morogoro.

Head and body measured 37 lage (83°? mm.). The smaller was

taken at Lumbo, and measured 23 inches (67-? mm.) in head and

body. Type from Zanzibar in Brit. Mus.

LACERTA VAUERESELLI (Tornier).

Torn. Zool. Anz. 1902, xxiv. p. 701.

Two specimens were obtained. One was shot at Ngong high

up on a tree-trunk; as this specimen is in the Nairobi Museum

at time of writing I am unable to give its sex or measurements.

Ngong Forest edge, 20.1x. 15.

The other is a male caught at Parklands Forest edge on 28.ix. 15.

It measures 6% inches over all (60°96 mm.) The tail is possibly

regenerated. ‘The frontal scale is of equal width along its length,

not broader anteriorly. It has nineteen femoral pores on the

right leg and eighteen on the left.

Nucras Emini (Blgr.).

Blgr. Ann. & Mag. N. H. (7) xix. 1907, p. 488.

Three specimens were caught in B.E.A. and G.K.A. In each

case they were running about in sandy paths or places. Their

measurements are as follows :—

Male. Kagiado, 28. xii. 15. 6 inches (68°80, regenerated).

Female. Bissel, 4.1.16. Gree, ald? Ss arnsem

Female. Longido West, 1.11.16. 63 ,, (65. 106 mm.).

LarasTIA LONGICAUDATA (Reuss).

Blgr. Cat. Liz. ii, 1887, p. 55.

Seven specimens in all. A young one at Voi, 17.vi.16.; a

half-grown one from Mbunyi, 15. v.16; and five adults, of which

four were females, from Dodoma, 8.xii.18. At no other place

did I see such fine specimens of this handsome lizard as at

Dodoma. Adl three localities were sandy with scattered thorn-

bush, beneath which the lizards quickly took cover ; owing to their

agility i in so doing. only a few specimens were taken, though the

creature was abundant at Dodoma.

The largest male measured almost 12 inches (100-202 mm.)

anil the largest female 124 inches (95°220 mm.). The male’s tail

appears to be intact. One of the Dodoma females had ten eggs In

the ovary. Remains of beetles were common in their stomachs ; :

one had a grasshopper, whilst another had gorged on a false-

spider (Solifugid), and the remains of a false-spider’s jaws were

Lo*

148 MR. A. LOVERIDGE ON

found in yet another lizard. How they manage to eat such an

unpleasant mouthful as a false-spider is difficult to comprehend.

LATASTIA JOHNSTONI (Bler.).

Blgr. Ann. & Mag. Nat. Hist. (7) xix. 1907, p. 292.

Sixteen specimens taken in shambas and open patches of waste

ground at Morogoro. As Hrenvias spekii was found in the same

spots, I had a little difficulty in distinguishing the species when

they were running about. JL. johnstoni has a trick, however, of

rushing in one direction and then facing about with a little leap,

so that it is looking in the direction from whence it came. The

tail is also noticeably longer, and has an illusory semi-transparent

pinkish effect which disappears after death. It was not nearly so

comion as //. spekii and more difficult to catch.

Colour notes made during life are as follows :—‘ Ground-colour

pale brown. Four parallel cream lines commence at parietals,

the outer pair extending forward along outer bordex of parietals

to the eyes, posteriorly they converge to form a single dorsal line

on anterior portion of tail, An upper pure white lateral line

arising in the sub-ocular passes over ear and is lost on tail. A

lower pure white lateral line commences in upper labial region,

passes through ear and over fore-limb, is interrupted by hind-limb,

but re-commences after and merges into the white on underside

of tail. Six or more yellow spots on side between upper and

lower lines whose continuity they may break, alternating black

bars and red blotches on sides. 4th to 8th upper labiais yellow.

Throat and underside of body china-white, with a good deal of

yellow towards the sides. Limbs mottled and striated.”

Largest male 8 inches (64140 mm.), largest female 77 inches

(60-140 mim).

ICHNOTROPIS CAPENSIS (Gray).

Bler. Cat. Liz. 11. 1887, p. 84.

One specimen taken at Delagoa Bay, 24.xi1.14. They were

not uncommon, darting about the reddish sandy soil among the

thorn-bush.

Eremias spext (Ginther).

Bler. Cat. Liz. ii. 1887, p. 84.

Forty specimens were collected at Mt. Siswa (19. vii. 15),

Mt. Margaret (21. vii. 15), Voi (17. vi. 15), Duthumi (20. ix. 16),

and Morogoro (v. d.).

The coloration of this species was as follows :—-‘‘ Ground-colour

pale brown. ‘Two cream lines starting from parietals converge

to form a single dorsal line just behind an imaginary line uniting

the fore-limbs where they join the trunk. This line is Jost on the

tail. Single dorso-lateral line commences at posterior border of

eyeand merges into the white underside of tail. Single pure white

lateral line originating in the sub-ocular and upper labial region

EAST AFRICAN LIZARDCS. 149

passes through ear and ends at hind-leg ; in some specimens this

line is broken up into short white dashes. Black bars and dashes

unite this parallel series of lines, and are also seen to a lesser

extent on the anterior part of tail. Sides marked with pale

green blotches. Upper and lower labials and under surface of

body are china-white.”

On December 13th, 1917, four eges were found in a female.

A Gerrhosaurus major lizard in captivity was seen to seize and

eat one of these Hremias, and while it was so occupied, a Gerrho-

saurus flavigularis in the same case came up and tried to take it

away. Specimens of this lizard were also found in the stomach

of a Harrier (Cirews macrurus), Morogoro, 28.11.18, and in a

Kestrel (Cerchneis tinnunculus), Morogoro, 4. xii. 17. In both in-

stances there were also lizards of other families in the stomach.

HOLASPIS GUENTHERI (Gray) 2

Blgr. Cat. Liz. 111. 1887, p. 118.

During an action at Matombo, 3.ix.16, I saw what I believe

to be this hzard coming down the trunk of a large tree. It was

a handsome creature with a blue tail and good deal of blue about

the back. On breaking camp two days later I again passed this

tree, and left the road to inspect. As I rode upI caught sight of

the lizard running up the trunk. As no specimen was collected

I cannot be sure of the species, however.

GERRHOSAURIDS.

GERRHOSAURUS MAgoR (A. Dum.).

BlersCaty lize, LSS oso. (21

Seven specimens inall were collected; their variations can best

be shown in the following table:

Index 7 : Measurements : ‘ é bei

letter, ©X- Locality. “T° ¢ B., Tail. Colour, Scale characters.

(a)... M. Lumbo. 240-315 Fulvous brown. Fronto-nasal not in cou-

tact with rostrat

(0) ceo 295 3 240-235 3 3 in contact

with rostral.

(@) coo NE . 200-267 Dark brown, on not in con-

spotted yellow. tact with rostra:.

(je Ee A 200-260 a r ees

Gane KF, ” 142-172 29 tb) oH) by)

F Dodoma. 210-217 2 cE) 7) 23

(9) «..

Tt will thus be seen that all the specimens with the exception

of “b” and “c” disagree with one of the specific characters of

G. major in that the fronto-nasal scale is not in contact with the

rostral. The Lumbo specimens were taken at almost the same

150 MR. A, LOVERIDGE ON

spot, and are obviously the same species. All the specimens agree

in having 10 longitudinal ventral scale-rows, 17-19 longitudinal

dorsal seale-rows, and 34 transverse dorsal scale-rows.

The colour and scale-character key given by Schmidt * breaks

down for the specimens (d) to (y), which, according to the key,

would fall under Gerrhosaurus grandis (Blgr.) of Zululand,

Both the Lumbo specimens were taken from holes in termite

heaps; m one instance two mungoose (Herpestes ivori) were

occupying the same burrow, all three creatures being found

huddled together at the end of the hole. Nearly all the Moro-

goro specimens were taken among the rocks bordering the river

at the south side of the town. The Dodoma specimen was taken

on a rocky kopje in desert country.

The examination of the stomach of one of the Lumbo specimens

revealed small beans and grass with a single leg from a_ beetle.

As already mentioned, one captive specimen seized and ate a

lizard (Hremias spekii). From one of the Lumbo specimens five

tapeworms were removed, each about a foot long.

GERRHOSAURUS NIGROLINEATUS (Hallow).

Bler. Cat. Liz. iii. 1887, p. 122.

Specimens possibly belonging to this species, and certainly

referable to it according to the key and descriptions in. the

‘Catalogue of Lizards,’ have been placed under G. flavigularis

(Gray) for veasons stated below.

GERRHOSAURUS FLAVIGULARIS (Gray).

Blgr. Cat. Liz. iii. 1887, p. 122.

A large series of Gerrhosaurus of this group were collected at

Nairobi, Moschi, Morogoro, Mkuyuni, Dar-es-Salaam, and Lumbo.

Also seen at Gonya, Handeni, and Kerogwe.

In his admirable paper on the ‘‘ Herpetology of the Belgian

Congo,” Schmidt? suggests that all South and East African

species of this group should be known as flavigularis flavigularis ;

whilst he proposes to retain the name of nigrolineatus for the

West African ferms which have a preponderance of nigrolineatus

characters, these to be known as flavigularis nigrolineatus. The

key which he applies for the distinguishing of the two races is

as follows :—

CC. Tympanic shield narrow; a dorso-lateral stripe ; dorsals in

D. Dorsal scales in a transverse row 20-26, mode 22 (South and

TBhaeie UNGAR) casopassnadoces oppmac sg oon ioder save soaccacweses | MOS IDs sana MUEH IS:

DD. Dorsal scales 24-28, mode 26 (Angola, Lower Congo).

subsp. nigrolineatus.

In our series of thirty-nine specimens only three have 24 scale-

* Schmidt, Bull. Am. Mus. Nat. Hist. xxxix. 1919, p. 519.

AA be

EAST AFRICAN LIZARDS. 151

rows, twenty-six specimens have 22 scale-rows, and ten have 20

scale-rows.

This character of flavigularis seems, therefore, a good one by

which to differentiate Hast African forms. In twenty-nine

specimens examined by Nieden he found only three specimens

with more than 24 dorsal scale-rows.

If we apply the relative position of the head-scales to the

present series thus,

Preefrontals in contact forming a long suture ....... icsseeee. Nigrolineatus,

Przfrontals separated, fronto- nasal touching frontal cee flavigularis,

we find no less than twenty-nine of the ee would be

referable to nigrolineatus, only five to flavigularis, whilst five are

intermediate in that the prefrontals are only barely in contact.

As has just been pointed out in the case of Gerrhosaurus major,

the relative position of these head-scales as a specific character is

of but little use in this genus.

If the character of the number of femoral pores be applied,

Hemoraliporese) Axor miOne es a seeee eee. ce: senor eeeeeenee see eee ee nILeO RO LUILEILALS

isiterrnvonen| jayoueelss 1S} OP MAIS 35 aco ong snaceodea sen onnsne sanceodsass sos0ss00;s00 jen BCeICHGTROR,

we shall again find that the large majority of the lizards are

referable to nigrolineatus. Not having had the opportunity of

going into the matter more thoroughly or examining Central and

West African series of these lizards, I do not like to express an

opinion, but where both nigrolineatus and flavigularis ave found

overlapping in ‘so many localities it seems a pity not to unite

them as a single species.

The largest: male taken measured 183 inches (156°311 mm.,),

the largest female 172 inches (137°305 mm.).

The coloration of the young specimens was generally more

vivid than in the adults. The following colour notes were made

on capturing a young speciman at Lumbo :—- “ Centre of back

occupied by broad chocolate-brown band, bordered on either side

by a black line one scale in width, on the outer side of which

again. is a sea-green or yellowish line. Both lines disappear

about half-way along tail. Sides vertically streaked with black

and sea-green, usually two of the former to one of the latter.

Belly white with er eamy tinge. Fore-legs brown, mottled black

and sea-green, hind-limbs spotted with pale yellow. Tail brown

with indistinet markings except on the basal portion where the

markings of the back persist.” Adult males show a great deal of

red or pink laterally on body and tail ; this disappears usually in

preserved specimens.

At Morogoro (4.1.17) a female was taken with four white

egos in ovary; these measured % inch long. A young male

83 inches in length was taken at the same place (19.1.17). Of

this specimen a note was made:—‘“ No gaudy markings; the

yellow lines are pale almost to whiteness and the black marks are

unnoticeable except by close examination.”

‘

152 MR. A. LOVERIDGE ON

Grasshoppers were the chief article of diet ; there was the one

interesting attempt to take a lizard (Hremias spekii) out of the

mouth of its larger relative (G@. major) already noted.

At Morogoro (7.1.17) an 18-inch specimen was eaten by a

Sand-Snake (Psammophis sibilans). At the same place (28.1. 18)

a young one was taken from the stomach of a Harrier (Circus

macrurus). At Dar-es-Salaam (24. vi. 18) a Kingfisher (Halcyon

orientalis) was shot, in whose stomach was found a Gerrhosaurus

measuring 91 mm. in length. !

ScINCIDA.

MABUIA MACULILABRIS (Gray).

Blgr. Cat. Liz. 111. 1887, p. 164.

Only four specimens of this lizard were taken. One from

Mombasa, two from Morogoro, and one from Duthumi; the

latter measured 9% inches (73°178 mm.), whilst the larger Moro-

goro specimen measured 94 inches (89°152 mm.).

MABUIA BREVICOLLIS (Wiegm.).

Blgr. Cat. Liz. ii. 1887, p. 169.

Six specimens taken at Kagiado (28. x11. 15) and Longido West

(1.16). Of these, the Longido specimens were all young ones,

which 1 caught one by one as they emerged from their refuge in

a termite heap. They measured 44 plus 41 mm., 43 plus 43 mm.,

43 plus 40 mm., 43 plus tail (injured). In these young specimens

the markings are very distinct and the side stripes are continued

across the back, uniting with their fellows on the opposite flank.

In a half-grown specimen from Kagiado measuring 65 plus 71 mm.

there is an interruption by an unmarked scale-row along the

dorsal median line. In the adult from the same locality the

markings have receded to the flanks, where they are just distin-

guishable. This specimen measures 8% inches (130°83 mm., tip

missing), and is an almost uniform dark brown. Both the

Kagiado specimens were taken on the same termite heap from

whose openings they emerged. The adult is infested with

acarines on the ventral scales.

MABUIA MEGALURA ( Ptrs.).

Blgr. Cat. Liz. iii. 1887, p. 195.

A large number were collected at Nairobi, one at Longido

West, and sixteen at Lumbo. Of the latter. the largest male

measured 8 inches (55:145 mm.) and the largest female just over

112 inches (65°235 mm.). Colour during life as follows :—

“‘ Above pale bronze. <A pair of black zig-zagging lines (caused

by borders of scales being black-edged) commence in scapular

region and end in pelvic region, though faintly persisting on

upper surface of tail in the form of a double row of black specks.

Upper labials white; the white persists in the shape of a narrow

EAST AFRICAN LIZARDS. 153

band along the side; this band is only one scale wide and vanishes

on the tail. Throat and under surfaces white.” One Nairobi 2

specimen has longitudinal striations on whole of under surface.

A female taken on September 20th, 1918, at Lumbo held four

eggs 8xX6mm. Another female taken at Parklands, Nairobi, on

22. vii. 15 had four eggs 8 or 9 by 6, irreguiar in shape and

containing small embryos.

It seems to me that the Lumbo specimens have mueh longer

tails than the B.H.A. and G.E.A. specimens. The habit of the

latter is much stouter both in body and thickness of tail. Their

colour is much darker also. Unfortunately the series from

Nairobi is not available, with the exception of twelve specimens ;

the largest of these measures 80 mm. in head and body and

122 mm. in tail, which is intact. Another 2 55+165mm., ¢ 55

+163, 2 55+162.

The Lumbo specimens were found rushing about on sandy

ground, which is very sparsely grown with clumps of grass.

Nairobi specimens on the other hand were coilected in grass up

to one’s knees, either on the plains or near the Parklands

Forest edge. They clambered with great agility about the tops

' of the grasses. Possibly the different kind of country where

they were obtained explains the differences in appearances and

the relatively shorter tail length of the grassy upland forms.

Spiders, a little grass (probably accidentally introduced), and

parasitic worms were found in the stomach of a male at Lumbo.

The worms have been identified as immature Physaloptera.

MABUIA QUINQUETHNIATA (Licht.).

Blgr. Cat. Liz. 111. 1887, p. 198.

Two males only obtained at Dodoma, 8. xii.18. The larger

measured 82 inches (927128 mm.) and the smaller just under

84 inches (907113 mm.). Colour during life :—‘‘ Pale bluish, with

faint coppery traces of lines on the back. Neck striated with

bright yellow and blackish lines. Preserved specimens do not

show these so well.” Both specimens were living beneath boulders

on rocky kopjes.

MaBvia varia (Peters).

Blgr. Cat. Liz. ii. 1887, p. 202.

Twenty-seven specimens collected at Nairobi, Longido West,

Kongwa, Morogoro, Duthumi, Lumbo, and Delagoa Bay. Was

also seen at Mbunyi and Kahe. The largest specimens came

from Nairobi, though Lumbo specimens had longer tails than

either B.H.A. or G.H.A. specimens.

Largest Nairobi. Male 6% inches (65°90 mm.).

a a Hemales62 seu corgon mms):

seplmmibo. » Male, \ 67 .se5.84(05).02) mm...)

Bi as Female 62 ,, (63°103 mm.).

154 MR. A. LOVERIDGE ON

A constant feature in the coloration of the Lumbo specimens

was the black specklings on the lower labials and chin region

which was only found, and then but slightly, in two specimens

(Morogoro) from other localities. The two specimens reterred to

were taken on the Uluguru Mts., 3000 ft. above Morogoro; they

differed from the specimens collected elsewhere in the very dark

ventral surface which is usually white.

The colouring of Lumbo specimens noted at the time was as

follows :—‘ Above of a pale or dark bronze, back variegated with

black spots, sometimes tipped with white. In one male three

somewhat indefinite white dorsal lines. White stripe commences

at nostril, unites with white upper labials at end of mouth, passes

beneath eye and through ear, and continues along side, where it is

particularly clear and distinct, as far as hind-leg, where it ends.

Distinct reddish tint in lumbar region and base of tail. The

whole under surface of body and tail pure white. Throat white,

freckled at chin and lower jaw region with jet black spots.”

The following notes were made on present females :—

Palla tnen ILS Kongwa. 7 eggs containing minute embryos,

eges 8X6 mm.

iby savateyllicy: Nairobi. 4 eggs containing large embryos, eggs

12x8mm. Embryos 15+10mm.

Hl ivan 8: Lumbo. 5 eges measuring 9 X 6 mm.

P s Bick ae * 10x ?mm.,

ih Tee i 10x10 mm, con-

tained large embryos.

99 3)

Masur striara (Peters).

Blgr. Cat. Liz. 11. 1887, p. 202.

Twenty-six specimens were collected at Nairobi, Morogoro,

Duthumi, Dar-es Salaam, and Lumbo. This lizard was seen at

almost every place through which we passed, as it is so common

and unmistakable a large series was not collected. It adapts

itself to any situation, arboreal or terrestrial. I have seen a

pregnant female basking on a palm leaf; the Dar-es-Salaam

specimen was obtained running down a palm-tree trunk. The

grass-huts of natives are a favourite spot, but it adapts itself to

the stone-built dwelling of the white man, emerging from holes

in the masonry to bask on window-sills or on the galvanised-iron

roof. In one house in which the writer lived they used to make

a great racket in the roof-guttering, where they could chase each

other. On putting on my slippers on one occasion, I found one

in the toe, where it had evidently retired for the night, as it was

evening. Rockeries and rubbish-tips afford good refuges, and

they fall easy victims when found inside old tins in the latter

situation, which is often the case,

The largest male, 8? inches (97°125 mm.), was taken at Lumbo,

also the largest female, 93 inches (108130 mm.). The colour varies

a good deal. At Handeni I observed some large specimens basking

EAST AFRICAN LIZARDS. 155

on the ruins of a hut; they were unusually spotted between the

dorsal lines. I marked one down and caught it beneath a beam:

to my surprise it had no spots. It was just possible the speci-

men [ was originally pursuing had got away and I had captured

another; on the other hand, the blonches may be a sign of health

and vigour, and disappear when the lizard ig frightened. A

female was taken at Morogoro with a salmon-coloured throat ;

it was also much spotted dorsally. At Nairobi (13. vii. 19), the

weather being very cold, two Skinks were seen upon the wall, the

one having chased the other up; both fell off the wall and, being

sluggish, were easily captured. One was a male (85°109 mm.) and

had the throat mottled brown; the other, whose sex was not

determined, had the throat mottled orange.

The following notes were made on pregnant females :—

28. viii. 15. Nairobi. Ovules small.

AOpe5 Ie s 7 large young, about 24 inches

in length.

24.x11.14. Delagoa Bay. 4 young, 1} to 13 inches, and 1

bad egg.

16.xu1.16. Morogoro. Ovules nell.

At Morogoro (23. xii.16) a striped Skink was seen running

along with a nauseous gaudy grasshopper in its mouth. Sitting

after sunset one day (27.1.17.) by the river-bank, a Skink came

out of a hole just beside me, and snapped the head off a millipede

which was also out for an evening stroll. Grasshoppers, a beetle,

and a Jand-snail’s shell were found in the stomach of a Nairobi

specimen (5. ix. 19).

On three occasions specimens of this Skink were found in the

stomach of hawks at Morogoro :—(1.) Kestrel (Cerchneis tinnun-

culus), 4.1.18; (i1.) Harrier (Buter b. rufiventris), 31.1.18 3 (i1i.)

Circus macrurus, 28. 11.18.

Some cats are very fond of harassing these Skinks. At Nairobi

(5.1x.19) a cat brought one into the house; twice she released

and recaught it, yet the lizard did not drop its tail. I held the

cat up by the tail until she released the Skink, and, though she

had chewed it, when dropped, it ran away and escaped into the

garden. Somehow she managed to again find it, and brought it

into the house in a dying state, its tail still intact.

Nematode worms (Oochoristica sp.) were found in the stomach

of the Dar-es-Salaam specimen in large numbers.

LyGosoMA SUNDEVALLII (Smith).

Very large numbers of this snake-like Skink were collected.

For the present paper fifteen specimens from Nairobi, Negari

Mtoni, Moschi, and Morogoro were available, and a series of over

fifty specimens collected at Lumbo during July. 1918.

A manure-heap is one of the best collecting-grounds for this

156 MR. A. LOVERIDGE ON

Skink; some of the finest specimens were found among the stones

of a camp incinerator which was being demolished ; beneath

stones, especially where there is an ant or termite nest, and in

accumulations of garden rubbish are other spots favoured by the

species.

The tail of this Skink is so often reproduced that it is difficult

to obtain full-grown specimens with the original tail. Some

record lengths were, however, obtained :—

Largest Morogoro. Male P (140°? mm.). Tail reproduced.

¥ a Female 2 (132°? mm.). a3 .

na 7 8 inches (105100 mm.). Tail intact.

» Lumbo. Male 7 > (97°80 mm.). 5, probably reproduced.

% 55 5 7t ,, ( 95°90 mm.). PING te

A an Female 74 ,, (10684 mm.). », probably reproduced.

Average of 50 Lumbo specimens 52 inches (80°61 mm.).

Tails intact and reproduced. The 50 Lumbo specimens consisted

of 23 males and 27 females, so that the proportion of the sexes

is fairly equal; all were collected in an area under 300 square

yards.

The variability of the coloration of this species almost bafiles

description. The tendency in Morogoro specimens was to be

heavily spotted with black on a ground-colour of purplish brown,

a very handsome form. The Lumbo specimens inhabiting a

sandy soil were more often of a light brown colour. Notes made

at the time read: ‘“ Very variable. Usually plumbeous above and

dirty white below. Underside of tail as often mottled as not.

Some specimens mottled all over upper surface with black and

white; black spots oblong, white spots round or oval. Others

again are plain above, but mottled on the sides and tail.” The

uniform dirty white of the under surface is noticeable in these

Lumbo specimens when compared with those obtained at Moro-

goro, which are generally mottled on the under surface of head

and body as well as tail.

Three females were found with developing eggs :—(i.) 2 eggs

7x6 mm; (ii.) 4 eggs 12xX8 mm.; (ill.) 4 eggs 15x8 mm.

All Lumbo (three very young specimens taken) :—(i.) Morogoro,

1.11.18, 76 mm.; (ii.) Lumbo, vii. 18, 85 mm.; (i1.) Morogoro,

Ib mn Ike, DIL sean

Freshly emerged bluebottles were more often found in their

stomachs than any other food; specimens from the incinerator

afore-mentioned were particularly well fed on these. Bluebottle

pupee were also found. Pupz and ordinary flies (Muscide),

beetle, lizard’s tail, lizard’s scales. The two specimens found

with lizard remains in their stomach had probably been feeding

on Ablepharus wahlhergi, which was abundant in the same patch

of ground.

Natives always kill these inoffensive creatures, supposing from

their snake-like aspect that they are poisonous.

EAST AFRICAN LIZARDS. We7r

LycGosoMA FERRANDII (Bler.).

Blgr. Ann, Mus. Gen. (2) xviii. 1898, p. 718.

Seven specimens in all; six of these from Longido West have

been examined by Mr. Boulenger and referred to this species.

The measurements are as Rolla cs: though almost all the speci-

mens have reproduced tails :—

Dodoma. Female 91:36 mm. Longido. Male 56:46 mm.

Longido. Male 90:40 mm. as oO) mm.

Re $ 62°62 mm, A Bag ASS miagine

_ oe 59°59 mm,

The 118 mm. specimen was caught under rather unusual

conditions. After a night of heavy rain I was summoned at

6 A.M. to the tent of one of the sergeants to see a small snake

(Lycophidium jacksoni) labouring to swallow this Skink. It was

the only snake of this species captured. Another man told me

that he had killed a snake with tiny legs in his blankets; on the

battered remains being produced it proved to be this species.

The other specimens were all obtained under stones. These

Longido specimens are of a uniform nut-brown colour above,

creamy or yellowish beneath.

ABLEPHARUS BOUTONI Var. PERONII (Coct.).

Blgr. Cat. Liz. 111. 1887, p. 347.

Seventy-one specimens collected at Mombasa, Dar-es-Salaam,

and Lumbo. This little lizard has adopted a marine life; it is a

remarkable sight to see it running over the rocks, which a

moment before were washed by waves; for company it has the

peculiar fish Periophthalmus and crabs of many species. It flies

before the incoming wave and presumably manages to avoid a

wetting. It seeks refuge when pursued in the many crevices of

the rock. It is extremely agile and difficult to capture.

Measurements of the largest specimens from each locality :—

Mombasa. Male 3 inches (42°35 mm.). Tail regenerated.

es Female 43 ,, (43°60 mm.).

Dar-es-Salaam. Male 43 ,, (45°70 mm.).

i Female 43 4, (47°65 mm.).

Lumbo. Male 42 ,, (4862mm.). Tail regenerated.

Female 3¢ = ,, (50°46 mm.).

Coloration very variable. Some Lumbo specimens, both young

and old, almost black, but underlying markings usually dis-

tinguishable on close examination. Greenish or olive- bronze ;

two light lateral (almost dorsal) stripes start at nostril. Upper

surface of tail marked with white dots in lines; these were absent

on regenerated tails. The Dar-es-Salaam specimens were much

more brown or copper y> and with the markings more distinct

than in the Lumbo specimens.

Five females collected at Dar-es-Salaam (11.iv.18) had each

dr 99 be)

158 MR. A. LOVERIDGE ON

two eggs in ovary; four of these batches were in an early stage of

development and almost round, measuring 5 mm., 5 mm., 9 mm.,

and 11 mm. in diameter; in the fifth were developing embryos,

which measured 11 x 6 mm., and were oval in shape.

Sea-slaters 10 mm. in length were found in the stomach of a

102-mm. specimen ; sandhoppers 9 mm. in length in the stomach

of a 107-mm. specimen. It seemed extraordinary that the lizards

could swallow such large prey. Flies were found in the stomachs

of a good many. A

Tapeworm measuring 32 min. in length was found along with

the sea-slaters in the stomach of the 102-mm. specimen just

referred to.

ABLEPHARUS WAHLBERGII (Smith).

Bler. Cat. Liz. ui. 1887, p. 350.

Eighty specimens were collected at Nairobi, Longido West,

Morogoro, Mkuyuni, and Lumbo. The largest male was from

the last locality, measuring 4 inches (40°59 mm.), though another

specimen with renewed tail measured 2 mm. longer in the body.

Largest female was from Mkuyuni and measured 42 inches

(43°67 mm.).

The coloration of Lumbo specimens was as follows :—‘ Pale

copper above, reddish tinge on tail. Dark copper band commen-

cing at nostril, passes through eye, above fore-legs, and along side

to hind-legs, where it disappears. A few irregular white lines in

region of ear originating in white upper labials. Belly and under-

side of tail a transparent brick-red.” This reddish appearance

of tail was only seen in specimens collected after the middle of

September; it may be a sign of the breeding-season or again

of locality, for most of the specimens collected between July and

September were collected at a different spot. The Nairobi and

Morogoro specimens were much darker in colour, bronze rather

than copper.

The species frequents grassy places, particularly where there is

much garbage or fallen leaves. They apparently prefer moist

spots beneath trees ; large numbers were discovered in uprooting

stumps and clearing the ground for making camps.

Two females collected in Oetober 1918 at Lumbo had each

two eggs measuring 6 X 2 mm. in ovaries. ‘Two females collected

at Mkuyuni (1.ix.16) and Morogoro (1.11. 18) had each six eggs

measuring 7 x 4 mm. and 9x 5 mm. respectively.

Principal food was white ants; a fly, beetle larva, and field

cockroach were also taken from the stomachs of specimens.

A specimen caught at Handeni (27.vi.16, lost in transit) by

the neck whirled its tail round very rapidly, and then making

it suddenly rigid, caused it to break off; the tail exhibited con-

siderable vitality, wriggling and jumping about for some time.

One of these lizards was found in the stomach of an Egret

(Bubuleulus ibis) at Morogoro (14. x11.17); as already mentioned,

EAST AFRICAN LIZARDS. 159

seales and a tail fragment, probably belonging to this species,

were taken from the stomachs of two Skinks (Lygosoma sunde-

valli). Three were found in the stomach of a snake (Chlorophis

neglectus), Nairobi (17. vii. 18).

SCELOTES EGGELI (Tornier).

Torn. Zool. Anz. xxv. 1902, p. 700.

Eight specimens of this Skink which was described by Tornier

from Usainbara, G.H.A.,were taken at Lumbo in July and August

1918. The coloration during life as noted at the time was as

follows :—‘‘ Copper-coloured above, becoming plumbeous on tail ;

dirty bluish white below; the two separated by a very dark

brown or blackish lateral band, well defined above but merging

into the bluish white below. Throat sometimes spotted.”

Largest male measured 332 inches (63°32 mm., tail short and

regenerated), largest female 44 inches (67°38 mm., also regene-

rated). Smaller specimens with uninjured tails show that the

tail should equal the length of head and body.

In ovaries of two females taken at Lumbo in July 1918 were

each two eggs measuring 7 X 5 mm.’

Text-fioure 1.

Scolecoseps boulengeri.

. SCOLECOSEPS, gen. nov.

Among the lizards collected at Lumbo were seven specimens

of a limbless burrowing Skink referable to no known genus.

Generic description. Characters as in Melanoseps, but nostril

pierced in the very large rostral, with whose posterior border it is

connected by a horizontal cleft, as in Acontias. ;

Locality. South East Africa.

SCOLECOSEPS BOULENGERI, sp.n. (Text-fig. 1.)

Specific diagnosis. Snout conical, strongly projecting, length

of the rostral a little more than one-fifth that of the head.

Internasals in contact, sometimes very narrowly, separating

rostral from fronto-nasal, which is twice as broad as long. Frontal

equal to or but little larger than fronto-nasal. Interparietal

160 MR. A, LOVERIDGE ON

sub-cordiform, notch towards snout, larger than any other head-

shields. Parietals narrow, band-like, narrowly in contact behind

apex of interparietal. Two supra-oculars, no supra-ciliaries.

Five or six upper labials, first largest, third entering the orbit.

Hye distinguishable.

Mental very large, its posterior border corresponding with the

suture between rostral and first upper labial,

Scales hexagonal, broader than long, in eighteen longitudinal

rows at mid-body, in thirteen rows at base of tail. Anal divided.

Tail less than half the length of head and body, ending in obtuse

point.

Body flesh-coloured, with eighteen longitudinal brown stria-

tions corresponding with the scale-rows. Snout paler above and

beneath, more or less free from mottlings. Tail darker by

reason of convergence of thirteen striations, in some specimens

almost blue-black. Reproduced tail-tips flesh-coloured or white

and extraordinarily like snout.

Type locality. Site of British camp at Lumbo, which is situated

on the mainland three miles from Mosambique Island, P.H.A.

Measurements of specimens collected :—

(a) Male. 90°16 mm. ‘Type in British Museum.

(6) Female. 65°25 mm. Type in British Museum,

(d) Female. 95°-41mm. 4 eggs in ovary, 11. vu. 18.

(e) ry 95:25 mm.

(Ff) 39 92-43 mm.

(9) ees 61-24 mm.

ANELYTROPIDA.

FEYLINIA CURRORI (Gray).

Bler. Cat. Liz. 11. 1887, p. 431.

A single specimen of this aberrant Skink was found in a bottle

in a German house at Morogoro. It measured 4 inches

(92:10 mm.).

CHAM ZLEONTIDA.

CHAMALEON GRACILIS (Hallow).

Bler. Cat. Liz. 11. 1867, p. 448.

A single male from Longido West (22.11. 16) measures 7? inches

(100-96 mm.). Colour during life:—“ Pale green, with dark green

saddle-like markings bordered by black spots.” Jmmediately after

death “pale green became dark, dark green became pale, black

spots turned orange.” The creature was blind in one eye when

found, having suffered some accident to the eye.

CHAMLEON DILEPIS (Leach).

Bler. Cat, Liz. 111. 1887, p. 450.

Collected at Voi (6.v.16) and Mbunyi (10. v.16) in B.E.A,,

EAST AFRICAN LIZARDS. 161

at Gonya (29.v.17) and Morogoro (1916-1918) in G.H.A., and

Lumbo (1918) in P.H.A. Specimens which I believe to belong

to this species were taken at Tsame, Handeni, and Msiha in G.H.A..,

but were lost in transit. Over 90 specimens were collected in

all. Many of the Morogoro specimens might be referable to

Ch. dilepis isabellinus (Giinther), but as there is a good deal of

intergradation, and sub-specificity of isabellinws is somewhat

doubtful, I refer them all to Ch. dilepis.

The following are some of the measurements :—

Female. Voi. Snout to tail 73 in.(142:115 mm.).

Female. Mbunyi. pe 9F5 ,, (130°100 mm.).

Male. Gonya. %, 65 ., (90°80 mm.},

Male. Morogoro, r 133 ,, (largest of 23 speci-

mens).

Male. 55 © ll ,, (138°144 mm., average

of 23 specimens).

Female. A 3s 143 ,, (largest of 26 speci-

mens).

Female. a: i 123 ,, (158-161 mm., average

of 26 specimens).

Both sexes. ye ‘3 111 ,, (148°153 mm., average

of 50 specimens).

Female. Lumbo. . 113 ,, (145-140 mm., average

of 13 specimens).

At Morogoro the sexes were very evenly balanced, but at Lumbo

no males were taken at all; thirteen females were taken between

July and October.

The number of eggs produced at a time is enormous. The Voi

specimen (6.v.16) contained 44; the Mbunyi female (10. v. 16)

28; at Morogoro between end of February and early in March

the largest number of eggs found were 48, 44, 43, and 40

respectively. These eges when nearly ready for laying are

almost spherical, and measure from 9-10 mm. in diameter. On

January Ist, 1917, after very heavy rains, the sun shone out

brightly. I took two very young chameleons on shrubby growths

not more than a foot from the ground and quite half-a- mile away

from each other. These young have a ridiculous appearance, the

head being out of all proportion to the body ; the occipital lobes

are scarcely developed ; their coloration was much brighter than

in the adult. Both measured less than 3 inches; other very

young ones were taken in February and March. Oviposition

takes place between March and June apparently, which coincides

with the rains; no enlarged ovules were found after April at

Morogoro, or in the thirteen Lumbo females which were collected

between July and October.

The chief diet of the species is grasshoppers; I have also seen a

captive specimen take a large black field-cricket. The following

have been found in their stomachs :—Cockroaches, a praying

Proc. Zoou. Soc.—1920, No. XI. a

162 MR. A. LOVERIDGE ON

mantis, flies, bluebottles, rose-beetles, a fairly large scarab, remains

of many species of smaller beetles, and a millipede. Other finds

which can scarcely be classed as articles of diet include portions

of the chameleon’s own cast skin, half a nutshell which was pro-

bably too bulky to pass out, and in a specimen which was found

dying there were four fragments of mica—three pieces measured

8x9 mm.and one 8x11 mm.; doubtless all had split off from

one piece which had adhered to the chameleon’s tongue when it

was feeding. During a shortage of insects I fed strips of Bulbul

flesh toa chameleon ; it took them readily enough, but afterwards

disgorged them again.

I rarely found these chameleons on bushes, but usually met

with them crossing the road ; several of them were taken ascending

the trunks of trees. Their movements are sedate, and each step

appears to be well meditated before being taken. They sway

gently from side to side when walking. When molested or

picked up, it assumes the policy of frightfulness: the occipital

lobes are raised, the mouth gapes widely to show the red interior,

the throat is dilated so that the orange-coloured interstitial skin

is seen between the black scale-rows, and a startlingly sudden

lunge forward is made—sometimes an actual bite if the creature

is sufficiently enraged. The teeth are blunt and conical, and only

once on the many occasions on which I have been bitten have I

known this species to draw blood.

The native has a holy horror of them, and even boys who will

capture and handle the most venomous snakes, cannot bring

themselves to touch an uncanny chameleon. “It spits at you,”

they say, “and cannot you see it has a bad eye?” This superstition

is shared by the uneducated Dutchman. JI recollect on. one

‘occasion, when the column halted for ten minutes one day, I picked

up a chameleon, which truly horrified a young Dutchman beside

me: he implored me to putit down. His argument was: “‘ They

are poisonous, because if you put some pipe-oil in their mouths

they will die. You can always tell a poisonous snake by this test,

for the harmless species are not affected by the oil.” He told me

he knew of a woman who died from a chameleon bite. I

put my little finger into the chameleon’s mouth, and let it chew

vigorously for a few seconds so that its teeth marks could be

plainly seen. He said it was evidently a young one, and was not

the least disturbed in his beliefs.

Their only enemies at Morogoro as far as I know were the

Boomslangs (Dispholidus typus). On four occasions I found or

heard of these snakes falling out of trees with a chameleon: the

chameleon is evidently an awkward mouthful. I fed chameleons

to captive Boomslangs and the snakes took them, but not before

there was a contest between them; the chameleons went through

the frightfulness tactics already mentioned, which caused the

snakes to start back. A large brown Boomslang was shot at

Lumbo with a chameleon in its stomach.

EAST AFRICAN LIZARDS. 163

No parasites were found in the stomachs of any of the speci-

mens examined.

CHAMELEON DILEPIS ISABELLINUS (Giinth.) ?

Giinther, P. Z. 8. 1892, p. 556.

As already stated, many of the Morogoro specimens might be

referred to this subspecies ; so it is with considerable hesitation I

apply this name to a female from Gulwe (28.iv.17) measuring

101-101 mm. and a female from Dodoma (8. xii. 18) measuring

105°106 mm., which, while typically isabellinus, I believe might

find their counterpart in the long series of seventy specimens

collected at Morogoro.

CHAMELEON BITENIATUS (Fisch.).

Blgr. Cat. Liz. i. 1887, p. 452.

Hight specimens were collected at Longido West (ii. 16) and on

the Longido-Moscehi trek. Colour in life:—“ Light brown or khaki;

a light lateral stripe commencing at the eye disappears towards

the base of the tail; a lower stripe commencing on the upper

labials continues along the side to the hind-leg; two small

saddle-like markings on anterior part of back were of an ochre

tint, and there were three blotches of the same colour connecting

the upper and lower stripes.”

Six of the eight specimens were females, the largest measuring

64 inches (88°78 mm.), largest male 52 inches (75:68 mm.). Four

of the females contained eggs, from seven to tenin number. The

largest were 7 mm. in diameter. Probably they would be laid in

March. a

CHAMELEON H@HNELII (Steind.).

Steind. Sitz. Ak. Wien, 1891, p. 307.

There are several examples in the Nairobi Museum of this

chameleon which were collected by the writer at Kabete (20. ii. 15)

and West Kenia (23.xi.15). The largest male from the last-

named locality measures 85 inches (110-100 mm.) and the largest

female 52 inches (73°63 mm.). The size of the casque in these

‘Specimens varies a good deal.

CHAMZLEON JACKSONI VAUERESCECE (Tornier).

Torn. Zool. Jahrb. Syst. xix. 1903, p. 176.

This is the common Three-horned Chameleon of Nairobi; it

differs from Ch. jacksoni (Blgr.) in that the females are three-

horned like the males. A very large series was collected at

Nairobi in 1915 and 1919. Unfortunately these are not avail-

able for the present paper as they are in Nairobi, except four

specimens and some notes made on a few of the others.

164 MR. A. LOVERIDGE ON

Meek, in his paper on the “ Batrachians and Reptiles from

British East Africa,’ makes reference to this subspecies under

the name of Chameleon jacksoni (Blgr.). He refers fifteen

specimens collected in Lukenya Province to this species, yet

says: ‘The males are easily distinguished by having the rostral

and two occipital horns of about equal length. These cephalic

projections on the female are short; the occipital ones are usually

very short, in these specimens never more than half the length of

the rostral horn.” This is then followed by a table of measure-

ments in support of the theory. By occipital he means pre-

orbital horns.

There is nothing to support this in the series collected at. -

Nairobi, of which measurements were taken. Only the measure-

ments of eleven specimens are given below, as there is no doubt

about the sex of these specimens; measurements were taken of

twice this number, but not being available for sexing, they are

not taken into account :—

Sexdoek eee 3s M. Mey Mee iis as = BN BYR Rs ae

Length of head and

THOGKF sidcctucdeoodce MUS} HO). Bh 90 65 132 118 117 116 100 £75.

Length of rostral

1NVOHIN) by jnoe eoB BMG 00 Ly Wp a7 We ff NK I. Alby BR 1S 5

Length of pre-

orbital horn ... 18-17 18 18-17 17 7 Gg) Hs) Das BE 1s

Where two lengths are given for the pre-orbital horns it is.

because they are of unequal length, and the measurement of the

right horn is then given first. It will be seen that the greatest

variation is found in the respective horn lengths. In two of the

females the rostral and pree-orbital horns are of the same length,

as is the case in most males; in one specimen (3809) the preae-

orbital horns are much longer than the rostral, whilst in another

(4563) the pre-orbitals are like tiny thorns only 1-5 mm. long.

One sometimes finds specimens with injured horns apparently

in process of being regenerated. Natives are, I believe, often

responsible for this condition of affairs. They are very afraid of

these reptiles, and so, if a young man has courage enough to seize

one of the creatures and cut off its horns with a knife, it is counted

greatly to his credit. I believe this is done so that the horns.

may be presented to his lady-love to thread on her necklace as

proof of his devotion.

The largest specimen taken was a female 14/ inches long

(132:128mm.). This specimen had also the largest number of eggs,

no fewer than forty being found in the ovary; these measured

20x20 mm. (17. vii.19). Thirty-six were found in another

EAST AFRICAN LIZARDS. 165

specimen, whilst the lowest number found was eleven measuring

4x4 mm. (10. v. 15).

Most specimens were found during the early rains of March-

April 1915, when the following notes were made of the use of the

horns in fighting, ete. :—

March 31st, 1915; Nairobi.—I have half-a-dozen live chame-

leons on branches of Hucalyptus, which are tied to the cords of the

electric lights and about five feet from the ground.

Whilst sitting at the table writing, I hear d the sound of a moth

flapping its wings, and, glancing round, saw a large yellow under-

wing (Meenas fullonica 2), twice as large as the common English

Yellow Underwing, in one of the Euealyptus branches. Supposing

it to have got caught in a spider's web, I seized a killing-bottle ;

on reaching the spot, however, I found that it was caught by the

right-hind wing by one of the chameleons. Being large and

strong it flapped vigorously: the chameleon, biding its time,

eulped it down in the intervals between its str uggles until only

a small portion of the body of the moth remained to view.

Meanwhile, the second chameleon on that branch had been

eyeing its companion with evil intent. As soon as it perceived

that the captor was in difficulties, it hurried across the intervening

twigs, and pausing only to take aim, shot out its tongue and caught’

one of the now feebly flapping wings; having fot this into its

mouth, it commenced a tug-of-war jow] to jowl, and thus succeeded

am wrenching the whole moth from its companion’s jaws. The

poor moth’s “fur” was flying in the air, and the first chameleon’s

mouth was woolly with it—this was all the share of the spoils

that it was destined to receive.

There are three electric-light cords with branches tied to them,

and on each are two of these chameleons. More than two are

not tolerated; introduce a third, and the two original inhabitants

hurry towards it open-mouthed, striding along faster than on any

other occasion. As a preliminary to fighting, the opponents face

one another, swaying their bodies from side to side; and if the

intruder does not turn tail at this and cast himself to the ground,

as is generally the case, he is at once attacked. One of the

chameleons was very ingenious, placing his chin to the branch,

which was of course vertical, so that his rostral and pre-orbital

horns point directly downwards; he thus advanced upon the foe

as it were with fixed bayonets: by this means he generally

succeeded in sweeping his opponent off, but sometimes the latter

would seize one of the horns with its fore: foot, and a tussle would

begin.

They would bite each other hard. and in one instance the one

chameleon mounted the other’s back, and digging its claws in

continued the attack from this vantage point. So strong was its

grip that I had the greatest difficulty in separating the com-

batants. Several times have I seen one seize the other’s “arm” in

very human fashion, and then butt in with its armoured head.

166 MR. A. LOVERIDGE ON

This use of the horns interested me greatly, as I had hitherto

regarded the horns as an extravagant growth such as is common

in beetles.

One chameleon in throwing itself off the branch was injured,

and lay on its side; the hind-limbs and tail were paralysed.

When put on the branch, it dragged them after it very helplessly.

In one minute from the time of its fall the tail and hind-limbs

and posterior part of the body were almost white, the rest of the

body being dark green verging on black. As it did not recover

I chloroformed it three minutes later.

April 1st, 1915.—Another of the chameleons fell or threw

itself down and was killed; the posterior half of the body went

white immmediately, and the creature never stirred again.

Sometimes one will jump down six or more times in a single

morning, and so I conclude that these two accidents occurred

through the ehameleons falling on their backs. When given a

grasshopper one of the chameleons held the kicking fore-legs in

its “hands” ; another, which was given five ant- lions in succession,

made use of its “ hands” in very human fashion for pulling off

the gauzy wings which were flapping about its Jaws.

CHAM#LEON MELLERI (Gray).

Bler. Cat. Liz. iii. 1887, p. 472.

Two specimens of this giant chameleon were taken. A female

at Mkuyuni (31. viii. 17) measured 21 inches (273-261 mm.), and

a male at Morogoro (1. x. 18), tail mutilated, 238mm. Hxtruded

tongue measured 17 inches.

My attention was drawn to the female by the horse-guard, who

found it wandering in the grass; its colour then was yellow and

green, pale and dark shades of both. Placed under a pot it.

became dark green, and then almost black with wrath. Suspended

by the tail it would turn, and grasping its hind-legs with its fore-

legs, climb up its own tail. Its claws were sharp enough to make

my bare arm bleed as it crawled up to my shoulder.

The second specimen was brought by a boy who caught it in

the hills behind Morogoro; it led well enough in a large cage

until a Lemur was introduced into the same cage, which already

contained a 12-ft. Python. During the night the Lemur chewed

the chameleon’s tail to a rag; the second night I caught it doing

the same thing, so chloroformed the chameleon. I was surprised

that a creature like a Lemur could tackle so big a chameleon

whose bite must be pretty severe.

RHAMPHOLEON BREVICAUDATUS (Mats.).

Matschie, Sig. B. Ges. Naturf. 1892, p. 107.

A female was taken at Morogoro (28. xi.17) measuring

2+ inches (40°16 mm.) over all. It contained five eggs of oblong

EAST AFRICAN LIZARDS. 167

shape measuring 9x5 mm. In captivity it took bluebottles

readily.

RHAMPHOLEON KERSTENII (Peters).

Blgr. Cat. Liz. 111. 1887, p. 175.

Two females, the largest, Voi (19. vi. 16), measuring 33 inches

(62:28 mm.). ‘The other, collected at Gonya (29. v.16), measures

just over 3 inches (51:26 mm.). <A third specimen, probably of

this species, was caught crossing the road at Handeni, but lost in

transit.

The Gonya specimen was caught holding on to the edge of a

fragment of bark with its right side pressed to the bole of

astump. Every peculiar angle of its head and body seemed to

assist in the deception—the colouring was perfect. Two yards

away a second specimen was found crawling out of the stream,

into which it had evidently been tumbled by natives engaged in

clearing bush and undergrowth. It subsequently escaped.

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ABNORMAL FEATURES IN THE PERITONEUM OF A RACCOON. 169

12. On Abnormal Features in the Peritoneum of a Raccoon.

By C. F. Sonnrac, M.D., Ch.B., Anatomist to the

Society.

[Received February 6, 1920: Read February 10, 1920. }

(Text-figures 18 & 19.)

The conditions present in the peritoneum of this particular

Raccoon are extremely interesting. They consist of :—

1. Absence of the Foramen of Winslow. ;

2. The presence of a communication between the greater and

lesser sacs on the inner side of the upper half of the left kidney.

3. Subdivision of the lesser sac of peritoneum by a vertical

gastro-pancreatic sheet. ‘The gastric and hepatic branches of the

coeliac axis artery run to the liver and stomach in this.

4, Subdivision of the lesser sac by transverse peritoneal bands

running from the pancreas to the hilum of the spleen. Through

this the splenic vessels run.

5. The presence of bands connecting the great omentum to the

abdominal viscera.

6. The presence of a bare or non-peritoneal area in front of

the left kidney.

The Lesser Omentum (text-fig. 18, no. 4).

The lesser omentum ran from the inner surface of the lower

end of the cesophagus, the lesser curvature of the stomach, the

upper border of the first part of the duodenum, and the outer

border of the upper half of the second part of the duodenum to

the inferior surface of the liver and the lateral abdominal wall.

It was also connected below to the duodenal mesentery. To join

the latter it passed in a slanting direction from the outer to the

inner surface of the duodenum. ‘The line it follows is shown in

text-fig. 18, no. 26.

The attachment to the liver is along a line beginning on the

inferior surface of the cystic lobe to the left of the fundus of the

gall-bladder, and runs along the inferior surface of the bladder,

being divided to embrace it, the portal fissure, the ridge on the

inner surface of the inner division of the right lateral lobe, and

the inferior borders of both divisions of the right lobe.

The caudate lobe of the liver (text-fig. 18, no. 2) is seen shining

through the omentum, and the common bile-duct, portal vein

(text-fig. 18, no. 25), hepatic artery, and inferior vena cava (text-

fig. 18, no. 24) are included within its layers. At no point can a

Foramen of Winslow be detected in this region.

The lesser omentum appears, in this case, to be formed of two

layers, which appear Y-shaped on cross-section, and the common

bile-duct (text-fig. 18, no. 3) is placed at the junction of the three

limbs. The two lateral limbs embrace an angle open to the left

(text-fig. 18, no. 5), and the arrow no. A in text-fig. 18 points

170 DR. C. F. SONNTAG ON ABNORMAL FEATURES

into it. Perhaps there has been some inflammatory process at

work here, for the bands connecting the great omentum, and

described below, are doubtless pathological.

Text-figure 18.

ORwN

| ae

Th 1) /LAGe

Ee =

ines ee

SL Sh.

a i, — ah

linn Dy

The peritoneum viewed trom the front.

The great omentum has been divided along the lines L.S.O. of text-fig. 19, and the

edges of the four layers are shown above at 7. The left kidney has been

pulled outwards and the transverse meso-colon (18) and descending colon

pulled down. The following figures are not described in text. 1. Diaphragm.

19 and 27. Lleum.

IN THE PERITONEUM OF A RACCOON. ey hi

The gastric attachments of the lesser omentum and the gastro-

pancreatic band (text-fig. 18, no. 6, and text-fig. 9, G.P.B.) fuse

on the lesser curvature and first part of the duodenum. The

latter band draws down the lesser curve of the stomach, so the

plane passing through the two curves meets the vertical at an

angle of 45° (text-fig. 19).

The Great Omentwne.

The posterior layers of the great omentum (text-fig. 18, no. 10),

in passing to the transverse colon, cover the pancreas, and have

a sharp free left border, which runs with a coneavity to the left,

from the hilum of the spleen (text-fig. 18, no. 11) to the transverse

colon. It overlaps the upper pole of the left kidney, but is *7 cm.

from the hilum of the latter. It is met by a triangular sheet of

peritoneum (text-fig. 18, no. 13), the apex of which is attached

to the anterior surface of the lower half of the left kidney.

There are two pockets between the triangular sheet and the

great omentum, and the arrows C and D in text-fig. 18 point into

them. Another sheet of peritoneum (text-fig. 18, no. 12) passes

from the inner border of the left kidney to the poster ior parietal

peritoneum.

Between the free edge of the great omentum and the last-

named peritoneal sheet there is an opening through which the

index-finger can easily be passed into the lesser sac, and the arrow

B in text-fig. 18 points into it. It is bounded in front by the

great omentum, to the outer side by the left kidney, and behind

by the peritoneum of the posterior abdominal wall. It is

bounded above by a mass of tissue formed by the great omentum,

the gastro-splenic ligament, and the pancreatico-splenic band

(text-fig. 18, no. 8).

The triangular sheet of peritoneum (text-fig. 18, no. 13) also

meets the part of the great omentum forming the transverse

meso-colon on the left side of the latter, and the ileal mesentery

(text-fig. 18, no. 28) meets it on the right at the same level, so a

peritoneal cross is formed. There are two pockets between the

mesentery and the meso-colon, and the arrows E and F in text-

fig. 18 point into them.

The Peritoneal Connections of the Pancreas.

The right limb of the pancreas lies within the duodenal loop,

and contained between the two layers of its mesentery. The

upper border of the body of the pancreas is connected to the

lesser curvature of the stomach by a gastro-pancreatic sheet (text-

fig. 18, no. 6, and text-fig. 19, G.P.B.), which passes behind the

posterior surface of the stomach. This sheet is continuous with

the duodenal mesentery on the right and the pancreatico-splenic

band on the left. It subdivides the lesser sac into two compart-

ments, and pulls down the lesser curvature of the stomach.

The celiac axis artery (text-fig. 18, no. 9) passes into the

172 DR. CG. F. SONNTAG ON ABNORMAL FEATURES

Text-figure 19.

V.P.

Lateral view of the peritoneum.

G.O. Great omentum; G.P.B. Gastro-pancreatic sheet; G.S. Greater sac; L.S.A.

Anterior compartment of lesser sac; L.S.O. Lines across which pieces of peri-

toneum are removed (see text-fig. 18, no. 7); L.S.P. Posterior compartment

of lesser sac; P. Pancreas; P.P. Parietal peritoneum; P.S. Plane passing

through curvatures of stomach; S.J. Small intestine; S.O. Lesser omentum;

T.C. Transverse colon; T.M.C. Transverse meso-colon; V.P. Vertical plane.

Arrows :—

Al. Passing up through lesser sac from abnormal foramen (arrow B. in text-fig. 18) ;

A2. In posterior compartment of lesser sac; A3. In anterior compartment of

lesser sac.

The peritoneum between S.O. and G.P.B. is thickened and represents a fusion of S.0-

and G.P.B.

IN THE PERITONEUM OF A RACCOON. W7E

pancreas, where it divides into the gastric, hepatic, and splenic

branches, the gastric and splenic veins pass into the pancreas,

and the portal vein is formed at the point where the splenic vein

emerges from the pancreas.

The Lesser Sac of Peritonewm.

The lesser sac of peritoneum extends from the diaphragm

above to the apex of the great omentum below, and it is sub-

divided into two compartments—a small anterior and a large

posterior one by the gastro-pancreatic band (text-fig. 19, L.S.A.

and L.S.P.).

The anterior wall is formed from above down by the peri-

toneum covering the posterior surface of the liver, the gastro-

hepatic omentum, the peritoneum covering the posterior surface

of the stomach, and the posterior of the two anterior layers of the

large omentum. The posterior wall is formed from below,

upwards by the inner of the two posterior layers of the great

omentum, the transverse colon, the transverse meso-colon, and

the posterior parietal peritoneum from the pancreas up to the

diaphragm.

The gastro-panereatic band carries the peritoneum, covering

the pancreas, up to the lesser curvature of the stomach. Below

it lies the small anterior sub-space (text-fig. 19, L.S8.A.), and above

it the large posterior one (text-fig. 19, L.8.P.). When the finger

is carried into the sac it can be introduced into either compart-

ment, but to enter the posterior one it must go above or below

the pancreatico-splenic sheet.

The anterior compartment has the following boundaries :—

1. Above and in front—the posterior surface of the stomach.

2. Below and in front—the great omentum.

3. Above and behind—the gastro-pancreatic band.

4. Below and behind—the great omentum, transverse colon,

and transverse meso-colon.

The posterior compartment has the following boundaries :—-

1. Anterior—the liver, small omentum, stomach, and gastro-

pancreatic band.

2. Posterior—the parietal peritoneum.

Tn text-fig. 19 the arrow Al runs vertically through the lesser

sac, and arrows A2 and A3 point into the two subdivisions.

The Peritonewm round the Left Kidney.

The left kidney has a strong peritoneal capsule (text-fig. 18,

no. 29), which surrounds its outer part, and there is a non-

peritoneal bare area in front (text-fig. 18, no. 30), but the capsule

is strong behind. It is connected to the left Fallopian Tube

174 ABNORMAL FEATURES IN THE PERITONEUM OF A RACCOON.

({text-fig. 18, no. 15) by a peritoneal sheet (text-fig. 18, no. 16),

and to the posterior parietal peritoneum. On the surface of the

bare area (text-fig. 18, no. 30), the left ovarian vessels cross the

kidney (text-fig. 18, no. 31), and pierce the capsule on its outer

border to reach the ovary.

Omental Adhesions.

The great omentum filled the abdominal cavity, but it has been

shortened for descriptive purposes in text-fig. 18. When it was

gently raised, it was seen that delicate filiform bands connected

it to the abdominal viscera. These are indicated by Roman

numerals in text-fig. 18. Bands 1 and 2 were tight, but all the

others were loose.

The band number | ran from the front of the great omentum

across the front of the stomach to the anterior surface of the

small omentum. Bands 2 and 7 ran from the free edge of

the omentum to the inner surface of the liver and the outer

border of the spleen respectively. Bands 3 and 6 ran from the

under surface of the great omentum; the former ended in the

mesentery of the duodenum, and the latter on the left extremity

of the pancreas. Bands 4 and 5 gradually opened out on the

free edge of the great omentum; the former terminated in

the mesentery of the ileum, and the latter on a nodule on the

apex of the urinary bladder (text-fig. 18, no. 17). On each side

of the bladder-wall a tortuous ridge, containing an artery, ran

down from the afore-mentioned nodule.

These bands are probably the result of inflammation.

Duodenal Bands.

The outer surface of the duodenum is attached by bands to the

parietal peritoneum (text-fig. 18, no. 22), and the mesentery of

the descending colon (text-fig. 18, no. 20). Between the bands

and the duodenal mesentery is a pocket (text-fig. 18, no. 9).

ON ABNORMALITIES OF ABDOMINAL ARTERIES OF A PANDA. 175

13. On Abnormalities of the Abdominal Arteries of a young

Panda. By C. F. Sonnrac, M.D., Ch.B., Anatomist

to the Society.

[Received February 20, 1920: Read March 30, 1920.]

(Text-figure 20.) ©

The animal in which the peculiarities described below occurred

was born in the Society’s menagerie on June 6th, 1919, and died’

on December 12th, 1919. The abnormalities involved the cceliac,

superior mesenteric, and renal arteries. The first one was

unique, but the others are occasionally met with.

The Celiac Artery.

The coeliac artery was abnormal in its origin, course, termina-

tion, and some of its branches.

It arose from the left side of the abdominal aorta, instead of

from the front, just above the superior mesenteric artery. It

passed upwards and to the left, behind the stomach and above the

pancreas, towards the cesophagus, describing a wide curve with

its convexity to the left. On reaching the esophagus it passed

behind it and entered the left pleural cavity through the cso-

phageal opening in the diaphragm. Within the left pleural sac

it exhibited a free loose part measuring 1°3 em., and then it

passed into the left lung at its antero-inferior angle. Within the

lower lobe of the left lung, to which it is confined, it runs upwards

parallel to the pulmonary vein, 2. e., it described a course which

is curved with the convexity to the left. As it passed through

the left lower lobe it rapidly diminished in calibre till it ended at

the upper border of the lobe as a fine thread.

The vessel had, therefore, two curves with their convexities

to the left—a wide one in the abdomen, and a narrow one

within the lung.

When the roots of the lungs were dissected, it was seen that

the right one received two bronchial arteries from the thoracic

aorta, but the left one received none at all. Consequently, the

celiac artery must be regarded as a nutrient vessel to the

pulmonary tissue. It is evident, however, that it does not supply

all the lung for it is confined to the inner part of the left lower

lobe (see text-figure), and the branches which it gives off are very

small. I was unable to detect nutrient vessels to the upper lobe

and the outer part of the lower lobe of the left lung. There was

no trace of an anastomosis between the cceliac and esophageal

arteries which frequently make an arterial network on the

surface of the lung.

Branches :—The*cceliac artery gave off branches in the abdo-

men and thorax, as follows :—

1. Pancreatic artery which soon divided into two small twigs.

2. Splenic artery.

176 DR. C. F. SONNTAG ON ABNORMALITIES OF THE

3. Gastric artery which divided into two branches. The upper

one anastomosed with the cesophageal twigs, and the lower one

with the pancreatico-duodenal branch of the superior mesenteric

artery. The two branches diverged at right angles from the

parent trunk, and ran along the anterior surface of the stomach

just below the lesser curvature.

4, Qsophageal artery which supplies the lower end of the gullet.

It broke up into a brushwork of terminal vessels which anasto-

mosed with one another and with the upper branch of the gastric

‘artery. There was no connection between these vessels and any

within the thorax.

Text-figure 20.

PULMONARY | rower

WEIN I LOBE

LEFT

LUNG

DIAPHRAGM

OESOPHAGEAL ARTERY-___Aye

GASTRIC ARTERY..__£

SPLENIC ARTERY.__

BRANCH oF \B

SUPERIOR MESENTERIC

ARTERY

PANCREATIC

ARTERY

The Coeliac Axis Artery.

5. Pulmonary twigs which were given off from both sides of

the vessel within the lower lobe of the left lung. These vessels

were very small and soon divided into two.

No branches were given off from the part between the ceso-

phageal opening in the diaphragm and the point of entrance of

the vessel into the lung, and there was no hepatic branch at all,

and the hepatic blood-supply came from the superior mesenteric

artery instead.

It is difficult to give an explanation of the conditions described

ABDOMINAL ARTERIES OF A YOUNG PANDA. 77

above, and the source of these abnormalities may be due to an

error of development or a pathological cause. Perhaps the bron-

chial arteries to the left lung have been occluded by reason of

some disease and the cceliac artery has taken their place. If

there were an anastomosis between the bronchial, cesophageal,

and cceliae vessels, and the former had been occluded, the latter

might take their place and pass in from the surface to supply

the lung. In time the latter would enlarge and appear as in this

specimen.

i have referred to many books and papers on comparative

anatomy and embryology, but I have not found any mention of

a similar condition. Chauveau*, after describing some pecu-

liarities of the gastro-pulmonary anastomoses, says, ‘‘ There are

other varieties of which it is unnecessary to speak”; but it is

unlikely that he would dismiss such a condition so summarily if

he had seen it.

The Superior Mesenteric Artery.

The unusual feature of the superior mesenteric artery was the

great development of the pyloro-duodenal branch, in order to

take the place of the hepatic branch of the celiac artery.

This branch passed upwards and to the right to reach the

pyloro-duodenal junction. It passed bebind it and then curved

to the left below the liver, giving off branches to the right and

left lobes of the liver, and the gall-bladder. None of hese vessels

entered the portal fissure of the liver.

The Renal Arteries.

The right renal artery soon divided into two after its origin

from the abdominal aorta, and the left one exhibited a very

tortuous course. These are, however, not abnormalities, but

merely extreme degrees of a condition common among the

Carnivora.

The reader is referred to page 171, where another peculiarity

of the cceliac axis occurs.

* Chauveau, A. ‘The Comparative Anatomy of the Domesticated Animals,’

page 613.

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EXTERNAL CHARACTERS OF THE RATEL AND WOLVERENE. 179

14. On the External Characters of the Ratel (JJellivora)

and the Wolverene (Gulo)*. By R.I. Pocoog, F.R.S.,

HZ:

[Received April 13, 1920: Read April 13, 1920.]

: (Text-figures 14-18.)

ConTENTS. Page

Bat rear liVi POV Soecanneec conan eee a NPR a eC ese Secon ace Boar LTD

Hxternal form Ao OPER ONERE REE nedctotcanaueinc. lel:

Meg alent se aoa neler a Een aes ance ICEL

The Feet .. Rr ieee nctobon wd tol

The Anus and the External ‘Genitalia, Oo aah 2 eee 184

Conclusion .. q PER sna oes, fey

Introduction.

Writing on Jellivora in 1902 (Zool. of Egypt, Mammalia,

p- 245), de Winton remarked :—‘ A glance at the generic names

mentioned in the synonymy of the African species will show that

great uncertainty has existed as to the true relationship of the

animal. Even in the latest text-books it has been placed among

the badgers, while in truth it is nothing but a giant weasel

modified for digging and quite closely related to /ctonym.”

Without admitting the truth of the last proposition, it is

unquestionably the case that the majority of authors, judging

from the structure of the feet and the general form of the body,

followed Gray in ciassifying Mellivora in the group, family, or

subfamily, as the case may be, typified by dZeles. De Winton, on

the contrary, rejecting the external characters and relying upon

the teeth and skull, placed it in the subfamily Musteline, which

comprised the following genera :—ZJephitis, Conepatus, Galera,

Galictis, Mellivora, Ictonyx, Mustela (now Martes), Putorius (now

Mustela), Peecilogale, Lyncodon, and Gulot. It would have been

very dificult to define the Musteline, as thus constituted, and

de Winton, perhaps wisely, made no attempt to doso. That

question does not concern me now. The point to which attention

may be drawn is the placing of Jellivora in the same group as

Gulo.

Although no authors appear to be very clear about the precise

position of Gulo, its kinship with Mustela and Martes has been

generally admitted. The latest opinion on the subject is that of

Mr. G. 8. Miller, who proposed to make it the type of a distinct

subfamily, Gulonine, equivalent to the Musteline, Meline, and

Lutrine, these four subfamilies comprising the genera of

Mustelide of Western Europe (Cat. Mamm. Western Europe,

pp. 341 and 432, 1912).

* The facts recorded were based upon the examination of fresh material in the

Society’s Prosectorium.

y+ Weber (Die Saug. p. 537, 1904) followed de Winton in classifying Mellivora

and Gulo in the Mustel: ine.

LOK

180

MR. R. I. POCOCK ON THE EXTERNAL CHARACTERS

Text-figure 14,

A. Side view of head of Galo.

B. Front view of rhinarium of the same.

C. Piece of the pinna of the same, showing the bursa, with its anterior

lamina (a) turned forwards and its posterior lamina (7) turned back-

wards.

T). Side view of head of Mellivora.

EK. Front view of rhinarium of the same,

All figures, except.C, X 3.

OF THE RATEL AND THE WOLVERENE. 181

The claim of Gulo to rank as a subfamily may be set aside for

the present, but it appears to me that Miller’s guess at the

affinities of the genus with J/ellivora, expressed in the following

passage, is very wide of the mark. He wrote (op. cif. p. 433) :—

“The subfamily Gulonine, consisting of the genus Gulo alone,

is well characterised by its peculiarities of skull, teeth, and

external form. Though usually regarded as a near relative of

the Musteline, the genus more probably finds its true affinities

in the African genus Mellivora.” This means that in Muiller’s

opinion Gulo is more nearly allied to JMellivora than it is to

Mustela or Martes. The main purpose of this paper is to refute

that idea.

External forn,

Apart from being heavily and powerfully built and provided

with comparatively short tails, the two genera are quite unlike

in shape. Afellivora essentially resembles Meles or Taxidea, beng

low on the legs, plantigrade, broad across the back and flat

along the spine, the body being rather depressed than com-

pressed. The form is that of a terrestrial fossorial beast, without

power to leap and with running capacity reduced almost to the

Carnivore minimum. ‘The hair is everywhere short, sleek, and

coarse.

Gulo stands comparatively high on the legs, is digitigrade,

comparatively narrow across the back, and arched along the

spine, the body being compressed rather than depressed. The

form is that of a terrestrial and arboreal beast, capable of running

at some speed and of leaping to a moderate extent. ‘The hair is

everywhere, except on the face, soft, furry, and mixed with wool.

The Head.

The top of the head in JMellivora is high and arched ; in Gulo

it is low and flat. (Text-fig. 14, A, D.)

The tufts of facial vibrisse are alike in the two genera

in the sense that they are the same in number and _ position

as in typical predatory Carnivores like the Canide, Viverride,

Mungotidz, and most Mustelide. (Text-fig. 14, A, D.)

The rhinariwm of Mellivora recalls that of Jeles in having a

well-defined area encircling the nostrils below and laterally,

although this area is considerably shallower than in that genus.

Also it is not continued inferiorly as a philérwm dividing the

upper lip. In Gulo the rhinarium is like that of Canis, being

continued inferiorly as a distinct philtrum dividing the upper

lip, and the nostrils are encircled laterally by an area of naked

nm) skin continuous with the philtrum in front. (Text-fig. 14,

B, E.

The external ear in Mellivora has been described as absent

The truth is that there is no definite laminate pinna standing

away from the head, the cavities of the ear being merely

182 MR. R. I. POCOCK ON THE EXTERNAL CHARACTERS

surrounded above and behind by a thickening of the integument.

The supratragus (plica principalis) is a simple oblique ridge, the

tragus, antitragus, and other inferior ridges are hardly apparent

and there is no trace of the bursa (text-fig. 14, D). In Gulo the

ear is quite normally developed, the pinna standing away from

the head as a mobile lainina. The cavity of the ear is larger than

in Mellivora, the supratragus is a larger ridge with a semiglobular

thickening, the tragus and antitragus and the normal ridges in-

ternal to them are well defined, and the bursa is well developed,

its posterior wall consisting of a semioval lamina arising behind

the margin of the pinna (text-fig. 14, A, C).

The Feet.

The fore foot of Mellivoraw is provided with long, powerful,

blunt fossorial claws The digits are short and unevenly spaced,

Text-figure 15.

A. Lower side of right hind foot of Mellivora.

B. Lower side of right fore foot of the same.

the distance between 1 and 2 being considerably greater than

the distances between the others. Digits 2, 3, and 4 are rather

tightly tied together by webbing, which extends more than half-

way along the digital pads. Digit 5 has more freedom of

movement. ‘The plantar pad isa large, irregularly semicircular

mass, with its four elements ill defined. It is followed by two

carpal pads separated from it by a deep groove. The external

carpal pad is a large mass; the internal is much smaller and

Text-figure 16,

OF THE RATEL AND THE WOLVERENE,. 183

differs in its smoothness from the rest of the pads, which are

coriaceous. The whole of the under side of the foot back to and

including the carpal pads is hairless. (Text-fig. 15, B.)

=/ Zig Ss NO

BF SZ G Yl:

ZG Z seas YP yi fff,

LL Ly Cy SEZ

The hind foot, allowing for its greater length, greater narrow-

ness, and short claws, is very similar to the fore foot; but the

pads of digits 3 and 4 are fused proximally, and there is a

single large metatarsal pad, narrower behind than in front and

extending along the middle line half-way between the plantar

B. Lower side of right fore foot of the same.

A, Lower side of right hind foot of Gulo.

Xt.

x}.

184 MR. R. I, POCOCK ON THE EXTERNAL CHARACTERS

pad and the heel. Its posterior edge is ill defined and the

integument of the foot behind it is naked and wrinkled. (Text-

fig, 15, A.)

The feet* of Gulo are totally different. Except that the fore

foot is shorter and broader and is provided with carpal pads, the

two are very much alike and may be described together. The

digits are longer than in J/edlivora, and are tolerably evenly spaced

and widely separable; the pollex and halluxsare relatively less

reduced than in Mellivora and the webs tying the digits together

are wide, and the middle of the edge of each is approximately on

a level with the proximal end of the digital pads, which are well

defined, oval in shape, and, like the plantar pads, coarsely striate.

The claws are alike in size and shape on the two feet, being

moderately long, curved, and sharp. The plantar pads are much

reduced antero-posteriorly. The four so-called interdigital

elements are well defined and connected by narrower strips, the

whole forming an irregularly shaped, curved, transverse band,

the concavity of the curve facing the carpus and tarsus. The

median largest element of this pad has its anterior edge emar-

ginate. Some distance behind the plantar pad on the fore foot

there are two small, reniform, carpal pads, the outer as large as

a bean, the inner as large as a pea. The digital and plantar

pads are to a great extent overlapped by hair and the carpal pads

are entirely concealed; but there is no trace of a metatarsal

pad on the hind foot. Except for the pads the whole of the

under side of the feet is covered with hair, which on the digits

and webs and behind the plantar pad is soft and woolly; but on

each side of the carpus and along the inner side of the metatarsus

the hairs are coarse and form a stiff bristly brush. (Text-fig. 16,

AG Be)

The Anus and the External Genitalia.

The anus in Mellivora is sunk in the centre of a circular area

of radially corrugated skin which folds over it and conceals it,

the upper and lower margins of this area meeting to forma

transverse rima—the condition very closely resembling that of

the Mungotide. The two anal glands are of great size, and

discharge copiously a suffocating fluid exactly as in the Skunks

(Mephitis, Conepatus), Zorilles (Ictonyx), Grison (Grisonia), and

Teledu (/ydaus) (text-fig. 17, A, B). In Gulo the anus is super-

ficial and not insunk, being merely surrounded by an area of

naked skin as in Canidee, Felidee, and the majority of Carnivora.

The anal glands are of normal size+, and discharge a yellow

fluid, the odour of which is musteline, but not so pungent as in

the Polecat (ustela putorius).

The anal and genital region in Mellivora is naked or scantily

hairy ; in Gulo it is thickly furred.

Gulo.

* Boas (Zool. Anz. xxxiv. p. 532, 1909) figures the hind foot, stripped of hair, of

+ Gray’s statement that they are absent is wrong.

OF THE RATEL AND THE WOLVERENE. 185.

Text-figure 17.

A. Anal area of Mellivora 3, with the root of the tail above, showing the

closed orifice of the anal sack.

B. The same partly dissected to show tbe anal glands. a.s., anal sack

spread open with a bristle passed through the duct of the anal gland

and emerging at the side of the anus; a.g., anal glands, on the right

side entire, on the left side opened to show the reservoir and the

bristle; sc., scrotum.

The penits.—In both the genera the prepuce, as in all Mustelide,

is situated far in advance of the scrotum, and the penis is pro-

vided with a long stout baculum. In Mellivora the baculum *

* This bone was described by Gilbert (Morph. Jahrb. xviii. p. 817, 1892).

I redescribed and figured it in 1918 (Ann. Mag. Nat. Hist. (9) i. p. 311, figs. c-m

p. 309). The baculum of Gulo was figured and described by Pohl (Jena Zeitschr.

xlv. p. 385, 1909).

186 MR. R. I. POCOCK ON THE EXTERNAL CHARACTERS

is relatively short and stout, very thick at its proximal end, and

grooved throughout its length below. The apex is sharply

upeurled and transv ersely expanded. into a hollowed, somewhat

basin-shaped disk, the rim of which is interrupted in front by a

channel, which is continued as a groove down the front of the

upeurved termination of the bone. This structure is perfectly

symmetrical in all its parts. (Text-fig. 18, B, E.)

Text-figure 18.

A. Anal and genital area of Gulo 9; a., anus with orifice of anal gland

on each side; v., vulva.

B. Lateral view of penis of Mellivora.

C. The same of Gulo.

D. Front view of the tip of the baculum of Gulo.

EK. The same of Vellivora.

In Gulo the baculum is relatively long and thin, without any

thickening at its proximal end and grooved beneath only at its

distal end, where it is gradually and lightly upcurled and decidedly

compressed. The apex bears a short, moderately high, com-

pressed, longitudinal crest, beneath which is a pair of short,

condyle-like processes separated by a deep cleft continuous with

the groove behind them. These processes are not symmetrical

and the crest is not upright. (Text-fig. 18, C, D.)

OF THE RATEL AND THE WOLVERENE. 187

The vulva.—I have no notes on the female generative organs

of Mellivora; but in Gulo the vulva is a naked piriform pro-

minence a little below the anus, from which it is separated by a

band of hair, and the orifice is a vertically elongated slit. (Text-

fig. 18, A.)

Conclusion.

From the characters above described it is evident that there

is no particular resemblance in any respect between JMellivora

and Gulo. The differences, on the contrary, are profound.

Miller’s suggestion, therefore, of kinship between the two genera

must be dismissed, and it appears to me that the evidence on

_ this head supplied by the skulls confirms that of the external

characters here discussed.

To what genera of Mustelide, then, are Gulo and Mellivora

related? Gulo, in my opinion, might be described broadly but

with much truth as a gigantic heavily-built Marten (J/artes).

I can find nothing in thestructure of the skull and teeth opposed

to the view that these two genera are related, and tolerably

closely related*. There are also no differences of moment

between them in the structure of the ear, of the rhinarium, of

the upper lip, of the anus, and of the external genitalia ; and the

feet of Gulo are little more than broad, short editions of those of

Martes tT, the claws, pads, disposition of the digits, hairiness of

the soles, etc., being strikingly alike in the twoanimals. Finally,

it does not appear to me that Gulo differs much more from

Martes than Martes differs from J/ustela or Vormela, the three

genera which constitute the subfamily Musteline as defined in

Miller’s volume. If this be true, the subfamily Gulonine can

hardly be considered a defensible group.

Mellivora is much more difficult to classify. The position

assigned to the genus by authors will depend upon their views

regarding the plasticity of the skull and teeth as compared with

the plasticity of the ears, feet, and other external organs described

in this paper. I cannot agree with de Winton that the genus fs

closely related to Jctonywx, and I doubt its near affinity with the

South American genus Galera; and although the feet and

rhinarium and general form are very like those of Meles, and the

pouched anus occurs in both the genera, the structure of the

skull and teeth should, I think, exclude Wellivora from a place

in the Meline, despite the heterogeneity of that subfamily as

constituted in the current text-books. The best way of dealing

with the genus at present seems to me to follow Gill in making

it the type and sole representative of a special subfamily, the

Mellivorine,

* The tip of the baculum is also asymmetrical in structure in Martes and

Charronia as it is in Mustela.

+ For figures and descriptions of the feet and ears of Martes martes and

M. foina, see my paper on these two species (Proc. Zool. Soc. 1914, pp. 1062-1068).

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ol dtive ytiah Leet St ae peal kL da Peon OF ‘wil a

ia daa ath. dutoddly howe A a) Aili” see tat A ee: {6

elt fian seule fh, looqaarbt ell eat d psa ascot leevouvoy fete see Pantene

ali to auintodrits aths eA TR dels. iLicet tH giser HMA EER Hradopiae at i

ory fy Fe AEE Wy eh sah 13 at aa Ke: tobe wh tT rd eh rand fetsiy’ tue

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witthtety to saw dead ad Po lofted dgor'or Sit aRF int

a tiotdae ast Tt watloPod fin OP atone Hivkorey diy tee

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nid yg) fesgih lith,

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fa er)

Piss : 2 grey Ta hws te 1). ree arn ot Bast: aa

NCH ROS pg ol iP TRAN rel) eae

oe”

ON ADDITIONS TO THE MENAGERIE. 189

EXHIBITIONS AND NOTICES.

February 10th, 1920.

Prof, KE, W. MacBripz, F.R.S., F.Z.S., Vice-President,

in the Chair.

Mr. R. I. Pocock, F.R.S., read the following Report on the

Additions made to the Society’s Menagerie during the months of

November and December, 1919 :—

NOVEMBER.

The registered additions to the Society’s Menagerie during the

month of November were 170 in number. Of these 83 were

acquired by presentation, 37 were deposited, 11 were exchanged,

35 were purchased, and 4 were born in the Menagerie.

The following may be specially mentioned :—

1 Mongolian Wild Horse (Hquus prjevalskit), deposited on

November 5th.

1 Lion (felis leo), from Senegambia, purchased on November 8th.

2 Cape Hyrax (Procavia capensis), from §, Africa, purchased

on November 10th.

A collection from India, including 1 Axis Deer, 5 Indian

Porphyrios, 1 Sarus, 2 Common Cranes, 3 Burmese Peafow],

presented by David Ezra, on November 4th.

A collection of Reptiles from California, including a Desert

Iguana (Dipsosaurus dorsalis), a Red-Ringed Snake (Coronella

zonata), and a Western Ring-necked Snake (Coronella amabilis),

all new to the Collection, presented by Dr. Cecil E, Reynolds.

DECEMBER.

The registered additions to the Society's Menagerie during

the month of December were 96 in number. Of these 38 were

acquired by presentation, 36 were deposited, 18 were purchased,

and 4 were born in the Menagerie.

The following may be specially mentioned :—

2 Tigers (Melis tigris), $ 2, and one Caracal (Felis curacal),

3, from India, presented by Alfred Hzra, Esq., V.P.Z.S., on

December 30th.

1 Spotted-necked Otter (Lutra maculicollis), new to the

Collection, from Sierra Leone, purchased on December 23rd.

4 Migratory Hamsters (Cricetulus migratorius), new to the

Collection, from Kazvin, N. Persia, presented by P. A. Buxton

on December Ist.

6 Mariqua Sun-birds (Cinnyris mariquensis), new to the

Collection, from 8. Africa, deposited on December 21sé.

190 PHOTOMICROGRAPHS OF ACARI.

2 Sun-Bitterns (Hurypyga helias), from South America, pur-

chased on December 16th.

1 Matamata Terrapin (Chelys jimbriata), from the Rio Negros,

presented by Dr. A. Bremner.

Mr. A. J. Euwes, F.R.S., communicated a letter relative to

the condition of the herds of Moose in Yellowstone Park.

Mr. D. Seru-Smiru, F.Z.8., read a letter from Messrs. Tullis,

Russell & Co., drawing attention to an interesting case of

response of Sparrows to colour, in which the birds appeared to

be peculiarly attracted to a Paper-making machine when paper

of a dark blue tint was being run off.

Mr. E. G. Boutencer, F.Z.S., Curator of Reptiles, exhibited

living specimens of a remarkable new Land-Tortoise, Zestudo

loveridgii, recently discovered by Mr. A. Loveridge in the neigh-

bourhood of Dodoma, East Africa, and described by Dr. G. A.

Boulenger, F.R.S. This Tortoise differs from all previously

described Lind-Tortoises in the carapace being quite flat, in the

so-called “shell” being perfectly soft, and in the complete

absence of ribs, costal and neural bony plates. Mr. Boulenger

stated that quite young examples differed from the specimens

exhibited, their carapace being dome-shaped and possessing ribs

and all other bones present in previously described Tortoises.

In referring to its habits, it was stated that the animal, having

been deprived of the protection of a bony “shell,” had taken to

living much after the manner of a Lizard, inhabiting holes in

the rocks, and that, according to Mr. Loveridge, great difficulty

is experienced in extracting the Tortoise from these holes, owing

to its habit of distending itself, and thereby wedging itself firmly

in the rock-cavity.

Mr. F. Marvin Duncay, F.Z.8., exhibited and made remarks

on a series of photomicrographs of Acari from the lungs of

Macacus rhesus, illustrating the larval, nymph, and adult stage

of the Acarid. He stated that both young and adult Rhesus

Monkeys appeared to be infected, but so far as his observations

had gone, the presence of the Mites in the lungs had in no case

been the cause of death. So far eggs had not been observed in

the vesicles formed by the presence of the Mites, though serial

sections of adult Acarids had shown the egg in an advanced state

of development, pointing to the probability of this stage being

to)

completed within the body of the female.

PHOTOMICROGRAPHS OF A FEMALE MOLE-FLEA, 191

Mr. R. H. Burne, F.Z.8S., exhibited some skeletons of fcetal

Mammals prepared by ‘Tadpoles, and drew attention to the

possible advantage of such a method over the more laborious

process by hand.

February 24th, 1920.

A. Smita Woopwarp, Hsq., LL.D., F.R.S., Vice-President,

in the Chair.

Mr. R. I. Pocock, F.R.S., read the following Report on the

Additions made to the Society’s Menagerie during the month of

January 1920 :—

The registered additions to the Society’s Menagerie during

the month of January were 109 in number. Of these 24 were

acquired by presentation, 17 were deposited, 61 were purchased,

5 were received in exchange, and 2 were born in the Menagerie.

The following may be specially mentioned :—

2 Hybrid Black and Brown Bears (Ursus americanus 3 X

U. arctos 9), born in the Menagerie on January Oth.

1 Persian Gazelle (Gazella subgutturosa 9 ), from Mesopotamia,

presented by the Ist Battn. lst Highland Light Infantry.

5 Bennett’s Wallabies (Macropus bennetti, 3 $, 29) (Tas-

mania), received in exchange on January 31st.

1 Naked-throated Bell-bird (Chasmorhynchus nudicollis), from

Brazil, purchased on January Ist.

2 Soft-shelled Land-Tortoises ( V’estudo loveridgii), from Dodoma,

E. Africa, presented by Arthur Loveridge, January 26th.

Mr. F. Martin Duncan, F.Z.8., exhibited photomicrograplis

of a female Mole-Flea (Hystrichopsylla talpe), and drew attention

to the presence of two spermathece, a feature in the anatomy of

the genital organs of this flea which appeared to have hitherto

escaped notice. Several specimens had been submitted to him,

and in all the females this characteristic feature was present.

He also pointed out that clinging to the abdomen of the flea, by

means of their sucker-discs, were a number of hypopial nymphs

of one of the Tyroglyphide. He had not at present been able

to determine to which species they belonged, but it was inter-

esting to note their attachment to the flea inasmuch that it was

occasionally taken in the nests of Wild Bees, and might act as a

dispersal agent of the Acari.

eee

192 ON ADDITIONS TO THE MENAGERIE.

March 16th, 1920.

Prof. E. W. MacBripg, F.R.S8., F.Z.8., Vice-President,

in the Chair.

Mr. R. J. Pocock, F.R.S., read the following Report on the

Additions to the Society's Menagerie during the month of

February 1920 :—

The registered additions to the Society’s Menagerie during

the month of February were 92 in number. Of these 16 were

acquired by presentation, 19 were deposited, 52 were purchased,

1 was received in exchange, and 4 were born in the Menagerie.

The following may be specially mentioned :—

2 Golden-crested Penguins (Catarrhactes chrysolophus), from

South Georgia, new to the Collection, purchased on February

10th.

A collection of 8. African Reptiles, including 4 White-throated

Monitors (Varanus albigularis) and 2 Derbian Zonures (Zonurus

derbianus), purchased on February 25th.

3 Beaver-Rats or Coypus (M/yocastor coypus), born in the

Menagerie on February 6th.

1 Spotted Cavy (Calogenys paca), born in the Menagerie on

February 29th.

Mr. E. G. Boutencer, F.Z.8., exhibited and made remarks on

a Frog with a duplicate foot.

Prof. J. P. Hitu, F.R.S., exhibited and made remarks on an

Embryo obtained from a Kangaioo recently living in the Society’s

Menagerie.

March 30th, 1920.

A. Suita Woopwarp, Esq., LL.D., F.R.S., Vice-President,

in the Chair.

Sir Frank Cotyrer, K.B.K., F.R.C.8S., exhibited and made

remarks on a series of photographs of skulls of Macacus rhesus,

showing pathological conditions of the teeth.

Prof. H. Maxweti Lerroy, F.Z.S., exhibited photographs

attesting the existence of Egret Farms in Sind.

ON SNAKES FROM EAST AFRICA. 193:

Mr. R. H. Burne, M.A., F.Z.8., exhibited a series of Pigs’

mandibles from the New Tye bails, showing overgrowth of tite

lower tusk owing to removal of the maxillary tusk.

April 13th, 1920.

A. Smita Woopwarp, Esq., LL.D., F.R.S., Vice-President,

in the Chair.

Mr. R. I. Pococs, F.R.S., exhibited and made remarks on two-

specimens of Fournier’s Hutia (Capromys pildrides), now living

in the Society’s Menagerie.

Mr. R. I. Pococn, F.R.S., gave an exhibition, illustrated by

lantern-slides, to show the differences in external characters.

between the Ratel (Mellivora) and the Wolverene (Gulo). He

pointed out that Mr. Gerrit Miller’s suggestion that the two

genera were related was unsupported by the facts.

April 27th, 1920.

A. Smita Woopwarb, Hsq., LL.D., F.R.S8., Vice-President,

in the Chair.

The Secretary read the following Report on the Additions to

the Society’s Menagerie during the month of March 1920 :—

The registered additions to the Society’s Menagerie during

the month of March were 106 in number. Of these 20 were

acquired by presentation, 64 were deposited, 15 were purchased,

3 were received in exchange, and 4 were born in the Menagerie.

The following may be specially mentioned :—

1 Lion (Felis leo), from Sennar, presented by the 8th Battn.

of the Hampshire Regt., on March 24th.

1 Eland (Zaurotragus ory#), born in the Menagerie on

March 15th.

2 Barbary Sheep (Ammotragus lervia), from Morocco, presented

by H.M. The King, on March 16th.

Mr. Artuur LovertpGe exhibited and made remarks on a

collection of Snakes which he had obtained in East Africa during

the years 1915-1919.

Proc. Zoou. Soc.— 1920, No. XILI. 13

194 ON A SPECIMEN OF AMBLYSTOMA.

Miss L. E. CuresMan, F.E.S., exhibited and described a series

of lantern-slides illustrating the life-history and habits of the

Ichneumon-fly, Rhyssa perswasoria.

Dr. P. Cuatmers Mircuett, F.R.S., exhibited and made

remarks on a series of photographs, taken by Sir H. A. Byatt,

K.C.M.G., in German East Africa, of the rare Abbot’s Duiker

(Cephalophus spadiz).

Mr. D. Suru-Smuirn, F.Z.S., exhibited a series of lantern-slides

showing the display of the male Monaul Pheasant (Lophophorus

impeyanus).

Mr. L. Hoesen, M.A., B.Sc., exhibited a specimen of Ambly-

stoma, the metamorphosis of which had been brought about by

one month’s feeding with Ox thyroid, and drew special attention

to the precocious transition to the Amblystoma-type of pigmen-

tation during the metamorphosis.

No. 199.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

February 10th, 1920.

Prof. E. W. MacBrips, F.R.S., F.Z.S., Vice-President,

in the Chair.

Mr. R. I. Pocock, F.R.S., read a report on the Additions made

to the Society's Menagerie during the months of November and ~

December, 1919.

Mr. A. J. Etwes, F.R.S., communicated a letter relative to

the condition of the herds of Moose in Yellowstone Park.

Mr. D., Seru-Smuirn, F.Z.S., read a letter from Messrs. Tullis,

Russell & Co., drawing attention to an interesting case of

response of Sparrows to colour.

Mr. R. I. Pocock, F.R.S., exhibited and made remarks on

two photographs of a Chinese Serow (Capricornis argyrochcetes).

Mr. E. G. Bounencer, F.Z.S., Curator of Reptiles, exhibited

living specimens of a remarkable new Land-Tortoise, Testudo

loveridgit, recently discovered by Mr. A. Loveridge in the neigh-

bourhood of Dodoma, East Africa, and described by Dr. G. A.

Boulenger, F.R.S. This Tortoise differs from all previously

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent's Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications; but it may be obtained on the

day of publication at the price of Sixpence, or, if desired, sent post-free for

the sum of Six Shillings per annum, payabie in advance.

described Land-Tortoises in the carapace being quite flat, in the

so-called “shell” being perfectly soft, and in the complete

absence of ribs, costal, and neural bony plates. Mr. Boulenger

stated that quite young examples differed from the specimens

exhibited, their carapace being dome-shaped and possessing ribs

and all other bones present in previously described Tortoises.

In referring to its habits, it was stated that the animal, having

been deprived of the protection of a bony “ shell,” had taken to

living much after the manner of a Lizard, inhabiting holes in

the rocks, and that, according to Mr. Loveridge, great difficulty

is experienced in extracting the Tortoise from these holes, owing

to its habit of distending itself, and thereby wedging itself firmly

in the rock-cavity.

Mr. F. Marvin Duncan, F.Z.S., exhibited and made remarks

on a series of photomicrographs of Acari from the lungs of

Macacus rhesus, illustrating the larval, nymph, and adult stage

of the Acarid. He stated that both young and adult Rhesus

Monkeys appeared to be infected, but so far as his observations

had gone, the presence of the Mites in the lungs had in no case

been the cause of death. So far eggs had not been observed in

the vesicles formed by the presence of the Mites, though serial

sections of adult Acarids had shown the egg in an advanced state

of development, pointing to the probability of this: stage being

completed within the body of the female.

Mr. R. H. Burye, F.Z.S8., exhibited some skeletons of fetal

Mammals prepared by Tadpoles, and drew attention to the

possible advantage of such a method over the more laborious

process by hand.

Dr. C. F. Sonnvaa, Ch.B., F.Z.S., exhibited and made remarks

on a series of black-board drawings and Jantern-slides illustrating

several unusual features in the peritoneum of a Raccoon.

Mr. H. R. Hoge, F.Z.8., read a paper entitled “On some

Australian Opiliones,” and pointed out that the genera and

species described belonged to the suborders Palpatores and

Laniatores—the Palpatores being represented by the genera

Pantopsalis and Macropsalis of the family Phalangiide and the

Laniatores by genera of Triwnobunide and Tricwnonychide. In

the case of the Phalangiide he had been able to establish that

long mandibles were a male and short mandibles a female

character.

Dr. C. F. Sonnrac, Ch.B., F.Z.S., communicated a paper on

the “ Larynx and Cisophagus of a Common Macaque, exhibiting

several unusual Features.”

In the absence of the Authors, Messrs. Rownanp E. Turner and

James WATERSTON, their paper on “ A Revision of the Ichneu-

monid Genera Labiwm and Pecilocryptus” was taken as read,

The next Meeting of the Society for Scientific Business will be

held on Tuesday, February 24th, 1920, at 5.30 p.m., when the

following communications will be made :—

The SECRETARY.

Report on the Additions to the Society’s Menagerie during

the month of January, 1920.

E. G. Boutencer, F.Z.S.

On some Lizards of the Genus Chalcides.

N.S. Lucas, M.B., F.Z.S.

eS a ee oe

Report on the Deaths in the Gardens in 1919: with Notes

on BRickets and Avian Hnteritis.

Srantey Hirst, F.Z.8.

Revision of the English Species of Red Spider (Genera

Tetranychus and Oligonychus).

The following Paper has been received :—

CG. Forster Cooper, M.A., F.Z.8.

ede eed elon eau eee

The Anthracotheriide of the Dera Bugti Deposits in

Baluchistan.

The Publication Committee desire to call the attention of

those who propose to offer Papers to the Society, to the great

increase in the cost of paper and printing. This will render it

necessary for the present that papers should be condensed, and

be limited so far as possible to the description of new results.

Communications intended for the Scientific Meetings should

be addressed to |

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Society oF Lonpoy,

Recent’s Park, Lonpon, N.W. 8.

February 16th, 1920.

No. 200.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON,.*

February 24th, 1920.

A. Smita Woopwarp, Esq., LL.D., F.R.S., Vice-President,

in the Chair.

Mr. R. I. Pococn, F.R.S., read a Report on the Additions

made to the Society's Menagerie during the month of January,

1919.

Mr. R. I. Pecocx, F.R.S., exhibited and made remarks on a

photograph of a young Pigmy Hippopotamus, and drew attention

to the striking difference in the shape of the ear, which is lenger

and narrower than in the adult animal.

Mr. F. Martin Duncan, F.Z.8., exhibited photographs of a

female Mole-Flea (Hysirichopsylla talpe), and drew attention to

the presence of two spermathece, a feature in the anatomy of

the genital organs of this flea which appeared to have hitherto

escaped notice. He also pointed out that clinging to the abde-

men of the flea by means of the sucker-discs were a number of

hypopial nymphs of one of the Tyroglyphide.

Mr. E. G. BouLencer, F.Z.8., communicated a paper dealing

with the geographical distribution and classification of some

* This Abstract is published by the Society at its offices, Zoolocical Gardens,

Regent's Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subseribe to the Publications ; but it may be obtained on the

day of publication at the price of Sixpence, or, if desired, sent post-free for

the sum of Sex Shillings per annum, payable in adyanee,

Lizards of the genus Chalcides. Chalcides occellatus was stated

to be divisible apart from the typical form into seven varieties

or subspecies. It was of interest from the evolutionary point of

view that these forms were connected in such a manner that. it

was possible to trace every link in the chain from a stout type

with 40 scales round the body from Morocco toa slender type

with only 22,scales from Abyssinia.

Dr. N.S. Lucas, F.Z.8., read a Report on the Deaths in the

Gardens in 1919, and made some observations on Rickets and

Avian Enteritis.

In the absence of the Author, Mr. Stantey Huirst’s paper on

“The Revision of the English Species of Red Spider (Genera

Tetranychus and Oligonychus),” was taken as read.

Professor W. N. F. Woopnanp gave a résumé of Mr. D. R.

BHATTACHARYA'’s paper on “The Aortic Ligament in Indian

Fishes.”

The next Meeting of the Society for Scientific Business will be

held on Tuesday, March 16th, 1920, at 5.30 p.m., when the

following communications will be made :—

R. I. Pococx, F.R.S.

On the External Characters of South American Monkeys.

C. F. Sonntag, M.D., Ch.B., F.Z.S.

The Comparative Anatomy of the Tongues of the Mam-

malia :—I. General Description of the Tongue.

—

The following Paper has been received :—

Ue See ED OND BLS.

On Abnormalities of the Abdominal Arteries of a young

Panda.

The Publication Committee desire to call the attention of

those who propose to offer Papers to the Society, to the great

increase in the cost of paper and printing. This will render it

necessary for the present that papers should be condensed, and

be limited so far as possible to the description of new results.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Society or LONDON,

ReGENtT’s Park, Lonpon, N.W. 8.

March 2nd, 1920.

No. 201.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

March 16th, 1920.

Prof. E. W. MacBripg, F.R.S., F.Z.S., Vice-President,

in the Chair.

My. R. I. Pocock, F.R.S., read a Report on the Additions to

the Society’s Menagerie during the month of February, 1920.

Mr. E. G. Bounencer, F.Z.S., exhibited and made remarks on

a Frog with a duplicate foot.

Prof. J. P. Hit, F.R.S., exhibited and made remarks on an

Embryo obtained from a Kangaroo recently living in the Society’s

Menagerie.

Mr. R. I. Pocock, F.R.S., read a paper, illustrated by lantern-

slides, on the external characters of the South American Monkeys,

and showed the variations in the range of structure of the ears,

nose, hands, and feet and external genitalia.

Dr. C. F. Sonntac, F.Z.8., communicated his paper on “The

Comparative Anatomy of the Tongues of the Mammalia,” and,

having first outlined the plan which would be followed in his

series of comparative studies, proceeded to describe the different

divisions of the tongue and the physical characters of each. He

demonstrated by diagrams and lantern-slides the different forms

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent's Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications; but it may be obtained on the

day of publication at the price of Sixpence, or, if desired, sent posi-free for

the sum of Six Shillings per annum, payable in advance.

which the papillz and openings of Wharton’s Ducts can assume

among the Mammalia, and exhibited specimens illustrating the

shapes and colours of the tongue, and arrangements for cleaning

the teeth.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, March 30th, 1920, at 5.30 p.m., when the

following communications will be made :—

Sir Frank Coryer, K.B.E., F.R.CS.

Exhibition of skulls of Wacacus rhesus.

©. F. Sonnac, M.D., Ch.B., F.Z.S.

On Abnormalities of the Abdominal Arteries of a young

Panda.

ArtHur LOVERIDGE.

Notes on East African Lizards collected 1915-1919, with

Descriptions of a new Genus and Species of Skink, and a new

Subspecies of Gecko.

A. M. ALtTson.

The Life-history and Habits of Two Parasites of the

Blowfly.

The following Papers have been received :-—

Arraur WILEY, F.R.S., F.Z.8.

An Apodous Ama calva.

H. A. Bayuis, M.A., and Ciayton, Lane, M.D.

A Revision of the Nematode Family Gnathostomide.

W. J. Daun, D.Sc., F.Z.8.

The Onychophora of Western Australia.

The Publication Committee desire to call the attention of

those who propose to offer Papers to the Society, to the great

increase in the cost of paper and printing. ‘This will render it

necessary for the present that papers should be condensed, and

be limited so far as possible to the description of new results.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Society or Lonpon,

ReceEnt’s Park, Lonpon, N.W. 8.

March 23rd, 1920,

No. 202.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

March 30th, 1920.

Dr. A. Suir Woopwarp, LL.D., F.R.S., Vice-President,

in the Chair.

Sir Frank Cotyrer, K.B.E., F.R.C.S., exhibited and made

remarks on a series of photographs of skulls of Macacus rhesus,

showing pathological conditions of the teeth.

Prof. H. Maxwenut Lerroy, F.Z.S., exhibited photographs

attesting the existence of Egret Farms in Sind.

Mr. R. H. Burne, M.A., F.Z.8., exhibited a series of Pigs’

mandibles from the New Hebrides, showing overgrowth of the

lower tusk owing to removal of the maxillary tusk.

Dr. C. F. Sonnrac, Ch.B., F.Z.8., read a paper on “ Abnor-

malities of the Abdominal Arteries of a Young Panda.”

In the absence of the Author, Mr. A. Loverince, his paper

on “ Kast African Lizards collected in 1915-1919, with Descrip-

tion of a new Genus and Species of Skink and a new Subspecies

of Gecko,” was taken as read.

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent's Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the * Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications; but it may be obtained on the

day of publication at the price of Sixpence, or, if desired, sent post-free for

the sum of Sta Shillings per annum, payable in advance.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, April 13th, 1920, at 5.30 p.m., when the

following communications will be made :—

ArtHur WILEY, F.R.S., F.Z.S.

An Apodous Amia calva.

H. A. Bayuis, M.A., and Crayton Lang, M.D.

A Revision of the Nematode Family Gnathostomide.

W. J. Dakin, D.Sce., F.Z.S.

The Onychophora of Western Australia.

A. M. Autson.

The Life-history and Habits of Two Parasites of the

Blowfly.

The Publication Committee desire to call the attention of

those who propose to offer Papers to the Society, to the great

increase in the cost of paper and printing. This will render it

necessary for the present that papers should be condensed, and

be limited as far as possible to the description of new results.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Society oF Lonpon,

Recent’s Park, Lonpon, N.W. 8.

April 6th, 1920.

No. 203.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

April 13th, 1920.

Dr. A. Suira WoopwarbD, LL.D., F.R.S., Vice-President,

in the Chair.

Mr. R. I. Pocock, F.R.S., exhibited and made remarks on two

specimens of Fournier’s Hutia (Capromys pilorides), now living

in the Society’s Menagerie.

Mr. R. I. Pococs, F.R.S., gave an exhibition, illustrated by

lantern-slides, to show the differences in external characters

between the Ratel (Mellivora) and the Wolverine (Gulo). He

pointed out that Mr. Gerrit Miller’s suggestion that the two

genera were related was quite unsupported by the facts.

In the absence of the Author, Prof. ArtHurR WIittey, F.R.S.,

his paper on “ An Apodous Amia calva” was taken as read.

Mr. H. A. Baytis, M.A., and Lt.-Col. Crayton Lanz, M.D.,

gave a résumé, illustrated with lantern-slides, of their paper on

“A Revision of the Nematode Family Gnathostomide.”

Prof. H. Maxweti Lerroy, F.Z.S., communicated a paper by

Mr. A. M. Aurson on “The Life-history and Habits of Two

Parasites of Blowflies.”

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent’s Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications ; but it may be obtained on the

day of publication at the price of Sixpence, or, if desired, sent post-free for

the sum of Sar Shillings per annum, payable in advance.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, April 27th, 1920, at 5.30 p.m., when the

following communications will be made :—

F. F. Larpuaw, M.A., F.Z.S.

Contributions to a Study of the Dragonfly Fauna of Borneo.

Part IV.—A List of the Species known to occur in the

Island.

R. Broom, M.D., D.Sc., C.M.Z.8., F.RB.S.

On some new Therocephalian Reptiles from the Karroo

Beds of South Africa.

The Publication Committee desire to call the attention of

those who propose to offer Papers to the Society, to the great

increase in the cost of paper and printing. This will render it

necessary for the present that papers should be condensed and

be limited so far as possible to the description of new results.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Society or Lonpon,

ReceEnt’s Park, Lonpon, N.W. 8.

April 20th, 1920.

No. 204.

ABSTRACT OF THE PROCEEDINGS

ZOOLOGICAL SOCIETY OF LONDON.*

April 27th, 1920.

Dr. A. Suirq Woopwarp, LL.D., F.R.S., Vice-President,

in the Chair.

The Sucrerary read a Report on the Additions to the Society’s

Menagerie during the month of March 1920.

Mr. Artur LoverimpGe exhibited and made remarks on a

collection of Snakes which he had obtained in East Africa during

the years 1915-1919,

Miss L. K. Currsman, F.E.S., exhibited and described a series

of lantern-slides illustrating the life-history and habits of the

Ichneumon-fly, Rhyssa persuasoria.

Dr. P. CHAuMers Mircyett, F.R.S., exhibited and made

remarks on a series of photographs, taken by Sir H. A. Byatt,

K.C.M.G., in German East Africa, of the rare Abbot’s Duiker

(Cephalophus spadiz).

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent's Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications; but it may be obtained on the

day of publication at the price of Sixpence, or, if desired, sent post-free for

the sum of Six Shillings per annum, payable in advance.

Mr. D. Srers-Smitg, F.Z.8., exhibited a series of lantern-slides

showing the display of the male Monaul Pheasant (Lophophorus

Impeyanus ).

Mr. L. Hoasen, M.A., B.Sc., exhibited a specimen of Ambly-

stoma, the metamorphosis of which had been brought about by

one month’s feeding with Ox thyroid, and drew special attention

to the precocious transition to the Amblystoma-type of pigmen-

tation during the metamorphosis.

In the absence of the Author, Mr. F. F. Larptaw, M.A., F.Z.S.,

his paper on ‘‘ Contributions to a Study of the Dragonfly Fauna

of Borneo.—Part IV. A List of the Species known to occur in

the Island,” was taken as read.

Dr. C. W. AnprREws, F.R.S., gave a réswmé of Dr. R. Broom’s

paper ‘‘On some new Therocephalian Reptiles from the Karroo

Beds of South Africa.”

The next Meeting of the Society for Scientific Business will

be held on Tuesday, May 11th, 1920, at 5.30 p.m., when the

following communications will be made :—

W. J. Daxtn, D.Sc., F.Z.8.

Fauna of Western Australia.—III. Further Contributions

to the Study of the Onychophora.

C. Forster-Coorer, M.A., F.Z.8.

Chalicotheroidea from Baluchistan.

W. T. Catman, D.Sc., F.Z.S.

Notes on Marine Wood-boring Animals.—I. The Shipworms

(Teredinide),

The Publication Committee desire to call the attention of

those who propose to offer Papers to the Society, to the great

increase in the cost of paper and printing. This will render it

necessary for the present that papers should be condensed and

be limited so far as possible to the description of new results.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOoLocicaL Socrery oF Lonpon,

ReGENt’s Park, Lonpon, N.W. 8.

May 4th, 1920.

Sek pe

eculncks

Exhibitions and Notices (continued). ‘

: age

Miss L, E. Curzseman, ¥.E.S. Exhibition of, and remarks upon, a series of lantern-slides

illustrating the life-history and habits of the Iehneumon-fly, Rhyssa persiasoria .... 194

Dr. P. Cuaummrs Mircnury, O.B.E., M.A., LL.D., D.Sc., ¥.R.S. Exhibition of, and

Yemarks upon, photographs of Abbot’s Duiker (Cephalophus spadix) .............. 194

Mr. D. Sera-Surrn, F.Z.8. Exhibition of, and remarks upon, lantern-slides showing the

display of a male Monaul Pheasant (Lophophorus impeyanus) .....0 eee rvecsces 194

Mr. L. Hoesen, M.A.,B.Se. Exhibition of, and remarks upon, a specimen of Amblystoma. 194

PAPERS.

. A Revision of the Ichneumonid Genera Labiwm and Pecilocryptus. By Rowianp E.

—

Turner and James Warerston. (Text-figures 1-11.) 1.2... .....ceeseceerecers 1

2. Description of the Larynx and Cisophagus of a Common Macaque (Macacus fascicu-

laris) exhibiting several abnormal Characters. By Dr. C. F. Sonnrag, Ch.B., F.Z.S8.,

Anatomist tothe Society... (Vext-figures 1-5.) 0... ccs cs cua devs dwatecs ce 27

3. Some Australian Opiliones. By H. R. Hoce, M.A., F.Z.S. (Plates I-III.) ........ 81

4. Revision of the English Species of Red Spider (Genera Tetranychus and Oligonychus).

PIP OTANM AYERS Dr CL Oxt-MaUees t=O.) “Fae aia'os «a yi Ha ae Weave uae eat ie se aa he ns 49

5. On the Aortic Ligament in Indian Fishes. By D. R. Buarracnarya, M.Sc., Zoological

Department, The Muir Central College, Allahabad, India, U.P. (Plates I. & II.

and Text-figures 1-5.) ...... She Ah SB DEIR IAG Ciena Bic 55-45 oatah ooR uae 61

6. On some Lizards of the Genus Chalcides. By E. G. Bourmncer, F.Z.8. (Text-

HERO ee ee I ON cet vepist cs cir ence seen rs lg. 'c: wky! win: fn jas 0" Olgcy aves Mauna aoe ee deren smear eoea ae: 77

7. Report on the Deaths in the Gardens in 1919. By Narnaniet 8. Lucas, M.B., F.Z.8.,

Hashologish to the Society. .(Wibh.4-Charts.) 2/0. «s/s as melee ces eben oe eee © 85

8. An Apodous Amia calva. By Artuur Wruuny, F.R.S., F.Z.S., McGill University,

IN IAB Sey Ame peter BN 2s ee el cara bs cla wht acgnene: xiao pea en a SEEN CASE eee etic)

9. On the External Characters of the South-American Monkeys. By R. I. Pocock,

RBS er = (ext toured a petatet fore wcaaio eiiays tel arelejel o eetatentyatane Bes Screen Mee RCI 91

10. The Comparative Anatomy of the Tongues of the Mammalia.—I. General Description

of the Tongue. By Dr. C. F. Sonntac, M.A., Ch.B., F.Z.8., Anatomist to the Society.

(Gilet trotme ssl le) ances cetera (ornlasietaiis sive x co) sitavers vatshav a alompregete ve tabsbetamemeesceelsvcncie sare. coo scl: 115

11. Notes on Hast African Lizards collected in 1915-1919, with Descriptions of a new

Genus and Species of Skink and new Subspecies of Gecko. By Arravur Loynrinesr.

(exch str enuinen lye ree auaices cesta olin tian at'+/eaia s/o sieve’) hele ecoleln ounfopehelatematedaysitsver sici'c's lee on siete 131

12. On Abnormal Features in the Peritoneum of a Raccoon. By C. F. Sonnrac, M.D.,

Ch.B., F.Z.S., Anatomist to the Society. (Text-figures 18-19.) .................. 169

13. On Abnormalities of the Abdominal Arteries of a young Panda. By C. F. Sonntag,

M.D., Ch.B., F.Z.S., Anatomist to the Society. (Text-figure 20.) ............65.. 175

14. On the External Characters of the Ratel (Mel/ivora) and the Wolverene (Gulo). By

Rel. Pocock, B-Rus., F.ZiS,)-° (Vext-figures 14-18.) eee eles eo ee ce cee ee 179

Alphabetical List of Contributors ..-..... ... SEES CoS 08 3 one eee Goa ie wht

bndiars @E IMMER NON = aie ok Seo eee eIG. etic Do Ot Clb Oo Bee eeane eae Sere xi

PLATES.

1920, Pars I. & II. (pp. 1-194).

Plate Page

Hoge: it

Tie} Australian Opiliones) 22): <2 v.00 ss ne == ae eet 31

III.

het ones a | Aortic Ligament in Indian Fishes ............ 61

NOTICE,

The ‘ Proceedings’ for the year are issued in four parts, paged consecutively,

so that the complete reference is now P. Z. 8, 1920, p.... The Distribution

is usually as follows, but on account of abnormal conditions Parts I. & II. are

issued together :—

Part I. issued in March,

ale % June.

eae b bb “3 September.

Vis rte December,

‘ Proceedings,’ 1919, Parts III. & IV. (pp. 227-499), were published

together on February 25th, 1920.

The Abstracts of the ‘Proceedings,’ Nos. 199-204, are

‘contained in this Part.

PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

AQVOLOGICAL SOCIETY

OF LONDON,

1920, pp. 195-656,

with 14 Puatres AND 143 TEx'r-FIGURES.

PRINTED FOR THE SOCIETY,

SOLD AT ITS HOUSE IN REGENT’S PARK.

LONDON:

MESSRS. LONGMANS, GREEN, AND CO,

PATERNOSTER ROW.

Las 8

OF THE

COUNCIL

ANID OPE LC Ris

OF THE

ZOOLOGICAL

SOCIETY OF LONDON.

1920.

Patron.

His Masgesty Tur Kina.

COUNCIL.

His Grace Tat Duke or Buprorp, K.G., F.R.S., President.

Tue Hon. Cecit Barina, M.A.

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

Lr.-Cot. S. Monckton Copr-

MAN, M.D., F.R.S.

Cuartes Drummonp, Esq.,

Treasurer.

Hues §. Guapsrone, Esq.,

M.A., F.R.S.E.

Sir Srpney F, Harmer, K.B.E.,

M.A., D.Sc., F.R.S., Vice-

President.

Pror. James P. -Hitu, D.Sc.,

F.R.S., Vice-President.

WitiiAM Huntsman, Esq.

Pror. Ernest W. MacBrips,

D.Se., LL.D., F.R.S.,

President.

Vice-

Cou. Sir Henry McManon,

C26. MiGs Ke@ alan:

K. G. B. Mrapr-Watpo, Esq.,

Vice-President.

P. Cuatmers MircHe.y, Esq.,

©. BE. MAS I Scr Tape

E.R.S., Secretary.

Te Haru or Onstow, O.B.E.

Major Arserr Pam.

AvpRIAN D. W. Poutocr, Esq.

His Grack THe Duke or

Ruttanp, K.G.

Tue Marquis oF Stico, F.S.A.,

Vice-President.

Masor RicHarp 8. TAytor.

A. Trevor-Barrve, Esq., M.A.

AntHony H. WinGcrIELD, Esq.,

Vice-President.

PRINCIPAL OFFICERS.

P, Coaumers Mircneut, C.B.M., M.A., D.Se., LL.D., F.B.S.,

Secretary.

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

Resident Superintendent of the Gardens.

D. Seru-Surru, Curator of Birds and Inspector of Works.

Epwarp G. BouLtencrer, Curator of Reptiles.

Miss L, KE. Curesman, F.H.S., Curator of Insects.

Prof. R. 'T. Lurper, D.Sc., M.D., Director of Prosectorium.

Dr. C. F. Sonnac, Ch.B., Anatomist.

Dr. N. 8. Lucas, M.B., Ch.B., Pathologist.

Dr. G. M. Vevers, M.R.C.S., L.R.C.P., Hon. Parasitologist.

F. Martin Duncan, F.R.M.S., Librarian.

F. W. Bonn, Accountant.

W. H. Coun, Chief Clerk.

LIST OF CONTENTS.

1920, pp. 195--656.

EXHIBITIONS AND NOTICES.

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

Menagerie during the month of April, 1920............

Mr. R. I. Pocock, F.R.S. Exhibition of, and remarks

upon, a mounted specimen of a pale variety of the

White-bearded Gnu (Connochetes albojubatus) ......

Miss Joan B. Procrer, F.Z.S. Exhibition of, and remarks

upon, a living epecimen of the tailed Batvachian

| Spelerpes fuscius Bonaparte ............2:000eecceccnreateeds

Prof. J. E. Duerpen, F.Z.S. Exhibition of, and remarks

upon, a series of lantern-slides illustrating the sexual

display and nesting habits of the Ostrich ...............

Prof. R. T. Lerrer, D.Se., M.D., F.Z.S. Exhibition of

lantern-slides illustrating the experimental trans-

mission of some Helminth infections.................,+..

Dr. P. Cuanmers Mrrcwett, C.B.E., M.A., LL.D., D.Sc.,

F.R.S. An Account, illustrated with lantern-slides,

of his recent Aeroplane Trip from Cairo to Tabora...

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

Menagerie during the month of May, 1920............

Prof. J. E. Durrpen, F.Z.S. Exhibition of, and remarks

mponwarseriesiol Ostrich CoP... aqaeereea- ocr a

Dr. R. J. Tinuyarp, M.A., F.L.S. An account of the Life-

hisbomyotsthie: Diya omilliy: rsa yaeeusci-eeeeer eis. healer

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

Menagerie during the months of June, July, August,

AMG MSCS UMSTEAD! fe. .vbic slc\veereis cedueemtqeeetnn <esiees was

Page

437

438

Dr. P. Coatmers Mircueny, C.B.E., M.A., LL.D., D.Sc.,

F.R.S. Exhibition of, and remarks upon, a Double-

tated Tazeardy 2.42 saccan te eee meee Rte mee ce one ec eee

Mr. R. I. Pococx, F.R.S., F.Z.S. Exhibition of, and

remarks upon, the skin of the groin of an exainple of

Ur age apis CULO ts. 1 ere Renner en aur cee nee eee

Mr. E. G. Boutenerr, F.Z.8. Exhibition of, and remarks

upon, living specimens of MWectwrus ........c0..0es00000

Mr. T. A. Barns, F.Z.8. An Account of his recent

Expedition through the Forests of Africa in search

of (Gorilla add! Olkcaprh Steerer ities ne

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

Menagerie during the month of October, 1920 ......

Mr. J. 'T. Cunnincuam, M.A., F.Z.S. Exhibition of, and

remarks upon, a specimen of the Leech Tvrocheta,

recently found in the Society’s Gardens..................

Mr. F. Martin Duncan, F.R.M.S., F.Z.S. Exhibition of,

and remarks upon, a series of Cinematograph Films

of Animals in the Society’s Gardens.....................

Page

655

656

LG.

18.

19.

PAPERS.

. The Life-History and Habits of two Parasites of Blow-

flies. By A. M. Aurson. With an Introduction by

Prof. H. Maxwett Lerroy, F.Z.S. (Text-figures

T= 20a \piawed: Meee b.wt Reed tl Mle eR) na

A Revision of the Nematode Family Gnathostomidae.

By H. A. Bayuis, M.A. Oxon., Assistant in the

Department of Zoology, British Museum (Natural

History), and Cxiayron Lane, M.D.Lond., Lt.-Col.

I.M.S. (vet.). (Text-figures 1-40; Plates I.—VIII.)..

Contributions to a Study of the Dragon-fly Fauna of

Borneo.—Part IV. A List of Species known to

occur in the Island. By F. F. Uarpuaw, M.A.

(Cantab.), E°Z:5. (Cext-fisures 1=4))) IR Be

On some new Therocephalian Reptiles from the Karroo

Beds of South Africa. By R. Broom, M.D., D.Sc.,

HERES OUMeEZES en (lext-tioures) 19") ee pe saeeene ee

Chalicotheroidea from Baluchistan. By C, ForsrEr-

Coorer, M.A., F.Z.8., Superintendent of the

University Museum of Zoology, Cambridge. (Plate

The zm “ADepsiem thei etsy a(S) Wapie meme et aanat conabghnoaseeeoor

. Fauna of Western Australia,—III. Further Contri-

butions to the Study of the Onychophora. The

Anatomy and Systematic Position of West Australian

Peripatoides, with an account of certain histological

details of general importance in the study of Peri-

patus. By Wm. J. Dakin, DSc. EZ.8., Hus:

Professor of Biology, University of Western Australia.

(Pilates T= Vidic... Acar vawntanne Dt: Re MRO Rea eee!

. Notes on Marine Wood-boring Animals.—I. The Ship-

worms (Teredinidae). By W. IT. Canman, D.Sc.

(Mexteiiotumes! TW) i. ali wc ame prencen eget cscs nrnsectete

. Report on Entozoa collected from Animals which died

in the Zoological Gardens of London during eight

months of 1919-20. By G. M. Vevers, M.R.CS.,

L.R.C.P., F.Z.8., Beit Memorial Research Fellow,

Demonstrator in Helminthology at London School of

Tropical Medicine, and Hon. Parasitologist to the

Zoologicalssocietyeor Hondo... ite. ceasccee esse sess se

Page

195

311

343

367

391

ie)

a |

Sills

(Su)

isu)

. Ona Collection of Tailless Batrachians from Hast Africa

made by Mr. A. Loveridge in the years 1914-1919.

By Miss Joan B. Procrer, F.Z.8. (Text-figures 1-4.)

On the Type-specimen of “una holsti Boulenger. By

Miss Joan B. Procrmr, F.Z.5. (Text-figure 5.) ......

On the External and Cranial Characters of the Huropean

Badger (JMeles) and of the American Badger ( Taaidea).

By R. I. Pocock, F.R.S. (Text-figures 19-25.) ......

. Onsome Results of Ligaturing the Anterior Abdominal

Vein in the Indian Toad (Lufo stomaticus Liitken).

By W. N. F. Woopianp, D.S8e.(Lond.), F.Z.S.,

Indian Educational Service, Senior Professor of

Zoology Muir Central College, Allahabad, U.P.,

India.<qd(@exteiigureydt) ideas sae. Hy, M2, Re ES

. Some Observations on the Structure and Life- History

of the Common Nematode of the Dogfish (Scylliwm

canicula). By J. H. Luoyp, M.Sc., F.Z.8S., Assistant

Lecturer and Demonstrator in Zoology at University

College; Cardiif, -(lext-tiguras 13n)\ets..o. ee

- On the Sexual Phase in certain Indian Naidide (Oligo-

cheta). By Haru Ram Meura, M.Sc., Professor of

Zoology, Hindu University, Benares. (Text-

figures 1-3.)

29. Observations on the Flight of Flying-Fishes. By H.

H. Hanky, M.A., Sc.D., Agra, India. (‘Text-figures

1g) een mater mre Peni. iat Mi Ou een ae aa eee

. Observations on the Life-History, Biology, and Genetics

of the Lady-bird Beetle, Adalia bipunctata Mulsant.

By Onréra A. Merrirr Hawkes, M.Se.(Birm.), B.Sc.

(Tiong 2 ae RRhe 2 ge cde ee eee kiero

Filariid Worms from Mammals and Birds in the

Society’s Gardens, 1914-1915. By C. L. Boutenezr,

M.A., D.Se., F.Z.8., Professor of Zoology, University

of the Punjab, Lahore. (Text-figures 1-12.) .. ......

. The Fauna of the African Lakes; a Study in Com-

parative Limnology with special reference to Tangan-

yika. By Wm. A. Cunnineton, M.A., Ph.D., F.Z:S.

(Lexte mounds: 12 )ing4.....iec cee eee sseriaesceeee-els nok. Renee

3. Descriptions of the Adult, Larval, and Pupal Stages of

a New Mosquito from Lord Howe Island, 8. Pacific.

By Henry F. Carrer, Liverpool School of Tropical

Medreme: ( Vext-founeswdGan) pesca taiacsseen eens

44]

449

457

467

475

491

507

Vil

Page

34. The Life-History and Habits of the Yellow Dung-Fly

(Scatophaga stercoraria) : a possible Blaw-Fly Check.

By G. 8. Correrent. With a Preface by Prof.

Maxwet. Lerroy, F.Z.8. (Text-figures 1-14.) ...... 629

35. Remarks on the Respiratory Movements of Nectwrus

and Cryptobranchus. By A. Witnny, M.A., D.Sc.,

LG Eve It ae ASS re ae oes Sc eens ee ee 649

Alphabeveal list ot Contributors...) eeeee eee eee Vill

prndaxero tei SGTAtTOMS 4 oe c.20 6 .Ninsosacoscneea demeanor aes xiii

TETAS KESTER 4, ANE EG aA rk SNE SIR SUT UR eye Dats uk pane te RV

ALPHA BHA RCA, List

OF TIE

CON Resa a Ouk’S,

With References to the several Articles contributed by each.

(1920, pp. 195-656.)

Aurtson, A. M.

The Life- History and Habits of two Parasites of Blow-

Flies. With an Introduction by Prof. H. Maxwe.i

iEPRoY, Hi4is. “((Chexttiounesil 2 Oecd ..sncgeee cence ae:

Bayuis, H. A., M.A. (Oxon.), Assistant in the Department

of Zoology, British Museum (Natural History), and

Lang, Ciayron, M.D.(Lond.), Lt.-Col. I.M.S. (ret.).

A Revision of the Nematode Family Gnathostomide.

(Platesm- Valle ;) Mest higires ta 08) Bere. ee sen pean

Bovu.encer, C. L., M.A., D.Sce., F.Z.S., Professor of Zoology,

University of the Punjab, Lahore.

Filariid Worms from Mammals and Birds in the

Society’s Gardens, 1914-15. (Text-figures 1-12.).........

Bouxencer, EH. G., F.Z.8., Curator of Reptiles.

Exhibition of, and remarks upon, living specimens of

Necturus ........ RM pee UL yh 2. 8 Ue a a a ee a

Broom, R., M.D., D.Sc., F.R.8., C.M.Z.S.

On some new Therocephalian Reptiles from the Karroo

Beds of South Africa. (Text-figures Oh cane troansenes

Page

195

49]

Cauman, W. T., D.Sc.

Notes on Marine Wood-boring Animals.—I. The Ship-

Worms (Teredinide). (Text-figures 1-11.) ...............

Carrer, H. F., Liverpool School of Tropical Medicine.

Descriptions of the Adult, Larval, and. Pupal Stages

of a New Mosquito from Lord Howe Island, South

aveitiee ua @ excites) U3.) 4. oc. 5e ee oat:

CorTrerELt, G, 8.

The Life-History and Habits of the Yellow Dung-Fly

(Scatophaga stercoraria); a possible Blow-Fly Check.

With a Preface by Prof. H. Maxwett Lerroy, F.Z.S

Beret tienes I A Na ooh aactectcisviovyni neq oe eRe ae

Cunninenam, J. T., M.A., F.Z.8.

Exhibition of, and remarks upon, a specimen of the

Leech Z'rocheta, recently found in the Society’s Gardens.

Cunnineton, Wm. A., M.A., Ph.D., F.Z.S.

The Fauna of the African Lakes; a Study in Com-

parative Limnology with special reference to Tanganyika.

(Resethivoumes: TRO ESS 020) Ly TS SAE Rie)

Daxty, Wm. J., D.Sc., F.Z.5., F.L.S., Professor of Biology,

University of Western Australia.

Fauna of Western Austvalia.—IIJ. Further Contri-

butions to the Study of the Onychophora. The Anatomy

and Systematic Position of West Australian Peripatoides,

with an account of certain histological details of general

importance in the study of Peripatus. (Plates I.-V.)...

Duerpen, J. E., F.Z8.

Exhibition of, and remarks upon, a series of lantern-

slides illustrating the sexual display aud nesting habits

GUE SNS OME CIN Se oe Re lak ou koe eae a ane

Exhibition of, and remarks upon, a series of Ostrich eggs

Proc. Zoou. Soc.—1920. b

Page

629

656

507

439

Duncan, F. Martin, F.R.MLS., F.Z.S., Librarian to the

Society,

Exhibition of, and remarks upon, a series of Cinemato-

gvaph Films of Animals in the Society’s Gardens .........

Forsver-Coorer, C., M.A., F.Z.8., Superintendent of the

University Museum of Zoology, Cambridge.

Chalicotheroidea from Baluchistan. (Plate I.; Text-

HUMES ESL) a. okorieine tec see REE oracle oe bee tee

Hankin, KE. H., M.A., Sc.D., Agra, India.

Observations on the Flight of Flying-Fishes. (Text-

HUGE 12), (Sith dcrda scion Ce eee ok Lo: 4 ee

Hawkes, OnkrA A. Merrirr, M.Sc.(Birm.), B.Sc.(Lond.).

Observations on the Life-History, Biology, and

Genetics of the Lady-bird Beetle, Adalia lipunctata

Mralisamt: eiccuisds aig ncticamet Jee aa Renee ie oo oe agate eae

Laipiaw, F. F., M.A.(Cantab.), F.Z.S.

Contributions to a Study of the Dragonfly Fauna of

Borneo.—Part IV. A List of Species known to occur in

the dislands) ((Text-fisunes tl —49) oe ieen ces: 4-6... cceeeee

Lane, Crayton, M.D.(Lond.), Lt.-Col. I.M.S.(vet.).

See Bayuis, H. A.

Lerroy, H. Maxwe tt, F.Z.8.

See Autson, A. M.

See Correreny, G. 8.

Lriprr, Re Ty, Ds): M.DEERZS:

Exhibition of lantern-slides illustrating the experi-

mental transmission of some Helminth infections.........

656

467

AT5

Ball

Lioyp, J. H., M.Sc., F.Z.S., @Assistant ‘Lecturer and

Demonstrator in Zoology at University College,

Cardiff.

Some Observations on the Structure and Life-History

of the Common Nematode of the Dogfish (Sceylliwm

canicula). | Ckext-—hounesil—3: )\s:AJsetee ae. ena el:

Meura, Haru Ram, M.8c., Professor of Zoology, Hindu

University, Benares.

On the Sexual Phase in certain Indian Naidide

(Oligochzeta)\ ((lext-fisumes 123°) 0 ease te

Mircuenn, P. CHaumers, C.B.E., M.A., D.Se., LL.D.,

F.R.S., Secretary to the Society.

Report on the Additions to the Society’s Menagerie

dunner the monthror “Agoril, UO 20 ne cnc eacere earn

An Account, illustrated by lantern-slides, of his recent

Aeroplane Trip from Cairo to Tabora ...............0....050:

Report on the Additions to the Society’s Menagerie

dunes the momthyot Mian. NO2O) se. 2. a aang

Report on the Additions to the Society’s Menagerie

during the months of June, July, August, and September,

Exhibition of, and remarks upon, a Double-tailed

OMAR WITT ak eA ee Oe ee ce MRE ARGC Us Ra ar er

Report on the Additions to the Society’s Menagerie

Huninerune mmlontbnon, October, 1920 soe ce eaeate scm acs.

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

Resident Superintendent of the Gardens.

On the External and Cranial Characters of the

European Badger (Meles) and of the American Badger

(Tamidea). (Text-figures 19-25.)..........:0e:ceesee eset eeeees

Page

449

423

X11

Page

Pocock, R. T. (cont.).

Exhibition of, and remarks upon, a mounted specimen

of a pale variety of the White-bearded Gnu (Connochetes

CLO OI RUDGE AUS): aM ti ce Aes oh aed ee gubtesrrsadG hates 437

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

of an example of Tragelaphus bustoni.:..........0..0.0.0000- 659

Procter, Miss Joan B., F.Z.S.

On a Collection of Tailless Batrachians from East

Africa made by Mr. A. Loveridge in the years 1914-1919.

(Bext=toutesy (EA Is. Meee enemy trees ahi Gee Cae 41]

On the Type-specimen of Rana holsti Boulenger.

(Mexb=fie ure 0)": 1 Syke ee ee RRR 2 ode eee 42]

Exhibition of, and remarks upon, a living specimen of

the tailed Batrachian Spelerpes fuscws Bonaparte ......... 437

TintyarD, R. J., D-Sc., MAL Wiss:

An account of the Life-History of the Dragonfly...... 439

Vevers, G. M., M.R.C:S., L.R.C.P., F.Z.8., Hon. Parasito-

logist to the Society.

Report on Entozoa collected from Animals which died

in the Zoological Gardens, London, during eight months

Of “POTD 219208 Fen Sets ee ee ae salen titee AN st 405

Winy, A., M.A,, D.Se., F.R.S., E.Z.8.

Remarks on the Respiratory Movements of Wecturus

ETUC 7a pLOUNETLCH US» Ue) Ra \oreme tee ame e ee sae Ae. or ie 649

Wooptann, W. N. F., D.Sc.(Lond.), F.Z.8., Indian Educa-

tional Service, Senior Professor of Zoology, Muir

Central College, Allahabad, U.P., India.

On some Results of Ligaturing the Anterior Abdominal

Vein in the Indian Toad (Bufo stomaticus Liitken).

(Mex ttre ure de). acces ah cee meee ce eee cele © 44)

INDEX OF ILLUSTRATIONS.

Acanthocheilonema diacantha, Figs. 3, 4,

pp. 499, 496.

-—— gracile, Figs. 5, 6, 7, pp. 497, 498-

African Lakes, Table to illustrate rich- |

ness of Fauna, Fig. 2, p. 594

Alopecopsis atavus, Figs. 4, 6, pp. 347,

349, 550.

Alysia manducator, Figs. 1-12, pp. 199,

201-203, 205, 206, 208, 209, 212.

Arthroleptis stenodactylus, Big. 1, p. 414.

Branchiodrilus hortensis, Figs. 1, 3,

pp. 459, 463.

Bufo stomaticus, Fig. 1, p. 445.

Chalicotheroidea, Figs. 5-7, pp. 864, 360.

Copera atomaria, Fig. 4, p. 304.

Curve showing growth of knowledge of |

Tanganyika fauna, Fig. 1, p. 593.

Diplotriena diuce, Figs. 9, 10, pp. 501,

502.

—— flabellata, Fig. 11, p. 503.

— tricuspis, Fie. 8, p. 500.

Echinocephalus multidentatus, Pls. VI.-

VIIL., p. 245; Figs. 33-56, pp. 286,

287, 290.

southwelli, Pls. 1V.-V., p. 245;

Figs. 30-32, pp. 282-284.

Figs, 25-29, pp. 279--281.

uneinatus, Fig. 24, p. 276.

Proc. Zoou. Soc.—-1920.

spinosissimus, Pl. 1V., p. 245; |

Filaria aramidis, Fig. 12, p. 504.

—— subcutanea, Figs. 1, 2, pp. 492, 493.

Flying-fish, Flight of, Figs. 1, 2, pp.469,

fo.

Gnathostoma spinigerum, Pls. VIL,

VIIL., p. 245; Figs. 37-40, pp. 293-

295.

Ictidosuchus longiceps, Bigs. 1-3, pp.844—

346.

Megaliaalus loveridegii, Big. 4, p. 418.

Meles metes, Wigs. 19-25, pp. 425, 427,

429, 451, 432, 434, 435.

Melittobia acasta, Wig. 20, p. 240.

Moschorhinus kitchingi, Figs. 7, 8,

pp. 351, 303.

Nais pectinata var. inequalis, Figs. 1, 2,

pp. 459, 460.

Nasonia brevicornis, Figs. 13-20, pp. 217—

220, 222, 225, 226, 240:

Peripatoides, Pls. I.-V., p. 367.

Phyllotillon naricus, Pl. 1., p. 357 ;

Figs. 1, 2, pp. 858, 360.

Proleptus scillicola, Figs. 1-3, pp. 451,

452, 454.

Rana holsti, Fig. 5, p. 421.

Rappia platyrhinus, Fig. 3, p. 416.

puncticulata, Fig. 2, p.415.

X1V

Rhinocypha moultont, Figs, 1, 2, pp. 329,

330,

Scatophaga stercorwia, Figs. 1-14, pp.

631-645.

Schizotherium ? pilgrimi, Pl. 1., p. 857 ;

Figs. 3, 4, pp. 862, 3538,

Spiroxys contorta, Pls. 1.-II., p. 245;

Figs. 1-3, p. 250.

gangetica, P1,I1., p. 245; Figs. 4-6,

p. 202.

Tanganyika fauna,Curve showing growth

of knowledge of, Fig. 1, p. 593.

Tangua anomala, Pl. IV., p. 245;

Figs. 16-20, pp. 265-267.

INDEX OF ILLUSTRATIONS.

Tanqua diadema, Pls. I11.-1V., p. 245;

Figs. 21-23, p. 269.

—— tiara, Pl. 1Il., p. 245; Bigs, 7-15,

pp. 29-261, 263, 264.

Tawidea americana, Figs. 19, 20, 22-25,

pp. 425, 427, 481, 482, 454, 435,

Teinobasis hirbyi, Fig. 4, p. 387.

Teredo mannii, Figs. 2, 3, pp. 395, 396.

navalis, Big. 1, p. 393.

Whaitsta platyceps, Fig. 9, p. 354.

Aylotrya australis, Figs. 6-8, pp. 899-

401.

capensis, Figs. 9-11, pp. 401-403.

saulit, Figs, 4, 5, p. 399.

INDEX.

1920.—Pages 195-656.

| New names in clarendon type.

Systematic references in italics.

(z.8.L.) indicates additions to the Society’s Menagerie. |

Acanthocephala, 410.

Acanthocheilonema diacantha, 494.

gracile, 496,

Aciagrion borneense, 336.

Adalia bipunctata, 475.

Aischnide, 312.

Aischnine, 312.

Athertide, 582.

Aithriamanta gracilis, 325.

Agriocnemis femina, 336.

rubescens, 336.

Agrion, Legion, 335.

Agrionoptera insignis, 321.

——- sexlincata, 321.

Alluroidide, 574.

Alopecopsis atavus, gen. et sp. nov.,

347.

Alysia manducator, 199.

Ammotragus lervia (4%, 8. u.), 437.

Amphieschna grubauert, 314.

peranmpla, 314,

Amphicnemis louise, 336.

madeleneé, 336.

martini, 330.

remiger, 336.

wailacet, 336.

Ampullariide, 543.

Anubantide, 529.

Anacieschna jaspidea, 316.

Anastillorhina augur, 230.

Anax gutiatus, 316.

Ancylide, 545,

Ancyracanthus, 309.

Anisoptera, 312.

Ara auricapilla (4,8. L.), 694.

Argulus incisis, 564.

Arthroleptis minutus, 414.

stenodactylus, 414.

wahlbergr, 414.

—— whytii, 414.

Anthropopithecus troglodytes (4. Ss.

437.

Anthrothreptes collaris (4.8. u.), 654.

Ascaris spherocephala, 271.

Atyid@, 559.

Azunua australis, 317,

vittigera, 318.

Bdelloida, 580.

Bison americanus (2,8. L.), 659,

Bosminide, 568.

Boulengerina stormsi, 520.

L.)

Brachydiplax chalybea chalybea, 322.

Brachygonia oculata, 322.

-—- ophelia, 32

XVI

Brachyura, 556.

Branchiodrilus hortensis, 468.

Branchiopoda, 571.

Branchiura, 568.

Breviceps mossambicus, 420.

Bubo carens, 420.

lacteus (4.8. u.), 437.

Bufo reqularis, 420.

stomaticus, 441.

Bungarus fasciatus (4.8. u.), 65+.

Burmagomphus vermiculatus, 317.

Caconeura collaris, 339.

—— dorsalis, 339.

gracillima, 340.

hoset, 339.

—— hyperythra, 889.

—— interrupta, 389.

lansbergi, 339.

moultoni, d40.

—— peramenda, 339.

verticalis, do9.

Calliphora erythrocephala, 196, 198,

230.

vomituria, 198.

Callosciurus notatus (z.8.L.), 654.

vittatus (Z. 8. L.), 695.

Calopterygine, 3205.

Caracinia harterti, 325.

Cancroma zeledont (4. 8. L.), 654.

Canis gubatus (z. 8.u.), 656.

- thous (4. 8. L.), 656.

Cariama cristata (4. 8. b.), 655.

Carposhaga finschi (4. 8. u.), 656.

muellert (4. 8. L.), 656.

——- pinon (4. 8. L.), 656.

Cassina senegalensis, 419.

Centropagide, 559.

Cephalophus dorsalis (Z. 8. u.), 656.

—— marwelli (4.8. u.), 656.

Ceriagrion bellona, 335.

cerinorubellun, 390.

Cervus xanthopygius (4. 8. u.), 653,

Cestoda, 408, 585.

Chetonotus formosus, 582.

pusillus, 582.

Chamepelia pallescens (4. 8. L.), 698.

INDEX.

Chamepelia rufipennis (4. 8. L.), 698,

Characinide, 524.

Chiromantis petersvi, 413.

—— werampelina, 413.

Chlorogomphine, 312.

Chonopeltis inermis, 564.

Chrysomia macellaria, 230.

Chrysomitris mexicanus (4. 8. L.), 693.

Chrysotis virenticeps (4. 8. L.), 699,

Chydoride, 568.

Cichlide, 526.

Ciliata, 590.

Cissolopha sanblasiana (2. s. u.), 653.

Cladocera, 567.

Clarotes laticeps, 488.

Clupeide, 524.

Coccinella variabilis, 487.

Coelenterata, 585.

Celiccia borneensis, 334.

campiont, 333.

—— flavostriata, 338.

— macrostigid, 333.

—- membranipes, 339.

— nigrohamata, 333.

—— octogesima, 333.

Colinus texanus (4. 8. L.), 658.

Colobus abyssinicus (4. S. L.), 654.

Connochetes albojubatus, 437.

Copera atomaria, 334.

Corduliine, 317.

Cracticus cassicus (4. 8. L.), 696.

Cratilla lineata, 321.

metallica, 321.

Crioceris asparagi, 486.

Crocodilus cataphractus, 519.

niloticus, 919.

—— stamensis (7%. 8. L.), 439.

Crocothemis servilia, 323.

Cryptobranchus, 649.

Cycloderma frenatum, 521.

Cyclopide, 560.

Cyneélurus gubatus (Z. 8. L.), 438.

Cynomyia cadaverinia, 230.

Cypride, 569.

Cyprinide, 525.

Cyprinodontide, 526.

Cyrenide, d01.

Cytheride, 56d.

INDEX.

Daphnide, 567.

Dendrolagus ursinus (4. 8. L.), 036.

Devadatta argyroides, 332.

Dicheilonema horridum, 499.

Diplacodes trivialis, 323.

Diplotrizna diuce, sp. n., 001.

flabellata, 502.

tricuspis, 499.

Dipsadomorphus dendrophilus (4, 8. u.),

654.

Disparoneura analis, 340.

-—— gurantiaca, d40.

Dolops ranarum, 564.

Dorylaimus macrolaimus, 577.

Drepanosticta rufostigma, 340.

Dyromys nitidula (2. 8. .), 653.

Dysphea limbata, 327.

lugens, 327.

Echinocephalus, 273, 27d.

— multidentatus, sp. n., 285, |

288, 289.

—— southwelli, sp. n., 283, 285, 288.

spinosissimus, 277, 283, 288.

striatus, 289.

-—— uncinatus, 275, 277, 288.

Elaphocephalus, 306.

Hlephas maxinius (4. 8. u.), bo.

Kpallagine, 326.

Ergasilide, 560.

Eucopepoda, 559.

Felis tigris (z. 8. u.), 694.

Filaria aramidis, sp. n., 503.

subcutanea, 492.

Flying-Fishes, 467.

Francolinus spilogaster (4. S. L.), 409.

Gabianus pacificus (Z. 8. L.), 653.

Gastrotricha, 581. 3

Glossoscolecide, 574.

Glypholycus bicolor, 520.

Gnathostoma, 291, 292.

—— accipitri, 305.

gracile, 300, 301, 303, 304,

hispiduni, 298, 300, 303, 304.

xvil

Gnathostoma horridum, 300, 303,

304.

—— pelecani, 301.

shipleyi, 301.

spinigerum, 293, 298, 302, 304.

turgidum, 301, 305, 304.

Gnathostomide, 245.

Guathostomine, 255, 256.

Gomphidia karschi, 317.

—— maclachlani, 317.

Gomphine, 317.

Gossea pauciseta, 582.

Grayia ornata, 520.

-—— tholloni, 520.

Gynnolemata, 539.

Gynacantha basiguttata, 315.

bayadera, 316.

demeter, 315.

—— dohrni, 315.

—— hyalina, 316.

machlachlani, 316.

Gyrator hernaphroditus, 584.

Harpacticide, 559.

Helieschna crassa, 318.

—— idé, 313.

stmplicia, 314.

uninervilata, 3)4.

Hemicordulia assimilis, 317.

Hemistoma ( Alaria) alatum, 438.

Hemisus marmoratum, 420.

Herpeton tentaculatum (4.8. .), 439.

Heterogomphus icterops, 317.

sumatranus, 317.

Hippopotamus amphibius (2. s.t.), 653.

Hirudinea, 575.

Hydrachnida, 572.

Hydrobasileus croceus, 324.

Hydrobiide, 543.

Hydrocherus hydrocherus (4.8. L.), 438.

Hydrophantide, 572.

Hygrobatide, 572.

Hyleothemis clementia, 320.

Hylambates bocagii, 419.

Johnstoni, 420.

Hylobates lar (4. 8. u.), 487.

Hylocichla alicie (2. 8.u.), 653.

Hypotenidiu striata (4.8. u.), 487.

Xvill

Lchthydiwm macrurum, 582.

Ictidosuchus longiceps, sp.

343.

Tetinus acutus, 317.

—— decoratus, 317.

—— melenops, 317.

Idionyx dohrni, 319.

Jagoria buhri, 313.

-— modiglianti, 312.

peciloptera, 313.

Lamprocolius australis (%. 8.u.), 698.

Lamprocorax sulaensis (a. 8. u.), 656.

Larus occidentalis (7%, 3. 1.), 658.

Lathrecista asiatica, 321

Lepidoderma hystrix, 582.

sguamatum, 582.

Lepidosirenide, 524.

Leptogomphus semperi, 317.

williamsoni, 317.

Lerneide, 560.

Lestes wallace, 340.

Libellagine, 328.

Libellutide, 317.

Libelluline, 319.

Limneide, 545.

Lineschna polli, 312.

Lucilia cesar, 198, 230.

—— serteata, 198, 230.

Lybius torquatus (%. 8. 1), 439,

Lyriothemis biappendicuiata, 320.

- cleis, 320.

Mabuia quingueteniata (2.8. t.), 437

Macrogomphus albarde, 317.

chelifer, 317.

decemlineatus, 317.

quadratus, 317.

Macronia barneensis, 318.

-—— cincta, 318.

—— cemgulata, 318.

—— euterpe, 318.

—— gersteckeri, 318.

—— westwoodi, 318.

Macromidia fulva, 319.

Macropus brunti (2.8. u.), 656,

Maerothricide, 568.

INDEX.

Macrura, 554.

Mastacembelide, 529.

Mastigophora, 589.

Mutronoides cyancipennis, 326.

Megalivalus loveridgti, 418.

Megapodagrion, Legion, 382.

Megascolecide, 574.

Meles, 423, 483.

Melittobia acasta, 231.

Metaphya micans, 519.

Micromerus aurantiacus, 331.

hyalinus, 332.

seniopacus, ddl.

—-- sticticus, 332.

Momotus venezuele (z.8.u.), 658.

Mongos caffer (4. 8. u.), 655.

Monhystera filleborni, 477.

similis, D77.

—— vulgaris, 577.

Mononchus macrostoma, 577.

—— tenuis, D177.

Monostoma pumilo, 488.

Mormyride, 5 ae,

Moropus elatus, 3

307, 309.

Moschorhinus kitchingi, gen.

sp. noy., Jol.

Musca domestica, 198,

Mutelide, 552.

230.

Naia bungarus (Zz. 8, u.). 694,

tripudians (4.8. L.), 6B4.

Naidomorpha, 574.

Nais pectinata var. inequalis, 458.

Nannophya pygmea, 322.

Nasonia brevicornis, 216.

Necturus, 649.

Nemathelmia, 405.

Nematoda, 408, 576.

Nesoxenia lineata, 320.

Neurobasis chinensis, 320.

Neurothemis disparilis, 525,

—-. fluctuans, 328.

—— terminata, 323.

Oda dohrni, 520.

Oligocheta, 573.

Onychargia atrocyaind, 3356

Onychothermis culminicola celebensis,

324,

—— culininicola culniinicola, 324.

Orchithemis pruinans, 320.

—— pulcherrima, 320.

vanthosoma, 320.

Orogomphus dyak, 312.

splendidus, 312.

Ortalis macealli (2.8. L.), 653.

Orthetrum chrysis, 322.

—— glaucum, 321.

—— pruinosum clelia, 321.

sabina, 321.

—— testaceum, 322.

Oryx leucoryx (2.8. 1.), 438.

Ostracoda, 564.

Palemonide, 55d.

Pantala flavescens, 325.

Papio anubis (2.8. u.), 654.

Pelomedusa galeata, 521.

Pentastomida, 571.

Pericnemis stictica, 337.

Peripatoides gilesii, 367.

—— leuckarti var. occidentalis, 367.

—— occidentalis, 507.

woodward, 367.

Peripatus, 367.

Perodicticus potto (z.8. u.), 655.

Phorinia grenlandica, 198.

regina, 230.

Phrynobatrachus boulengeri, 413.

natalensis, 413.

—— ranoides, 413.

Phrynomantis bifasciata, 420.

Phylactolemata, 539.

Phyllotillon naricus, 357,

Pipilo crissalis (2.8. u.), 653.

Pitangus rufipennis (z. 8. L.), 680.

Planaria tanganyika, 588.

Planorbide, 545.

Platycnemis, Legion, 535.

Platyhelmia, 405.

Platysticta, Legion, 340.

Ploima, 578.

Podolestes chrysopus, 332.

—— orientalis, 332.

Pollenta stygia, 220.

Polypteride, 524.

INDEX. X1X

Porifera, 586.

Pornothemis serrata, 820.

Potamarcha obscura, 321.

Potamonide, 557.

Proleptus scillicola, 449.

Protoneura, Legion, 338.

Protosticta kinabaluensis, 340.

versicolor, 340.

Protozoa, 588, 589.

Pseudagrion inicrocephalum, 335.

—— pruinosum, 330.

Pseudagrionoptera diotima, 323.

Pseudophea basalis, 326.

impar inequipar, 827.

--—— subcostalis, 326.

— subnodalis, 326.

—— tricolor, 326.

Pycnosoma rufifacies, 230.

—— varipes, 230.

Physaloptera constricta, 272.

Rana delalandii, 412.

holst?, 421.

—— mascareniensis, 412.

nutti, 412.

ornata, 413,

——- oxyrhynchus, 412,

Raphisma inerniis, 328.

Rappia argus, 417.

—-— cinctiventris, 417.

——: fulvovittata, 417.

—— granulata, 415,

—— marmorata, 417.

—— platyrhinus, sp. n., 416.

puncticulata, 415,

Rhinagrion borneense, 332.

elopure, 333.

Rhea americana (2. 8. t.), 439.

Rhinochetus jubatus (2. s. t.), 439.

Rhinoneura villosipes, 331.

Rhizopoda, 58%.

Rhizota, 580.

Rhyothemis aterrima, 324.

obsolescens, 325.

pygmea, 324.

—— phyllis phyllis, 324.

—— rufa, 323.

— triangularis, 325.

Rhinocypha biseriata, 328,

XX *

Rhinocypha cucullata, 329.

eximia, 329.

— humeralis, 329.

—— kharschi, 328.

-—— moultoni, 328.

——. stygia, 328.

sp. A, 329.

, sp. B, 330.

Rotifera, 578.

Sarcophaga aurifrons, 230.

Scardafella inca (4. 8. L.), 653.

Scatophaga stercoraria, 629.

Schilbe mystus, 438.

Schizotherium pilgrimi, sp. n., 362.

Scyllium canicula, 449.

Serranide, 526.

Sidide, 567.

Sieboldius japonicus, 317.

Siluride, 525.

Spelerpes fuscus, 437.

Spheriide, 551.

Spinus pinus (z. 8. L.), 658.

Spiroxyine, subfam. n., 247.

Spirorys, 248.

contorta, 249, 251.

gangetica, sp. n., 201, 253.

Spongillide, 587.

Stenagrion dubium, 338.

Stenostoma gilvum, 588.

Zeucops, 583.

stuhlimanni, 588.

Sternotherus derbianus, 521.

—— nigricans, #21.

sinuatus, 521.

Struthio molybdophanes (2. 8. L.), 439.

Tanqua, 256, 258.

anomala, 268.

diadema, 268, 271.

—- tiara, 259, 264.

Tardiyrada, 573.

Taxidea, 423, 433.

Teinobasis kirbyi, 337.

rajah, 338.

Teredo, 392.

——— mannit, 39d.

navalis, 392.

norvagica, d9-+.

INDEX.

Tetracanthagyna brunnea, 315.

degorst, 315.

plagiata, 314.

waterhousei, 315.

Tetrathemis flavescens, 319.

irreqularis hyalina, 319.

Thalarctos maritimus (x. Ss. u.), 655.

Tholymis tillarga, 325.

Tiaride, 544.

Tilapia nilotica, 488.

Tiphobiide, 544.

Tramea limbata, 324.

Trematoda, 407, 584.

Trichoglossus rosenbergi (z. 8. L.), 656.

Trilobus graciloides, S17.

Trithemus aurora, 323.

Jestiva, 324.

Trocheta, 656.

Tropidonotus olivaceus, 520.

Turbellaria, 5895.

Turtur decipiens (2. s. u.), 439.

isabellina (2. s. L.), 439.

roseigriseus (7%, 3. L.), 439.

Tyriobapta kukenthali, 322.

laidlawi, 322.

= torrida, 322.

Unionide, 551.

Urolestes metanoleucus (z. 8. L.), 439.

Urothemis signata insignata, 325,

Vestalis amena, 326.

berylle, 526.

Vipera lebetina (2. s. u.), 655.

russelli (z. Ss, L.), 654.

Vireo olivacea (2. 8. L.), 653.

Viviparide, 543.

Vortex quadridens, 584.

Whaitsia platyceps, 354.

Xenopus levis, 522.

muellert, 522.

Aylotrya, 397.

—— australis, sp. n., 397.

—— cabensis, sp. n., 402.

Zygonyx iris, 324.

Zygopterda, 825.

Zyeomma petiolatum, 325.

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EXHIBITIONS AND NOTICES.

The SucruTary. Report on the Additions to the Society’s Menagerie during the month

of April, 1920 ...... CO IAGIN OOH AY Bnd aaGe i bod Wala Matag Mens Sos eo oo aay on LS oc

Mr. R. I. Pococx, F.R.S8. Exhibition of, and remarks upon, a mounted specimen of a

_ pale variety of the White-bearded Gnu (Connochetes albojubatus) .....-.-.+.+..4.

Miss Joan B. Procter, F.Z.8. Exhibition of, and remarks upon, a living specimen of the

tailed Batrachian Spelerpes fuscus Bonaparte, born May 8th, 1920 ..............

Prof. J. E. Duzrpey, F.Z.S. Exhibition of, and remarks upon, a series of lantern-slides

illustrating the sexual display and nesting-habits of the Ostrich ................

Prof. R. T. Lurper, F.Z.8. Exhibition of lantern-slides illustrating the Experimental

transmission of some Helminth injections ................6 a Sailalsiuies ei stateiedl res kei

Dr. P. Cuatmers Mitcunut, 0.B.E., M.A., LL.D., D.Se., K.R.S. An Account, illustrated

with lantern-slides, of his recent Aeroplane Trip from Cairo to Tabora .,.......-

The Szcretary. Report on the Additions to the Society's Menagerie during the month of

Miietry, LOD O iiss) oes atarteyes aietare talon ets) oe ate ties laity tare ciatinto tte Cie Were/ ei wit ots Joe states eee

Prof. J. E. DuErpsn, B.ZS. Exhibition of, and remarks upon, a series of Ostrich eggs ..

Dr. R. J. TILLYARD, M.A., F.L.S. An account of the Life-history of the Dragonfly ....

Page

437

487

438

439

Contents continued on page 3 of Wrapper.

PROCEEDINGS

OF TIE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF TIIE

ZOOLOGICAL SOCIETY OF LONDON

PAPERS.

15. The Life-History and Habits of two Parasites of Blow-

flies. By A. M. Atrson. With an Introduction by

Prot. H. Maxwetu Lerroy, F.Z.S.*

[Received March 29, 1920: Read April 13, 1920.]

(Text-figures 1-20.)

INDEX.

Page

AGS OMG NGCOUMbOLUneIStaNt OLE Wiotk: » 0. ...ce eel ete eneEne nn nLOG

Breeding Methods ......... sesbostanel Ss}

Life-History and Habits ae Bia sith siaovdunaeen ans, end Eigse booties 199

Lite-History and Habits of Nasonia brevicornis Ashm.and Hosts ... 216

Super-parasitism, or Accidental Secondary Parasitism .................. 230

Refrigerating Experiment ............... eta ey cticanl hZ4O

Conclusions upon the iBlaorkornle Importance ee ae Daiiucntor

JEL Aa BVNGL Ty (DRBDIAD AAS HENSIN ees Se odd seston vabanasericantcs cotoonaboceosanbe © ell

PS) DUS AUTEN orelgh a iehodnatce<c oat Mecca ea RIA HE MERREEN Ce Scio as oGdwsv olan hp esweene week be

Aceeraedements) SCA SC nan Ae EERE? oP cicindl dance cis an aeRO I}

INSRNS CES Joobenebancoacedd noo Sete CeR ee MeERErE man Enee ostoon odomuauoacstsocoabee tn: alee

INTRODUCTION.

This paper deals with part of the work initiated after a visit

to Australia, where the lack of any means of control of the Sheep

Blow-flies is paimfully evident. As some of these probably came

from England, it seemed desirable to study the natural checks

* Communicated by Prof. Lerroy.

Proc. Zoou, Soc.—i920, No. XV. 14

196 MR. A. M. ALTSON ON THE LIFE-HISTORY AND

that keep down the pest here: and the accidental discovery at

the Society’s Gardens of three important checks gave the work a

start. In this communication two Hymenopterous parasites are

discussed: other parasites are under investigation.

Acknowledgments are due to the Australian Commonwealth

Government for a small grant to assist the work.

H. M. Lerroy.

A SHorr ACCOUNT OF THE START OF THE WORK.

About the last week in June Prof. Maxwell Lefroy gave the

writer a tin containing approximately 1500 Dipterous puparia

to breed out. These had been forwarded by Miss Cheesman, the

Society's Assistant Curator of Insects.

The contents of the tin were divided into five lots, and put into

an equal number of jars containing damped soil. The jars were

covered with muslin. (At the outset, it should be explained that

occasional damping of soil was necessitated by the fact that the

soil was kept in sacks in the laboratory, and consequently any

moisture originally in it soon evaporated. )

Adult flies began to emerge a week later, and continued to do

so for a period of six or seven days. They were identified as

Calliphora erythrocephala Meig.

The jars containing the empty puparia and a large number

of intact puparia were then put aside. They were occasionally

examined, and on 14th July the first jar inspected was found to

contain an active Hymenopteron, which upon closer observation

appeared to be a female Braconid; the examination of the other

four jars showed that three of them also contained specimens of

the Braconid, Asthe remaining jar did not contain any Braconid,

the contents were emptied and carefully examined; all empty

puparia were separated from those which were still intact, and

six of the latter were opened, with the result that two were found

to contain fully-formed flies which had failed to emerge, another

a shapeless, smelling, moist mass—an atrophied fly-nymph—and

the other three contained Braconids, one in an early pupal state,

the others fully-formed adults apparently ready for emergence,

free of the pupal skin, and wings fully expanded ca

With the appearance of this parasite, breeding experiments

were commenced. The Braconids were released in a muslin-

covered glass cylinder, which was placed in a large tray containing

soil. Food was put in.

Blow-fly larvee and eggs were obtained by exposing meat at the

Zoological Gardens and on the baleony of the Imperial College of

Science, South Kensington, and a test made to ascertain if the

Braconids would parasitize the larve. The test was satisfactory,

and one female began ovipositing one minute after the admission

of the larvee, and in 25 minutes five females were at work.

* Both insects crawled out of their opened puparium ; one at once passed the

meconium and discharge, the other did not: both were males. The former lived a

normal period, the latter was dead on the second day.

HABITS OF TWO PARASITES OF BLOW-FLIES. 197

Numbers of the parasite were emerging from the jars at this

time, anda large cage was brought into use to contain them. As

it was desirable to obtain as many adults as possible, a search was

made at the Society’s Gardens at the spot where the original

material was found, and a further supply of puparia obtained.

Whilst sorting out this material preparatory to putting the intact

puparia into jars, a number of Chalcids were observed crawling

about on the soil. Most of these were captured. A closer scrutiny

of the puparia disclosed minute holes in several and Chalcids

emerging from some of them; it was also noticed that a few

puparia showed that Braconids had emerged from them. In view

of the appearance of this Chalcid, each puparium—they were

of a Calliphora species and presumably erythrocephala, the same

species as the flles—was placed in a separate tube. Both para-

sites continued to emerge from this material daily for a period of

twelve days.

As no Chaleids appeared from the original material, which had

been in the jars for nearly a month, it was evident that they

had carried out their attack after the original material had been

vemoved; and as this consisted entirely of puparia, it seemed

reasonable to assume that this Chalcid parasitized the pupal

stage. This supposition proved correct, and will be referred to

later.

A constant supply of blow-fly material for parasitization was

maintained by putting small receptacles contaiming meat out m

the open and placing it under control as soon as eggs had been

deposited on it. No opportunity occurred at this time to cbtain

meat blown by flies of definite species under control; but with

this object in view, small quantities of the larve from each lot of

blown material were segregated and allowed to develop into

ailalts, when their Species were determined, and they were then

placed under control in separate cages.

In the hope that the Braconid was still about, and in order to

obtain large numbers of them, receptacles containing soil and

meat were put out in the vicinity of the spot from which the

original material was obtained; this, Miss Cheesman kindly

undertook to do, and the receptacles were left in the open until

the blow-fly larvee had commenced pupation, when the material

was removed to the laboratory to be bred out. It was at once

observed that the large parasite was still about. It was seen

to come to the receptacles and to attack the larve. Whilst

this was being done, Prof. Lefroy put receptacles out at Heston

to see if the parasites could be obtained in this locality. Only

the Braconid parasite appeared from the latter place.

All lots of material obtained from Regent’s Park and Heston

were bred out to observe if any parasites other than the Braconid

and Chalcid made their appearance: none did.

By the end of August large numbers of the parasites were in

the cages in the laboratories and extensive breeding operations

in progress, and every effort was made to maintain a constant

14*

198 MR. A. M. ALISON ON THE LIFE-HISTORY AND

supply of active parasites so that by the end of the year there

would exist a large stock of hibernating material.

It was not until October that the writer took specimens of the

parasites to Mr. J. Waterston at the Natural History Museum

for identification. The Braconid was identified as Alysia mandu-

cator Panz. and the Chalecid as Masonia brevicornis Ashm.

Mr. Waterston kindly brought to the writer’s notice a paper by

Graham-Smith (1) in which these parasites were referred to. Up

to this time no reference to the Braconid was known, and that

the Chalcid should prove to be Wasonia brevicornis was not

anticipated, especially as the illustrations in Australian publica-

tions in the writer’s possession depicted a female of this species

walking with an extruded ovipositor, a position which it only

assumes in death.

BreEepine Mrrnops.

After the first lots of the parasites had completed their work,

the several species of blowflies, which had been obtained from the

meat exposed in different localities, were segregated.

The species obtained were :—

Blue Boublescn cece <-)<seaeeeer Caliiphora erythrocephala Meig.

be 067 <, FEL ie Cae pee * vonutoria Linn.

British Sheep Maggot-fly ... Lucilia sericata Meig.

Green Bottle. ..cc.. sccsneeanes 4 cesar Linn.

Green) Bottle ic. .xc.oncnteeeer Phormia grentandica 7tt.

In addition to these, the Common House-fly, Musca domestica

Linn., which is constantly bred at the College, was available.

Upon Prof. Lefroy’s advice as to the best medium in which to

breed the larve, ox liver was used, and found very satisfactory.

A somewhat important point in this connection was that liver

could be procured easily as it did not fali within the scope of the

‘Meat Rationing Order.”

The liver was cut into small pieces, placed in glass receptacles

and put into the cages containing the different species of ‘ blow-

flies.” As soon as it had been blown—that is, eggs deposited on

it—the receptacle was withdrawn and emptied into a glass dish,

in which additional pieces of liver were put when the eges

hatched. The receptacles were refilled and put back into the

cages when more eggs were required ; this method was continued

as long as larvee or puparia have been required. The glass dishes

with the newly-hatched larve were placed in large trays of

18 inches diameter and 4 inches height, and which contained finely

sifted sand or soil to a depth of 2 to 3 inches. It was found that

the full-grown larvee invariably crawled out of the glass dishes,

and after roaming about on the sand, proceeded to disappear and

pupate.

This method of breeding the blow-flies was successful with all

species except Lucilia sericata; the original lot of these were

obtained from the fleece of a “struck” sheep. A piece of sheep’s

skin was obtained consisting of the tail and the whole of the anal

HABITS OF ''WO PARASITES OF BLOW-FLIES. 199

area with a quantity of feces adhering to it, and was placed on a

piece of liver on fine damped soil in a tray; artificial heating was

applied, and the species was successfully reared by this means.

Several methods of presenting the larve for parasitization were

tried, the original object being the attainment of those conditions

in which they would be an easy prey for the Braconid’s attack.

It was found that free larvee severely damaged the females, so

that it became necessary to reproduce conditions as nearly natural

as possible.

In the case of the Chalcid, host puparia—in a proportion of

20 per female—were placed in small receptacles in the cages and

left there for ten days for parasitization. The puparia were

obtained from the trays by sifting; this rough handling did no

appreciable damage.

The breeding operations, were in progress from July to the

end of December ; from July to the middle of September ali lots

of parasitized material were reared to maintain supplies of parent

generations for the cages; after this period the parasitized

material was exposed on the balcony of the College to hibernate

and form the necessary stock for shipment.

Lirs-History AND Hasirs oF ALYSIA MANDUCATOR PANzZ.,

AND Hosts.

Introductory.— Alysia manducator Panz. was selected by

Latreille as the type of his genus Alysia, and belongs to the group

EHxodontes of the family Braconide.

It should be noted that the following account is mainly based

upon observations of this insect in captivity; opportunities to

observe it under natural conditions were very limited.

The Length of the Life-cycle.—TVhe length of the life-cycle from

egg to adult is, under suitable conditions of temperature, from

33 days and upwards, with a mean average of 52 days, but varies

considerably for some reason even amongst those of the same

parent and under the same conditions. ‘Table I. shows this

extraordinary variation, and refers to nine lots of material para-

sitized in the laboratory. Graham-Smith (1) records an instance

in which the life-cycle took only 25 days.

The Hgg—The egg (text-fig. 1) is very small, cylindrical,

tapering, and broadly rounded at one extremity, broadest at the

Text-figure 1.

Kee of A. manducator, from ovary. Size 675X150 mm. X61. Original.

other? with the micropyle narrowly attached and having the

appearance of a large protuberance. It is just visible to the

200 MR. A. M. ALTSON ON THE LIFE-HISTORY AND

TaBLE I.—Showing variations in length of life-cycle.

Alysia manducator.

Dates Nos. of Jars containing parasitized material and | Daily

of first and dates when parasitized. Temp. to

last SS eT ; 2 — | nearest

of adults. |17/7/19)18/7/19 18/7/19 19/7/19 21/7/19)21/7/19 31/7/19|31/7/19) 6/8/19 | Cent.

eee yes ce area ia

INES, 3 sooone A | 25

JOY egauoa Ie 24

PAD eaaene | 22

2 eerie 22

D2 Bee 19

Bind sees | 22

DA Peas 19

735). esas | 18

QEi alse. 18°5

Dipsenss 175

AS) 35 16

PS noma 19

G)0eanaden 175

EY esciare 19

Sle, DW coces: 17

Dok aise | 18°5

Saeco | 19

eee | 21°25

5 eafeielele | 24.

Crawest A f 22'5

(jeanne 21

Sikes | * i | 22°5

OP apaeen | 1 24

De doses | 25

11 ddogan | vy | { A QT

AD. v a + Mone 27

1a eons | | A 19

1 ae v Vara 18

Moa tease: | As || aly

16 4 V7

eet : 20

Ci | : | 21

LO sii | | 18

DON a: | | | 18

Die | | | | 18

20) hes | | | 18°5

2B eaten aoe

24 } | 20

2oneee | \ee | | 21

Beran: 20

Bilao | 20

AD) sence | 20

CAS ite | 19°5

BO Hine | | | | 195

Octane | | | | 20

Dra sae: | | Bas 20

Shyer | | | | 20

Ae eine eee — 195

Bikers | ages 20

Ge schjees | eu | | 20

i | | 20

ghnee | | | 19

Ouse | | | 20

THO) ad | | | | | | 20

eee | | 19°5

aes | | | | Vv | 20

Thus, contents of No. 1 Jar took to complete cycle 33-58 days; No. 2, 51-56 days; No.3,

50-57 days; No. 4, 51-56 days; No. 5, 53-55 days; No. 6, 49-55 days; No. 7, 44-68 days;

No. 8, 45-57 days; No. 9, 40-69 days: giving a general average of 463-59 days with a mean

average of 52.

HABITS OF TWO PARASITES OF BLOW-FLIES. 201

naked eye. It is translucent white, with a smooth glossy surface.

The time required for the ege to hatch varies, and in an investi-

gation into this point active first instar larve were found in the

hosts, which had pupated 66 hours after the eggs were deposited,

the temperature during this time being 20°C. But, that the

ege hatches in from 30 to 50 hours generally is obvious from the

condition of the host pupa, which is either in a state of histo-

lysis or an early stage of histogenesis ; and also by the occasional

distorted appearance of the host puparium. The egg is deposited

in any part of, the larval host, and this lack of discrimination

accounts for the non-appearance of the parasite on occasions. If,

for instance, the egg was lodged amongst muscles, it would be

crushed by the movements of the host larva; this point is

mentioned again under Oviposition.

The Larva.—YVhe first instar larva (text-fig. 2) differs in

appearance from the later instars, particularly in the shape and

appearance of its head. which is a thickly chitinized brownish

capsule fitted with a stout pair of mandibles, and has a pair of

small protuberances on the dorsal surface, the antenne. It

Text-figure 2.

First instar larva of 4. manducator. Camera-lucida drawing, partly

reconstructed. 41. Original.

consists, apparently, of 14 seginents including the head; the

ing Orenern terminates Ina “ Pale itee appendage, the sucloranravsl

vesicle. A similar organ in J/icrogaster, and other endo-parasitic

Hymenoptera, Gatenby (5) concludes is respiratory in function.

That this is the function of this organ in A. manducator appears

very probable, and it presumably functions by means of osmosis.

Since the larva sometimes hatches in the body of its larval

host before histolysis has begun, its head seems well adapted to

enable it to move in the host pupa if it hatches—-as it usually

does—during the process of histolysis or that of histogenesis.

What part of the host forms the food of the various instars

has not been observed, but it must consist of liquid or semi-

liquid, and in the first instar would probably be the body-fiuid

and fat-bodies; and the later instars would feed on those

systems which have ceased breaking down or building up after

the original attack of the first instar larva. That the larva

sometimes hatches before the host pupates is evidenced by the

fact that a puparium containing the parasite occasionally bears a

202 MR. A. M. ALLTSON ON THE LIFE-HISTORY AND

distorted appearance—indicating the severance of a muscle—or is

shrivelled and excessively wrinkled (text-fig. 8).

The first instar larva, with the exception of its brownish head,

is translucent white and glabrous throughout ; the integument is

very soft and easily damaged.

Text-fig. 3 illustrates a larva of the second instar, in which the

capsule head has disappeared and has given place to one the type

of which remains constant in the later instars. The particular

Text-figure 3.

Second instar larva of 4. manducator. Greatly magnified. Original.

larva illustrated was removed from a Calliphora sp. puparium,

and was placed in a watch-glass containing three macerated fly-

nymphs with a few drops of water added to reduce the density,

and with the aid of a microscope it was kept under observation

for more than half-an-hour whilst feeding. The larva remained

quite motionless during this time, its mouth-parts alone were

constantly working; no movement of the mandibles was observed,

but the extensive lip-like labium was seen to be continually

moving with a “rippling” action whilst the liquid was being

absorbed. The abdominal vesicle has proportionately shortened.

The mid-intestine now shows up clearly, swollen with food and

giving a pale yellow coleur to the larva.

The intermediate instars show no superficial difference from

the second, beyond the shortening of the “tail”-like abdominal

vesicle and the increase in the size of the larva and corresponding

swelling of the mid-intestine.

Text-figure 4.

AMA

Full-grown larva of A. manducator. Greatly magnified.

Length 4:25 mm. Original.

The last instar larva (text-fig 4) differs from the preceding

instars in t!:at ~-with the exception of the cephalic and posterior

HABITS OF TWO PARASITES OF BLOW-FLIES. 203

sezments—the integument is covered with fine sete and scattered

sensory hairs (text-fig. 5). The mouth-parts of this stage are

well defined (text-fig. 6). The posterior appendage has almost

disappeared ; easily visible just under the integument and scattered

about in the abdominal region are large white particles. The

spiracles, which are of a very simple structure, number nine pairs,

and appear on the second thoracic and first eight abdominal seg-

ments. There appears, on either side, a small main trachea with

segmental branches.

Text-figure 5. Text-figure 6.

Text-fig. 5.—Sete and sensory hairs on portion of integument of full-grown larva

of A. manducator. Greatly magnified. Original.

Text-fig. 6—Head of full-grown larva, partly reconstructed, camera-lucida drawing.

X 25. Original.

During all the larval stages the mid-intestine is closed caudally,

and it is not until the emergence of the adult insect that the

residuary and undigested food-matter is voided.

The last instar larva has, on several occasions, been observed

feeding upon the liquid remnants of its host, having at some stage

pushed the trachez of the host to the sides of the puparium ;

the latter being lined with the fly-nymph’s pupal skin. It

seems only reasonable to assume that chitin, represented by the

integument of the fly pupa, and the main tracheal trunks of

the blow-fly larva which remain constant in histolysis do not

form part of the larval diet. This postulation, if correct, would

exempt the Braconid larva from the necessity of eating its own

cast larval skins.

Before the pro-pupal stage, the final instar larva—which by

this time completely occupies the puparium of its host—spins a

silken cocoon, which either adheres (a) to the thick chitinized

walls of the puparium, or (6) to the stretched integument of the

fly pupa, and which generally is caused to adhere to the walls of

the puparium. In (@) the absence of the fly-pupal skin would

be accounted for by the first instar larva having started its

attack before the host larva had settled down tc pupate or

before histogenesis had started, and (6) after histogenesis had

started. So that the texture of this cocoon or lining to the

puparium is not always the same, the external portion of it—that

204 MR, A. M. AL'TSUON ON THE LIFE-HISTORY AND

which is in contact with the fly puparium— may be silk or pupal

integument, and in those cases examined the tracheal system

was visible either outside the silk or between the silk and pupal

integument; no cast skins however were detected, the search

was not very thorough. Sometimes the cocoon with the fly-

pupal integument shows the outline of the fly-nymph’s legs ete.,

indicating late hatching of the egg and after histogenesis had

proceeded some time.

The spinning of this cocoon raises an interesting point as to

the means whereby the larva, which appears to fill the puparium,

is able to move about within it, to accomplish the task of covering

every part of the inside, and also of absorbing the liquid remains

of the fly-nymph, taking into consideration that its intestine is

now filled with a semi-solid mass of undigested fly-pupa. It has

been observed that the spinning of the cocoon reduces the size of

the larva to some extent, but observation has not definitely

established the means adopted; some larve have been found

contracted into a small compass as though able to revolve within

the puparium and thus change their direction, whilst others

appear to be working in the manner common amongst silk-

spinning Lepidopterous larvee. The latter method is the most

probable, and is supported by the appearance of the larva.

The lning strengthens the puparium considerably, and is a

sure indication of the presence of the parasite in its last stages.

It is weakest at the anterior end of the host puparium, where the

chitinized mouth-parts of the blow-fly larva form an obstruction.

The length of larval life in the different istars appears to vary

considerably, but uo details were obtained; in one instance,

however, a full-grown Jarva was found in a puparium 18 days

after abe egg was deposited. ‘This is the shortest period recorded.

On the other hand, some intact puparia, which were from a lot

parasitized on 6th August, were opened on 26th November,

and 13 were found to contain full-grown healthy larvee, 112 days

after the eggs were deposited. The final istar larva is the stage

where the great variations in the life-cycle occurs. The time

taken to pass from this stage to that of the pupa varies from a

few days to many months. This point has been observed by

Graham-Smith (1), who also observed in 1915 that from material

parasitized in 1914, “the individuals emerging in the spring

were much larger than those which emerged in the autumn.”

This and the observations made during the course of this work

have caused the writer to form the opinion that among the

factors respousible—beyond the question of food-supply—is that

of the size of the host puparium. The integument of the

puparium of an undersized blow-fly larva is much thinner than

that of the full-grown larva, consequently the parasite within the

former is more susceptible to the variations of the temperature,

which alone would retard development.

In size the full-grown Jarvee vary considerably, but this is

naturally dependent upon the size of the host.

HABITS OF TWO PARASITES OF BLOW-FLIES. 205

The pro-pupal stage, 7. e. after spinning the cocoon, may last a

few days or several months. In two specific instances observed

the pro-pupa changed to the pupa, the one 2 days after it was

exposed in the puparium, the other 5 days after.

Some hibernating larvee from the stock have been observed in

puparium which they had not yet lined, although four months

had elapsed since the date of oviposition.

The Pupa.—The pupa (text-fig. 7) is loose within the cocoon,

and is capable of slight movement. The meconium now appears

to become slightly compressed, due to the process of meta-

morphosis.

Text-figure 7.

Lateral and ventral aspect of pupee of 4. manducator. Greatly magnified.

The figure on left a few days older than that on right. Original.

During the whole pupal stage the afore-mentioned large white

particles are visible, at fist scattered about in the abdomen and

later can be seen between the tergites and sternites.

The pupa, which is glabrous throughout, is, at first, creamy

white with reddish-brown eyes and ocelli, in a few days the head

and thorax turn grey, gradually becoming black; meanwhile, the

sternites and tergites—widely separated over the distended

abdomen—begin to turn grey, and the eyes and ocelli darken;

the legs and antenn™ similarly begin to show the coloration of the

adult, likewise the mouth-parts. ‘The last larval skin splits across

the head dorso-ventrally backwards, and then apparently slips

back to the apex of the abdomen, from which it is disengaged by

movements of the abdomen and antenne of the male or the

ovipositor of the female. In no instance has the larval exuvium

been found attached to the pupa, but lying in a crumpled mass

beyond the apex of its abdomen.

The length of the pupal stage was, in two specific instances,

found to be 7 and 10 (lays respectively with a mean temperature

206 MR. A. M. ALTSON ON THLE LIFE-HISTORY AND

of 21° C., but lengthens considerably in a low temperature ;

although this will not delay indefinitely the emergence of the

adult, which either emerges upon the slightest increase in the

temperature, or, in the event of a steady and prolonged

decrease, dies within the puparium.

The Adult.—The adult emerges from the host puparium and its

cocoon by breaking away that part directly in contact with the

head (text-fig. 8). It is furnished with a very powerful pair of

mandibles (text-fig. 9, a) beautifully adapted for the work in

hand. It breaks away the cocoon and puparium by an outward

movement, and does not bite the obstruction. Text-fig. 9, b

illustrates the outline of a section through the centre of a man-

dible, and shows its scoop-like appearance ventro-dorsally. In

Text-figure 8, Text-figure 9.

Text-fig. 8—Shrivelled and wrinkled puparium from which 4. manducator has

emerged, Greatly magnified. Original.

Text-fig. 9.—(a) Left mandible of adult. (b) Section through centre of mandible

; of adult. 41. Original.

the outward movement of the mandibles a small fracture appears

on the puparium, generally between the second and fourth

segment, gradually increasing in size until the perfect insect has

created an aperture large enough to enable it to escape. Obser-

vation has failed to disclose any use for the mandibles other

than breaking out of the host puparium, which operation, when

in progress by several insects, has been distinctly audible by

putting the ear over the mouth of the jar in which they were

confined. It is worthy of note that the mechanism of the man-

dibles must be diametrically opposed to that of most insects,

but upon consideration it seems obvious that by no other means

could so large a parasite escape from the puparium, which presents

to it a concave surface.

HABITS OF TWO PARASITES OF BLOW-FLIES. 207

The aperture made in the puparium is usually just large

enough to allow the insect to extricate its head and antenne,

thorax and legs, and part of the wings and the base of the abdo-

men, when by the pressure exerted against the distended

abdomen, the voiding of the meconium is assisted, and is either

accompanied or followed by a white discharge—the white particles

previously referred to,—which instantly dries and has a cement-

like appearance. This substance, upon analysis, has been found

to contain sodium hydroxide, but whether some of this is used

to soften the silk has not been established. The compressed

meconium, which is a deep black-brown rod, is completely

enclosed in a sac of integument, which resists boiling in caustic

potash, giving the impression that it is chitinized and probably

the larval mesenteron, which during metamorphosis has become

closed at the anterior end and compressed.

In some instances the exit-hole is made too Jarge, and the

adult appears with its abdomen still distended ; then by constantly

passing the tarsi of the hind-legs along it, it exerts sufficient

pressure to enable it to void the meconium: this method is not

always successful, and results in one or two days in the death of

the insect. Flight is impossible with the meconium unvoided,

although vain efforts to rise have been observed. In some

instances the transverse cut made with the mandibles is extended

so far around the puparium that, on emerging, the insect com-

pletely breaks off the top of the puparium. Hmergence generally

takes place from the cephalic end of the puparium, but in several

instances it has been observed that this was accomplished from

the posterior end.

The wings are always fully extended before the adult attempts

to break out of the puparium. Only in a very few cases has it

been observed that the wings have become damaged and torn by

the jagged edges of the emergence hole; this condition is generally

accompanied by injuries to the soft integument between the

abdominal piates, resulting iu the death of the insect in one to

two days.

Examination of intact puparia some time after the emergence of

the parasites has shown that for some reason unascertained—not

due to temperature—a small number never succeed in escaping,

whilst a few have been found with the cocoon-lining too thick to

extricate themselves.

Nothing of the pupal exuvium of those insects, which void the

meconium whilst escaping. can as a rule be seen; it is generally

buried under it, whereas the others are usually found with it

attached to the apex of the distended abdomen. Males emerge

before the females. The greatest emergence of males is from

3 to 4 days earlier than the females.

The Sexes.—Size :—The size of the adults varies considerably,

but is, of course, relative to the size of the host. The measure-

ment of several discloses a range of from 24 mm. to 64 mm. in

length.

208 MR. A. M. ALTSON ON THE LIFE-HISTORY AND

General appearance.-—The wales (text-fig. 10) are easily

distinguished from the females by their long antenne, which are

approximately equal to the over-all length of the insect from the

frons to the apex of the wings when in repose, and which extend

beyond the apex of the abdomen. The ovipositor is barely visible

when the insect is looked at from above, but can be seen when in

a lateral aspect. Both sexes are shining black, with rufous legs.

Text-figure 10.

Male A. manducator. Greatly magnified. Original.

Courting and Mating.—TYhis takes place very shortly after

emergence, but never occurs unless the meconium has been voided

some time. When seeking a mate (in the cage) the male moves

vapidly along; violently swaying the antenne and vibrating the

wings, which are opened as in flight. Mating lasts about

8 minutes, but was not frequently observed. ‘The sex attraction

is only evident in the male, and when newly-emerged females

were admitted to the cage, males eagerly sought them. The

female evinces little interest in the male and can easily repel it.

The males make no distinction between virgin and fertilized

females.

Flight.— Both sexes are capable of sustained flight. This has

been observed in the cage, and on fine days it appeared as if all

were “on the wing.” Any which escaped from the cages or

when handling were generally instantly out of reach and through

the open skylights. Those observed around baited receptacles

in the open appeared to come from all directions; this is con-

firmed by Marshall (quoted under Oviposition).

HABITS OF TWO PARASITES OF BLOW-FLIES. 209

Ovipositicn.—Females do not begin ovipositing until a day or

so after emergence, and irrespective of whether they have mated

or not; a few have been observed trying to oviposit before

voiding the meconium.

The ovaries are large and well filled with eggs. A dissection

of the ovaries of 12 females gave an average of 366 eggs per

female. The greatest number counted in a single female was 416.

Graham-Smith (4) records a female with ovaries containing

“Cat least 549 eggs.” Table II. gives the details of the count of

the contents of individual ovaries :-—

TasLE [1.—Contents of Ovaries of 12 females.

No. | No. | No. |

| i | 7

| No.| No.| No.| No.| No.| No. | No. | No.| No. :

| TCO 4) ala |) Ie Total. | Average.

| i.) 2) 8.) 4.) 6. Gall 7a 148.1 9.

| |

; 412| 375 | 408 | 368 | 352 | 328 | 363 | 416 | 362

25 338 347 4394 | 36616

Oviposition takes place in the larva of the blow- fly (text-fig.11);

half- to full-grown larvee are usually selected. In the cage the

Text- figure 11.

Female 4. manducator ovipositing in larva of Phormia grenlandiea.

Greatly magnified. Original.

chemotropic effect of carrion was—when the atmosphere was not

charged with the odour—almost instantaneous upon the females -

( = Duin 5 ceed 0 2 Sy)

they became violently agitated, swaying their antenne, and then

might proceed to clean themselves, particularly the antenne

and the abdomen ventrally in the region of the Ovipositer, and fly

to the carrion, or reverse the procedure and fly over and around

the receptacle containing it and the larve, and then alighting near

z i fo) :

by, go through the cleaning process. It is, of course, obvious

210 MR. A. M. ALTSON ON THE LIFE-HISTORY AND

that the sense-organs in the antenne and the palps of the

ovipositor had suddenly received the odour of the carrion and

stimulated the insect to oviposit. The males are also attracted

by earrion—apparently a sexual tropism to enable them to locate

the females. Marshall (3) observes:—‘ They ” (both sexes) ‘scent

the aroma of carrion at a surprising distance, as I once had

occasion to observe in watching the remains of a dead rook, upon

which they descended in constant succession, apparently from the

sky, like vultures. The males generally alighted on blades of

grass close to the attractive object, as if to wait for their

partners, without interrupting them in their unsavoury occu-

pation.”

As soon as the females have alighted, they begin their search

for the larve, and if the initial effort to insert the ovipositor is

successful, they remain depositing eggs until they die, either from

exhaustion or because their task is completed. A few have been

observed to leave the larve and go in search of food and not

return the same day.

Only one egg is deposited in each larva by an individual

female, at least, under conditions which are approximately

natural—i. e., where the larva has means of escape by disappearing

into the meat or earth.

The ovipositor is inserted diagonally under the integument of

the larva when the attack takes place in the thoracic or

abdominal regions. The larva makes every effort to escape,

wriggling and squirming, and frequently damaging the female.

These frantic efforts to escape are put an end to by the effect of a

poison injected with the insertion of the ovipositor. The

immediate effect of the insertion of the ovipositor is to cause the

larva to vomit and void. The poison produces a paralyzing

effect, and causes the larva to contract and then lie motionless

whilst the egg is deposited. This operation varies in time, and

may take 30 seconds to 5 minutes*—the more exhausted the

female is, the longer it takes.

The “temporary paralysis” of the larva lasts from one to two

minutes, so that when an exhausted female delays the with-

drawal of the ovipositor the larva endeavours to release itself ;

normally though, when the ovipositor is withdrawn, it is still

motionless, and remains so for some seconds. The female then

moves off in search of another victim. , The first movements of

a larva recovering from the “temporary paralysis” are to extend

itself to its full length, and then, still shghtly under the influence

of the poison and the unpleasant ovipositor, it goes through a

series of extraordinary convulsive and constrictive movements,

which ripple the integument either from the cephalic to the

anal end or vice versa, as though it was endowed with intelligence

* One female, which had not oviposited in more than a dozen Jarve, was observed

to retain the ovipositor within a larva for 21 minutes, and as the initial dose of

poison did not suffice to keep it motionless, further doses were apparently

administered each time it moved. Needless to say, this larva died.

HABITS OF TWO PARASITES OF BLOW-FLIES. 211

and was trying to squeeze and crush the ege*. As soon as this

ceases it hastily disappears into the soil or carrion, apparently

stimulated by its experience to pupate. The foregoing obser-

vation, naturally, only refers to larve free of the carrion or on

top of it (otherwise they could not have been observed), but those

attacked which are partially buried in it do not appear to behave

in the same manner; they are surrounded by food and hidden

from the light.

The first 20-30 larvee attacked will be “‘ paralyzed” instantly,

and then the poison apparently becomes less rapid in its effect ;

whilst it has been noticed that a female which rests for some

time and feeds, seems to renew the effectiveness of its poison.

An individual female—as has been already noted—does not

normally attack the same larva twice. In this connection a

series of experiments were carried out under conditions as nearly

natural as was possible, and by employing one female at a time

it was observed that each larva parasitized, either free of the

carrion or partially embedded in it, made every endeavour to get

into the soil after it had recovered from its “‘ temporary paralysis ”

either directly or through the carrion. It appears to be definite

that effective oviposition stimulates the larva to pupate. How-

ever, in cases in which two or more females inject the same larva

the second one coming upon it just when it begins to move

or encountering it on its way to escape,—its death ensues within

24 to 48 hours. Its organs appear to disintegrate, the dead

larva gradually darkens until it turns black; dissections disclosed

little else than a thick dirty putrid liquid. In warm weather,

with a temperature of about 22°C., the dead larva dries up

within 6 or 7 days. The majority of these over-parasitized larvee

never succeed in getting into the soil.

A female does not attack a motionless larva, but may prod it

with the ovipositor, usually causing some movement with

unpleasant results for the larva. It is also in this way that a

larva just recovering from “temporary paralysis” becomes

a victim to over-parasitism.

On several occasions the writer has observed a moving larva

stop suddenly and lie motionless upon the approach of a female,

although it may have been touched by no more than one leg of

the latter. This behaviour is obviously due to the larva having

been previously attacked by a female, but whether it was attri-

butable to chemotropism, which is most probable, has not been

established. The predominant odour was that of carrion.

Females which have been ovipositing for some time and are in

a filthy condition from the carrion and putrefactive juices, will

endeavour to insert their ovipositor into any object which they

happen to feel moving, with the result that frequently two or

more can be seen together, “jabbing” their ovipositors amongst

each other’s legs in a vain effort to get it firmly fixed.

* The writer, as previously stated, is of the opinion that many of the eggs are

damaged by this movement, particularly if located in or amongst muscles.

Proc. Zoou. Soc.—1920, No. XV. ey

PALLY? MR. A. M. AL'TSON ON THE LIFE-HISTORY AND

The ovipositor, which is extruded, is inserted in different parts

of the larva’s anatomy. Generally, when the larva is free—i.e.,

on the surface of the carrion, under it, or on the surface of the

soil,—it is inserted about the Ist to 3rd abdominal segments ; very

small females attack the cephalic end or the anal plate, particu-

larly in the case of full-grown larve, probably owing to their

inability to get astride the victim, as is usuai with the larger

females. When the larva is moving in the carrion and the

movement is perceptible, the female endeavours to insert the

ovipositor through it into any part of the larva, whilst a female

coming upon a larva disappearing into the soil or carrion will

attack the apex of the abdomen or anal plate, and sometimes is

unable to bring the victim to a standstill before the ovipositor

has disappeared into the cavity; whilst, on the other hand, a

larva emerging from the carrion is attacked in the anterior

region.

Whilst the insect is ovipositing, the palpi or “feelers” (text-

fig. 12), which in repose form lateral sheaths to the poison and

Text-figure 12.

Palp of ovipositor of A. manducator. W.P., wiping-pad. X 61. Original.

piercing-blades, are constantly in use for locating the moving

larva. The ovipositor is withdrawn from a victim with a distinet

jerk in order to disengage the barbs of the piercing-blades, and

by means of the powerful muscles attached to the hinge-like

continuation of the blades, it instantly springs back between the

palpi, the apex of the former alighting at the base of the latter,

so that whilst the sternites are assuming their normal position of

repose, the point of the ovipositor is cleaned by the ‘‘wiping-pads ”

(text-fig. 12, W.P.) of the receding palpi. These “ wiping-pads”

are on the inside of the basal portion of the palpi, extending

HABITS OF TWO PARASITES OF BLOW-FLIES. 2illess

approximately half of the length, and consist of soft dirty white

integument roughly in folds.

Length of Period of Oviposition.—Females, whether freshly

emerged or otherwise, and which had or had not mated, once

allowed to oviposit, were dead the next day if they had been

constantly ovipositing, or when resting and feeding occasionally

lasted three to four days ; in one instance a female lived five days

ovipositing in about 50 larvee daily.

Progeny of Single Females.—No extensive data were obtained

on this point, but in the laboratory the.average was 33:74 per

female. (See Percentage of Parasitization.)

Parthenogenesis.—Unmated females reproduced males ; whether

the latter were fertile was not investigated.

Proportion of the Sexes.—Vhis appears te be about equal on

the whole in individual lots of material parasitized in the

laboratory, although one sex may considerably predominate.

The figures given under this heading are details of emergence

from three lots of material which were parasitized by free

Braconids in Regent’s Park, and give the following result: 199g

and 294 9. No other details of emergence from outside material

were compiled. Graham-Smith (4) gives, substantially, the

following details for ‘“‘Spring” and “ Autumn” batches which

emerge in 1916 from material parasitized in the autumn of

1915. The sum total for both batches are 2891 ¢ and 749 9,

clearly indicating arrhenotokie, a proportion, which the writer

ventures to suggest may, in this instance, be due to the immediate

proximity of host-infested carcases when the females of the

autumn batch of 1915 emerged.

Length of Life of Adult.—In the laboratory the length of the

life of adults was not very long. Observations showed that, in

the case of two freshly-emerged males which were not permitted

to mate, one lived 25 days, the other 31 days; two unfertilized

females not allowed to mate or oviposit lived 33 and 38 days;

two females and two males confined together, the former not

being allowed to oviposit, the males lived, the one 18 days, the

other 23 days; the females, one 25 days, the other 33 days. And

as stated oviposition rapidly ends the female’s life. In each of

the foregoing tests ample supplies of food were given, but the

insects were confined in glass jars, which did not give much

opportunity for fight and the atmosphere was permeated with

the odour of carrion. The length of life in the open or in

unpolluted air would doubtless be longer.

The length of life within the main cage appeared to be much

shorter during the months of November, December, and January

in spite of a mean temperature of 20°C. Activity was always

greatest on fine days, and particularly when the sunshine was

directly upon the cage.

Food.—When the breeding operations began, the adults were

fed on sugar diluted with water in a ratio of 1 to 5, but owing to

the ‘‘ Rationing Restrictions,” honey had to be substituted, and

15*

214 MR. A. M. ALTSON ON THE LIFE-HISTORY AND

diluted in the same proportion proved to be the better food, and

was occasionally varied with water only. Observation in the

open failed to discover any of this species feeding, and in this

connection Marshall (8) only observes “ both sexes likewise fre-

quent flowers for the purpose of feeding.” It is probable that

“honeydew ” constitutes a source of food, as is the case with

many Hymenopterons, but was not tested with this species.

Seasonal Abundance.—From 14th July, 1919, to the end of

January 1920, with a few days’ exception, there have been active

adults in the main cage; the breeding, which, to begin with, was

almost a daily occurrence, caused such an overlapping of

generations as to bring about this result. Graham-Smith (4)

observes :—‘* The parasite, however, is abundant throughout the

season, for it was observed attacking fly larva from 30th May to

Ist November, 1916.” From the observations of the writer, these

insects were present in large numbers from July to October 1919

in Regent’s Park; and as the original stock bred in the

laboratory appeared on 14th July, and taking the average period

of the life-cycle as 52 days (see Table L.), this gives 24th May as

the date upon which the parents of this generation were deposit-

ing eggs; this therefore confirms the above observation.

Details of Hmergence.—Accurate sex details of the daily emer-

gence from certain lots were kept from 18th August to 20th

September, after which date these had to be abandoned, the

breeding operations at this date occupying all available time.

Details of daily emergence for the period 3rd to 12th September,

corresponding to the period of greatest activity shown in Table I.

and affecting laboratory-bred material from the jars No. 1, No. 2,

No. 3, No. 4, No. 5, & No. 6 and from No. 7 for one day shown

in Table I., are given below in Table IIT. :—

Tasie ITI.— Details of Daily Emergence.

PasNox 8 918 2/8 Pd Bid Pld Bs Vis Pls Qs Y| Totals.

Pega erlgn) eto (et cle ee tema ame Geet eeD gen Gee

for one day.| — 3 9 13} 38) 42) 34) 31) 42) 158

Dates. Females.

Sept. 3rd | 4th | 5th | 6th | 7th | 8th | 9th | 10th|11th|12th) 214

These figures merely indicate the early appearance of the males.

During the period covered by Table Ill. the average daily

contents of the main cage was over 300 individuals of both sexes ;

this figure allows for the withdrawal of ovipositing females and

admission of daily emergences from all sources.

Percentage of Parasitization.—Graham-Smith (1), referring to

two lots of material attacked by ‘“‘free” A. manducator in 1914

and bred out in 1915, states :—‘“ In the former it is 60 per cent..

HABITS OF LWO PARASITES OF BLOW-FLIES. 215

in the latter nearly 90 per cent. That the latter figure is a true

index of the extent of Braconid infection in the original stock of

pupie is shown by the fact that 9 per cent. of the pupz in the sun

tin were not parasitised.” Of two lots of puparia from which—

the parasite emerged during 1916, this observer found (4) :—

“« At least 25 per cent. of the puparia” from one lot ‘and 57 per

cent. of those” from the other ‘were infected with A. mandu-

cator.’ Summarizing his observ: poe on emergence of this

parasite in 1915-16-17, he says (4):—Of the 1G 028 puparia

collected in the autumn of 1914— 15 6 from sunny and shady

situations 7041 or 43 per cent. were infected with A. manducator,

while of the 4787 puparia collected during the summer months

of 1916-17 only 508 or 10 per cent. were infected.”

From two lots of material collected from Regent’s Park on

27th July, and where parasitization by ‘“ free” Braconids occurred

under conditions similar to those above, the writer found the

percentage of parasitism to be 23°12 per sent, and 52-32 per cent.

respectively. These lots were both brought in at a time when

the larvee had begun to pupate, they were examined in November,

previous to this ‘time emer gence had ceased. The figures given

are derived from the number of puparia from which “A. mandu-

cator had emerged, and from those intact puparia in which dead

adults, dead pupee, and active and dead larvee of the parasite were

found, as against the total number of puparia in the receptacles,

including those from which flies had emer ged, or failed to

emerge, and those containing atrophied fly- nymphs. It would

be of interest to know the number of females responsible for

these figures. Dead females were always found in the receptacle,

but no reliance can be placed on their number.

In laboratory-bred material, in ten lots, it was found that the

percentage of parasitization ranged from 15-04 per cent. to 48°99

per cent., the latter figure being due to nine females. But the

outstanding feature of this examination was, that it took a total

of 39 females to reproduce definitely 1306 individuals exclusive

of over-parasitized larve—an average of 33:74 each, or only

9°89 per cent. of their average egg-capacity. This result can but

be attributed to confinement and to the bad ventilation of the

cage, also to the fact that the parasite’s sense-organs, which are

normally stimulated to bring about the inclination to oviposit,

would have been dulled and deadened by the ever-present odour

of carrion in the laboratory. Under better caging conditions

and when living in an atmosphere unpolluted with the stench of

the putrefactive juices of the carrion, better results would

certainly be obtained.

Hibernation.— A. manducator hibernates as a full-grown larve.

This was observed under natural conditions by Graham-Smith

(1 and 4), and is supported by the condition of the stock material

and by the Refrigerating Experiment.

Attraction to Light.—Avtificial light is very attractive, and

caused the insects to crowd on that side of the cage nearest to it.

216 MR. A. M. ALTSON ON THE LIFE-HISTORY AND

Hosts.—The species of larve used in this work and from which

A. manducator Panz. was successfuily bred, are as follows :—

Lucilia sericata Meig.

5, cesar Linn.

Phormia grenlandica Ztt.

Calliphora erythrocephala Meig.

Ns vonitoria Linn.

No preference was shown for any individual species, the

odour of the putrefactive juices being the chemotropic stimulation

to oviposit; and therefore it is probable that other carrion-

feeding cyclorhaphous larvee would be found to be suitable hosts.

In this connection Marshall (3) states:—‘‘ They have been

reared from various maggots, as Lucilia cesar L., Cyrtoneura

stabulans Fall., Hydrotea dentipes Fab.”

Lirre- History anp Hasirs or VAsSON/A BREVICORNIS ASEM.,

AND Hosts.

Historical.—Nasonia brevicornis Ashm. was first described by

Girault and Saunders, 1909 (6). They bred it from the puparia

of the Common House-fly (J/usca domestica Linn.) at the Illinois

Entomological Station at Urbana in 1908. It has since been

reported from Chili and India, and was bred in 1911 from the

puparia of C. erythrocephala Mg. by Graham-Smith (1) at

Cambridge, England. In November 1913 it was bred from the

puparia of Pycnosoma rufifacies by McCarthy (2) at the Govern-

ment Sheep-fly Experiment Station at Yarrawin in New South

Wales, and a few days later discovered at Longreach, Central

Queensland (7). As already stated, page 197, it was first bred

by the writer from the puparia of C. erythrocephala collected

from Regent’s Park, London, in July 1919.

Introductory.—In the following account of the life-history and

habits of this insect, the writer freely quotes from the writings

of the previous observers, adding here and there observations of

his own. In doing this the writer desires to put forward as

general and collected an account of this important insect as

possible ; further, it may be observed that the writer had already

made an independent study of this subject before specimens were

identified and the original description of Girault and Saunders (6)

brought to his notice. Full acknowledgment is made to these

references, which for the sake of the continuity of the account

are mostly signified by reference numbers only.

The Length of the Life-cycle.—This is entirely dependent upon

the temperature, and its influence affects the full-grown larval

stage the most.

The various observers differ as to the period of the life-cycle,

but this can be accounted for by the probable differences in

temperature when the observations were made. Girault and

Saunders (6) give 224 days as the average in the spring and

15 days in autumn; McCarthy (2) 11 days in summer; Froggatt

HABITS OF TWO PARASITES OF BLOW-FLIES, 217

(7) 11 to 14 days; Froggatt (8) 15 days. Under laboratory

conditions with a mean temperature of 20° C. it was found to

average 21 days.

The Egg.—The egg is translucent white, with a smooth glossy

surface, cylindvical, slightly tapering to one extremity and

broadly rounded at both ends (text-fig. 13). The newly-deposited

egg measures from °30 to ‘35 mm. in length and :11 to 14 mm.

wide at broadest part (8). The eggs are placed in clusters under

the shell of the puparium and upon the surface of the pupal

integument of the developing fly, the latter occasionally being

punctured by the ovipositor. ‘They are found in clusters of

Text-figure 13.

Eee of WN. brevicornis, 26 hours old. Size ‘125 *350 mm.

x41. Original.

2 to 12 or more or singly, and are situated on any part of the fly-

nymph, but are commonly found in the depression of the

junction of head and thorax, or thorax and abdomen. ‘The eggs

appear to be coated with a trace of some sticky substance, which

causes them to adhere together, and to the covering integument

of the fly-pupe” (8). The egg undergoes a slight increase in

size as the development of the embryo progresses. The duration

of the egg-stage varies considerably ; those observed ranged from

30 to 74 hours.

The Larva.—\mmediately upon hatching, the young larve start

feeding. They puncture the pupal skin of the host with their

mandibles, and with these firmly attached to the enveloping skin

of the pupa, proceed to absorb the body-fluids of their host.

They remain about the same position until full-grown. As the

larve develop, the host is gradually absorbed, and consequently

shrinks inversely to the growth of larve. “As a rule, the

remains of a parasitised host—the fully-formed pupa—is a flat,

scale-like mass, apparently consisting of the ventral shell of the

pupa and that of the head; for example, the thece of the eyes,

legs, and wings are discernible, and the remains are not much

shrunken so far as the original length is concerned. In the

ease of Cynomyia cadavernia, in one puparium infested with

21 larve of the first spring generation the parasites were all

attached to the dorsal surface of the host from the pronotum to

the tip of the abdomen; these parasitic larvee were nearly full-

grown. But in another puparium of the same host, in which

13 larve were found, their attachment to the host appeared to

be haphazard, and the host-pupa was considerably shrunken,

especially in width” (6). No evidence of larval predaceousness is

forthcoming, although dead larve are frequently found; these

218 MR, A. M. ALTSON ON THE LIFE-HISTORY AND

are sometimes found with living ones, but usually all are dead.

The dead larva is always discoloured and appears to have shrunk

to the size of the meconium, so that they are then quite hard.

The number of larval instars have not been worked out, but

the general appearance of the larve is constant. The larva

(text-figure 14) is a typical vermiform hymenopteron, broadest in

the centre and tapering towards the extremities. It consists of

apparently 14 segments. It has nine consecutive pairs of

Text-figure 14.

Full-crown larva of N. brevicornis. X25. Original.

to) to)

spiracles, of a very simple structure, on 2nd and 3rd thoracic .

and Ist to 7th abdominal segments. The integument is smooth,

glabrous, very thin and easily damaged; it appears to secrete

some sticky fluid which entirely bathes the integument, and

which enables it to ‘‘ grip” the surface over which it is moving.

An examination of the larval integument discloses minute pores

scattered about the surface and presumably the source of this

“sticky” fluid. Jarvee have been frequently found in a puparium

from which a number of adults have emerged with portions of

pupal exuvia attached, sometimes completely covered with it.

The mid-intestine is closed candally during the entire larval

development, and the waste matter within it is brown. This

gives the larva a dirty white appearance. The slightest puncture

made in anactive larva about the centre of its body will cause the

contents of the intestine to exude instantly, leaving the larva

white. The period of development of the larve occupies about

7-10 days from the time of hatching, with a pro-pupal stage of

1-3 days.

A few hours before casting its last larval exuvium, the contents

of the mid-intestine is voided. This appears to indicate that the

mesenteron and proctodeum become connected before the pupa is

formed. ‘The meconial discharges of this parasite, found

scattered through the host puparium, are brownish yellow or

dark olive-gveen in colour, and consist of small conglomerations of

round pellets, or are sometimes in irregular chains, like some

bacteria, but are never single, solid pieces. .... ” (6).

The length of larval life is very variable.

The Pupa.—After voiding the meconium the larva remains

motionless—unless disturbed by a fellow-oceupant of the host

HABITS OF TWO PARASITES OF BLOW-FLIES. 219

puparium—with its posterior segments surrounded by the me-

conial discharge, which dries rapidly. So that when the larval

skin bursts and recedes, the pupa (text-fig. 15), which does not

entirely free itself from the larval exuvium, consequently becomes

anchored to the meconium. This has been observed by emptying

the larval contents of a puparium into a small glass tube.

Text-figure 15.

Pupa of WN. brevicornis. Greatly magnified. Original.

The duration of the pupal stage is variable. ‘This was

obtained in one ease only. A larva pupated during the night of

Sept le, MOOS re. 2. The resulting adult female emerged

at 10 a.m. Sept. 23, 1908, making a pupal stage of approximately

54 days. ‘The average length of this stage for the first spring

generation (17 cases) was 9 days (May 14-23, 1909)” (6).

Another observer states:—‘‘The pupal stage occupies about

five days” (8). Those pupz observed by the writer ranged from

8-12 days. ‘‘ When first formed, the pupz are yellowish white,

the eyes garnet, with some duskiness at the caudal edges of

the abdominal segments soon afterwards; the mandibles, legs,

antenne, and wing-pads gradually become dusky, and about

48 hours previous to eclosion, the head, thorax, and abdomen, in

succession, begin to show dark colour, the head and thorax

together becoming a deep black before the abdomen shows very

much colour, and then, after about 6 hours, the latter turns

gradually but rapidly black. About 20 hours before eclosion the

colour is jet-black, which just preceding emergence changes nearly

to the colour of the mature adult. At eclosion the adults are

fully coloured.” (6.)

The pupal exuvium, which is very stiff, is a golden brown, and

does not recede as in the Braconid, but appears to be broken up

in parts; pieces which retain the outline of the antenne are

frequently found loose; similarly, pieces from the head and legs

can be found; the abdominal portion seems thinner, and is usually

attached to the apex of it when the insect emerges.

220 MR. A. M. AL'TSON ON THE LIFE-HISTORY AND

The Adult.— Emergence of the Adult.—‘‘In general, it may be

stated that the adult parasites emerge from the host puparium

through from 1 to 3 cireular holes, situated variously, usually in

the dorsal or dorso-lateral aspect; and when more than one exit-

hole, the two or three are usually scattered or widely separated.

The manner of emergence does not differ for sex. The exit-hole

varies in diameter from 0°75 to 1:50 mm.; it is usually larger and

single when the host is Musca or Chrysomyia and smaller when

Phormia, though this difference may be more apparent than real.

Individual exit-holes may of course vary considerably in shape ;

for rarely it may involve the whole of one end of the host

puparium, and is then relatively very large and irregular. The

margins of the exit-holes are always jagged or serrate, showing

that. the adults gnaw their way out....... In regard to the time

of emergence, the males usually emerge from “2 to 20 hours

earlier than the females, a few emerging some hours previous to

the simultaneous emergence of the majority, but there is con-

siderable variation in individual cases. Thus some males may be

the last to emerge, but the tendency is for them to emerge earlier

than females.” (6.)

In the laboratory the number of exit-holes was not limited

to three, four and five have been seen, and the exit-holes, con-

sequently, dorsal, ventral, or lateral (text-fig. 16).

Text-fgure 16.

Puparium with two exit-holes latero-ventrally. VV. brevicornis.

Greatly magnified. Original.

Both sexes emerge with the wings fully expanded.

Lianvee Lene been fe in puparia from which—several weeks

or Its had emerged. If this takes place

under el tanaitiontt it would appear that these have very

HABITS OF TWO PARASITES OF BLOW-FLIES. 221

little chance of completing their development, and would probably

be killed by mites or some other enemy.

Many attempts were made to observe the initial process of

making the exit-hole, but without success. However, on one

occasion a female was observed enlarging an exit-hole. A few

days previously this particular puparium had been enclosed in a

glass tube stopped with a cork. The tube was under observation

with the aid of table binoculars to watch the movements of some

adults of both sexes which had emerged. There were two exit-

holes in the puparium, dorsally, with centres about 3 mm. apart,

one being much larger than the other, and through the former

all the large females in the tube must necessarily have escaped.

Suddenly a female's antenne were noticed to issue from the small

exit-hole, and were violently agitated. The upper part of the

epicranium could be seen against the inside of the puparium ; the

head was then moved, as was evident by the changing positions

of the waving antenne; eventually, after the scapes had several

times come into contact with the serrated edges of the exit-hole,

they were withdrawn; the female could then be seen changing

its position. This accomplished, the head began to appear latero-

ventrally through the hole, the right eye coming up first;

gradually more and more of the head appeared still inaintaining

its latero-ventral aspect, until the edge of puparium was slightly

oblique to the junction of the mandibles. The right eye and

gena were then above the surface of the puparium, the antenne

still inside. The female now began to bite the puparium, the

right mandible coming down on it from the outside, the left

working up on it from inside. This process continued in a leaf-

eating-caterpillar-like movement—semicircular forwards and

backwards— until the hole was made large enough to enable the

female to escape. No effort was made to test the size of the

hole; the insect continued to bite until, suddenly ceasing, it

began changing its position again until it was ventral side up

inside the puparium, then, moving forward and bending upwards,

the antenne, head, anterior legs, thorax, middle and posterior

legs, and abdomen appeared successively, until the female was on

the surface of the puparium, erect. It immediately hegan elean-

ing itself: the apex of the abdomen bore a portion of the pupal

exuvila, which came away with a minute white meconial discharge.

A few seconds later the female was found by a male.

In the laboratory, when the puparia are unburied in soil, these

discharges can be seen scattered all over them, and appear as

minute white strings.

The Sexes.—Size :—The females vary from 1:0 mm. to 2°30 mm.,

the males from 0°60 mm. to 2-0 mm. (6).

General appearance.—-Both sexes are distinguishable to the

naked eye. The females have large wings extending beyond the

apex of the abdomen, whereas the males (text-fig. Wy ) are semi-

apterous; their ond nena wings do not extend farther than

the third abdominal segment.

222 MR. A. M. ALTSON ON THE LIFE-HISTORY AND

“...the female is a rich dark green, the abdomen nearly

black, the head and thorax dorsad, reflecting brassy scintillations,

in some lights entirely brassy ; in the ventral aspect and also the

lateral, the thorax appears bluish; the antenne dark, the scape

and pedicel a rich brown, the legs mostly brown with darker

femora. On the contrary, the males appear a brilliant bright

metallic green, reflecting brassiness, the antenne and legs light

yellowish brown, the wings small, clouded, with a soiled appear-

ance; the ventral aspect is the same, but with slight traces of

metallic bluish. The male is noticeably more brilliant and bright

than the female, which is somewhat sombre.” (6.)

Text-figure 17.

Male NV. brevicornis. Greatly magnified. Original.

Variations of the foregoing description of the type species

were observed in the females; with hght brown or yellowish

brown femora, not dark or clouded; general colour deep metallic

blue ; fore-wings not naked in the basal triangle of wings below

the submarginal vein, the costal margin with more bristles. Of

six such specimens mounted for microscopical examination, no

uniformity was found in the venation. ‘The number of sete

in basal area of fore-wing varied from 2-5, position also varied.

Bristles on costal margin also varied, as many as 14 were

observed.

Courting and Mating.—‘‘ Courting in this insect is not a com-

plex habit. It follows almost immediately after emergence, at

least in confinement. Where «a number of both sexes are

gathered together, all recently emerged, the males and females

ave constantly in motion, the former active, seeking the females ;

the antenne of both sexes also constantly in motion, held inclined

upward in the natural position, giving quick, jerky, wavy move-

ments. When one individual meets another, the antenne simply

touch whichever portion of the body presents itse’f first, and the

two turn aside and pass on; or if they happen to be individuals

of opposite sex and (apparently) the occasion is suitable—which

en CE PEPE Ma aS pace Ne

en ee ee

HABITS OF TWO PARASITES OF BLOW-FLIES. 223

is most often the case immediately following emergence—the male

hastily climbs upon the back of the female, runs forward, and grasps

her head with the fore-feet, usually at the lateral aspect of the eyes

or sometimes at the cheeks; the intermediate feet grasp some

portion of the thoracic pleura, usually at the mesothorax, and the

hind-feet take hold along the sides of the abdomen or the edges

of the flat wings. The legs are not stretched out or used for

embracing the body of the female, but the hold is taken by the

feet alone, and the position of the male is not strained, but rather

that of the natural position of rest. His body is parallel with

and above the body of the female and projects beyond (cephalad)

it, so that the head is between the upturned antenne of the

female and stretched over hers, his abdomen reaching to a point

above the third abdominal segment or to a point opposite to the

distal end of the marginal vein of the fore-wings, upon which it

actually rests. Having quickly attained this position, the male

senses the antenne of the female with his own, and immediately

begins suit in earnest by rubbing his head up and down against

the inner (mesal) surfaces of the flagella of the female, which

are held up in a V-shaped position, at the same time holding

the scapes erect and apart and the flagella back, pointing laterad

at right angles to the scape and at every downward movement

bringing the scapes together; this movement of the head is

accompanied by a corresponding ‘ petting” movement of the

female flagella against the cheeks of the male. The up and

downward movements of the head are regular and continued for

from 5 to 10 seconds, each completed movement occupying

slightly less than a second of time; and they are alternated with

a period during which the head of the male is motionless and _ his

antenne sensing these of his mate, either by touching both of

their tips to the tips of her antenne, or else by stroking them up

and down; the mandibles, maxille, and labium with both pairs

of palpi are themselves in almost constant motion, but, so far as

observed, they play no part as organs of sensation, with the possible

exception of the maxillary palpi. Sometimes the male rubs but

one of the flagella of the female, turning the head to one side.

No other movements than these are observable, but there is some

variation in the occurrence of either of the two movements

described, and also in the number of times they are repeated

before sexual union is permitted by the female. The male may

be received coldly; he may make the movements without

attempting union, or after alternating them three or four times,

he may attempt union without success, and then run forward to

repeat the actions, and this may continue as long as the female

permits, either resulting successfully or unsuccessfully. In the

presence of other females, if received coldly, the male soon tires,

leaves and seeks another mate. In order to attempt union, the

male has to reverse his position, and run back to the tip of

the abdomen of the female, where he usually reaches over the

tips of the wings and senses with the antenne, quickly turning

924 MR. A. M. ALTSON ON THE LIFE-HISTORY AND

and reaching around again with the tip of his abdomen to gain

entrance into the vaginal orifice. Or, on the other hand, he may

simply back quickly to the caudal end of the female and attempt

union. In one case,.... coition lasted for fourteen seconds,

in another for ten seconds. Mating is promiscuous for both

sexes.” (6.)

During the breeding operations, when females were scarce—

being occupied with host puparia,—the writer has frequently

observed an unresponsive female crawling about with several

males upon her, the lower ones holding on dorsally and laterally,

and those above holding on to each other. Sometimes the female

with this load on her would try to climb the side of the cage.

This effort always ended in the whole party falling to the bottom.

On three specific occasions the number of males thus congregated

numbered 10, 7, and 11.

The males have some curious habits. When the puparia from

which adults have commenced to emerge aie lying unburied on

soil, a male can be observed to take possession of one, and either

stand on it waiting for a newly-emerged female to appear, or he

will enter it and periodically make an appearance to seek a mate.

Generally the males remain in the immediate vicinity of the

puparia from which emergence is taking place, and running over

and avound these in search of females, they are continually

meeting each other ; then they start to fight. This usually takes

place by means of the antenne and fore legs, resulting frequently

in the loss of a few joints of the flagella by one or both antago-

nists. During these scuffles there were sometimes three or four

participants, and then it frequently happened that a puparium

became dislodged, with comical results if it was occupied by a

‘domesticated ” male: he would sally forth and attack the first

within reach; generally one in no way responsible for his dis-

conifort. From a short distance the whole area occupied by the

males seemed to consist of combatants.

Females only resorted to fighting when disturbed during

oviposition ; seldom did males venture into the dishes containing

host puparia for parasitization.

Flight.—Uhe females, in spite of their large wings, are only

capable of flying short distances, in a very jerky manner, about

6 feet at a time at most. Before starting on a flight, they usually

indulge in a few preliminary movements of their wings. Their

most customary method of progress is to crawl.

The males are quite incapable of flight, and the only occasions

on which they have been observed to use their wings are when

courting and fighting.

Oviposition.—This takes place within a host puparium, prefer-

ence being shown for those between 24 to 72 hours old—that is to

say, after histogenesis has started and the developing fly-nymph

is covered in its pupal skin and free of the puparium except

where attached anteriorly and posteriorly by the trachee. A

female will not oviposit in a puparium containing an atrophied

HABITS OF TWO PARASITES OF BLOW-FLIES. 225

fly-nymph, neither one containing a fully-formed fly which for

some reason has died or failed to emerge, nor will it oviposit in

an empty puparium from which the occupant or occupants have

emerged, although efforts to induce some to do so were made by

burying that portion from which emergence took place; its

behaviour with such puparia is initially the same as with a

healthy puparium, but after examination and possibly an attempt

to insert the ovipositor, it crawls off in search of others. ‘To

enable it to distinguish between those puparia which contain

sustenance for its progeny and those which do not, the female

must not only possess very delicate sense-organs, but apparently

uses the point of the ovipositor, as will be shown later.

Before attempting to insert the ovipositor, a female spends

some time critically examining the puparium, crawling over and

around it, with her head ‘aclinet towards it, constantly waving

the eee with which she frequently touches it. Hewines

apparently, satisfied herself that it is a fit object for attack, she

bends the abdomen so that the apex touches the puparium, and

then with the tactile hairs upon the palpi and abdomen, and with

the point of the ovipositor, she proceeds to prod it until a position

is located—such as a groove in the contracted integument of the

puparium—through which to make a puncture. The ovipositor

is then held in the minute groove, and the apex of the abdomen

springs back to a position in which it assumes almost its natural

shape, although diagonally opposed to the puparium, and in doing

this the full ‘length of the ovipositor is exposed (text-fig. 18);

Text-figure 18.

Female WV. brevicornis ovipositing; 1st ES Greatly magnified.

Original.

this has hitherto been hidden in its recess along the ventral plates

of the abdomen. The female now endeavours to pierce the

puparium—not always with success, in which case she moves off

to another spot and repeats the process,—which she appears to do

with a slight rotatory and up-and-down movement of the ovi-

positor, aceom panied with frequent twitching of the antenne, a

constant movement of the trophi, a slight twitching of the apex

of the abdomen, and a general appearance of slight 1 movements as

if her entire strength was being exerted upon the task. As the

ovipositor gradually disappears into the puparium, the position of

226 MR. A. M. ALTSON ON THE LIFE=HISTORY AND

the hind-legs is sometimes changed, and they are placed further

apart so as to lower the body in rhythm. During this movement

the abdomen gradually assumes an extraordinary lateral aspect

until, when the ovipositor is completely inserted, its appearance is

similar to that depicted in text-fig. 19. The flexibility of the

sternites is remarkable. Sometimes, after the ovipositor has

Text-figure 19.

Ate \)

Li ” iii

Female WN. brevicornis ovipositing; 2nd position. Greatly magnified.

Original.

My G

Ys %,

been inserted its full length, it is partly withdrawn, and appears

to be moving slowly in a circle, giving the impression that at this

time the eggs are being discharged so as to lie together in a

cluster but not on top of each other. Eventually, by lifting the

abdomen till it recovers its position as shown in text-fig. 18,

and then lifting the thorax with the hind-legs, it disengages the

ovipositor, which instantly springs back to its normal hidden

position.

Generally a minute drop of clear liquid appears at the point

of insertion of the ovipositor directly it is withdrawn, but the

female moves backwards over the hole until it is located by her

palpi or antenne, and then appears to “suck up” the liquid

The drop of liquid is not always visible, and the nature of it has

not been determined.

One observer says :—‘‘ This liquid is probably used in the first

case as a lubricant by bathing the styles as they work on the

sheath while puncturing” (8). Another :—‘“ The liquid is either

a fluid resembling in its nature a synovial fluid, or else it had

been acting as a lubricant for the styles; the former is the more

probable ” (7). Another observer says :—‘ In many cases it (the

hole) became covered with a white mycelium-lke growth the

nature of which we have not determined” (6). That the fune-

tion of the liquid is to seal the hole seems probable. Only in

three instances amongst material parasitized in the open have

mites (undetermined) been found inside intact puparia inhabited

by developing Chalcids, and in one case a pupa had been partially

devoured.

The time occupied in ovipositing 1s variable, ranging from a few

seconds to half-an-hour. A few specific instances are quoted :—

“The deposition of an egg observed at 9.45 pP.m., Sept. 14,

required 16 minutes; the host was Phormia regina. Another

HABITS OF TWO PARASITES OF BLOW-FLIES. 227

observation made at i 15 a.m. the same day showed that the act

required 8 minutes; the host puparium was that of Musca

domestica;.... A female confined at 9.20 A.m., Sept. 10, deposited

into puparia of the Phormia at 9.32 a.m.and 1.20 p.m. the same

day. One confined at 10 a.m. the same date with two puparia ef

the same host oviposited at once.” (6.)

The position of insertion of the ovipositor is very variable.

Any segment laterally, dorsally, or ventrally is attacked, in

captivity.

One female may insert her ovipositor more than once in the

same puparium. One puparium may be attacked by several

females in turn; sometimes two will be seen at work at the

same time.

Length of Period of Oviposition.—When amply supplied with

host puparia, the females appear to live from 3-4 weeks. The

first generation from material obtained from Regent’s Park

were still actively at work, whilst their own progeny were

emerging and had begun attacking hosts.

Pime alapsing between Hmergence and Production.— When host

puparia are available, oviposition takes place within « few hours.

In three specific cecenecs observed a Girault and Saunders

oviposition took place in 24, 103, and 3 hours after emergence.

Progeny of Single Females The number of eggs deposited by

a female varies. ‘In three instances one female placed in a tube

with fifteen pupz parasitized the whole, with the exception of

two pupe which had decayed. The total number developing

from the one parasite in the first case was 140, in the second 148,

and the third 96. (The numbers emerging from each pupa varied

from 15 to 1 in these experiments.)” (8.)

In the case of two females each confined separately with host

puparia, the first with 130, the second with 86, their respective

progeny numbered 21 males, 57 females, total 78, and 38 males,

65 females, total 103. The first female parasitized 17 puparia,

the second 22 (6).

Parthenogenesis. —‘On September 27th, 1908, 12 virgin females

of brevicornis, reared separately from puparia of Phormia regina

and in no instance accessible to males, were confined separately,

each in a small gelatine capsule with a single known healthy

puparium of Phormia; on September 29th at 11.30 a.m., in three

cases females were observed ovipositing; on October 15th, 1908,

the progeny of two of the virgin females emerged as follows :—

17 males....and 15 males..... Other emergences did not

occur, but in three instances the larvee of the parasites were

found in the host puparia, all dying, however. Hence this

parasite is parthenogenetic.” (6.)

Proportion of the Sexes.—-Under this heading Girault and

Saunders tabulate the numbers and sexes of 7369 specimens dealt

with by them. In lot No. 3 (of this table) they bred 710 males

and 786 females; these were from puparia collected “from a

single host lot from a decomposed cadaver, city dumping-

Proc. Zoot, Soc,— 1920, No, XVI. 16

228 MR, A. M. ALTSON ON THE LIFE-HISTORY AND

grounds.” In lot No. 5 they bred 228 males and 116 females

from ‘ host puparia in fecal matter, miscellaneous.” These lots

represent the most complete figures for material parasitized

under natural conditions, and clearly indicate parthenogenetic

reproduction.

Length of Life of Adult.—As previously stated, females kept

amply supplied with host puparia and a little food lived from

3-4 weeks. ‘The males confined with these females only live

7 or 8 days.

In Australia, under similar conditions and in the cooler

weather, they are stated to live from 4 to 6 weeks (7).

When confined in glass eylinders without food or host puparia,

they were found to live from 4 to 6 days, the males dying

first (6 and 7), and when confined in glass cylinders with food and

no host puparia, they were found to live from 18 to 20 days (7).

“The difference in the length of the life of the wasps kept in a

confined space, and of those active ely at work in the cages, may be

due partly to the direct effects of confinement; but in the writer’s

opinion, it is due more to over-copulation of the females in the

confined spaces” (7).. This may be partly accountable, but the

females appear to be quite capable of resisting the advances of

the males, and the writer suggests that the undetermined drop

of liquid which sometimes exudes from the punctured puparium,

and which all observers agree 1s generally ‘‘ sucked up” by the

female, may contain nutriment. This would explain why those

confined with food and host puparia to deposit eggs in, live longer

than those with food and no opportunity to deposit eggs. This

view is further supported by the fact that (under breeding

conditions) the females seldom left the dishes containing host

puparia for food.

It has, however, been ascertained that individuals of both

sexes can live for some time with very little food. In the case

of three females (presumed to have mated) each was confined

separately in tubes within 14 hours of emergence; and into each

tube a minute drop of food was placed on the glass by the aid of

a needle; no further supply was given. The first female was

dead on 15th day, the second on 19th day, the third on the 21st day.

Of three males confined separately in tubes (none of them had

had an opportunity to mate) and given a similar initial drop of

food, one managed to escape on the 7th day through a fissure

in the cork, another died on 14th day, and the last on 22nd day,

No test was made with virgin females.

Food.—In the laboratory the same food as was used for the

braconids and flies was given to these, but very infrequently and

only a very little at a time, otherwise they would get stuck in it;

and if their wings came in contact with it, they seemed quite

incapable of extricating themselves. How the semi-apterous

males obtain food under natural conditions is not known; and

taking into consideration their habit of remaining in the imme-

diate vicinity whence they emerge, it is very probable that most

HABITS OF TWO PARASITES OF BLOW-FLIES. 229

never obtain any. The females have a better chance with thei

short jerky flight.

Hifective Parasitism.— Under this term Girault and Saunders

refer to an interesting phenomenon, and one which was also

observed by the writer. It is, that the attack of Vasonia can be

effective upon the host pupa within 15 to 24 hours prior to the

moment when the adult fly would have emerged, the length of

the period being approximately such time as is required for the

depositing, dev elopment, and hatching of the egg, and which in

the instances observed by the writer would be from 30 to 74 hours.

In a specifie instance, and one which must be about the extreme

limit of effectiveness, a puparium was noticed to be cracked

along the crease at the anterior end, but not sufficiently to have

allowed a fly to emerge; it was opened, and found to contain

seven larve feeding upon a perfect pupal fly which had cracked

its enveloping skin anteriorly and the puparium, but which had

failed to get any further, its organ of locomotion—the ptilum —

apparently having lost its efficiency gradually as the larvee fed.

Percentage of Parasitization.—The writer has no figures rela-

tive to the percentage of parasitization under natural conditions,

and the data relative to laboratory conditions cannot be taken as

a true index of the capacity of Vasonia, as the number of host

puparia presented for attack were worked out on the basis of

20 per female, and the numbers of the latter m the breeding-jars

or main cage were only approximately ascertained.

Upon oierenee to Girault and Saunders the following is

found :—“ Further, the local abundance of this parasite is ind1-

cated by the fact that in at least a portion of the experiment just

mentioned, a portion selected at random, the percentage of

parasitism was as high as 90 per cent. We have evidence to

show, on the other hand, that this percentage of mortality of the

host was by no means general, but was considerably lower on

the average for this season of the year.” (6.)

Hilhomnati ar —This parasite hibernates as full-grown larve

within the puparia of its various hosts. _ It emerges in the spring,

the earliest recorded emergence, in England, being the end of

April (4).

Attraction to Light.—The females are freely attracted to artificial

light, the males very slightly.

Hosts.—‘The different species of puparia given to Wasonia

brevicornis to parasitize, and from which it was successfully bred

are as follows :—

Musca domestica Linn.

Calliphora erythrocephala Meig.

is vomitoria Linn.

Phormia grenlandica Att.

Lucilia cesar Linn.

» sericata Meig.

16%@

Tsu) MR. A. M. ALISON ON ‘THE LIFE-HISTORY AND

The hosts reported from America by Girault and Saunders (6)

ae -—

Musca domestica Linn.

Calliphora erythrocephala Meig.

Phormia regina Linn.

Lucilia cesar Linn,

» sericata Meig.

Chrysomia macellaria Fabry.

Cynomyia cadaverinia Desv.

Sarcophaga species ‘‘ K” (Spee. nova).

The hosts reported from Australia by Froggatt, Jun. (7) are as

follows :—

Pycnosoma rufifacies. “ Mostly in the field.”

ws varipes. ‘*To a lesser extent in the field.”

Anastellorhina augur. :

Pollenia stygia. (

Lucilia sericata. ‘In the laboratory.”

Calliphora erythrocephala.

Sarcophaga aurifrons.

SUPER-PARASITISM, OR ACCIDENTAL SECONDARY PARASITISM.

That Nusonia brevicornis might be a “ Hyperparasite” upon

Alysia manducator originally oceurred to the writer, when the

second lot of puparia were collected from the spot at Regent? s Park

whence the original Braconid parasitized material was obtained,

and when it was found that this Chalcid was emerging from it.

However, no Braconid lining to the puparia was obser ved, at the

time, in any from which the Chalcids had emer ‘ged; and as the

initial breeding operations showed that the latter-—then undeter-

mined—bred freely from healthy host puparium, it was assumed

that “hyperparasitism ” was not the rédle of NV. brevicornis ;

although it appeared reasonable to suppose that the Chalecid

might act accidentally as a secondary parasite in the early stages

of development of the Braconid—that is, during the first or

second instar—and when the greater part of the fly-nymph still

existed. That secondary parasitism in this early stage could

take place appears probable; and it does not require a great

stretch of imagination to conceive the race for life within a ;

. : . . -

puparium, in the fly-nymph of which an early instar Jarva of the :

Braconid is slowly feeding, when suddenly one by one a dozen, §

more or less, Chalcid larvee commence feeding operations from

the outside of, and through the enveloping integument of the fly-

nymph. That the result of such an unequal contest would be

against the Braconid, it seems only reasonable to assume; and

that the latter would eventually be killed by the Chalcids biting

into it. No efforts were made to investigate this point; it

nppeared too obvious a potentiality, and to soon the remnants

IABITS OF TWO PARASITES OF BLOW-FLIES. 231

of a fly-nymph for evidence of the existence of the Braconid of

which the chitinized head-capsule of the first instar or mandibles

of the second would be the sole initial evidence of its existence,

weuld require time, which the writer at that time could not

spare.

Later—in October—a reference was found to this subject,

Graham-Smith (1) on pp. 532—4 and on p. 537, in which another

Chaleid, MJelittobia acasta Wk. (9), is definitely stated to act as a

hyper- parasite * on A. manducator, and a passing reference to

NV. brevicornis occurs and to the existence of numbers of then

undetermined Chaleid larvee in Braconid-lined puparia. Again,

in November, further reference (Graham-Smith) (4) was found to

this subject, and this time referring to the presence of JV. brevi-

cornis reared from puparia with the silk lining of A. manducator.

This observer states :—‘ Puparia collected in ube autumn of 1916

and kept outside yielded in May 1917 38 males and 36 females.

Some of these autumn puparia had been parasitized by A. mandu-

cator..... Some of these autumn puparia remained intact, and

were dissected in March 1918. Of these, 28 were found to

contain living Chalcid larve, and were kept in tubes in a warm

room. Nineteen of these 28 showed infection with A. manducator.

Adult WV. brevicornis, 11 males and 48 females, emerged in May

1918, a year and a half after the puparia were collected.”

The writer therefore decided to carry out an investigation into

this mmportant matter; and as the subject is of considerable

interest, the various experiments are given in detail.

To begin with,it might be mentioned that the point previously

referred to—i. Cs, secondary parasitism in the early stages of the

Braconid larve—was not investigated, for the reason stated.

The following two experiments were made to verify the above

quoted statement :—-

KHeperiment No. 1.—A large tube 6 inches x 1| inch was used.

Into this was placed—on 25th November—one normal size Calli-

phora puparium, in which a small fracture of the pupal case was

made so as to disclose the Braconid lining, and therefore denoting

the presence of either a full-grown larva, pro-pupa, or pupa.

Three fertilized and recently-emerged females of WV. brevicornis

were then released in the tube, and on the inside of it a small

quantity of focd was smeared. The tube was kept under occa-

sional observation for four days, during which time none of the

females were seen to oviposit, but could be seen crawling over

the puparium, carrying out the usual critical examination.

After 29th November no further observations were made. On

17th December—21 days later—the tube was opened; the three

females were dead, and the puparium was then carefully opened

under binoculars. In it was found one active full-grown Chalcid

larva, two dead and dried-up larve apparently nearly full-grown,

and two small shrivelled masses of newly emerged larve. The

* There appears to have been nothing else in the tins where this oceurred except

puparia containing individuals of A. manducator.

Zon MR. A. M. ALTSON ON THE LIFE-HISTORY AND

Braconid was found to be in the full-grown Jarval stage, and was

only partially dried up. In endeavouring to move the active larva,

the integument was punctured by the needle, and its unvoided

waste instantly exuded. ‘This caused the writer to form the

opinion that the four dead larve of the Chalcid might possibly

have met their death by a similar agency—the sete and sensory

hairs (text-fig. 5) on the integument of the full-grown Braconid

larva.

Hxperiment No. 2.—Another examination into this point was

therefore carried out. On 30th December four normal-sized

puparia, which when fractured disclosed the Braconid lining, were

placed separately in glass tubes Nos. 1-4, and into each two

fertilized Nasonia females were admitted, fmid a smear of food

made on the inside of each tube. They were then corked.

On 31st December—the next day—tube No. 3 was found to

contain an active d. manducator male. The male was released

in the Braconid cage, the Vasonia females were replaced by new

ones, and another puparium—a small one—obtained and put into

tube 3. Occasional examination of the tubes was maintained,

and, when necessary, fresh smears of food made. On 9th January

both females in tube No. 2 were dead. On 10th January, tube

No. | was found to contain an active A. manducator female; the

Nasonia females were still active. A new puparium was put into

the tube, and that from which the Braconid female had emerged

was examined. Notbing bearing any resemblance to Chalcid larvee

or eggs was found ; the pupariam contained the Braconid larval

exuvium, pupal exuvium, the meconium, and white cement-like

discharge. On 11th Febr uary tube No. 4 was found to contain

an active female Braconid ; both Vasonia females were dead. ‘The

puparium was examined, and besides its normal contents, two

small masses were found adhering to the lining but which were

not distinguished. The Vasonia females in tubes Nos. | and 3

were observed to be dead. On 10th March the contents of tubes

1, 2, and 3 were examined. No.1 puparium was opened, and

found to contain a dead and shrivelled larva of A. manducator,

and attached to it was a dead Chaleid larva, and loose in the

puparium two living Chalcid larve. No, 2 puparium contaimed

a dead and shrivelled larva of A. manducator with four dried and

shrivelled Chaleid larvee—very small ones—and all adhering to

their host. The puparium in No. 3 tube was examined, and

towards the cephalic ventral end of it a slit was observed, through

which part of the head of an adult Braconid could be seen, as

though it had tried to emerge. It was then observed whilst

opening the puparium that, at the point where the writer had

made a fracture mid-dorsally in the first place, this had either

been carelessly done, or the female—for such it was—had extended

it in her efforts to emerge when bringing the necessary pressure to

bear upon the cephalic end in order to get her mandibles into it,

as she would cause the anterior half to bend with her, so that it

acted as if hinged, and thus frustrating her efforts to emerge.

HABITS OF TWO PARASITES OF BLOW-FLIES. 230

No signs of Chalcid infestation was observed. The female may

have died within a few days of the start of the experiment.

The foregoing experiments show that, under certain conditions

—-and with fractured puparia-—Vasonia can super-parasitize

A. manducator \ined and inhabited puparia, and in two

instances three larve reached full growth.

Whilst experiment No. 2 was developing, experiments into

other directions were undertaken.

It has been previously stated that blow-fly Jarve upon

recovering from “ temporary paralysis” are stimulated to pupate

and to escape from the females of 4. manducator. It was there-

fore decided to ascertain to what depth such larvee worked their

way into the soil.

Experiment No. 3.—A wooden box with a sliding lid was

requisitioned. One end was removed, and the sliding lid sawn

across into strips an inch wide. Thus when placed upright with

the remaining end as the base of the box, the original base

formed one side and the sliding lid in strips the other, with the

original sides as ends.

This box was then gradually filled with soil which, during the

building up of the requisite depth, was twice subjected to running

water to damp it. Seven inches of soil were eventualiy put in.

On the same day—9th January, 1920—pieces of liver upon which

fly-larvee had been feeding were placed on the surface of the soil

in a compact mass. One female A. manducator was then caught

in a tube from the main cage. ‘he mouth of the tube was then

held over the liver until the female was stimulated to descend

upon it. As soon as she got on to it, she started examining it for

larve. 25 of these, about three-quarters to full grown, were at

hand; they were placed near the female, one at a time, so soon

as she had oviposited into each of them. In three instances she

attacked twice, having failed to come in contact with the sub-

stitutes, and in each case the latter were withdrawn and given

to her a second time. It took 57 minutes to get the 25 larve

parasitized. The female was moved back into the tube and

given a smear of food on the cork. She was allowed to rest in

the tube for 20 minutes. In the meanwhile another 25 larve

had been collected. The female was now released again, and the

same process started, but she only attacked one of these larvee ;

and although others were placed near her, she could not be

induced to attack any more, and after 26 minutes the experi-

ment was given up for the day. The remaining 24 larve of the

second lot were returned to their breeding dish, and the female

caught and left in the tube with food.

No further action was taken for the next two days. On 12th

January, 1920, the experiment was continued. The above female

was found dying; another was therefore removed from the cage,

and at 2.30 p.m. it was set to work on 25 larve as before, but

at 3.25 p.m. it suddenly became dark, and the female refused to

oviposit any more. Jn this time—55 minutes—she had dealt with

234 MR. A. M. ALTSON ON THE LIFE-HISTORY AND

20 larve. The remainder, five, were returned to their dish, and

the female placed in a tube. The surface of soil was on this date

quite dry.

On 14th January the contents of the box were examined, but

in removing the first inch strip of the “side,” some of the dry

soil fell out and the dead larva with it (this is recorded as 3"

in the table); greater cave was then exercised in removing the

other strips. The result is shown for convenience in Table LV.

Taste LV,.—-Depth of Soil penetrated by parasitized larvee.

| [eae

Dates! iin lal lites. |e 22 Sale arerameeialime fltas h aeepee 7

| | | | | |

(nya taeneeenn ee Blea NS ath Gal Ne Mesa cele | 39

| | | | | a

Motaly oe esis Mole 01) Ghee AeaeerEl O.) |. 1Og| cota al oneke

* Dead. + 1 dead.

When removing the soil, which was done by gradually scraping

it off with the edge of a steel rule, it was found that at 2? inches

down the soil was caked and still damp. This coagulation was

no doubt due to running the water into the box on very dry soil.

Seven of the puparia were broken in scraping the soil off.

Haperiment No, 4..—Another experiment of the above descrip-

tion was carried out, but in this instance the soil was spread out

into a tray and damped before putting it into the box. The

same wooden box was used, and filled with this soil to a depth

of 7 inches. The experiment was started in the morning of

22nd January, 1920. Pieces of liver were placed on the soil,

and one female 4. manducator allowed to emerge from a tube on

to it; 25 larve were then given separately for oviposition. ‘The

female effectively dealt with these in 45 minutes; she was then

removed and another female taken from the cage. The second

female was also given 25 larvee, and disposed of these in an hour

and a quarter. Over-parasitization was observed to take place

in one larva in this experiment.

On 27th January, 1920, five days later, the contents of the box

were examined by the same process of scraping the soil off; it

had not coagulated in this case, and only two puparia were broken.

The result of the experiment is given in Table V.

Table V. shows that 19 larvee went deeper than any in No. 3

experiment, there being an absence of caked soil.

Both experiments Nos. 3 and 4 showed that the majority of

Braconid parasitized larvee reached 2 inches or more under the

surface of the soil.

Three experiments were made to ascertain the burrowing

capacity of WV. brevicornis. Nos. 5 and 6 were started before the

Pee ey ee ee

HABITS OF TWO PARASITES OF BLOW-FLIES. Dia

Taste V.—Depth ot Soil penetrated by parasitized larvee.

| cal Sok Ea |

Depth ...j4°")3" 2" 1" 14" }13" | 12" |o"! 92" 9x" | oa" lanl gun | gam | gar igi 40" | Totals.

oe | I |

arved.!.., —|—3* 3}, — | 1 —]1lj/— —] 1 fif—}ir}—-r-—)

| | |

Bupa sleet te | led A 8 ke |) Ee Py al 39

(ie -| hea | -- | ae

(opalSieercs eae Been 1 | ey ea ay iy ee teh aie zn 50

* 2 dead. + 2 dead.

writer was in possession of the information gained in experil-

ments Nos. 3 and 4.

Hxperiment No. 5.—A cylindrical glass jar was used. It was

filled with 5 inches of damped soil ; on this was placed a piece of

liver with some larve feeding upon it. They were mostly full

grown. ‘This experiment was started on 20th December, 1919.

On 22nd December it appeared that the larve had ceased feeding

and had gone into the soil to’ pupate. The liver was therefore

removed, and under it two puparia were found; no others were

visible. The liver was replaced. 18 fertilized domme lee of Nasonia

brevicornis were then released in the jar, which was at once

covered with bolting silk, and this smeared with food. ee

Ist January, 1920, adult C. erythrocephala were emerging ; 1

Chalcids could be seen, so no further supplies of food were given.

The flies were left to die, so that nothing should be ‘stircled:

Adults continued to emerge up to dth January ; afew days later

they were all dead. On 10th January, as no living adults could

be seen, neither flies nor parasites, the two puparia on the

surface were removed, and the contents of the jar emptied and

sorted. No attempt was made to gauge the depth to which the

larve had crawled. The jar was found to contain 47 empty

puparia and an equal number of dead flies. 14 intact puparia

were found; these were opened, and yielded four puparia con-

taining Chaleid larve; the remainder consisted of dead fully-

formed flies or atrophied fly-nymphs with no sign of Chaleids.

The two puparia from the surface were filled oath Chaleid larvee.

Of the Chalcid females, only 15 were collected ; the other three

were not searched for a second time in the soil.

Experiment No. 6.—This was another burrowing test, in which

20 puparia were placed at various depths ranging from two at

5 inches to seven on the surface, and were situated near the glass

side so as to be visible to the observer. Thesoil used was very dry

and lumpy, with a certain amount of fine dust due to the crumpling

of the lumps. Four females and one male were released in the

jar. Mating was observed. The experiment started on Ist

January 1920; and on 20th February, when the contents were

emptied and the intact puparia opened—there were only four of

these, flies had emerged from the rest,--none bore any signs

236 MR. A. M. ALTSON ON THE LIFE-HISTORY AND

of Chalcids. ‘This experiment, therefore, was a complete failure.

The only point of interest was that the two flies placed at a depth

of 5 inches reached the surface.

Experiment No. 7.—Vhis was also a burrowing test. It was

started on 30th January, 1920. The same jar was used as in

No. 4. It was filled with 2 inches of damped soil, and on this

93 newly formed puparia were placed; above the puparia 2 inches

of damped soil was run in. (The writer was now in possession of

details of tests 1 and 2.) On the surface of the soil 12 more

puparia were scattered, and a piece of liver upon which larvie

had been feeding was also put in. The liver covered five of the

surface puparia. The liver was used to reproduce, on a small scale,

natural conditions. 20 fertilized females were released in the jar

on the same day; it was covered with bolting silk and smeared

with food. The jar was kept under observation. From 31st

January to 9th February some of the females could be seen at

work on the surface puparia. From 10th February to 13th they

appeared to be dying. On the latter date C. eryihrocephala began

to emerge, and continued to do so for several days. On 18th

February all flies were dead. On 19th February the contents

were examined. The 12 surface puparia were opened, and all

found to contain Chalcids in advanced or early pupal stage, and

a few as larve. 72 dead Calliphora were found on the surface.

The 93 puparia at a depth of 2 inches below surface were then

turned out, and 72 empty puparia were found; the remainder,

21, were still intact, and were opened and examined. No signs

of the Chalcids were found; the contents of these intact puparia

were dead flies in various stages.and atrophied nymphs.

Experiments Nos. 4 and 6 indicate that female Vasonia, even

when confined in a small space with host puparia covered with

soil loosely run in, are incapable of burrowing toany extent; and

in this connection the following observation of Froggatt (7) 1s

quoted :—“ It is generally amongst the pupe of Pycnosoma

rufifacies, and to a lesser extent P. varipes, that this species of

Chalcid wasp is found actually at work in the field. This is

largely due to the habits of the larve of these two species, which

do not crawl away from a carcase to pupate unless the remains

have been much disturbed, but pupate either just under the edge

of the remains or else affix themselves to the bones, wool, or other

portions of the carcase, and thus they are more easily found than

the other species, all of which generally crawl a considerable

distance from the carcase, and scatter...... Nodefinite reason

can yet be assigned to the apparent distaste of the wasps for the

pup of Ophyra nigra and Pycnosoma varipes in the laboratory.

In the field the pups of the latter species are practically always

found to be parasitized.”

Further, Vasonia does not bear the appearance of a burrowing

insect ; its head, viewed dorsally, is wider than any other fart

of its body, whereas Melitiobia acasta females (specimens of

which Mr. Waterston kindly gave the writer) have a narrow head

HABITS OF TWO PARASITES OF BLOW-FLIES. 237

sloping backwards ventrally, and which is attached to a tapering

thorax, narrowest at the back of the head, and certainly better

adapted for burrowing in loose soil.

The last point investigated was an examination of all the

puparia contained in two jars, of parasitized material obtained

in the open. ‘The contents of these jars, it should be explained,

were the result of exposing meat on soil ina glass receptacle at

the Society’s Gardens, with the object of obtaining additional

supplies of the Braconid. ‘The collection of the contents of this

receptacle was delayed until 14th August, and when sorting out

the intact puparia from those from which blow-flies had emerged,

the writer came across a few dead female Vasonia as well as dead

female A. manducator. It was therefore assumed that a double

infestation had taken place. At the time, an attempt was made

to differentiate between the puparium of one or other of the

parasites, but both were in too early a stage to disclose any

difference when subjected to transmitted light. The material

was consequently left unsorted, and emptied into two jars to

await developments. Vasonia began to emerge on 31st August,

and in order to give these an opportunity of escaping as they

emerged, the linen covers of the jars were replaced by pieces of

mosquito netting, through the mesh of which the females were

able to escape. Both jars were placed inside a large glass cylinder.

This was used as the cage, and in it were put dishes with supplies

of blow-fly puparia for the Chalcids. On 3rd September

A. manducator began to emerge; the Chalcids were still doing so.

This continued until 29th September. This state of affairs neces-

sitated constant handling of the jars to release the Braconids, so

that the contents beeame disturbed; and although it was not

observed at the time, it now transpires that some of the Vasonia

females never left the jars, but proceeded to super-parasitize the

Braconids in their pupavia, and, further, some of those—the

progenitors of the above—found dead in the receptacle when

the material was collected probably did the same.

When all the puparia had been collected from the jars, they

were sorted under three headings:—(A) Those from which

A. manducator had emerged, (B) those from which Vasonia had

emerged, and (C) those which were intact. Those from which

blowflies had emerged had already been removed, and the number

is not known. The numbers under the three headings were :—

(A) 195, (B) 50, (C) 506. Those under (A) were put aside, those

under (B) were dissected for evidence of successful super-para-

sitism, and those under (C) were dissected and classified under

several headings as follows :—

(1) Atrophied fly-nymphs. )

(2) Dead fully-formed flies. |

B Zeerrcoud seg pe signs of Chalcid attack.

? 9 f

(5) ‘ my adults. |

(GO) Actives \, larve. J

238 MR, A. M. ALTSON ON THE LIFE-HISTORY AND

(7) Unlined puparia, with active Chaleid larvee. (These had,

superficially, fly-remains.)

(8) Braconid lined puparia, with dead Chaleid lanvee. (The

Braconids were in all stages, pup predominating.

(9) Braconid lined puparia, A naiine Chaleid larve. (These

Braconids were in all stages, pupee predominating. )

(10) Braconid lined puparia, with some dead Chaleid imagines

and larvee. (These consisted, with one exception, coe at

least one male.)

The examination of these puparia showed that where super-

parasitism had taken place, with the exception of a very few

instances-—the numbers were not noted,—the puparium concerned

was a very small one, about the size of J. domestica, aud con-

sequentiy, in the case of a blow-fly, a puparium with an exception-

ally thin integument. And, further, that the attack of the

Chaleid larve had taken place in the abdomen of the Braconid

where this was an advanced pupa, the only part where soft

integument exists, due to the distention of the Bee

consequent upon the presence of the meconium (text-fig. 7) :

therefore, seems conclusive that a Masonia female is able .

distinguish between a hard surface and one yielding to pressure,

by means of the apex of her ovipositor. It is worthy to note

that Vasonia appeared to be most effective when the Braconid

was a pupa—that is to say, in the identical stage of development

to that of its normal dipterous hosts.

The numbers under the heading (B) were (1) 33 puparia, with

Chaleid exit-holes and unlined by the Braconid, presumed pri-

mary parasitism ; and (2) 17 with Chaleid exit-holes and definite

super-parasitism. In each instance the puparium had been lined

by the Braconid. A feature of these 17 puparia was the position

of the exit-hole: in 3 it was made in or about the middle of the

puparium, in 6 at the apex of the anterior end, and in 8 at the

apex of the posterior end. In only one instance had two efforts

been made to escape. This occurred in one of the 3; a small some-

what elongated aperture had been made in a position diametrically

opposed to the exit-hole used. A point of great interest in the

writer’s opinion, with regard to the position of the exit-hole in

the above, is that in 6 the Chaleid got out at a point where the

Braconid lining i is weakest-—that is, where it is run over the pro-

truding chitinized mouth- -parts of the late blow-fly larva; and

in 8 the Chaleid got out at the next weakest point—that is,

where the lining is run over the blow-fly larva’s posterior

spivacles.

The figures under heading (C) are not given in detail; those

for sub- headings (1) to (7) do not materially bear on the subject of

the investigation, but for 1 and 2 they were very high, and mainly

due, in the writer’ Ss opinion, to the unshaded position of the glass

receptacle and consequent evaporation of any moisture in the

Wiese

MRIS 2 Mere 3

HABITS OF TWO PARASITES OF BLOW-FLIES. 239

soil, which was aided by the delay in removing the contents.

Whereas the figures for (8), (9), and (10) do bear on the subject ;

they are as follows :—

(8) 69 Braconid lined puparia, with dead Chaleid larvee.

(9) 15 és 5 0 with active* Chaleid larvee.

(10) 23 5 Bi bs with some dead Chaleid ima-

gines and larve.

The outstanding point in this instance of super-parasitism is

that most of it did not take place in the open, so far as (B) (2)

and (C )) (8), (9), and (10) are concerned ; it ae took place in

the jars in the laboratory, and in proof of this postulation, the

puparia could not have been lined by the full-grown larve

by 14th August, or the adults would have emerged before the

3rd September, and further super-parasitized Braconids would

have been in the larval stage if it had taken place in the open,

whereas the contrary was the case.

To conclude: it has been shown in experiments Nos. | and 2

that under certain circumstances Vasonia is capable of super-

pavasitizing a Braconid larva within its lined puparium ; and in

experiments Nos. 3 and 4 that blow-fly larve attacked by the

Braconid are stimulated to escape and pupate, and that the

majority of them—76 per cent.—reached 2 inchés and more

below the surface of the soil; and in experiment No.5 Nasonia was

unable to burrow to the depths reached by the majority, 93 per

cent. of the unparasitized blow-fly larvee; and in experiment No.7

definitely failed to reach any one of 93 puparia 2 inches under

the surface, which is consistent with their behaviour in the field.

And, it must be pointed out, that in each instance of super-

parasitism reported by Graham-Smith (1 and 4) it appears that the

puparig bsequently ascertained to contain the Braconid

primavily—were collected and moved from a “sun” tin to a

“shade” tin or vice versa; and in the case reported by the

writer, emerging Nasonia ‘remained in the jars containing

Braconid parasitized puparia, which were no doubt dislodged De

the frequent handling of the jars when releasing Braconids. So

that in none of these instances were the conditions even

approaching natural.

It therefore appears that if super-parasitism takes place in

nature, it is not very frequent and then only accidental, and that

where it does take place in a puparium of a full-grown blow-fly

larva which has been reinforced by a normal Braconid lining,

it is doubtful whether the small, short, and rectangular mandibles

(text-fig. 20, a, b,c) of Nasonia will enable it to escape; whilst

in the case of the female Welittchia acasta, this is aided by the

distinctly acute mandible (text-fig. 20, d) with its one large outer

tooth.

* A few of these were found singly in the puparia, and are the largest Nasonia

larvee seen by the writer.

240 MR. A. M. ALTSON ON THE LIFE-HISTORY AND

REFRIGERATING EXPERIMENT.

In order to ascertain if the parasites would be able to undergo

six weeks—the length of sea voyage to Melbourne—in the re-

frigerator of a ship, a few of the parasites ete. were given a test.

Prof, Blackman was approached, and kindly gave the writer

permission to use a small cupboard in a refrigerating machine in

his department.

Text-figure 20.

Mandibles of Nasonia brevicornis male: (a) left; (b) right; (c) right, im profile,

outer tooth on top. 160. Original. (d) Mandible of female Melittobia

acasta; only females of this species emerge. After Waterston. (d) is not

proportionate with (a), (b), and (c).

As other experiments were going on at the time, it was necessary

to use a burying medium in which no insect life existed, and

clean sand was therefore used.

A large glass jar was used as the receptacle, and sand to a

depth of 3 inches was run into it as a foundation. A few

puparia containing A. manducator, N. brevicornis, Calliphora

erythrocephala, and a few larve of the latter species were each

buried in sand in separate glass tubes left open; these were then

pushed into the 38-inch foundation of sand to keep them vertical.

- The jar and contents were put into the chamber on 28th

November. The constant temperature was reputed to be 2° C.

On 15th January—6 weeks and 6 days later—the jar was

removed to the laboratory and the contents examined.

One Braconid, a male, had emerged, on what date is unknown;

HABITS OF TWO PARASITES OF BLOW-FLIES. 241

another Braconid puparium was found with a dead adult half-way

out of it, the remainder were intact; one of these was opened

and found to contain a living Braconid larva. The Chalcid-

infested puparia were all intact ; one was opened and contained

living larve. The Calliphora pupavia were intact, and as the

outline of advanced fly-pupa could be seen in each, they were left

so; the Calliphora larve were all dead. The contents, except

the latter, were returned to the jar in their respective tubes, and

it was placed about 3 feet from a Bunsen burner. On 16th

January one female Calliphora emerged, and in the next four

days all had emerged. The Chalcids commenced emerging on

30th January, and continued to do so for some days. The balance

of the Braconids began to emerge on 12th February.

To summarize: both parasites were able to stand the test when

put into the refrigerator in their respective full-grown larval

stage—that is, the customary hibernating condition.

CONCLUSIONS UPON tHE Economic IMPORTANCE OF

ALYSTA MANDUCATOR PANZ. AND NV ASONIA BREVICORNIS ASHM.

Tt is obvious that, in discussing this subject, the writer is only

able to treat of it from a theoretical standpoint. But in doing

so he is influenced by a desire to place on record some opinions

which have been formed during the course of this work, and some

of which gain considerable support from the observations and

writings of those who have dealt with these parasites before him.

It would appear that both parasites working in conjunction

would form an excellent combination, the one attacking the

larval stage, the other the pupal. And between them—it may

be assumed—a beneficial influence would be exerted in the control

of the Blow-fly.

The occasional and accidental super-parasitism on the part of

N. brevicornis is but an assurance of the maintenance of the

“Wave Law”*. It is actually a twofold assurance, because

although Vasonia may kill the Braconid, it may itself die in the

course of development or become entrapped—upon reaching the

imaginal stage—within the silk-lined puparium of the blow-fly

larva.

Individually, Alysia manducator is considerably superior to

Nasonia brevicornis as an enemy of the Blow-fly. The former is

vastly superior in flight—in both sexes—a most important factor

where artificial distribution is concerned; and it is also so from

the greater number of hosts it can injure and eliminate in the

course of breeding; the ratio is approximately 40: 1. Its egg

capacity is nearly four times greater than the Chalcid, and equal

to and in some cases greater than that of several species of blow-

flies. In length of imaginal life it is about equal to the Chaleid ;

the period of development from egg to imago is longer, and in

individuals of the same parent a considerable variation takes

* Pointed ont to the writer by Prof, Lefroy.

242 MR. A. M. ALTSON ON THE LIFE-HISTORY AND

place, but where artificially propagated, this would be a great

advantage in ensuring a continual overlapping of generations,

and so obtain the benefit of its activity throughout its season.

On the other hand, the females only of WV. brevicornis are

eapable of flight, and which—so far as has been observed—

consists of long jumps of 6 feet or so at a time, its commonest

method of progression being to crawl. ‘The males are semi-

apterous and quite incapable of flight, and although they are of a

domesticated ‘‘ temperament,” remaining in the vicinity whence

they emerge, they are very liable to overlook their natural

functions whilst endeavouring to obtain mastery over rivals, and

consequently the unfertilized females would, parthenogenetically,

produce a colony of males, possibly isolated and incapable of

finding mates or doing any damage to blow-flies. For these

reasons, it seems obvious that MNasonza cannot prove to be a

decisive factor in biow-fly control unless they are constantly

distributed in large numbers over small areas. Whereas, if they

were artificially propagated and distributed on these constant

refuse-heaps found in and around populous areas, and near

dung-heaps in town and country stables—whence to be farther,

unwittingly, distributed,— Vasonza would, on account of its

semi-social and ‘ domesticated” habits, prove a most important

factor in the control of ‘‘The Common House-or Typhoid-fly and

its allies,” which are constantly breeding in such stercoraceous

places and where host puparia would be within crawling distance

of the Chaleid.

SuMMARY.

(1) The breeding operations and accumulation of supplies of

Alysia manducator Panz. and WNasonia brevicornis Ashm. com-

menced in July and ceased in December 1919.

(2) Alysia manducator oviposits in the larve of several carrion-

feeding Diptera. Only one parasite emerges from each host.

puparia. Over-parasitism kills the larva. The mean average of

the life-cycle is 52 days, and as short as 25. Both sexes are

capable of sustained flight, and lived over a month in captivity.

Average percentage of parasitism over three years was 43 per

cent., observed by Graham-Smith (4). Average contents of

ovaries 366 eggs for 12 females.

(3) Nasonia brevicornis oviposits in the puparia of several

species of stercoral and carrion-feeding Dipterous larve. From

one to 62 individuals have been found in single puparia parasi-

tized in captivity. The length of the life-cycle ranges from 11 to

221 days in different countries. Only the’female can fly, and then

only very short distances, and can live, whilst ovipositing, from

4 to 6 weeks, but fora considerably less period without host puparia.

The male remains near the vicinity of emergence, where its life

is spent in fighting and mating. Average progeny in the case of

five females—observed by Girault & Saunders and McCarthy—

was 113 per female, and affecting on the average only 16-4 hosts,

HABITS OF TWO PARASITES OF BLOW-FLIES. 243

(4) Nasoma can act as an accidental secondary parasite upon

A. manducator if and when puparia containing the latter are

within its limited reach.

(5) Both parasites in their hibernating stage—7. e., full-grown

larvee—ean successfully withstand over 6 weeks at 2° C.

(6) Alysia manducator appears to be a more important parasite

as a natural control for the Blow-fly than Nasonia brevicornis,

which appears to be more effective as a natural control of Diptera

which constantly breed in permanent refuse- and garbage-heaps,

and where hosts would be within crawling’ distance, such as

primarily the Common House-fly, Musca domestica.

ACKNOWLEDGMENTS.

To Miss Cheesman, Curator of the Insect House of the

Society’s Gardens, Regent’s Park, the writer is indebted for

the original lots of material from which the parasites emerged,

and ‘for her assistance in procuring additional supplies of both

blowflies and parasites.

To Mr. J. Waterston, Natural History Museum, the writer’s

thanks are due for identifying the parasites. Acknowledgments

are also due to Mr. C. Gunns, Head Laboratory Assistant at the

Imperial College (Zoology Dept.) for assistance in maintaining

supplies of cages, breeding and feeding materials.

REFERENCES.

1. Granam-Smitu, G. S.—‘‘ Observations on the Habits and

Parasites of Common Flies.” Parasitology, June 1916.

2. Froeearr, W. W.—Miscellaneous Publication No. 1716.

“The Sheep Maggot-fly (Calliphora rufifacies) and its

Parasite.” Agricultural Gazette of New South Wales,

Feb. 2, 1914.

3. Marsuaut, Rev. T. A.‘ A Monograph of British Braconide,”

Part V. The ‘Transactions of the Entomological Society

of London, 1894.

4, GRAHAM-SmitH, G. S.—‘‘ Further Observations on the Habits

and Parasites of Common Flies.” Parasitology, Oct. 1919.

5. Garensy, J. Bronri.—‘‘ Notes on the Bionomics, Embryo-

logy, and Anatomy of Certain Hymenoptera Parasitica.”

Journal of the Linnean Society, No. 224, June 30, 1919.

6. Grrautt, A. A., and SaunpeErs, G. E.—‘‘ The Chaleidoid Para-

sites of the Common House- or Typhoid-fly (Musca

domestica Linn.) and its Allies.” Psyche, Dec. 1909 and

Feb. 1910. ;

7. Frocearr, J. L.—‘‘* An Economic Study of Nasonia brevi-

cornis, « Hymenopterous Parasite of Muscid Diptera.”

Bull. Ent. Research, March 1919.

. Froeearr (1915).—Extracted by Graham-Smith, G. 8. (1).

Warerston, J.—“‘ Notes on the Morphology of Chalcidoidea

bred from Calliphora.” Parasitology, vol. ix. No. 2,

Feb. 26, 1917.

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GNATHOSTOMIDA.

ON THE NEMATODE FAMILY GNATHOSTOMIDA. 245

16. A Revision of the Nematode Family Gnathostomide.

By H. A. Bayuis, M.A. Oxon., Assistant in the

Department of Zoology, British Museum (Natural

History), and Cuayron Lanz, M.D. Lond., Lt.-Coi.

EME SSiGet

[Received March 13, 1920: Read April 13, 1920.]

(Submitted for Publication by permission of the Trustees of the British Museum.)

(Text-figures 1-40; Plates I.-VIIL.*)

Family GNATHOSTOMID AE.

Cheiracanthidea Diesing (1861, p. 615).

Gnathostomide Railliet (1895, p. 1268).

Oxyuride (in part) Railliet and Henry (1916, p. 114).

Grathostonnde Skrjabin (1916, p. 972) [misprint].

Heterakide (in part) Seurat (1918, p. 25).

This family of nematodes was founded by Railliet to include

only the genus Gnathostoma Owen. Though the family-name

appears constantly in text-books on medical and veterinary

helminthology, a comparative study of the genera and species

which appear naturally to fall into cluse relationship with

Gnathostoma spinigerum, the type-species, has never, to our

knowledge, been undertaken. Such relationships have, in some

cases, been hinted at by previous writers. The genus Vanqua

was tentatively referred to the family by Leiper (1908), while

certain species of Hchinocephalus have been referred by v. Linstow

to Cheiracanthus, which isa synonym of Gnathostoma. One other

genus, Spiroays Schneider, 1866, the relationships of which have

never been decided with any precision§, we now propose to

include in this family, an inclusion which appears to necessitate

a division of the family into two subfamilies. Certain other

forms seem, from their existing descriptions, to be closely

related (see p. 305), but in the absence of opportunities for re-

examination of specimens we are obliged to be content with the

mere suggestion of their affinities.

The chief family characteristic is the possession of a pair of

large, fleshy, trilobed, lateral lips. Each lip is provided externally

with three papille, while internally its cuticle is thickened and

* Communicated by the SECRETARY.

+ For explanation of the Plates see p. 310.

{ For tamily diagnosis, see p. 247.

§ Railliet and Henry (1916) place it among the Oxyuride, with Labiduris as its

nearest ally. Seurat (1918) regards it as forming, with Camallanus, a subtamily,

Camallanine, of the family Heterakide.

246 MR, H. A. BAYLIS AND LT.-COL. CLAYTON LANE ON

frequently raised into tooth-like prominences in the form of

longitudinal ridges, which either meet or interlock with those

of the other lip.

The esophagus, in all the genera, is of a simple club-shaped

type, increasing gradually in diameter from before backwards,

muscular throughout, and without specially modified regions such

as a bulb, gizzard, or glandular portion. In some species it is of

considerable length, measuring up to about one-fifth of the total

body-length. The usual valves are present at its opening into

the intestine.

A. pair of cervical papille, usually not prominent, is always.

present in both sexes, and the tail of the female is provided with

a small pair of later. al papillee.

The male possesses more or less well-developed caudal ale, and

two spicules are always present. The latter may be equal, sub-

equal, or markedly unequal in length. In the majority of cases

they have a characteristic ornamentation.

The species of all the genera are parasitic in the alimentary

canal of their hosts, usually in its anterior part, and show a

greater or less tendency to adopt a habit of burrowing in the

tissues. Some of them are not infrequently found buried. com-

pletely in the stomach-wall, where tumours tend to be formed

round them at the expense of the host. Others, while not

penetrating to this extent, obtain a very firm hold by burying

their heads in the mucous membrane. One genus (Gnathostoma)

has been found as a rare, and probably abnormal, parasite of

man, its habitat in this case being the subcutaneous connective

tissue and not the alimentary canal.

As indications of possible affinities outside the family, the

following points are of interest :—

(1) Tn Spirosxys the caudal ale of the male are developed into

“bursa” closely resembling that of Physaloptera, in that it is

sone anteriorly across the ventral surface of the body.

(2) The genus Hartertia Seurat, 1915, appears, in some respects,

to form a link between Spiroxys and such genera as Physaloptera

and Habronema.

(3) In Gnathostoma the ventral surface of the caudal region,

in the male, is partly covered with spinous processes, which may

be compared with the low papilliform processes that cover this

region in Physaloptera.

(4) The occurrence, throughout the family, of paired lateral lips

suggests close affinity with the Spiruroidea or Spiruridz (see foot-

note, p. 247).

It may be noted here that Stephens (in Fantham, Stephens,

and Theobald, 1916) places Gnathostoma, Tanqua, and Rictularia

in this family. Wedl (1862) also placed Rictularia near the

Cheiracanthidea. Hall, however (1916), places it among the

bursate nematodes.

THE NEMATODE FAMILY GNATHOSTOMID#, 247

Family Diagnosis.

_GNATHOSTOMID® Railliet, 1895.

Spiruroidea *(?): with two large, trilobed, lateral lips, having

the etiticle of their inner surfaces thickened and usually raised

into longitudinal tooth-lke ridges which meet or interlock with

those of the other lip. Tail of male with more or less well-

developed lateral ale and two spicules. The vagina runs forward

from the vulva before giving off the two or four uterine

branches. Eggs with Me shells, ornamented externally with

fine eranulations.

Key to Subfamilies of Gnathostomide.

A. Cuticle behind the lips distended into a head-bulb

by foar submedian ballonets, a process from each

ot which extends into the body-cavity beside the

cesophagus as a cervical sac... 2. Gnathostomine (p. 254). .

B. Head-bulb, ballonets, and cervical sacs absent ......... Spiroxyine.

SPTROX YIN A, subfam. n+

These Gnathostomid are without head-bulb, cervical sacs, or

ballonets. Their lips are characteristic. A deep cleft parts from

the body the whole thickness of the dorsal and ventral edges of

each, while the pulp of the middle lobe is separated from the

pulps of the adjacent lobes by indentations so deep as to produce

an a a which has been aptly likened by Schneider (1866,

p. 125) to the club on a playing-card (PI. I. fig. 2, Pi. II. fig. 7).

Hach lobe carries a papilia, the subdorsal and Subventral papillee

having conspicuous and the lateral papille inconspicuous termin-

ations. The cuticle supporting the inner surface of each middle

lobe is greatly thickened, and projects anteriorly beyond the edge

of the lip as a sharp tooth-like prominence.

The cuticle of the tail in the male is expanded laterally into

alz and ventrally into vesicular swellings, which recall the

ventral fusion of the ale in Phy ysaloptera. ‘There are eleven pairs

of caudal papille ~, of which two pairs are definitely ventral, one

pair lying in front of, the other behind, the cloacal opening; the

other nine are more lateral. Of these lateral papillae Nos. 2

and 5 are situated more ventrally than the others; six are post-

anal and three preanal, aud in general the distension of the

* Hall (1916) makes Railliet’s superfamily Spiruroidea a synonym of Orley’s

family Spiruride, of the superfamily Filarioidea. If this system of classification is

followed, there is no group of higher than family rank, embracing nematodes with

paired lateral lips, to which the present family can be assigned. We therefore use

the name of the superfamily Spiruroidea in this sense. °

+ For subfamily diagnosis, see p. 248.

£ Nory.—Throughout our descriptions and in our figures we have numbered

the caudal papille of the male worms consecutively from the extremity of the tail

forwards—the first pair, or “‘ No. 1,” bemg that nearest to the tip.

248 MR. H. A. BAYLIS AND LY.-COL. CLAYTON LANE ON

cuticle tends to cause their terminations to lie at the bottom of

funnel-shaped depressions. Strong oblique ventral muscles occupy

the whole distance between the cloacal opening and the anterior

termination of the ale, and may depress part of the preanal

surface into the semblance of a sucker. ‘There are two subequal,

delicate, tapering spicules without flanges.

Tn the female the tail, when seen from the side, has the dorsal

surface convex and the ventral concave, and there is a notch on

its ventral surface just anterior to the tip. The vulva lies near

the middle of the body, whence the muscular vagina runs

anteriorly before dividing into two thinner-walled uteri, the one

running towards the head and the other towards the tail. The

egg-shell is thin, colourless, oval, with fine external stippling,

a ciear space separating it from.the granular contents.

The worms are found attached to, or buried in, the wall of the

stomach of tortoises in Europe, Asia, and Africa, and possibly

America.

The subfamily contains only the genus Spiroxys.

Subfamily Diagnosis.

SPIROXYINA.

Gnathostomide : without head-bulb, ballonets, or cervical sacs ;

lips wide, with a narrowed base and a trefoil-shaped pulp, and

having the internal cuticle of the middle lobe much thickened,

the point of the thickened portion projecting anteriorly as a sharp

tooth; caudal cuticle of the male expanded into lateral ale and

a preanal vesicular swelling and bearing two pairs of ventral

papille, one in front of and one behind the cloacal opening, and

nine pairs of lateral papille of which six ave postanal and three

preanal, and which, by reason of the cuticular distensions, fre+

quently appear to lie at the bottom of funnel-shaped depressions ;

spicules delicate, tapering and subequal; vulva near the middle

of the body; the vagina running anteriorly; the two uteri

opposed; ova with thin, colourless, stippled shell, from which the

unsegmented granular contents are separated by a space.

Generic Diagnosis.

SPIROXYS* Schneider, 1866.

Spiroxys Schneider (1866, p. 125).

Spiroxis Schneider (1866, p. 29) [misprint].

Spiroptera (in part) Rudolphi (1819, pp. 25 & 242).

? Ascaris (in part) Rudolphi(1809, p. 193; 1819, pp. 25 & 242).

? Physaloptera (an part) Leidy (1856, p. 53).

The genus has the characters of the subfamily.

Genotype: Spiroxys contorta (Rud., 1819).

* Hor key to species, see p. 249. For measurements, see Table I., p. 253.

THE NEMATODE FAMILY GNATHOSTOMID®. 249

Key to Species of Spiroxys.

AY Wobestof the lips umarmed|.............5.-..-c-e.--s-sesseeseeee © Se CONLOTLG.

B. Each of the three lobes of each lip carries close to

either edge of its inner surface a sharp cuticulartooth. iS. gangetica (p. 251).

1. Sprroxys conrorra * (Rud., 1819). (Text-figs. 1-3; Pl. I.

figs. 1-4; Pl. IL. fig. 5.)

Spiroptera contorta Rudolphi (1819, pp. 25 & 242).

Spiroxys contorta Schneider (1866, p. 125).

v. Linstow (1909, p. 58, figs. 22 a, 22 6).

Spiroxys contor tus Railliet and Henry (1916, p- 114).

? Spiroxys contortus (Rud.) Seurat (1918, p. 23).

@ Ascaris testudinis Rudolphi (1809, p. 193; 1819, pp. 25 &

242).

? Physaloptera contorta Leidy (1856, p. 53).

We have examined two sets of specimens of this nematode

belonging to the collection of the British Museum (Natural

History), one from Siebold’s collection (which had been removed

from the stomach of Hmys orbicularis [#. ewropea|) small,

without eggs, and presumably immature; the other mature

and labelled ‘‘from a water tortoise.” They have no specific

differences.

The worms are much twisted. Schneider describes the body

as being always bent towards the ventral aspect. This appears to

be at least usually the case, many of our specimens having an

elbow-like bend in the neck-region. The head end (Pl.

figs. 1 & 2) is very slender. Thence the diameter of the body

gradually i increases, nor does it again diminish till close to the anus.

Except for characters common to the genus the lips are unarmed.

In dorsal (or ventral) view (Pl. I. fig. 1) the tip of the dorsal

(or ventral) lobe, particularly its pulp, is hooked posteriorly,

while its anterior edge is rounded off.

In the male the caudal ale (Pl. I. figs. 3, 4; Pl. IT. fig. 5) are

well-marked; the amount of their distension, and of that of the

ventral cuticle between them, varies, but there appears to be

regularly a considerable swelling of the latter just before the ale

cease anteriorly (Pl. IT. fig. 5). The ventral papille are sessile

(Pl. I. fig. 3). The spicules (Pl. I. figs. 3, 4; Pl. Il. fig. 5) are

long, slender, tapering, and transversely striated, have the ap-

pearance of being hollow, and end in a very fine point (text-fig. 1).

They do not carry alz, as was supposed by Schneider.

-In the female the ventral caudal notch is relatively coarse

(text-fig. 2). The female organs (text-fig. 3) have the generic

characters.

In introducing the name Spiroptera contorta, Rudolphi (1819)

mentions it as synonymous with Ascaris testudinis. He had

* For specific diagnosis, see p. 261.

250 MR. H. A. BAYLIS AND LT.-COL. GLAYTON LANE ON

previously (1809, p. 198) given as his authority for the latter

name Braun (in Schneider, 1789), but expressly mentions that he

has been unable to obtain the appendix in which this name

appears. We also have failed. Since Stiles and Hassall (1905)

do not mention this name in deaiing with this genotype, we have

presumed that it is a nomen nudum and have disregarded it.

Text-figure 1.* Text-figure 2.

O:'/ mm.

O-3 mim.

Text-fig. 1.—Spiroxrys contorta. ‘Terminal portion of spicule.

Text-fig. 2.—Spiroxys contorta. Tail of female; lateral view.

Text-figure 3.

ut.

iS,

Lp ee

CET ISD

CUE EIT GSS

COUTTS

Spirowys contorta. Female genital organs. (Mature specimen.)

It is necessary to mention Leidy’s (1856, p. 53) Physaloptera

contorta as a possible synonym. His description is as follows :—

“ Body capillary, most narrowed anteriorly with the posterior

four-fifths spirally contorted; white, with the intestine brown.

Lips prominent, constricted from the body, trilobate. Tail short,

conical, acute, in the male with narrow ale each furnished with —

five funnel-shaped pores.

* Wor explanation of lettering, see p. 310.

THE NEMATODE FAMILY GNATHOSTOMID®. 251

“Leneth of the female six lines to an inch; breadth to one

quarter-of a line; male from one-Half to three-fourths of the size.

“Frequent in the stomach of Hmys serrata, Hmys reticulata,

Cistudo carolina, and Kinosternwm pennsylvanicum, adhering to

the mucous membrane in the same manner as Physaloptera

constricta” (vide infra, p. 272).

The general appearance, the shape of the lips, the “funnels”

of the caudal papille in the male, and the habitat and hosts all

suggest that this species is either identical with or closely alhed

to Sau oxys contorta, but since the ty pe of Leidy’s species cannot

now be traced, the question must remain an open one.

With regard to the Spiroxys contortus (Rud.) of Seurat (1918),

from the African tortoise, Clemmys leprosa, we tind the deserip-

tion of this form somewhat ditticult to understand. Jf our

interpretation of it is correct, Seurat’s worm differs from the

Huropean species in the following points :— ~

(1) the presence of an internal tooth on each lobe of each lip;

(2) the presence of a pair of large papille on the middle lobe

of each lip, instead of one small papilla ;

(3) the presence of an adanal, lateral pair of genital papillee

in the male;

(4) the presence of an accessory piece in the male.

Specific Diagnosis.

Sprroxys conrorva (Rud., 1819).

A slender Spiroxys; the lips without special armature, the tips

of the dorsal and ventral lobes, especially their pulp, hooked

posteriorly; spicules ending in a very fine point; caudal notch

of female relatively coarse.

_ 2. SPIROXYS GANGETICA*, sp. n. (Text-figs. 4-6; Pl. II.

figs. 6-10.)

We have examined two batches of nematodes belonging to this

species, the one collected by Mr. Southwell from a host indentified

by him as 7rionya gangeticus, and the other collected by one of

us (C. L.) from a tortoise killed in the Ganges delta, probably

Trionyx gangeticus. The two sets of specimens are specifically

identical.

This worm is stouter than the genotype. The head end tapers

much more gradually than the tail and is usually bent atan angle.

Close to either edge and near its base each lobe of either lip bears

a cuticular tooth on its inner surface (Pl. II. figs. 6 & 7).

Each lip, that is, has six teeth in addition to that characteristic

of the genus. In dorsal (or ventral) view the inner face of the

dorsal (or ventral) lobe is flattened against its fellow of the other

lip, the cuticular teeth appearing from beneath near the angles

* Wor specific diagnosis, see p. 253.

p g ’

252 MR. H. A. BAYLIS AND LY’.-COL. CLAYTON LANE ON

so formed (Pl. II. fig 6). The male has well-developed caudal

ale (Pl. II. figs: 8 & 9), the lateral papille being correspond-

ingly buried. The ventral papille are stalked and cup-shaped

or forked at their ends. The lateral papille are more widely

separated than those of the genotype. The spicules are orna-

mented with granulations tending to arrange themselves in

Text-figure 4.

0O-5mm.

Spiroxys gangetica. Tail of female: a, ventral view; 6, lateral view;

c, lateral view of tip, at a higher magnification, to show the “ notch.”

Text-figure 5. Text-figure 6.

Text-fig. 5.—Spiroxys gangetica. Female genital organs.

Text-fig. 6.—Spirovys gangetica. Ovum.

transverse rows, and each has a blunt tip surmounted by a clear

chitinous cap (PI. IT. fig. 10).

In the female the tail is relativ ely move slender than in the

genotype both in lateral and in ventral views, while the notch

near its tip and the hook formed by it are penton iy delicate

(text-fig. 4,c). The generative or eans (text- fig. 5)and eggs (text-

fig. 6) are of the generic type.

THE NEMATODE FAMILY GNATHOSTOMID.

Specific Diagnosis.

SPIROXYS GANGETIGA.

A stoutish Spiroxys; each lip carries a special armature of six

fine sharp teeth, one on either edge of each lobe;

the flattened

inner surfaces of the dorsal and ventral lobes are applied to those

of the opposite side; spicules end in a blunt point; caudal notch

in female very fine.

Taste I.—Measurements of the Species of Spiroxys.

alle measurements in Tull engiie es. 8)

S. contorta. S. gangetica.

Length 15-25 2074-30 | 39-43 48-50'3|

Thickness ...... 0°5-0°'7 0°45-0°6 | 0°75 0°85-0'95|

Lips: dorso- ventral dhametert: 011-013 0713 0711-013 013 |

Lips: length 0°05-0°06 —_-0°06—-0°07|0°14-0°16 0°16-0°2 |

Distance between striations of body . 0:008 07004 | 0°0125 0°013-0°0175)

Distance from head-end to cervical papillee... 0°65-1-0 — 1-2-1275 1:125-1: 5 |

s 5 oy WSTARESTHINNE? ‘sos con one 0'4.-0°6 0'5-0°7 \0°75—0°825 08

53 5 » end of esophagus 19-36 2°4-3°0 39 4 to 43

A 5a » excretory pore ... 0°45-0°75 0-6 0:925—-1-15 09-10

Length of tail 0°3-0'4 05-0755) 0°35 0°65

2° —

Length of spicules roach .. 2°37-3°0 as ae Se oF

Distance from tip of tail to vulva ..... — 8-2-13°6, — 20-27

_» caudal i papille (2 ) = Not seen.) — 0°32

Size of ova (maximum) JOE: — 007X005) == 0:06 0°04

Taste II.—Hosts and Distribution of the Species of Spiroxys.

a : : i

Species. | Host. | Locality. Recorder. |

mh pe eet : | ae |

es Fs | : |

» . » % | Schneider (1866).

|B. europea. | |

Emys { Vestudo | orbicularis. 4 | vy. Linstow (1909).

[B. lutaria.]

Emys { Testudo | orbicularis. 3 | Baylis & Lane

| (LE. europea. | (present paper). |

| Emys {Testudo| orbicularis. 5 ss 5 |

| (* Water-Tortoise.’’) |

| Hmys serrata. N. America. | Leidy (1856).

[ Recorded as Physa-

Emys reticulata.

Cistudo carolina.

Kinosternum pennsylvanicum.

loptera contorta. ||

S. gangetica. |

Trionyx gangeticus.

Baylis & Lane

(present paper).

uH|

254 MR. H. A. BAYLIS AND LT.-COL. CLAYTON LANE ON

GNATHOSTOMIN A *, subfam. n.

The members of this subfamily are characterized by the

possession, immediately behind the lips, of a cuticular swelling,

which we term the “ head-bulb.” his head-bulb always con-

tains four subglobular, submedian, membranous structures, for

which we use the expression ‘“ballonets.” Hach of these is

connected by a narrow neck with one of the four organs which

have been called by several authors ‘cervical glands,” but which

we prefer to name “cervical sacs.” The latter are elongated,

apparently unicellular, thick-walled, hollow, blind organs ‘hang:

ing freely into the body-cavity at ‘the sides of the cesop! hagus.

Externally they are provided with a coat of fibrils showing a

spiral arrangement (text-fig. 17) and probably contractile. The

protoplasm lining the sac internally probably secretes a fluid into

the lumen.

The cuticle of the head-bulb may be simply raised into

transverse ridges or pronounced. striations with projecting

posterior edges; or the striations may be beset with rows of

chitinoid hooks, strongly resembling the ‘ rose-thorn” type of

hooks found in certain Cestodes, such as Dipylidiwm. The

hooks are embedded by means of their own roots in the cuticle,

and are not merely outgrowths of the cuticle itself.

The external surface of the head-bulb may be unaffected by

the presence of the ballonets within; or may be indented

between the ballonets so as to give an external expression of

the internal structure. Between the ballonets, sometimes push-

ing in the ballonet-membranes before them, certain muscles run

up to the lips, crossing the cavity of the head-bulb obliquely in

such.a way that when contracted they serve to shorten the

head-bulb in an antero-posterior direction. The function of the

head-bulb is probably to act as a burrowing-organ or holdfast.

The worms presumably first bury their heads in the tissues of

the host while the head-bulb is in a deflated condition, and then

the cervical sacs, contracting, force thei: contained fluid into the

ballonets, thus distending the head-bulb. The cuticular stria-

tions and hooks are additional structures rendering the whole

apparatus more effective.

These various structures, with the exception of the ballonets,

have been noted by previous writers, but quite a different inter-

pretation has usually been put upon them. Fedchenko (1872),

working on Gnathostoma hispidum, seems, according to our

translator, to have believed that the matter inside the head was

secreted by the four cervical sacs, and he thus comes nearly to

our own conclusion. He notes the structureless necks into

which the sacs pass, but has observed no actual connection

between the saes and the head-bulb. On the other hand, the

prevailing interpretation is due to v. Linstow, who describes

* For subfamily diagnosis, see p. 255.

THE NEMATODE FAMILY GNATHOSTOMID. 209

(1893) in the same species an elaborate arrangement by which

ducts from the two cervical sacs of each side unite into a common

duct, each common duct ,piercing the corresponding lip and

opening on its outer surface by a conspicuous pore 0:013 mm.

in diameter. The same author (1904a) has also deseribed

similar arrangement of ducts in Tanqua tiara.

von Ritz (1900), for Gnathostoma hispidum, and Leiper (1909),

for G. siamense, follow the very circumstantial description of

v. Linstow.

The ballonets, or rather their contents, seem to be referred to

by v. Linstow as the ‘“‘ Marksubstanz” of the head-muscles, but

their essential nature, as closed membranous sacs, has escaped

the attention of all these observers. Now, in viewing the head

of one of these worms as a transparent object, it is easy to

mistake the muscles that run across the head-bulb for continua-

tions of the cervical sacs, or ducts connecting them with the

exterior. One of us (Baylis, 1916) fell into this error in the

vase of Vangua, and suggested the presence of pores on the

anterior surface of the “striated swellings” (7.¢., the head-bulb).

This is the chief mistake into which other observers have

probably fallen. The ‘ pore” on the outer surface of the lip

described by v. Linstow in Gnathostoma corresponds in position

to the lateral papilla which we find throughout the subfamily.

This papilla sometimes terminates in a little “dimple” of the

cuticle, which might give it the appearance of a pore; further-

more, when seen in transverse section, as in v. Linstow’s case,

the large nerve running up to the papilla might easily give the

impression of a duct. The central portion of the nerve then has

a reticulate appearance, and under a low power rather suggests the

lumen of a duct.

We have been able to trace the cervical sacs into the ballonets,

but are unable to find any external opening, in series of sections

of Echinocephalus and of Tangua (text-fig. 10), and there seems

to be no reasonable doubt that the arrangement is similar in all

the genera and species.

Subfamily Diagnosis.

GNATHOSTOMINA.

Gnathostomide : with a cuticular head-bulb provided either

with marked transverse striations or with rows of backwardly-

directed hooks, and containing four membranous, submedian

ballonets, the cavity of each of which is in communication with

one of four elongated, blind, cervical sacs hanging freely in the

body-cavity.

Habitat (adult): stomach or intestine (exceptionally other parts

of the body) of fishes, reptiles, and mammals.

Type-genus: Gnathostoma Owen, 1836.

256 MR. H. A. BAYLIS AND LT.-COL. CLAYTON LANE ON

Key to Genera of Gnathostomine.

A. Head-bulb furnished with transverse cuticular ridges

. . . =}

having sharp, baclandlyapeieenae edges, but

without hooks ............ aneee boson von. cpap node NANG iMtale

B. Head-bulb armed with eraneverse rows of recurved

hooks.

Gin Vroyalhy wayAATNCEL Gon cceeseceoanse cisssrcsee.. Lchinocephalus (p. 278).

b. Body partially or Mahal aan ay Backeeraiye

GREORG| SIONS! opyecdeprephenses cagueossnssnsonosnen | MDCANONAOUDG! (is ZS).

TANQUA* R. Blanchard, 1904.

Ascaris (in part) v. Linstow (1879, p. 320).

a Stossich (1896, p. 52).

Heter ‘akis (in part) v. Linstow (1904 a, p. 97).

Otenocephalus T v. Linstow (1904 a, p. 102).

Tanqua R. Blanchard (1904, p. 478).

Tetradenos v. Linstow (1904 6, p. 301).

% « (1906) p73).

This genus is characterized by having the large cuticular

head- bulb unarmed, but marked with coarse and very distinct

transverse striations, having their posterior edges projecting so

as to give the outline of the bulb a more or less serrated appear-

ance in optical section.

A former brief account given by one of us (Baylis, 1916) of

this genus requires some correction in the light of further work.

Thus the lips are definitely lateral, and the statement formerly

made to the contrary (following the erroneous orientation of

von Linstow) is withdrawn.

The lips ave curiously asymmetrical, each being tw isted slightly

towards the dorsal or ventral side. in ‘sneh a way that the teeth

of the middle lobes cross each other like the blades of a pair of

scissors. The dorsal and ventral lobes of each lip do not, as was

formerly supposed, bear each a single tooth, but each is bilobed

on the inner surface so as to form two teeth: There are thus

five teeth on each lip, and these interlock with those of the |

opposite lip, giving a very close bite, which enables the animal

to obtain a firm grip of the tissues of the wall of its host’s

stomach.

The head-bulb is divided by longitudinal indentations into

eitlrer two (a dorsal and a ventral) or four (submedian) portions.

In the interior of the bulb are contained the four ballonets with

delicate membranous walls, the cavities of which are in com-

munication with those of the four cervical sacs. When the

head-bulb is divided into four swellings, each swelling contains

one of the ballonets; while in the forms which have only two

swellings, two ballonets are contained in each.

* For generic diagnosis, see p. 258.

+ Not Ctenocephalus Kol., 1857 (dipteron).

THE NEMATODE FAMILY GNATHOSTOMIDA. SST

The cuticle immediately behind the head-bulb always shows an

invagination forming more or less of a “collar” round the neck.

In one species this invagination is so highly developed that the

worm can withdraw its entire head into a prepuce-like sheath ‘of

cuticle. In species in which the collar is less highly developed,

portions of the host’s tissues are not infrequently found caught

between it and the back of the head-bulb, so that the collar

appears to function as an accessory apparatus for enabling the

worm to secure itself.

The cuticle of the body is thick and tough, and is marked with

fine transverse striations, but otherwise smooth. The cervical

papille are usually situated at about one-fourth of the length of the

cesophagus from the anterior end. The excretory pore is situated

at about the same level, or a little in front of it, on the ventral

side. The cesophagus is usually rather long, and the cervical

sacs relatively short, extending beside it to not more than

one-third of its length.

The tail of the male is curled towards the ventral side, and

is provided with strong oblique muscles for this purpose. The

caudal ale are rather well-developed, extending from a little in

front of the cloacal aperture to near the tip of the tail. There

are always eight pairs of caudal papille, which vary very little

in arrangement in the different species. The papille are of

different sizes, the largest being always those of the third, fifth,

and seventh pairs. The fourth and sixth pairs are always small,

and more ventral in position than the others. The cloaca opens

at about the level of the fifth pair, and consequently the fourth

and sixth pairs form a small, ventral, circum-anal group very

much like that seen in Spirowys. All the large papille in Tanqua

show an extremely characteristic structure, having a very large,

swollen, basal portion and a finger-shaped termination.

The spicules are equally characteristic. They are tubular,

eylindrical, and equal in length, and always present a rasp-like

appearance, owing to a covering of minute irregular granulations.

Only the extreme tip is smooth, and this is usually of smaller

calibre than the shaft of the spicule, forming a little rounded or

conical projection.

As regards the female organs, the vulva is situated in the

posterior half of the body. The vagina runs forward fora longer

or shorter distance before opening into the uterus. The latter

may consist of two or four branches. When two branches

are present, they are opposed —7.é., one runs forward and the

other backward. When four are present, three turn forward ~

and one backward. Various swellings, functioning as recepta-

cula seminis or as egg-reservoirs, occur in the course of the

uterine tubes, but their position does not seem to be constant.

In fact, the oeneral shape of the uterus varies so greatly in

different specimens of the same species that the expansions of

the tubes appear to be of a temporary nature, and cannot be

made use of for systematic purposes.

DHS) 5 MR. H. A. BAYLIS AND LT.-COL. CLAYTON LANE ON f

It is a curious fact that the number of uterine branches in

this genus appears to be in direct correlation with the number

of external divisions of the head-bulb. In the forms parasitic

in snakes, where the head-bulb is composed of two portions

only, the uterus has two branches; while in 7’. tiara, where there

are, in the adult, four swellings on the head-bulb, there are also

four uterine branches.

Nothing is known of the development or life-history of any

of the species of Zanqua. The hosts, so far as is known, are all

carnivorous reptiles, and belong to semi-aquatie genera; and it

is not unnatural to suspect that there may be an intermediate

host which lives in or near fresh water, and is eaten by the final |

host. «

Generic Diagnosis.

Tangua* R. Blanchard.

Gnathostomine : head-bulb coarsely striated transversely, un-

armed, divided externally into two or four swellings containing

the ballonets. Body unarmed. Each lip with five teeth, inter-

locking with those of the other lip. Cuticle behind the head-bulb

forms a more or less pronounced collar or invagination. ‘Tail of

male with well-developed ale and eight pairs of papille, of which

the largest are the third, fifth, and seventh, The fourth and

sixth pairs small, ventral, and forming a cireum-anal group. The

larger papille have a large swelling before the finger-shaped

termination. Spicules equal, tubular, rasp-like, with smooth

tip. Vuiva in posterior half of body; vagina running forward

from the opening. Uterus consists of two opposed branches, or

of three anterior branches and one posterior. Hggs oval, with

thin shell ornamented with fine granulations. Embryos not

fully-formed at the time of laying.

Habitat: stomach of semi-aquatic lizards (Varanide) and

semi-aquatic snakes (Zropidonotus, etc.).

Genotype: 7. tiara (v. Linst., 1879).

Key to Species of Zanqua 7.

A. Head-bulb with four swellings. Uterus with four

branches, three anterior and one posterior. Parasitic

In) sem-agquatie Lizards (\Vianamidee) i ereer ene ere se -see eles T. tiara (p. 259).

B. Head-bulb with two swellings. Uterus with two branches,

opposed. Parasitic in semi-aquatic snakes.

Ge Head retractileymeererc eee ea eee eee ee eee an enc Lemon (D208) k

ba Headinotmebactilet: -mrc-ne essere eeecete rae erties saiseen meen CLIZOTIZCULCA (DayAta 40)

* For measurements, see Table II1., p. 270.

+ For doubtful species, see p. 271.

THE NEMATODE FAMILY GNATHOSTOMID. 259

1. Tanqua TraRA* (v. Linst., 1879). (Text-figs 7-15; Pl. III.

figs. 1J-13.)

Ascaris tiara v. Linstow (1879, p. 320; pl. v. fig. 1).

- Parona (1898, p- 114).

Otenocephalus tiara v. Linstow (1904 a, p. 102; pl. ii. figs.

23-27).

Taunqua tiara R. Blanchard (1904, p. 478).

a » Leiper (1908, p. 189).

», (in part) Baylis (1916, p. 224; text-fig. 1).

Tetradenos tiara v. Linstow (1904 b, p. 30 i

- e . (1906, p. 173).

The specimens from Z'ropidonotus asperrimus, previously re-

ferred by one of us (Baylis, 1916) to this species, have now been

Text-figure 7.

Tanqua tiara. The head; dorsal (or ventral) view.

redetermined as 7. anomala (v. Linst.) (q. v.), T. tiara being

restricted to the forms from various species of Monitor lizards

(Varanide). The description of the caudal papille of the male of

T. tara then given was correct, though the specimen figured -

(J. c., text-fig. 2, p- 226) was not 7’. tiara, but 7. anomala, which

in this respect is hardly distinguishable from it. The structure

of the female genital apparatus was not completely elucidated,

and will be given here in greater detail.

*“ Wor specific diagnosis, see p. 264.

Proc. Zoou. Soc.—1920, No. XVIII. ao 18

260 Text-figure 8.

Lp.

<o e.

$./7. p. , [ Dy ==

an })

aN \ [/

Ls bead

O-/mm.

Za ai

Ae Re

| | ia

Uy, ai

Tanqua tiara. The head; lateral view.

Text-figure 9.

Tanqua tiara, Head of young specimen; dorsal (or ventral) view.

O-lmm.

Text-figure 10.

Tanqua tiara. Transverse sections through the anterior region. A, at about the

middle of the head-hulb; B, at the back of the head-bulb; C, at the level

of the cuticular collar.

Text-figure 11.

Text-figure 12.

“iY

cd. p s

Text-fig. 11.—Tanqua tiara. ‘Terminal portion of spicule.

Text-fig. 12—Tanqua tiara. Tail of female; a, lateral ; 6, ventral view.

O'/ M77,

O-3mm

262 MR. H. A. BAYLIS AND LT.-COL. CLAYTON LANE ON

The species appears to be widely distributed in India, Ceylon,

the Malay Archipelago, Australasia, and Africa, and shows little

local variation except in size.

The lips (text-figs. 7 & 8; Pl. III. fig. 11) are large and thick,

and somewhat flattened in ‘front as seen in a dorsal or ventral

view (text-fig. 7). In very young specimens the shape of the

lips is more conical (text- fig. 9). When the lips are viewed from

the dorsal or ventral side (text- fig. 7), there is seen on the edge

of each tooth a little pointed projection which appears to be the

expression of a ridge running along the inner surface of the lip,

and recalling the dentigerous ridges met with in certain

Ascaridee

The hende bulb is large, and is distinctly divided (Gn mature

specimens) into four swellings by iongitudinal depressions 1 in the

mid-dorsal, mid-ventral, aia mid-lateral lines (text-figs. 7 & 8;

Pl. ILI. fig. 11). In very young examples (text-fig. 9), measuring

from 5 to 7 mm. in length, and in which the lips are not yet

fully formed, the head-bulb shows only two swellings, situated

dorsally and ventrally. The transverse striations on the head-

bulb (in the adult) are discontinued at the depressions which

separate the four quarters of the bulb.

The collar-like cuticular invagination behind the head-bulb is

well-marked. The cesophagus occupies about one-fifth of the total

length. The cervical sacs are only wbout one-quarter of the

length of the esophagus, or less.

In the male the fifth, or adanal, pair of caudal papille

(Pl. IIT. figs. 12 & 13) is the largest. The intervals between the

second and third, and between the seventh und eighth, pairs are

longer than the rest. The spicules have a small, smooth,

rounded tip, just in front of which the diameter of the spicule is

slightly reduced (text-fig. 11).

In the female the tail (text-fig. 12) is short, straight and

conical. The female genital apparatus is highly characteristic.

The vulva is situated towards the posterior end of the body,

within the last quarter of the total length. The internal organs

consist essentially of a vagina and four uterine .tubes*, leading

to four ovaries. The four divisions of the uterus are invariably

arranged on the same general plan, three of them passing off

towards the anterior and one towards ‘the posterior end. A

considerable amount of variation appears to exist, however, in

their mode of origin. Sometimes (text-fig. 13) they appear to

originate separately from a common narrow stem which is con-

tinuous with.

and sooner or later turns back more or less parallel with the

vagina; then a little further forward one branch comes off and

runs forward ; and, finally, the common stem bifurcates at its

*In this connection it is interesting to note that another nematode from a

Monitor (Physaloptera varani Parona, 1890 (=P. quadrovaria Leiper, 1908}) also

has a four-fold division of the uterus.

st, the posterior branch comes off

263

THE NEMATODE FAMILY GNATHOSTOMID.

(avqiauez mo uaUTToeds SunoX) ‘suBs1O [eID eTVUa “7.127 oNUnT,

eng

De a oe

h sll La

Hn ie

‘PL otnsy-7xo], .

(erpuy wor momdeds orngeyy) ‘suBS10 [eZIHesS oRIUay *Y.1927 DNbUDT,

Wei) fo

lap

SUTogtN

apg.

} /, /

“an

"Cy OINST-4Xoq,

264 MR. H. A. BAYLIS AND LT.-COL. CLAYTON LANE ON

anterior end to form two parallel branches which also run

forward.

At the opposite extreme we have observed cases (text-fig. 14)

where the vagina runs into a more or less oblong, swollen

uterine sac, which gives off a single branch at its posterior end

and three branches side by side anteriorly. Between these

extremes of variation there appear to be intermediate forms

(text-fig. 15)in which the point of origin of the first of the three

anterior branches shows a greater or less tendency to approach

that of the other two. The shape of the common uterine

chamber varies considerably. We are unable to recognize in

Text-figure 15.

ut. Gee

Tanqua tiara. Female genital organs. (Young specimen from Nigeria.)

these varieties in the form of the female organs any of systematic

value, since we have found the same range of variation in

material from both Indian and African sources, and since the

extremes are, as has been stated, connected by intermediate

forms.

Specific Diagnosis.

TaNQua TIARA (v. Linst.).

Head-bulb divided in the adult into four swellings, each con-

taining a single ballonet. Head not retractile within the

cuiicular collar. Vulva within the last quarter of the body.

Uterus with three anterior branches and one posterior.

Habitat : stomach of semi-aquatic lizards (Varanide).

For list of hosts, see p. 271.

2. TANQUA ANOMALA * (vy. Linst., 1904). (Text-figs. 16-20;

PIS Ve stre. 4%)

Heterakis anomala v. Linstow (1904 a, p. 97; pl. i. figs.

10-11).

Tangua anomala Baylis (1916, p. 229).

1’. tiara (in part) Baylis (1916, p. 223; text-fig. 2).

Of this species we have now, through the kindness of Dr.

J. Pearson, of the Colombo Museum, had the opportunity of

* Wor specific diagnosis, see p. 268.

THE NEMATODE FAMILY GNATHOSTOMID2E. 265

re-examining two of the original specimens. These were, unfor-

tunately, both females, no male being available, but we have

been able not only to confirm our suspicions as to the species

belonging to the genus Z’angua, but also to compare the type-

specimens with our own examples from Indian snakes, and to

satisfy ourselves that these belong to the same species.

This form appears to be very variable in size, while constant

in other characters. The extreme size-variations we should

certainly have been inclined to regard as being at least sub-

species, had we been able to find any definite anatomical

Text-figure 17.

Text-figure 16.

Sse

C.S./1.

O-S m7,

= oe -

ob. ve yp ae fe

Te YUy tye ee

sco

Lee

TATU

Olt,

-, es

[ina

| i

Og || Pearce o> aoue noe

Text-fig. 16..—Tanqua anomala. Head and cesophageal region ; lateral view.

(The cervical sacs are unusually short in this specimen.)

Text-fig. 17.—Tanqua anomala. Anterior end; nearly ventral view.

differences in support of this view. The type-specimens in the

Colombo Museum, from TZropidonotus piscator in Ceylon, are

among the smallest, while the largest examples we have seen

came from a snake of another genus (/omalopsis buccatw) in

Siam.

The anterior part of the body (text-fig. 16) is tapering. The

lips (text-figs. 17, 18) are large and prominent, and more conical

in shape, when viewed from the dorsal or ventral edge, than

266 ON THE NEMATODE FAMILY GNATHOSTOMID#.

those of ;7. tiara. Some of the worms were received, in spirit,

Text-figure 18. |

O03 min.

Tangua anomala. Anterior end; lateral view.

Text-figure 19.

Tanqgua anomala. Tail of male; ventral view.

still firmly attached by the lips to the stomach-wall of the host,

and it requiredga considerable pull to remove them.

(uauttoads cuno xX )

*SUB.GLO [RILUaS O[vUlOY “YpypMmoun pnbuny,

WU Of

‘0% OINSY-4xXaq,

268 MR. H. A. BAYLIS AND LT.-COL. CLAYTON LANE ON

The head-bulb (text-figs. 16-18) is, relatively to the size of

the worm, very small. It consists of two hemispheres separated

laterally but continuous dorsally and ventrally, and is thus

comparable with that of very young examples of 7. tiara. The

striations are very well-marked.

The cesophagus occupies from one-twelfth to one-eighth of the

total length. The cervical sacs (text-figs. 16 & 17) are relatively

short, extending through, roughly, the first quarter of the length

of the cesophagus.

There is nothing in the number, shape, or arrangement of the

caudal papille of the male (text-fig. 19; Pl. LV. fig. 14) to dis-

tinguish this form from 7’. tiara. The preanal ‘ sucker-like

organ,” referred to by v. Linstow (1904 @), appears to us to have

been simply the terminal portion of the intestine, seen in optical

transverse or oblique section by transparency through the body-

wall. We can find no superficial sucker-like organ on the ventral

surface.

The vulva is situated at about one-third of the body-length

from the posterior end. The vagina is short, and opens at right

angles into the uterus. Thelatter has two branches, which usually

proceed at once from the termination of the vagina to run

anteriorly and posteriorly respectively. In a young specimen

from 7 -opidonotus asperrimus (text-fig. 20), the “muscular vagina

appeared to be very short, and continued as a common uterine

tube for a short distance before the divergence of the two

branches.

Specific Diagnosis.

TANQUA ANOMALA (v. Linst.).

Head-bulb divided in the adult into two swellings, situated

dorsally and ventrally, each containing two ballonets. Head not.

retractile within the cuticular collar. Vulva at about the

junction of the middle and last thirds of the body. Uterus

with two opposed branches.

Habitat: stomach of semi-aquatic snakes (Tropidonotus, etc.).

For list of hosts, see p. 271.

3. TANQUA DIADEMA* Baylis, 1916. (Text-figs. 21-23; Pl. III.

fig, Los Ply tig. 6.)

Tungua diadema Baylis (1916, p. 227; text-figs. 3, 4).

This species, which is now redescribed with some slight correc-

tions, is readily distinguished from the preceding form from

snakes by the prepuce-like fold of cuticle within which the

head-bulb and lips can be completely retracted. The cuticle,

especially anteriorly, is often very thick and wrinkled, reminding

one of the condition frequent in Gnathostoma.

* For specific diagnosis, see p. 271.

THE NEMATODE FAMILY GNATHOSTOMID A. 269

The head-bulb (text-fig. 21) is divided into two hemispheres,

as in JZ’. anomala. The lips (text-fig. 21) are rather large

relatively to the head-bulb. The csophagus occupies about

one-tenth of the total length. The cervical sacs are short.

In the male the tail (PI. Lil. fig. 15; Pl. LV. fig. 16) has

rather wide ale. The caudal papille are similar in number and

arrangement to those of the other species. The second pair from

the tip of the tail, however, are relatively small in some specimens,

and do not always show the basal swelling to a very marked

extent. The spicules are suddenly narrowed at the extremity to

form a little, conical, smooth tip (text-fig. 22).

Text-figure 21. Text-figure 22,

The head, with the cuticular sheath removed ;

Text-fig. 21.—Tanqua diadema.

dorsal (or ventral) view.

Terminal portion of spicule.

Text-fig. 22—Tanqua diadema.

Text-figure 23.

0-5 mm.

at.

ate ina@ant

Ls aes

LT LOTT AN fe =

hiitiaieaiss ipiavtai au att AMtgALLL

Spi

CPA

Ai See Secreai|

es stints Heath

SERS SUD ASL age eels

Tanqua diadema. Female genital organs. (Mature specimen.)

In the female the tail is tapering and sharply pointed. The

vulva opens in the middle third of the body. The vagina (text-

fig. 23) is short and thick. It opens at right angles into the

uterus, the two branches of which at once pass off in opposite

directions, without further subdivision.

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a “puapyip WE, ; ] “‘p)VWoUd Wh) | “DADV “TD Sadat Sian cere saroadg

(‘sa.1go mT] rut UL SIUd WAIN SvoUt iv)

‘pnbuny, Jo sowadg oy Jo syuomMamsveyy — TT] ITaV,

THE NEMATODE- FAMILY GNATHOSTOMIDA.

Specific Diagnosis.

TANQUA DIADEMA Baylis.

Head-bulb divided in the adult into two swellings, situated

dorsally and ventrally, each containing two ballonets.

Head

retractile within the Cnnbulae collar, which forms a prepuce-like

sheath.

opposed branches.

Habitat :

“intestines ”

latus, in Brazil.

Tasie 1V.—Hosts and Distribution of the Species of Tangua.

T. tiara

T. anomala

T. diadema

Species.

[? T. tiara] ...

yy ae

Vulva in middle third of “éhe body. Uterus with two

(stomach) of a an Felicops angu-

Host.

Varanus “ ornatus”

(probably V. albigularis).

Varanus salvator.

bb} 33

[Hydrosaurus bivittatus.| |

Varanus gouldii.

Faranus bengalensis.

| Varanus niloticus.

3” 39

. LES

Ve Syne WB exanthematicus.

sp.] |

Locality.

eee

| beurre

| Cevlon.

India.

| Hed. Malay

States.

Australia or

New Guinea.

| Ceylon.

| White Nile.

Gold Coast.

Zanzibar.

Northern

Nigeria.

Tropidonotus piscator.

39 29

Tropidonotus asperrimus

[=Z. quincunciatus|

(local form of ZT. piscator). |

| Homalopsis buccata.

| Water-snuke.”’

“Snake.”

| Helicops | Uranops| angu-

latus.

‘Ceylon.

India.

| Ceylon.

Siam.

India.

| oo)

Brazil.

Recorder.

_v. Linstow (1879).

| Parona (1898).

_v. Linstow (1904 a).

| Baylis & Lane

(present paper).

Leiper (1908).

Parona (1898).

|v. Linstow (1904. @).

| Leiper (1908).

eae ts (1916).

"Baylis (1916)

_ (specimens deter-

mined by Leiper).

| v. Linstow (1904).

| Baylis & Lane

| (present paper).

‘| Baylis (1916) [re-

cordedas 7. tiara].

| Baylis & Lane

| (present paper).

oy)

Baylis (1916).

Doubtful Species of Zanqua.

ASCARIS SPHHEROCEPHALA Rud., 1809.

Rudolphi (1809, p. 188).

On account of certain resemblances Wena the characters of

this species, as described by Rudolphi, and those of the worms

of the Gnathostomid group, we mention it here for the sake of

Allo MR. H. A, BAYLIS AND LT.-COL. CLAYTON LANE ON

completeness. It is quite uncertain to what genus it belongs,

but if Rudolphi’s account of the head-bulb is correct 1t seems to

approach more nearly to Vanqua than to any other. On the

other hand, its host (a fish) and its habitat (the spiral valve of

the intestine) suggest very strongly that it may have been an

Echinocephalus (q.v., p. 273). Rudolphi distinctly states, how-

ever, that the head-bulb is transversely strivted and composed of

two hemispheres, and he makes no mention of any armature

of hooks or spines.

The species is briefly diagnosed as follows :-—

“ Ascaris: membrana capitis utringue semiorbiculari, striata ;

cauda obtusiuscula.”

The following is a rough paraphrase of the further account

(originally in Latin) of the worm, which was found in the lower

part of the intestine of the Sturgeon, Acipenser stwrio, in the

spiral valve.

Worms an inch long, slender, white. Head spherical to naked

eye, separated from the body by a constriction, and inflexed.

Under the microscope a hemispherical membrane is seen on

either side, transversely striated, giving the head a spherical

shape. Mouth smali, surrounded by three large, bluntly conical

lips, joined at the apex, but not at the base, by lateral membranes,

*‘s0 that they become obscured.” Body smooth, attenuated at

both ends, more so anteriorly. Tail rather blunt. Lips of vulva

prominent, in third quarter of body. Aiimentary canal like that

of Asearids. Eggs very peculiar, greatly elongated, blunt-ended

(oblong-elliptical), clear at one end.

Rudolphi says (as we understand him) that on account of the

general shape of the body and of the lips the worm belongs to

the “ Ascarides,” but that the arrangement and “ obscuring” of

the lips [sc. by the ‘‘ membrane” referred to] and the peculiar

eggs mark it off as something different. His description suggests

that the ‘three lips” he saw were the three tooth-bearing lobes

of one lip, seen from the side by transparency, and the ‘*mem-

brane” joining them at the apex the cuticle of the outer or more

lateral portion of the lip.

PHYSALOPTERA CONSTRICTA Leidy, 1856.

Leidy (1856, p. 53).

“Body white, with the brown intestine shining through,

cylindrical to within a short distance of the extremities, incurved ;

anterior extremity with one or two constrictions, and abruptly

inflexed. Lips large, lateral, constricted from the body, each

trilobate. Tail of female incurved, abruptly conical and acute ;

of the male alated, with the ale narrow, long, and turgid.

Length of female 1? inches, breadth two-fifths of a line; male

half the size. .

“Found frequently in the stomach of Zropidonotus sipedon,

with the anterior extremity of the body hooked through the

THE NEMATODE FAMILY GNATHOSTOMIDA. Dies

mucous membrane, and very tightly adhering by means of the

one or two constrictions.”

The form thus described by Leidy so strongly suggests a

Tanqua, both on account of several points in the brief descrip-

tion and on account of its host and habitat, that we feel obliged

to inelude it in our account of the genus. It is impossible, how-

ever, without further investigation, to assign it to a definite

position.

ECHINOCEPHALUS * Molin, 1858.

Echinocephalus Molin (1858, p. 154).

fe Pe LSOle pats | 1):

Cheiracanthus (in part) v. Linstow, in Shipley and Hornell

(1904, p. 100).

os (in part) v. Linstow, in Shipley and Hornell

(1905, p. 54).

Molin’s original generic diagnosis was as follows :—

“* Caput discretum, echinatum ; os orbiculare, terminale, magnum,

imerme, vel armatum, corpus cylindricum, inerme, vel echinatum ;

vagina penis dipetala ; [anus lateralis + |.—Avium et piscium endo-

parasita.”

Under this generic name two species were originally enume-

vated by Molin, #. uncinatus and H. cygni. The latter was

regarded as a species wquirenda, leaving FE. uncinatus the un-

disputed type-species. 4H. cygni belongs to the genus Hystrichis,

and is, in fact, a synonym of H. pachycephalus Molin. Into the

affinities of Hystrichis it is beyond our present scope to enter,

but in spite of the rather strong superficial resemblance borne by

some of its species to some of the Gnathostomide, we regard it as

being of quite a different type from this group. Molin’s generic

characters for Hchinocephalus seem, however, to have been based

in part on the characters of ZH. cygni, and for this reason it

becomes necessary to emend the diagnosis, besides adding to it

from our own observations (see p. 275). Thus the “os orbiculare”

and the statement that the genus is parasitic in birds as well as

in fishes belong rather to Hystrichis than to Echinocephalus.

Molin (1861) rightly suggests the relationship of this genus

to Cheiracanthus (2. e., Gnathostoma). It shows equally great

resemblances to Zanqua, and may, in fact, be looked upon as a

Tanqua-like form complicated by the development of rows of

spines or hooks along the cuticular ridges or “striations” of the

head-bulb.

The lips are, in most cases, very similar in plan to those of

Tangua, and show a similar interlocking arrangement of the

internal, tooth-like, cuticular ridges. In one form, however, the

arrangement is complicated by the multiplication of the teeth on

* Wor generic diagnosis, see p. 275.

7 Added in 1861.

274 MR. H. A. BAYLIS AND LT.-COL. CLAYTON LANE ON

the dorsal and ventral lobes of the lips, so that each lip bears

some 20 teeth altogether.

The head-bulb forms a continuous ring surrounding the base

of the lips; it is never, in the species hitherto met with, divided

externally into two or four swellings, as in Zangua, though it

still contains the same four ballonets connected with the cervical]

sacs. ‘The latter sometimes end in a little button-like appendage

(text-figs. 30, 33, a).

The size of the hooks on the head-bulb, the number of rows,

and the number of hooks in each row, show considerable variety,

and afford useful specific characters.

‘The body is smooth, as in Z’anqua, the cuticle being, as a rule,

finely striated transversely. The neck-papille are rather pro-

minent. Molin’s statement (1861) that the anus is lateral is

entirely erroneous. The excretory pore is very inconspicuous,

and is always to be found at about the same level as the cervical

papille.

The tail of the male is coiled ventrally, and is provided with

slight cuticular ale, into which a series of paired caudal papille

project. Of these there are always eight pairs, as in Tangua, but

the arrangement of the pairs differs somewhat in the different

species. One constant feature is the long space separating the

eighth or most anterior pair from the rest. Some of the papille

are rather large, with a swelling at the base recalling that seen

in Tanqua, but never so well-developed. The spicules are very

characteristic. They are tubular and usually rather long and

slender, the left spicule being apparently always a fraction of a

millimetre longer than the right. Instead of the broken, external,

rasp-like markings seen on the spicules of Z'angua, we find m

Echinocephalus an irregular transverse striation, which only

breaks up into a rasp-like pattern near the tip. The markings

appear to be internal, the outer surface being smooth and trans-

parent. The tip of the spicules consists of a thickening of the

smooth outer layer only, and may be sharply pointed or somewhat

blunt in different species.

The tail of the female is short and bluntly rounded at the tip.

The vulva is invariably situated near the posterior end of the

body, within a very short distance of the anus. The long vagina

runs forward, and it seems to be characteristic of the genus that

it forms a single coil in its course before opening into the uterus.

The latter consists of a wide oblong egg-sac, which becomes very

voluminous in the gravid female, and two horns which run for-

ward from its anterior end. The ova have a finely granulated

surface, and are of a similar oval shape to those of Zanqua.

They do not contain fully-formed embryos when laid.

The adults are found in the intestine (usually in the spiral

valve region) of sting-rays (Trygonide and Myliobatide). There

is evidence that the development is indirect, requiring an inter-

metiate host in the form of one of the bivalve molluses upon

which the final hosts feed. The larve of one species, at least,

THE KEMATODE FAMILY GNATHOSTOMIDA. 275

are found occupying cysts in the tissues of such bivalves as the

pearl-oyster (see p. 277), where they appear to select chiefly the

adductor muscle of the shell as their habitat.

Generic Diagnosis.

HicHINOCEPHALUS* Molin.

Gnathostomine: head-bulb armed with transverse rows of

hooks; not externally divided into swellings, but containing

four ballonets internally. Body unarmed. No cuticular collar

behind the head-bulb. ‘Tail of male with slight ale and eight

pairs of papille, the most anterior pair always separated by a

long interval from the rest. Spicules slightly unequal (left

longer than right), tubular, long and slender, marked with

transverse striations. Vulva near posterior end of body. Vagina

long, opening into a wide uterine sac, which gives off two br anches

anteriorly. Eggs oval, with thin shells ornamented with fine

eranulations. Hmbryos not fully-formed at the time of laying.

Habitat (adult): intestine (usually in spiral valve region) of

sting-rays and other Hlasmobranch fishes.

Genotype: H. uncinatus Molin, emend. Baylis and Lane.

Key to Species of Ychinocephalus

(excluding /. striatus Mont.).

A. Head-bulb with not more than six rows of hooks...... FE. uncinatus.

B. Head-bulb with more than six rows of hooks.

a. Dorsal and ventral lobes of lips each with two

teeth.

a’. Head-bulb with 15 to 18 rows of about 150 to

200 hooks each ...... .. HH. southwelli (p. 283).

b’. Head-bulb with 30 to 40 vows of v ery numerous

hooks ...... .... LH. spinosissimus (p.277).

b. Dorsal and ventral ‘lobes of ‘lips ‘each with a

number (eight or more) of teeth..................... E. multidentatus (p.285).

1. EcHiInocePHALus uncINATUS T Molin, 1858. (Text-fig. 24.)

Echinocephalus uncinatus (in part) Molin (1858, p. 154).

(in part) Molin (1861, p.311; pl. xi.

SESW.) (6))

Cheiracanthus uncinatus v. Linstow, in Shipley and Hornell

(1904, p. 100; pl. ii. figs. 41, 44, 45-48).

Echinocephalus gracilis Stossich, in Shipley and Hornell (1906,

p. 89).

The original description of this, the type-species of the genus,

appears to us to have been based on two distinct species, both of

which we have been able to recognize among our material. The

head-bulb is described by Molin as having about 30 rows of small

hooks in the male, and only six rows “of larger hooks in the

27 7)

* For measurements, see Table V., p. 288.

+ For specific diagnosis, see p. 277.

Proc. Zoou. Soc.—1920, No. XIX. 19

276 MR. H, A. BAYLIS AND LT.-COL. CLAYTON LANE ON

female. These two forms are also clearly figured in his later

paper (1861). The size, number, and arrangement of the hooks,

in our material, appear to give good and constant specific

characters, and we feel little hesitation in concluding that Molin’s

male ‘ wneinatus ” corresponds to what will be described below as

E. spinosissimus v. Linstow, while his female is a distinct form.

We are, therefore, faced with the necessity of deciding which of

Molin’s two forms is to be designated as the type of £. wncinatus.

A good and appropriate specific name already existing for the

form with 30 or more rows of hooks, we have thought it advisable

to retain it; and we feel justified in selecting Molin’s female

form, with 6 rows of hooks, as the true H. uncinatus *.

Our own material is scanty, consisting of one larval individual

found encysted in a Pinna and three very young specimens from

a sting-ray, Myliobatis nieuhofi. As none of these individuals

Text-figure 24.

Man mR: >

Pn an Ww S&S

wal 9 a

Any OMA 7:

Ppp opennpo.h.-

Echinocephalus uncinatus. Anterior end of larval specimen; lateral view.

contain fully-developed sexual organs, our description must

remain incomplete.

The lips are not yet fully developed in any of the specimens,

and we are unable to describe their structure. The head-bulb

(text-fig. 24) is armed with 6 rows of hooks, each row containing

between 40 and 50. The spines increase in size from before

* The question of nomenclature is somewhat further complicated by the fact that

Shipley and Hornell appear to have submitted larval forms of a species of Hehino-

cephalus to both von Linstow and Stossich, and these two authorities held different

opinions as to their identity. von Linstow ascribed them to Mbolin’s: species

uncinatus, while Stossich created for them a new species, gracilis. We cannot,

however, find anything in Stossich’s (in Shipley and Hornell, 1906) remarks upon the

larvee to justify this step, and as the specimens are definitely stated by both

authorities to have 6 rows of hooks, we regard the name gracilis as a synonym of

wneimatus.

THE NEMATODE FAMILY GNATHOSTOMIDA. DRE

backwards. The neck-papille were not seen. Caudal papille

could not be detected, nor was the position of the vulva yet

visible. The cuticular ‘striation is extremely fine and indistinct.

As regards the occurrence and life-history of this species, it

seems fairly clear that it requires an intermediate host, in the

shape of a bivalve mollusc, for its transference into the alimentary

canal of its final host. Molin’s original adult material was found

in Trygon brueco in the Adriatic. von Linstow (in Shipley and

Hornell, 1904) states that it also occurs in 7. pastinaca. The

larval stages are found encysted in the tissues of bivalves, usually

in the adductor muscle of the shell. Thus they have been found

fairly frequently in the pearl-oyster (see Shipley and Hornell,

1904, pp. 101-102), and we have now recorded their occurrence

in Pinna sp. In the pearl-oyster the larve are occasionally

found “entombed in the nacreous lining of the shell” (Shipley

and Hornell, 1904), where their shape is said to be wonderfully

preserved.

In addition to the bivalves, the trigger-fishes, Balistes mitis

and B. stellatus, are recorded by Shipley and Hornell (1904) as

hosts for the later larval stages. It is not quite clear whether

these fishes are regarded as a necessary second host of the parasite

or not. The species of Vrygon are said to devour both the Balistes

and the oysters, but the presence of the worms in Balistes may

have been abnormal. In these fishes they were found not only in

the alimentary canal, but also in the peritoneum and connective

tissue, in which they were thought to burrow by the help of the

head-bulb, which was seen in both inflated and deflated conditions.

Specific Diagnosis.

EicHINOCEPHALUS UNCINATUS Molin, emend. Bavlis and Lane.

Head-bulb with six rows of hooks, each row containing from

AQ) to 50 hooks.

For list of hosts, see p. 289.

2. KGHINOCEPHALUS SPINOSISsIMUS * (v. Linst., 1905). (Text-

figs. 25-29; Pl. LV. figs. 17, 18.)

Lchinocephalus wneinatus (in part) Molin (1858, p. 154).

(USIoN i gos SUIS jolly erat.

figs. 5, 6).

Cheiracanthus spinosissimus v. Linstow, in Shipley and Hornell,

(1905, p. 54, pl., figs. 12, 13).

This specific name was proposed by von Linstow for a form

having 30 to 33 rows of hooks on the head-bulb, each row

containing some hundreds of hooks. The host of the type-

specimens was Myliobatis aquila, from the Gulf of Manaar.

9 2? “ID 9?

* For specific diagnosis, see p. 283.

IS)

278 MR. H. A. BAYLIS AND LY'.-COL. CLAYTON LANE ON

We believe, as has been stated above, that the male form

described by Molin under the name of Ychinocephalus uncinatus

is referable to this species. The host, in this case, was Zrygon

brucco. Among our own material there are specimens from the

sting-rays, Trygon walga and Urogymnus asperrimus, which

appear to belong to the same species.

From their size (15°7 mm. in Jength) von Linstow’s examples

seem to have been immature, and few details are given of their

internal stucture. We will therefore give a fuller description

based upon our own material, chiefly upon the specimens from

Urogymnus.

The worms are rather long and slender in general appearance.

The largest female in our possession is not quite mature.

The lips (text-figs. 25 & 26) are large, very prominent and

massive, each consisting of an outer portion bearing the three

papille, and internally a rounded middle, and a dorsal and a

ventral triangular, tooth-bearing lobe. These lobes meet those

of the other lip and guard the mouth. The middle lobe is

without teeth, while the other lobes are each provided with an

internal thickening of the cuticle which is produced into tooth-

like ridges (text-fig. 28). The teeth interlock, as in Zangua,

with those of the opposite lip. The posterior surface of the

dorsal and ventral lobes bears a row of small tooth-like serrations

(text-figs. 25, 26, ser.).

The head-bulb (text-figs. 25, 26) is distinctly marked off from

the neck, and is visible to the naked eye as a little knob. The

hooks (text- -fig. 27) are very minute and very numerous. They

are arranged in from 30 to 40 transverse rows, each row containing

a very lars ve number (several hundreds) of hooks, The rows are

not always complete circles, but sometimes break off suddenly,

while dichotomous branching of the rows is not infrequent.

The cuticular striation on “the body is Tela fine. von Linstow’s

statement that the wsophagus is only ;. of the body-length

appears unintelligible, unless it is a misprint for 4-1, and even

in this case it does not agree very well with our own measure-

ments.

Of the eight pairs of caudal papille in the male (PI. IV. fig.

17), those of the first pair are small and lateral, the second pair

more ventral. Pairs 3 to 6 form a group on either side of the

anus, the 6th pair being apparently adanal. The 5th pair is

situated somewhat more laterally than the rest. These four pais

are placed close together, but a longer space separates pairs 2

and 3and pairs 6 and 7. The 8th pair is separated by a very

long interval from the 7th, and stands quite far forward, these

last two pairs being both pre eanal. All the papille, except the

small first pair, have a somewhat swollen base, not unlike

the strueture seen in Zanqua, though less strongl y developed.

The stout spicules (Pl. IV, fig. 17) are very slightly unequal in

length, the left being the longer. Hach spicule terminates in a

smooth, conical point (Pl. IV. fig. 18).

THE NEMATODE FAMILY GNA‘

HOSTOMID2

Text-figure 25,

RAI

Yn!

RN Ry

WARM A

FAN IMUME MAAN AY

(UAC

LOA OURS pA

fi ! intel WHat atin

Heidt UCU OA

AY i.

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PCM OC i

(tC COE EE OSE UPDATER RNY

' rey 1 '

; PAULA

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AUN ALD UVM UTEEO tipi

ce UA COE NEL EP ME pps

Say rit WUD COSTED OD Wy

OO a

tite

etl

C.S./2.

Eehinocephalus spinosissimus. The head; dorsal (or ventral) view.

Text-figure 26.

A WANE THE wy

MU \\)

" VANTANLET BENNY VED TERNS =

NEA VUE UAT

wet

\N i MATOS

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\\\\ RU SAT Watt HH

STAI BY

NUE ct g tt EESTI TUT Cray ay UND PEKT UU E VYE Asn

O-3Smm.

Hehinocephalus spinosissimus. The head; lateral view.

280 MR. H. A. BAYLIS AND LY.-COL. CLAYTON LANE ON

Text-figure 27.

O-OS mm.

Echinocephalus spinosissimus. Hooks: a, in profile; 6, in surface view.

Text-figure 28,

c.€

O-/mm.

Lichinocephalus spinosissimus. 'ransverse.section through the lips. (The section

is somewhat oblique, passing through the middle lobe and only one outer

lobe of each lip.)

THE NEMATODE FAMILY GNATHOSTOMIDA. 281

The vagina (text-fig. 29) including the unpaired portion of the

uterus, runs forward for nearly 5 mm. before giving off the two

Text-figure 29.

vt,

Echinocephalus spinosissimus.

Posterior end of female; lateral view, showing genital organs.

uterine branches. Weare unfortunately unable to describe the

ova, none having been seen in a fully-developed condition in our

specimens.

Text-figure 50.

TARA

Fa ittiiManatanns Na

iyi Wn SHAHN

PATNA TAN

Hitt

etitit

ce ia

(cut (Uae! wt

SN Wis Me

c.S.

Lichinocephalus southwelli. Head and cesophageal region ; lateral view

THE NEMATODE FAMILY GNATHOSTOMID. 283

Specific Diagnosis.

ECHINOCEPHALUS SPINOSISSIMUS (v. Linst.).

Head-bulb with 30 to 40 rows of hooks, each row containing

several hundreds. Dorsal and ventral lobes of lips bear two

teeth each.

For list of hosts, see p. 289.

3. ECHINOCEPHALUS SOUTHWELLI*, sp. n. (Text-figs. 30-32;

Pl. IV. fig. 19; Pl. V. figs. 20-23.)

This species, which we name in honour of Mr. Southwell, to

whose kindness we are indebted for this and much of our other

material, was also found in Urogymnus asperrimus. It is a form

intermediate in several features between LZ. spinosissimus and

a third species (1. multidentatus) to be described below, and all

these three species were found together in the same individual

host.

The present species is rather shorter and considerably more

slender than #. spinosissimus.

The lips (PI. IV. fig. 19; Pl. V. fig. 20) are rather squat,

and their dorsal and ventral lobes bear only two teeth each, as in

L. spinosissimus. he head-bulb (text-fig. 30), however, is much

Text-figure 31.

cd.p

O:/ mm.

Echinocephalus southwelli. Caudal extremity of female; ventral view.

more like that of the next species, H. multidentatus, though

smaller and less flattened antero-posteriorly. ‘There are from 15

to 18 rows of hooks, which have a maximum length of 0°035 mm.,

and are thus intermediate in size between those of #. spinosissi-

mus and H, multidentatus. The number of hooks in each row is

also intermediate, there being, as nearly as can be estimated, from

150 to 200.

The cuticular striation on the body is rather coarse. The cervical

sacs are nearly as long as the cesophagus. In the male, the first

pair of caudal papille (Pl. V. figs. 21, 22), near the tip of the tail,

* For specific diagnosis, see p. 285.

a. a. A

ale; lateral view,

Posterior end of fem

Text-figure 32.

SS)

Echinocephalus southwelli.

‘sans.

‘ showing genital o1

THE NEMATODE FAMILY GNATHOSTOMIDS,. | 285

are quite lateral; the 2nd, 5th, 6th, 7th, and 8th form a longi-

tudinal series, while the 3rd and 4th are a little nearer to the

mid-ventral line. Pairs 2-5 forma group rather close together,

of which 2 and 4 are small papille and 3 and 5 relatively large.

The intervals between 5 and 6 and between 6 and 7 are about

equal, and greater than those between the posterior pairs, while a

much longer interval separates 7 and 8.

The spicules (Pl. V. figs. 22, 23) are rather slender. A little

before the tip each spicule is narrowed and then expanded some-

what, before terminating in a conical point.

Tn the female the tail (text-fig. 31) is very short. The vagina

(text-fig. 32) leads into a very wide uterine sac.

Specific Diagnosis.

ECHINOCEPHALUS SOUTHWELLI Baylis and Lane.

Head-bulb with 15 to 18 rows of hooks, each row containing

about 150 to 200. Dorsal and vential lobes of lips bear two

teeth each.

For list of hosts, see p. 289.

4, ECHINOCEPHALUS MULTIDENTATUS*, sp.n. (Text-figs. 33-36 ;

Pl. VI. figs. 24-27; Pl. VII. fig. 28.)

This is the third species obtained from the sting-ray, Uro-

gymnus asperrimus. In general naked-eye appearance it closely

resembles /. southwelli, but 1s rather shorter and proportionately

stouter. The head-bulb (text-fig. 33) is very conspicuous, and

gives the worm much the ‘appearance of a small nail.

The lips (Pl. VI. figs. 24, 25) are well-developed, but not

prominent. In structure they are more complex than in any of

the other forms studied by us. The outer portion bears the usual

three papille, and the inner portion is, as usual, trilobed. The

cuticle of the opposed inner surfaces of the dorsal and ventral

lobes is raised into a number (from 8 to 11 on one lobe) of tooth-

like ridges. These teeth are not constant in number or arrange-

ment, and are not always symmetrical on the two lobes of the

same lip, as may be seen in Pl. VI. fig. 25. Whether the teeth

of the two lips can be interlocked is uncertain, though their

appearance suggests that this is probably the case.

The head-bulb (text-fig. 33) is very wide compared with the

neck which follows it, and is somewhat flattened antero-posteriorly.

There are from 11 to 13 rows of large hooks (text-fig. 34). Hach

row may be estimated to contain some 100.

The cervical sacs are, as in 1. southwelli, of approximately the

same length as the esophagus, both terminating at a point a

little more or less than 2 mm. from the anterior extremity.

* For specific diagnosis, see p. 289.

286 MR, H. A. BAYLIS AND Ll.-COL. CLAYTON LANE ON

The esophagus is thus very short as compared with that of

EL. uncinatus and E. spinosissimus. The neck- papule are

prominent.

Text-figure 33.

aw Oe oe.

Pee ny paies

SSA PPO PP IDOL f “OO Gd TTS

pe Aah HEM FEB RG NAA

nant VW GROVE DD Dp, HADI

jai ie IN

DIVVVVD II Pp pps, :

yt MAKE mee WI 99%

sau nena Dae

| \

Lichinocephalus multidentatus. Head and cesophageal region; lateral view. (The

triangular opening in the head-bulb is due to an artificial splitting of the

cuticle.)

Of the eight pairs of caudal papille in the male (PI. VI.

figs. 26, 27), the first, as usual, is lateral, while the rest are more

ventral in position. Pairs 1 to 5 appear to be postanal, 6 to 8

THE NEMATODE FAMILY GNATHOSTOMID. 287

Text-figure 34.

= = ue

a " By

O-/mr7 : yt

Echinocephalus multidentatus. Hooks: a & 6, various profile and surface yiews ;

e, a strip of cuticle with hooks, from a macerated specimen.

Text-figure 35.

Echinocephalus multidentatus. Posterior end of female; lateral view.

preanal. Pairs 3, 7 and 8 are conspicuously larger than the

remainder. There is a long space between Vand 8. Nos. 4and 5

are small papille, one pair, which we call 4, being im a line with

Nos. 2 and 3, while No. 5 is displaced laterally and posteriorly so

as to lie at the same transverse level as No. 4. The papille

project into a very feebly developed ala on either side. The

MR. H. A. BAYLIS AND LT.-COL. CLAYTON LANE ON

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THE NEMATODE FAMILY GNATHOSTOMID

spicules are rather slender.

smooth and rather blunt.

A. 289

Their tips (Pl. VIL. fig. 28) are

The vagina (text-figs. 35, 36) passes into a voluminous uterine

sac.

Specific Diagnosis.

EICHINOCEPHALUS MULTIDENTATUS Baylis and Lane.

Head-bulb with 11 to 13 rows of large hooks, each row contain-

ing about 100. Dorsal and ventral lobes of lips

teeth each.

bear 8 or more

TasLe VI.—Hosts and Distribution of the Species of Hchinocephalus.

Recorder.

| Molin (1858).

y. Linstow, in Shipley

& Hornell (1904).

Baylis & Lane

(present paper).

Shipley & Hornell

(1904).

ch)

} -Y] 32

Baylis & Lane

(present paper).

y. Linstow, in Shipley

& Hornell (1905).

Molin (1858)

[recorded as

Hi. uncinatus |).

Baylis & Lane

(present paper).

Species. Host. | Locality.

| “i cathe

| EB. uncinatus (adult) | Trygon brucco. Adriatic.

| Bs Trygon pastinaca. [P]

(immature) | Myliobatis niewhofi. Ceylon.

- Balistes mitis. e

| 5 Balistes stellatus. Pe

(larva) | Margaritifera marga- “3

ritifera. | M. vulgaris. |,

as Pinna sp. | *

jee he TT SA eee SW

| E. spinosissimus-..... | Myliobatis aquila. | Ceylon

| | ike

| Trygon brucco. | Adriatic.

| i}

| |

| Trygon walga. |. Ceylon.

| Urogymnus asperrimus. 5

E. southwelli ......... | Urogymnusasperrimus. Ceylon.

E. multidentatus .... Urogymnus asperrimus. Ceylon.

Et. striatus ............' Seyllium sp. Peru.

i? (OV aneeaeeee Aétobati narinari. Loyalty Is.

Monticelli (1889).

Shipley (1900).

5, EcuINnocePHatus srriatus Monticelli, 1889.

Monticelli (1889, p. 71).

? Shipley (1900, p. 560; text-fig. KE).

The name £. striatus was given by Monticelli to some specimens

from the stomach of Seylliwm sp., from: Pay

ta, Peru. Sub-

sequently Shipley doubtfully referred to the same species some

Text-figure 36.

ut:

cd. p.

Echinocephalus multidentatus. Posterior end of female; ventral view,

showing genital organs.

THE NEMATODE FAMILY GNATHOSTOMID A, 291

worms from the intestine of a four-spined sting-ray, Aétobatis

narinart, from Loyalty Islands. The name #. striatus, however,

seems to be a nomen nudum, and we have no means of identify-

ing the species, though we mention it here for the sake of com-

pleteness. Shipley (. c.) gives a figure of the head of the form

referred to by him, and in this floure some 14 rows of hooks are

indicated, so that his species would seem to approach closely to

our &. southwelli, aud 1s possibly identical with it.

GNATHOSTOM A* Owen, 1836.

Gnathostoma Owen (1836, p. 125).

Cheiracanthus Diesing (1838, p. 189).

A Diesing (1839, p. 221) [not Chetracanthus Diesing

of v. Linstow, in Shipley and Hornell (1904, p. 100) nor

(1905, p. 54) J.

Filaria (in part) Schneider (1866, p. 98).

Cheiranthus v. Linstow (1893, p. 202) [misprint].

Gnathostomum Mitter (1912, p. 150).

The members of this genus are stout worms with the characters

of the subfamily. The head-bulb is, according to the state of

contraction of the contained muscles, globular or somewhat

flattened antero-posteriorly, and bears in the known species from

eight to twelve transverse rows of simple hooks, like those of

x chinocephalus, set on a cuticle which shows no external trace of

the four underlying ballonets. Some of the anterior and posterior

rows may be obscured, the first by the inrolling of the anterior

part of the head-bulb, as the result of contraction of the internal

muscles, the second by the partial retraction of the head-bulb

into the neck and the resulting interposition of the densely set

scale-like spines which cover this part of the body. These are set

more or less alternately in transverse rows and have their free

eilges indented to varying degrees, so that they come to possess

shar p points of varying shape ‘and number, Towards the middle

of the body the spines become simple and either continue as such

to the posterior end or progressively diminish in size and finally

disappear, leaving the posterior part of the body naked. The

excretory pore has not been detected, a failure easily understood

when one considers the dense covering of spines and the minute-

ness of the pore in other genera of the subfamily.

The male has lateral caudal ale each sustained by four large

papillz and by the tip of the tail, which has the general appearance

of, and has sometimes been counted as, an unpaired terminal

papilla (Schneider, 1866, p. 86 and text-fig.; v. Linstow, 1895,

p- 200) te. TL): Between the ale are two pairs of small, sessile,

ventral papilla. The male has two unequal spicules and no

accessory piece, although v. Linstow (1893, p. 206, fig. 13) has

described, lying ventr alliy to the spicules, a body which he seems

to regar d as such, calling it a “ Stiitzapparat.”

* For generic diagnosis, see p. 292.

Proc. Zoo. Soc:-—1920, No. XX. 20

292 MR. H. A. BAYLIS AND LT.-COL. CLAYTON LANE ON

The female has a blunt tail; the vulva probably always lies

behind the middle of the body; the vagina is long; there are two

uteri; the eggs have a thin colourless shell, marked by a very

fine external stippling, and have at one pole an appearance

as of a watch-glass-shaped cap due to a thinning of the shell

combined with a recession of the shell-membrane, and providing

a means of exit for the embryo. It has, however, sometimes

been described as a polar thickening of the shell (Schneider,

1866, p. 98; v. Linstow, 1893, p. 207).

The genus (vide Table VIII., p. 304) has a wide distribution

in a number of carnivorous hosts. Its normal habitat is clearly

the wall of the stomach. The genotype acquires an added

interest in that it has been found in man (in the subcutaneous

tissue). The presence of certain species in the lumen of the gut

we attribute to their having been parasitic in some host which

was devoured by the animals from which they were actually

recorded. Regarding one record of a Gnathostoma from the

colon judgment must be suspended (vide p. 305).

Generic Diagnosis.

GNATHOSTOMA * Owen.

Gnathostomine: head-bulb armed with simple hooks, the

ballonets giving no external evidence of their presence; body

armed with cuticular spines, anteriorly scale-like with the free

edges incised into points varying in number and shape, more

posteriorly becoming less subdivided and finally appearing as

simple spines, which either continue as such to the posterior end

or disappear, leaving the hinder part of the body unarmed; the

male with unequal spicules and four pairs of large lateral and

two pairs of small ventral caudal papille ; vulva behind the middle

of the body; vagina long; uteri two in number; ovum with thin

colourless shell, a marked thinning at one pole causing a weak

spot through which the embryo escapes.

Habitat: Normally the gastric wall, usually of carnivorous

mammals,

Genotype: G. spinigerum Owen, 1836.

Nove.—The name Grathostoma has been placed on the official

list of generic names by the International Commission on

Zoological Nomenclature 7.

Key to Species of Grathostoma.

A. Body completely clothed with spines ............... wa. G. hispidum (p. 298).

LB. Spines clothe only the anterior half or two- reds of

the body.

a. The spines immediately behind the head-bulb comb-

like, having four points of about equal length...... G. spinigerum (p. 293).

* For measurements, see Table VII., p. 302.

+ Opinion 66 (Smithsonian Institution, Washington).

THE NEMATODE FAMILY GNATHOSTOMID®. 293

b. The spines immediately behind the head-bulb leaf-

like, having five points, one at the tip and two

aikornae Gliese GNCLE- ",-p sseundongoceaedeobtebeoodmeesnsdacdoses | Gro OAUCHO {Qo BOM).

G. horridum and G. turgidwm are too insufficiently described

to be capable of inclusion in a key.

1. GNATHOSTOMA SPINIGERUM * Owen, 1836. (Text-figs. 37-40;

Pl. VII. figs. 29-32; Pl. VIII. figs. 33-38).

Gnathostoma spinigerum Owen (1836, p. 125).

Cheiracanthus robustus Diesing (1838, p. 189) [nomen nudum].

Cheiracanthus robustus Diesing (1839, p. 222; pl. xiv.

figs. 1-7, pl. xvi. figs. 1-24).

Cheiracanthus socialis Leidy (1859, p. a

Filaria radula Schneider (1866, ae 98; By [Ole alae), 8) aia 8) ))).

Cheiracunthus siamensis Levinsen (1889, p. 3233; pl. vu.

figs. 9-14),

Gnathostoma paronat Porta (1908, p. 8).

Gnathostomum spinigerum Mitter (1912, p. 150; pl. v.).

The contradictory and incomplete character of existing

descriptions of this nematode, and the consequent unnecessary

multiplication of species, fully justify its redescription.

Text-figure 37.

Sara AG io FST

Fee t us ~ ZS

y G +E VuvVvFey oh s)~. SSS -

He LEE Ly NE Meus VN SSN IBS Ss.

b YVYNAVYY WaEES

Secures YAVVYVY WAGES

ZS Hho meas Vuyy ae VV VY ae NY

ss at S ~\

2 fe) sae ho fa foes a e \

L O-3 717.

Gnathostoma spinigerum. Anterior end; dorsal (or ventral) view (surface).

The specimens now described are from /elis pardus and were

received in part from the Veterinary College, Belgachia, Calcutta,

where they were collected by the late Mr. 8. N. Mitter, and in

part from the Parel Laboratory, Bombay. In each case the

worms had produced fatal perforation of the gastri¢ wall.

In this species the cuticle lining the inner aspect of each lobe

of either lip is longitudinally thickened into a ridge which meets

its fellow of the opposite side. In our specimens the head-bulb

* For specific diagnosis, see p. 298.

20%

294 MR. H. A. BAYLIS AND LT.-COL. CLAYTON LANE ON

(text-figs. 37, 38; Pl. VII. fig. 29) carries from 8 to 11 rows of

hooks (text-fig. 39). The comb-like spines immediately behind

the head (text-fig. 40, a) carry on their straight distal edges four

short points. These rapidly give place to three-pointed spines

(text-fig. 40, 6), the middle point of which is typically the longest.

Spines of this type extend posteriorly to beyond the termination

Text-figure 38.

A Aan Sd

y AVA ANASS SA

YY aaa?

Ny)

VV VV)

ZZ:

Gnathostoma spinigerum. Anterior end; lateral view (surface).

Text-figure 39.

Gnathostoma spinigerum. Hooks from the head-bulb: @, surface; 4, profile view.

of the cesgphagus. There follow progressively two-pointed and

single-peinted spines (text-fig. 40, c, d), which in turn diminish

in size so that at 8 mm. from the head-end they form mere

points protruding beyond the transverse striations of the cuticle.

‘These soon disappear, leaving the posterior part of the body

naked, except for the spines to be described below on the tail of

the male.

DOA

THE NEMATODE FAMILY GNATHOSTOMIDA. 295

The diameter of the cesophagus immediately behind the head

is 0°125 mm.; its widest diameter is 0°6 mm. and lies at about

0-6 mm. from its termination.

The cervical papille (Pl. VIT. figs. 30, 31) are coarse, rounded

knobs with their terminations directed posteriorly, each lying in

a small bald area among the spines.

In the male the four large caudal papille on each side (Pl. VII.

fig. 32; Pl. VIII. fig. 33) lie close together in the caudal ale.

No. 1 is the smallest, the others being all of about the same size

but varying in relative proportions according to the aspect from

Text-figure 40.

ay ole /

y Vi 6

0-05 1m.

— /

}

¥,

Gnathostoma spinigerum. Body-spines: a, from the neck, immediately behind

the head-bulb; 6, from the csophageal region; ec, from behind the

cesophagus (abont 4 mm. from the anterior end); d, from about the middle

of the body.

which they are viewed. ‘Their nerves are particularly large and

obvious. Near the bases of Nos. 1 and 2 of these papille he the

two pairs of small ventral papille. The cloacal opening lies in

front of these.

On the greater part of the posterior 0°8 mm. of the ventral

surface of the caudal extremity the cuticular striz are closely set

with small spines (Pl. VII. fig. 32; Pl. VIII. fig. 33), the general

296 MR. H. A. BAYLIS AND LY.-COL. CLAYTON LANE ON

direction of whose points 1s away from the cloacal opening.

There exists, however, a bald Y-shaped area whose narrow stem

reaches from the tip of the tail to the anus and whose broad

arms stretch thence laterally and anteriorly as far as the most

anterior of the lateral papille.

The short right spicule (Pl. VIII. fig 34) is wide at the base

and narrows somewhat abruptly about its middle, continuing

narrow thence to its rounded point. The long left spicule has

the same general shape as the right except that the shaft narrows

at about the junction of the anterior and middle thirds, and that

the middle third has at least the appearance of being the narrowest

part. When the large spicule is extruded it is seen that it is

hollow-looking, faintly striated, with a fine colourless outer iayer

which thickens to form the extreme point of a slight terminal

expansion (Pl. VIII. fig. 35

In the female, the vulva, shaped as a slight tranverse slit, opens

into a vagina with a general anterior direction. In the specimen

examined its first 2 mm. was strongly muscular and narrow, the

beginning being markedly, tortuous; the next 0-5 mm. was dilated

and full of eggs; then followed a narrow muscular loop running

posteriorly and * dorsally nearly to the level of the vulva and

returning on itself to enter another short dilatation close to the

first one; thence it ran forward as a narrow muscular tube taking

a nearly straight course to the posterior end of the cesophagus.

Tmmediately after turning posteriorly at this point it was found

broken and the continuation could not be discovered. The course

traced measured 11°5 mm. Owen’s (1836, p. 126) account of his

dissection gives it a course of over 25 mm. before dividing into

the two uteri. ‘The tail of the female is, in a lateral view

(Pl. VIII. fig. 37), rounded dorsally and flattened ventrally,

while in a ventral view (Pl. VIII. fig. 36) its end is bluntly

rounded and carries close to the tip a pair of unusually massive

caudal papille. It is clear that a collapse of the cuticle about

these papille might readily produce the “three-lobed” appearance

which Levinsen (1889) figures (vide infra, p. 297).

The ovum has a thin colourless shell with a very fine granu-

lation on its outer surface and the usual polar cap. In the

female examined the ova contained fully-formed embryos, some

of which were found in the act of escaping through the thinned

pole (Pl. VIII. fig. 38).

The justification for the correctness of the list of synonyms

given above is to be sought in the following lines and in

Table VII., page 302.

The original description of Gnathostoma spinigerum (Owen,

1836) based on specimens removed from tumours in the stomach-

wall of a young tiger which died in the London Zoological

Gardens, corresponds, so far as details are given, with that just

written above, except that he described the armature of the head-

bulb as similar to that of the body and noted only one spicule and

four pairs of papille, apparently three of the large lateral ones

She PS

THE NEMATODE FAMILY GNATHOSTOMIDA. 297

ani the sessile pair behind the cloacal opening. He found also

small worms, possibly young individuals of the same species,

5 lines long *.

Diesing (1859, p. 222) in describing Cheiracanthus robustus

noted its closeness to Gnathostoma spinigerum, but, accepting as

correct Owen’s description of the head and spicule, he, with

ditiidence, separated the two forms. It is probably not doubted

at the present time that Owen and Diesing were working on the

same species. In his subsequent description of Diesing’s material,

v. Drasche (1883, p. 126) described four pairs of largé and three

pairs of sessile papillee.

Under the name of Cheiracanthus siamensis Levinsen (1889)

described a single poorly preserved immature female nematode

which had been removed from an abscess in the breast of a young

native woman in Siam. He was able to deal with external

characters only. The distribution and shape of the spines on the

head and body were identical with those of Gnathostoma spini-

gerum. In his Latin synopsis (J. c. p. 325) he uses the expression

“ Corpus in partem caudalem trilobatwm desinet.” His figure of

the ventral aspect of the tail shows, however, that what actually

existed was merely a slight compression of the lateral outline of

the tail posteriorly to the anus, a condition which does not

correspond to the idea conveyed by the term ‘“ three-lobed.”

Leiper (1909, p. 70) has described a male of Gnathostoma

siamense which, as the context indicates, came from a sub-

cutaneous swelling in a native of Siam. The lips were large and

fleshy, measuring “1:5 by 0°5 mm.” [?0°15 by 0:05 mm.] and

each bearing two papille witha median protrusion between them.

The cephalic hocks measured 0-015 by 0:005 mm. Spines covered

the anterior four-fifths of the body, the most anterior having

three digitations, the most posterior one only. The ejaculatory

duct was 1-Dmm.long. Later Leiper (1911, p. 18; 19138, p. 281),

after re-examining Diesing’s original examples of Chetracanthus

robustus (which, however, he speaks of as having come from the

leopard), reaches the conclusion that the male form obtained from

man corresponds exactly with the male of Cheiracanthus robustus,

which, he remarks, “is acceptedly the same as Gnathestoma spint-

gerum.” In no respect, indeed, do the forms from the sub-

cutaneous tissue of man differ, except m the matter of maturity,

from those from the stomach of Felidae, but must be considered

merely as individuals which have strayed into an unnatural

habitat in an unnatural host (cf. Leiper, 1909, p. 80).

Cheiracanthus socialis Leidy (1859, p. 53) was found in cavities

in the thickened stomach-wall of the mink (J/ustela vison). There

is in its size, in the character of its lips, head and hooks, in the

shape and distribution of the spines on the body, in the internal

organs, so far as described, and in the genital papille, nothing to

* Owen did not describe the males as 5 mm. long and the females as twice that

length, nor the tail of the female as trilobed (vide Stephens, in Fantham, Stephens

and Theobald (1916, p. 385)).

298 MR. H. A. BAYLIS AND Lit'.-COL, CLAYTON LANE ON

distinguish it from Gnathostoma spinigerum. It does not seem

reasonable to consider the name otherwise than as a synonym,

pending re-examination of the original material.

Gnathostoma paronai Porta (1908, p. 8) 1s a name based on a

single badly preserved female so opaque that no internal structure

was made out. Its meagre description is in every way applicable

to Gnathostoma spinigerum. It was found free in the intestine of

Rattus [Mus] rajah. Jts unusual habitat and poor condition

suggest that it was in reality a moribund parasite of some animal

eaten by tlie rat.

Schneider (1866, p 98) described from the gastric wall of

Paradoxurus philippinensis a parasite, Filaria radula, with the

general external appearance of Gnathostoma spinigerum. He also

notes particularly that the egg-shell was finely stippled and

thickened at one pole, but detected only three pairs of caudal

papille in the male. He refused to identify his specimens

with Gnathostoma spinigerum, partiy on account of their different

geographical distribution and partly because the tail-papille,

as he believed them to be situated, had an arrangement which

he associated with the genus Filaria. These reasons for separating

it from G. spinigerwm cannot be accepted as cogent, nor are there

any cogent ones to be found in the description.

Specific Diagnosis.

GNATHOSTOMA SPINIGERUM Owen, 1836.

Gnathostoma: eight to eleven rows of hooks on the head-bulb ;

posterioly-directed spines cover the anterior half or two-thirds

of the body, the anterior being comb-like, with four subequal

points, while the three-pointed spines have typically the middle

point the longest; in the male, small spines with the points directed

away from the cloacal opening cover most.of the ventral aspect of

the posterior 0°8 mm. of the body; right spicule three or four

times as long as the left; tail of the female, in ventral view,

uniformly rounded, with very massive papille.

For list of hosts, see p. 304.

2. GNATHOSTOMA HisPprpUM * Fedchenko, 1872.

Gnathostoma hispidum Fedcheuko (1872, p. 106; pl. xv.).

Cheiracanthus hispidus v. Linstow (1893, p. 201; pl. vil.

figs. 1-16).

Cheiranthus hispidus v. Linstow (18938, p. 202) [misprint].

The description which follows is based partly on a translation

which we have privately obtained of the essential parts of

Fedchenko’s Russian paper T, in which he describes material from

* For specific diagnosis, see p. 300.

+ The principal contents of the paper are rendered more accessible through its

Latin summary, and through an abstract of it in German by Leuckart (1873).

THE NEMATODE FAMILY GNATHOSTOMID. 299

the wild pig of Turkestan and the domestic pig of Hungary, and

partly on descriptions of material from Hungarian pigs by

Csokor (1882), v. Ratz (1900), and vy. ee (1893), and from

Rouen pigs by Ciurea (1911). A paper by Strése (1892),

and one by Collin (1893) in which he claimed to have found ifs

species in the ox in Berlin, we have been unable to consult. The

last is of minor importance in that Collin later authorised

Wolfthiigel (1912) to withdraw this statement, since, in the cir-

cumstances under which the material was received, he could not

exclude the pig as the possible host. In these descriptions

variations from the subfamily and generic characters detailed

above (vide pp. 254, 291) have, as we believe incorrectly, been

described by v. Linstow and Ciurea.

The anterior body-spines (vide Table VII., p. 802) have sub-

equal points, which accordingly, asin Gnathostoma spinigerum,

terminate at about the same level. More posteriorly the median

point tends to jnaeonue the longest (Fedchenko), the others

gradually disappearing, so that at about the posterior end of the

cesophagus the spines “are simple but long. They extend as such

over the rest of the body except, judging by v. Linstow’s figures,

the large caudal papille of the male. Of these there are four pairs,

No. 1 being the smallest and separated by an interval from the

other three, which he close together and are of about the same

size (Ciurea). Of sessile papillee Fedchenko figures a pair between

the bases of Nos. 1 and 2 of the larger ones; v. Linstow omits

these but describes a similar pair close to one another and anterior

to the cloacal opening, while Cinvea, in addition to the two pairs

which we have described as genevic characters, finds three more

pairs, two in front of and one behind the cloacal opening. By alk

these authors the termination of the tail is counted and described

as an unpaired median papilla.

The vulva lies in the middle of the body (Fedchenko) or some-

what behind (v. Linstow) or in front of (Csokor, Ciurea) this

point. Csokor’s description of the cesophagus suggests that the

anterior part of his specimens was much egnunaceel which may

explain the anterior position of the valva in them. We have

disregarded these statements in the description of the family.

Ciurea and v. Linstow agree in describing the egg as having a

hyaline appendage or wart-shaped structure at one pole; the

former describes the outer surface of the shell as showing small

dimples, the latter finds it smooth. It is stated by v. Linstow

that in his specimens embryonic development had begun, and

that at the vulva there were a number of many- nucleated cells,

which have, however, in his figure an appearance very suggestive

of ova.

Except for Collin’s statement, later withdrawn, the described

habitat (Table VIIL., p. 304) has always been the stomach-wall

of the pig, penetration into which may be partial or complete,

the worms in the latter case (Fedchenko) lying between the

gastric tunics.

300 MR. H. A. BAYLIS AND LT.-COL. CLAYTON LANE ON

Specific Diagnosis.

GNATHOSTOMA HiIsprpuM Fedchenko, 1872.

Gnathostoma: nine to eleven rows of hooks on the head-bulb ;

posteriorly directed spines cover the whole of the body, the

anterior being comb-like, with seven points, and shorter than

the more posterior. The left spicule is twice as long as the right.

For list of hosts, see p. 304.

3. GNATHOSTOMA HORRIDUM (Leidy, 1856).

Cheiracanthus horridus Leidy (1856, p. 53).

Under the name of Cheiracanthus horridus Leidy described

from the stomach, presumably from the stomach lumen, of

Alligator mississippiensis four females, 22 inches long and a line

and a half thick, with the body “ eylindr ical, incurved, posteriorly

subclavate, obtuse; anteriorly covered with palmate plates

furnished with as many as eight spines and degenerating

posteriorly to simple spines.” It is uncertain whether this last

expression implies that the spines reached the posterior end of

the worm.

The want of mention of any burrowing and the fact that the

host was a reptile and predatory are in favour of the belief that

these worms were really parasites of some host devoured by the

alligator; while, with the possible exception of size, there is no

characteristic by which this worm can be distinguished from

Grathostoma hispidum. Regarding this last point it is probably

not disputable that a dead or dying worm commonly increases

in size as its muscles relax and decomposition begins.

Gnathostoma hispidum and G. horriduwm are not, however, here

described as synonyms, partly because no certain conclusion is

possible from the evidence, and partly because to do so would

necessitate the substitution of a name based only on females very

imperfectly described for one based on specimens which have

been investigated with considerable thoroughness. No specific

diagnosis of G. horridwm is attempted.

4, GNATHOSTOMA GRACILE (Diesing, 1838).

Cheiracanthus gracilis Diesing (1838, p. 189), nomen nudwm.

Ks bs Diesing (1839, p. 2255: pl. xiv. figs. 8-11;

pl. xvii, figs. 1-20).

os v. Drasche (1882 2, p. 126; pl. ix. figs. 1-2).

[ Not Echinocephalus 9 gracilis Stossich (in Shipley and Hoe nell,

WO gs teS))4 |

Apart from its length (vide Table VII. p. 302) this nematode

has the following specific characters. ‘The spines on the anterior

part of the body are leaf-lke, with a maximum of five points,

one at the tip and the others along the lateral edges, two on

THE NEMATODE FAMILY GNATHOSTOMIDA. 301

each border. Hach cervical sac displays a constriction near its

posterior end, a condition, however, which v. Drasche (1882,

p. 126) looks upon as a temporary local contraction; the spicules

are stouter than in “Chetracanthus robustus”; the tour large

lateral caudal papille of the male are so arranged that there is

a considerable interval between Nos. 1 and 2; the egg is figured

as being without a polar cap, and the habitat was the intestinal

canal of a fish which reaches a length of 15 feet and is presumably

carnivorous.

The great length and unusual habitat suggest, as in the case of

Gnathostoma horridum, that the real host was some mammal which

was devoured by the ‘ Pirarucu.”

Specific Diagnosis.

GNATHOSTOMA GRACILE (Dies., 1838).

Gnathostoma: anterior body-spines leaf-like, with five points,

one at the tip and two along each edge.

5. GNATHOSTOMA TURGIDUM Stossich, 1902.

Gnathostoma turgida Stossich (1902, p. 13).

This species is based on a short description of the external

characters of two poorly preserved females. Stossich is convinced

that it is distinct from Gnathostoma spinigerwm, basing his con-

viction on the statement that the discoidal head-bulb has from

10 to 12 rows of spines, the body is cylindrical and tapers in

both directions, and its spines are of varying shape. There is no

further information beyond the details to be found in Table VIL.,

p. 302, and Table VIII., p. 304. The name is likely to be a source

of future confusion. Since there is nothing specifically distinctive

in the description no specific diagnosis is possible.

Species which have been attributed to the Genus Gnathostoma.

GNATHOSTOMA SHIPLEYI Stossich, 1900.

Gnathostoma shipleyi Stossich, in Shipley (1900, p. 560, fig. G).

Lictularia paradoxa v. Linstow (1903, p. 272; pl. xviii. fig. 5).

Acuaria pelagica Seurat (1916, p. 785, figs. 1-5).

Seuratia shipleyi Skrjabin (1916, p. 971).

Seurat’s careful description of the female of this species dis-

closes neither cervical sacs nor ballonets. The absence of these

and of trilobed lips excludes the worm from the Gnathostomide.

GNATHOSTOMA PELECANI (Chatin, 1874).

Sclerostoma pelecant Chatin (1874, p. 6; pl. viii. fig. 12; pl. ix.

figs. 1, 2).

Gnathostoma pelecant Skrjabin (1916, p. 972).

302 MR. H. A. BAYLIS AND LY.-COL. CLAYTON LANE ON

Taste VII.—Measurements of

(All measurements

far i |

| G. spinigerum.

= |

: iy | Diesing, ;

Described by ...... L Owen. Dajardin, =| ~~ Schneider. Leidy. Levinsen.

| v. Drasche. |

Under the name of {| Gnathostoma Cheiracanthus Filaria Gnathostoma Cheir.

¢ spinigerum. robustus. radula. sociale. siamensis.

| 3. ?. 3 2. a

J UEREMID se peontnnee one Gouee eae eae 99 °° 18 25 10 to 12 10 to 12 25 31 9

Thickness (maximum) 2 25 | 2 il 115 1

Irene throtiread lyase. eet eirn ere |

Leneth of head- bulb . Saeubabaee sal

Transverse diameter of head-bulb

Length of ballonets ....... |

Distance from head-end to ter- 2Q|

mination of cervical sacs ... §

Length of neck of cervical saes |

| Breadth of lips Bre we

Number of rows of hooks on |

head-bulb.. sade 6 or 7 6 or 7 | se 8

Leneth of hooks on head-bulb .. | ae | aN

Breadth of hooks on head-bulb... | ae | Br

Distance between rows of hooks|

on head-bulb | a6 ne

Distance between rows of body-| |

spines ..... baal no | an aiate

Portion of body covered by ) anterior {

| __spines . a) | two-thirds.

| Maximum number of. points on) |

body-spines ...... 5 4. 3 3 inn

| Distance from head. end to. cervi-

cal papillee nat snae ae

Distance from head- end toi nerve-

Dee from head-end to ter- 2 |

mination of csophagus ..,... §

WeONerilee OE ipl soonccsccosaeudeasece ss |

| |

- |

Length of spicules........... ...... ‘

| !

Distance from tip of tail to 2 8 |

PV Als eigen since eee AN

Ova (maximum measurements) .

the Species of Gnathostoma.

in millimetres.)

THE NEMATODE FAMILY GNATHOSTOMIDA.

303

ie los CS G. G,

Cop Snider. | gracile. | horridum. | turgidum.

uy, ob Ee z

Leiper. Baylis & Lane. v. Linstow. Csokor. Ciurea. Wy Re Leidy. | Stossich.

Cheir. Gnathostoma i

siamensis. spinigerum. i

relies | ies | ae ae Tia oa |

eas 6-4 to || lo to 22 tO Gyn eel 9 to 2 to ae .

Wei liatere| ins || ia | 35 | 2 | 31 | 9c? | Mase, 36 5 ae

: t | LEIS TO | IAD |G < | ss

0-4 0°32 OFS || @FSY/ |

28) ae or42 | ! |

0°525 |

0°55 to 075 |

0-6

ie ais 0°25

va Gi) 28) ro RS ‘

1 18 v| 20 0°88 : O07 | |

460 03 | || !

0719 |

oe 15 9 to 0:2 | | |

0°25 | |

8 8 to 11 leet. 12 | 10 to 12

0:015 0°025 | 0°029 | 0°018

0°005 0°01 ah ee 007) | |

0°023 | | |

‘ 0-017 ! 0:22 | |

anterior anterior i} é ' anterior

5 heule two-thirds. |, ule: half.

4 7 ai 9 | 5 8

| |

0°7 to 0°75 |

05 | |

315 | 34 || 1/55 |

LORI ema tO of 343 | 65 | |

Si | total length. |

O22 | O15 || : 0312

ae z e A {stout 2 is

R. ithe R. L. 1 Ri. Longer. |Short’r.| |

: Ne 2°3 to O46 | : am eee ss | |

iil O4 2-63 08 | O-4 | 0°88 | 1:29 0°32 | ! | |

4. | Somewhat pe ong | Tn front i | |

| behind middle. Fiddle thirds (Goodale:

0:06 X 00385 = ||}:«0'072 X 0039s! GO0741 x 0:0418 |!

| |

302

Under the name of {

CEI ian mnontincnon snags enatencotn, ben

—Thiekness (maximun)

TUGHY pe UNIO Celi Gh Cline yarns mvennnstees

Length of head-bulb

Tangverse diameter of head-bulb

Length of ballonets

Distance from head-end to Py

mination of cervical sacs...

Length of neck of cervical sacs .

Breadth of lips

Number of rows of hooks on

hhead-bulb...

Lenath of hooks on hend- ‘bulb...

| Breadth of hooks on head-bulb..

Distimce between rows of hooks

on head=bwhb 1.0...

Distance between rows of Dody-

Sie

Dintaine an weatateatilt to fis.

mination of eS. i}

Length of tail

ven en eee

Length of spicules —

| Dist tinee a ba of fol

MR. H. A. BAYLIS AND LY?.-

COL. CLAYLON LANE ON

Taste VII.—Measurements of

(All measurements

Owen.

Gnathostoma

spinigerun.

6 or7

two-thirds.

Diesing,

Dajardin,

v. Draschie.

Cheiracanthus

robustus.

10 to 12

6 or 7

G. spinigerun.

Schneider. Leidy. Levinsen.

Filaria Gnathostoma Cheir.

radula. sociale. siamensis.

é. 2 or

10 to 12 25 31 9

2 1 115 1

; 8

3 3

THE NEMATODE FAMILY GNATHOSTOMID.2,

the Species of Gnathostoma.

in millimetres.)

G. hispidum.

i gracile.

Leiper. Baylis & Lane. v. Linstow. Csokor, Ciurea Diesing,

. iv. Drasche.

Cheir. Gnathostoma |

siamensis. spinigerun. |

| Ape. s sien

es fe we 16 to | 29/ he fee é. Peal

se rs 0) B) 22 to 19to | 32t ;

Wd» ‘ to1675| 183 18 25 25 Se neal ee 36

5 é : 1:18 to | 1°78 to 2 d

06 19 12 1:38 1:85 2 25 = | 5 )

Ord 0°32 ae oe 0°37 0:37 |

0°23 5c O42

0525

05 to 0°75

Doo 0°25

(| 1:9 to ‘ d

1 18 v| 2-0 0°88 ' 07

See 03

019 }

elo to ()

025

8 8 to 11 9 to 11 12

0015 0°025 07029 0018

0005 0-01 a 0-007 )

0°023

igs 0-017 0:22

unterior anterior

+ half two-thirds. whole. ci )

4 7 el gr ieee

O°7 to O75 !

(0s)

315 | 34 1 Be .

to to 343 | 65

3°5 4 total Teeth

O22 | O15

303

horvidum,

Leidy.

tengidumn,

—_————___

Stossich,

10 to 12

half,

304

MR. H. A. BAYLIS AND LT.-COL. CLAYTON LANE ON

Immature forms, 3 mm. long, without sexual organs, were found

by Chatin encysted in the subcutaneous tissue, and immature

females, not less than 4 mm. long, encysted in the subscapular

air-sac, of a pelican (Pelecanus onocrotalus).

showed four transverse rows of hooks.

unequal length, in the position of the cervical sacs.

is made of any spines on the body.

The rounded head

There were six tubes of

No mention

These features necessitate

the provisional inclusion of the species in the Gnathostomide,

but, if the details are correct, exclude it from any of the genera

that we have described.

Taste VIII.

Habitat, Hosts and Distribution of the Species of Gnathostoma.

Parasite.

G. spinigerum ...

Habitat.

| Gastric wall.

Subcutaneous

tissue.

Intestine.

G. hispidum

G. horridum

G. gracile

G. turgidum

Gnathostoma sp.

Gastric wall.

Stomach.

Host.

Felis tigris.

Felis sylvestris.

[F. catus fer. |

F. catus.

FF. pardus.

India.

F. concolor. Brazil.

Mustela vison. N. America.

Paradoxurus philip-| Philippines.

pinensis.

Canis familiaris. Pim «

- a India.

eee cust |

Homo sapiens. Siam.

Rattus rajah. Island of Menta-

wel.

Sus scrofa ferus. Turkestan.

Sus scrofa domesticus., Hungary.

ty) 3) 22

39 33 »

33 33 39

a ha

Alligator mississip- | N. America.

piensis.

Intestinal canal.

LF]

Wall of large

intestine.

Arapaina gigas | Brazil.

[= Sudis gigas

= Vastres cuvieri]..

| :

Didelphis azare. | Argentine.

| Aon

Leontocebus sp. | French Guiana.

| Diesing (1889).

Cobbold (1879).

|v. Linstow (1893).

Mitter (1910).

Mitter (1912).

Baylis & Lane

(present paper).

Leidy (1859).

Schneider (1866).

Mitter (1912).

Levinsen (1899) ;

Leiper (1909).

Porta (1909).

Fedchenko (1872).

Czokor (1882). |

v. Ratz (1900).

Leidy (1856).

Diesing (1839).

Stossich (1902).

Weinberg & Bri-

mont (1909).

THE NEMATODE FAMILY GNATHOSTOMIDA. 305

GNATHOSTOMA ACCIPITRI Skrjabin, 1915.

We have been unable to consult a paper in which, under this

name, Skrjabin has described a parasite from an eagle in

Turkestan. We have no data at all on which to base any

comment.

GNATHOSTOMA sp., Weinberg and Brimont, 1909.

These authors give (1909, p. 104) a detailed report on certain

lesions produced in the large intestine of a “'Tamarin” (a small

monkey) in French Guiana by parasites which they identify as

Gnathostomes. The habitat mduces a natural doubt as to whether

the parasites may not have been (Hsophagostomes, and a study

of the text does not completely dissipate this, since no actual

description of the worms is given. The following s sentence occurs:

“Muni d’un nombre considérable d’ épines et de lamelles chitineuses

trés solides, le Gnathostome enfonce profondément son extrémité

céphalique, et peut ainsi amener, par ce seul moyen mécanique,

une rupture de la paroi intestinale.” It is not clear whether the

expressions used here refer to Gnathostomes in general or to

these worms in particular, so that it appears wise to suspend

judgment as to the systematic position of these parasites till they

have been properly described.

For Cheiracanthus uncinatus and Cheiracanthus spinosissimus

see Lchinocephalas (supra, p. 273), to which genus they belong. }

Genera doubtfully to be attributed to the Gnathostomide.

ANCYRACANTHUS Diesing, 1839.

Ancyracanthus Diesing (1839, p. 227).

Ancryacanthus v. Linstow (1893, p. 205).

Genotype: Ancyracanthus Patani Diesing (1839, p. 227;

Dibesive dies 21—27 5 pl. xvas figs. 1-20).

Aneyr acanthus pinnatifidus v. Drasche(1884, p. 111;

pl. iv. figs. 6-11).

These nematodes are characterized by the fact that the head

bears four appendages set cross-wise, two springing from each of

the lateral lips. Hach appendage is pinnate and is connected

with a cervical sac. Furthermore, from the attachment between

appendage and sac springs a long conical process, nearly as long

as and lying close beside the latter. Diesing and von Drasche

are in disagreement regarding the existence of a communication

between the cavities of the sac and of the appendage, which,

taking into consideration the complicated branching character of

the iatter, is not surprising. It is stated by von Drasche that

the cervical sacs have no spiral layer.

Habitat: gut of Podocnenis expansa and P. tracaxa.

306 MR. H. A. BAYLIS AND LT.-COL. CLAYTON LANE ON

Enaeepeeea de aa 1860.

Hlaphocephalus Molin (1860, p. 343).

Genotype: Llaphocephalus octocornutus Molin (1860, p. 344).

y. Drasche (1884, p. 113;

pi. ii. figs. 21-23).

This genus and species, based on a single female from Ara

| Psittacus| macao, ave characterized by the possession of four

cervical sacs which are without the internal processes found in

Ancyracanthus. Asin that genus, there are four freely-projecting

external appendages, each of which, however, possesses an

external process shorter than itself. The cuticle of the body

carries closely-set spines.

Whether <Ancyracanthus pinnatifidus and Llaphocephalus

octocornutus do or do not belong to the Gnathostomide can only

be ascertained after further investigation, but it is not impossible

that the structures just mentioned correspond to the ballonets

and cervical saes of the Gnathosiomine. If this be so, the

difference between the subfamily and these two forms presumably

lies in an exuberant development, in the latter, of the ballonets,

unconfined by a stout overlying covering. In the forms

unquestionably belonging to the subfamily such confinement

within a uniform bead-bulb is complete in Gnathostoma and

Hehinocephalus, less complete in Tanqua anomala and T'. diadema,

aud least so in Yangua tiara, in which last the prominences

corresponding to the four underlying ballonets are sometimes

strikingly distinct. It is not difficult to imagine that further

exuberance and external subdivision or indentation of the cuticle

covering the ballonets might well produce those external and

internal modifications which have been described in these two

genera.

be) 99

Nove

Our best thanks are due to Mr. T. Southwell, formerly Director

of Fisheries for Bengal, Bihar and Orissa, for supplying us with

much of the material upon which the work has been based,

expecially as regards the genus Hehinocephalus.

As vegards the nomenclature of the hosts, we are indebted to

Mr. Oldfield Thomas, F.R.S., for verifying and correcting the

names of all the mammals mentioned in the paper; and to

Mr. C. Chubb, Dr. G. A. Boulenger, F.R.S., and Mr. C. Tate

Regan, F.R.S., for performing the same kind office in the cases of

birds, reptiles, and fishes respectively.

For the Lye ea of some excellent serial sections we are

indebted to the skill of Mr. Cecil Gunns, of the Zoological

Department, Imperial College of Science.

For all errors and omissions we accept full responsibility.

The figures have all been drawn to scale with the aid of the

Abbé camera lucida.

The type-specimens of the species described as new are in the

British Museum (Natural History).

THE NEMATODE FAMILY GNATHOSTOMIDZ. 307

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22 33

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308

Lerper, R. T.

LeucKkart, R.

Levinsen, G. M. BR.

Lriystow, O. von.

” Lb}

’ 2

22 3

” 2

Mirrer, S. N.

2 ”

Motrin, R.

Monricett, F. 8.

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Porta, A,

MR. H. A. BAYLIS AND LT.-COL, CLAYTON LANE ON

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THE NEMATODE FAMILY GNATHOSTOMIDA. 309

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bb) bb)

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*SCHNEIDER, J. G.

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39 2)

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Hornet, J.

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aly

310

ON THE NEMATODE FAMILY GNATHOSTOMIDA.

Explanation of Lettering of Teat-figures and Plates.

a., button-like appendage of cervical

Sac.

b., ballonet.

b.c., cavity of ballonet.

e., collar.

c.p., cervical papilla.

e.s., cervical sac.

c.s.n., neck of cervical sac.

c.t., cuticular thickening.

ed.p., caudal papilla.

e.p., excretory pore.

@.8., EZE-SAc.

h., hooks.

h.b., head-bulb.

h.m., head-muscele.

1., lip.

EXPLANATION

Mie. 1. Spiroxys contorta.

2. 3) ”

3. ’ ”

A. cy) 29

5. b) 33

l.p., lateral papilla.

l.p.n., nerve of lateral papilla.

my7., Nevve-ring.

ces., cesophagus.

r.g., rectal glands.

S., spines.

ser., serrations.

s.m.p., submedian papilla.

s.m.pn., nerve of submedian papilla.

sp., spicule.

sucker-like depression.

teeth.

termination of tail.

uterus.

vulva.

vagina.

OF PLATES I-VIII.

The head ; dorsal (or ventral) view.

The head; lateral view.

Tail of male; ventral view.

Tail of male; lateral view.

Tail of male; lateral view, at a higher magnification,

(Right spicule broken).

to show the papille.

The head; dorsal (or ventral) view.

ie of $6) The head; lateral view.

8. x 5 Caudal extremity of male; ventral view.

9. ; 3 Caudal extremity of male; lateral view.

10. rr i Terminal portion of spicule.

11. Tanqua tiara. The head; nearly dorsal view.

12. 55 45 Tail of male; ventral view.

13. A - Tail of male; lateral view.

14. Tangua anomala. Tail of male; lateral view.

15. Tanqua diadema. Tail of male; ventral view.

16. ss Tail cf male; lateral view.

17. Echinocephalus spinosissimus. Tail of male; nearly ventral view.

18. a - Terminal portion of spicule.

19. Hchinocephalus sowthwelli. The lips; dorsal (or ventral) view.

20. 45 5 One of the lips: lateral view. The cuticular

thickenings are seen by transparency.

2 “5 Tail of male; ventral view.

22. 3 es Tail of male; lateral view.

23. > ij Terminal portion of spicule.

24. Echinocephalus multidentatus. The lips; dorsal (or ventral) view.

25. Fr < One of the lips; lateral view. The teeth

are seen by transparency.

26. = Tail of male; ventral view.

27. Sy a Tail of male; lateral view.

28. i 3 Terminal portion of spicule.

29. Gnathostoma spinigerwm. Anterior end: lateral view (optical section).

30. ; i Surface view of one of the cervical papille,

surrounded by spines.

31. 9 Profile view of one of the cervical papillee.

32. 3 5 Tail of male; ventral view. On the left-hand

side of the drawing most of the spines

have been omitted.

33. 3 5 Tail of male; lateral view.

34, re ws The spicules.

35. 5 4s Terminal portion of left spicule.

36. 9 i Tail of female; ventral view.

BY/- 53 3 Tail of female; lateral view.

38, a. Ova, some with embryos escaping,

ON BORNEAN DRAGONFLIES. Balt

17. Contributions to a Study of the Dragonfly Fauna of

Borneo.—Part IV. A List of Species known to occur

in the Island. By F. F. Latptaw, M.A.(Cantab.),

E.ZS.

‘Received April 9, 1920: Read April 27, 1920. |

(Text-figures 1-4.)

INDEX. Page

Teeainibrod chr OMla.Haeairecnt cosas serase is dseccniciee eee ede ae Le

PILI St OF OPECIES ie ete esses ecienl hs he chs Ween aL,

III. Some Remarks on Geographical Distribution ...... 341

I. Introduction.

I have attempted in the following list to catalogue all species

of Dragonflies which have been recorded from Borneo, or which

are otherwise known to me to occur in the island.

In the case of some species I have added notes that may, I

hope, be of service to others who may deal with this fauna.

In addition to records already available in the literature of the

order O.lonata, I have been able to compile my list from the

examination of the large amount of material sent me by Major

J.C. Moulton whilst Curator of the Sarawak Museum.. Some of

this material has already been dealt with, as noted in the

references to literature; and Major Moulton’s type specimens

are, unless otherwise noted, deposited in the British Museum.

Where possible, paratypes have been sent to the Sarawak

Museum.

I have also been able to examine a collection sent to me nearly

twenty years ago by Dr. C. Hose from the Baram district.

References to specimens examined deal in every case with

Major Moulton’s material unless otherwise stated.

His localities are in the territory of Sarawak, with the following

exceptions, which are in British North Borneo :—Mt. Kinabalu,

Tampassuk (River), and Khotabelud.

In order to avoid undue prolixity in the list, I have given

references chiefly, if not entirely, to faunistic and systematic

papers. For the Libelluline I have thought it necessary to quote

only Dr. Ris’s Monograph of the subfamily, referred to in the

sequel as “ Ris, Cat. Coll. Selys, Libell.” (‘Catalogue Systématique

et Descript. Collections Zoologiques du Baron Edm. de Selys-

Longchamps,’ Fasc. ix.-xvi., Libelluline). The publication of the

Monograph was completed in 1919, and it contains a very full

synonymy of all the Bornean species.

For the Gomphine:and Chlorogomphine. I give reference only

to my previous paper on the group (these Proceedings, 1914,

pp. 51-63, pl. i.), where citations of recent papers are to be found,

Slee MR. F. F, LAIDLAW ON

For species of other groups I quote, where available, Kirby’s

‘Synonymic Catalogue of Neuroptera Odonata’ (1890) as “* Kirby,

Cat. Odonata,” and, in addition, references to later papers. Of

these, Fase. xvi. of the ‘ Catalogue of Collections du Baron Edm.

de Selys-Longchamps,’ dealing with the Corduline, by Mavtin,

is referred to as “ Martin, Cat. Coll. Selys, Cordul.” ; whilst Fasc.

Xvill.xx. of the same Catalogue, dealing with the Alschnine,

are referred to as ‘‘ Martin, Cat. Coll. Selys, Adschn.”

I have in certain cases given measurements of the specimens

noted, in each case of the abdomen “abd.” and hinder-wing ‘h. w.”

In the case of the male, after the measurement of the abdomen,

the sign + followed by another measurement refers to the length

of the anal appendages.

Where the venation of the wings is discussed, I have used the

revised nomenclature proposed by Tillyard in his book, ‘The

Biology of Dragonflies.’

An asterisk placed before the name of a species signifies that

I have not seen a specimen of that species from Borneo. ,

Parentheses around the names of authors placed after specific

names in this paper are used in accordance with Article 23 of

the International Rules of Nomenclature (Eroe. 7th Int. Cong.

Boston, 1907, p.-44 (1912)).

Il. List of Species.

ANISOPTERA.

ALSCHNIDA,

CHLOROGOMPHIN2.

1. OroGomMPHUS DYAK Laidlaw.

Orogomphus dyak Laidlaw, Proce. Zool. Soe. London, 1914,

pp. 99-60, pl. 1. figs. 4-7.

235,292 9. Mt. Matang. 3¢ $6. Mt. Merinjak, 2200 ft.,

28. v. 14.

. OROGOMPHUS SPLENDIDUS Selys.

Or ogomphus splendidus Laidlaw, Proc. Zool. Soc. London, 1914,

pp- 60-61, pl. 1. fig. 8.

29 2. Sarawak.

LHSCHNINE.

*LINE[SCHNA POLLI Martin.

Lineschna polli Martin, Cat. Coll. Selys, Aischn, pp. 136-137,

(figs. 133, 134), pl. iii. fig. 9.

4, JAGORIA MODIGLIANI (Selys).

Oligoeschna modighiantti Kirby, Cat. Odonata, p. 86.

BORNEAN DRAGONFLIES. 315}

Doleschna elacatura Needham, Bull. Amer. Mus. Nat. Hist.

23, p. 143, fig. 3.

? Jagoria peciloptera (pars) Karsch, Entom. Nachr. xv. p. 239

(3). |

Jagoria elacatura Martin, Cat. Coll. Selys, Adschn. p. 135.

Jagoria modighianti Kriger, Stettin. Entomol. Zeitg. 1898,

pp. 238-259, 327; Martin, Cat. Coll. Selys, Adschn. p. 130

(figs. 126, 127); Ris, Ann. Soc. Entomol. Belg. pp. 240-242.

229. Matang Rd.

I have followed Ris in the synonymy of this species.

* Wings suffused with golden brown throughout.

@. Abd. 40 mm., h. w. 42 mm.

5. *JAGORIA PGCILOPTERA Karsch.

Jagonia peciloptera Karsch, Entom. Nachr. xv. 1889, p. 238;

Kiiiger, Stettin. Entomol. Zeitg. 1898, pp. 327-328; Martin,

Cat. Coll. Selys, Aischn. pp. 132-133 (fig. 129).

6. *JAGORIA BUHRI Forster.

Jagoria buhri Forster, Insecten-Borse, 1903; Martin, Cat.

Coll. Selys, Adschn. pp. 131-132 (fig. 128).

This species recorded from N. Borneo by Dr. Forster appears te

me to be very closely related to that immediately preceding it,

T have not seen examples of either of these two last species.

7. HELLESCHNA IDH (Brauer).

Amphieschnu? ide Kirby, Cat. Odonata, p. 93.

Helieschna ide Kriiger, Stettin. Entomol. Zeitzg. 1898,

pp. 323-324; Martin, Cat. Coll. Selys, Aischn. pp. 164-165

(fig. 166); Ris, Ann. Soc. Entomol. Belg. lv. 1911, pp. 242-243

(fig. 10).

ie Metane! ids 2.2 ll lo kuching, 4.1.98. 16.

Buntal, iii.12. 19. Matang Rd.,4.i:11. 19. Saribas, xi. 1900.

1 @. Kuching, iv. 03. .

$6. Abd. 524+7mm., hw. 52mm. @. Abd. 57+mm., h. w.

53 mm.

The females each have four large spines on ventral plate, the

median pair being larger than the lateral pair. The apex of the

plate is strongly decurved, and the margin carries smaller spines

on either side of the four larger ones. From the description and

figures (Martin, Joc. ci. figs. 163, 166) I cannot separate this

species from the next.

8. *Herrmscuna CRASSA Kriiger.

Heliceschna crassa Kriiger, Stettin. Entomol. Zeitg. 1898, pp-

324-325 ; Martin, Cat. Coll. Selys, Aischn. pp. 162-163 (fig. 163).

314 MR. F. F. LAIDLAW ON

9, *HELIASCHNA UNINERVULATA Martin.

Helieschna uninervulata Martin, Cat. Coll. Selys, Auschn. pp.

163-164 (figs. 165, 168).

This species, described originally from Borneo, occurs also in

Lower Burma. (1 6, coll. E. B. Williamson.)

10. *HeLimscHna sIMPLICIA (Karsch).

Amphicschna simplicia Karsch, Ent. Nachr. xvii. 1891, p. 308;

Karsch, Ent. Nachr. xviii. 1892, p. 250.

Helieschna simplicia Karsch, Ent. Nachr. xix, 1893, p. 195 ;

Martin, Cat. Coll. Selys, Aischn. pp. 161-162 (fig. 162).

11. *AMPHIASCHNA PERAMPLA Martin.

Amphiceschna perampla Martin, Cat. Coll. Setys, Aischn. p. 115

(figs. 108, 109).

12. AMPHIHSCHNA GRUBAUERI Forster.

Amphiceschna grubauert Forster, Insekten-Borse, xxi. (1904)

pp. 1-3 (sep.); Martin, Cat. Coll. Selys, Aischn. pp. 115-116

(fig. 110); Laidlaw, Journ. Straits Branch R. Asiat. Soc. lxiii.

1912, p. 94.

1 2. Mt. Batu Lawi, 24. v.11.

The arculus in the fore-wings of this specimen is of a primitive

character. The upper part is very oblique and readily seen to

be formed by M,_,, M,, which separate before M, is met by the

scarcely thickened cross-vein which forms the lower part.

The upper part of the hinder-wing is much less oblique.

The costal, subcostal, median, and cubito-anal spaces in both

fore- and hinder-wing are opaque, dark brown to a level of about

one cell before the arculus; the anal area of both wings tinged

with orange-brown to about the same level. Beyond the nodus

the wings are tinged with orange-brown almost to the apex.

The ventral plate curves upwards at its apex and carries about

8 small irregular spines. Its ventral surface has a deep conical

depression to receive the points of the terebre.

Abd. 62 mm., h.w. 63 mm. Pt. 2 mm.

(The measurement given by me (loc. cit.) of 70mm. for the

hinder-wing was an error.)

13. TerRACANTHAGYNA PLAGIATA (Waterhouse).

Tetracanthagyna plagiata Kirby, Cat. Odonata, p. 94; Kruger,

Stettin. Entomol. Zeitg. 1898, pp. 287-288; Martin, Cat. Coll.

Selys, Aischn. pp. 145-146 (figs. 144, 145).

OF Walkussivies:

Ris (loc. cit.) suggests that the next species, separated from

plagiata on account of the colouring of the wings of the female, is

identical with it ; and that 7’. plagiata has two forms of female, one

with a costal stripe and transverse band of brown on the wings

BORNEAN DRAGONFLIES, 315

(7. plagiata type), the other with costa] stripe only (7. vittata

type). No male with transverse bands has been recorded.

Structurally the females of the two forms are identical, and

Ris’s view is in all probability correct. In this case 7. vittata

McLach. will be a synonym of 7’. plagiata Waterh.

: ©. Abd. 68 mm., b.w. 80 mm.

[TErRACANTHAGYNA virraTA McLach. ]

Tetracanthaygna vittata Mclachlan, Trans. Entomol. Soc.

London, 1898; Martin, Cat. Coll. Selys, Adschn. pp. 144-145

(fig. 143); Ris, Ann. Soc. Entomol. Belg. lv. 1911, pp. 243-244.

IL @, il Qe linea, 78

See remarks under T. plagiata.

The female of this pair is possibly the largest and bulliest mdi-

vidual of all living Odonata on record. The pair in full flight

must have afforded a splendid sight to any entomological

enthusiast.

6. Abd. 70+7'5 mm., h. w. 72 mm.

©. Abd. 70 mm.. h. w. 83 mm.

14, *TerrRacANTHAGYNA DEGORsI Martin.

Tetracanthagyna degorsi Martin, Bull. Soc. Entom. France,

1895; McLachlan, Trans. Entomol. Soc. London, 1898; Kriiger,

Stettin. Entomol. Zeitg. 1898, p. 288; Martin, Cat. Coll. Selys,

Aischn. p. 147.

15, *TerRACANTHYGNA WATERHOUSEL McLach.

Tetracanthagyna waterhouse: McLachlan, Trans. Entomol. Soc.

London, 1898; Martin, Mission Pavie, } Névropteres, ( sep.) p. 14;

Martin, Cat. Coll. Selys, Adschn. pp. 143-144 (fig. 142).

16. TErRACANTHAGYNA BRUNNEA MecLach.

Tetracanthagyna brunnea Mclachlan, Trans. Entomol. Soe:

London, 1898; Martin, Cat. Coll. Selys, A’schn. pp. 146-147

(fig. 141).

Tetracanthagyna pla giata Laidlaw (nec Waterhouse), Proc.

Zool. Soc. London, 1902, p. 79.

(The specimen recorded by me, loc. cit. 1902, from the Malay

Peninsula belongs here and not te 7. plagiata.)

1 9. Sarawak: h. w. 66 mm., abd. 55 mm.

17. GYNACANTHA DEMEYER Ris.

Gynacantha demeter Ris, Ann. Soc. Entomol. Belg. liv, WO,

pp. 245-246 (fig. 12).

16,192. Matang Rd., 22.11. 20.

18. GyYNACANTHA DOHRNI Kruger.

Gynacantha dohrni, Kruger, Stettin. Entomol. Zeitg. 1898,

pp. 277-287, fig. p. 280; Martin, Cat. Coll. Selys, Atschn.

316 MR. F. F. LAIDLAW ON

pp. 199-200 (fig. 204); Ris, Ann. Soc. Entomol. Belg. lv. 191],

pp. 244-246 (fig. 11).

23 6. Saribas. 1g. Quop, 19.11.14. 19. Mt. Merinjak,

23.v.14. 29 2. Matane Rd.,1l.iv.12. 19. Kuching, 14.1. 96.

The collection also contains 3 2 Q which I have not been able

to identify with certainty. They are from Tabekang, 13. v. 14,

and Kakus, vi. 1913. In these specimens the wings are suffused

with orange-brown. The material at my disposal is unfortu-

nately insufficient to permit a satisfactory handling of the genus.

19. *GYNACANTHA BASIGUTTATA Selys.

Acanthagyna basiguttata Kirby, Cat. Odonata, p. 95.

Gynacantha basiguttata Kriiger, Stettin. Entomol. Zeitg. 1898,

pp. 277-284, fig. p. 279; Martin, Mission Pavie, Neévropteéres,

(sep.) p. 14; id, Cat. Coll. Selys, Aschn. pp. 192— 193 (Giles SIs

[2 fig. 107 4. aoliemi Kriiger, see Ris, loc. cit.]; Ris, Ann. Soe.

Entomol. Belg. lv. 1911, pp. 246-247 (fig. 13).

20. *GYNACANTHA BAYADERA Selys.

Gynacantha bayadera Selys, Ann Mus. Civ. Genova 2. x. (xxx.)

1890, p. 51; Kriiger, Stettin. Entomol. Zeitg. 1898, pp. 277-283,

fig. p. 280; Martin, Mission Pavie, Névropteres, (sep.) p. 14;

Martin, Cat. Coll. Selys, Aaschn. pp. 196-197 (fig. 200); Ris,

Ann. Soc. Entomol. Belg. lv. 1911, pp. 244-245;-Ris, Nova

Guinea, xiii. Zool., Livr. 2, pp. 111-112.

21. *GYNACANTHA MACLACHLANI Kriiger.

Gynacantha maclachlan Kruger, Stettin. Entomol. Zeitg. 1898,

pp. 277-287, figs. p. 280 and p. 319; Martin, Cat. Coll. Selys,

Atsehn. pp. 200-201 (fig. 205).

22. *GYNACANTHA HYALINA Selys.

Acanthagyna hyalina Kirby, Cat. Odonata, p. 95.

Gynacantha hyalina Selys, Ann. Mus. Civ. Genova, 2. x. (xxx.)

1890, p. 50; Kriiger, Stettin. Entomol. Zeitg. 1898, pp. 277-283 ;

Martin, Cat. Coll. Selys, Aichn. pp. 198-199 (fig. 208).

23. ANAX Gurratus (Burm.).

Anax guttatus Kirby, Cat. Odonata, p. 84; Laidlaw, Proc.

Zool. Soc. London, 1902, p. 78; Martin, Mission Pavie, Névrop-

téres, (sep.) p. 14; Martin, Cat. Coll. Selys, Auschn. pp. 23-24

(fig. 17); Ris, Senckenbers Naturfors. Gesellsch. xxxiv. 1913,

p. 92

G5 Oo hit Wien O. sai, 4b,

24, *ANACIHSCHNA JASPIDEA (Burm.).

Anacieschia jasprdea Kirby, Cat. Odonata, p. 86; Kriiger,

Stettin. Entomol. Zeitg. 1898, pp. 274-275; Martin, Cat. Coll.

Selys, Afschn. pp. 30-31 (fig. 25).

| For

BORNEAN DRAGONFLIES. 317

GoMPHINA.

references to literature, see Laidlaw, Proc. Zool. Soc.

London, 1914, pp. 51-63: “Contributions to a Study of

the Dragonfly Fauna of Borneo.—Part Il. The Gomphine

and Chlorogomphine ” (plate i.). |

25.

36.

37.

Icrinus AcutUS Selys (MSS.).

. *ICTINUS DECORATUS Selys.

. IcrInus MELZNorS Selys.

. GOMPHIDIA MACLACHLANI Selys.

*GOMPHIDIA KARSCHI Selys.

SIEBOLDIUS JAPONICUS Selys.

*MACROGOMPHUS ALBARD#& Selys.

. MacroGoMPHUS DECEMLINEATUS Selys.

3. MACROGOMPHUS QUADRATUS Nelys. °

MicROGOMPHUS CHELIFER Selys.

* LEPTOGOMPHUS SEMPERI Selys.

LEPTOGOMPHUS WILLIAMSON! Laidlaw.

BURMAGOMPHUS VERMICULATUS Martin, subsp. insularis

Laidlaw.

38.

law.

HETEROGOMPHUS ICcTEROPS Martin, subsp. borneensis Laid-

39. HETEROGOMPHUS SUMATRANUS Kriiger.

Heterogomphus swmatranus Laidlaw, Proc. Zool. Soc. London,

1917, p. 232.

40.

LIBELLULIDA.

CoRDULIIN#.

HEMICORDULIA ASSIMILIS Selys.

Hemicordulia (%) assimilis Kirby, Cat. Odonata, p. 46.

Hemicordulia assimilis Laidlaw, Proc. Zool. Soc. London, 1913,

pp. 64-65; Martin, Cat. Coll. Selys, Cordul. pp. 11-12; Ris,

Nova Guinea, ix. Zool. 3, pp. 501-502 (fig. 21).

Al.

AZUMA AUSTRALIS (Hagen).

Epophthalmia australis Kirby, Cat. Odonata, p. 54; Martin,

Cat. Coll. Selys, Cordul. p. 63; Ris, Ann. Soc. Entomol. Belg. ly.

1911, pp. 248-250 (figs. 14, 15).

318 MR. F. F. LAIDLAW ON

42, *AZUMA VITTIGERA (Ramb.).

Epophthalmia vittigera Kirby, Cat. Odonata, p. 54; Martin,

Mission Pavie, Névropteres, (sep.) p. 8; id., Cat. Coll. Selys,

Cordul. pp. 62-63.

43. Macromia crncTa Ramb.

Macromia cincta Kirby, Cat. Odonata, p. 55; Kriiger, Stettin.

Entomol. Zeitg. 1899, p. 325; Martin, Cat. Coll. Selys, Cordul.

p- 68; Laidlaw, Proc. Zool. Soc. London, 1913, p. 69.

26 6. Baram.

Length of abdomen 49 mm., of hinder-wing 45 mm.

44, *MACROMTA BORNEENSIS Kriigev.

Macromia borneensis Kriger, Stettin. Entomol. Zeitg. i899,

pp. 330-332 ; Martin, Cat. Coll. Selys, Cordul. pp. 68— 69.

45. *MACROMIA GERSTAECKERI Kriger.

Macromia gerstaeckeri Kriiger, Stettin. Kntomol. Zeitg. pp. 335—

338 ; Laidlaw, Proc. Zool. Soc. London, 1902, pp. 76-78 ; Martin,

Cat. Coll. Selys, Cordul. p. 70.

46. MAcROMIA WESTWOOD! Selys.

Macromia westwoodi Kirby, Cat. Odonata, p. 55; Kriiger,

Stettin. Entomol. Zeitg. 1899, pp. 325-326; Martin, Mission

Pavie, Névroptéres, (sep.) p. 8; Laidlaw, Proc. Zool. Soc. London,

1913, p. 69; Martin, Cat. Coll. Selys, Cordul. p. 72.

1 ¢. Lio Matu, 30.x. 14.

Length of abdomen 42 mm., of hinder-wing 41°5 mm.

47, MAGROMIA EUTERPE Laidlaw.

Macromia euterpe Laidlaw, Proc. Zool. Soc. London, 1915,

pp. 26-29, figs. 1, 2.

266, 1@. Kinabalu. 16,19. Mt. Merinjak, 600 ft.,

24. v.14.

48, *MACROMIA CINGULATA Ramb.

Macromia cingulata Kirby, Cat. Odonata, p. 55; Martin,

Mission Pavie, Névropteéres, (sep.) p. 8; Martin, Cat. Coll. Selys,

Cordul. p. 70.

[Of the six Bornean species here recorded I have seen examples

of three only. The large female specimen referred to in a

previous paper (Proc. Zool. Soe. London, 1913, p. 68) is probably

a female specimen of Epophthalnvia australis Hagen.

Macromia euterpe Laidlaw appears to be closely allied to

M. westwocdi Selys. Both species are characterized by having

segments 1-6 of the abdomen of metallic-green lustre, whilst the

remaining four segments are black, save for the yellow basal

mark on 7.

On the other hand, the anal angle in the wings of the males are

BORNEAN DRAGONFLIES. 319

different in shape; that of J. ewterpe as shown in the figure

given (loc. cit. text-fig. 1,4) is rather deeply indented, that of

M. westwoodi much more nearly straight. But the readiest

means of discriminating the two species is afforded by the anal

appendages of the male. In the specimens of M. ewterpe I have

examined, the upper pair are in every case almost destitute of an

external tooth, and are but little shorter than the lower appendix.

In the case of J. westwoodi both the specimen before me and

that described by de Selys have the external tooth well

devoloped, and the lower appendage exceeds the upper pair in

length very considerably, projecting beyond them by nearly a

third of its total length when seen in profile.

For the rest I record M. cingulata Ramb. and M. gerstaeckeri

on Martin’s authority. |

49, MacromipiA FuLVA Laidlaw.

Macromidia fulva Laidlaw, Proc. Zool. Soc. London, 1915

pp. 29, 30, text-fig. 3.

1 g. Kinabalu. 1 9. Mt. Matang, 4. xi. 13.

The male is the type of the species and is in the British

Museum.

The female is very immature. The body-colour is a pale brown,

with darker bands at bases of abdominal segments 3-7, and there

is a brown basal spot on each wing extending nearly to aw,, the

vest of the wing being perfectly clear. Abd. 34 mm., h. w. 35 mm.

50. Iptonyx DouRNI Kriiger, subsp. BORNEENSIS.

Idionyx dohrni Kriiger, subsp. borneensis Laidlaw, Proc. Zool.

Soe. London, 1913, p. 67.

51. Meraprnya micans Laidlaw.

Metaphya micans Laidlaw, Sarawak Mus. Journ. No. 2, 1912,

pp. 65-67, pl. i.;,id., Proc. Zool. Soc. London, 1913, pp. 65-66

(Goll: hie 1s ls a)

The genus occurs in Borneo, New Guinea, and, I believe, in

New Caledonia.

LIBELLULINA.

Group I. (of Ris).

52. ‘TETRATHEMIS IRREGULARIS HYALINA Kirby.

Tetrathemis wregularis hyalina Ris, Cat. Coll. Selys, Libell.

p. 47.

2.6 6. Matang Rd., 25.11.10. 1 ¢. Retuh, 16.v.14. 1 6.

Limbang, 8.1x. 09.

53. *TETRATHEMIS FLAVESCENS Kirby.

Tetrathemis flavescens Ris, Cat. Coll. Selys, Libell.-p. 52.

1d. Sarawak. Autotype in Brit, Mus. 2 unknown.

320 MR. I. F. LAIDLAW ON ?

54, OpA DoHRNI (Kriiger).

Oda dohrni Ris, Cat. Coll. Selys, Libell. pp. 62-63 yo 25-28).

466,399. Matang Rd.

55, Hy L®orHEeMis CLEMENTIA Ris.

Hyleothemis clementia Ris, Cat. Coll. Selys, Libell. pp. 64-65

(figs. 29, 80); Laidlaw, Journ. Straits Branch R. As. Soe.

INO}, Ged, UDP, os Ss, jolla, ies, 3.

19. Mt. Batu Lawi (allotype). Sarawak Museum.

Group IL. (of Ris).

56. PoRNO?rHEMIS SERRATA Kriiger.

Pornothemis serrata Ris, Cat. Coll. Selys, Libell. pp. 92, 93

(fig. 64).

536,299. Matang Rd.

57. ORCHITHEMIS XANTHOSOMA Laidlaw.

Orchithemis xanthosoma Ris, Cat. Coll. Selys, Libell. pp. 1056-

1057 (fig. 615).

33636,229. Matang Rd. Type d in British Museum ;

allotype 2, coll. Selys.

58. ORCHITHEMIS PULCHERRIMA Brauer.

Orchithemis pulcherrima Ris, Cat. Coll. Selys, Libell. pp. 85-86

(figs. 54, 55).

19. Sadong Hill. 1¢,22 2. Matang Rd., 24.vi.09. 3d 3,

39 9. Baram (C. Hose).

*ORCHITHEMIS PRUINANS (Selys).

cena pruinans Ris, Cat. Coll. Selys, Libell. pp. 87-88

(figs. 56, 57).

60. LyRIorHEMIS CLEIS Brauer.

Lyriothemis cleis Ris, Cat. Coll. Selys, Libell. pp. 108-111 |

(figs. 78, 79, 80). :

1g. Limbang. 2466. Mt. Murud, xi-xii. 14. 1 9. Tatau.

1 g. Selindong, 12.vi-xi. (266,19. Baram; C. Hose.)

61. *LyRIOTHEMIS BIAPPENDICULATA (Selys).

Lyriothemis biappendiculata Ris, Cat. Coll. Selys, Libell. -

pp. 106-107 (figs. 71, 76, 77}.

62. NESOXENIA LINEATA (Selys).

Nesoxenia lineata Ris, Cat. Coll. Selys, Libell. pp, 126-128

(figs. 93, 94).

1 $. Matang Rd,

BORNEAN DRAGONFLIES. Bl

63. *LATHRECISTA ASIATICA (Fabr.).

Lathrecista asiatica asiatica Ris, Cat. Coll. Selys, Libell.

pp. 130-132 (figs. 95, 96, 97).

64, AGRIONOPTERA INSIGNIS (Ramb.).

Agrionoptera insignis insignis Ris, Cat. Coll. Selys, Libell-

pp. 137-138 (fig. 99). .

1g. Buntal, vi.1910. Thorax dark metallic green, lighter

markings almost lost. Abdomen with segments 3-8 scarlet-red,

each with apical, black ring; on 8 about one-sixth the length

of segment. Wings ‘suffused with golden yellow, especially at

apices; basal marks golden brown; on fore-wing se to aa’,

cu to Ac. Hinder-wing sc not reaching aw’,, cw to Ac.

Abd. 28 mm., h.w. 31 mm. Pt. 3mm.

65. AGRIONOPTERA SEXLINEATA Selys.

Agrionoptera sexlineata Ris, Cat. Coll. Selys, Libell. pp. 144—

145 (figs. 102, 103).

jel OS Baram, los se LO:

The female from Baram has the wings hyaline throughout,

and the red marking on the 7th abd. segment only.

66. CRATILLA METALLICA (Brauer).

Cratilla metallica Ris, Cat. Coll. Selys, Libell. pp. 152-153

(figs. 108, 109).

io. Min Murud, Sx.t4: 1S, 1 O. Matane Rds lit. 09)

67. CRATIDLA LINEATA (Brauer).

Cratilla lineata Ris, Cat. Coll. Selys, Libell. pp. 153-155

(Giese O 111):

68. PoTAMARCHA OBSCURA (Ramb.).

Potamarcha obscura Ris, Cat. Coll. Selys, Libell. pp. 156-157

(fig. 112).

69. ORTHETRUM GLAUCUM (Brauer).

Orthetrum glaucum Ris, Cat. Coll. Selys, Libell. pp. 233-234 ;

Laidlaw, Proc. Zool. Soc. London, 1915, p. 25.

70. ORTHETRUM PRUINOSUM CLELIA (Selys).

Orthetrum pruinosum ‘clelia Ris, Cat. Coll. Selys, Libel.

pp. 242-243.

Orthetrum clelia Laidlaw, Proc. Zool. Soc. London, 1915, p. 26.

71. ORTHETRUM SABINA (Drury).

Orthetrum sabina Ris, Cat. Coll, Selys, Libell. pp. 223-225

(figs. 133, 149).

322 MR. F. F. LAIDLAW ON

72. ORrHETRUM TESTACEUM (Burm.).

Orthetrum testaceum testacewm Ris, Cat. Coll. Selys, Libell.

pp- 234-236 ; Laidlaw, Proc. Zool. Soc. London, 1915, p. 26.

73. OrrHETRUM CHRySIS (Selys).

Orthetrum chrysis Ris, Cat. Coll. Selys, Libell. p. 237.

Group LV. (of Ris).

74, NANNoPHYA PyGmMaA Ramb.

Nannophya pygmea Ris, Cat. Coll. Selys, Libell. pp. 3847-348

(figs. 196, 197).

63646,2 2 9. Sarawak.

75. BRACHYGONIA OCULATA (Brauer).

Brachygonia oculata Ris, Cat. Coll. Selys, Libell. pp. 353-354

(figs. 203, 204).

3 6 3. 4th Mile, Rock Rd., Sarawak, 1909.

76. BRACHYGONIA OPHELIA Ris.

Brachygonia ophelia Ris, Cat. Coll. Selys, Libell. p. 304

(fig. 205).

1 Q. 4th Mile, Rock Rd., Sarawak, 1910.

77. TYRIOBAPTA TORRIDA Kirby.

Tyriobapta torrida Ris, Cat. Coll. Selys, Libell. pp. 355-356

(figs. 206, 208), and pp. 1120-1121 (fig. 643).

19. Sadong. 1 9. Matang Rd., vi. 09.

78. TYRIOBAPTA LAIDLAWI Ris.

Tyriobapta laidlawi Ris, Cat. Coll. Selys, Libell. pp. 1121-

1122 (fig. 644).

1 d. Sarawak (autotype). Coll. Ris.

79. TYRIOBAPTA KUKENTHALI (Karsch).

Tyriobapta kukenthali Ris, Cat. Coll. Selys, Libell. pp. 857-358

(fig. 209), and p. 1122.

3 6 6. Sarawak.

80. BRACHYDIPLAX CHALYBEA CHALYBEA Brauer.

Brachydiplax chalybea Ris, Cat. Coll. Selys, Libell. p. 363.

Brachydiplax chalybea chalybea id. op. cit. p. 1123.

1G. Santubang, 19.7 100 Sito. Baram (C7iose) sa ailice

Kotabelud, 17. viii. 13,

BORNEAN DRAGONFLIES. BoD

81. RApPHISMA INERMIS Ris.

Raphisma inernis Ris, Cat. Coll. Selys, Libell. pp. 370-371

(fig. 222).

35 6. Sarawak.

Group VI. (of Ris).

82. *DIPLACODES TRIVIALIS (Ramb.).

Diplacodes trivialis Ris, Cat. Coll. Selys, Libell. pp. 468-470

(figs. 293, 294).

83. *CROCOTHEMIS SERVILIA (Drury).

Orocothemis servilia Ris, Cat. Coll. Selys, Libell. pp. 539-542

(fig. 320).

84. *NEUROTHEMIS DISPARILIS Kirby.

Neurothemis disparilis Ris, Cat. Coll. Selys, Libell. p. 566.

85. NEUROTHEMIS FLUCTUANS Fabr.

Neurothemis fluctuans Ris, Cat. Coll. Selys, Libell. pp. 566-569.

1g. 4th Mile, Rock Rd., Sarawak. 1g,1@. Limbang,

2 Orville Orr NS Mierimiak. 13.s01 14, i Glee ibaram

(C. Hose).

86. NrUROTHEMIS TERMINATA Ris.

Neurothemis terminata Ris, Cat. Coll. Selys, Libell. pp. 569 -572

(figs. 328, 329, 334, 335).

266. Matang Rd. 16. Matang Mountain. 16. Bidi,

va.98. Ig. Buntal, 16.40.12; . l.d« Samarakamy 17v1. 10;

87. RuopoTHEMIS RUFA (Ramb.).

Rhodothemis rufa Ris, Cat. Coll. Selys, Libell. pp. 592, 593

(fig. 350).

1 9. Retuh, 16.v. 14.

Group VII. (of Ris).

88. PsEUDAGRIONOPTERA DIOTIMA Ris.

Pseudagrionoptera diotima Ris, Cat. Coll. Seiys, Libell. pp. 748-

749 (figs. 425, 426).

1. Sarawak.

89. TRItHEMIS AURORA (Burm.).

Trithemis aurora Ris, Cat. Coll. Selys, Libell. pp. 775-778

(fig. 442); Laidlaw, Proce. Zool. Soc. London, 1915, p. 26.

Doel eo. Kinabalu... 2.¢¢¢.. Ulu Akary Malas aS Mb:

Murud, 18. vii.14. 1 ¢. Samarakan, 17. vi. 10.

Proc. Zoou. Soc.—1920, No. XXIT. 22

324 MR. F. F. LAIDLAW ON

90. TRirHeMis Festiva (Ramb.).

Trithemis festiva Ris, Cat. Coll. Selys, Libell. pp. 796-799

(figs. 456, 457); Laidlaw, Proc. Zool. Soc. London, 1915, p. 26.

3 go. Kinabalu. 1 9. Mt. Merinjak.

91. ZyGonyx 1RIs Selys.

Zygonyx iris Ris, Cat. Selys, Libell. pp. 820-823 (fig. 478) ;

Laidlaw, Proc. Zool. Soc. London, 1915, p. 26.

264. Kinabalu. 1 9. Mt. Murud, 2. xi. 14.

92. ONYCHOTHEMIS CULMINICOLA CELEBENSIS Ris.

Onychothemis culminicola Forster, subsp. celebensis Ris, Cat.

Coli. Selys, Libell. pp. 835-836.

il @o Uaiam.

Under lip brown, upper lip and face brown, with faint metallic

blue reflexion; thorax brown, also with rather a metallic tinge.

Abdomen entirely scarlet; anal appendages—upper pair red at .

base, distal two-thirds black; lower appendage red, terminating

in a pair of fine black points. Wings lightly suffused with

yellow. Legs black. Abd. 32 mm.; h.w. 35 mm. Scarcely

fully mature.

93. *ONYCHOTHEMIS CULMINICOLA CULMINICOLA Forster.

Onychothemis culminicola culminicola Ris, Cat. Coll. Selys,

Libell. pp. 835, 836.

Group X. (of Ris).

94. HyDROBASILENS CROCEUS (Brauer).

Hydrobasilens croceus Ris, Cat. Coll. Selys, Libell. pp. 969-970

(fig. 562).

95, TRAMEA LiMBATA (Desjardins).

Tramea limbata Kirby, Cat. Odonata, p. 4.

Tramea translucida Kirby, loc. cit. p. 3.

Tramea limbata Ris, Cat. Coll. Selys, Libell. pp. 979-988

(figs. 563, 568, 569).

96. RHYOTHEMIS PHYLLI§ PHYLLIS (Sulzer).

Rhyothemis phyllis phyllis Ris, Cat. Coll. Selys, Libell. pp. 939—

940 (fig. 546, pl. v.).

97. *“RHYOTHEMIS ATERRIMA Selys.

Rhyothemis aterrima Ris, Cat. Coll. Selys, Libell. p. 953.

98. RuyvoruEmis pyemma (Brauer).

Rhyothenvis pygmea Ris, Cat. Coll. Selys, Libell. p. 955.

BORNEAN DRAGONFLIES. 325

99. RuyorneMis opsoLuscens Kirby.

LKhyothenvis obsolescens Ris, Cat. Coll. Selys, Libell. pp. 958-959

(ol. vine):

100. Rayornemis TRIANGULARIS Kirby.

Rhyothemis triangularis Ris, Cat. Coll. Selys, Libell. pp. 962-

963.

101. *ZyxomMMA PETIOLATUM Ramb.

Zyconumea petiolatum Kis, Cat. Coll. Selys, Libell. pp. 903-905

(lig. 523).

102. *THOLYMIS TILLARGA (Fabr.).

Tholymis tillarga Ris, Cat. Coll. Selys, Libell. pp. 913-915

(fics. 531, 532).

103. PANTALA FLAVESCENS (Fabr..).

Pantala flavescens Ris, Cat. Coll. Selys, Libell. pp. 917-920

(fig. 533).

104. *CAMAGCINIA HARTERTI Karsch.

Camacinia harterti Ris, Cat. Coll. Selys, Libell. pp. 928-929

(figs. 538, 539).

105. UROTHEMIS SIGNATA INSIGNATA (Selys).

_ Urothenis signata insignata Ris, Cat. Coll. Selys, Libell.

pp. 1024-1025.

106. *ArHRIAMANTA GRACILIS (Brauer).

Brachydiplax gracilis Kirby, Cat. Odonata, p. 22.

Brachydiplax melenops Kirby, loc. cit.

Athriamanta gracilis Ris, Cat. Coll. Selys, Libell. pp. 1032-

10338 (figs. 597, 598).

ZYGOPTERA .

CALOPTERYGINE.

107. NEUROBASIS CHINENSIS (Linn.).

Neurobasis chinensis Kirby, Cat. Odonata, p. 102; Forster,

Ann Soe. Entomol. Belg. xli. 1897, pp. 204-210; Kriiger, Stettin.

Entomol. Zeitg. 1898, pp. 74-75; Laidlaw, Proc. Zool. Soc. London,

1902, pp. 86-87.

Neurobasis chinensis chinensis Ris, Tijd. v. Entomol. Iviii.

LO oe paG.

22*

326 MR. F. F. LAIDLAW ON

108. MarrRoNoIpDES CYANEIPENNIS Forster.

Matronoides cysneip2nnis Forster, Wiener Entomol. Zeitg

1897, iii.; id., Ann. Soc. Entomol. Belg. xl. 1897. (sep.) pp. 1-5

Liillw, Jouva. Straits Branch R, Asiat. Soc. [63] 1912, p. 95

id., Proce. Zool. Soc. London, 1915, p. 30.

6 oo. Mb. Selinguid and Mt. Batu Lawi, 3500 ft., 30. v.11.

Var Sasi) Lao Met. Kinabalu, ix. 1913, up to 3000 ft.

109. VEsraLis AMaNA Selys.

Vestalis amena Kirby, Cat. Odonata, p. 103; Kriiger, Stettin.

Entomol. oe 1898, p. 75; Laidlaw, Proce. Tash Soe. iiidondon

1902, p. 87; id., Proc. Zool. Soe. London, 1915, pp. 30-31.

LIS So : ° . Mt. Kinabalu up to 3000 ft.

110. VesTALIs BERYLL& Laidlaw.

Vestalis berylle Laidlaw, Sarawak Mus, Journ. i. 6. 1915, p. 273.

1 g (autotype). Retuh. Specimen in Brit. Mus.

Wings hyaline, slightly tinged with yellow. A single row of

cells between Cu, and Cu,. Body metallic green. Legs black.

The species is remarkable for the great relative length of the

abdomen (70 mm.) as against 40 mm. for the hinder-wing.

EPALLAGINA.

111. PssuporH#A rRICOLOR (Selys).

Pseudopheu tricolor Kirby, Cat. Odonata, p. 109.

866. Retuh, Wyre is, 2 2 Oo. Saribas Sconce

Baram; C'. Hose).

112. PsrupoPH#A SUBCOSTALIS (Nelys).

Pseudophew subcostalis Kirby, Cat. Odonata, p. 109; Laidlaw,

Proc. Zool. Soc. London, 1915, pp. 32-33.

4g 6. Tampassuk, 19.viii. 13. 2 ¢ g. Saribas. 36 ¢. Mt.

Murud. 46 ¢. Ulu Akar.

113. PseupopH@aA suBNODALIS Laidlaw.

Pseudopheea subnodalis Laidlaw, Proc. Zool. Soe. Lado 1915,

p. dl.

13 6 ¢6,19% Mt. Kinabalu, ix. 13.

114. PseuDoPH#A BASALIS Laidlaw.

Pseudophea basalis Laidlaw, Proc. Zool. Soc. London, 1915,

p. 32.

4 $6. Kinabalu, 11-18. ix. 13.

BORNEAN DRAGONFLIES. 327

The males of these four Species are at first sight very Beal

alike. ‘They may be discriminated as follows :—

A. Opaque colouring of hinder-wing begins at nodus or one or

two cells before. Its margin runs transversely across the

wing; never a dark subcostal band extending to the wing-

base.

Abd. 35 + 1 mm., h. w. 27-5 mm. P. tricolor Selys.

B. Opaque colouring of hinder-wing begins about half-way

between wing-base and nodus. Its margin runs obliquely

outwards and backwards , and is rather irregular ; usually

a hyaline indentation about 4-6 cells deep between M,

and Cu,. Dark band in subcostal space of all four wings

extending to wing-base im adult specimens, reaching the

nodus in the fore- wing. Apex of fore-wing tipped with

dark brown from the middle of the pterostigma.

Abd. 30 + 1 mm., h. w. 25 mm. P. subcostalis Selys.

C. Opaque area of hinder-wing begins at about two-thirds of

the distance from the wing-base to the nodus, but area

between M,+, and Ris always hyaline wp to nodus. Dark

subcostal stripe not so well marked as in P. subcostalis,

and scarcely indicated on the fore-wing, which also has

much less opacity at its apex.

Abd. 35+1 mm., h.w. 28-30 mm. P. sabnodalis Laidlaw.

D. Hinder-wing opaque from the base to apex except for the

basal space, and for certain quite irregular areas which

In some specimens occur on the wings. Costal and sub-

costal spaces of fore-wing dark brown up to nodus,

Abd. 35+-1 mm., h. w. 27-28 mm. P. basalis Laidlaw.

Unfortunately I do not know the females.

These species probably all belone to the eroup P. variegata

i i y S S J

(Ramb.). The group occurs in all parts of the Sondaic area.

115. PsEUDOPH#A IMPAR INEQUIPAR (Selys).

Pseudophea inequipar Kirby, Cat. Odonata, p. 109.

43 <6. Tatau.

At most these are but a local race of the older species 7. enupar

(Selys) from Malacca and Sumatra.

Its nearest allies seem to be P. dispar (Ramb.) and P. frasert

Laidlaw from W. India, though the relationship is not very close.

116. DyspHma LUGENS Selys.

Dyspheea lugens Kirby, Cat. Odonata, p. 110.

26 6. Limbang River, iv.10. 1 9 (#). Retuh, 16. v.14.

The wings of the female specimen show in light hyaline brown

a colour-pattern identical with that of the male.

S$. Abd. 38mm.+1:°5mm.,h.w. 32mm. 9. Abd. 50 mm.,

h.w. 29 mm.

328 MR. F. F. LAIDLAW ON

117. DyspH#a LimpAta Selys.

Dysphea limbata Kirby, Cat. Odonata, p. 40.

3.6 ¢. Tatau. 1 3. Baram (C. Gose coll.; no date).

The male from Bavam belongs to the race semilimbata of Selys.

It is without the black costal line between the nodus and ptero-

stigma of the fore-wing. All the specimens have the hyaline

parts of the wings suffused with jel brown.

3. (Baram). Abd. 34 mm.+1-:5 mm., h.w. 30 mm.

6. Tatau. Abd. 35 mm.+1:5 mm., h.w. 30°5 mm.

LIBELLAGINA.

118. RHINOCYPHA BISERIATA Selys.

Rhinocypha biseriata Kirby, Cat. Odonata, p. 113,

Very closely related to 2. angusta Selys from Sumatra and

to R. perforata Perch. from Malacca and Lower Siam. The

group 1s Indo-Chinese and Sondaic in distribution.

119. RurnocypHa KARSCHI Kriiger.

Rhinocypha karschi Kruger, Stettin. Entomol. Zeitg. 1898,

pp. 83-85 ; Laidlaw, Proc. Zool. Soc. London, 1902, p. 90.

4 64. Limbang, ix. 09.

The species stands alone, without, so far as I know, near allies.

It is confined to the Sondaic area.

120. *RuiInocyePHA srvGtA Forster.

Rhinocypha stygia Forster, Ann. Soc. Entomol. Belg. xli. 1897,

pp. 210-211.

121, RetnocypHa moutront Laidlaw. (Text-fig. 1, a, b.)

Rhinocypha moultoni Laidlaw, Proc. Zool. Soc. London, 1915,

p- 39.

435,62 Q. Kinabalu, Sept. 1913.

The adult female of this species resembles that of 2. stygia

Forster very closely, to judge at least by Forster’s rather brief

description. But the fully adult male is so brightly coloured

about the body—much more so than the female—that I do not,

think it possible that stygia, which is entirely black about the

body, can be merely a very adult specimen of the same species.

The four males of mowléoni that I have been able to examine are

fully mature, and it is interesting to find that they retain on the

abdomen the colour-pattern characteristic of the teneral female,

which is lost in the mature female. For whereas the male

retains the paired dorsal spots of the abdomen from segment 2

to 9 as rich orange-red marks in addition to the yellow paired

lateral marks, thee dorsal marks are entirely lost in the fully

adult female, but are very conspicuous in newly-emerged females

as large ering -yellow areas covering about three- quariens of the

BORNEAN DRAGONFLIES. 329

dorsum of each segment from 2 to 8; fused at their bases with

the lateral system. (In the male the two series, dorsal and

lateral, are fused for their whole length from 2 to 6.) So that,

Text-figure 1, a, b.

Lateral view of colour-pattern of abdomen of Rhinocypha moultoni.

a, male; 6, immature female.

whilst not refusing to admit the possibility of &. stygia being

the extremely adult stage of R. mouwltoni, I do not think it at all

likely, and retain here the latter species as distinct.

122. RHINOCYPHA CUCULLATA Selys.

Rhinocypha cucullata Kirby, Cat. Odonata, p. 114.

6 6 go. Tatau. 5 ¢ ¢. Saribas.

An isolated species, confined apparently to Borneo.

123. *RuinocyPHA HUMERALIS Selys.

Rhinocypha humeralis Kirby, Cat. Odonata, p. 114.

124. *RHINOCYPHA EXIMIA Selys.

Rhinocypha eximia Kirby, Cat. Odonata, p. 114.

125. ReINOocYyPHA sp. A.

This species was identified for me by Martin. He has named

and described it in the forthcoming monegraph of the Caloptery-

gide in the Selysian collection, the appearance of which has been

very considerably delayed. As I do not wish to forestall him, I

note here only that the species appears to me to be related to

Forster’s R. aurulenteus from Buru, and perhaps more remotely

330 MR. F. F. LAIDLAW ON

to R. cucullata Selys. I have examined a series of specimens (d )

from Lio Matu and Tatau.

126. RurnocypHa sp. B. (Text-fig. 2.)

The remarks made on the previous species refer also to the

present one, which appears to be allied to 2. moultoni. 1 have

seen 5 specimens from Mt. Batu Lawi, all males.

Text-figure 2.

Lateral view of colour-pattern of abdomen of Rhinocypha sp. B.

[The primitive marking of the abdomen of the genus Lhino-

cypha, and perhaps of Micromerus, is probably a series of paired

dorsal marks, and outside these on either side a series of lateral

markings, the latter originally possibly a line extending along

the length of the segment, but now in many cases broken up into

anterior and posterior spots.

The species which in this paper I refer to as Rhinocypha sp. A

seems to preserve the primitive colouring to a fairly typical

extent; this is also the case in &. moultoni and in Rhinocypha

sp. B. Other species show various modifications of the colour-

pattern, which, it may be noted, may be either blue, orange, red,

ov yellow. Tilame, § maw Jay pucwilicia for example, in the males, only

the lateral colour-marks are retained, save on segments 8 to 9,

which have also dorsal markings. In 2. aurulentus Forster

from Buru only the lateral marks are retained; whilst in the

exquisite /?. iridea Selys from Burma only marks belonging to

the dorsal series are to be seen. Lastly, in some presumably

specialized forms—e. g., R. stygia Forster—these markings have

disappeared altogether. |

BORNEAN DRAGONFLIES. 33)

Diagnostic table for males of Bornean species of Rhinocypha.

A. All four wings with opaque areas.

a. Opaque area of hinder-wing with hyaline windows,

““mesothoracic triangle” present ........................ BR. biseriata Selys.

b. Opaque area of wings without hyaline windows.

1. Large antehumeral blue ray on thorax.

z. Terminal two-fifths of wings opaque ................ BR. humeralis Selys.

(3. Terminal fourth of wings opaque ..................... RB. ewimia Selys.

2. Antehumeral band of synthorax narrow.

y- Terminal opaque area of wings beginning at level

of prostigma in front; posterior tibiee not dilated ;

slight enlargement of posterior margin of second

abdominal segment ..........00...000eceecessereneses.. Rhinocypha sp. A.

6. Terminal two-fifths of wings opaque. Posterior

tibiz dilated, blue on anterior surface. Marked

enlargement of posterior margin of second abdo-

TIAN STNG 5c. cocuconenAbbesendabbbaRBoHbebHoonesencn | Jihe CoKeMaIG ISVS.

B. Hindez-wings only with opaque markings. Dorsal marks

Oi Boel, SSRN AAS) VAC, Jn ccoahaoasuubdepsnogesesesternenonsens Jee Uatenactelbo IOnnretee.

C. No opaque markings on wings.

i, ANlaloreavtnenyl seneera tomas OE &ionon aeodoasesdonseaospecdors sence: R. stygia Forster.

2. Dorsal markings on abdomen red, lateral markings

yellow.

a. Apex of clypeus marked with orange-yellow.

Lateral abdominal markings from segment 2-5

broken into an anterior line and posterior spot.

The anterior line is entirely fused with the dorsal

mark on these segments. From 6-8 the lateral

series is represented only by an anterior spot fused

1) Hn) Cloves | WHENY IS 4 Gor sndecuoocdcdbshebescesuapcdeeanacs Jeetrpoodtarod Enlennys

8. Apex of clypeus black.

Lateral abdominal markings present as a complete

band of rather irregular outline on segments 2-7,

only fused with the dorsal marks at its extreme

base, and not at allon segments 2-3, on which

segments the dorsal mark is small or absent ...... Rhinocypha sp. B.

127. RHINONEURA VILLOSIPES Laidlaw.

Rhinoneura villosipes Laidlaw, Proce. Zool. Soe. Londen, 1915,

pp. 33-35 (text-figs. 4, 5 a).

1 ¢. Mt. Kinabalu.

Autotype in British Museum.

128. MicROMERUS AURANTIACUS Selys.

Micromerus aurantiacus Kirby, Cat. Odonata, p. 115.

2 ¢ 6. Limbang.

129, MicROMERUS SEMIOPACUS Selys.

Micromerus semiopacus Kirby, Cat..Odonata, p. 115; Ris, Ann.

Soc. Entomol. Belg. lv. 1911, pp. 285-234 (fig. 1).

Noe luo Manu, G2 oro oUluy Akar.) sa. pIAe 29 ONO re ilcie

Matu.

aoe MR. F. F, LAIDLAW ON

130. Mtcromerus HYALINUS Selys.

Micromerus hyalinus Kirby, Cat. Odonata, p. 115.

96. Baram, x.1910. 2¢ 6. Tatau 3 dd ad, 30d

juv., 3 2 9. Saribas.

131. Mrcromervs sricricus Selys.

Micromerus sticticus Kirby, Cat. Odonata, p. 115.

3.66. Tatau.

TIncerte sedis.

132. DEYADATTA ARGYROIDES (Selys).

Devadatta argyroides Kirby, Cat. Odonata, p. 111.

This genus seems to me to be not very remote in venation

from the ‘* Legion” Megapodagrion.

Legion MEcAPpoDAGRION.

133, *PoDOLESTES ORIENTALIS Selys.

Podolestes orientalis Kirby, Cat. Odonata, p. 126; Kvriger,

Stettin. Entomol. Zeitg. 1898, pp. 98, 99.

134. PopoLEstEs CHRYSOPUS Selys.

Podolestes chrysopus Wirby, Cat. Odonata, p. 126.

1g¢,19 incop. Matang Rd., 28. x.09. 2¢ $. Baram, vii. 09.

26 46. 4th Mile, Rock Rd., Sarawak, 24.vi.09. 1 9. Matang

Rd., 24. vi. 09.

135. RHINAGRION BORNEENSE (Selys).

Amphilestes borneensis Kirby, Cat. Odonata, p. 126; Karsch,

Entomol. Nachr. xvii. (1891), no. 16, p. 2: Kriger, Stettin.

Entomol. Zeitg. 1898, pp. 137-138.

10 6 ¢,2 2 @. Lio Matu, 4.x1.14. 192. Kuching, 5. x1. 09.

3. Agrees fairly closely with the description of the type. The

yellow mark on the dorsum of the thorax is triangular rather

than oval, its base running quite transversely across the thorax,

and its inner side close to the mid-dorsal carina. In one specimen,

however, the anterior margin of the triangle is distinctly oblique,

so that the mark becomes more nearly oval. The colouring of

the ventral side of the thorax seems to vary with age; it is black

in the fully adult specimens, but on the younger it is yellow with

a smoky tinge.

Abdomen: segments 1--7 reddish brown, 1 with yellowish mark

on dorsum, 2-7 with black apical ring, and 2 with pale sub-

apical mark as well; 8,9 white, enamelled; 8 with a reddish

tinge of rather blue-white; 10 black.

Abd. 30 mm. + 1 mm., h. w. 21 mm.

2. Head black, but with yellow median mark on post-clypeus

and yellow margins around the eyes. Prothorax hght golden

BORNEAN DRAGONFLIES. BOO

brown above, paler below, with the posterior lobe whitish green.

Thorax golden brown above, passing to grey-white below. Tri-

angular marks on the dorsum as in the male.

Abdomen: segment 1 greenish white above, pale below : seg-

ments 2-7 reddish brown, progressively darker backwards, with

subterminal grey-green rings; segments 8-10 black, 8 with

lateral longitudinal whitish band.

Abd. 27 mm., h. w. 25 mm.

(For generic name, see Calvert, Proc. Acad. Nat. Sa. Phila-

‘delphia, 1913, p. 258.)

136. *RHINAGRION ELOPUR# (Selys).

Amphilestes elopure Kirby, Cat. Odonata, p. 126.

Legion PLATYCNEMIS.

137. Ca@Liccta ocroGESIMA (Selys).

Celiccia octogesima Kirby, Cat. Odonata, p. 128.

192. Matang Rd., 3.1. 10 (damaged).

138. Ca@sLiccIA MEMBRANIPES (Ramb.) (race NEMORICOLA

Laidlaw ?).

Celiccia nemoricola Laidlaw, Journ. Straits Branch R. Asiat.

Soc. [63] p. 95; id., Proc. Zool. Soc. London, 1915, p. 37.

Celiccia membranipes vace nemoricola Laidlaw, Proc. Zool. Soc.

London, 1917, pp. 230-231.

Common on Mt. Kinabalu.

139. CaLiccia FLAVosTRIAtA Laidlaw.

Celiccia flavostriata Laidlaw, Proc. Zool. Soc, London, 1917,

pp. 223-224 (figs. 1, 2).

26d. Mt. Merinjak, 21.v.14. 16d. Mt. Matang, 4. x11. 13.

Autotype in British Museum. Paratype in Coll. Mus. Sarawak

and in my own collection.

140. Ca@niccia cAmproni Laidlaw.

Celiccia campiont Laidlaw, Proc. Zool. Soc. London, 1917,

pp. 224-225 (figs. 3, 4).

iio LivoMiatiny olsen. 14)

141. Ca@iccia macrostiema Laidlaw.

Celiccia macrostigma Laidlaw, Proc. Zool. Soc. London, 1917,

pp. 225 227 (figs. 5, 6).

1d. Baram, 20.x.10 (1 9%. 19. x. 10).

142. CanicciA NIGROHAMATA Laidlaw.

Celiccia nigrohamata Laidlaw, Proc. Zool. Soc. London, 1917,

p. 228 (figs. 7, 8).

334 MR. F, F. LAIDLAW ON

[| Ca@sLICCIA BORNEENSIS (Selys). |

Celiccia borneensis Kirby, Cat. Odonata, p, 128.

(See under C. octogesima (Selys), Laidlaw, Proc. Zool. Soc.

London, 1917, p. 231.)

143, Coprra AvTomaRIA (Selys). (Text-fig. 3.)

Copera atomaria Kirby, Cat. Odonata, p. 129.

(See also Kriiger, Stettin. Entomol. Zeitg. 1898, pp. 103-107 ;

Ris, Tijdschr. v. Entomol. Iviii. 1915, p. 7, sep.)

Upper anal appendages of male about one-half length of lower

pair. Tibiee not dilated. In the adult male the thorax is bronze-

black above, with a few irregular yellow spots representing the

antehumeral band. The sides of the thorax are mottled black

and yellow, the legs are entirely red-brown. Abdomen black, with

yellow lateral markings on segments 1 and 2 anda basal bluish-

white ring on segments 3-6. Segment 10 and upper pair of anal

appendages white, lower pair of appendages black; distal half

marked with white dorsally.

Text-figure 3.

Prothorax of Copera atomaria seen from above.

Adult female coloured much as the male, but duller. The

posterior femora have a row of black dots along their dorsal

surface. The posterior thoracie margin of the female carries a

pair of small triangular projections directed upwards and for-

wards. These are much less acute than the corresponding

structures of the Sumatran species, C. acutimargo Kriiger.

6. bd) 28)4-10:75 mm hiw.el 7-aemme » C1. Abadia Ogmuny,

h. w. 19 mm.

Specimens from Lio Matu seem to have been taken in com-

pany with Caconewra verticalis Selys; at least in many cases I

found examples of both species papered together.

To what extent these Bornean specimens are to be regarded as

distinct from other allied forms of the genus it is at present

impossible to say. But I think it likely that a number of forms

of the vittata series of the genus will ultimately be characterized.

As I have seen only examples of one form from Borneo, I have

retained for it the Selysian name,

BORNEAN DRAGONFLIES. 339

Legion AGRION.

144. ONYCHARGIA ATROCYANA Selys.

Onychargia atrocyana Kirby, Cat. Odonata, p. 1395; Kruger,

Stettin. Entomol. Zeitg. 1898, p. 118; Ris, Nova Guinea, xiii.

Zool. 2, pp. 94-95 (fig. 13).

1d. Mt. Murnd, 18. xii.14. Very adult, lacking the iast four

abdominal segments.

145. CERIAGRION CERINORUBELLUM (Brauer).

Ceriagrion cerinorubelluny Kirby, Cat. Odonata, p. 154; Kriger,

Stettin. Entomol. Zeitg..1898, pp. 119-120; Ris, Abh. d.

Senckenberg Naturfors. Gesellsch. xxxiv. p. 019; id., Tijdschr.

vy. Entomol, lviii. 1915, p. 13 (sep.): Laidlaw, Rec. Indian Mus.

Xvi. part 11. p. 188.

26 6. 4th Mile, Rock Rd., Sarawak, 30.x1.09. 3¢ ¢. Retuh,

16.v. 14.

146. CeRIAGRION BELLONA Laidlaw.

Ceriagrion bellona Laidlaw, Sarawak Mus. Journal, 1. 1916,

p- 274.

236 6. Mt. Matang, xii.13. 1 ¢. Kinabalu, ix. 13.

Ab commences before the level of Ac. Wings in adult lightly

tinged with yellowish brown. Excision on hind-margin of tenth

abdominal segment A-shaped. Lower anal appendages (of male)

about twice as long as upper pair. Head orange-brown above,

greenish white below. Prothorax and synthorax coppery brown

above, yellowish green below. Abdomen: first segment coppery

brown, second to sixth carmine, seventh to tenth dull brown.

Abd. 28°5 mm., h. w. 20 mm.

147. PsEUDAGRION MICROCEPHALUM (Ramb.).

Pseudagrion microcephalum Kirby, Cat. Odonata, p. 153; Ris,

Supplement Entomol. no. 5, 1916, pp. 40-43 (figs. 13-19);

Laidlaw, Rec. Ind. Mus. xii. pp. 23-24 (fig. 1); Ris, Ann. Soc.

Entomol. Belg. 1911, p. 235.

1 2 (damaged). 4th Mile, Rock Rd., Sarawak.

Dr. Ris gives a full synonymy of this species, and a deseription

of several races characterized by the shape of the anal appendage

of the male (Ris, Suppl. Entomol. oc. cié.).

The Bornean species seem to be below the average in size.

148. PsEUDAGRION PRUINOSUM (Burm.).

Pseudagrion pruinosum Kirby, Cat. Odonata, p. 153; Kriiger,

Stettin. Entomol. Zeitg. 1898, p. 119; Laidlaw, Sarawak Mus.

Journal, ii. 1916, p. 275; Ris, Nova Guinea, xiii. Zool. 2,

pp. 97-98 (fig. 18).

356 MR. F. FB. LAIDLAW ON

149. ACIAGRION BORNEENSE Ris.

Aciagrion borneense Ris, Ann. Soe. Entomol. Belg. 1911,

pp. 234-235 (figs. 2-3).

1 2. Tabekang, 12. v.14.

Occurs also in the Malay Peninsula.

150. AGRIOCNEMIS FEMINA (Brauer).

Agriocnemis femina Ris, Supplement Entomol. no. 5, 1916,

pp. 22-26 (figs. 13, 17).

(For full synonymy of this species, see Ris, loc. cit.)

151. ARGIOCNEMIS RUBESCENS Nelys.

Argiocnemis rubescens Kirby, Cat. Odonata, p. 158; Ris, Abh.

d. Senckenberg Naturfors. Gesellsch. xxxiv. pp. 516-518.

(For this species also, consult Ris’s paper quoted here.)

to. Malimau, “Sox FO. ee, Mit. “Mermiyak. tay vei:

1 3g. 4th Mile, Rock Rd., Sarawak.

152. AMPHICNEMIS WALLACEI Selys.

Amphicnemis wallacei Kirby, Cat. Odonata, p. 157; Ris, Ann.

Soc. Entomol. Belg. lv. 1911, pp. 2386-237 (figs. 4, 5); Laidlaw,

Proc. Zool. Soc. London, 1913, p. 70.

1g. Baram.

153. AmpuicnEemis Louis# Laidlaw.

Amphicnemis louise Laidlaw, Proc. Zool. Soe. London, iSlhey,

p. 71, pl. iv. figs. 5, 5a; Ris, Tijdschr. v. Entomol. Ivii. 1915,

pp. 13-14.

154, AMPHICNEMIS REMIGER Laidlaw.

Amphicnemis remiger Laidlaw, Journal Straits Branch R.

Asiat. Soc. [63] 1912, pp. 96--97, pl. fig. 4; id., Proc. Zool. Soe.

London, 1913, p. 72.

1 3. Batu Lawi. 2 3 S (imperfect). Murud, 20. xii. 14.

155. AMPHICNEMIS MADELEN% Laidlaw.

Amphicnemis madelene Laidlaw, Proc. Zool. Soc. London,

1913, pp. 71-72 (pl. iv. figs. 6, 6@).

266. Kuching.

156. AMPHICNEMIS MARTINI Ris.

Amphicnemis martin Ris, Ann. Soc. Entomol. Belg. lv. 1911,

pp. 237-238 (fig. 6); Laidlaw, Proc. Zool. Soe. London, 1913,

pp. 72-73.

13. Limbang. 1. Mt. Merinjak, 22. xi. 14 (both imperfect).

And, in addition, I have before me 6 2 2 from Murud, 20.x1.14,

BORNEAN DRAGONFLIES. 337

and 1 @ from Limbang which I have not identified with cer-

tainty; also a damaged female specimen from atau which is

possibly identical with A. grazilis Kriiger from Sumatra; and,

lastly, 1 @ from Baram belonging to a different, and undeter-

mined, species.

I hope to pay more attention to this interesting genus when

opportunity offers.

157. PERICNEMIS STICTICA Selys.

Pericnemis stictica Kirby, Cat. Odonata, p. 158; Kruger,

Stettin. Entomol. Zeitg. 1898, p. 158; Laidlaw, Proc. Zool. Soc.

London, 1902, p. 386.

1 g. Limbang (in fragments).

158. Tetnopasis KirByt Laidlaw (?). (Text-fig. 4.)

Teinobasis kirbyi Laidlaw, Proc. Zool. Soc. London, 1902,

pp. 386-387.

Teinobasis superba Laidlaw (nec Selys), Proce. Zool. Soe.

London, 1917, p. 231.

1g. Lio Matu, 4.xi.14. (Specimen to be sent to British

Museum.)

Text-figure 4.

Lateral view of anal appendages of Teinobasis kirbyi, (?).

M, and Ms united from their origin for the length of one cell.

Upper branch of superior anal appendage short, about one-half

the length of lower branch, which is nearly equal in length to the

lower appendage. Ac hes much nearer the level of Ax, than of

Ax..

Head: upper surface entirely black; the upper hp polished,

but without metallic reflex ; lower lip grey-white; the rest of the

under and posterior surfaces greenish white.

Prothorax: black above, grey-white below.

Synthorax : black above, no trace of antehumeral stripes; dark

grey on the sides, a little pulverulent; paler below.

Abdomen: slender, but segments 7-10 distinctly stouter

338 MR. F. F. LAIDLAW ON

than 3-6. Bronze-black above, brownish white below. Anal

appendages (see text-fig. 4) brown tipped with black.

Legs: gvey-white, with black articulations and black lines on

the posterior surfaces.

Wings: vather smoky; pterostigma almost square, but its Inner

margin more oblique than its outer; dark brown, very finely

edged with lighter colour surrounded by a thickened black vein.

Px 15.

Abd. 40+°5 mm., h. w. 23 mm.

Segments 3-6 of the abdomen measure 29:5 mm.

Compavison of this specimen with Ris’s figures and examina-

tion with his diagnostic table (Ris, Nova Guinea, xiil. Zool. 2,

pp. 100-102, fig. 22) has enabled me to determine that this

species is not 7’. superba Selys, from which species it differs in

its smaller size, absence of antehumeral stripes, and shape of anal

appendages. My species, 7. kirbyi, is unfortunately very im-

perfectly known, but on the whoie the present specimen would

seem to resembie it closely in colouring and venation, and it may,

I think, be regarded as conspecific with it.

159. Ternopasis RAJAH Laidlaw.

Teinobasis rajah Laidlaw, Journ. Straits Branch R. Asiat. Soe.

[63] 1912, p. 97.

25 S. Limbang, 22. vi. 11.

Thorax orange-red above, with median bronze-green stripe.

Abd. 33 mm., h. w. 20 mm.

Autotype ¢ in British Museum. Paratype d, coll. Ris.

160. SrENaGRION DUBIUM (Laidlaw).

Pseudagrion (¢) dubium Laidlaw, Journ. Straits Branch R.

Asiat. Soc. [63 | 1912, pp. 97-98, pl., fig. 5.

Stenagrion dubiwm id., Proc. Zool. Soc. London, 1915, p. 39;

id. ops cub. NON. p. 251) (igaw ho):

Legion PROrONEURA.

[I propose the following arrangement of the Oriental species

grouped under the Selysian genera Disparoneura and Caconeura

(Allonewra Selys). It is, I think, natural, and has the advantage

of retaining many of the species in the original genera :—

A. Ac lies midway between aay and aa.

a. Posterior lobe of prothorax of female armed with

hook-like projections.

a, Ab meeting nerve descending from quadrilateral.

i. Cu, reaching hinder margin of wings beyond

half the wing-length ............................ Chloroneura Waidlaw.

ii. Cu, reaching hinder margin of wings before

Jail (eaves SyysmOVSR NSO VERE on yo5 oy geo we weaaseyuse caben ans

B. Vestige of Ab not meeting nerve descending

from quadrilateral, or absent altogether ......... Caconeura Kirby.

}. Posterior lobe of prothorax of female simple ......... Indoneura Laidlaw.

}3. Ac lies nearly at level of ax, or proximal to it ............ Risionewra Munz.

Disparoneura Selys.

BORNEAN DRAGONFLIES. 339

Of these genera: Chloronewra appears to be confined to penin-

sular India; Disparonewra has a westerly range, extending from

Africa to India and Burma and reaching its limit eastwards in

Borneo ; Caconeura, more specialized in some respects, extends

from Burma to Malaya, with one species in Ceylon ; whilst the

still more specialized Lisionewra is Papuan (see Laidlaw, Rec.

Ind. Mus. xiii. 1917, pp. 343-344), ]

161. CACONEURA DORSALIS (Selys).

Caconeura dorsalis Kirby, Cat. Odonata, p. 134.

Disparoneura dorsalis Laidlaw, Proc. Zool. Soe. London, 1913,

p- 75.

occ) bara O. ose). 1 6. Murud, VSaxaie las

162. CACONEURA VERTICALIS (Selys).

Disparoneura verticalis Kirby, Cat. Odonata, p. 154; Kriger,

Stettin. Kntomol. Zeitg. 1898, p. 114; Forster, Fasc. Malay.,

Zool. iv. Odonata, Part II., (sep.) p. 14; Laidlaw, Proc. Zool.

Soc. London, 1913, p. 75.

Ali Gao 2 @. Ino Matu, x1.14. 26 6, 12. Retuh, 14 vol4:

The presence of a vestige of Cu, is constant in all these

specimens.

163. CACONEURA HYPERYTHRA (Selys).

Caconeura hyperythra Kirby, Cat. Odonata, p. 134.

Disparoneura hyperythra Laidlaw, Proc. Zool. Soc. London,

ORS ps 76.

164. Cacon=uRA HOSEI (Laidlaw),

Disparoneura hosei Laidlaw, Proc. Zool. Soc. London, 1918,

pp. 76-78.

13. Baram (C. Hose), autotype, coll. F.¥.L. 56 6. Lio

Matu, 4.xi.14. (1 ¢. Coll. Brit. Mus.)

165. CAcoNEURA PERAMG@NA (Laidlaw).

Disparoneura peramenag Laidlaw, Proc. Zool. Soc. London,

1913, pp, 76-77 (pl. iv. figs. 8, 8 a).

166. *CACONEURA TNTERRUPTA (Selys).

Disparoneura interrupta Kirby, Cat. Odonata, p. 134,

167. *CacoNEURA COLLARIS (Selys).

Disparoneura collaris Kirby, Cat. Odonata, p. 134.

Disparoneura notostigma collaris Forster, Fasc. Malay., Zool.

iv. Odonata, Part IL., (sep.) p. 13.

*[CACONEURA LANSBERGI (Selys). |

Caconeura lansbergi Kirby, Cat. Odonata, p. 134.

Location somewhat doubtful.

Proc. Zoou. Soc.—1920, No. XXIII. 23

340 MR. F. F, LAIDLAW ON

168. *CacoNEURA GRACILLIMA (Selys).

Caconeura gracillima Kirby, Cat. Odonata, p. 134.

169. CaconEuURA MouLtont (Laidlaw).

Disparoneura moultont Laidlaw, Journ. Straits Branch R. Asiat.

Soc. [63] 1912, pp. 98-99.

lS. Batu Wawa, ev.

Upper lip pale yellow, with fine black margin, otherwise velvety

black above. Dorsum of prothorax and synthorax black, a fine

yellow lateral stvipe on the latter. Abdomen brownish black.

No vestige of Cu, ; Cu, reaches first cross-nerve after the quad-

rangle in the fore-wing, the second in the hinder-wing ;

14 postnodal costals in fore-wing.

Abd. 34 mm., h.w. 19 mm.

170. *DIsPARONEURA AURANTIACA Selys.

Disparoneura aurantiaca Kirby, Cat. Odonata, p. 154.

171. DispARoNEURA ANALIS (Selys).

Disparoneura analis Kirby, Cat. Odonata, p. 1384; Kriiger,

Stettin. Entomol. Zeitg. 1898, p. 111; Laidlaw, Sarawak, Mus.

Journ. ii. no. 6, 1915, p. 275.

1g. Baram, 29.11.10. 16. Saribas, 09. 16. Murud,

Vix es Tino Matus eexapl AS

Legion PLatysricra.

172. DREPANOSTICTA RUFOSTIGMA (Selys).

Platysticta rufostigma Kirby, Cat. Odonata, p. 133; Laidlaw.

Proc. Zool. Soc. London, 1913, p. 79 (pl. iv. fig. 9).

(For definition of the genus Drepanosticta, see Laidlaw, Records

Ind. Mus. xiii. part vi. p. 339 (1917).)

173. Prorostricra vERsicoLor Laidlaw.

Protosticta versicolor Laidlaw, Proc. Zool. Soc. London, 1913,

pp. 78-79.

174. Prorosricra KINABALUENSIS Laidlaw.

Protosticta kinabaluensis Laidlaw, Proc. Zool. Soc. London,

1915, pp. 37-38 (fig. 5, B).

Also 1 (imperfect), Mt. Merinjak, 26.v.14, belonging to

Drepanosticta, with the hinder margin of the thorax bilobed.

Apparently an undescribed species.

Legion LEstEs.

175. *Lusims WALLAcEr Kirby.

Lestes wallace Kirby, Cat. Odonata, p. 162.

BORNEAN DRAGONFLIES. 341

Lestes sp.

1S. Matang Rd., 21.viii.20 (lacking head and end of

‘abdomen).

Wings hyaline, but with smoky tinge, most marked at apex ;

petiolated up to level of Ac, which lies nearer Ax, than Ax,

M, commencing 83-9 cells distal to nodus m fore- -wing, 63-7

cells in hinder- wing. None of the sectors angulose except Cur

Pterostigma, large, ‘dilated, about four times as long as broad.

One supplementary sector and vestige of a second interposed

between M, and M,. Quadrangle broad, lower side twice as long

as upper side; outer angle acute.

This interesting species shows distinct affinities to the Indo-

Chinese Orolestes..

There are also amongst Major Moulton’s material some three

fragmentary specimens, all Agrionines, which are too imperfect

to describe. They appear to be unnamed species. One is a

Pseudagrion ; a second is either a Psewdagrion or belongs to an

allied genus; the third I Smee refer with certainty to any

genus, though again it may be related to Pseuwdagrion.

I have omitted from the list certain common and widely-spread

species which almost certainly occur in Borneo, but which have

never yet been recorded from the island. These would add some

ten species or so to the list.

Allowing for these broken or unrecorded species, we may, I

think, assume that the list includes from between 70 to 80 per

cent. of the total Dragonfly fauna of the island.

III. Some Remarks on Geographical Distribution.

Dragonflies are so important in this respect that a few notes

are not out of place.

It is difficult to subdivide the Oriental Region in any entirely

satisfactory way; but one may contrast the equatorial Dragonfly

fauna, ranging from about the equator to roughly 10° N,, with

the tropical fauna lying mainly between 10° N. and 25° N. The

equatorial fauna may be called the Malayan, the tropical (unless,

indeed, that can be shown to consist of more than one fauna of

co-ordinate importance) I label here the Indo-Chinese.

The Malayan fauna occupies as its main areas :—

The Sondaic area—i.e., the Malay Peninsula, Borneo,

Sumatra, and Java.

The Philippine Islands.

The Celebes.

Any detailed analysis of the fauna of Malaya would extend

this paper to undue length; hence I give only a very general

table, and prefix “so far as £ know” to the whole.

ate

B42 ON BORNEAN DRAGONFLIES.

TaBin of distribution of genera found in Borneo.

J. Genera confined to Borneo.

Lineschna, Pseudagrionoptera, Matronoides, Rhinoneura, Stenagrion.

II. Genera specially characteristic of Malaya or confined there-

to (several of these occur in Indo-China).

Tetracanthagyna, Amphieschna, Orchithemis, Pornothemis, Brachy-

gonia, Tyriobapta, Micromerus*, Dysphea, Devadatta, Podolestes,

Rhinagrion, Caconeura, Amphicnemis, Pericnemis.

III. Genera of Oriental distribution, with species confined to

Sondiac area.

Orogomphus, Onychothemis, Macromidia, Azuma, Idionyx, Jagoria,

Gomphidia, Leptogomphus, Macrogomphus, Microgomphus, Burma-

gomphus, Heterogomphus, Vestalis, Pseudophea, Rhinocypha, Cocliccia,

Copera, Disparoneura.

IV. Genera common to Oriental and Australian Regions, with

species confined to Sondaic area.

Agrionoptera, Neurothemis, Brachydiplax, Lyriothemis, Camacinia,

Aciagrion, Drepanosticta, Protosticta.

V. Genera mainly Papuan or Australian, with species confined

to Sondaic area.

Metaphya, Oda, Raphisma, Teinobasis.

VI. Paleotropical (a) or Holotropical (>) genera, with species

confined to Sondiac area.

(a) Ictinus, Tetrathemis, Rhyothemis, Aithriamanta, Pseudagrion,

Ceriagrion.

(b) Gynacantha.

VII. Special cases :—

Hyleothemis. Borneo, 8. India, Ceylon.

Helieschna. W. Africa, Malaya, Burma.

Macromia. Paleotropical, Palearctic, Nearctic.

Some negative characters showing contrast with Indo-Chinese

enon ees —

Absence of Cordulegastrine, relatively poor development of

genera of Calopterygine and of Brachytryon series

of Aischnine.

Relative scarcity of species of the following genera :—

Onychogomphus, Lestes, Ischnura, Agriocnemis.

Absence of Davidius, Anisogomphus, Cyclogomphus, Gomphus,

Pseudothemis, Sympetrum, Palpopleura sexmaculata,

Bradinopyga, Anisopleura.

Genera not mentioned in the above table have no species

peculiar to the Sondiac area.

* Micromerus has 13 species in Malaya; one, JZ. lineatus, widely spread in Inéo-

Chinese province ; and one, JZ. finalis, m Ceylon,

NEW SOUTH AFRICAN THEROCEPHALIAN REPTILES. 343

On some new Therocephalian Reptiles from the Karroo

Beds of South Africa. By R. Broom, M.D., D.Sc.,

Deliesa obi sess

[Received April 13, 1920: Read April 27, 1920.]

(Text-figures 1-9.)

INDEX. Page

Hevidoswuchus longeceps, Sp NOV. 2.0is..ec-cscienreos-sehaeese | O4O

PA OPECOPSUS MUALUS, SEN. eb SP. NOV. scvsecceses-aesenseseen: 347

Moschorhinus kitchingi, gen.'et sp.nov. .................. 851

Whaitsia platyceps Waughton .............00.ccekeeees. 854

Though more species and genera of Therocephalians are known

than of either of the other two large suborders of the Carnivorous

Therapsida—the Gorgonopsia and the Cynodontia,—the structure

of the skull and skeleton is much better known in these later

suborders. This is largely due to the fact that the Thero-

cephalians are mainly found in the Lower Beaufort zones, where,

owing to the petrological conditions, very fine complete spe-

cimens are rare. ‘The Gorgonopsiaus which are found mostly

in the Cistecephalus zone, and the Cynodonts which mainly occur

in the Upper Triassic zones, are usually represented by well-

preserved skulls with a matrix which can be fairly easily removed.

Recently I have been fortunate in discovering one or two very

interesting specimens of late Therocephalians formed in the Upper

Endothiodon and Cestecephalus zones, which add considerably to

our knowledge of the Therocephahan skull.

IcrimosucHus LONGICEPS, Sp. NOV.

In 1900 I discovered near Pearston the remains of a very

interesting type of ‘Therocephalian which I described under the

name Teneiaena we primevus. Until recently no other’ specimen

of either this genus or species has ever been discovered, and as

the type skull is very imperfect, a good specimen of this genus has

long been one of our principal desiderata. In December 1917,

I found at Bruintjeshooyte, between Somerset Hast and Pearston,

a good skull, which may be referred to the genus Ietidosuchus,

though a, distinct species from /. primevus.

The skuli is nearly complete, lacking only the incisor portion,

the oecipital condyle, the arches of the left side and the quadrate

region of the vight. The palate is much weathered on the left

side, but nearly perfect on the right.

As preserved, the skull measures in greatest length 135 mm.,

and when complete was probably 152 mm. Jong. The greatest

breadth is about 84mm. From the front of the orbit to the

base of the canine is 52 mm., and from the front of the orbit to

344 DR. R. BROOM ON SOME NEW

the front of the snout was probably about 75 mm. The inter-

orbital measurement is 22 mm., and the antero-posterior measure-

ment of the orbit is 82 mm.

The premaxille and septomaxille are lost. The nasals are long

and narrow, only slightly wider in front and behind than in

the middle region. ‘Vhe maxilla is unusually long in comparison

with its depth. It passes forwards a considerable distance 1m

advance of the canine and backwards much beyond the last molar.

In front of the main canine there are two small canines as

in Scaloposaurus, but the present type, besides differmg very

greatly in many other ways, differs in having the third tooth

developed into a powerful canine. The canine here measures

about 15 mm. in height and 5 mm. in antero-posterior length. It

is usually curved, and is remarkable for having no serrations

along its posterior border. ‘The three canines together measure

95 mm, Behind the main canine is a diastema of 6°5 mm.

followed by a series of 9 small cone-like molars, the whole molar

series measuring 25mm. In Jetidosuchus primevus there are ap-

parently only 8 molars, but as both the upper and lower Jaws are

Text-figure 1.*

Side view ot skull of Ictidosuchus longiceps Broom.

badly preserved, it is not improbable that there may also have

been 9 in the type species. ‘The molars of /ctidosuchus primevus

are considerably larger and closer together, but appear to be

otherwise similar to those of /. longiceps.

The prefrontal is a relatively large bone which forms the

anterior half of the upper orbital margin and extends down to

the middle of the anterior border. The lacrimal is considerably

less than half the size of the prefrontal, as will be seen in the

figure given.

The jugal is a very long slender bone forming most of the sub-

orbital bar and about half of the outer temporal arch. A strong

but short ascending process forms the lower half of the post-

orbital arch, articulating with the short postorbital bar of the

postorbital bone.

The frontals are relatively small, and only form a very small

part of the supraorbital margin. The hones are about the same

length as the orbit, and articulate in front with the nasals by

* For explanation of lettering, see p. 355.

SOUTH AFRICAN THEROCEPHALIAN REPTILES. 345

an irregular transverse suture, and posteriorly with the parietals

by a transverse serrated suture.

There is no postfrontal.

The postorbital is relatively small, forming the posterior half

of the upper orbital margin and the upper half of the postorbital

arch. It extends backwards by the side of the parietal to the

plane of the posterior border of the pineal foramen.

The parietals are relatively long and slender. In front they

are flattened out, and form a transverse suture with the frontals.

Posteriorly they form a sharp intertemporal crest, and at the

front of the crest is a large oval pineal foramen. From the

Text-figure 2.

ANYWAY

Upper surface of skull of Ictidosuchus longiceps Broom.

posterior part of the crest the parietals pass outwards, forming a

considerable part of the occipital crest.

The occiput is very imperfectly preserved. It is shallow and

broad, and from the upper margin of the foramen magnum the

bone passes upwards and forwards. The elements cannot be

clearly made out, though they are doubtless as in better-known

Therocephalians. The squamosals and quadrates are lost.

As already stated, the palate has not been fully cleared in front

as the matrix is required to support the teeth, but the structure

is sufficiently revealed in all the posterior part. A transverse

fracture through about the plane of the third last molar shows in

section the prevomers. Though very closely placed to each other

346 DR. R. BROOM ON SOME NEW

they are not anchylosed. Hach has an ascending thin plate and

a short descending plate, and from each bone a transverse plate

passes outwards which approaches and doubtless further back

meets the palatine. ‘The palatines as seen here in section are

slender curved bones which below meet the maxillaries, and on

passing inwards and then upwards again approach and possibly

meet the upper borders of the maxillaries.

The prevomers extend backwards a considerable distance

behind the plane of the Jast molars and meet the anterior ends of

the pterygoids. The palatines form the greater part of the bony

roof of the mouth, passing about as far back as the posterior end

Text-figure 3.

err m--

Palatal surface of skull of Ictidosuchus longiceps Broom.

of the maxilla. There is behind the palatine a large suborbital

vacuity as in Seylacosaurus, and, as in that genus, this vacuity is

bounded externally by the ectopterygoid. ‘The ectopterygoid is

moderately slender, but in front of the pterygoid process which

hes along the inner side of the mandibles it has a well-developed

descending process, which adds much support to the pterygoid

process.

The pterygoid is fairly similar to that already described and

figured by me in Seylacosawrus, and possibly some of the apparent

differences may be due to certain features being preserved in the

present specimen which were lost in the other. As in most

primitive reptiles, there is an anterior process which, passing

SOUTH AFRICAN THEROCEPHALIAN REPTILES. 347

inside of the palatine, meets the prevomer ; an outer descending

process which forms the posterior border of the suborbital vacuity

and has a long articulation with the ectopterygoid; and a long

posterior process, the inner part of which apparently articulates

with the basisphenoid, and the outer branch of which doubtless

passes back to meet the quadrate. Though the posterior process

is almost perfectly preserved, as the quadrate is lost the mode of

articulation is not seen. An interesting point about the ptery-

goid is that there are no teeth on it. Near the posterior part of

the anterior palatine process there is ine the middle line a

prominent median spur formed by the two bones meeting, and

immediately behind this is a moderately large median vacuity.

At the inner end of the outer process is a well-marked descending

spur, the exact length of which is unknown as it is broken off

near the base.

Above the outer branch of the posterior process is seen the

well-developed epipterygoid. This differs from that figured by

me in Seylacosawrus (Phil. Trans. 1915) in having a shorter and

broader upper portion and a differently shaped basal, which les in

the pterygoid as seen in the figure given.

The basicranial region is too much weathered to be worth

describing in detail.

ALOPECOPSIS ATAVUS, gen. et Sp. nov.

Aen, ee zy Os atte)

This new genus and species is founded on a nearly complete

skull discovered by me at New Bethesda, and it is of much

importance as revealing another new type of Therocephalian.

Text-figure 4.

Side view of skull of Alopecopsis atavus Broom.

The specimen is in rather indurated shale, and as the bone is

more friable than the matrix, complete development is difficult,

and it has been considered advisable to leave a fair amount of

matrix on the specimen. Enough has, however, been undertaken

to show most details of the structure. The whole of the right

side of the skull is preserved except a part of the quadrate and

squamosal regions, and the right mandible is in position. The

left side has lost the greater part of the jugal and squamosal,

348 DR. R. BROOM ON SOME NEW

and the left mandible has been completely detached and lost

before fossilisation.

The skull as preserved measures 198 mm. and was probably

about 202 mm. in greatest length. Lhe greatest width across

the squamosals has been. about 90 mm. From the front of

the snout to the front of the orbit is 90 mm. ‘The greatest

width of the snout in the canine region is 45 mm., and the

narrowest measurement between the supraorbital edges is 39 mm.

The premaxilla is relatively small, and is much overlapped by

the maxilla. It sends upwards between the nostrils a slender

internasal process to meet the nasal above. Most of the teeth

are lost, but portions of three are stil seen, and from the sockets

the number and size of all can be determined. There is evidence

of six incisors, of which the 6th is very small. The whole six

measure 24 mm. At the front end of the premaxilla are two

moderately large foramina for branches of sensory nerve——

presumably a branch of the second division of the Vth.

The septomaxila lies along the outer border of the nostril, and

passes backwards a short distance between the nasal and maxilla.

Between the septomaxilla and maxilla there is, as in other

previously known early carnivorous Therapsids, a large foramen,

the significance of which is at present unknown. In most feetal

mammals there is a large gland duct developed along the outer

side of the nasal cavity, which fiom its very early development 1s

evidently the remains of some structure once of much greater

importance. It seems not improbable that this large foramen,

which passes into the nasal cavity between the septomaxilla and

maxilla and is continued as a wide groove for some dist2nce

upwards and backwards along the upper border of the maxilla,

was for the lodgement of the glandular and possibly sensory organ

whose rudiment is met with in most mammals.

The nasal is long and narrow, but broader at each end than in

the middie. The shape and relations will be best understood

from the figures given.

The maxilla iy a powerfully developed bone of considerable

depth. There is a single large canine which unfortunately is not

well preserved on either side. In front of the elevation which

accommodates the root of the canine, the maxilla overlaps the

premaxilla as in most Theroceyhalians, and in the anterior part

of the bone are three fairly large foramina, presumably for

branches of the Vth nerve. In the canine region tne surface of

the bone is considerably pitted, probably by glands in the skin,

and there are a number of small openings, apparently for nerves.

The maxillary branch of the Vth nerve apparently passes into

the maxilla by a large foramen on the inner side of the bone

behind the canine, and passing forwards outside the canine, comes

to the surface through the numerous small foramina in the

front of the bone. Behind the canine there is a long slender

alveolar margin, which is remarkable in having no trace of molar

SOUTH AFRICAN THEROCEPHALIAN REPTILES. 349

teeth. In many skulls when no molars are found we suspect

that they may have been lost, but here this can hardly be the

case. Both maxille are well preserved and the dentary as well,

yet in none of the three bones is there any trace of molars.

Further, the alveolar margin is narrow and sharp, and would be

much too slender for the accommodation of molars large enough

to be serviceable to an animal of the size.

The lacrimal is fairly large, and forms a considerable part of

the facial surface, which is very smooth.

The prefrontal forms the anterior and upper quarter of the

orbital margin. It meets the frontal, nasal, maxilla, and

lacrimal bones.

The frentals are relatively small, and the two form most of the

Vext-figure 5.

Upper surface of skull of Alopecopsis atavus Broom.

slightly concave interorbital surface. In front each has a short

articulation with the nasal and a much longer oblique articulation

with the prefrontal, and as the prefrontal nearly extends back-

wards to the postorbital, the frontal only forms a very small part

of the orbital margin. Posteriorly the frontal has a long articu-

lation with the postorbital, and a short interdigitating suture

with the parietal mainly hidden by the postorbital.

There is no postfrontal.

The postorbital is a peculiarly twisted bone. Its inner end

lies against the narrow parietal crest, and from this, passing

350 DR. R. BROOM ON SOME NEW

outwards, it forms the back wall of the frontal region and the

anterior border of the temporal fossa. From the point where it

meets the frontal at the orbital margin it passes almost directly

backwards, forming most of the posterior half of the upper

orbital margin. It then passes downwards behind the jugal to

nearly the level of the lower side of the orbit.

The jugal is a large bone. It forms almost the whole of the

suborbital arch and about half of the postorbital. Posteriorly it

passes back below the squamosal to near the quadrate region.

The parietals are narrow, and form a median crest which in

front is overlapped by the postorbitals. The pineal foramen is

rudimentary.

The back of the skull is not well preserved, the bones being

very rotten, but so far as can be seen the structure does not differ

from that of typical Therocephalians.

Text-figure 6.

Section through skull of Alopecopsis atavus Broom, immediately behind

postorbital arch.

A section through the skull immediately behind the post-

orbital arch shows the relations of the parietals and postorbitals

above and the pterygoid below, with between them a thin

anterior process of the basisphenoid, and above them the feebly

developed element which I believe to be the true vomer, but

which most others call the parasphenoid.

The right mandible is fairiy complete and the dentary almost

perfect. In front there ave three incisors which together measure

9 mm., and theseare closely followed by a long procumbent canine

which has antero-posterior diameter of 8°5 mm. and a height of

something over 20mm. ‘here appear to be no serrations on either

the incisors or the canine. Though the alveolar margin of the

dentary is perfectly preserved, there is no trace to be seen of any

SOUTH AFRICAN THEROCEPHALIAN REPTILES. By) |e

molars. The dental formula of Alopecopsis atavus is thus the

very remarkable one :—i. $, c.},m. f=. As will be seen from

the figure I give, the dentary is very peculiarly shaped. Behind

the canine the upper border of the bone is deeply concave, so

that, even if there had been teeth, they could not have met molars

in the upper jaw unless they were exceptionally long. The back

part of the dentary is powerful, and there is a long but thick

coronoid process. The coronoid bone is well developed, not so

deep as and much thicker than the coronoid in Gorgonopsians

or Cynodonts. The back part of the jaw is not sufficiently well

preserved to admit of description, but itis relatively much shorter

than in Gorgonopsians and apparently more like that of Bawria.

MOscHORHINUS KITCHINGI, gen. et sp. nov.

This new genus and species is founded on a well-preserved

specimen discovered by Mr. James Kitching near New Bethesda

Road. It consists of the anterior two-thirds of the skull of

Text-figure 7.

Upper surface of front of skull of Woschorhinus kitchingi Broom.

a moderately large Therocephalian. The specimen is slightly

crushed, but the matrix is only slightly harder than the bone.

and it has been found possible to display almost every detail

of the structure of the palate.

The skull is manifestly that of a broad-headed short-snouted

form, the front of the snout being usually blunt and wide.

The premaxillaries are broad and powerful, and each carried

six well-developed incisors. As preserved, each bone has only

five teeth, but quite manifestly the 3rd is lost from the right

side and the 6th from the Jeft. The whole series measures 41 mm.

The first four incisors are much flattened and with the long axis

BE DR. R. BROOM ON SOME NEW

directed mainly antero-posteriorly. The posterior two incisors

are more rounded,

The septomaxillary is large, and forms the lower border of the

nostril, but does not extend far backwards between the nasal and

the maxilla. ‘The foramen, usually present between the septo-

maxilla and maxilla, is small.

The nasal bone is exceptionally large and broad. In front it is

crushed down on the nostrils and the details of structure cannot

be made out, but doubtless the arrangement is similar to that in

typical Therocephalians. Posteriorly the nasal meets the frontal

a little in front of the plane through the centre of the orbit.

The shape of the bone will be best understood from the figure.

The maxilla is a short but powerful bone. It overlaps the pre-

maxilla to between the roots of the 3rd and 4th incisors. It has

one large canine which measures 26 mm.x 12mm. Behind the

canine only a very small part of the maxilla shows on the palatal

aspect, and in the specimen there are no teeth, but on each side

there are indications of two old sockets with a possible third very

small one; and I think we may assume that there were three

molars, of which the 3rd was small. The three would measure

about 19mm. The upper dental formula would thus be 1.°, ¢.’, m.*

The prefrontal is relatively small, and wedged in between the

nasal and frontal above and the maxilla and lacrimal below.

The lacrimal is slightly larger than the prefrontal, and it forms

most of the front of the orbit. Near the middle of the bone and

on the orbital margin is a well-developed bony boss, and on the

inner side of this and near its upper end is a large lacrimal

foramen.

Very little of the jugal is preserved, but the part underneath

the orbit forms with the maxilla a very deep and powerful sub-

orbital arch.

The frontals are large and wide, as seen in the figure.

There is no postfrontal. The postorbital lies on the frontal

and parietal.

The parietal crest is narrow as in typical Therocephalans, but

the pineal foramen is rudimentary.

The palate 1s beautifuily preser ‘ved and remarkably interesting.

The palatal portion of the premaxillaries has not been displayed

as the matrix is required to support the fragmentary incisors,

The prevomers form a large part of the bony roof of the mouth.

In front they are ameby laset and form a wide plate between the

internal nares. At the back part of the nares they are much

constricted, but beyond this widen out again into a fan-shaped

structure which lies between the palatines and meets the ptery-

goids. In this back part a median suture between the two

prevomers is very distinctly seen.

The palatine is a large and powerful bone. Externally it has

a long articulation with the maxilla, and here the palatine is so

thickened that it forms more of the wide alveolar surface than

does the maxilla. In front and nearly on the plane of the back of

SOUTH AFRICAN THEROCEPHALIAN REPTILES. 353

the canine, where the maxilla passes up to form the vault of the

palate, the palatine lies closely against the maxilla except where

there is a large foramen, presumably for a nerve. Haughton

figures two foramina in a corresponding structure in Akido-

gnathus. Near the middle of the palatine is an oblique ridge.

which probably supported a soft palate and carried the internal

nares back to the anterior pterygoid region. The inner side of

the palatine articulates with the prevomer and the pteryg god.

The greater part of the posterior border forms the margin of the

suborbital vacuity. Further out isa moderately large articulation

with the ectopterygoid.

The ectopterygoid is an irregularly shaped bone, not unlike a,

bird’s quadrate. Anteriorly it articulates with the palatine ;

externally with the maxilla and probably with the jugal. Be-

tween the maxilla and the ectopterygoid is a small foramen.

Posteriorly there is a large flat articulation between the ecto-

pterygoid and the pterygoid. In front of the pterygoid process

there is an even larger ectopterygoid process. All the outer side

Text-figure 8.

Palate of Moschorhinus kitchingi Broom.

of the suborbital vacuity is formed by the ectopterygoid and

much of the posterior.

Only the anterior portions of the pterygoids are preserved.

From the broad deep lateral portions which articulate with the

ectopterygoids they pass forward only a comparatively short

distance to meet the prevomers and the palatines. In this region

each pterygoid sends down a thin vertical plate, which, uniting

with its neighbour, forms a median keel.

In general structure the palate agrees more closely with that in

Akidognathus, though the proportions of the bones are very

different. In Akidog gnathus the prevomers have, as in JMJoscho-

rhinus, a suture between them, and yet are anchylosed in front.

Moschor hinus differs in having no interptery ygoid vacuity so far

as can be seen in the specimen. If one be present, it must be

much further back than in Ahkidognathus.

354 DR. R. BROOM ON SOME NEW

There is another palate recently discovered with which a

comparison may be made—viz., Whaitsia platyceps. This new type

was recently described by Haughton, and though it differs greatly

from JMoschorhinus, the snout has so much superficial resemblance

that a comparison seems advisable, even if only to clearly differen-

tiate the types. Whaitsia is a large Therocephalian—one of the

largest known. It has a broad flat snout, and if it were not for

the. great difference in the palate, one might be inclined to think

that Moschorhinus was a near ally. Barther, in the type of

Whaitsia the number of incisors cannot be anal out, though

there are four in what is regarded as a co-type. The palate,

however, is so very unlike that of any form previously known

Text-figure 9.

Diagram of palate of Whaitsia platyceps Haughton.

that Mr. Haughton very wisely puts Whattsia in a new family—

the Whaitsidie. ;

Mr. Haughton has given an excellent description cf the type

and a fieure of both upper and lower sides. The description he

has given is so accurate that little need be added to it. But the

fisure he gives of the palate is not very clear, and a new and

independent figure of this unique palate may not be regarded as

superfluous.

If the palate, as I figure it, be compared with that of Moscho-

rhinus, it will be seen to differ in only two important points.

The suborbital vacuity, which in all typical Therocephalians is

large, is here practically closed, and the back part of the palate is

thus made to resemble slightly that of the Gorgonopsia. In

front, instead of there being a pair of large openings as in all

‘ SOUTH AFRICAN THEROCEPUALIAN REPTILES. 355

other known Therocephalians, there are four openings. If the

anterior part of the palate be compared with that in Jfoscho-

rhinus, it will be seen that the difference is that the anterior ends

of the palatines pass inwards, and meet the prevomers dividing

what were large internal nares into anterior and posterior

portions —the posterior alone being, as Haughton has suggested,

the internal nares. What has happened is exactly comparable to

what has happened with the external nares in Chameleon, where

the original opening becomes divided into two by the prefrontal

passing forwards.

Haughton discusses the affinities of Whaitsia with Thero-

cephalians, Gorgonopsians, and Cynodonts, and inclines to regard

it as a highly specialised Gorgonopsian, but it seems to me that

it is a true but aberrant Therocephalian.

References to lettering.

Ang. Angular; B.O. Basioccipital ; B.S. Basisphenoid; Dené. Dentary ;

He.P. Ketopterygoid or Transpalatine; Hp.P. Epipterygoid; Fr. Frontal; Ju.

Jugal; Mv. Maxilla; Na. Nasal; OpO. Opisthotic or Paroccipital; Pa. Parietal ;

Pal. Palatine; Pme. Premaxilla; PrF. Prefrontal; PoO. Postorbital; Pd.

Pterygoid ; P.Vo. Prevomer; Sma. Septomaxilla; Sg. Squamosal; WVo=Pa.8.,

Vomer=Parasphenoid.

Proc. Zoou. Soc.—1920, No. XXIV 24

P. Z. S. 1920, FORSTER- COOPER, PI. I.

Bale & Danielsson, Ltd.

CHALICOTHEROIDEA from BALUCHISTAN.

ON CHALICOTHEROIDEA FROM BALUCHISDAN. abv

19. Chalicotheroidea trom Baluchistan. By C. Forsrer-

Coopur, M.A., F.Z.8., Superintendent of the Uni-

versity Museum of Zoology, Cambridge.

[Received April 16, 1920: Read May 11, 1920.]

(Plate I.* and Text-figures 1-7.)

Dr. Pilgrim, in his report f on the fossil vertebrate fauna of

the Bugti Hills, has described two forms of Chalicotheres, of which

the smaller is represented by material too fragmentary to be

named, while to the larger he has given generic rank with the

name of Phyllotillon naricus.

My own collections have produced a small amount of further

material for study, fragmentary and in some cases much worn,

but sufficient to add in some points to our knowledge of these

two forms.

Holland and Peterson — in their very complete memoir on

Moropus elatus and the osteology of the Chalicotheroidea have

taken exception to the creation by Dr. Pilgrim of a separate

genus for lis species, Phyllotillon naricus. This objection is

reasonable, seeing that the chief character on which the genus

is founded is one which occurs in other genera of Chalicotheres.

Dr. Pilgrim states$ that “the difference is particularly well

shown in the structure of the upper premolars. ... An unworn

specimen of pm.3....shows that the large inner cusp was

united to the ectoloph by a double instead of a single crest.”

This statement, as Holland and Peterson point out, is open to

alternative interpretations—viz., either that there is a crest

running from each side of the inner cusp to the ectoloph, or that

each crest is double.

The former interpretation represents the actual condition of

the tooth, and is the meaning Dr. Pilgvim wished to convey||. As

this condition is fonnd in JJoropus it loses its value as a generic

character. On the other hand, to judge from Depevet’s figure of

Macrotherium grande 4, the fourth premolar does appear to differ

in the arrangement of the crests from that of the Bugti specimens

in that the anterior crest seems less developed in MW. grande, in

which form the anterior external cusp is a separate rounded

hillock. Gaudry’s figure of Schizotheriwm modicum shows both

* Wor explanation of the Plate, see p. 366.

+ Rec. Geol. Surv. Ind. x]. p. 67, and Mem. Geol. Surv. Ind. n.s. vol. iv. Mem. 2,

p. 33.

{ Mem. Carnegie Mus. vol. iii. no. 2, 1913.

§ Mem. loc. cit. p. 33.

|| Lhad the opportunity of consulting Dr. Pilgrim on this poimt during a short

visit made by him to Cambridge, and have his authority for making this statement.

| Arch. du Mus. d’Hist. Nat. de Lyon, xol. v. 1892, pl. iii. fig. 1.

24*

358 MR. C. FORSTER-COOPER ON

ridges, but the anterior seems to be weaker than the posterior,

while the preceding premolars seem to have no anterior ridges.

Chalicotherium sivalense is like Schizotheriwm in this respect,

according to Falconer’s figures in the ‘ Fauna Antiqua Sivalensis,’

plate Ixxx.

There are other differences from JJacrotheriwm in the shape of

the various teeth, so that there are some grounds for Dr. Pilgrim’s

separating them.

Whether or not the Bugti specimens can be placed in any

other described genus is even more difficult to decide. Our com-

bined material is: scanty and none too well preserved, so that

many characters remain unknown or at best uncertain, and for

the present it seems useful to retain the generic name of

Phyllotillon.

The material forming the basis of the present description

consists of :—

1, A series of upper teeth, pm. 3-4, m. 1-2, much worn but

giving the outline and general shapes of the teeth.

A moderately worn third and fourth upper premolar.

Three separate upper molars.

A fourth upper premolar only a little worn.

Fragmentary lower molars and lower jaws, sufficient when

combined to give an outline of the middle portion of

the mandible.

6. Some separate toe bones.

x — Co ho

The first and second specimens (text-fig. 1) supplement Dr. Pil-

grim’s figure * in giving a better preserved shape of pm. 3 and

Text-figure 1.

Phyllotillon naricus. 3rd and 4th premolars, Ist and 2nd molars. X 4,

in adding the shape of the second molar. The third premolar as

well as the fourth (this outline filled in from the second specimen)

have, roughly speaking, a square outline; the anterior outside

border is produced into a somewhat prominent style, more pro-

nounced than in Macrotheriwm and to much the same degree as

in Moropus. In proportions the teeth differ from other deseribed

* Mem. loc. cit. pl. xii. fig. 2.

CHALICOTHEROIDEA FROM BALUCHISTAN. 359

forms in the greater approximation of the length and breadth,

being square e rather than transversely elongated.

The upper figure in Plate L. represents the fourth upper

premolar in a slightly worn condition, and illustrates the two

ridges described by Pilgrim; in the centro-internal cusp and

broad internal cingulum it shows features characteristic of all

Chalicotheroids.

The molars are all elongated, and in this respect differ from

those of Macrotheriwm, where they are al] square, and from NVestort-

therium and Circotherium, where the molars change from square

in the front one to elongate in the third. They agree best with

those of Moropus, but carry the elongation to a still greater

degree, and represent the greatest modification in this respect.

The lower figure in Plate I. shows two right upper molars just

erupted, only the posterior half of the second being preserved.

The condition of this fragment is excellent. In shape and in

pattern they resemble Dépéret’s figures * of the milk teeth of

Macrotheriwm and also those figured by Pilgrim for this species 7.

They are, however, in all probability the permanent second and

third molars, being considerably larger than the measurements of

the milk teeth given by Pilgrim—i. e., the third molar is 48 mm.

long and 36 mm. wide in the front half as against 40 mm. and

31 mm. The only points in which these teeth differ from

Pilerim’s description of his specimen are that here the whole

external surface of the imetacone—i.e¢., from mesostyle to

metastyle—is considerably smaller in the third molar than in the

second, and that faint ribs are present on the external surface of

the metacone of the second tooth and quite absent on that of the

third. A corresponding rib, rather more clearly marked, is

present on the paracone external surface of the third molar,

and esa pe would be found in the other molars. ‘These ribs

are not shown in Dépéret’s figures {, and in Holland and Peterson’s

figures of JMoropus they are Shown only on the protocones of

all three molars, where, however, they appear to be strongly

marked.

Text-fig. 2 shows the outline of one of the fragments of mandible

which has the socket for three incisors, or the second and third

incisor and a canine—all apparently small-rooted teeth—and the

whole premolar-molar series except the last half of the third

molar. This is continued by another fragment (the dotted outline

in the figure) with the third and the second half of the second

molar. As these fragments coincide in size they may be con-

sidered as belonging to the same species.

The mandible thus reconstructed shows certain peculiarities.

Compared with the best-known Chalicothere (J/oropus elatus), as

figured by Holland and Peterson §, it has a much flatter lower

* Loe. cit. pl. 11. figs. 4 & 5.

+ Mem. pl. xii. fig. 3

10 ILO; Cie

§ Loe. cit. pl. li,

’

Phyllotitlon naricus.

the proportions are entirely different.

jaw is not known, but in the fragments at hand there is no sign

a slight one.

360

MR. GC. FORS'TER-COOPER ON

border

molar.

and is without the swelling under the anterior pre-

There is evidence, however, of a similar swelling further

forward, which in this jaw is just under the mental foramen.

This point marks the beginning of the symphysis, and the lower

ridge-like.

border in the region just behind the symphysis is sharp and

M. elatus.

There is apparently only one mental foramen instead

of several, but it is much larger in proportion than those of

The diastema between the incisor

and

premolar is small (civea 25 mm.), and the latter tooth seems to

have been very much reduced in size.

length of JZ. elatus, while the teeth are nearly as big, so that

The jaw is about half the

Text-figure 2.

2°

Fragment of mandible with dotted outline of another

fragment in continuation.

The hinder part of the

of a downwardly-turned angle, though Pilgrim’s figure * suggests

The flat lower border is a point of difference from J/aero-

therium rhodanicum and other forms in which this feature is

known, while the shortening of the anterior part is a point in

common with W. rhodanicum and Chalicotherium sivalense.

The subjoimed measurements show that the animal occurred in

two sizes, which possibly represents a sexual difference :—

* Loe. cit. pl. xii. fig. 4a,

Upper Teeth.

CHALICOTHEROIDEA FROM BALUCHITSTAN.

361

| Premolars. Molars.

Third. Fourth. First. Second. | Third.

| eA si KAS Gas eae Wa awesbes Parke 2 +

}

Length Breadth Length Breadth} Length | Breadth: | Length! Breadth : Length | Breadth:

| | | ifront hind} front hind) front hind!

- 93 ewe epee ne cole | | |

| | | |

23 2 aE NS 27 32 FOR 27 37 35 — = ==

| | | |

| | ? > 35) 48 «| 36 84 |

| fli ea Bal 80) |

Three separate molars | | 20 |

2 2nd or 3rd ; au SES |

| |

{|} 41 | 32 28

Lower Teeth.

19? ? 23 1 |) S10 A aKa 2 Ze R22 iP 22 ?

| | | |

ee lL ek Moen 28 2) | 16 182] 40love psete at mene ae

| | |

| 23 13 24 15 | | | |

| | | | | 4a |29 9

| | | | | | |

| | | | | 51 | 25 24]

| |

Measurements in millimetres.

A few toe bones found in these deposits are noticed at the end

They show certain characters of interest, but as

they may be the smaller toes of this species or the larger ones of

of this paper.

the following,

The main

tollows :—

characters of this form

they are of no specific value.

may

be summarised

ane Sint 0

Dentition 5-5 or 9° pm. 4, m.5- Upper premolars square.

Lower pm.2 small (pm.1 absent above and below). pm.3 & 4

elongate. All molars elongate, especially the second. Lower

border of lower jaw unusually flat: one mental foramen. Jaw in

section rather flat outside and slightly convex inside. Moderate

diastema in front of anterior lower premolar. There is thus a

mixture of various generic chaiacters, the balance, perhaps,

favouring a general resemblance to Moropus. The form is at

present insufficiently known, but there is enough to warrant the

retention of Dr. Pilgrim’s generic name until further material is

discovered,

362 MR. C. FORSTER-COOPER ON

SCcHIZOTHERIUM PILGRIMI, sp. lh.

Pilgrim also mentions the presence of a smaller form in these

deposits, indicated by some fragments too much worn for accurate

description. In the present collection are two unworn and _per-

fectly preserved upper molars (Plate I., central figure) of an

animal much smaller than those of the species just described,

and which may belong to Pilgrim’s undescribed species although

even a shade smaller than the latter.

‘The smaller size of these teeth suggests at first that they are

milk teeth, but there are certain arguments against this view,

which is not adopted here. The specimen fortunately has just

enough of the anterior part on the zygomatic arch to show the

relative position of the teeth in the maxilla (text-fig. 3). A

reference to Holland and Peterson’s figure * shows that in the

Text-figure 3.

Schizotherium? pilgrimi. 1st and 2nd upper molars in side view, showing their

position relative to the zygomatic arch. The position of the antorbital foramen

is marked with an *. Nat. size. Type-specimen.

milk and permanent molars of Moropus the last milk tooth lies

well in front of the anterior root of the maxillary process of the

zygoma, the first molar under the root of this process, and

the second under its posterior border. If the present specimen

is grown in a similar way, it supports the view that the teeth are

the first two true molars. The anterior tooth is slightly worn, the

second only just erupted. Beneath them there are cavities without

trace of successional teeth and too small to allow of them. The

teeth being fully formed and practically in wear, the germs of the

succeeding teeth would have been comparatively well developed.

Moreover, of two fragments of lower jaws one has the last milk

molar fairly well worn and the front part of the first lower molar

just erupted. One side of this specimen has been ground away,

and the section (text-fig. 4) shows the last milk molar with the

* Loe, cit. pl. li. figs. 1-2.

CHALICOTHEROIDEA FROM BALUCHISTAN. 363

section of the edge of the permanent succeeding tooth below it;

there is no germ below the next tooth, whose roots are much

larger and straighter. This tooth fits the front tooth of the two

upper molars mentioned above.

A second specimen shows two lower molars well worn, of which

the first is about the same size as the first true molars of the

previous specimen. The fragment is broken away underneath,

and the roots show that they belong undoubtedly to permanent

teeth.

Text-figure 4.

Schizotherium? pilgrimi. Fragment of lower jaw with last deciduous tooth and

Ist true molar. One side of the specimen is ground away, showing the edge of

the germ of the 4th premolar, marked with an *. The anterior root of the

deciduous tooth is in section, the level of the posterior root not being quite

reached by the grinding. The stout front root of the 1st molar is in section, the

posterior is partly in section and partly a cavity: 7. e. the lower half, where the

root has fallen out. Nat. size.

The upper dentition being the best preserved is here made the

type-specimen of the species.

Measurements (in millimetres) :—

Upper molars. Type-specimen.

1st molar. 2nd molar.

UOMO GN kee Sean. a... 21 25

Breadth, front half... 20 24

5 second half.. 20 23

Lower molars. Ist. 2nd. Ist. 2nd.

Wemothee Ftc stihl 18 — 18 22

Breadth, front half... 10 — 10 12

« second half... 2 — jah 12

Figured specimen (text-fig. 4). Second specimen.

The upper molars are square, and resemble those of Schizo-

theriwm modicum in size as well as in shape. The attribution of

the present species to the genus is tentative.

364 MR. C. FORSTER-COOPER ON

There is yet another specimen of a lower jaw with a much-

worn first molar and roots of the last premolar and second and

third molars. The first molar is 22 mm. long and about J2 mm.

broad, and is therefore larger than the other specimens. The

lower border of the jaw is extremely straight, as in Phyllotillon

naricus. In size it seems to correspond with Pilgrim’s unnamed

fragment. Whether it represents a sexual difference from

Schizotherium pilgrimi ov yet a third species remains unsolved.

Text-figure 5,

? Metacarpal or metatarsal ofa Chalicotheroid. Nat. size.

Until the anatomy of these animals, and indeed of all Chalico-

theres with the exception of Moropus, is better known, it is

impossible to feel sure of their generic position. A good deal

of interest lies in the condition of the feet and the gradual loss of

the first and fifth toes. According to Holland and Peterson *,

both should be present in the fore foot of the Schizotheriine. In

the present collection are several phalanges and one bone which

may be a lateral podial (text-fig. 5); it is just possible that it

may represent a fifth metacarpal or tarsal. Of the phalanges

three out of six show the co-ossification of the proximal and

median (text-fig. 6), which was evidently a common feature, as

Holland and Peterson describe it in Woropus. The free proximal

phalanges all show strongly-marked tubercles on the posterior

faces (text-fig. 7) for the flexor attachments. Some of the

fused bones show this feature, but to a less degree. They are

not to be found in corresponding bones of Macrotheriwm or

* Loe. cit. p. 201,

CHALIGOTHEROIDEA FROM BALUCHISTAN. 36 a

Chalicotherium in the British Museum collection, nov have they

been noted or figured elsewhere. It is not possible at present to

assign these bones to one or other of the Baluchi species.

Text-figure 6.

Co-ossified proximal and median phalanges of a Chalicotheroid,

lateral surface. XX 4.

Text-figure 7.

Median phalanx of a Chalicotheroid, posterior surface showing

the two tubercles. X 3%.

It is interesting to note that while remains of Anthracotheres

and Rhinoceroses of all kinds are very abundant in the Dera

Bugti deposits, Chalicothere remains are here, as elsewhere,

extremely rare. They seem to have been of solitary as well as

of strange habits, though recent discoveries seem to point to

Moropus having been to some extent gregarious.

Note.—Vhe specimens mentioned in this paper together with

all the other Chalicothere material from Baluchistan have been

added to the British Museum collections.

366 ON CHALICOTHEROIDEA FROM BALUCHISTAN.

EXPLANATION OF PLATE I.

Upper Figure.

Phyllotillon naricus. 4th upper premolar. Nat. size.

Lower FIcuReE.

Phyllotillon naricus. 8rd upper molar and part of 2nd, Nat. size.

CENTRAL FIGURE.

Schizotherium 2 pilgrimi. 1st and 2nd upper molars. Nat. size ; type-specimen.

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WEST AUSTRALIAN PERIPATOIDES.

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WEST AUSTRALIAN PERIPATOIDES.

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WEST AUSTRALIAN

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PERIPATOIDES.

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WEST AUSTRALIAN PERIPATOIDES.

ON THE ONYCHOPHORA OF WESY AUSTRALIA. 367

20. Fauna of Western Australia.—III. Further Contributions

to the Study of the Onychophora. The Anatomy and

Systematic Position of West Australian Peripatoides,

with an account of certain histological details of general

importance in the study of Bie pennas By Wm. J.

Dagny, D.Se., F.Z.S., F.L.S8., Professor of Biology,

University of Western iAristinailtad

[Received March 16, 1920: Read May 11, 1920.]

(Plates I.-V.*)

INTRODUCTION.

There are yet many details of interest in the structure of

Peripatus which remain problematic. The histology of the eye

and the structure of the trachee may be cited as examples ;

whilst even the presence of cilia, an unusual character for an

arthropod, has been doubted by some histologists.

In addition to general features, such as the above, there is

always the consideration that the anatomy of the different species

of Peripatus should be thoroughly well known, especially as in

many places there is a tendency for the advance of civilisation

and closer settlement to drive this somewhat rare animal to

extinction. It has been comparatively easy in Western Australia

for the author to procure specimens of this interesting group of

arthropods and the facilities for examining specimens being so

to speak on the spot, there seemed a likelihood that a close study

would reveal something new regarding the West Australian

species at least. Further support was lent to this view by reason

of the fact that one West Australian species, the first to be

discovered, had only been collected once many years ago, and was

very briefly and quite insutticiently described. Practically no

details of the anatomy were known.

In all, three species of Peripatus—or, perhaps more correctly,

Peripatoides—have been recorded from Western Australia,

namely :—

1. Peripatus leuckarti, var. occidentalis Fletcher, syn. Peri-

patordes oscidentalis Dendy.

2. Per ipatoides gilesit Spencer.

3. Peripatoides woodwardi Bouvier.

. Of these three, Peripatoides occidentalis was the first to be

collected. It was discovered about 160 miles south of Perth, at

* For explanation of the Plates, see p. 388.

368 PROF. W. J. DAKIN ON THE

Bridgetown, by a Mr. Lea and named by Fletcher (Proc. Linn.

Soc. N.S.W. 1895 (2) x.). Twelve years later, Peripatus was

found in another locality much farther north, in the hills at

Armadale, only about 18 miles from Perth. Specimens of these

collected by I Mr. H. M. Giles were sent to Professor Baldwin

Spencer. He found them to belong to a new species, and in a

short paper, published (12) in 1909, “named it P. g gilesit after the

collector. Specimens of the supposed third species had been

collected four years before this, ¢.e., in 1905, by a German

Expedition (The Hamburg Exped. of Michaelsen and Hartmeyer).

They were also collected near Perth and in the hills. These

specimens were sent to Bouvier, who wrote a detailed description

of the anatomy and discussed the rel: tionships of the species to

other known Australian forms. Bouvier’s paper (3) appeared in

te the same year in which Spencer's description of P. gilesti

vas published. The supposed occurrence of two species in the

same district and the fact that both descriptions appeared in

the same year aroused the curiosity of the present author, and

the result of his investigations (see Proc. Zool. Soe. London,

June 1914) confirmed his suspicions. Peripatoides woodward

and Peripatoides gilesii turned out to be one and the same species,

and since Spencer’s description was published some months before

Bouvier’s paper, the name PL. gilesii took precedence over

P. woodwardi. The fullest account of the species is, however,

to be found in the paper of Bouvier (8) under the name of

P. woodwardi. Now, since the previous publication of the writer

(Dakin, Proc. Zool]. Soc. 1914), an exploration to the S. W. resulted

in the collection of over 100 specimens of the Peripatus first

made known from West Australia, 72. e., Peripatoides occidentalis.

The consequence was that the entire question was reoperied,

About 100 specimens of P. gilesit were collected for the purpose

of a detailed examination and comparison of both species, the

northern and southern. The conclusion of this research was

rather surprising. In all the peculiarities that marked the

northern species (P. gilesii) the southern species agreed. The

difference between the two forms was so slight that we could not

regard them as more than varieties.

"Thus we have reduced the number of species of Peripatoides in

West Australia from three to one. The southern type must

retain the name Jeripatoides occidentalis—the northern form

should be known as Peripatoides occidentalis var. giles. Further

than this, however, a detailed investigation of the specimens has

shown that Bouvier’s description of the anatomy contains several

inaccuracies, some of which are decidedly important from the

point of view of comparisons. These inaccuracies are without

doubt excusable, for the number of specimens at the disposal of

Bouvier was small and the preservation could not have been all

that was to be desired.

ONYCHOPHORA OF WEST AUSYRALIA. 369

OCCURRENCE OF THE SPECIMENS OF PERIPATOIDES

IN WEST AUSTRALIA.

It is noteworthy that all the specimens of Peripatus taken up

to date have been found in the hills and at some little elevation

away from the coastal plains. On the other hand they have not

been found very far into the interior. They have been found in

situations which, at least during the winter months, are some-

what damp but not excessively so. There is no doubt that

Peripatus extends over a wide area in West Australia, but it is

very difficult to map out this area, for collecting is not exactly an

easy task and the animals have their usual eccentric local

distribution within if. For example, the author went out on

one oceasion to a small valley in the hilis where Peripatus was

believed to occur, An entire morning was spent searching

without signs of a specimen, and it was decided that after lanch

a new spot should be tried further ahead—the next little valley,

in fact. The next valley turned out, however, to be the one

where we had imagined ourselves all the morning, and we soon

found Peripatus there as on previous occasions. Here were two

little valleys only ten minutes apart, with the same vegetation,

the same amount of moisture, and presenting such a similar

appearance that one had been mistaken for the other. Yet in

the one we found many specimens of Peripatus, in the other,

none.

Specimens have up to date been captured at the following

places:—Lion Mill, Mundaring Weir, Armadale, Kelmscott,

Kalamunda (all specimens found at these places were supposed

to be P. gileszt), and at Jarvahdale and Bridgetown in the 8.W.

(P. occidentalis).

Most specimens bave been obtained at Mundaring Weir,

Armadale, and Jarrahdale. At the latter place 13 specimens

were found under a small piece of branch about 1 foot square,

and nearly 150 specimens were captured in three days. Very

few indeed have been found under stones or under bark or fallen

trees. The usual place is on the surface of the soil underneath a

small or large piece of wood (a fallen branch or part cf one).

Herethe ground is more or less damp and there is no grass. In

the same situation white ants are by far the most ubiyuitous

creatures, with nests of true ants frequently lending variety.

Large centipedes are often commen, but not usually with

Peripatus, which prefers, on the whole, the absence of ants,

centipedes, and millipedes from its particular sheltering log.

In the summer no specimens have ever been found and this

despite some very arduous work in the broiling sun. Peripatus

is, however, extremely sensitive to drought, and before the dry

season—November or December to April—it must find its way

either below the surface of the ground or far into the erannies

and cracks in fallen and decaying logs. As the breeding season

coincides with the dry season it is impossible to study the

370 PROF, W. J. DAKIN ON THE

embryology of West Australian Peripatus, unless specimens are

kept alive under artificial conditions in the laboratory through

the long summer.

The regions in the hills where Peripatus is found are baked

dry in the summer, and the temperature rises frequently to

104° F. in the shade in these valleys. The hidden recesses where

protection is sought must be somewhat moist, for on several

occasions the mere carriage of specimens for a few hours in a

cardboard box, with none or insufficient moist earth, has resulted

in the death of the animals through drying up. This is all

the more noteworthy, since half-a-dozen specimens will live for

days in a small glass tube. When suddenly exposed to the light

of day the animals remain motionless, but after a few minutes

they may move quite actively to get out of the light. Slime is

often ejected from the slime glands on touching the specimen

with the forceps or the finger.

EXTERNAL CHARACTERS.

CoLtour AND Parrern.

Both the varieties of Peripatoides occur in two rather different

colours, brown, and dark olive-green, and the numbers of each —

are practically equal. Between these two shades there are

specimens bearing various intermediate tints, and frequently the

brown specimens are so marked with dark brown (almost black)

that they have quite a variegated appearance.

Closer investigation with a dissecting binocular microscope

brings out the fact that the entire surface of the animal’s body is

covered with small papillae. The pigment in the brown specimens

is arranged as follows: There is an almost uniform brown back-

ground from which arise the papille above-mentioned. These

papille are either brown (a little darker than the background) or

jet black. Some of them, however, have a pale yellow area round

the base. The arrangement of the papille is responsible for the

dark patterns of the skin.

In a very dark brown specimen the effect is due to the back-

ground being much darker, the papilla remain the same, and the

pale yellowareas,as round the bases of certain black-tipped papille,

are conspicuous. If the background is still darker—almost

black—with a slight tinge of the brown in it, the effect of the

black papille, the pale yellow areas round some of them, and

the colour of the background is to produce the very dark olive-

green shade of some of the specimens.

The ventral surface is more free from papillze and the general

background is an almost clear white. In the bright brown

specimens, between the legs of each pair there are two patches

where the background is a pale grey—a patch to either side

of the middle line. On each patch there are minute black papille.

Between each successive pair of legs the background is more or

less tinged with pale orange brown and there are minute darker

ONYCHOPHORA OF WES! AUSTRALIA. Sel

orange papille. Hach ring here hears larger and more separated

papille which are quite white. They seem to correspond to the

white areas round papille of the dorsal surface.

In both varieties an extremely fine light median lne runs the

length of the dorsal surface. It may not appear so in all

specimens at a first glance. This, however, is due to the fact

that in the light brown specimens, where it seems clear, it runs

down the middle of a narrow band somewhat paler in colour than

the rest of the dorsal surface and rather free from papille. In

the dark specimens, on the other hand, this narrow band is

darker than the rest of the dorsal surface, and with the naked

eye or low power may be all that is observed. A higher magni-

fication will shew that a very fine light line runs through it

medially, as in the light-coloured specimens. :

The longitudinal band of the dorsal surface referred to above

is related to a slight depression, the fine white line being an

extremely narrow groove. The narrow band was noted by

Bouvier, but the fine median white line escaped his notice.

In both varieties there are always a number of clear white

papille between the successive legs at about the ventral margin

of the flanks. These correspond, however, to the large papille

of the dorsal surface and have merely lost the black pigment—

they are identical with the large clear papille of the ventral

surface.

The skin is thrown into a number of folds or ridges, as is usual

in Peripatus. There are about 14 of these between two similar

points opposite two successive legs. The ridges are to be seen

both dorsally and ventrally, but are not continuous round the

entire circumference of the body, for they are interrupted in the

middle of the dorsal surface by the longitudinal line. But for

this, however, some are continuous. Others arise between these

larger folds and do not run so far. Naturally those opposite the

legs are not continuous on to the ventral surface. The folds are

not all of the same width, but it can hardly be said that they ave

alternately wide and narrow in either variety. There is really

very little difference.

OrAt, PAPILLA.

The oral papille are to be found in their usual position—there

is nothing of particular interest to add with regard to them.

On many oceasions slime was shot out from the openings of

slime glands on these oral papille when the specimens were

touched. It was emitted in large quantities when the animals

were dropped on to water containing a little formalin, and also

when the specimens were narcotised with chloroform.

Tun Crrenatic Raion.

Bouvier (8) has described in considerable detail the rings or

ridges of the integument at the bases of the antenne and in the

Proc. Zoou. Soc,— 1920, No. XX V. 25

)

ia PROF. W. J. DAKIN ON THE

neighbourhood of the eyes. He recognises in certain Peripatus

species what is termed an ocular ring of papille, with part of it

differentiated to form a frontal organ. In others he states :—

“Vareceau oculaire sattenue, puis disparait assez brusquement

aprés avoir déerit un peu plus d’un demitour, mais a ce niveau

ou méme bien plus en dédans, l’arceau infra-oculaire prend une

remarquable prédominance, forme parfois un organe frontal et se

continue par Varceau spiral.” The northern variety (Bouvier’s

Peripatoides woodwardi) is supposed to be one of the forms most

typical of this group of species.

Several specimens of both varieties have been examined, and,

so far as can be made out, there is no difference greater than the

variations met with in either variety, between the two forms.

Tue Lees.

The number of legs appears to be very definite in the West

Australian Peripatoides, and no variations are to be recorded

amongst the individuals from any one area. There is, however,

a marked difference between the two varieties which inhabit the

northern and southern areas respectively.

Bouvier stated that Peripatoides gilesti vesembled P. suteri and

differed from all the other Australian Peripatoides in the possession

of 16 pairs of legs. At the same time it was stated to differ

from P. suteri in only having three pedal papillae, in which respect

it agreed with other Peripatoides.

Fletcher diagnosed the southern variety, P. occidentalis, as

possessing 15 pairs of walking-legs, but no further description

was given. ‘This is the difference between our two varieties,

and it is most constant. All the specimens from Mundaring,

Armadale, Kelmscott, ete., in the north, have 16 pairs of legs,

whatever be their size or sex. All the southern specimens

bear only 15 pairs of legs. All the legs are similar with the

exception of the 4th and 5th pairs in both varieties, from

which one may conclude perhaps that it is one of the posterior

pairs that is missing in the southern type.

Each leg is marked by rings bearing papille, but near the

apex on the ventral surface these papille have united to form

spinous pads, the middle one of which is somewhat wider than

the others. Now in the northern variety the 4th and 5th pairs

of legs differ from the others in the fact that the proximal spinous

pad is segmented, a small central segment being cut off from

two larger lateral lobes. This central segment takes the form of

a papilla upon which the duct of the excretory organ opens.

They are accurately figured by Bouvier, who remarks that the

urinary papille of the 4th and 5th legs are always independent

of the neighbouring parts of the proximal spinous pad, whereas

they are always adherent in Peripatoides suteri and sometimes in

P. orientalis ( P. leuckartii). We may now add that the condition

described above holds good in every detail for the southern

form too,

a ne ee

ONYCHOPHORA OF WEST AUSTRALIA. Be

THe MANDIBLES.

The character of these structures has been used very con-

siderably in systematic works, and consequently they have more

than a little interest for us here.

Each mandible consists as usual of two blades. ‘The outer

blade presents a single large projection, but no small teeth.

The inner blade is provided with a large principal tooth and a

number of smaller accessory teeth. According to Spencer (12)

there are four clearly marked and one minute accessory tooth,

whilst Bouvier remarks that, as in P. saéteri, P. nove zealandic,

and P. occidentalis, there ave five accessory teeth. In the course

of this present research the mandibles have been removed from a

large number of specimens, and examined, with the result that

whilst one can say that five accessory teeth are most common

upon the inner blade, there may be six or even seven. In the

latter case the extra teeth are very small. There is again no

difference to be noted between the northern variety and the

southern form previously known as P. occidentalis. ‘here is never

an accessory denticle at the base of the large tooth on the outer

jaw blade.

EXTERNAL SEXUAL DIFFERENCES AND CrurRAL PAPILLA.

There is no difference in the number of legs borne by the two

sexes as In Peripatus nove-britannie, although the two sexes are

to be distinguished by other characters associated with these

appendages, 7. e., the crural papille. The female aperture is

larger than the male aperture, as noted by Bouvier, and both

apertures ave found between the bases of the legs of the last pair.

Behind the aperture of the reproductive organ in the male, and

consequently just posterior to the last pair of appendages, are

two small openings which can be recognised by their slightly

tumid, pigmentless lips. These are the apertures of the anal

glands.

The Crural Papille ave the most distinctive features of the

male. Unfortunately there has been some confusion as to the

number present, and these structures have been taken as of

considerable systematic importance. Bouvier (3) gives the

following table for three males in his collection (the species

formerly known as P. woodwardi ov P. gilesti) :—

ucla ile Dh, Spel MERU Mi Poare bay. m1Ou iUllaloa alee Tea laeans

i Rt. iLL aa GTO Gn I PET Te apOT Og UGier

Sp. ¢

Lft. iL G0.) ea a eoeh Nee Ep eae AT a Tel clap lv (0) a

Reet 1 lo MOO ONO NON O Momo Olmo IO womor

Sp. d.

Lft. TT Sk Obid 1,00) 1h ont aOR emo

Hy eR i, TL ULE Ge ME aT) -GLA@ AGM Lae ub

sp.t4

Witt: OM eIeO! "0. 1) Voc Ace KOMEOME OM GANG

O5*

374 PROF, W. J. DAKIN ON THE

It will be noticed that there is a very large amount of variation.

Bouvier found, however, that in many cases the papille could

be discovered by a study of the anatomy. He concluded that

there was reason to believe that Peripatoides woodwardi possessed

erural papille for all the legs with perhaps the exception of the

last two pairs, although further investigation might show them

to exist there too.

In making a comparison with other Australian species of

Peripatoides, Bouvier states (relying on Fletcher's description of

P. occidentalis (7)) that our northern variety differs from all

other Australian forms in the possession of crural papillae on the

lst pair of legs. In fact, this is stated to distinguish P. woodwardi

from all other known species of Peripatus. This statement

renders an examination of the southern variety particularly

interesting.

Let us first take Bouvier’s northern type. We have found

that the mere presence or absence of crural papillae when examined

externally means almost nothing. Sections show always that the

papille are present on certain legs but that they may he either

invaginated (see P]. ITT. fig. 4, Cru. pap.) or protruded. Every

specimen examined has had different crural papille protruded.

It may either depend upon the fixative or the animal may pro-

trude certain papille at definite times. The preserved specimens

would then indicate the condition at the time of fixation.

The fact remains, however, that crural glands are present in

the male in every leg (PI. I. fig. 3, Cr.Gl.), and papille are to

be found on all of them too. This is a correction to Bouvier’s

otherwise excellent description, for he is not certain of their

presence on the penultimate pair. They ave rudimentary on this

pair, corresponding to the condition of the crural gland, which

is very small in these: legs (see Pl. I. fig. 3). The papilla is

well developed on the last legs, and its place is marked even

when withdrawn by a minute aperture with raised lips.

It is striking to find that the above description will answer

exactly for the southern form (P. occidentalis). Males have been

obtained with almost all the appendages showing protruded crural

papille. Sections indicate the presence of crural glands and

erural papille exactly as they are found in the Mundaring

specimens. We must emphasise in this connection the necessity

for the external examination of many specimens before one can

state how many crural papille are present. Thus we now have

two varieties (of one species) which are characterised by the

possession of erural papille on the Ist pair of limbs.*

* The above puts out of court Fletcher’s description of P. occidentalis in which

he states that the males haye white papilla on most of the legs, bat not on those of

the \st pair.

a |

ONYCHOPHORA OF WEST AUSTRALIA. 3

INTERNAL ANATOMY.

ALIMENTARY CANAL, ETC.

The alimentary canal presents nothing of exceptional inport-

ance in the way of differences from the conditions observed in

other species of Peripatus. The jaws have already been described.

A few words are necessary with regard to the muscles attached

to them. It is frequently stated that the muscles of the man-

dibles are the only striped muscles in Peripatus. <A careful

examination of all the muscles in the West Australian Peripa-

toides has been made, but so far as transverse striation is con-

cerned no differences can be made out between any of them. No

muscles bear cross-stripes, all appear smooth,

SALIVARY GLANDS.

Two well-developed salivary glands are present, opening by a

common duct ventrally into the mouth. The common median

duct is very short (see Pl. I. fig. 1) and gives rise almost at once

to two narrow tubes which run out at right angles to the long

axis of the body and then bend suddenly backwards when the

lateral body-wall is reached. ‘'[his portion (Pl. I. fig. 1, Sal.G.d.)

is non-glandular, and the walls consist of compact mole cells.

A marked change takes place when the ducts bend abr uptly back-

wards to run in the lateral compartiments in close proximity to

the nerve-cord; the ducts pass here into the glandular region

(Pl. 1. fig. 1, Sal.Gl.) of the salivary gland. The cells of this

part are often much vacuolated, and the nuclei are pushed to the

bases of the cells, where they le in close proximity to a thin

muscle and connective-tissue sheath (see Pl. II. fig. 5).

The length of the salivary glands is such that they extend back

$0 somewhere about the 7th or 8th pair of legs—that is to say,

just beyond the middle of the body.

THe TRACHES.

The respiratory organs of Peripatus have always been regarded

as of special note in view of the interesting relationships of the

Onychophora. It is somewhat surprising, then, to find that even

to-day there is some doubt as to whether a spiral fibre is present:

in the tracheal vessels. Other points are also uncertain.

The respiratory organs, as is well known, consist of trachee.

These are very delicate and of minute diameter. In the West

Australian Peripatoides they could only be made out with difh-

culty in preserved specimens, although more easily in sections.

They are, however, exceedingly clear when freshly killed speci-

mens are dissected under water and examined with a Zeiss

binocular dissecting microscope. Ample material has rendered

this mode of examination possible.

Distribution of Trachee.—As is well known, the trachex of

376 PROF. W. J. DAKIN ON THE

Peripatus arise in bunches from the bottom of little epidermal

pockets which may be termed stigmata or tracheal pits. For a

varying distance the delicate tracheal tubes run in a bundle an

more or less parallel to each other and without branching

gradually, however, the trachex separate off in large or hal

packets from the main bundle and radiate in different directions.

The trachez of these bundles in their turn gradually separate

until they run alone. As a consequence of this arrangement and

the fact that only the larger main bundles are readily visible

even when filled with air, each tracheal pit appears to give rise

to a little irregular rosette or star of trachez. That is to say,

this is the appearance when the inner surface of the body-wall is

examined. The arrangement is indicated in the illustration

(LE WE ies 19),

The tracheal pits are arranged somewhat irregularly, but

mainly in the manner indicated by Balfour in 1883 in Peripatus

capensis. ‘There are two irregular rows dorso-laterally on each

side, that is in the quadrants between the heart (Pl. II.

fig. 6, Ht.) (Qmid-dorsal line) and each longitudinal nerve (Pl. II.

fig. 6, N.C.). The more ventral series on each side appears to

comprise the larger bundles. On the ventral surface there are

also four longitudinal series of tracheal pits, two to each side of

the mid-ventral line (only two series, those to the left of the mid-

ventral line, are shown in fig. 6). The row next to the longi-

tudinal nerve on each side appears to comprise the larger bundles.

Itis difficult to say how many tracheal pits there are to a segment,

for the number appears to vary, and the smallest ones are not

easily seen. No attempt has been made to determine the full

number. Gaftron (8) states that there are about 75 per segment

in Peripatus edwardsi. We have counted over 32 without

trouble in segments of our Peripatoides.

There are some very large tracheal bundles in the head, supply-

ing the large nerve ganglia. Some of the largest of these arise

ventrally, and there seems to be a series of tracheal pits sur-

rounding the mouth-opening. Just behind the mouth there is a

large pit in the mid-ventral line. There are also large pits

ventrally placed and in the median line in front of the mouth.

Other large pits occur to the sides of the oral aperture.

Branching of the Tracheew.—A great deal of doubt has been

expressed as to the course of the traches in Peripatus. Thus,

in Balfour’s treatise on the Anatomy and Development of

P. capensis (1), the following statement occurs: ‘‘ Moseley states

that the trachez branch, but only exceptionally.” Balfour stated

that the tracheze were « extremely minute, unbranched (so fax

as 1 could follow them) tubes.” Sedgwick, in his article in the

Cambridge Natural History (10), states that the trachez “ appear

to branch but only exceptionally.”

Now the main trunks do not branch in the West Australian

Peripatoides, but if a piece of alimentary canal-wall in the

fresh state is mounted in salt solution and examined with

ONYCHOPHORA GF WEST AUSTRALIA. eal

jz oil-immersion lens, branched trachese can easily be found

(see Pl. UI. figs. 7 & 8).

The nerve- call. ganglia, alimentary canal, and in fact all the

organs are well supplied with trachee and branching is easily

discovered. But it is only when an oil-immersion lens is applied

to practically living tissues that the full extent of the tracheal

system becomes apparent. Pl. II. fig. 8 shows more distinctly

the manner of division of the tracheal vessels.

P|. II. fig. 7 is but a very small area of the alimentary canal-

wall indicating the course taken by the branching tracheal vessels

there. Attempts to follow out with certainty the fine termi-

nations of the branched trachee have so far met with little

success. They simply end, but whether the end has been seen

or whether still finer capillary tubes continue and penetrate

cells is uvknown.

The Structure of the Trachee.—Typical insect trachee are

elastic structures lined by an extremely delicate chitin layer

which is strengthened by a spiral fibre. The spiral fibre is said

to be absent from the fine capillary twigs. In large insect

trachee the spiral thickening is easily observed with a moderately

low power of the microscope. The largest trachez of Peripatus

are, however, of minute dimensions, and it is not surprising

therefore that uncertainty should have arisen as to whether they

presented the spiral so characteristic of other tracheate arthropods.

Balfour noticed something and was led to state that the trachez

exhibited a faint transverse striation which he took to be indica-

tive of a spiral fibre. No one seems to have gone beyond this

since, and Sedgwick (10) in 1910 restated it in the description :

““The traches are minute tubes exhibiting a faint transverse

striation which is probably the indication of a spiral fibre.”

Gaffron (8) remarks that it is questionable whether a spiral

fibre exists, the only indication being some fine cross striping seep

only with high powers.

This question has been solved, like certain others, through

the application of the oil-immersion lens to fresh material

mounted in salt solution. There is now no doubt but that the

delicate trachese of Peripatoides are strengthened by an exces-

sively minute but perfect spiral fibre (PI. II. fig. 9).

Tur CruRAL GLANDS.

Reference has already been made to the crural glands in the

section dealing with the external characters.

They ave particularly well developed in the males, where a pair

can be found for every pair of lege. With the exception of the

glands of the first and last pairs of legs, which are highly modified

in both the West Australian forms, the crural glands are entirely

contained in the legs. The external aperture is very distinct.

It is situated distally to the nephridial aperture on the ventral

surface of all the legs except the 4th and 5th, where the excretory

opening is found near the end of the appendage. The extremity

378 PROF. W. J. DAKIN ON THE

of the gland-duct which opens to the exterior is provided with

a swollen circular lip which forms the crural papilla. The

surrounding epidermis is retvactile and may be invaginated

to form a little Crural Pit. The crural papilla is then quite

invisible from the exterior, but this is merely a temporary con-

dition, and consequently there is no point in counting visible

crural papille in these animals unless the count is checked by

sections. With the exception of the last pair of glands, opening

on the last pair of appendages, the aperture of the gland leads to

a narrow duct bounded by small eubical cells. This duct runs

upwards to open into a large vesicle which extends distally and

occupies quite a large extent of the leg-cavity. The vesicle itself,

t.e., the crural gland proper (see Pl. IV. fig. 10) is lined by a

layer of small and.compact cubical epithelial cells with large nuclei

centrally placed. The glands are usually readily distimguished

by reason of the contents, which in sections stained with heema-

toxylin and eosin appear bright pink, Still more characteristic

is the fact that the contents are perfectly homogeneous and

non-granular.

The first pair of Crural Glands differs considerably from all the

rest. The duct opens in the same manner as in the succeeding

glands but instead of leading to a sac in the leg it passes into the

lateral cavity of the hody and opens into an elongated sac which

runs almost the entire length of the animal (see Piaeetioas,

Cr.Gl.'). This tubular gland can be easily picked out in transverse

sections, for it is always cut transversely and lies not far from

the lateral nerve-cord and below the salivary gland in sections

where this is also present. The structure of its wall is quite

characteristic.

These extraordinary crural glands of the first legs were dis-

covered in the West Australian LPeripatoides from the hills near

Perth by Bouvier. They are noted in his monograph as peculiar

to the species. Jt is important, therefore, to note that the same

feature is present in the southern variety—in short, this character

is peculiar to both West Australian varieties of Peripatoides.

The Crural Glands of the penultimate legs are extremely

minute. The duct is short and leads into a very small vesicle.

Bouvier was uncertain as to whether crural glands existed here

at all. He was unable to find them in his specimens.

The Crural Glands of the last pair of legs are very different

from all the others. The ducts pass direct from the legs into the

central body-cavity, where they lead into two somewhat wide

tubes which run forwards entangled in the coils of the gonoducts

(seeue] Mieitic rouC r.Gilli):

Crural glands and crural papillee are features of the male sex,

but in Willey’s account of the Anatomy and Development of

Peripatus nove-britannie (13) the following refevence occurs

to crural glands in the female: ‘‘ Wherever they occur (crural

glands) they are found only in the male except in P. capensis,

where they are said to occur in the female also (Sheldon).

ONYCTIOPHORA OF WEST AUSTRALIA, 379

Without denying their occasional existence in the female

P. capensis, | may say that I have failed to find them present,

and I doubt, on @ priort grounds, if they normally occur in the

female.”

In the paper (11) to which Willey refers the following state-

ment is made: ‘‘I have examined several legs of P. capensis,

both of males and females, and have found a crural gland in

every one except the first pair of legs.”

A careful search through sections of female Peripatoides of

Western Australia has shown that crural glands do oceasionally

occur in the female. They are not always present, however,:and

when found there is no regularity as to the legs containing them.

In any case they are not highly developed, although they have

the same vesicle with the same pink-staining homogeneous con-

tents found in the male.

Tur NeEpPHRIDIA.

A very careful study of the nephbridia of Western Australian

Peripatoides has been made, and well-preserved sections have eluci-

dated many points. It will be perhaps desirable to describe their

structure in detail, especially since some features, the presence of

cilia for example, ave now made known for the first time. Mention

of cilia occurs in all general descriptions of Peripatus, but only

to the effect that they are found in the generative ducts. The

cilia discovered in the nephridia of Peripatoides ave remarkably

well developed and of great length, reminding ore more of flagella.

The nephridia, with the exception of those of the 4th and 5th

pairs of legs, open on the ventral surface at the junction of

each leg and the body. Those of the 4th and 5th pairs of legs

are quite different from the others. They open on a special

papilla situated on the ventral surface of the respective legs (see

Pl. I. fig. 2, Neph.’). It is rather striking that with variations

in the anatomy of the Onychophora, and im particular with

considerable variations in the number of legs, there should be such

a constancy in regard to the position of these special nephridia.

They are of almost exactly the same form in the West Australian

Peripatoides as in Peripatws capensis, and similar enlarged

nephridia occur in other species.

Nephridia are found in all the pairs of legs without exception

in the West Australian Peripatoides.

Structure of a typical Nephridiwm.—Vhe aperture, which, except

in the case of the 4th and 5th nephridia, is unmarked by any

papilla, appears as a little crevice in the epidermis. This leads

into a short tube of minute diameter. This duct passes upwards

into the lateral compartment of the body and opens into a thin-

walled collecting vesicle (Pl. IV. fig. 11, Coll. Ves.). The cells

of the duct are small, cubical, or somewhat flattened, but the

vesicle is lined by a very delicate squamous epithelium of large

cells, the nuclei appearing distinctly some distance apart. The

vesicle might easily be mistaken for a split in sections were it not

380 PROF. W. J. DAKIN ON THE

for its constant presence and the ducts opening intoit. Following

the vesicle the nephridium is divisible into two marked sections,

a tubular portion and a terminal chamber. The tube describes a

rough circle and then turns abruptly on itself, so that its termin-

ation is close to the vesicle referred to above (see PI. LV. fig. 11).

The terminal chamber (PI. LV. fig. 11, Cee.) is now well known,

although missed by the first investigators, who believed that the

nephridia opened into the lateral compartment of the body.

The chamber lies partly above and partly posterior to the col-

lecting vesicie referred to above.

The first portion ef the nephridial tube internal to the

collecting vesicle is lined by a very delicate and characteristic

epithelium of large flat cells. As a consequence of the size of

the cells relative to the diameter of the duct it is possible to have

transverse sections with only two or three uuclei showing (see

Pl. LV. fig. 11). The greater part of the nephridium between the

terminal chamber and the collecting vesicle is built in this way.

The section, however, which actually opens into the terminal

chamber is very different. The wall of the nephridium becomes

thicker and far less delicate and is formed of a compact layer of

columnar epithelial cells (see Pl. IV. fig. 11, Cil.h.). These

cells are so crowded and the nuclei stain so distinctly that most

previous workers have noted the peculiarity. In fact this change

in the character of the cells has been taken as indicating the

passage from the ectodermal part of the nephiidium to the

mesodermal portion (see Glen, Q. J. M.S. 1918, vol. bxiii.).

Now it is the cells of this section of the nephridium which bear

the cilia (Pls. LII.-IV. figs. 11 & 12). These are so long that

after projecting from the cell they extend along the lumen of the

duet for a relatively considerable distance. It is extraordinary

that in many figures showing the structure of the nephridia of

Peripatus details of the histology are given at a high magnification,

yet no indication of cilia is presented.

Bundles of long cilia are very characteristic of renal cells,

although at the same time they are in the highest degree peculiar

for the arthropoda. The Annelid resemblances of Peripatus

are certainly heightened as a result of the examination of well-

preserved sections through the ciliated ducts of these nephridia*.

* Since writing the above I have been enabled to examine a copy of Gaffron’s

famous paper (8) on the Anatomy and Histology of Peripatus, in which the first

mention of the presence of cilia in this animal—in the Receptacula seminis—was

made. Looking through his description of the nephridia I found to my surprise

the following lines referring to the region where the duct opens into the coelomic

yesicle. It must be remembered that the vesicle was unknown at the time, and its

remains were supposed to be a funnel-like nephrostome opening into the body-cavity.

“By hesitzt wie der Trichter selbst, kleinzelliges, im Leben wahrscheinlich wim-

perndes Hpithel .... Gattron never indicates that he found cilia here nor are

any shown in his illustrations of this region. We must conclude that the remark

was merely a conjecture, probably suggested by the apparent resemblance to the

open nephrostome of an annelid. It is curious, however, that his successors who

have studied the nephridia have not commented on this. Hither the cilia are only

found in the West Australian Peripatoides or else my preparations must be par-

ticularly favourable ones.

ONYCHOPHORA OF WEST AUSTRALIA. 381

The Nephridia of the 4th and 5th pairs of legs (Pl. 1. fig. 2,

Neph.’). These nephridia differ from the others firstly im the

increased length of the tube between the distal collecting vesicle

and the opening on the leg. This is due to the fact that the

renal aperture is situated near the extremity of the appendage.

The vesicle presents the same structure as before. From the col-

lecting vesicle a long, tubular portion extends posteriorly within

the lateral compartment of the body. This section presents the

same type of wall as the corresponding section of the other

nephridia. Having reached somewhere about the second suc-

ceeding pair of legs the tube turns on itself and runs forward,

the two limbs being in close contact. The wall still presents the

large flattened cells. This section passes into the ciliated duct,

which is particularly well marked in these nephridia, and opens

into the ewlomice chamber, which lies close to the distal collecting

vesicle.

Tur REPRODUCTIVE ORGANS.

A somewhat detailed description of the reproductive organs of

the West Australian Peripatoides is rendered necessary owing to

the fact that Bouvier’s specimens of the northern variety were

not sufficient to allow of a complete and accurate account of the

anatomy. ‘his applies in particular to the female, in which con-

nection Bouvier states (3) ‘‘ L’appareil génital femelle ne présente

rien de particulier, si ce n'est Patrophie complete, ow a pew pres

complete, des receptacles séminaux.” The receptaculum seminis

is, however, weil developed. The explanation of the mistake

probably lies entirely in the state of preservation of the few

specimens available. We have had the good fortune to obtain

many specimens and to preserve them in many ways.

The Male Reproductive Organs consist fivst of the two testes,

which lie fairly far forwards entangled amongst the diverticula of

the slime glands (see Pl. I. fig. 3, Tes.) and dorsal or lateral to

the alimentary canal. These organsare tubular and from the wall

cells are cut off which are apparently the spermatocytes. These

do not develop into spermatozoa in the testes. Apparently the

development of spermatozoa takes place in the vesicula senvinalis

(see Photomicrograph, Pl. III. fig. 14).

This feature is to my mind rather interesting; yet it is one

which is never mentioned in text-books. It was naturally

thought at first to be quite a new discovery. As a matter of fact

it was seen by Gaffron many years ago and figured in his work (8).

Gaffron was struck also by the resemblance to the conditions in

the earthworm and actually writes: “ Etwas ehnliches findet

sich bekanntlich beim Regenwurm, wo ja auch die eigentlichen

Hoden den ‘Samenblasen’ gegenuber sehr zurucktreten und die

Weiterentwicklung der Spermatozoen in letzteren stattfindet.”

Each testis opens into a seminal vesicle (PI. I. fig. 3, 8.V.),

one usually lying somewhat in front of the other owing to the

382 PROF. W. J. DAKIN ON THE

crowded condition of the body-cavity. It is difficult to separate

the delicate ducts in preserved and consequently hardened spe-

cimens. ‘The seminal vesicle may, however, attain such a size

at a certain season of the year (September—October) as to fill up

most of the body-cavity where it occurs. This is well shown in

the illustration Pl. III. fig. 4, which is a photomicrograph of a

transverse section passing ‘through such a seminal vesicle. ‘The

wall of the seminal vesicle is rather delicate, being formed of

somewhat flattened epithelial cells (Pl. IV. fie. ier The vas

deferens leaves the seminal vesicle at the opposite side from the

entrance of the testis or testis duct. (It is impossible to divide

that portion of the reproductive organs beyond the seminal vesicle

into regions.)

The first portion of the vas deferens following the seminal

vesicle is lined by almost cubical cells ae LV. figs. 15, 15).

This leads imperceptibly into a section (by far the longest) the

wall of which is of flattened cells, the epithelial inyer being, how-

ever, supported by a layer of longitudinal and circular Tnusele

fibres. ‘This section of the vas deferens coils about, entangled

with its fellow of the opposite side and also with the terminal

portions of the accessory reproductive glands.

What we may term the vas deferens of the right side con-

tinues its course to the left of the alimentary canal right away

on towards the posterior extremity of the animal (see PI. I.

fig. 3, Vas.def.r.). Just in front of the male opening it dives

under everything, even the two nerve-cords, and, reaching the

right side, it continues its way anteriorly again.

‘This extraor dinary difference from the course of the other vas

deferens (Pl. I. fig. 3, Vas.def.l.) is always met with. It was

indicated by Bouvier, but his dvawing is not quite accurate.

Eventually, somewhere about the 4th or 5th leg from the pos-

terior extremity, both vasa deferentia join up to form a very wide

terminal unpaired tube. The first portion of this wide duct has

thin walls, the epithelium consisting of flattened cells (Pl. IV.

fig. 16). Asthe reproductive apeniane is reached the walls become

thicker (Pl. V. fig. 17) owing to a gradual development of trans-

verse and longitudinal muscles. The terminal portion (rl. V.

fig. 18) is very muscular, the walls being quite thick. It is

probably extrusible. This last section turns over to the right

side of the alimentary canal and passing underneath the right

nerve-cord reaches its opening to the exterior (see PI. I. fig. 3).

As Bouvier pointed out, the vast chamber formed be the

commencement of the unpaired duct is usually filled with a mass

of spermatozoa, the whole taking the form of a convoluted cord

(Pl. LV. fig. 16, Sp.). There is no sign of any chitinous envelope.

It will be seen that the unpaired section of the reproductive

duets formed by the union of the two vasa deferentia is the region

for the storage of spermatozoa and thei massing into sper mato-

phores—not “the so-called seminal vesicle, where | spermatogenesis

takes place.

ONYCHOPHORA OF WEST AUSTRALIA, 383

From the fact that sperms are found throughout the ducts of

the female I have no doubt that females are impregnated thr ough

the vaginal aperture.

Accessory Ducts, ete.—The crural glands of the last legs of the

male are modified as already pointed out. The minute duct,

which opens in the usual place, runs into the central. body-cavity

and enlarges to form a thin-walled vesicle which runs forwards

as far as the antepenultimate leg or thereabouts. It walls and

contents (see Pl. LY. fig. 10) are similar to those of the normal

crural glands of the anterior limbs.

Two other aécessory glands, the anal glands, open in close

proximity to the male reproductive opening, and slightly posterior

to it. From each opening a narrow duct passes laterally under

the nerve-cord and then turns dorsally and inwards and for-

wards, gradually widening until a rather wide sac is produced

(Bly letiegs), Am) Gie):

These glandular sacs are rather prominent in transverse

sections near the posterior end of the animal (Pl. V. figs. 19

and 20, An.Gl.), and are easily picked out by reason of the

intensely vacuolated cells which form their walls. The contents,

too, are very granular and deeply staining (see Pl. V. fig. 21).

The Reproductive Organs of the Female-—The ovaries of the

West Australian Peripatoides agree with those of the other

Australian forms and with the Cape and New Britain species in

having thin walls, so that the developing eggs come to Hea

freely) in the central division of the body-cavity (Pl. I. fig

Ov.). From each organ an oviduct (Ovid., Pl. V. tig. 22) mor

forwards for a short distance to open into a large and fully

_ developed receptaculum seminis by two ducts (Pls. I.,V. figs. 2, 22,

Rec.Sem.). It is true that the receptaculum may reach a much

larger size in September—November, but it is none the less a

well-defined permanent structure at all times.

The oviducts, which open into the two ovaries, unite for an

extremely Shane distance at their origin and then separate again

(Pl. V. fig. 22). Their course is ‘eu amidst the acinaol vinnie

of the slime-clands for a short distance to the point where each

communicates with a receptaculum seminis. This first seetion of

the oviduct is characterised by somewhat thick walls, the epi-

thelial cells are deep and crowded together, the lumen of the duct

usually appears restricted (see PI. We fig. 23),

As Willey noted in the case of Peripatus novee-britannice, the

walls of this first section of the oviduct differ from these of

the rest of the genital duct. Willey (13) termed this portion the

infundibulum, and remarked that the striking contrast between

the infundibula and the ovarian tubes seen in P. nove-britananie

has not been remarked in other species.

The infundibula of the West Australian Peripatoides do not

differ materially from those of P. nove-britanniw. So far,

384 PROF, W. J. DAKIN ON THE

however, as the ovarian tubes are concerned, I have been able to

distinguish peritoneal investment, tunica muscularis, and germinal

epithelium, and, the thickness of the wall increases as we pass to

the infundibulum (Pl. V. fig. 24). I should be inclined, on the

whole, to doubt the possibility of the ovarian tubes not being

strictly homologous structures throughout the genus—a suggestion

of Willey’s

The receptacula seminis (Pl. V. fig. 22, Rec.Sem.) are two

thin-walled bags, each of which communicates with the infun-

dibulun of its le by two short ducts. The same condition is

met with in other species of Peripatus where the receptaculum

is present.

The preparations which I have at my disposal show that cilia

are present in the tubes connecting the infundibulum with the

receptaculum (see Pl. V. fig. 25). This is the position in which

cilia were first discovered in Peripatus by Gaffron (8). It must

be noted that these cilia are much shorter and less distinct than

those described earlier in this paper as occurring in the nephridia.

The two uteri do not differ essentially in histological structure

from the infundibula. They possess a well-developed muscu-

lature — transverse and longitudinal fibres surrounding the

epithelial wall. These uteri pass forwards for a short distance »

and then return, usually one to each side of the alimentary canal

(Pl. I. fig. 2, Ut.). They meet posteriorly quite near the ex-

texnal aperture and a median and very short vagina (PI. I.

fig. 2, Vag.) leads to the exterior. During the summer months

each uterus presents the appearance of a string of sausages.

This is due to the chain of developing eggs or embryos con-

tained within it. The eggs are of very large 1 size indeed, and the

uterus is swollen considerably round each and constricted between

them. ‘The West Australian examples of Peripatoides bring their

reproductive organs to maturity during the winter, which is the

only period of feeding and activity in general. Fecundation

probably takes place about August to October. The species is

viviparous. . [For further reference see notes on Spermatogenesis

and "Reprod uction in a following paper. |

SuMMARY OF CHARAOTERS DIAGNOSTIC OF THE West AUSTRALIAN

PERIPATOIDES.

There are two West Australian varieties of Peripatus—sub-

gen. Peripatoides. ‘They agree in external characters and in

anatomy, with the exception that whilst the northern form

possesses constantly 16 pairs of legs, the southern variety has only

15 pairs. It is not considered advisable to separate these two

forms as distinct species. ‘They can only be considered varieties.

The first to be discovered and named was the southern variety,

which was termed Peripatus leucharta Sing., var, occidentalis, by

Fletcher in 1895.

Since that date, however, this western form has been raised to

ONYCHOPHORA OF WEST AUSTRALIA. 385

specific rank by Dendy (5), and is now recorded as Peripatoides

occidentalis. Thevediscovery of the typical form and the demon-

stration that its anatomy is almost identical with that of the other

form which Bouvier recognised as quite distinct from all other

Australian Peripatoides endorses Dendy’s action. The northern

variety was termed P. woodwardi by Bouvier, but we have

shown (4) that this name lacks priority, the name P. gilesii

having been previously given by Baldwin Spencer. The position

is, therefore, that the only West Australian species of Peripatoides

is Peripatoides occidentalis, the southern form being the original

one named, the northern is thus to be known in future as

P. occidentalis var, gilesit.

Previous diagnoses of P. occidentalis are to a large extent

incorrect.

The Diagnosis of the Species is as follows :—

(1) Specimens fall into two colour series, in one of which dark

ereen-black predominates, in the other a brown-red. (2) The

legs number 16 pairs in the var. gilesii, 15 in the typical form.

(3) The third pedal ring (or spinous pad) is usually slightly

narrower than the first, and the intermediate ring a little larger

than in the other Peripatoides; the rudiments of the 4th ring

are practically invisible. (4) There are no accessory teeth on the

outer blade of the mandible, and 5 or 5 on the inner blade.

(5) The urinary papilla of the 4th and 5th legs are each on a

separate segment of the first pedal ring. (6) The crural papillee

are present on all the legs of the male, but may not be obvious

owing to retraction. (7) Crural glands are present opening on

all the legs; those of the Ist pair are very long and extend

almost the entire length of the body in the lateral compartments.

Those of the last pair of legs are also long and run forwards

entangled with the reproductive ducts in the central body-cavity.

All the other crural glands are contained in the legs. Those of

the pair of legs preceding the last are very small. (8) The male

reproductive organs may extend forwards as far as the 6th or

7th pair of legs from the posterior extremity. The two vasa

deferentia after leaving the vesicule seminales in which sperma-

togenesis takes place run a tangled course. That of the right

side runs almost to the extreme posterior end and to the left side

of the body, it then curves under both nerve-cords and runs for-

ward on the right side to meet its fellow and form an unpaired

duct at about the level of the 4th or 5th pair of legs from the

posterior end. The proximal portion of this duct is very wide

and very thin-walled and forms a reservoir where spermatozoa

accumulate in tangled “cordons.” It passes gradually into a

short muscular ejaculatory duct. (9) The ovaries are dorsal in

position and extend forwards from near the posterior end,

oceupying the hinder third of the body at the breeding-season.

. The walls of the ovaries are thin, and the eggs when ripe appear

386 PROF. W. J. DAKIN ON THE

in consequence to le freely in the central cavity of the body—

in reality the ovary-wall projects in the form of very delicate

follicles. (10) Receptacula seminis are present, each communi-

eating with the oviduct of its side by two ducts which are

ciliated.

AFFINITIES OF THE WEST AUSTRALIAN

PHERIPATOIDES.

Bouvier discusses at some length the affinities of his Peripatoides

woodwardi. His account requires bringing up to date owing to

the corrections necessary by reason, first, of the increased know-

ledge of this northern variety, and, second, owing to the infor-

mation now brought forward regarding the anatomy of the

southern form known to Bouvier as a distinct species. According

to Bouvier the West Australian Peripatus is remarkable for its

multiple affinities. It 1s supposed to resemble P. sutera of New

Zealand by the presence of 16 pairs of legs and by the absence of

an accessory tooth on the outer blade of the mandible. At the

same time it is distinguished from this species by most other

characters and resembles P. leuckartii* in the following :-—

1, alternation of tegumentary folds; 2, reduction of the pedal

papille to three; 3, the relative dimensions of the rings of the

soles; 4, the multiplicity of the crural glands; 5, the analogy of

the anal glands; 6, a certain resemblance in the unpaired portion

of the male gonoduct.

It is, however, different from all other species in the possession

of crural glands on the first legs—glands of enormous length.

It is supposed to be unique in the constant presence of 16 pairs

of legs with 3 papille. Its unpaired male duct is supposed to

be really like no others. In short, it is supposed to present a

mixture of primitive characters with others indicating a long

evolution. These may be classified as follows :—

Primitive Characters. Advanced Characters.

1. Crural glands opening on 1. No teeth on outer blade of

each leg. mandible.

9. 16 pairs of legs. 2. Pedal papillae reduced to

three.

3. Seminal receptacles atro-

phied.

4, Character of male gono-

duct.

I do not consider that we can lay much stress upon affinities

which are only indicated by the presence of 16 versus 15 pairs of

* P. leuckartii is taken as the correct name for the Ges PonuEoe of the

7 2 “ @ ape ni sae A :

East, usually designated P. orientalis by Bouvier. For a discussion on the nomen-

clature of this species see Dendy (Q. J. M.S. vol. xlv, p. 388, and Zool. Anz. 1906,

pp. 175-177).

ONYCHOPHORA OF WEST AUSTRALIA. 387

legs or the presence or absence of an accessory tooth on the outer

mandibie-blade. Surely some such change as this could have

occurred as a mutation over and over again. In any case the

West Australian species occurs in two forms, one with 16 and

one with 15 pairs of legs, so that the resemblance to P. suteri

does not hold good.

As a matter of fact Peripatoides occidentalis approaches most.

closely Peripatoides leuckurtii, and in addition to the resem-

' blances noted by Bouvier we may add that receptacula seminales

are present in both forms. The species is, however, very distinct

from all other Peripatus species, and thus from all the other

Australian species (which is not surprising seeing that the two:

Peripatus regions are separated by over two thousand miles, the

greater part of which is country quite uninhabitable by Peripatus)

in the presence of extraordinarily long crural glands opening on

the first pair of legs. The northern variety is the only known

Peripatus in Australia having 16 pairs of legs.

SUMMARY OF RESULTS OF GENERAL IMPORTANCE.

(i. €. probably applicable to most if not all species of

Onychophora.)

I. The cells of a certain part of the so-called nephridium—

that which opens into the terminal ccelomic vesicle—bear

long and well-developed cilia. Thus cilia occur in the

excretory ducts of Peripatus as well as in the repro-

ductive organs.

Ii. Crural glands do sometimes occur in the female, but do

not seem to possess any ducts,

III. The tracheze of Peripatus possess a characteristic spiral

supporting fibre.

This has been a disputed question for many years.)

IV. The trachee of Peripatus, although running a separate

course for some distance from the tracheal pit, eventually

branch (see Pl. IT. figs. 7 & 8).

V. Spermatogenesis does not take place in the testis but in

the seminal vesicles. This discovery, made originally

by Gaffron, seems to have been lost sight of in most.

descriptions of Peripatus,

LITERATURE CITED IN TEXT.

1. Batrour.—Anatomy and Development of Peripatus capensis.

Q. J. M.S. 1883.

2. Bouvrer.— Monographie des Onychophores. Ann. des Sci.

Nat.(9) 2&5. Paris, 1905 & 1907.

3. Bouvrer.—Onychophora. Die Fauna Sid-West Australiens,

Bd. ii. Lief. 18. Jena, 1909.

Proc. Zoou. Soc.—1920, No. XX VI. 26

388

00 NID

10.

Lie

12.

13.

Fig.

PROF. W. J. DAKIN ON THE

Daxtn.—The Onychophora of West Australia. Proc. Zool.

Soe. London, 1914.

Denpy.—On the Oviparous Species of Onychophora.

Q. J. M.S. vol. xlv. p. 388.

Denpy.—Zool. Anz. Bd. xxx. Nr. 6, 1906, pp. 175-177.

FiLercuerR.—On the Specific Identity of the Australian

Peripatus, ete. Proc. Linn. Soc. N.S. W. 1895, vol. x.

Garrron.—Zoolog. Beitriige (Schneider). Breslau, 1883.

SEDGWICK.— Various papers on Peripatus. Reprinted in

vol. iv. Studies from the Morphological Laboratory of the

University of Cambridge. 1889.

SepDGwick.— Article on Onychophora in Cambridge Natural

History.

SueLpon.—Anatomy of Peripatus capensis, etc. Q.J.M.S.

vol. xxviul. 1888.

BaLpwin Spencer.—Proc. Roy. Soc. Victoria, vol. xxi. (N.s.)

pt. 2, 1909.

Wixitey.—Anatomy and Development of P. nove-britannice.

Zoological Results. Part I. 1898.

DESCRIPTION OF FIGURES.

Puates 1.—V.

Peripatoides occidentalis.

Dissection, showing slime gland, alimentary canal, and salivary gland.

x dg.

Dissection, female, showing reproductive organs and nephridia. X 53.

Dissection, male, showing reproductive organs and crural glands. X 53.

Photomicrograph. ‘Transverse section, showing large seminal vesicle

lying above alimentary canal and erural papilla retracted.

Longitudinal section. Salivary gland. X 400.

Inner face of body-wall in freshly dissected specimen, showing arrange-

ment of main trachese. The piece extends across the two vential

nerve-cords, and laterally up one side and beyond the mid-dorsal line.

The positions of three legs are shown (N). X 20.

7. Alimentary canal-wall under low power, showing trachee branching.

x 96:

8. Tracheal branching. XX 640.

9. Tracheal tube, showing spiral thickening. X 800.

10. Section through crural gland. X 160.

11. Nephridium in longitudinal section, showing cilia. XX 240.

12. Photomicrograph of ciliated part of nephridium.

13. Opening of vas deferens into vesicula seminalis, section to show structure

of wall. >< 200.

14. Photomicrograph of section through vesicula seminalis, showing sper-

matozoa and spermatogenesis.

15. ‘Transverse section. First part of vas deferens. >< 200.

16. Median part of vas deferens, with mass of spermatozoa. X 200.

17. Terminal thick-walled part of vas deferens. X 200.

18. Longitudinal section. Ductus ejaculatorius. > 200.

19, Diagrammatic transverse section in plane of last pair of legs. X 24.

DX Bw

20. Diagrammatic transverse section in plane where vas deferens crosses below -

nerve-cords posteriorly. X 24.

21. T.S. Part of wall of anal gland of male. X 240.

22. Diagram showing connections of receptacula seminis with oviducts, and

_ ovaries. X 200.

23. 'T.S. Infundibular region of oviduct. X 200.

24. L.S. Wall of ovary and oviduct. X 240.

25. T.S. Duct of receptaculum seminis. X 280.

An.Gl.

‘Ci. R.

Coe.

Coll. Ves.

Cr.Gl.

CrGl/

Cr.G1.””

Crur.pap.

ONYCHOPHORA OF WEST AUSTRALIA.

389

EXPLANATION OF LETTERING.

Anus.

Anal gland.

Ciliated portion of nephri-

dium.

Ccelomic cavity.

Collecting vesicle.

Crural gland.

Crural gland of Ist leg.

Crural gland of last leg.

Crural papilla.

Cuticle.

Epidermis.

Heart.

Position of leg.

Nerve cord.

Nephridium.

Ditto of 4th and 5th legs.

Or. Pap.

Oy.

Ovid.

Ph.

Rec.Sem.

Rect.

Oral papilla.

Ovary.

Oviduct.

Pharynx.

Receptaculum seminis.

Rectum.

Salivary gland.

Salivary gland duct.

Seminal vesicle.

Slime gland.

Spermatozoa.

Testis.

Uterus.

Vagina.

Vas deferens.

. Vas deferens of right side.

Vas.def.1.

Ditto of lett side.

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ON MARINE WOOD-BORING ANIMALS. 391

Notes on Marine Wood-boring Animals.—I. The Ship-

worms (Teredinide). By W.'T. Cauman, D.Sc.

(Submitted for Publication by permission of the Trustees of the British Museum.)

[Received April 27, 1920: Read June 1, 1920.]

(Text-figures 1-11.)

The specimens discussed in this paper were collected, for the

most part, on behalf of a Committee appointed by the Institution

of Civil Engineers to inquire into the deterioration of structures

exposed to sea-action. This Committee, at my suggestion, re-

quested its correspondents at various seaports to send in speci-

mens of animals damaging the timber of harbour works. The

result has been to get together a collection of very considerable

importance, both from the point of view of the practical engineer

and from that of the scientific zoologist. A set of the specimens

will be placed in the Museum of the Institution of Civil

Engineers, and the remainder have been presented by the

Committee to the British Museum (Natural History). I desire

to express my sense of obligation to the members of the Com-

mittee, more especially to the Chairman, Sir William Matthews,

K.C.M.G., and the Secretary, Mr. P. M. Crosthwaite, as well as

to the various harbour engineers named below, by whom the

specimens were collected and preserved.

In dealing with the Teredinide I trespass with reluctance on

the domain of the malacologists. It is only the impossibility of

finding a student of Mollusca ready to undertake the description

of the collection that leads me to publish these notes, to which,

however, the very accurate figures drawn by Miss G. M. Wood-

ward may give some permanent value. I am indebted to my

colleagues, Mr. B. B. Woodwazd and Mr. G. C. Robson, for much

guidance Avil help in exploring the literature and in examining

the Museum collections of Mollusca.

Many writers have commented on the difficulties that stand in

the way of a systematic study of the Teredinide. The lack of

agreement as to the characters to be regarded as generic is

strikingly shown in the recent synonymy of several species, while

the inconstancy of specific characters drawn from the valves of

the shell was commented on long ago by Forbes and. Hanley

(Hist. Brit. Mollusca, i. p. 87 (1848)). These dithiculties I cannot

pretend to have solved, but some general considerations suggested

by study of my mater ial may be worth recording.

The changes in form of the shell-valves during growth seem to

have received little attention, although several writers mention

the obvious fact that the number of the striz on the anterior

and antero-median divisions of the valves increases with age.

Together with this, however, there goes on a resorption of the

392 DR. W. T. CALMAN ON

posterior margin, beginning on the dorsal side, just behind the

hinge-knob, and extending downwards. The rapidity and extent

of this erosion appears to differ in different species. Incon-

spicuous, as a rule, in 7’. navalis*, it becomes very marked in

certain tropical species. For example, in many specimens of

Teredo mannii, mentioned below, the auricle and nearly the whole

of the postero-median area have been removed, while the antero-

median (vertically striated) area occupies the greater part of the

surface of the valve. It may be suggested as a possibility that

the absence of extensive erosion in most specimens of 7. navalis:

is due to the fact that this is a short-lived and indeed almost an

annual species, the individuals rarely surviving the winter, while

the much larger 7’. mannii may be longer lived, the individuals

perhaps surviving for several years in the warmer waters which

it inhabits. From the practical point of view it would be very

important to ascertain the duration of life and the rate of growth

in the different species.

Genus TEREDO Linn.

Hedley (Proc. Linn. Soc. N. 8S. Wales, xxiii. 1898, p. 92}

regarded the presence of a ‘‘cup-shaped mantle which ......

surrounds the bases of siphons and palettes” as the chief

distinctive character of a genus to which he applied at first the

name Calobates of Gould, and later (Mem. Austral. Mus. Sydney,

iii. 1899, p..508) Nausitoria (i. e. Nausitora) of Wright. Hedley

states that the type of the genus Teredo, “according to the

figures of Forbes and Hanley and other writers,” entirely lacks

this structure. ‘The accompanying figure (text-fig. 1, A) is taken

from a well-preserved specimen from the estuary of the Thames,

for which I am indebted to Dr. W. M. Willoughby, Medical

Officer of Health for the Port of London. This specimen appears

to be referable, without doubt, to the typical 7. navalis Linn.

It will be seen that the base of pallets and siphons is surrounded

by a fleshy collar or fold of the mantle, entirely similar to that

found in Zeredo mannii and various other species which Hedley

refers to Vausitora or Calobates.

TEREDO NAVALIS Linn. (Text-fig. 1.)

Teredo navalis Linneus, Syst. Nat. ed. x. 1758, p.651; Forbes

and Hanley, Hist. Brit. Moll. i. 1848, p. 74, pl. 1. figs. 7, 8,

pl. xviii. figs. 3,4; Gatliff and Gabriel, Proc. R. Soc. Victoria,

EXViil. (n.s.) 1915, p. 117.

? Teredo pedicellata Quatrefages, Gwyn Jeffreys, Brit. Conch.

ili. 1865, p. 174, and v. 1869, pl. liv. fig. 3.

* Gwyn Jeffreys’ description of the “var. divaricata” of T. norvagica, the

“var. occlusa”’ ot T. navalis, and the analogous varieties of other species, as well as

the specimens named by him in the Norman collection, suggest that these varieties.

are based on unusually old specimens, in which the antero-median area occupies a

much larger portion than usual of the surface of the valves while the auricle has

been almost completely removed.

MARINE WOOD-BORING ANIMALS. 393

Locality.—Simon’s Town, South Africa. Specirnens forwarded

by Lieut. L. H. A. Shadwell, R.N.V.R., Officer in charge of

Works, H.M. Dockyard. From Blue Gum timber, Ordnance

Jetty, E yard; from Pitch Pine, Old Ordnance Jetty, E yard ;

and from creosoted Danzig, A, No. 3 slip.

Remarks.—Teredo navalis and T'. pedicellata have both been

recorded by Gatliff and Gabriel from Victoria, but, so far as I

know, neither has been recorded from South Africa.

Some of our South African specimens agree very closely, as

regards the structure of the pallets, with specimens from

Alderney in the Norman collection determined, apparently by

Gwyn Jeffreys, as 7’. pedicellata. Jeffreys himself remarks that

Text-figure 1.

Teredo navalis Linn.

A. Siphons and associated structures in a specimen from the estuary of the

Thames at Gravesend. ‘The fleshy collar at the base of the siphons has

peen divided and reflected, showing the imsertion of the right pallet from

the inner side. 3B. Pallet of a specimen from Simon’s Town. Actual

length about 5 mm.

“this is not a satisfactory species,” and the identification of his

specimens with the form described by Quatrefages appears to

be largely conjectural. Quatrefages’s description (Ann. Sci. Nat.

Zool. ser. 3, xi. 1849, p. 26) contains little to suggest it except

the statement that the pallets are coloured dark brown. In the

Alderney specimens and in some of those from South Africa

(text-fig. 1, B). the stalk of the pallets may be as long and less

than one-fourth as wide as the blade. The blade is flattened on

the inner and convex on the outer surface, with the basal half

calcified, white, and nearly opaque. The distal half is mostly

394 DR. W. 1. CALMAN ON

composed of a more or less translucent horny material varying in

colour from yellowish to dark brown, within which the distal ead

of the calcified part is seen to project as a rounded cone. On the

outer surface, the central area of this horny part is occupied by

a calcified plate, rectangular or irregularly rounded in outline,

which reaches the distal but not the lateral margins. ‘The distal

end of the pallet is concave or notched, with a more or less deep

central conical pit.

The structure described above is most easily seen in the

smaller specimens from South Africa, measuring about 40 mm.

in length of body and having the pallets about 5 mm. long. The

Alderney specimens are a good deal smaller. In some South

African specimens of about the same size, however, the distal

calcified plate cannot be seen, and in some larger specimens the

basal calcification, instead of penetrating into the interior of the

horny part, extends up along its lateral margins. In the largest

specimen of all, in which the body is a foot long, the pallets are

wholly calcified, with the distal portion slightly yellowish but

not horny. This large specimen does not appear to differ in any

but the most trivial details from Huropean specimens referred

to 7. navalis. The valves of the shell afford no distinctive

characters.

In the absenee of any more satisfactory definition of the

oe species 7’, pedicellata, all our South African specimens

may be referred to 7. navalis.

TEREDO NORVAGICA Spengler.

Teredo norvagicus Spengler, Skriv. Nat. Selsk. Kigbenhavn, il.

H. 1, 1792, p. 102, pl. i. figs. 4-6, B; Forbes and Hanley, Hist.

Brit. Moll. i. 1848, p 66, pl. 1. figs. 1-5; (norvegica) Calman,

Marine Bor‘ng Animals, Brit. Mus. Nat. Hist. Economic Series,

No: 10; L995 p.95 fig. 2, pda tigeo:

Teredo bruguierti Delle Chiaje, Memorie &e. iv. 1829 (1830),

. 32, pl. 54. figs. 6.12, 13; Suter, Manual N. Z. Moll. 1913,

Oe OSE jalls Thy figs. 7 a-d; Gatliff and Gabriel, Proc. R. Soe.

Victoria, XXViil. (n.s .) 1916, p. 118, pl. xi. figs. 9 & 12.

Remarks.— Suter recorded this species from Auckland, where,

however, he considered that it was in process of being displaced

by Xylotrya saulit (i. e. XY. australis). It is therefore of interest

to note that the latter species alone occurs in the collection which

we have received from Auckland.

Suter has adopted Delle Chiaje’s name for this species on the

ground that Spengler’s was “not binomial,’ and he is followed

by Gatliff and Gabriel. It is true that, in the part of Spengler’s

memoir which deals with the genus Zeredo (but not in that

dealing with Pholas), the specific name is followed by a comma,

not by a full stop. Those who consider this an adequate reason

for displacing a name long in use and widely known will, no

doubt, continue to refer to this species as 7’. bragwierit.

MARINE WOOD-BORING ANIMALS. 395

TEREDO MANNII Wright. (Text-figs. 2 & 3.)

Kuphus mannii Wright, Trans. Linn. Soc. xxv. pt. 3, 1866,

p. 565, pl. lxv. figs. 1-8; Hedley, Rec. Austr. Mus. ii. 1899,

p. 134.

Nausitoria manni Hedley, Rep. Austr. Ass. Adv. Sei. viii.

1901, p. 248.

Locality.—Brisbane, Queensland. Specimens forwarded by

Mr. H. A. Cullen, Engineer for Harbours and Rivers. From

Jetties &e. in river 5 or 6 miles from Moreton Bay, in Ironbark

(Zucalyptus paniculata) and Pine (Araucaria cunningham).

Remarks.—Of this species, already recorded by Hedley from

Cooktown, in the north of Queensland, the holotype, from

Singapore, is in the Museum collection. Unfortunately, it has

been allowed to dry, but it has been possible, by soaking it in

water, to extract the valves and pallets and to restore the siphons

Text-figure 2.

Teredo mannii (Wright), from Brisbane.

A. Outer, 3. Inner surface of pallet. C. Siphons and associated structures.

The fleshy collar has been divided and the right pallet removed.

to something like their original form. In all the characters thus

ascertained the specimens now recorded from Brisbane show a

close resemblance to the holotype.

The most characteristic feature of the species is the form of

the pallets (text-fig. 2, A & B). . The blade or expanded portion

is roughly erescentic in shape. The concave distal edge has a

convexity in the centre, giving it somewhat the outline of a

cupid’s bow, and the convexity is usually, but not always, incised

by a/ narrow median notch. The inner surface of the blade is

flat, the outer excavated distally; along the margin of the

excavated area is a border of dark brown horny substance which

is continued along the distal margin of the inner surface. The

396 DR. W. 1. CALMAN ON

remainder of the surface of the blade is chalky white, and the

material composing it envelops the distal end of the more trans-

lucent stalk and ends in a sharp irregular line*.

The valves of the shell in all the specimens J have examined

show, to an unusual degree, the effects of secondary absorption

already referred to. This is marked, not only in the dorsal

region, posterior to the articular knobs, as in the valves of more

normal Teredinide, but along the whole of the posterior margin

as far as the ventral articular knob. In most specimens the

whole region of the auricle has disappeared and the greater part

of the postero-median region. In some specimens, as in the

Text-figure 3.

Teredo mannii (Wright), from Brisbane.

Valves of shell. A. Right, B. Left valve of a spec!men in which the eroded area

(seen in B) only occupies the upper margin, while the lower part of the

auricle still persists. C. Right valve of a specimen of about the same

sizé in which the erosion extends down the posterior margin as far as

the lower point of the shell and the auricle has been entirely removed.

holotype, this latter region is represented by a narrow border

along the greater part of the hind margin; in one of the

specimens figured (text-fig. 8, C) it has been entirely removed

* The extremely close resemblance, pointed out by Wright, between the pallets.

of this species and those of Kawphus arenarius as figured by J. E. Gray is very

surprising, if the statements as to the widely different habitat of the last named

species are correct.

ee ee

MARINE WOOD-BORING ANIMALS. aoT

except for a small piece near the ventral angle. In another

specimen (text-fig. 3, A, B), in which the lower part of the

auricle still remains, it extends only a very little below the level

of the anterior division of the valve; on the inner surface it

is not defined anteriorly, passing with quite unbroken surface

into the postero-median division.

The siphons (text-fig. 2, C), of which the ventral, or inhalent,

slightly exceeds in diameter the dorsal, or exhalent, are separate

quite to the base. The fleshy collar which surrounds them is of

considerable size, and, in the contracted state of the preserved

specimens, conceals the siphons for about half of their length.

Genus Xytorrya J. E. Gray.

Xylotrya (Leach MS.) J. E. Gray, Proc. Zool. Soe. 1847,

p. 188,

Whatever Leach’s XYylotrya may have been (the reference by

Menke, Syn. Méth. Moll. 2nd ed. 1830, p. 121, and the description

by J. E. Gray, Syn. Brit. Mus. 44th ed. 1842, p. 76, suggest that

it was the genus now known as Xylophaga), the name appears to

have acquired validity only when Gray in 1847 referred to it the

Teredo bipalmulata of Lamarck.

The species of the genus are for the most part sharply

differentiated from those of Zeredo by the segmented blade of

the pallets. This blade is composed of a series of hollow cones

successively ensheathing one another and arranged on a central

axis forming a continuation of the stalk. The only approach to

a transition between the two genera that I have seen is found in

Nausitora dunlopei Wright, in which the ensheathing cones are

very numerous and closely set, and appear, in the solitary type-

specimen which I have examined, to be partly consolidated on

the inner surface. They thus come to resemble the lamine of

which the blade is built up in some, at least, of the species

of Veredo, differing from them, however, in their more regular

arrangement *.

Many of the species referred to this genus have been only 1m-

perfectly described, and still more imperfectly figured. Possibly

the two species to which new names are applied below may be

identical with forms already named, but they are certainly

distinct from any in the Museum collection.

XYLOTRYA AUSTRALIS, sp.n. (Text-figs. 6, 7, & 8.)

Calobates saulii Hedley, Proc. Linn. Soc. N. 8. Wales, xxi.

1898, p. 94, figs. 7—9.

Nausitoria saulii Hedley, Rep. Austr. Ass. Adv. Sci. vii. 1901,

DeLeon plac Tesi.

* Cf. Fischer, Journ. Conchyl. v. 1856, p. 131.

398 DR. W. T. CALMAN ON

Leredo (Xylotrya) saulii Suter, Man. N. Z. Moll. 1913, p. 1021,

pl. lv. fig. 8, a, 6; Gatliff and Gabriel, Proc. Roy. Soc. Victoria

(n. 8.) xxvill, 1916, p. 121, pl. xiii. fig. 11.

Nec Nausitora saulii Wright, Trans. Linn. Soc. xxv. 1866,

p. 067, pl. Ixv. figs. 9-15.

Localities.— Brisbane, Queensland. Specimens forwarded by

Mr, EK. A. Cullen, Engineer for Harbours and Rivers. From

Jetties &e. in river 5 or 6 miles from Moreton Bay, in Ironbark

(Lucalyptus paniculata) and Hardwood (£. maculata).

Auckland, New Zealand. Specimens forwarded by Mr. Hamer,

Engineer to the Auckland Harbour Board. From Kauri and

Black Butt timber.

Remarks.—Wright states that the type-specimens of his

Vausitora saulii, which were presented to the British Museum by

Miss Saul, came from Port Phillip, Australia, and it is perhaps

this statement rather than any very exact correspondence with

his description or figures that has led Australasian naturalists to

apply the name to the species common in Australian and New

Zealand waters. It appears, however, that Wright’s statement

was in error. The specimens in the Museum collection labelled

as “Types” and presented by Miss Saul are stated, on the label

and in the Register of Mollusca, to be from Callao, Perut. It is

true that none of the valves or pallets can be definitely recognised

as the original of any one of Wright’s figures, but there are

two characters in which these specimens agree closely with his

account; the auricle shows, on the inner surface, a series of

conspicuous curved ridges indicated in Wright’s pl. Ixv. fig. 10,

and several of the pallets have the outer surface worn away so

as to expose the ‘‘central core-like body” mentioned in the

description (p. 568) and shown in pl. Ixv. fig. 15.

These type-specimens, however, appear to me to be specifically

distinct from those I have examined from Brisbane and Auckland,

which undoubtedly belong to the species called WV. saulii by

Hedley and other Australasian naturalists. The chief differences

may be briefly stated as follows :—

XY. saulii (Wright) (text-figs. 4 & 5). Dorsal outline of valve

sloping steeply without break into the upper margin of the

auricle, which exceeds half the total depth of the valve and

descends on the hind margin for more than half the distance

from the anterior notch to the ventral edge. The auricle is

marked with rather widely spaced lines of growth which, espe-

cially on the inner surface, appear as strong curved ridges. The

anterior border of the auricle on the inside overlaps as a narrow

band and is closely appressed to the inner surface. The pallets

(text-fig. 5) have the segments strongly calcified and closely set,

the average interval being estimated at not more than one-fifth

* Suter (Man. N.Z. Moll. 1913, p. 1022) mentions Callao among the localities

for the species, but states that the type is from Port Phillip. 1 do not know the

source of his information.

MARINE WOCGD-BORING ANIMALS. 399

of the width. The distal edges of the segments are acutely

V-shaped (this may be partly but not altogether due to their

being worn away), and no trace of serration can be seen in any of

Text-figure 4,

Aylotrya saulii (Wright). Syntype from Callao.

A. Right, B. Left valve of shell.

Text-figure 5.

Xylotrya saulii (Wright). Syntype from Callao. Pallet.

Text-figure 6,

Xylotrya australis, sp. n. Syntype from Auckland.

A. Right, B. Left valve of shell.

the specimens. The stalk is smooth, The soft parts are un-

known. The valves measure up to 7'3 mm. in length and a little

less in depth.

400 DR. W. I. CALMAN ON

X. australis, sp. n. (text-figs. 6, 7, & 8). Dorsal outline form-

ing a distinct angle or concavity at the base of the auricle, which

is not more than half the total depth of the valves and descends

on the hind margin for not more than half the distance from the

anterior notch to the ventral edge. The lines of growth on the

auricle are much more closely set and less prominent, and are not

at all conspicuous on the inner surface. The anterior border of

the auricle on the inside overlaps as a broader band which is

Text-figure 7.

Xylotrya australis, sp.n. Syntype from Auckland.

A, External surface of pallet. B. Single segment of

the pallet, further enlarged.

generally slightly raised from the inner surface. The pallets

(text -fig. 7) have the segments thin and fragile and more widely

spaced, the average interval being estimated at about one-third

of the width. ‘The distal margins of the segments are concave

or obtusely V-shaped, with a delicate membranous border, at the

base of which the calcified portion shows a series of coarse and

somewhat irregular serrations which become very conspicuous in

dried specimens. The stalk is smooth and shorter than thre-

MARINE WOOD-BORING ANIMALS. 401

‘times the width of the distal part. The siphons (text-fig. 8) are

adherent for two-thirds of their length in preserved specimens.

Text-figure 8.

Xylotrya australis, sp.n. Syntype from Auckland.

Siphons and associated structures from the right side. The fleshy collar

has been divided and reflected, the pallets remaining attached.

Text-figure 9.

Xylotrya capensis, sp.u. Syntype from Simon’s Town.

A. Right, B. Left valve of shell.

The largest complete specimen (from Auckland) is about 35 em.

long in the preserved state. The valves measure up to 13°5 mm.

in length and about the same in depth.

402 DR. W. T. CALMAN ON

XYLOTRYA CAPENSIS, sp. n. (Text-figs. 9, 10, & 11.)

Locality. Simon’s Town, South Africa. Specimens forwarded

by Lieut. L. H. A. Shadwell, R.N.V.R., Officer in charge of

Works, H.M. Dockyard. From Blue Gum timber, Ordnance

Jetty, E yard, and from Pitch Pine, Old Ordnance Jetty, E yard.

Description.—Valves of shell (text-fig. 9) with dorsal outline

not steeply sloping posteriorly, where it is defined from the

auricle by a shallow concavity. The auricle is very deep, exceed-

ing half the total depth of the valve and extending downwards

Text-figure 10.

Aylotrya capensis, sp.n. Syntype from Simon’s Town.

A. Distal portion of pallet. DB. Stalk of pallet. C. Single segment

of pallet, further enlarged.

for about half the distance from the anterior notch to the ventral

edge. The lines of growth on the auricle are rather closely set

and not conspicuous on the inner surface. The anterior border

of the auricle on the inside overlaps as a broad band, which is

distinctly raised from the inner surface. The pallets (text-fig. 10)

have the segments thin and fragile and closely set, the average

interval being estimated at one-sixth of the width. The distal

margins of the segments are regularly crescentic (on the outer

surface) with a broad striated membranous border but without

MARINE WOOD-BORING ANIMALS. 403

any trace of serration. On each side the border is produced as

a long filament which extends beyond three or four segments in

front. The stalk is minutely roughened and, in the specimen

measured, nearly five times as long as the width of the distal

part. The siphons (text-fig. 11) are adherent for at least five-

sixths of their length in preserved specimens.

The largest complete specimen is about 30 em. long in the

presen ae state (excluding the pallets). The valves measure

12°5 mm. in length by 11°5 mm. in depth. The pallets are about

46 mm. long.

Text-figure 11.

) va \\

)y)

Aylotrya capensis, sp.n. Syntype from Simon’s Town.

Siphons and associated structures from left side. The fleshy collar

has been divided and reflected, the pallets remaining attached.

Remarks.—In the structure of the pallets, especially in the

elongated peduncle, the broad, closely-set segments, and the long

lateral filaments into which they are produced, this species differs

from all those of which I have seen specimens. Blainville’s

USSU of the pallets OP AC pennatifera, with the segments

‘pourvues de chaque cdté d’un long cil” suggests a comparison

with our species. Specimens in ihe Museum collection referred

to XY. pennatifera, however, differ widely, having the segments

but little wider than the stalk and bearing a fringe of filaments

in place of the striated membranous border.

pS)

Proc. Zoo, Soc,—1920, No. XX VII.

we “wel

is aie

' posit

‘deadaiite

ay Ws Pr Bt

ON ENTOZOA FROM ANIMALS WHICH DIED IN THE GARDENS. AO5

22. Report on Entozoa collected from Animals which died

in the Zoological Gardens of London during Hight

Months of 1919-1920. By G. M. Vuvurs, M.R.C.8.,

L.R.G.P., F.Z.S., Beit Memorial Research Fellow,

Demonstrator in Heiminthology at the London School

of Tropical Medicine, and Honorary Parasitologist to

the Zoological Society of London.

[Received June 1, 1920: Read June 1, 1920. |

During the past eight months I have made an attempt to

examine systematically for Entozoa all animals dying in the

Gardens, and have attended post-mortem examinations of four

hundred animals for this purpose.

Before the body was actually opened, a microscopical exami-

nation of the feeces was made, whenever practicable, for ova and

embryos of Entozoa which would give some indication of the

parasites harboured, and would direct attention to the particular

regions for special search.

Whether this preliminary investigation gave a positive or

negative result, a subsequent search of all organs was carried out.

I have also applied this method of diagnosis to living animals

in the Gardens, and these examinations have in some cases given

positive results. On the death of the animal the diagnosis has

been confirmed by the discovery of the adult parasites; for

example, the Cylichnostomes recorded from the Grevy’s Zebra in

the accompanying chart were detected in this manner.

Of the four hundred animals examined 76 or 19 per cent. were

found to harbour parasites.

The Entozoa found fall into the following Phyla and

Classes :—

Number

of Species Percentage.

found.

Cestodne eer ie 21-40

PLATYHELMIA. {

TM ReITHAEHHOC EY Scekcasecene of 10-00

Nematode. ssseaea se: 45 64:40

NEMATHELMIA. {

Acanthocephala ...... 3 4-20

Sot ile 70 100:00

Tn all cases of Nematoda and Acanthocephala there was a pre-

ponderance of female forms. In four eases females only were

found. ‘There were 13 animals which harbouzed more than one

species of parasite. In a Leopard Cat (Hels bengalensis) as many

as five different species were found.

The material afforded a valuable opportunity of determining

27°

406 MR. G. M. VEVERS ON ENTOZOA FROM ANIMALS

the length of life of parasites of various groups in their hosts.

Very little reliable information has been gathered on this im-

portant point. The evidence given by the incidence of parasites

which have intermediary hosts is, of course, more trustworthy

than that of forms which have a simple life-cycle. Im the latter,

infection is accumulative, and may either be acquired in the

paddock or be brought into the Gardens on food: thus nine

examples of Gastrodiscus egyptiacus were found in a Grevy’s

Zebra which had been in the Gardens for six years. This parasite

normally occurs in Africa, and requires as an intermediary host

a freshwater molluse (Cleopatra bulimoides) which has not been

recorded from Europe. There can be no doubt, then, that the

specimens found had actually lived in the Zebra since it came

from Africa, and were, therefore, over six years old.

In the same Zebra were a number of species of Bursate

Nematodes, some of which have been recorded both trom Afriea

and Kurope. The life-cycle here is a simple one. That it is not

possible to draw trustworthy conclusions in such a case is well

illustrated by the findings in an Onager (Zquus_ onager) which

died quite recently. A number of the same species of parasites

which occurred in the Zebra were found in this Onager, which

had been born in the Gardens.

Two of the species of Cylichnostomes in the Grevy’s Zebra had

not been previously recorded, and it is possible that these were

originally imported, but the infection may have been renewed

in the paddocks. In this connection it is noteworthy that a

Chapman’s Zebra which died last year, after nine years in the

Gardens, had only species of Bursate Nematodes which occur in

European Horses.

We have, as another example of the contaminative group which

might accumulate in the Gardens, an apparently unrecorded

species of Atractis in the Elephant. Many specimens of this

Nematode were found in the Indian Elephant which died in

December last and which had been in the Gardens for twelve

years. Recent examination of the feces of the Elephant living

in the next paddock showed that this one also is heavily infected,

The minute but fully mature females of Atractis are passed

from time to time in the feces, and these contain embryos so far

advanced as to have the adult form. If these embryos are

discharged by the mother worm in the intestine of the host,

it is yonesive ble that they might attain sexual maturity almost

immediately, and would then provide an exception to the general

rule that parasitic worms do not produce a second generation of

adult forms within the body of their definitive host.

The following points of especial interest were noted in

individual species of Parasites :—

Two specimens of Gnathostoma spinigerum were obtained from

the stomach of a Leopard Cat (Felis bengalensis). A dissection

of the head-parts of a still living worm showed that the neck-

elands are hollow and contractile, ‘and contain a fluid which plays

WHICH DIED IN THE ZOOLOGICAL GARDENS, 407

a part in altering the size of the head, thus supporting the view,

as to the function of the ‘ ballonets,” recently put forward in a

paper read before this Society by Baylis and Lane*.

A microscopical examination of the feeces of the same Leopard

Cat showed many minute Nematode embryos. A similar exami-

nation of the stomach contents gave the same result, but no

adults could be found in this or any other of the neighbouring

organs. However, in the mucosa of the cesophagus and naso-

pharynx the same embryos were present, but here each was

coiled up in an exceedingly thin membranous shell. ‘The pre-

sence of these viviparous eggs in the nasopharynx led to the

discovery of the adult worms in a most unusual position, for the

frontal sinus was next explored, and here large numbers of a

species of Synthetocaulus were found. So far as we have been

able to ascertain, this species is new to science, but it is closely

allied to S. rufescens, which occurs occasionally in the lungs and

air-passages of the Sheep in Europe.

It is of interest to note that for some time before death the

animal suffered from “ fits” and was often seen to lose its balance

and fall. These “ fits,” and loss of equilibrium were no doubt

due to the presence of Synthetocaulus in the frontal sinus.

Further examination of the feces from the same

showed many Trematode ova, which were recognized as those of

Paragoniinus westermanni. The lungs were then searched, and

four specimens of the adult fluke found. The number of eggs in

the feces was exceedingly large considering the few adults which

gave rise to them.

IT am indebted to Professor R. IT. Leiper for his invaluable

assistance and advice on a number of the more intricate points

arising in the course of the above inquiry.

aninal

List of Parasites found, with their Hosts.

TREMATODA.

Length of

Genus. Species. Host. time in

Gardens.

+Gastrodiscus aeyptiacus Grevy’s Zebra. 6 years.

(Cobbold, 1876). (Africa.)

Railliet, 1898.

+Notocotyle triserialis (2) Netta rufina. 1 week.

(Diesing, 1839). (India.)

Diesing, 1850.

Paragonimus westermanni Felis bengaleusis. 6 months.

(Leuckart, 1889). (India.)

Stiles, 1900.

Platynosoma _illiciens Khai phastos 3 months.

(Braun, 1901). erythrorhynchus.

Looss, 1907. (S. America.)

* P.Z.S. 1920, p. 245.

+ Denotes that this Parasite has not been recorded before from this Host.

408 MR. G. M. VEVERS ON ENTOZOA FROM ANIMALS

Genus.

Species.

Macrodera

formosum

Nicoll, 1911.

naja (Rud. 1819).

Looss, 1899.

Host.

Length of

time in

Gardens.

Zamenis flagelliformis.

(S. America.)

Tropidonotus natrix.

(Britain.)

CrsroDA.

Cyclophyllidea.

Tenia

Davainea

Hymenolepis

(Echinocotyle.

Hyracoteenia

Oplnotenia

Tetrabothrius

crassicollis

Rud. 1810.

goura

Fuhrmann, 1909.

paucitesticulata

Fuhrmann, 1909.

sp. Inq.

sp. nov.

sp. 10V

villosa

(Bloch, 1872).

Wolfth. 1899.

sp. ing.

)

procavize

Beddard, 1912.

sp, ing.

cylindraceus

(Rud. 1819).

Diesing, 1850.

Pseudophyllidea.

Dibothriocephalus sp. inq.

Ascaris

Genetta genetta.

(Spain.)

Goura coronata.

(New Guinea.)

(2) Calcenas nicobarica.

(Nicobar Islands.)

Fringilla celebs.

(Britain.)

Schizorhis concolor.

(S. Africa.)

Caccabis chukar.

(Syria.)

Casuarius

uniappendiculatus.

(New Guinea.)

Casuarius

uniappendiculatus.

(New Guinea.)

Tetrax tetrax.

(Britain.)

(4) Quelea quelea.

(S. Africa.)

Procavia capensis.

(S. Africa.)

Crotalus atrox.

(Cent. America.)

(2) Larus glaucus.

(Europe.)

Conepatus proteus.

(Argentine.)

NEMATODA.

osculata Otaria californiana.

Rud. 1819. (North Pacitic Ocean.)

holoptera (8) Testudo ibera.

Rud. 1819. (S. Europe.)

sp. ing. Casarea casarca.

(Europe.)

7 months.

5 months.

6 months.

1 week.

6 months.

3 months.

10 months.

6 months.

9 months.

4. months.

3 months.

5 months.

7 years.

2 years.

6 months.

1 year.

5 years.

+ Denotes that this Parasite has not been recorded before trom this Host.

WHICH DIED IN THE ZOOLOGICAL GARDENS.

409

Length of

(Rud. 1819).

(Esophagostomum apiostomum

Willach, 1891.

Ancylostomum conepati

Uncinaria

Hemonchus

Syngamus

Solanet, 1911.

criniformis

(Goeze, 1782).

sp. ing.

contortus

(Rud. 1803).

bronchialis

(Muhlig, 1884).

Cylichnostomum imparidentatum

(Poteriostomum).

+Cylichnostomum goldi

Quiel.

Boulenger, 1916.

+(@sophagodontus robustus

Giles, 1892.

+Triodontophorus intermedius

Sweet, 1909.

(S. America.)

(7) Macacus rhesus.

(India.)

Conepatus proteus.

(Argentine.)

Vulpes vulpes.

(Britain.)

Felis lynx.

(Thibet.)

Hippotragus equinus.

(Africa.) ,

Casarea casarca.

(Europe.)

Chapman’s Zebra.

(Africa.)

Chapman’s Zebra.

( Africa.)

Chapman’s Zebra.

(Africa. )

Chiapman’s Zebra.

(Africa.)

Genus. Species. Host. time in

Gardens.

Ascaris sp. inq. Spheniscus demersus. 3 weeks.

(S. Africa.)

Belascaris mystax Felis bengalensis. 6 months.

(Zeder, 1800). (India.)

Leiper, 1907.

Toxascaris sp. inq. Vulpes lagopus. 2 weeks.

(Syria.)

Porrocecum crassum Grus communis. 9 years.

(Deslongchamps, 1824). (Hurope.)

Raill. et Henry, 1912.

Contracecum — spiculigerum Phalacrocorax carbo. 4 months.

(Rud. 1819). (Britain.)

Raill. & Henry, 1912.

Oxysomatium brevicaudatum Anguis fragilis. e

(Zeder, 1800). ( Britain.)

Heterakis vesicularis Phasianus torquatus. 2 weeks.

(Dujardin, 1845). (China.)

Heterakis vesicularis Ceriornis satyra. 7 years.

(Dujardin, 1845). (India.)

+Cucullanus microcephalus Chrysema seripta rugosa. 1 month.

(Dujardin, 1845). (West Indies.)

Gnathostoma spinigerum Felis bengalensis. 6 months.

Owen, 1836. (India.)

Ascaridia sp. Inq. Centropus rufipennis. 10 days.

(India.)

Ascaridia lineata (2) Ocyphaps lophotes. 4 years.

(Schneider, 1836). (S. Atrica.)

Physaloptera _— retusa ‘Tupinambis teguexin. 1 month.

18 months

(approx.).

2 years.

5 months.

1 week.

5 years.

9 years.

br}

+ Denotes that this Parasite has not heen recorded hefore from this Host.

410 on

ENTOZOA FROM ANIMALS WHICH DIED IN THE GARDENS.

Length of

Genus. Species. Host. time in

Gardens

+Strongylus edentatus Chapman’s Zebra. 9 years.

Looss, 1901. (Afvica.)

Stronvgylus vulgaris Chapman’s Zebra. 3

Looss, 1901.

}Probsmayria vivipara Grevy’s Zebra. 6 years.

Ransom, 1907.

+Cylichnostomum nassatum var. parvum

Strongylus

Strongyloides

Synthetocaulus

Oxyuris

Filaria

Setaria

Diplotrizna

Trichocephalus

Dispharagus

yAtractis

Yorke & Macfie, 1918.

vulgaris

Looss, 1901.

intestinalis

Grassi, 1883.

sp. Inq.

equi

Schrank, 1788.

longicollis

Schneider, 1866.

gracilis

Dujardin, 1845.

sp. 1nq.

tricuspis

(Hedschenko, 1879).

Raill. & Henry, 1909.

affinis

Rud. 1801.

dispar

Rud. 1801.

squamatus

(v. Linstow, 1883).

sp. noy.

Grevy’s Zebra.

Felis bengalensis.

(India.)

Felis bengalensis.

(India.)

Chapman’s Zebra.

(Atrica.)

Testudo graca.

(Europe.)

Ateles grisescens.

(S. America.)

Hippotragus equinus.

(Hast Africa.)

Acridotheres ginginianus.

(India.)

Ovis vignei.

(India.)

Macacus rhesus.

(India.)

Phalacrocorax carbo.

(Britain.)

EHlephas indicus.

(India.)

AGANTHOCEPHALA.

Kchinorhynchus claveeceps.

Echinorhynechus) sp. ing.

J ) ST q

gen. Inq.

(Echinorhynelius) sp. ing.

gen. inq.

Chrysema scripta rugosa.

(America.)

Callicebns moloch.

(S. America.)

Leontocebus ursulus.

(S. America.)

6 months.

9 years.

6 months.

2 months.

1 week.

6 years.

13 months

(approx.).

4, months.

12 years.

1 month.

3 weeks.

+ Denotes that this Parasite has not been recorded before from this Host.

ON TATLLESS BATRACHIANS FROM EAST AFRICA. 411

23. On a Collection of Tailless Batrachians from Hast Africa

made by Mr. A. Loveridge in the years 1914-1919.

By Miss Joan B. Procrer, F.Z.S.

[Received May 19, 1920: Read June 15, 1920.]

(Text-figures 1-4.)

This collection, made during the war, consists of examples of

33 species, two of which are new, representing the families

Ranide, Hngystomatide, and Bufonide*. Mr. Loveridge has

presented the types of Rappra platyrhinus and Megalixalus

loveridgii to the British Museum, together with specimens of the

little-known species <Arthroleptis stenodactylus Pfeft., Rappia

punticulata Pfeff., and many others.

IT am greatly indebted to Mr. Boulenger for much kind

assistance in the working out of this collection.

List of Localities.

The localities fall into three divisions :—

1. Brivis Hast AFRICA.

Nairobi.

Parklands, Nairobi.

Besil, about 50 miles south of Nairobi.

Donya Sabuk, 30 miles from Nairobi.

Thika, about 50 miles from Nairobi.

Kagiado, about 100 miles due west of Makindu, which is on

the Port Florence-Mombasa Railway.

Kedong Valley, about 100 miles from Nairobi, and south of

Escarpment Station on the railway.

Kenia Forest.

Tumu Tum, on the road from Thika to Mt. Kenia.

2. Ex-German Hast Arrica (now Tanganyika Territory).

Gonya, about 50 miles south of Kilima-Njaro.

Longido West, about 50 miles west of Kilima-Njavro.

Amani, 40 miles from Port Zanga.

Dodoma, about 110 miles west of Morogoro, on the Dar-es-

Salaam—Tanganyika Railway.

Kongwa, about 20 miles north of Mpapua.

Morogoro, about 100 miles west of Dar-es-Salaam on the

railway.

Tulo, )

Duthumi, J

* Pipide should also have been represented, but the large series of Xenopus

collected were lost in transit.

50 and 60 miles south of Morogoro.

4132 MISS JOAN B. PROCTER ON TAILLESS

Mhonga, about 20 miles east of Morogoro.

Dar-es-Salaam.

3. PortucursE EKasr AFRICA.

Lumbo.

RANIDZA.

RANA, 8. str.

1. Rava nurti Bler.

Nairobi. Morogoro. Longido West.

35 specimens. *harer age length of adult males 55 mm., of

adult females 70 min. ; largest female 83 mm.

Snout variable both in length and in degree of acumination.

Subgenus ToMoprerNa.

9. RANA DELALANDII Bibr.

Kagiado. Ex-G.H.A.

4 specimens. Length of adult females 41 and 45 mmm.

A fine white ver selena line in one specimen ; a linear vertebral

groove in three specimens.

Subgenus PrycHADENA.

3. RANA OXYRHYNCHUS Sund.

Thika. Nairobi. Morogoro, Duthumi. Gonya.

72 specimens, Average length of adult males 40 mm., of

adult females 55 mm.

An extremely variable species. Snout equal to or twice as

long as diameter of eye, rounded or acutely pointed, sometimes

very projecting. Length of tibia 14 to 1% times in length of

body, more often 13 times in females. Posterior corner of sht-

like opening of male’s vocal sac in line with lower border of arm.

No light vertebral band.

4, RANA MASCARENIENSIS D. & B.

Nairobi. Gonya. Ex-G.H.A.

15 specimens. Average length of adult males 47 mm.; females

half-grown.

Hind limb very variable in length, tibio-tarsal articulation

See anywhere between eye and “tip of snout ; length of tibia

13 to 1? times in length from snout to vent in males, 1? to 2

times frat females. Vocal sac of male in line with upper border

of arm; 11 half-grown females have rudimentary vocal sacs.

Markings very regular, the first two series of spots coinciding

with the first two glandular folds, and often bisected by them :

almost all specimens have a broad light vertebral band, and

often a still lighter fine vertebral line im acldition.

though nearly full-grown.’—A. LovertpGE, Field Notes.

€

BATRACHIANS FROM EAST AFRICA. 413

Subgenus HipEBranprTiA.

5. RANA ORNATA Ptrs.

Ex-G.H.A.

l specimen, 9. Length 56 mm.

Exquisitely marked with series of dark, large, elongated spots,

broad temporal bands, barred limbs; throat and breast mottled

with dark brown, with the characteristic paired Y-markings.

CHIROMANTIS.

6. CHIROMANTIS XERAMPELINA Ptrs.

Ex-G.H.A.

1 specimen, probably a female. Length 66 mm.

7. CHIROMANTIS PETERS Bler.

Dodoma. Ex-G.H.A.

3 specimens. Length 50, 55, and 59 mm.

Inner fingers with a mere rudiment of web, outer differing

from those of the type in being 7 instead of 3 webbed.

PHRYNOBATRACHUS.

8. PHRYNOBATRACHUS NATALENSIS Smith.

Kagiado. Nairobi. Longido West. Morogoro.

29 specimens. Average length of adults 30 mm.

An extremely variable species. ‘Toes sometimes 3 instead of/}

webbed. Skin perfectly smooth or extremely warty; all male

specimens from Nairobi are of the latter description, but three

females from the same locality are smooth. A wide white

vertebral streak in three specimens.

9. PHRYNOBATRACHUS RANOIDES Bley.

Morogoro. HEx-G.H.A.

2 specimens. Length of larger 33 mm.

10. PHRYNOBATRACHUS BOULENGERI de Witte.

Morogoro. Duthumi. Gonya. Tulo. Ex-G.H.A.

61 specimens. Average length of adults 26 mm.

Skin perfectly smooth or warty. Markings variable, three

forms with and one without vertebral bands; five specimens

have a very broad vertebral band, two havea broad one, and two

have a fine white vetebral line. These four forms are shown

in figures of P. natalensis *, P. Z.S. 1907, pl. xxii.

* Boulenger, ‘‘ Second meee on the Batrachians and Reptiles collected in South

Africa by Mr. C. H. B. Grant.

414 MISS JOAN B. PROCYER ON 'TAILLESS

11. ARTHROLEPTIS WAHLBERGII Gthr.

Morogoro, Amani.

2specimens. Length 25 mm.

Dark without distinct markings; dermal ridge along the

vertebral line. Back of one specimen covered with minute

tubercles.

12. ARTHROLEPTIS WHYTII Blegr.

1 specimen. Length 35 mm.

Pale brown above, with a dark subtriangular marking between

the eyes; a fine dermal ridge along the vertebral line.

13, ARTHROLEPTIS MINUTUS Bler.

Kenia Forest. Nairobi.

33 specimens. Average length 17 mm.; large specimens 22 mm.

The majority of the specimens are dark olive, without distinct

markings; four have a fine, and two a broad white vertebral

line.

14. ARTHROLEPTIS STENODACTYLUS Pfeff.

Morogoro. Duthumi. Ex-G.H.A.

16 specimens. Average length of adults 32 mm.

This species, described by Pfeffer * from a single female from

Kaihengo, is new to the collection of the British Museum.

Text-figure |.

Hand of male and female Arthroleptis stenodactylus.

Underside, X 2.

Head broader than long; canthus rostralis obtuse; loreal

region oblique, concave; interorbital width greater than that of

upper eyelid ; tongue with a conical papilla; tympanum distinct,

usually 3 diameter of eye. First and second fingers equal in

length, third 15 times length of second in females, 13 to 2 times

in males ; tips of fingers and toes somewhat swollen but not

* Jahrb. Hamb. Wiss. Aust. x. (1898) Taf. 1. fig. 11.

BATRACHIANS FROM EASY AFRIGA, 415

dilated ; tees with a rudiment of web; inner metatarsal tubercle

as long as or slightly shorter than inner toe; no outer tubercle.

Tibio-tarsal articulation reaches the eye or slightly beyond;

length of tibia 3 times its breadth, twice in length of body.

Skin smooth. Reddish brown or olive-grey above, with the

dark vertebral markings characteristic of the genus; a fine white

vertebral line in five, a broad one in three specimens; barred

limbs; whitish beneath, sometimes with a dark throat: a few

spots on lower lip. A fine linear vertebral ridge in two

specimens. Males with an internal vocal sac. F

RAPpPIA *,

15. RAPPIA GRANULATA Blgr.

Ex-G.H.A.

6 young.

16. RAppPrIiaA PUNCTICULATA Pfeff.

Morogoro. Ex-G.H.A.

7 specimens. Length 22 to 32 mm.

Text-figure 2.

Rappia puncticulata. Nat. size.

Interorbital space varies from 14 to 2 times width of upper

eyelid. Fingers with a rudimentary web, toes ? webbed.

Length of tibia 21 to 24 times its breadth, 33 to 4 times in

length of body.

The specimen figured by Pfeffer is of a uniform colouring,

* “ Rappia, sp. A Tree Froz was found impaled on the spike of an Aloe, in

company with sundry grasshoppers which formed the larder of a Shrike.”°—

A, LoverrDGE, Field Notes.

416 MISS JOAN B. PROCTER ON TAILLESS

with a light dark-edged lateral band passing round snout and

ending on the sacral region; it has also a fine dark vertebral line.

Several young in the British Museum collection have similar

markings, but the seven noted above differ considerably. In

these the light lateral band is heavily bordered with black, the

enclosed dorsal area and upper surfaces of tibia, tarsus, and

forearm irregularly spotted with black or dark grey ; they have

also a large white dark-edged spot on the heel. The markings

of the smallest specimen are so light and indistinct that it more

resembles the type specimen.

17. RApPIA PLATYRHINUS, sp. n.

Nairobi.

1 specimen, ¢.

Head small, broad as long, moderately depressed ; snout flat,

markedly tr umncate, as long as diameter of eye; canthus rostr alls

feebly marked ; loreal region oblique, concave; nostril at end of

Text-figure 3.

Rappia platyrhinus. Nat. size.

snout; interorbital space nearly twice width of upper eyelid ;

tympanum hidden. Fingers long, ; webbed, second longer than

first; toes slender, fully webbed : subarticular tubercles soft,

flat, small; inner metatarsal tubercle also soft and small ;

outer one. ‘Tibio-tarsal articulation reaches middle of eye;

length of tibia 5 times its breadth, 13 times in length from snout

to vent. Skin smooth above, granular below. Uniformly

brownish-grey above, finely speckled with black pigment cells

which are slightly concentrated along canthus rostralis. Male

with internal vocal sac and adhesive subgulay cise,

BATRACHIANS FROM EAST AFRICA. 417

Measurements in millimetres.

SMOUMULCOMMEMO Mc... 5s... ge. ae eee 28

Eteadieeee. 9

Width of eae 9

SMOWb) esses eM IRM Reo 2c 3

Tnterorbital raed Sab eg gs ae ean oa)

1st rayeyer airs ch Ue mn MM _ 2°5

Did oe

SES uh a ara ace Get ele a Nm aS et EN 5

ABE) IN de Soe SLA Se ae RE es ELS 4A

FPS olan lini O Pere er ert. ca 5 sisters ociame ye eee 43

COICTEVE | PG ksanns Seine Sete em Sm Emre Sema Ratan 14

TING egal ceiatl cie tena ny ee aR eR Rc, 19

The broad truncate snout combined with the very short web

between the fingers are the distinguishing features of this

species.

18. RApPIA CINCTIVENTRIS Cope.

aol Ex-G.E.A.

2 specimens. Length 31 and 20 mm.

Male from NATobi greyish ; female from Bx-G.E.A. pale

brown, with a subtriangular dark marking; both have two paired

series of small aneibene le on head and shoulders. The male

differs from the normal in having a subgular dise.

19. Rappra FuLYOVITrATA Cope.

Duthumi. Morogoro.

30 specimens. Average length 22 mm.

Body exceptionally elongated and depressed. Light purplish

brown, with two paired dorsal streaks of a darker shade but

indistinct.

20. RAPPIA MARMORATA Rapp.

Besil. Nairobi.

5 specimens. Length of largest 34 mm.

Male specimen ‘aon Nemobs is black, with three wide white

dorsal bands and finely spotted sides and limbs; two females

from the same locality are uniform grey, speckled with black

above, salmon-pink beneath, especially hinder side of thighs;

two females from Besil are uniform pale grey.

21. RAppPiaA ARGuS Ptrs.

Morogoro. Dar-es-Salaam.

3 specimens. Length 32 and 35 mm.

Two are light brown above; a white, black-edged band passing

round snout, , through eye to temple or back of “head ; : the third

has several ocellar spots on the back, but no canthal band. The

absence of the characteristic spots is unusual, but there are

418 MISS JOAN B. PROCTER ON 'TAILLESS

two similar specimens in the British Museum collection. In the

spotted form the canthal band is usually present and continued

down the side of the body.

MEGALIXALUS.

22. MEGALIXALUS LOVERIDGII, sp. n.

Morogoro,

1 specimen, 2.

Head small, depressed, broader than long; snout rather

pointed, as long as diameter of eye; canthus rostralis rounded ;

Text-figure 4.

Megalixalus loveridgii. Nat. size.

loreal region feebly oblique; nostril near end of snout; inter-

orbital width greater than that of upper eyelid; tympanum

just distinguishable on right side, 3 diameter of eye. Fingers

long, 3 webbed, the web extending as a fringe to discs of third

and fourth, second longer than first, third twice length of first ;

toes entirely webbed; discs small and round; subarticular

tubercles small, soft, feebly prominent ; inner metatarsal tubercle

small, soft, oval; outer one minute. Tibio-tarsal articulation

reaches eye; length of tibia 43 times its breadth, 24 times in

BATRACHIANS' FROM BAST AFRICA. 419

length of body. Length from snout to vent 3} times length of

head. Skin smooth, dotted with minute white tubercles each

bearing a minute black spine, on upper surfaces of head, body,

forearm, tibia,@and hinder side of tarsus. Smooth beneath ;

throat granular ; - some small warts at corners of mouth. Pale

brown above, with a faintl y marked band, commencing on snout

and progressively widening on the back. Uniformly speckled

on upper surfaces with black pigment cells, which are slightly

concentrated along the canthus rostralis; lower surfaces pale

brown.

Measurements ur millimetres.

SIMOMIG FOR VEIL Any ages dadccs. te ocean mene er 36

TEC GL AB a a eetials COU. ita tu Se Mane ae pence a Ee 3 1]

WATCH MMORMINE AGM Geet iiosec. nek le sce, ee IE

STA GLUE kb acicae ocis a e a A RMR S TER Voy AR aie, 45)

Ilniermorlomeall wild Sooonnedsnesoccnonconsonoos 3°D

LUSih EINIEN Dobos ocnsn Copenne We Au std saaradiecaemee 4

DAG Wie Neh oo RARE ee Meee men reese ae 5

3 ME RE ieee tet ore oe te os, ute stscnintelarae See ER 8°5

ANG Ng Me PoP io sala Qo apecsn LU erepa ts ascites 6

eTenratclrlintviallo peewee verseistsics as coats sare toe a

REIL OCs Urea AA Re MEM aM ORIE MR fe 1 8 2 i 18

Olimar ee resets tat ei ence aktia o Nats Rr De

The closest affinity of this species is I. fornasinii Bianconi,

which it resembles in dermal characters, but which differs in

having the fingers } to 3 webbed and shorter, especially the

third, Spite is i times length of first. It is alco less elongate

in habit, the length from snout to vent being little over 3 Annas

the length of the head.

CASSINA.

23. CASSINA SENEGALENSIS D. & B.

Nairobi. Ex-G.H.A.

3 specimens. Length 29, 39, and 40 mm.

Male specimens from Nairobi have a vertebral and two paired

series of dark elongated spots; female from Ex-G.E.A. has five

unbroken dark bands. ‘The former have the tips of the digits

much swollen, almost amounting to small discs as in Hylambates ;

specimens in the British Museum collection from the same

locality show the same variation.

HYLAMBATES.

24, HyLamBatses Bocaci! Gthr.

Nairobi.

3 specimens. Length 26, 31, and 35 mm.

Skin slightly granular above, with a linear vertebral groove.

Upper parts uniform dark erey.

Proc. Zoou. Soc.—1920, No. XX VIII. y 28

490 ON TAILLESS BATRACHIANS FROM BAST AFRICA

HYLAMBA'TES JOHNSTONI Blor.*

Mhonga. Ex-G.H.A.

6 specimens. Length 27 to 45 mm.

ENGYSTOMATIDA.

PHRYNOMANTIS.

96. PHRYNOMANTIS BIFASCIATA Smith.

Ex-G.H.A. Lumbo.

3 specimens. Length 38, 40, and 48 mm.

Dark purplish brown with pink markings, agreeing with var, A

of the British Museum Catalogue.

BREVICEPS.

97. BREVICEPS MOSSAMBICUS Ptus.

Morogoro. Kongwa. Dodoma. Luimnbo.

15 specimens. Length 29 to 48 min.

HeEmMIsvs.

28, HeMIsus MARMORATUM Ptrs.

Gonya. Morogoro. ;

8 specimens. Length of largest 35 mn.

Length of tibia varivb! e, going from 23 to 3 times in length

from snout to vent. ‘The usual occipital fold is Abeentr in

three specimens, and a linear vertebral groove is present in three

specimens.

BUFONIDS.

BUFO.

29. Buro recuLAris Reuss.

Nairobi. Kedong Valley. Donya Sabuk. Tumu Tumu.

Longido West. Morogoro, Duthumi. ‘ulo.

27 specimens. Average length of adult 80 mm.

Every specimen has Glas: a fine light vertebral line or a

sunken line in the skin; one female has its thighs marbled with

red, but there are nove of the beautiful red forms figured in

Mr. Boulenger’s Second Report on the Grant Collection 7. The

parotoids ave very variable in length, sometimes 4 times as long

as broad, and greenish in colour.

30. Buro cARENS Smith.

Nairobi

1 specimen. Length 76 mm.

%* “At Mhonea, when my boy brought me my mule which had been grazing in a

patch of kathr-corn, I was ‘delighted to find my saddle already occupied by a large

frog, whilst another was ensconced in the nose-bag.”—A. Loverrp@n, Field Notes.

Dp?

a Pants UG, falls SO, .

ON THE TYPU-SPECIMEN OF RANA HOLSTI. 431

, TAO

24. On the Type-Specimen of Rana holsti Boulenger.

By Miss Joay B. Procrsr, 77.2.8.

[Received June 1, 1920: Read June 15, 1920. ]

(‘Text-figure 5.)

In “A Note on Bambina, the Dagger-Froe”*, Mr. G. K.

Noble states that the type-specimen of Rana holsti (Brit. Mus.

92 9. 3.19), which is a female, hag “a, well-developed dagger.”

This is not the case. The preepollex of this specimen terminates

- in a knob-like tubercle not unlike the subarticular tubercles of

the fingers. Even considerable pressure cloes not cause the

protrusion of the bone within. In order to ascertain whether

this female would have been able to expose a ‘‘ dagger,” I have

dissected the prepollex of the vight hand, and find that the

Text-figure 5.

Right hand of Rana holsti. Underside, nat. size.

Dissected to show termination of bone of praepollex, and distal phalanx

of first finger. (T'ype-specimen.)

bone, although undoubtedly very pointed, terminates over a

millimetre short of the tip of the dermal tubercle; moreover,

the tubercle has a thick wall, to the ianer side of which the tip

of the dagger is attached by soft connective tissue, forming a

pad between it and the skin which shows no sign of perfo-

ration. The bone is not curved, as stated in Mr. Van Denbure’s

description 7.

Mr. Boulenger’s description t:—‘‘ A very prominent knob

(rudiment of pollex) on inner side of first finger” is perfectly

correct, and there was clearly no ‘ oversight” of the dagger on

his part, since such a weapon does not exist. This mistake on

+ Proce. Cal. Ac. Sci. vol. iii. 1912, p. 197.

+ P.Z.S. 1913, p. 1023.

28*

439, ON THE TYPE-SPECIMEN OF RANA HOLSTT.

the part of Mi. Noble is difficult to account for, as his idea of

the type-specimen was apparently derived from Dr. Stejneger*,

who says: ‘Il have had the privilege of examining the unique

type-specimen, but I have nothing to add to Boulenger’s ori-

ginal description”; a most accurate figure of the left hand is

given by hin.

It was therefore quite a reasonable suggestion to make, even

if now proved to be incorrect, that the dagger is a secondary

sexual character of the male as the “aiguillons cornés et cadues

qui arment le doigt interne de certains Leptodactylus V Amérique

et qui servent a renforcer ’amplexus pendant l’accouplement” 7.

In Tympanoceros newtoni Bocage, for another instance, the male

is armed with a powerful, protruding dagger on the inner side of

the first finger, whilst the female has none whatever. This

point, however, is set at rest by Mr. Van Denburg, who says

‘the dagger is fully developed in adults of both sexes. Our

collection includes females which contain eggs nearly ready for

laying. These [sie] are armed with spurs as large and formidable

as are to be found in males.” This shows, therefore, that the

“dageer” may, or may not, protrude externally in FR. holst,

unless, which is unlikely, the females described by Van Denburg

and Noble belong to a distinct, closely allied species.

* Herp. Jap., Bull. U.S. Nat. Hist. Mus. No. 58, 1907, p. 105, fig. 84.

+ Boulenger, C. R. Ac. Sci. 1918, vol. clxv. pp. 987-999.

ON THE EUROPEAN AND AMERICAN BADGERS. 423

25. Onthe External and Cranial Characters of the Huropean

Badger (Meles) and of the American Badger (Tawidea)”.

By R..1. Pocock, F.R.8.

| Received May 22, 1920: Read June 15, 1920.]

(Text-figures 19-25.)

ConTENTS. Page

Mit ROCUIE bio Mee tscach ans cc wat he cosa acon eee 423

AVI ew Eliericlesieran arene eres aia tes a alonisiewsicitaren nisvehe ne Sea ee aL

CARN a DUCE ca ede, eh ae I ence eh ANS)

MnewAmalandeGenitalwAveds ene tse kererteee seen EAS

Nhe. (Sikqmll eaycel WI teetiNay en chocaboeedassaaayechcobscauonnoonee . “s3i4

WONCIUSTONS eee eee ea nee tO

Introduction.

Many descriptions have been published of the Huropean and

American Badgers +, and the wide divergences between them in

the structure of the skull and teeth were long ago insisted upon

by Baird ; but although attention has been drawn to some of the

differences in external characters, it seems that dried skins have

been in all cases the only material available for the purpose. So

far as I am aware, no author has had the opportunity hitherto of

instituting a comparison between the genera based upon fresh

material; and no one appears to have questioned the right of

Tauxidea to be included in the same subfamily as Meles. Hven

Gray ¢, who split his family Melinide (= Melince of many recent

authors) into the five tribes—Melina, Mellivorina, Mephitina,

Zorillina, and Helictidina, ranged Tuwidea alongside Meles, being

evidently of opinion that the kinship between these two genera

is closer than the kinship between J/eles and Arctonyc.

As will appear in the sequel, the outcome of my comparison

between the external characters—supplemented by cranial and

dental characters—of the two types is to suggest that the like-

nesses between them are superficial, adaptive, and due to

similarity of habits, and that the differences between them do

not justify their relegation to the same tribe or subfamily. It

will be remembered that MJellivora was also formerly assigned

to the Meline on account of its badger-like build and feet; but

the tendency of modern opinion is to regard the genus as a

* The facts recorded in this paper are based upon specimens examined at the

Society’s Prosectorium.

+ The most exhaustive and most recent description of the skull and teeth of

Meles known to me may be found in the ‘Catalogue of the Mammals of Western

Europe’ Ly Miller. he external characters, based upon an examination of dried

skins, are, however, briefly dismissed. Coues gave a long and on the whole accurate

description of Tawidea in his volume on Fur-bearing Animals, 1877.

t Cat. Carnivorous etc. Mammalia, 1869, pp. 120-121. Gill (Smithsonian Mise.

Coll. xi. pp. 64-66, 1872) adopted Gray’s subdivisions, but converted the tribes of

Melinidz into subfamilies of the Mustelide. Coues followed Gill.

424 MR. R. I. POCOCK ON THE EXTERNAL CHARACTERS

specialized member of he Musteline or to place it na subfamily

apart *. :

The Head.

The forehead is higher and rounder in Meles than in Tawidea.

The ear of Meles is modérately large with. tolerably evenly

rounded edge. It is simple in structure, the bursa being sup-

pressed; the tragus is small and the antitragus scarcely

developed, ‘The supratragus (plica principalis) is of average size

but not valvular, merely presenting a hemispherical thickening.

The ear of Zaaidea does not differ from that of J/eles in any

important particular apparently, although the. lamina is less

salient and its free edge is not continued inferiorly so far towards

a point beneath the intertragal notch (aditus inferior).

The facial vibrisse in Meles ave reduced by the suppression of

the interramal tuft, and the superior genal tuft is at most repre-

sented by one short bristle at least in the specimens examined.

The mystacial and submental vibrisse are moderately well deve-

loped, one of the latter on each side being exceptionally long; the

inferior genal tuft is represented by one or two bristles behind

the corner of the mouth and the superciliary tuft by two or more

over the eye. In Vaxidea the tufts are normal in number and

situation, the genal tufts being represented by about three

bristles, the upper being some distance below the level of the

eye; but the interramal tuft has only about two short bristles.

The nose of Meles is produced and snout-like, and overlaps the

under jaw considerably. The rhinariwm is exceptionally large ;

its upper surface is naked as far back as a line behind the

posterior ends of the nostrils. The anterior surface forms a

wide, deep, flat disc, without trace of a median groove. The

inner expanded portion of the nostril is large, the outer forms a

long narrow slit extending horizontally to the lateral edge of the

rhinarium. The infranarial portion is exceptionally deep and

well developed both mesially and laterally; its inferior edge is

convex, but varies in the degree of convexity, and is sometimes

proluced into a point in the middle line; but there is no philtrum

and the upper lip is hairy across the middle and uncleft.

Judging from descriptions, the rhinarium of Arctonyx, which

has been compared to that of a pig, resembles tolerably closely

the rhinarium of Meles.

The nose of Taaidea is less developed and less snout-like than

that of Meles. It does not overlap the lower jaw to the same

extent, and is not so deep from the summit of the rhinarium to

the edge of the upper lip. The rhinariwm itself also differs from

that of Meles in being covered above with hair nearly up to its

anterior edge, in having an anterior median groove, and in

* See my paper on Mellivora and Gulo (PR. Z.S. 1920, pp. 179-187).

yr Figured and described by Boas, Ohrknorpel der Séug. p. 150, pl. xxi. fig. 221

(1912). :

pe) |

OF THE EUROPEAN AND AMERICAN BADGERS. 4

Text-figure 19.

EE ZZ

NN, : LLLP i ff

Pe i,

—

A. Side view of head of Taxidea americana.

B. Rhinarium and upper lip of the same, from the front.

C. Rhinarium of the same, from above.

D. Side view of head of Meles meles.

E. Rhinarium and upper lip of the same, from the front.

F. Rhinarium of the same, from above.

(In B and E the rhinarium and upper lip are represented in the same plane,

so that the lip is not foreshortened.)

426 MR. Rk. 1. POCOCK ON THE BXTERNAL CHARACTERS

having a shallow infranarial portion on each side. As in Meles,

however, there is no philtrum, the upper lip being continuously

hairy and without median groove.

The Feet.

The fore feet of J/eles, as is weil known, are essentially

fossorial, the claws being of great length and far surpassing those

of the hind foot. The digits are united by integument be-

yond the proximal end of the digital pads, and are susceptible

only of shght separation. Digits 2, 3,4, and 5 are subequally

spaced, but digit 1 (pollex) is more widely separated, smaller, and

set farther up the foot than digit 5. The digital pads are not

well defined proximally, and the space between them and the

plantar pad is quite naked. The plantar pad is wide, as wide

approximately as the foot, and imperfectly four-lobed; the

pollical lobe is small. Behind the plantar pad there is a large

naked area, with a tuft of hair in the centre; and at the upper

or proximal end of this naked area lie two carpal pads, one on

each side and separated by a moderately wide space; the outer

of these two pads lies near the margin of the carpus and is

larger than the inner. They vary to a certain extent in size and

distinctness. ;

The hind foot is much narrower than the fore foot and has

much shorter claws. The Ist digit (hallux) is small and set

higher up the foot than digit 5, which is itself a little higher

than digit 2. The digits are only slightly separable, and are

webbed as in the fore foot, except that digits 3 and 4 are

closely united, the fusion sometimes extending to the very tip of

the digital pads, although usually these pads are separated to a

small extent at their distal ends. Asin the fore foot, the space

between the digital pads and plantar pad is quite naked, and the

plantar pad is large, as wide as the foot, and indistinctly lobed.

Behind it there is a large, naked, triangular area, pointed behind,

which is mostly covered by the two metatarsal pads, which are

sometimes separated in the middle line, sometimes fused, and

are separated from the plantar pad, at least in the middle, by a

narrower or broader naked area. Behind the metatarsal pads

the lower surface of the foot is covered with hair.

Hodgson’s illustrations* of the feet of J/eles lewewrus attest

their similarity to those of Meles meles, and unpublished

sketches of the feet of Arctonya by this author show that they

resemble the feet of MJeles in general features. Perhaps the

plantar pads are a little narrower and more decidedly trilobate,

and no inat of hair is shown on the area between the plantar and

carpal pads; but two carpal pads are shown on the fore foot

and two metatarsal pads in the centre of a naked area on the

* Journ. Asiatic Soc. Bengal, xvi. pl. 11. (1897). It may be noted that on this plate

the sketch of the hind foot of Helictis nipalensis is labelled Urva cancrivora, and

that of the latter is similarly labelled Helictis nipalensis.

§:

Are

OF THE EUROPEAN AND AMERICAN BADGERS. 427

hind foot, but this area is larger than in eles and the hairy area

up to the heel is shorter.

The fore foot of Vawidea resembles that of JJeles in general

Text-figure 20.

A. Right hind foot of Me/es eles.

B. Right fore foot of the same.

C. Right hind foot of Taxidea americana.

D. Right fore foot of the same.

x s

form, in the length and strength of the fossorial claws, and in

the nakedness of the area between the digital and plantar pads ;

498 MR. R. I. POCOCK ON 'THE EX'TERNAL CHARACTERS

but it differs in many structural details. The piriform digital

pads are much larger and better defined along their proximal

margin, and those. of the second, third, and fourth digits are

united by webbing extending past Me middle of each, these three

digits being closer together than the second is to the first or the

fourth to the fifth, the latter bemg nearly at the same level as

the first. Also the entire foot is wider as compared with its

length, and the plantar pad is much narrower and does not occupy

the whole width of the foot. It is very imperfectly divided into

four lobes. ‘The area behind it on the inner (pollical) side of the

foot is partially overgrown and overlapped by hairs; on the outer

side it is naked, and on the naked area a little way behind the

plantar pad but towards the middle line is a single, rather small,

hemispherical carpal pad, representing the inner or radial carpal

pad of Meles. This pad is partly overlapped and, according to

Coues, is sometimes overgrown by hair (‘ Fur-bearing Animals,’

p. 266).

Similai differences, so far as the larger size of the digital pads

and the greater width of the: ‘digital postion of the foot are con-

cerned, are observable between the hind feet of the two genera ;

but the third and fourth digits of Taaidea are not so closely

united, there being a definite, though narrow space between the

inner proximal ends of the pads. The plantar pad is very

different in ZVawidea. It is irregularly cordate in shape and

about as long as wide, and its lateral margins do not nearly

extend to the edges of the feet behind the first and fifth digits.

There is, moreover, no trace of metatarsal pads, the hairs of the

metatarsal area reaching down to the proximal margin of the

plantar pad.

The Anal and Genital Areas.

In Meles, as is well known, the anus is sunk in a shallow

depression, varying apparently to a certain extent in depth

according to the individual. Between this and the base of the

tail there is a deep subeandal pocket, partially divided into a

right and left deeper portion by a vertical partition. The

inferior margin of this pouch is a transverse lamina of integument,

forming the partition between it and the shallower circumanal

depression. The skin of the subcaudal pouch itself is hairy and

glandular *, and secretes copiously a sticky but not particularly

foul-smelling fluid which stains the surrounding integument and

hairs black. The true anal glands do not discharge directly into

this subeaudal pouch, but just within the orifice of the anus as

in all Mustelide. I have verified the existence of this pouch in

the Japanese Badger (JZ. anakuma), and, according to M. Edwards,

it is present in the Tibetan species (J/. lewewr us). It is also

present in the Oriental genus Arctonye, as recorded by Evans in

* As fully described by Chatin, Ann. Sci. Nat. (5) xix. pp. 106-109, pl. vii. figs.

66-67 (1874).

OF THE EUROPEAN AND AMERICAN EADGERS. 429

the following passage :—‘ [there] is a caudal pouch directly under

the origin of the tail,... but quite distinct from, and wholly

unconnected with, the anus or genital organs. The sac is formed

by duplicate folds of the common integument, having a lining of

naked membrane, secreting a brown unctuous matter, not unlike

cerumen, or wax of the ear” *.

Text-figure 21.

eee

eae a

NG ome

_A. Rear end of Mees meles, male, showing the subcaudal and anal pouches

distended nearly to the fullest extent.

B. The sane of the female, but with the pouches rather less distended

transversely.

Gairdner supplements this account as follows :—‘ Two scent

glands were found discharging into the postcaudal pocket. The

secretion was brownish yellow and the hind parts were stained

by the flow, and the stench so pervaded the beast that the coolies

were unable to eat it’ >.

In the male of J/eles the hairy scrotum is situated just below

the rim of the circumanal sac, which, except in the middle line,

is covered with short hairs. The baculum has been figured and

* Journ. Asiatic Soc. Bengal, viii. pt. i. p. 408 (1839).

+ Journ. Nat. Hist. Soc. Siam, i. no. 4, p. 253 (1915). From the passage quoted

it appears that the secretion of the glands of Arctonyx is much stronger in smell

than that of Meles. Meles has the habit, observable in Mongooses and Civets, with

analogous glands, of rubbing the secretion on objects so that the scent is dis-

seminated.

430 MR. R. I. POCOCK ON THE EXTERNAL CHARACTERS

described by Blumenbach* and Pohl?. It is about 4 inches

long and slightly inerassate at the base, flattened and grooved

beneath throughout its length and carinate above in its proximal

half, then flattened and depressed, with a median dorsal groove

up to the tip, which is straight or slightly upturned and expanded

laterally into a roughened dise with semicircularly curved free

margin. ‘This apex is perfectly symmetrical, and an elongated

slit perforating the bone behind the tip suggests that the latter

results from the fusion of two short terminal processes.

In the female the area around the genitalia is smooth; the

genital orifice is a little below the naked rim of the circumanal

sac, and opens at the summit of an inferiorly expanding groove,

which ends in an angular prepuce, forming a glandular space round

the small clitoris, which is strengthened with a small bone,

In Taxidea there is no trace either of the deep pouch imme-

diately beneath the tail or of the shallower depression in which

the anus is sunk. The anus, on the contrary, is protuberant,

and in profile view stands away from the base of the tail above

and from the perineal region below like a hemispherical mound #.

The anus opens just below the centre of this elevation, and the

two anal glands, about the size of a hazel-nut, open within the

orifice, the ducts traversing a definite papilla as in Mephitis.

The secretion is colourless with a sweetish, not unpleasant

musteline odour.

Below the anal prominence there is in the female a long naked

perineal area, terminating inferiorly in a piriform prominent

vulva, with the orifice above and a somewhat acuminate clitoris

below. On each side of the vulva, a little below the level of the

orifice, there is a glandular pocket about 6 mm. deep, from the

bottom of which arise a few sete, each planted in a shallow

pit.

Thus the anal and genital areas of the female Zaxidea differ

profoundly from those of Jeles §.

T have had no opportunity of examining a male Zawidea; but,

* Handbuch vergl. Anat. 1824, p. 476.

+ Jena. Zeitschr. xly. p. 385 (1909).

+ Coues’s statement (tom. cit. p. 267) that “the perineal region shows, imme-

diately beneath the root of the tail, a large transverse fissure leading into the

peculiar subcaudal pouch of the Meline”’ is erroneous; and the error arose probably

trom the examination of dried skins, which were apparently all the material available

for examination, judging from the bottom paragraph on p. 68 of the volume cited.

§ It is possibie, however, that the ditference in the size and situation of the

genital orifice in the specimens examined may be more apparent than real. The

examples of IMJeles were wild caught animals, one of which was known to have

produced young before capture. ‘The example of Tawidea, on the contrary, was

received from New York as an adult specimen in 1910, and died, when an old

animal, in Dec. 1918. Of her history previous to her arrival in London I know

nothing, but she never bred nor was seen to pair with the mate after coming to the

Gardens; and it may be that the small size and low position of the genital orifice

and the consequent length of the perineal area are attributable to failure of copulation

and parturition,

OF THE EUROPEAN AND AMERICAN BADGERS. 431

according to Coues, there is a well-developed baculum. He

describes it as “4 inches long, clubbed at one end, compressed,

and with a shallow sulcus in the continuity ; the other end bent

Text-figure 22,

So NG

A. Rear end of Meles meles, female, with the subcaudal and anal sacs closed.

xe.

B. The same of Taxidea americana, female. X 5.

C, Lateral view of ano-genital area of Tavidea, female, showing the prominent

anus and the clitoris with its lateral glandular pit partly opened. X $.

D. The lateral gland of the clitoris of the same, opened to show the set at the

bottom.

KE. Clitoris of same, elevated to show the glandular pits closed.

F, Anus of same, spread open to show the papille of the anal ¢lands.

432 MR. R, I, POCOCK ON THE EXTERNAL CHARACTERS

nearly at a right angle, abruptly and irregularly flattened and

grooved ” (tom. ct. p. 269).

This description is not very intelligible, and it is doubtful if

the describer knew either the proxim: al from the distal extremity

or the dorsal from the ventral surface; but I infer that the bone

is compressed, grooved throughout its extent below, thickened at

the base and hooked at the apex, but whether the curvature of

the hook is directed upwards or downwards does not appear ;

and whether the apex is symmetrical or asymmetrical is also

unknown.

Skull and Teeth.

The skull of Meles meles was fully described and illustrated by

Miller; that of YVaaidea was figured and described by Coues.

Text-figure 23.

aaacat

A. Upper view of the skull of Taxridea. X 3 approx.

B. The same of eles.

Elliot also reproduced photographs of it *, and Baird pointed out

some of the differences between the two genera in the crania and

teeth.

* Wield Columb. Mus, ii. p, 320 (1901).

OF THE BUROPHAN AND AMERICAN BADGERS.

433

In the following table the principal differences are placed side

by side for comparison :—

MELEs.

Muzzle

Zygomata

“Blongated, comparatively uar-

|Stronely salient behind orbit |

row, with prominent pre-

Treille! Intraorbital fora-

men large. above the anterior

portion of the upper molar

and behind the carnassial.

Short and broad,

TAXIDEA.

with short

premaxille. | Infraorbital

foramen small, above the

the posterior base expanded

laterally and posteriorly con-

siderably beyoud glenoid.

| anterior portion of the

| upper carnassial.

Moderately salient behind

orbit; its posterior base not

expanded laterally and pos-

teriorly beyond glenoid.

Brain-case.....)...

Occipital area ... |

Upper surface sloping posteri-

orly ; lateral walls rounded,

converging behindzygomata.

Sagittal crest high.

Much narrower than zygomatic

width. Mastoids compara-

tively narrow and elongated,

aes downwards and for-

rards beneath the auditory

meatus and lower than the|

glenoid. — Basioecipito-sphe- |

noidal plane inclined up-

wards from foramen. |

Moderately

inflated, ssarcely

below the plane of the occi-

pital condyles and uot ex-

tending torwards to the

glenoid.

Upper surface hardly sleping

posteriorly; lateral walls

gradually divergent from

orbits to occiput. Sagittal

crest LO or absent.

Almost as wide as zygomatic

width. Mastoids greatly

expanded but short, not

projecting below auditory

meatus and about on a leve’

with the elenoid. Basiocci-

pito-sphenvidal plane hori-

zontal.

Much inflated, a long way

Lelow the plane of the occi-

pita! condyles and abutting

against the glenoid.

Rr OTAMINGE 2.2 ...2.. |

For. vot. concealed and open-

ing alongside fur. lac. ant. ;

for. ov. considerably in ad-

vance of fur. lac. med.

For. vot. not concealed, open-

ing separa'ely from and be

neath for. lac. ant.; for. ov

just in front of for lac.

med.

Teeth

| Upper

carnassial comment,

tively small. le-s than 3 area

of melar, Molar irregularly

four-sided, with two roots

imbedded in cheek and two

large cusps exposed in lateral

view of skull; crown with

one median longitudmal

ridee of cusps. Te car-

nassial with heel ahout as

large as the anterior portion,

and hollowed im the middle

with two external and two

iniernal but no median

cusp; in the anterior portion

of the tooth the anterior

cusp is smaller than the main

cusp, which is well im ad-|}

vance of the inner cusp.

Upper carnassial enormous.

larger than molar. olay

equilaterally triangular.

with one root imbedded im

cheek and one cusp exposed

in lateral view of the skull.

Crown with two BES WISIENK

rows of tubercles. Lowe

carnassial with eel about

3 the area of the anterio

portion, with one external

one inedian, and one pos:

terior cusp forming a trans

verse line; in the anterio

portion of the tooth th.

anterior cusp Is as large a:

the mam cusp, which is m

the same transverse line as

the inner cusp.

434 MR. R. I. POCOCK ON THE EXTERNAL CHARACTERS

Text-figure 2

Taxides.

A. Posterior view of the skull of Tawidea. X % approx.

C. The same of eles.

B. Inferior view of posterior portion of skull of Tawidea (f.0., foramen

ovale).

D. The same of Jfeles.

Conclusions.

Tn view of the nature and number of the differences between

Meles and Tawidea in skull and teeth, it seems no exaggeration to

say that the resemblances between the genera in those particulars

are only such as entitle them to a place in the family Mustelide.

Unquestionably the skull of Z'aaidea presents a greater likeness

to that of Mellivora than to the skull of J/eles ; but it is, in my

Pee ee ee

OF THE EUROPEAN AND AMERICAN BADGERS. 435

opinion, by no means certain that this likeness involves close

aftinity, since the two genera differ considerably in the structure

of the two posterior maxillary teeth and in the development of

the pinna of the ear, of the pads on the feet, etc.

Text-figure 25.

A. Posterior maxillary teeth of Taxidea. Nat. size.

C. The same of Meles.

B. Posterior mandibular teeth of Tawxidea.

D. The same of eles.

Pending an examination of Mydaus and Helictis*, which I

have not seen, I propose to restrict the subfamily Meline to the

genera JJeles and Arctonyx. With these limitations the JJeline

* This genus, as already stated, was severed from the Meline both by Gray and

Gill.

Proc. Zoou. Soc.—1920, No. X XIX. 29

436 ON THE EUROPEAN AND AMERICAN BADGERS.

may be briefly distinguished as follows from the Taxiidine, a new

group which, for the present, contains Taaidea alone :—

a. A well-developed subcaudal pouch; rhimarium with very deep

infranarial area ; plantar pads wide, carpal and metatarsal pads

comparatively large, the latter on a naked area behind the

plantar pad; upper carnassial much smaller than quadrilateral

molar; lower ecarnassial with enormous heel etc. .................. MWeline.

b. No subcaudal pouch; rhinarium with shallow infranarial area ;

plantar pads narrower; carpal pads much reduced, hind foot

hairy down to plantar pad, metatarsal pads suppressed ; upper

carnassial larger than triangular molar; lower carnassial with

Govier memnnrely SoMa WES Ss —soaonscoosooseaannonnsabonaacanqnanssoocns LleBgaidorGe

SEXUAL DISPLAY AND NESTING-HABITS OF THE ostricn. 437

EXHIBITIONS AND NOTICES.

May 11th, 1920.

Prof. J. P. Hitt, F.R.S., Vice-President, in the Chair.

The Secretary read the following Report on the Additions

to the Society’s Menagerie during the month of April, 1920 :-—

The registered additions to the Society’s Menagerie during the

month of April were 118 in number. Of these 39 were acquired

by presentation, 15 were deposited, 28 were purchased, 32 were

received in exchange, and 4. were born in the Menagerie.

The following may be specially mentioned :—

1 Chimpanzee (Anthropopithecus troglodytes), from West Africa,

purchased on April 21st.

2 White-handed Gibbons (Hylobates lar), from Rangoon,

received in exchange on April 15th.

2 Barbary Sheep (Ammotragus lervia), born in the Menagerie

on April 11th.

1 Milky Eagle-Owl (Bubo lacteus), from Rhodesia, purchased

on April 19th.

1 Banded Rail (Hypotenidia striata), from India, new to the

Collection, purchased on April 28th.

1 Five-banded Lizard (Mabuia quinqueteniata), from Rhodesia,

new to the Collection, deposited on April 18th.

On behalf of Messrs. EB. Gerrard & Sons, Mr. R. I. Pocock,

F.R.S., exhibited » mounted specimen of a pale variety of the

White-bearded Gnu (Connochetes albojubatus), shot by Capt.

Keith Caldwell, R.A., F.Z.S., in Masailand, and pointed out that

apart from the general pale yellowish-brown tint, the variation

affected different parts of the body in different ways, the neck-

mane, the long hairs on the face, and the tail-tuft, which are

normally black, being dirty white, whereas the bands on the

body, making the brindled pattern, which are also normally

black, were brownish red.

Miss J. B. Procrer, F.Z.S., exhibited and made remarks upon

a living specimen of the tailed Batrachian, Spelerpes fuscus

Bonaparte, born on May 8th, 1920.

Prof. J. EK. Durrprn, F.Z.8., exhibited and made remarks

upon a remarkable series of lantern-slides illustrating the sexual

display and nesting-habits of the Ostrich.

438 THE SECRETARY ON ADDITIONS 'lO THE MENAGERIE,

June 1st, 1920.

Sir Srovny F, Harmer, K.B.E., F.R.S., Vico-President,

in the Chair.

In the absence of Prof. R. 'T. Lerpsr, F.Z.8., his exhibition of

Jantern-slides illustrating the Experimental transmission of some

Helminth infections, was deseribed by Dr. Vevers. Specimens

of Schilbe mystus, Olarotes laticeps, and Tilapia ailotica infected

with encysted trematode Jarve, and found near Cairo, were

shown. By feeding young wolves, bred in captivity, with these

fishes Prof. Leiper sueceeded in rearing the adult worms in

enormous numbers. The cysts of Schilhe mystus gave rise to

Hemistoma (Alaria) alatun of the dog, and those of Clarotes

laticeps to Monostoma pumilio Looss, i. pwmilio, which would

appear to be more closely related to Zocotrema than to Monostoma,

is normally a parasite of the Pelican and Kite in Kgypt. That

it can be reaved in enormous numbers in Wolves undermines the

conception of “ plysiological species” applied by Looss to certain

trematode infections.

Dr. P. Coatmers MrrcHext, F.R.S., gave an account, illustrated

with Jantern-siides, of his recent Aeroplane Trip from Catro to

Tabora, and deseribed the character of the country passed over

and the birds and mammals seen.

June 15th, 1920.

Prof. EK. W. MacBrips, F.R.S., Vice-President,

in the Chair.

The Secretary read the following Report on the Additions to

the Society’s Menagerie during the month of May, 1920 :—

The registered additions to the Society’s Menagerie during the

month of May were 398 in number. Of these 63 were acquired

by presentation, 188 were deposited, 130 were purchased, 4 were

received in exchange, and 13 were born in the Menagerie.

The following may be specially mentioned :—

MAMMALIA.

1 Cheetah (Cynelurus jubatus), from Kilosso, Tanganyika

Territory, presented by C. MacMahon, Esq., Assistant Political

Officer of Kilosso, on May Ist.

2 Capybaras (Hydrocherus hydiocherus), bred in England,

purchased on May 12th.

1 Arabian Oryx (Oryx lewcorya), from Central Arabia, depo-

sited by H.M. The King on May 14th.

ON THE LIFE-HISTORY OF THE DRAGONFLY. 439

AVES.

2 Somali Ostriches (Struthio molyidophanes), purchased on

May 8th.

2 White Rheas (Rhea americana), purchased on May 12th.

1 Kagu (Rhinochetus jubatus), from New Caledonia, purchased

on May 4th.

4 Long-tailed Shrikes (Urolestes melanoleucus) and 1 Black-

collared Barbet (Lybius torquatus), from South Africa, deposited

on May 22nd. New to the Collection.

4 Isabelline Turtle-Doves (Zurtur isabellina), 2 Rosy-grey

Turtle-Doves (Turtur vroseigrisews), € Dongola 'Turtle-Doves

(Turtur decipiens), from North-East Africa, deposited on May Ist,

and 2 Spotted-bellied Francolins (francolinus spilogaster), pre-

sented by Major Maurice Portal, F.Z.S., on May Ist. All new

to the Collection.

REPTILIA.

1 Siamese Crocodile (Crocodilus siamensis) and 2 Tentacled

Snakes (/erpeton tentacultwm) from Siam, the latter new to the

Collection, presented by Dr. Malcolm Smith, F.Z.8.

Prof. J. H. Durrpen, F.Z.8., exhibited and made remaiks upon

a sertes of Ostrich eggs.

broke do. Tmovarp, M.A. D.Sc, Fos) bsi.S:, gave an

account, illustrated by lantern-slides, of 'The Life-history of the

Dragonfly, with special reference to Australian forms Dr,

Tillyard dealt first with the structure of the female ovipositor,

and showed the correlation between the habit of laying eggs in

the tissues of plants and the elongated form of the eggs, on the

one hand, and that of laying them freely in the water, the eggs

in this case being of a much more rounded form, ‘I he develop-

ment of the embryo and the hatching of the larva were next

dealt with; the creature that hatches from the egg is not an

active larva, but a sheathed pronymph, whose existence lasts but

a few seconds, and from which the active young larva emerges in

its turn, representing actually the second larval instar of other

insects. The various types of larve found in the two suborders

Anisoptera and Zygoptera were next shown, and a series of

slides dealt with the interesting larval specialisations in the

gizzard, the prehensile labial mask, and the rectal and caudal

ells.

No. 205.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

May 11th, 1920.

Prof. J. P. Hitt, F.R.S., Vice-President,

in the Chair.

The Secretary read a Report on the Additions to the Society’s

Menagerie during the month of April, 1920.

On behalf of Messrs. HE. Gerrard & Sons, Mr. R. I. Pococr,

F.R.S., exhibited a mounted specimen of a pale variety of the

White-Bearded Gnu (Connochetes albojubatus), shot by Capt.

Keith Caldwell, R.A., F.Z.S.,in Masailand, and pointed out that

apart from the general pale yellowish-brown tint, the variation

affected different parts of the body in different ways, the neck-

mane, the long hairs on the face, and the tail-tuft, which are

normally black, being dirty white, whereas the bands on the

body, making the brindled pattern, which are also normally

black, were brownish red.

Miss J. B. Procror, F.Z.S., exhibited and made remarks on a

living specimen of the tailed Batrachian, Sipelerpes fuscus Bona-

parte, born on May 8th, 1920.

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent’s Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Hellows who subscribe to the Publications ; but it may be obtained on the

day of publication at the price of Sixpence, or, if desired, sent post-free for

the sum of Six Shillings per annum, payable in advance.

Prof. J. E. Durrpen, F.Z.S., exhibited and made remarks on

a remarkable series of lantern-slides illustrating the sexual dis-

play and nesting-habits of the Ostrich.

Dr. W. J. Daxry, F.Z.8., communicated his paper on “ Fauna

of Western Australia.—lII. Further Contributions to the Study

of the Onychophora”; and illustrated his remarks by lantern-

slides and microscopic specimens, showing points in the structure

of Peripatus.

Mr. C. Forstzr-Coorrer, M.A., F.Z.8., gave a résumé of his

paper ‘‘ Chalicotheroidea from Baluchistan,” illustrated by speci-

mens and lantern-slides.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, June Ist, 1920, at 5.30 p.m., when the

following communications will be made :—

Dr. G. M. Vevers, F.Z.S.

Report on the Entozoa collected from animals which died

in the Gardens during the past nine months.

Prof. R. T. Leper, F.Z.S.

Exhibition: Experimental transmission of some Helminth

infections.

Dr. W. T. Catman, F.Z.S.

Notes on Marine Wood-boring Animals.—I. The Shipworms

(Teredinide).

The SECRETARY.

Notes on an African Trip, with lantern illustrations.

The Publication Committee desire to call the attention of

those who propose to offer Papers to the Society, to the great

increase in the cost of paper and printing. This will render it

necessary for the present that papers should be condensed and

be limited so far as possible to the description of new results.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Society OF Lonpon,

Recent’s Park, Lonpon, N.W. 8.

May 18th, 1920.

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No. 206.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

June ist, 1920.

Sir Sripney F. Harmer, K.B.H., F.R.S., Vice-President,

in the Chair.

Dr. G. M. Vevers, F.Z.S., communicated his “‘ Report on the

Entozoa collected from Animals which had died in the Society’s

Menagerie during the past Nine Months.”

In the absence of Prof. R. T. Lerprer, F.Z.S., his exhibition of

lantern-slides illustrating the Experimental transmission of some

Helminth infections, was described by Dr. VEVERs.

Dr. W. T. Cauman, F.Z.S., gave a résumé of his paper ‘“‘ Notes

on Marine Wood-boring Animals.—I. The Shipworms (Tere-

dinidee).”

Dr. P. Coatmers MitcHet1, F.R.S., gave an account, illustrated

with lantern-slides, of his resent Aeroplane Trip from Cairo to

Tabora, and described the character of the country passed over

and the birds and mammals seen.

* This Abstract is published by the Society at its offiees, Zoological Gardens,

_ Regent’s Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications ; but it may be obtained on the

day of publication at the price of Sixpence, or, if desired, sent post-free for

the sum of Six Shillings per annum, payable in advance,

The next Meeting of the Society for Scientific Business will be

held on Tuesday, June 15th, 1920, at 5.30 p.m., when the

following communications will be made :—

The SECRETARY.

Report on the Additions to the Society’s Menagerie during

the month of May, 1920,

Prof. J. EH. Durrpen, F.Z.S.

Exhibition of, and remark on, Ostrich eggs.

Miss Joan B. Proctsr, F.Z:8.

1. On a Collection of Tailless Batrachians from East Africa

made by Mr. A. Loveridge in the Years 1914-1919.

2. On the Type-Specimen of Rana holsti Boulenger.

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

‘On the External and Cranial Characters of the European

Badger (J/eles) and the American Badger (Tuaidea).

R. J. Trtyarp, M.A., F.LS.

The Life-History of the Dragonfly.

The following Papers have been received :—

J. H. Luovp, M.Sc., F.Z.8.

Some Observations on the Structure and Life-History of the

common Nematode of the Dogfish (Scylliam canicula).

‘

Wm. A. Cunnineton, M.A., Ph.D., F.Z.8.

The Fauna of the African Lakes ; a Study in Comparative

Limnology, with special reference to Tanganyika.

W.N. F. Wooptanp, D.Sc., F.Z.S.

On some Results of ligaturing the Anterior Abdominal Vein

in the Indian Toad (Lu/o stomaticus Lutkin).

The Publication Committee desire to call the attention of

those who propose to offer Papers to the Society, to the great

increase in the cost of paper and printing. This will render it

necessary for the present that papers should be condensed, and

be limited so far as possible to the description of new results.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Society or Lonpon,

Reegent’s Park, Lonpon, N.W. 8.

June 8th, 1920.

No. 207.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON,*

June 15th, 1920.

Prof. EH. W. MacBrips, F.R.S., F.Z.S., Vice-President,

in the Chair.

The Secrerary read a Report on the Additions to the Society’s.

Menagerie during the month of May, 1920.

Prof. J. KE. Duerpen, F.Z.S., exhibited and made remarks upon

a series of Ostrich eggs.

Miss Joan B. Proctor, F.Z.8., gave a réswmé of her papers.

(1) ‘*On a Collection of Tailless Batrachians from Kast Africa

made by Mr. A. Loveridge in the Years 1914-1919,” and

(2) ‘On the Type-specimen of Kana holstt Boulenger.”

Mr. R. I. Pocock communicated his paper ‘‘ On the External

and Cranial Characters of the European Badger (eles) and the

American Badger (Tawidea).

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent’s Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications ; but it may be obtained on the

day of publication at the price of Simpence, or, if desired, sent post-free for

the sum of Sta Shillings per annum, payable in advance.

Dr. R. J. Trutyarp, M.A., F.L.S., gave an account of the

Life-History of the Dragonfly, with special reference to Austra-

lasian forms, and illustrated his remarks by a series of lantern-

slides.

This Meeting closes the Session 1919-1920. The next Meeting

of the Society for Scientific Business will be held on Tuesday,

October 19th, 1920, at 5.30 p.m. A special notice will be

circulated early in October.

The following Papers have been received :—

J. H. Luoyp, M.8c., F.Z.8.

Some Observations on the Structure and Life-History of the

common Nematode of the Dogfish (Scyllium canicula).

Wm. A. Cunnineton, M.A., Ph.D., F.ZS.

The Fauna of the African Lakes; a Study in Comparative

Limnology, with special reference to Tanganyika.

W.N. F. Wooptanp, D.Sc., F.Z.S.

On some Results of ligaturing the Anterior Abdominal Vein

in the Indian Toad (Bufo stomaticus Lutkin).

Haru Ram Meura, M.Sc.

On the Sexual Phase in certain Indian Naididz (Oligocheia).

The Publication Committee desire to call the attention of

those who propose to offer Papers to the Society, to the great

increase in the cost of paper and printing. This will render it

necessary for the present that papers should be condensed, and

be limited so far as possible to the description of new results.

Cominunications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZooLocica Socrery or Lonpon,

Recent’s Park, Lonpon, N.W. 8.

June 22nd, 1920.

15.

16.

Lig

18.

19,

20.

21.

22.

25.

PAPERS.

The Life-History and Habits of two Parasites of Blow-flies. By A.M. Anrson. With

an Introduction by Prof. H. Maxwexu Lerroy, F.Z.8. (Text-figures 1-20.) ......

A Revision of the Nematode Family Gnathostomide. By H. A. Baynis, M.A. Oxon.,

Assistant in the Department of Zoology, British Museum (Natural History), and

Crayton Lanz, M.D.Lond., Lt.-Col. I.M.S. (ret.). (Dext-figures 140; Plates I-

TPIS) ae SAU a a a Aa ar eg Age tts ct ae Yet

Contributions to a Study of the Dragonfly Fauna of Borneo.—Part IV. A List of

Species known to occur in the Island. By F. F, Larpuaw, M.A. (Oantab.), F.Z.8.

RSet hon eay At pe aero gweia ah aicaiala'e ale wcin So sin alsa cls ao Ot aEM ete una a Maat aimee, eG

On some new Therocephalian Reptiles from the Karroo Beds of South Africa. By

Re-BRooM, M.D. DSe., HRS., O.M.Z.8. (Text-figures 1—9:) 2.) S).c5. cece se ees

Chalicotheroidea from Baluchistan. By C. Forstar-Ooormr, M.A., EZS., Super-

intendent of the University Museum of Zoology, Cambridge. (Plate I. and Text-

EWG S17) ge Nene eee AG anne DEERE oe Om naa eR Ean GUce ac cee ao eemee

Page

245

311

307

Fauna of Western Australia.—III. Further Contributions to the Study of the |

Onychophora. The Anatomy and Systematic Position of West Australian Peripa-

‘toides, with an account of certain histological details of general importance in the

study of Peripatus. By Wm. J. Dakin, D.Sc., F.Z.S., F.L.S8., Professor of Biology,

University of Western Australia. (Plates I.-V.) ..... RANE NCiC NTRS te Mee art ee

Notes on Marine Wood-boring Animals.—I. The Ship-worms (Teredinide). By

W. T. Cauman, D.Sc. (Text-figures 1-11.) ........4.. RE SUIT cant eae e

Report on Entozoa collected from Animals which died in the Zoological Gardens of

London during Hight Months of 1919-1920. By G. M. Vevzrs, M.R.C.8., L.R.C.P.,

‘F.Z.8., Beit Memorial Research Fellow, Demonstrator in Helminthology at London

School of Tropical oe and Hon. Parasitologist to the Zoological Society of

Ibe OM op gide,.cb de Gentes SO tag UA UM EA MOn am cena noon bop SCO uuunse de oo

. On a Collection of Tailless Batrachians from Hast Africa made by Mr. A. Loveridge

in the years 1914-1919. By Miss Joan B. Procter, F.Z.S. (Text-figures 1-4.)

On the Type-specimen of Rana holst Boulenger. By Miss Joan B. Procrzr, BR Z.8.

(UES TETRIS) Ota Bc GAH s Be Nn SIA ein EA RMe CEO ERO. OO OD DAT ea omni cerets

On the External and Cranial Characters of the European Badger (Meles) and of the

_American Badger (Tawidea). By R. I. Pocock, F.R.S. (Text-figures 19-25.)

405

41]

PLATES.

1920, Parr III. (pp. 195-439).

Plate

Bayurs and Lane: 1.)

oS } GiimthoshOmid ey sfc s oi veces ac oie ere oe

VI.

VII.

VIII. /

Forster-Coorer : I. Chaliaotherciden from Baluchistan .

Daxin : He)

iT] :

TIL. oe Australian Peripatoides

v.)

Page

3oT

NOTICE.

The ‘ Proceedings’ for the year are issued in fowr parts, paged consecutively,

so that the complete reference is now P. Z. 8. 1920, p.... The Distribution

is usually as follows, but on account of abnormal conditions Parts I. & II. are

issued together :—

Part I. issued in March.

SS aiadale * June.

yoo LI september;

LVR December,

‘ Proceedings, 1920, Parts I. & IT. (pp. 1-194), were published

together on July 21st, 1920.

The Abstracts of the ‘ Preceedings,’ Nos. 205-207, are

contained in this Part.

PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

ZOOLOGICAL SOCIETY

OF LONDON. | a

1920. (a0m 14 1991 >

& .

“oral mus av ie

PART IV.

contarntine Paces 441 to 656, witn 40 TExt-FiGuRES.

DECEMBER 1920.

PRINTED FOR THE SOCIETY,

SOLD AT ITS HOUSE IN REGHNT'’S PARK.

LONDON :

MESSRS. LONGMANS, GREEN, alt CO.,

PATERNOSTER ROW.

[Price Twelve Shillings. |

LST OR CON TENTS,

1920, Parr 1V. (pp. 441-656).

EXHIBITIONS AND NOTICES.

Page

The Sucretary. Report on the Additions to the Society’s Menagerie during the months —

of June, July, August, and September, 1920 .. ....c eee ce ee ee eee eee eee eee 653

Dy, P. Craters Mitcunit, O.B.H., M.A., LL.D., D.Se., FE.R.S. Exhibition of, and

remarks upon, a Double-tailed Lizard .......... 0s. e ee cee ee ce eee eee eee ee | 655

Mr. R. I. Pococx, F.R.S. Exhibition of, and remarks upon, the skin of the groin of

Tragelaphus bwxtoni csv. s cere cece ce cece teen ee eee cece eees eee eee nee eens 655

Mr. E. G. Bouzenesr, F.Z.8. Exhibition of, and remarks upon, living specimens of

Necturus.. 0. reece see e nce e eee eee eer e ee eee ees wi eieileidate foialeke a cele eae 655

Mr. T. A. Barns, F.Z.S. An account of his recent journey through the Forests of Africa

in search of Gorilla and Okapi 2... sce cece cece ee ne ee ee estes eee poe Ge 656

The Sucrerary. Report on the Additions to the Society's Menagerie during the month of

Oatober, TOQO. 2S coe asks tate mite wteiote Ramevelevertoaiagela, alelelleiace Wisje cle cline tiniest ae COED

Mr. J. T. Cunninquam, M.A., F.Z.S. Exhibition of, and remarks upon, a specimen of the

Leech Trocheta, recently found in the Society’s Gardens ............+--++++-+00.. 656

Mr. F. Martin Duncan, F.Z.8. Exhibition of, and remarks upon, a series of Cinemato-

graph Films of Animals in the Society’s Gardens ......s. ee sess eee sees eee ee 656

Contents continued on page 3 of Wrapper.

ZOOLOGICAL SOCI

KTY OF LONDON.

Yurs Society was founded in 1826 by Sir Sramrorp Rarriss,

Mr. J. Sasine, Mr. N. A. Vigors,

and other eminent Naturalists,

fer the advancement of Zoology and Animal Physiology, and for the

introduction of new and curious subjects of the Animal Kingdom,

and was incorporated by Royal Charter in 1829.

Patr

HIS MAJESTY

COUN

On.

THE KING. fan

OIL.

HIS GRACH THE DUKE OF BEDFORD, K.G., E.R.S., Provitour Lis

Tas Hon. Cucit Barina, M.A.

Atrrep H. Cocks, Esa., M.A.

Lr.-Cot. 8. Monexron-Corrman,

M.D., F.R.S.

Cuartrs Droummonp, Hsa.,

Treasurer.

Hueu 8. Gransronu, Ese., M.A.,

F.LR.S.E.

Sir Sipyey F. Harmer, K.B.E.,

Mew DSc. 7H Ris. Vice-

President.

Pror. Jamzus P. Hitt, D.Sc.,

F.RAS., Vice-President.

Wittiam Hunrsman, Ese.

Pror.HanustW. MacBripg,).sc.,

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

Cot. Sin Haney McManon,

G.C MG Kee:

H. G. B. Meapu-Wanpo, Hse,

Vice-President.

P. Cmatmers Mrrcnent, Hse,

CBE. MAS DiSe:, Jul De

F.RS., Secretary.

Tur Hart or Onstow, O.B.E.

Masor Atperr Pam.

Aprian D, W. Pottock, Hse,

H.G. Tar Duxeor Rurtann, K.G. |

Tus Marquis or Suigo, F.S.A.,

Vice-President.

Masor Ricowarp S. Taytor.

A. Trevor-Barrys, Hse., M.A.

Awrpony H. Winertezp, lise.,

Vice-President.

The Society consists of Fellows, and Honorary, Foreign, and

Corresponding Members, elected according to the By-Laws. It

earries out the objects of its foundation by means of its collection

of living animals, by its Library, and by its Scientific Publications.

The Office of the Society, Regent’s Park, N.W.8, where all com-

munieations should be sent, addressed to “The Secretary,” is open

from Ten till Five, except on Saturdays, when it closes at OnE p.m.

The Library, under the superintendence of Mr. F. Martin Duncan,

F.Z.8., F.R.M.S. is open daily (except Sunday) from Ten a.w, till

Five p.m.; on Saturdays, Ten a.m. till One p.m.

The Library is closed from Good Friday to Easter Monday, and

upon all other Bank Holidays. It is also elosed annually for

cleaning purposes during the whole month of September.

The Meetings of the Society for General Business are held in

the Meeting Room at the Society’s Office on the third Wednesday

of the month at 4.30 p.m. except in September and October.

The Meetings for Scientific Business are held in the Meeting

Room at the Society’s Office fortnightly on Tuesdays, except in

July, August, September, and December and January, at half-past

Five o'clock p.m.

The Anniversary Meeting is held on the 29th of eel or the

nearest convenient day, at Four p.m.

The Society’s Gardens are open daily from Nine o’clock until

Sunset. Mr. R. I. Pocock, F.R.S., F.L.S., is the resident Super-

intendent and Curator of Mammals, Mr. D. Seth-Smith is Curator

of Birds and Inspector of Works, Mr. EH. G. Boulenger is Curator of

Reptiles, Miss L. K.Cheesman, F.E.S., is Curator of Insects. Appli-

cafiens for anatomical material or facilities for work in the

Prosectorium should be addressed to Dr. R. T. Leiper, Director of

the Society’s Prosectorium.

TERMS FOR THE ADMISSION OF FELLOWS.

Frrirows pay an Admission Fee of £5, and an Annual Contri-

bution of £3, due on the Ist. of January, and payable in advance,

or a Composition of £45 in lieu thereof; the whole payment,

including the Admission Fee, being £50.

No person can become a Frttow until the Admission Fee and

first Annual Subscription have been paid, or the annual payments

have been compounded for.

Fruitows elected in November and December are not liable for

the Subscription for the vear in which they are elected.

PRIVILEGES OF FELLOWS.

Frttows have Personal Admission to the Gardens upon signing

their names in the book at the entrance gate, and may introduce

Two Companions daily.

The Wire or Hussanp of a Fetrow can exercise these privileges

in the absence of the Fellow.

Until further notice, Frtnows will receive 40 undated Green

Cards, available on any Sunday or week-day up to the end of

February of the year following the year of issue, and 20 White

Cards available on any week-day up to the same date. Twenty

of the Green Cards may be exchanged for a book containing two

Orders for each Sunday in the year. ‘Twenty White Cards may

be exchanged for a book of dated Week-day Orders, each Order

available for any day during the week except Sunday. Special

ehildren’s tickets are no longer issued, but the Green and White

Cards are perforated, and each half is valid for a Child under twelve

years of agé. It is particularly requested that Fellows will sign

every ticket before it goes out of their possession. Unsigned tickets

are not valid.

Frttows are not allowed to pass in friends on their written

order or on presentation of their visiting cards.

Frtrows have the privilege of receiving the Society’s ordinary

Publications issued during the year upon payment of the additional

Subscription of One Guinea. This Subscription is due upon the

Ist of January, and must be paid before the day of the Anniversary

Meeting, after which the privilege lapses. Frntows are likewise

entitled to purchase these Publications at 25 per cent. less than

the price charged to the public. A further reduction of 25 per

cent. is also made upon all purchases of Publications issued prior

to 1881, if above the value of Five Pounds.

Fetiows also have the privilege of subscribing to the Annual

Volume of ‘The Zoological Record,’ which gives a list of the Works.

and Publications relating to Zoology in each year, for the sum of

One Pound Ten Shillings. Separate divisions of volumes 39 onwards.

can also be supplied. Full particulars of these publications can

be had on application to the Secretary.

Fentows may obtain a Transrerasie Ivory Ticknr admitting

two persons, available throughout the whole period of Fellowship,

on payment of Ten Pounds in one sum. A second similar ticket

may be obtained on payment of a further sum of Twenty Pounds.

Any Fritow who intends to be absent from the United Kingdom

during the space of at least one year, may, upon giving to the

Secretary notice in writing, have his or her name placed upon the

“dormant list,” and will then be called upon to pay an annual

subscription of £1 only during such absence, but after three years

must make a further application to be retained on that list.

Any Frrntow, having paid all fees due to the Society, is at liberty

to withdraw his or her name upon giving notice in writing to the

Secretary.

Ladies or Gentlemen wishing to become Fellows of the Society

to)

are requested to communicate with “The Secretary.”

P. CHALMERS MITCHELL,

Secretary.

Regent’s Park, London, N.W. 8.

December, 1920.

MEETINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON

FOR

SCIENTIFIC BUSINESS.

1921.

TUESDAY, FEBRUARY ...... 8 and 22.

MARCH ......... 8 and 22.

Be ZACP RIL Aaa nee Dd and 19.

i NAY os eens 10 and 24.

bi JUNE cleo. ft

The Chair wiil be taken at half-past Five o'clock precisely.

ZOOLOGICAL SOCIETY OF LONDON.

LIST OF PUBLICATIONS.

Tne scientific publications of the Zoological Society of London

are of two kinds—“ Proceedings,” published in an octavo

form, and “ Transactions,” in quarto.

According to the present arrangements, the ‘‘ Proceedings”

contain not only notices of all business transacted at the sciea-

tific meetings, but also all the papers read at such meetings

and recommended to be published in the ‘‘ Proceedings” by

the Committee of Publication. A large number of coloured

plates and engravings are issued in the volumes of the

“ Proceedings,”’ to illustrate the new or otherwise remark-

able species of animals described therein. Amongst such

illustrations, figures of the new or rare species acquired in a

living state for the Society's Gardens are often given.

The “Proceedings”? for each year are issued in four parts,

paged consecutively, during the months of March, June,

September, and December. From January 1901 they have

been issued as two half-yearly volumes, indexed separately.

An “ Abstract of the Proceedings’ is published by the

Society on the Tuesday following the date of the Scientific

Meeting to whichit refers. It is issued along with the “‘ Pro-

ceedings,” free of extra charge, to all Fellows who subseribe to

the Publications, but it may be obtained on the day of publi-

cation at the price of Sixpence, or, if desired, sent post free

for the sum of Six Shillings per annum, payable in advance.

The ‘‘ Transactions” contain such of the communications

made to the Scientific Meetings of the Society as, on account of

the nature of the plates required to illustrate them, are better

adapted for publication in the quarto form. They are issued

at irregular intervals.

Fellows and Corresponding Members, upon payment of

a Subscription of One Guinea before the day of the Anni-

versary Meeting, are entitled to receive the Society’s

Publications for the year. They are likewise entitled to

purchase the Publications of the Society at 25 per cent. less

than the price charged to the Public. A further reduction

of 25 per cent. is made upon purchases of Publications

issued prior to 1881, if they exceed the value of Five

Pounds.

Fellows also have the privilege of subscribing to the

Zoological Record for a sum of One Pound Ten Shillings

(which includes cost of delivery), payable on the Ist of July

in each year; but this privilege is forfeited unless the

subscription be paid before the 1st of December following.

The following is a complete list of the publications of the

Society already issued. !

TRANSACTIONS OF THE ZOOLOGICAL SOCIETY OF LONDON.

Ato. 20 vols. and Index. Pricete Price to the

Fellows. Public.

Vols. I-IV. (out of print).

Vol, Y., containing (67) Plates (ea(862=Go\in 2. 5) 4.93 Pe EY)

PONT LS! 5 92 Pp) co (Chelte=69) SU Oe 2 IO). ©

Bee VLL, f 73 1 wan SOOO \ie es LO! A Ole eyanles onan)

ap NBUUES s 82 » «+ (1872-74) 7853 IP) Uk 0)

Siu Wil Da 5 99 moe (IStO=T iw Bae it GB Ge 2 ©

Xi, x 95 Feed CLONE) mere ae LO) OMS ey 0

Index! Vols. Xs) Quy. eee Gis33—/O) en. 10 an OOO

Vol. -XI., containing 97 Plates .. (1880-85) .... 912 0 12 16 0

poe eee GTOTE 3 65 oo (Use) soeR Hats © 74 0

Ay ee GUN, A 62 oa (SID) Segaeon ie pada

PE NOION SC ist AE i 8 CISSIE-9S) Seay ae on @ HO. ©

a ae ot VES :, 52 ye de (SOS 4901) ae aie des. (6: Ge lee)

oe saan ates my so, CODISISOD agus) 1 ALO

og oC WE, ai Wace iy oo (CSS =IS08) oe oS CNS Aa)

» VIIL., yp) (4d jp tame LOO MERCER) 5 "4 SO) 5 8 0

Sp INO NC 5) ano woo (EOSICIO) se low @ pedis EN (0)

Bae Oe pa Wed eo) py OLOIPETOTS) ORES te ©

y XXI].—Part 1. (¢ Plates & 12 Text- fod)

GwinswhlG 2b ono g wala ay 3 0 0

now avie to offer for sale, at the reduced price of £30, sets of Vols. V._XVI. inclusive, and

In consequence of a re-arrangement of the stock of the ‘‘Dransactions,’ the Society isf

fos papers, of which a list can be supplied, at about one-fourth their published price.

PROCEEDINGS OF THE COMMITTEE OF SCIENCE AND

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LIGATURING THE ANTERIOR ABDOMINAL VEIN IN INDIAN TOAD. 44]

26. On some Results of Ligaturing the Anterior Abdominal

Vein in the Indian Toad (Bufo stomaticus Liitken).

By W. N. F. Wooptanp, D.Sc. (London), ¥.Z.8.,

Indian Educational Service, Senior Professor of Zoology,

Muir Central College, Allahabad. U.P., India.

[Received June 1, 1920: Read October 19, 1920. ]

(Text-figure 1.)

Preliminary Statement.

Most of the lower Vertebrata differ from the Mammalia in

that the liver has conveyed to it a quantity of venous blood which

has traversed the tissues of the legs and pelvic region, in addition

to the venous blood which, as in Mammals, is derived from the

gut-walls and contains the digested food products. This fact that

in the lower Vertebrata the liver receives a portion of ordinary

non-gut venous blood has not received the attention which its

possible significance deserves. So far as I know*, only one

author has ever offered an explanation, and this solely appertained

to the coccygeo-mesenteric vein of Birds. Owen in 1841+ made

the following observations: “The venous system of the kidneys

is so arranged in birds that the blood can be distributed either to

the portal system by the mesenteric vein [7.e. the blood brought

to the kidneys by the femoral veins ean flow posteriorly through

the substance of the kidneys in the so-called hypogastric veins

and so enter the coceygeo-mesenteric vein], or to the pulmonary

system by the vena cava and right side of the heart, according to

the degree of rapidity with which the pulmonary or portal

systems of veins are respectively supplied, or in other words,

according to the activity with which the circulation in each

of these systems may he going on at two different periods....

This disposition has been erroneously supposed to indicate that

the urine was secreted from the venous blood in birds, as

in reptiles and fishes; but the end attained by the venous

anastomoses in question bears a much closer relation to the

peculiar necessities and habit of life of the bird, and, so far as

I know, has not hitherto been explained. There is no class of

animals in which there may be, at any two brief and consecutive

periods of existence, a greater difference in the degree of energy

and rapidity with which the respiratory functions are performed

than in birds. When the bird of prey, for example, stimulated

by a hungry and an empty stomach, soars aloft and sweeps the

air in quest of food, the muscular energies are then strained to

the utmost, the heart beats with the most forcible and rapid

* The author has not had access to literature in India.

+ “On the Anatomy of the Southern Apteryx,” Trans. Zool. Soc. London,

vol. ii. 1841.

Proc, Zoou, Soc,—1920, No, XXX, 30

A42, PROF. W. N. F. WOODLAND ON LIGATURING THE

contractions to propel the current of blood along the systemic

arteries, and the pulmonary vessels require the greatest possible

supply of blood to serve the heart with the due quantity of

arterialized fluid: the digestive system, on the other hand, is in

a state of repose, and we may conceive the portal circulation to

be at its lowest ebb. Suppose the Hagle to be glutted with his

quarry and reduced to a state of torpor; the animal functions

ave now at rest, but the organic powers concerned in the assimi-

lation of the food are in full play, and the portal or hepatic

circulation is as active as was the pulmonary a short time before.”

And Owen further adds that ‘the anastomosis of the pelvic veins,

in being the means of conveying common venous blood into the

liver, goes to prove that the blood of the vene porte does not

require any peculiar preparation by circulation in the spleen or

other viscera to fit it for the secretion of bile.’ This explanation

seems plausible, especially when we reflect that the common

assumption made in nearly all modern text-books to the effect that

the blood always flows anteriorly in the so-called hypogastric veins

(also sometimes called the “renal portal” veims) of the bird

is almost certainly wrong, it being, on the contrary, more than

probable that the blood in these veins always flows posteriorly *,

as conjectured long ago by Jacobson (1817), Jourdain*, and other

authors. Butthis explanation evidently does not apply to animals

like Amphibia and Reptilia, which are notoriously sluggish and

yet pour into their livers a much greater proportion of non-gut

venous blood than birds. Also in Mammals, which most resemble

Birds in the alternating activity of the respiratory and portal

systems, a communication between the post-renal and portal veins

does not exist.

Now a supply of ordinary non-gut venous blood to the liver

may signify (1) that the venous blood is to enable the liver to

obtain a greater supply of water than it would otherwise receive,

or (2) that the liver is, in part, in these forms, an organ of

excretion and supplementary to the kidneys. The first supposition

naturally occurs to one when Amphibia are in question, since a

toad or frog certainly never drinks water by mouth, but always

absorbs it by the belly and thigh skin, and this water pre-

sumably is taken to the liver by the anterior abdominal vein

(and to the kidneys by the renal afferent veins), and this is

probably the case in all Amphibia. Though in Fishes, Reptilia,

and Birds there is no certain evidence of cutaneous absorption

of water, yet it is well known that Snakes and Lizards frequently

evince a desire to lie in water, and Fishes, Crocodiles, and Turtles

of course habitually live in it. It may also be remarked that the

anterior abdominal vein, or its equivalent, usually has factors from

% his is a subject I hope to investigate in the near future. It is almost certain

that these so-called hypogastric veins of birds are the homologues of the “ pelvic”

veins of Amphibia and Reptiles and not of the “renal portal” veins, and if this be

so, the blood must flow posteriorly in them.

+ M.S. Jourdain, “ Recherches sur la Veine Porte Renale,’ Annales des Sciences

Nagurelles, 4 ser., Zoologie, Tome xii, 1859, p. 134.

ANTERIOR ABDOMINAL VEIN IN THE INDIAN 'TOAD. 443

the walls of the urinary bladder, which, afterali, is a possible source

of water. Finally, it is noteworthy that, for some reason or other,

Reptiles and Birds (even when aquatic) conserve their water very

eavefully, as we may realize when we note the semi-solid character

of their urine*, the absence of sweat and mammary glands and

their non-feetal rearing of offspring ; whereas Mammals, on the

other hand, are very prodigal with water in all these respects.

Hence it is possible that, comparatively little water being taken

into the gut, the liver “arranges for” an accessory supply of

blood in order to satisfy its water requirements.

As to the liver, regarded as an organ of excretion in lower

Vertebrata, I have no evidence.

That in the Common Frog, however (and therefore in all other

animals with anterior abdominal veins), the anterior abdominal

vein supply to the liver is not essential to the life of the animal

is proved by the abnormalities occasionally found’, in which the

anterior abdominal opens into a pre-caval instead of into a

hepatic-portal vein, the animal apparently being normal in all

other respects.

The Ligaturing of the Anterior Abdominal Vein in the

Common Indian Toad.

While in india in 1915 and 1916, I determined to ascertain the

effects of ligaturing the anterior abdominal vein, though it is

evident that this operation will not give results comparable with

those to be found in the abnormalities just referred to, because

in the latter the blood-flow in other veins is not interfered with,

while in a toad with a ligatured anterior abdominal vein the

blood, which would otherwise flow through this, is foreed to pass

along the two renal afferent veins—the result being increased

blood-pressure in the renal afferent veins and interference with

the renal arterial circulation £.

In all I performed thirty-four experiments. The method I

finally adopted was to anzsthetize the toad with ether, cut through

the belly skin and the underlying muscular body-wall for two or

three centimetres, ligature the anterior abdominal vein in two

places and remove the portion of the vein in between. Both the

body-wall and the skin were sewn up with sterilized silk thread

* Sharpe’s statement (Amer. Jour. Physiology, vol. xxxi.) that the water of the

urine of birds is absorbed in the rectum, a thick paste of uric acid remaining, is in

all probability incorrect—the semi-solid urine of birds and reptiles is found in this

form in the ureters, and is so secreted by the kidneys. See Appendix in my paper

“On the ‘ Renal Portal’ System (Renal Venous Meshwork) and Kidney Excretion

in Vertebrata,” shortly to be published.

+ Woodland, W.N. F., Zool. Anzeiger, Bd. xxxv. 1910, p. 626. Also O’Donoghue,

C. H., ibid. Bd. xxxvii. 1911, p. 36.

~ See my paper “On the ‘Renal Portal’ System (Renal Venous Meshwork) and

Kidney Excretion in Vertebrata,’ Parts I. and II., shortly to be published. The

results of double perfusion and other experiments prove that as the pressure in the

renal afferent veins increases, the arterial flow becomes retarded, and above a certain

point is stopped altogether. Ligaturing the anterior abdominal vein in the Indian

toad more than doubles the amount of blood in each renal afferent vein.

30%

444 PROF. W. N. F. WOODLAND ON LIGATURING THE

and the wound cleansed with weak carbolic acid, carbolic ointment

being rubbed on the surface. While the wound was healing, 1

covered it with a pad of cotton wool, protected with a waterproof

sheet, the four corners of which were drawn out and tied anteriorly

above the scapulz and under the animal’s throat, and posteriorly

over its back. The animals were given an occasional bath in

shallow distilled water (the skin haying previously been well

cleaned), but were normally kept dry, the pad protecting the

wound from urine and fecal matter in the basin.

Out of 11 toads operated on as above described (save that L

only ligatured the anterior abdominal vein in one place) in 1915,

one lived for ten days, one for fourteen days, one a day short of

eight weeks, and two others were killed after eight weeks and

eleven weeks respectively. 1 shall only record my examination

of the three long-period survivors, all of which were operated on

on October Ist, 1915, and the wounds healed by October 12th.

A day or so later they were all active and feeding well.

One, as already stated, died on November 26th, one day short of eight weeks

after the operation. It was very thin, and had been ill during the previous week.

It weighed at death (after subtracting weight of food in gut) 21°5 gms. The heart

was normal in size (heart ratio =229°0*) ; the liver a trifle large (liver ratio=21°5 *).

The liver was very diseased, being full of small cysts; the spleen was much enlarged

and also full of cysts, and the fat-bodies very minute. Sex not recorded. ‘Tie

anterior abdominal vein was found to be well ligatured and was quite empty; on the

other hand, the renal afferent yeims and post-caval were very large, and the two

kidneys (quite healthy) were gorged with venous blood and therefore rather dark in

colour. It is important to note that a new anterior abdominal vein had not been

formed. The kidneys, after as usual being slightly squeezed and all attached vessels

removed, weighed together 0°230 gm., and were therefore apparently enormously

enlarged (kidney ratio=93'5*, body-weight taken at death). .

The toad (a male) which I killed on November 28th—eight weeks and one day

after the operation—was perfectly healthy, being active and feeding well, and all the

internal organs in perfect condition. In this toad I found to my surprise that @ new

vein had been formed posterior to the ligature and entering the liver, also two or

three small new veins coming from the muscles of the anterior ventral body-wall and

opening into the principal new vein (text-fig. 1, B) 7. The body-weight (atter weight

of food in gut subtracted) was 21°9 gms. The heart ratio was 199°3*; the liver

(weighing exactly 1:0 gm.) ratio=23°9*; the kidney ratio=1648%. ‘The kidneys,

therefore, were about normal in size.

The remaining toad (a male) 1 killed on December 18th—cleven weeks after the

operation—and this also was in perfect health mside and out. In this toad also a

new vein had been formed to allow the ligatured anterior abdominal to become

functional. The body-weight was 30:0 gms. The heart ratio=229'0*; the liver

ratio=19°6 *; the kidney ratio=157°9*. The kidneys, therefore, were

about normal

in size.

‘ ody

aoe. eee —for the heart (all vessels cut off and

all blood washed out and dried) in these toads is 216°8 (determined in 29 toads); the

normal ratio for the liver (squeezed and dried) was 284 (18 toads); the normal ratio

for the kidneys (wt. of both kidneys, attached ureters and vessels being removed)

was 237°5 (82 toads). J must add, however, that in two other lots of toads the

average kidney ratios were 155°2 (10 toads) and 159°3 (6 toads). The kidney ratio

(also the heart and liver ratios) shows great variations, not connected with the time

of year or with sex.

+ Compare the “ rapid formation of a collateral circulation so that the blood could

get round the ligature to the liver” (Starling, ‘ Principles of Human Physiology,’

1912, p. 858) when in a mammal the portal vein was ligatured off from the liver and

made to open into the posterior vena cava. In this case, however, the liver is

deprived of blood from the gut and not merely of an additional supply of ordinary

yenous blood, as in the toads with ligatured renal afferent veins,

* The normal (average) ratio—

ANTERIOR ABDOMINAL VEIN IN THE INDIAN TOAD. 445

Tn 1916 [ was unfortunate in this experiment, possibly owing

fo my predilection for giving the animals a bath every morning,

but probably also owing to the fact that in each operation I

ligatured the anterior abdominal vein in two places and eut out

a portion about half a centimetre in length in between. Out of

23 toads operated on only five survived for more than eleven

days. That this was due largely to the mode of operation and

not solely to the ligature of the anterior abdominal vein is shown

Text-figure 1.

A B

A. Small regenerated anterior abdominal vein in a toad which was killed four weeks

after the operation. LG, positions of the two ligatures, the intervening

portion having been cut out in the operation. NV, new vein formed.

B. Large regenerated anterior abdominal vein in a toad killed eight weeks and one

day after the operation, which merely consisted of the vein being ligatured

in one position. GB, gall-bladder ; other letters as in A.

I cannot guarantee the absolute fidelity of these figures, since they have

been copied from rather rough sketches made when I dissected the toads.

by the fact that out of ten contro! toads in which an identical

operation was performed, save that the exposed anterior abdominal

vein was not ligatured, nine of them did not survive for longer

than eleven days, and only one for six weeks and two days.

OF the five survivors of the actual operation in 1916, one which survived eleven

days had a kidney ratio (in all these ratios, unlike those of 1915, the weight of the

body is that taken at date of operation) of 119°7; another which survived twelve

days had a similar kidney ratio of 134°2; another survivor of two weeks and one day

had a kidney ratio of 95:8. Only one toad completely survived the operation, and

this was a female which I /illed four weeks after the operation. ‘The animal was

then in perfect health inside and out, and in this case I found that a new vein had been

formed to enable the anterior abdominal vein to continue to function (text-fig. 1, A).

The animal weighed 23:4 gms. on the date of operation (July 22nd), but when killed

446 PROF. W. N. F. WOODLAND ON LIGATURING THE

on August 19th only weighed 14°6 gms. (after allowing for weight of food in gut).

All internal organs were quite healthy, and the renal afferent veins were very large,

due of course to the small size of the newly-formed portion of the anterior abdominal.

The heart had apparently become much reduced in size, or was naturally small, the

ratio being (weight of body at date of operation) 325°0 ; the liver was also extremely

small (ratio=67'2); and the kidneys were below the average size (ratio=260'0).

Summarizing the results of these experiments, we may conclude

that the ligature of the anterior abdominal vein caused either

the death of the animal or the re-formation of the anterior

abdominal vein: in no case did an animal survive in a healthy

condition for a considerable length of time without a functional

anterior abdominal vein.

Conclusions.

The fact that a new anterior abdominal vein was always formed

in those toads which survived the operation described is no proof

that the supply of blood to the liver by this vein is essential to

the animal, because, as I have already pointed out, this feature is

most certainly due to the fact that the arterial circulation in the

kidneys is interfered with under the conditions of these experi-

ments, and that is a sufficient reason for the formation of a new

anterior abdominal vein.

We have also seen that frogs can live without the blood in the

anterior abdominal vein being added to that in the hepatic portal

vein, and this fact by itself is good evidence for the view that the

non-gut blood is not essential to the animal’s welfare.

Two questions remain : (1) why should an anterior abdominal

vein exist ? and (2) why should it normally open into the hepatic

portal vein? The answer to the first question I have already

indicated in a paper* published in 1906. In this paper I

contended that in animals with “ portal” kidneys the flow of

blood through the primitive posterior cardinal veins is considerably

hindered by the kidney tubules invading the lumina of the two

veins and subdividing them up into coarse networks of sinusoids

(Shore +, Minott), and that the anterior abdominal vein is formed

as an alternative route to relieve the congestion consequent on the

formation of the “renal portal” system. Judging from recent

measurements of the relative diameters of the renal afferent and

anterior abdominal veins in Bufo stomaticus and Rana temporaria

respectively, I find§ that in the Indian toad about three-fifths of

the venous blood from the legs flows to the heart via the anterior

abdominal vein and about two-fifths via the two renal afferent

veins, and that in the frog (2. temporaria) about one-half of the

blood flows by each of these two routes, from which we may

conclude that the resistance to flow of the blood offered by the

liver capillary system is in the toad about one-third and in the

frog about one-half of that offered by the renal venous meshwork

(‘renal portal” system) of each kidney. From other evidence$

* Woodland, W. N. F., Proc. Zool. Soc. London, 1906, p. 886.

+ Shore, T. W.. Jour. Anat. Physiology, vol. xvi. (u.s.) 1901.

* Minot, C.S., Proc. Boston Soc. Nat. Hist. vol. xxviii. (10) 1898, p. 265.

§ See Part Il. of my paper “On the ‘ Renal Portal’ System ete.,” shortly to be

published.

ANTERIOR ABDOMINAL VEIN IN THE INDIAN TOAD. 447

I have also concluded that the resistance offered to the blood

traversing the hepatic portal system is very little more than that

experienced by the blood when the anterior abdominal vein opens

directly into a pre-caval vein, and this is supported by the fact

that in such abnormalities (in which the anterior abdominal opens

directly into a pre-caval) the renal afferent veins are of about the

normal size. ‘The renal venous meshwork in both kidneys then

offering considerable resistance to the passage of blood from the

hind limbs and tail, I again suggest that this blood has sought an

additional path by means of which it can reach the heart without

traversing the “renal portal” systems, and this suggestion is in

complete agreement with the view of morphologists that the

anterior abdominal vein of Amphibia and Reptilia is a “new”

structure, and has nothing in common with the vein which it so

often resembles—viz., the umbilical vein.

Since we have provisionally concluded that the blood in the

anterior abdominal vein is of no use to the liver, and that the

opening of this vein into the hepatie portal vein offers practically

no more resistance to the passage of the blood than if it opened

directly into a pre-caval, the answer to the second question must

evidently be to the effect that the anterior abdominal vein opens

into the hepatic portal vein merely because it is more convenient,

the hepatic portal vein being more accessible than the pre-caval,

and offering, as we have seen, but little more resistance to the

flow of the blood. Occasionally, however (in ‘ abnormal” frogs),

the ancestral connection of the anterior abdominal vein with a

pre-caval vein is retained. It may beadded that in some animals

(e.g-, im most Elasmobranchs) the posterior cardinal sinuses are

so little broken up by the relatively small kidneys (Vialleton*)

that a bypath for the blood (anterior abdominal vein) is

unnecessary.

* Vialleton, M. L., “ Caractéres lymphatiques de certaines veines chez quelques

Squales.” C. R. Hebdom. des Séances de la Soc. Biol. Paris, Tome liv. 1902.

LIFE-HISTORY OF THE COMMON NEMATODE OF THE DOGFISH. 449

27. Some Observations on the Structure and Life-History

of the Common Nematode of the Dogfish (Seylliwm

canicula). By J. H. Luoyp, M.Se., F.Z.8., Assistant

Lecturer and Demonstrator in’ Zoology at University

College, Cardiff.

[Received May 15, 1920: Read November 2, 1920.]

(Text-figures 1-3.)

Introduction.

In the following pages an account is given of the parasitic

Nematode, Proleptus scillicola, together with some observations on

its life-history. The work was carried out, partly in the Zoolo-

gical Department of the University of Birmingham and. partly

at the Marine Biological Laboratory at Plymouth, towards the

end of 1914 and the beginning of 1915, but until now I have

been unable to write up my results for publication, owing to my

absence on military service.

J should like here to express my gratitude to Professor F. W.

Gamble for his kindly criticism and advice during the course of

this work. My thanks are also due to the Board of Studies in

Zoology of the University of London for the use of their table

at Plymouth, and to the Royal Society for providing a grant

which enabled me to procure the necessary material.

A description has been thought desirable because, although

the worm is exceedingly common, no complete account of its

anatomy exists, and its life-history has never been worked out.

Proleptus scillicola occurs in the alimentary canal of Scylliwm

canicula from the mouth to the pyloric constriction. It has also

been reported from S. catulus (5), S. stellare (1), Raia clavata (1),

and Raia circularis (1). I have only had the opportunity of

examining one Scyllium catulus, and this specimen was absolutely

free from infection.

Other species of the same genus under a different generic

name have been reported and briefly described as follows :—

Spiropterina inflata (8) Linstow, attached to the wall of the

stomach of Scylliwm immoratum; Spiropterina afr icana (6)

Linstow, from stomach of Anguilla sp. (%); Spiropterina elegans

(12) Orley, from the stomach of Heaacanthus griseus; Spiro-

pterina dacnodes (4) Creplin, from the cesophagus of Raia clavate

and the stomach of Squalus mustelus.

The worms are white in colour, and occur either free or

attached within their hosts; when attached, it is always by the

tail-end.

The females are considerably larger than the males, from which

they are easily distinguished by the tail, which in the male is

coiled.

450 MR, J. H. LLOYD ON THE STRUCTURE AND LIFE- HISTORY

Historical.

Some confusion appears to exist as to the correct generic name

of the Common Nematode of Seylliwm canicula. The first generic

name assigned was Proleplus by Dujardin (5) in 1845. He

(distinguished two species-—Proleptus acutus and Proleptus obtusus,

but as he gives neither description nor diagram of Proleptus

obtusus, 1b may be considered invalid.

In 1860, Molin (11) described a nematode under the name of

Listiocephalus dacnodes, which should apparently ve referred to

the genus Proleptus, as it differs considerably from the remaining

species of /istiocephalus and appears to be wrongly placed.

During the following year (1861) Van Beneden (2) described the

worm under the generic name of Spiropterina.

Previous to Van Beneden placing it in a separate genus, it

was provisionally assigned by Creplin (4) to the genus Spiroptera,

chiefly because its head-end bore some slight resemblance to the

Spiropteras of Valpa, Hrinaceus, etc.

In 1870, Van Beneden (1) mentioned the worm under the

generic name of Coronilla, and distinguished three new species :—

Coronilla scillicola, Coronilla robusta, and Csronilla minuta.

Neither the new genus nor the new species were described, but

six drawings were given, which referred indiscriminately to two

of the species—viz., Coronalla robusta and Coronilla seillicola.

Spiroptert ia appears to be the generic name by which the

worm is most commonly known, but Linstow (7) has put forward

a strong argument in favour of the name Pr oleptus on the ground

of priority. For this reason I shall use the generic name /’ro-

leptus, but as Dujardin’s species acutus evidently refers to a

distinct species which occurs in Seylliwnr catulus, and his species

obtusus is difficult to determine, I propose to retain Van

Beneden’s specific name seillicola, aud shall describe the worm

under the name of Proleptus scillicola.

In addition to the foregoing, it has been briefly described by

Linstow (9). My observations differ from those of Linstow in

several respects, and his diagrams are inaccurate.

MorpHouoey.

(a) Haternal Characters.

The body is surrounded by a very thick, transparent cuticle,

which is finely ringed transversely. Proleptus scillicola possesses

extraordinary vitality. I have kept specimens alive in normal

salt solution for over 17 weeks, whilst Linstow (9) records the

fact that “specimens which had been in Miiller’s fluid for

48 hours arrived in a living condition.”

The best results for whole mounts were obtained with Looss’s

fixative. Worms killed in this manner are straightened consider-’

ably and retain their transparency. Perenyi’s fluid proved to be

the best fixative for animals which were to be sectioned, whilst

Para-carmine was the most effective stain.

OF THE COMMON NEMATODE OF THE DOGFISH. 45]

The body is slightly attenuated towards the anterior end, and

the head (text-fig. 1) is characterized by the presence of a

peculiar cuticular collar. It was probably the presence of this

collar which suggested to Van Beneden his generic name

Text-figure 1.

‘bn

(O06:

N.R.

Head of Proleptus scillicola.

LETTERING OF FIGURES.

A, Anus. Oes. (sophagus.

Al.C. Alimentary canal. Ov.D. Oviduct.

C. Cuticle. hae 58 Papille.

CG: Cuticular collar. | i, Rectum.

M0. Cloaca. _ R.BIW. Right bursal wing.

C.W. Cuticular wing. RG. Rectal gland.

Lh, Intestine. | SaSe Small spicule.

LT. Lips. |. (i Uterus.

LBW. Lett bursal wing. | Va. Vulva.

LAS. Large spicule. | WV. Vagina.

N.R. Nerve-ring.

Coronilla. There are two rounded, protruding, lateral lips, each

of which bears a single conical tooth. Linstow (9) mentions in

addition ‘“interiorly a small pointed cone,’ but I have failed to

find any evidence of this either in whole mounts or sections.

The excretory pore is exceedingly small, and is situated in the

452 MR. J. He. LLOYD ON THE STRUCTURE AND LIFE-HISTORY

mid-ventral line, almost as far distant posteriorly from the nerve-

ving as the latter is from the anterior end.

The “tail” of the male (text-fig. 2) is coiled usually in a

single ring, but occasionally into a spiral of two rings. There

is an egg-shaped bursa, to right and left of which the cuticle is

drawn out into wing-like expansions, supported by eight costal

papillz on each side. Van Beneden (2) states that there are six

or seven papille on each side. These papilla are pedunculate in

form and are arranged symmetrically in two rows.

There is a single papilla at either end of each row ; between

the extreme papillz are six others arranged in three pairs. One

Text-figure 2.

Lateral view of tail of male Proleptus scillicola, showing large and small

spicules and costal papille.

par is situated in the region of the cloaca, and the two remaining

pairs are posterior to it. The pre-cloacal papillae are somewhat

longer than the remainder. Linstow (9) bas figured three

papille on each side of the cloaca, but after careful examination

of many bursee, I have failed to discover the third papilla.

Two copulatory spicules of unequal length are present, the left

being approximately five times as long as the right. ‘The average

length of the longer spicule is 2°125 mm. and of the shorter

“41 mim.

The longer spicule is curved towards the distal end, and at the

beginning of the curvature there is a prominence on the convex

OF THE COMMON NEMATODE OF THE DOGFISH, AD3

side. Linstow (9) has stated that the larger spicule is bent into

a hook at its distal end, but [ have not observed this arrange-

ment in any of the worms I have examined. It is covered with

numerous transverse markings, which give it a striated appear-

ance under a fair magnification, and Nenana to a point at its

distal end. Both spicules are ensheathed in cuticle.

The “tail” of the male ends bluntly, and in the neighbourhood

of the spicules the cuticle is raised into rows of rounded eleva-

tions on the ventral surface.

The tail of the female (text-fig. 3) is bent at an angle to the

body and is somewhat attenuated. In the region of the vulva

the cuticle increases considerably in thickness cn the ventral

surface and forms a pad.

Measurements in the case of Proleptus scillicola are almost

valueless, as there are wide variations not only in the sizes of

individuals, but also in the proportions of different parts. The

male varies considerably less than the female.

Mature females vary in length from 32°17 mm. to 59 mm., and

the tail measures from °395 mm. to ‘583 mm. The proportion

of tail: total length varies from 1:75:47 to 1:146°6. Linstow

has given this proportion as 1:38°7.

The distance from the vulva to the tail-end varies from

°887 mm. to 1:06 mm., and the proportion of ‘ Vulva to tail

end”: total length varies from 1: 39-7 to 1:53:07.

The males vary in length from 31:73 mm. to 36°2 mm. The

tail measures from 1°03 mm. to 1°56 mm., and the proportion of

tail: total length extends between 1:19°66 and 1:30°8. Lin-

stow’s proportion is given as 1:21°6. It would appear that

Linstow has based his proportion on measurements of a single

worm of each sex.

(b) Internal Anatomy.

The alimentary canal is normal, consisting of an esophagus,

intestine, and rectum.

The osophagus is of unequal thickness, being approximately

twice as broad behind the nerve-ring as it is in front of at.) In

the female it varies in length from 3°69 mm. to 5°86 mm., and

the Rroporlion of csophagus : total length varies from 1:8°17

to 1:11:52. In the male the cesophageal length fluctuates between

3-6 mm. and 4- 53 mm., and the proportion of cesophagus : total

length from 1:7°32 to 1: 8°81.

In es section the lumen of the alimentary canal is tri-

radiate anteriorly, but towards the posterior portion of the

intestine it becomes tetra-, penta-, or hexa-radiate.

Rectal glands occur Around the anterior end of the rectum in

both sexes. They are four in number and are symmetrically

arranged.

The female reproductive system is typical, and the vulva opens

to the exterior, a short distance in front of the anus, on the

ventral surface. The eggs are developed from a polynucleated

454 MR. J. H. LLOYD ON THE SLYRUCTURE AND LIFE-HISTORY

mass of protoplasm, and acquire distinctness as they approach

the oviduct. ;

The ova vary in length from 47°5 m to 52, and in breadth

from 28°75 w to 34:1 uw. The thickness of the shell varies from

6°25 « to 6-52 4. Asa rule a coiled-up larva is formed within

the shell before the “ eggs” are extruded from the uterus.

It is possible to burst the “eggs” under a cover-glass and set

free the contained larvae, which average 201 w in length. They

are slightly attenuated towards the head-end, which is charac-

terised by two protruding lips as in the adult, but there is no

cuticular collar. The tail ends in a point, and is bent at an angle

to the remainder of the body, but exhibits no sexual differen-

tiation.

Text-figure 3.

Tail of mature female Proleptus seillicola.

The male reproductive system is of the usual type.

The excretory system consists of two canals, which pursue a

sinuous course, one in each lateral line. From each canal

a single branch descends to the ventral surface, where they unite

just before reaching the excretory pore.

The nervous system consists of a well-defined cireumeceso-

phageal ring which gives off numerous branches both anteriorly

and posteriorly.

Life- History.

The literature dealing with the life-history of Proleptus scilli-

cola is scanty. In 1865, McIntosh (10) described ‘“‘The Tre-

matode Larva and Ascaris of the Carcinus menas.” He

discovered two specimens of his so-called Ascaris, one of which

was lost, but his description of the other in some respects agrees

with that of Proleptus scillicola,

OF THE COMMON NEMATODE OF THE DOGFISH. 455

In 1870, Van Beneden (1) stated that “Dr. MeIntosh has

found asexual nematodes in the ‘liver’ of Carcinus menas,

which appear to be the young of Coronilla (Van Beneden).”

Again, in 1875, Van Beneden (8a) stated that “The ordinary

crab of our coasts, Carcinus menas, is the vehicle of a nematode

which becomes a Coronilla robusta in the stomach of a ray.”

Vaullegeard (14) in 1896 veported the larva of Coronilla

robusta (Van Beneden) from a number of crustaceans, but stated

that it was rare. In addition to Carcinus manas, he discovered

it in Portunus marmoratus Leach, Hyas araneus Linneeus, and

Pagurus bernhardus Linneus, but found only oac example in

each of the last three hosts. Later he found a dozen nematode

larve in Portunus depurator Pennant (=P. marmoratus Leach).

Linstow (9) mentions the discoveries of McIntosh and Vaulle-

geard, and states that the latter distributed the preparations of

the larve, which indubitably belong to the genus Spiropterina

(Van Beneden).

Heperimental,

Following up the suggestion of Van Beneden, I have attempted

to infect the Common Shore-Crab, Carcinus menas, with the

larvee of Proleptus scillicola.

The results so far have been disappointing, as I have succeeded

in obtaining only one larva, which I may reasonably assume to

have developed from eggs, taken in with mature females of

Proleptus scillicola, on which the crabs were fed.

During the course of these experiments I have dissected

100 crabs, of which 59 were utilised in the feeding experiments

and the remainder in control experiments. Of the 59 crabs used

in the feeding experiments, 8 were infected with Proleptus larve,

43 larve being found, Two of the 41 crabs used in control

experiments were infected, six larvee being found.

Owing to the fact that all the larvae, except one, found in the

erabs used in the feeding experiments were as old or older than

those found in the controls, 1 am bound to conclude that they

occurred as the result of natural infection.

The larva which I presume to be the result of my feeding

experiments measured 1:85 mm. The length of the cesophagus

was °35 mm., and the proportion cesophagus: total length was

1:5:28. There was no cuticular collar present, but the tail

was bent at an angle to the remainder of the body.

The older larve examined varied in length from 10°16 mm.

to 20 mm., and the length of the cesophagus from 1:79 mm.

to 2°68 mm. The proportion cesophagus: total length fluctuated

between 1: 5:09 and 1: 8°29.

In the majority of these older larvee the head-end was sur-

rounded by tie cuticular collar which is so characteristic of the

adult worm. Genital organs were absent, as was also differen-

tiation at the tail-end.

456 LIFE-HISTORY OF THE COMMON NEMATODE OF THE DOGIISH.

I have dissected a number of specimens of Pagurus bernhardus,

but have not found any larvee of Proleptus scillicola in them.

Conclusions.

The above experiments show that the larve of Proleptus scilli-

cola undoubtedly occur in Carcinus menas, but my failure to

produce an artificial infection seems to point to the fact that

Carcinus is not the true intermediate host.

It is possible, however, that Carcinus menas may play some

part in the infection of the Dogfish. Seylliam canieula does not

normally occur at a less depth than thirteen fathoms, whereas

Carcinus mcenas rarely occurs below a depth of three to four

fathoms. During the breeding-season, however, Dogfish come

close inshore, and it is probable that some of its infection takes

place at this season.

Literature.

1. Bunepen, P. J. vAn.—“ Les Poissons des Cotes de Belgique,

leurs Parasites et leurs Commensaux,” plate 38. Meém.

Acad. Se. Bruxelles, 1871.

2. BrenepEN, P. J. van.—‘‘ Mémoire sur les Vers intestinaux.”

Jompt. Rend. Acad. Sc. Paris, Suppl. 1861, pp. 270-271.

3. Brenepen, P. J. vAn.—Les Commensaux et leurs Parasites,

1875, p. 206.

38a. Benepen, P. J. vAN.—Animal Parasites and Messmates.

Translation of 3. 1st English edition. London, 1875.

4. Creprin.— Spiropterina dacnodes.” Archiv fiir Naturgesch.

1851, p. 308.

Dusarpry.— Histoire des Helminthes, 1845, p. 105,

Linstow, Dr. O. von.—Mitteilungen aus der Zoologischen

Sammlung des Museums fiir Naturkunde in Berlin.

1. Band, 2 Heft.

7. Linstow, Dr. O. von.—Archiy fiir miky. Anat. und Entwick.

Band 60, 1902.

8. Linsrow. Dr. O. von.—Archiv fiir Naturgesch. Berlin, 1890.

Don

Band 1, Heft 3, pp. 180-181, Tab. x. figs. v.—viii.

Linstow, Dr. O. von.—Achiv fiir mikr. Anat., Band 58.

Bonn, 1901, pp. 191-194, Tab. ix. figs. 27-30.

. McInrosu, W. C.—‘‘The Trematode Larva and Ascaris of

the Carcinus menas.” Q.J.M.S8. vol. vi. (new series),

1865, pp. 201-204.

11. Monin, R.—Sitz. k. Akad. Wiss. 1860, pp. 512-513.

12. OrtEy.—‘‘ Die Entozoen der Haien und Rochen.” Termés-

zetrajzi Fiizetek, vol. ix., Budapesth, 1885, pp. 97-126

& pp. 216-220 (Spirop. elegans).

13. Srosstcu.—Note Elminth. ‘Trieste, 1893.

14. Vautiecnarp, A.— Bullet. Soc. Linnéenne de Normandie,

A sér. t. x., 1896, pp. 50-53,

(

ON THE SBPXUAL PHASE IN INDIAN NAIDID&. Ad

28. On the Sexual Phase in certain Indian Naididee (Oligo-

cheta). By Haru Ram Méura, M.Se., Professor of

Zoology, Benndn University, Beans: ae

[Received June 15, 1920: Read November 2, 1920. |

(Text-figures 1-3.)

I have recently collected in the neighbourhood of Agra a large

number of examples of the following species of Naididee and

Tubificide, many of which are fairly common there :—

Nais pectinata var. inequalis Stephenson.

Nais communis Piguet, var. punjabensis Stephenson.

Hemonais laurentii Stephenson.

Cheetogaster orientalis Stephenson.

Chetogaster punjabensis Stephenson.

Devo linosa Leidy.

Pristina longiseta Ehrbg.

Branchiodrilus hortensis Stephenson.

Branchiura sowerbyi Beddard.

As is well known, the Naididze usually reproduce asexually

by fission, and in many species the genital organs have never yet

been described. As Stephenson remarks (3), if such descriptions

“were available throughout the group, it can hardly be doubted

that we should be able to judge better of the affinities of genera

and species, and consequently to improve our classification ; since

the diagnoses of species and genera, and the scheme of dassté-

cation, ‘depend at present to an unduly large extent on one single

set of characters, the form and distribution of the sete.” I

therefore give an account of the sexual organs in two of the

above species, WVais pectinata vav. ineoorlis and Branchiodrilus

hortensis ; though the organs have been described in certain

other species of Vais, we have as yet no account of them in any

species of the genus Lranchiodrilus.

All the species of Naididee which have been observed by

Stephenson to become sexual in Lahore, considerably further

north than Agra, do so from February to May; the rains are

there later and scantier than further south, and May, June, and

sometimes July, before the rains appear, when the ponds are dry

and the ground baked hard, represents the most unfavourable

season of the year for pond-life. In Europe these worms would

seem usually to enter on the sexual phase in the autumn, before

the rigours of winter. In Agra I found the sexual specimens

deser hea below in the autumn—in this part of the country the

rains are abundant from the latter part of June to September ;

the ponds begin to dry up in October, and the cold weather

* Communicated by J. SrmeireNnson, 1).Sc., F.Z.S.

Proc, Zoou. Soc.—-1920, No. XX XI. 31

458 PROF, LWARU RAM MELLRA ON THE

appears, as in Kurope, to be the unfavourable season. Whether

the sexual phase makes its appearance in spring or autumn, there-

fore, it seems to be a measure of protection against approaching

adverse conditions; the ova, quiescent or developing slowly

within the cocoon, are probably able to withstand such conditions

better than the adult animal.

In the case of Vais pectinata var. inequalis, atter the attain-

ment of the full sexual phase, the alimentary canal in several of

my specimens was seen to degenerate; the sime phenomenon

has been noticed by Stephenson (8, 4) in Dero limosa and

Temonais lawrentit.

Many specimens of branchiura sowerbyi, Dero limosa, branchio-

drilus hortensis, and Hemonais laurentii were found living

together in the mud of a pond near Sikandra. A similar curious

association of Branchiura sowerbyi and a species of Branchio-

drilus has been noted by Beddard (1) in the Victoria regia tank

in the Royal Botanic Society’s Gardens in Regent’s Park ; and

Branchiura sowerbyi, Branchiodrilus hortensis, and a species of

Dero have been found associated by Stephenson under’ natural

conditions at Lahore (2). These three are among the few genera

of Oligocheeta which possess gills; in Hemonais lawrenti, the

fourth worm which I found in the association, though there are

no gills, the vascular system is, for one of the Naidide, particu-

larly highly developed.

NAIS PECTINATA Var. INZQUALIS Stephenson.

Since Stephenson had only spirit specimens at his command,

I prefix a short account of some features of the general anatomy

of this worm.

My specimens were larger than Stephenson’s, their usual

length being 8-10 mm., but when the worms are fully extended

it may reach 15-18 mm. The colour is light reddish brown.

The prostomium is bluntly conical. The worms exhibit active

wriggling movements. The number of segments varies consider-

ably—trom 40 to 95.

The ventral sete are 4-6 in a bundle, usually 5; the length of

those in the anterior segments (ii.-v.) is 97-105 pw, of the rest

88-93 w. The dorsal hairs were 306-332, and the dorsal

needles 106-112 4; there is a slight indication of a nodulus on

the latter.

The penial setie (text-fig. 1) are the modified ventral sete of

the sixth segment. They are 4-6 in a bundle, 98-105 in

length, and are somewhat swollen near the tip, which is usually

not forked although it is slightly hooked. Only two sete were

noted as being bifid at their free end, and in these the prongs

were short. blunt, and of equal length. The whole bundle has

somewhat the appearance of a fan, and arises to the inner side of

the male genital aperture.

Ordinarily the dorsal sete begin in the sixth segment, as

SEXUAL PHASE IN INDIAN NAIDIDA. 459

usual in the genus. In sexually mature specimens, however,

they begin in segment viii.; in one specimen there were needle

sete only in the dorsal bundles of viii., but no hairs; in a few

the sete only began in ix.; in two cases the sete were seen to be

thrown off from segment viii. when a cover-glass was gently

placed over the worm,

The body-cavity contains a large number of rounded corpuscles,

brownish in colour, and in addition there are a few colourless

corpuscles filled with refractile granules. Ccelomie corpuscles

were more numerous in specimens which had been kept in the

laboratory for several days. They are few in the first six

segments.

Text-figure 1.

A. Penial seta of Nuis pectinata var. inequalis. X 540.

B. Penial seta of Branchiodrilus hortensis. > 540.

Text-fig. 1 drawn by camera lucida.

The pharynx occupies segments 111.—v., and is diffusely covered

by a small amount of chloragogen pigment, which extends right

up to the prostomium. In a transverse section the cavity has

the appearance of an inverted I’, owing to the presence of a

median dorsal diverticulum ; this diverticulum and the dorsal

wall of the pharynx are ciliated. On the upper and lateral

surfaces of the pharynx are a number of pyriform cells, their

narrow ends resting on the surface of the pharynx; these are

arranged in groups of three, four, or more, and the groups are

separated by strands of muscle which pass upwards from the

pharynx to the body-wall. The cells are about 30 in length

and 9 » in thickness ; they stain deeply with hematoxylin. The

whole pharynx much resembles that of Hamonais laurentii (4).

The gut is not distinctly differentiated into cesophagus, stomach,

and intestine ; it is somewhat larger in segments vi.—vili., then

narrow as far as xiv., after which it is continued as a fairly broad

tube for some distance. Its epithelium is ciliated. The anus, is

alt

460 PROF. HARU RAM MEHRA ON THE

dorsal. Strong ciliary movements were seen in the posterior part

of the gut, the direction being forwards; antiperistaltic con-

tractions were also noticed to be taking place over some length

of this part of the tube.

The blood is yellowish, and without corpuscles. The dorsal

vessel lies on the left side of the alimentary canal near the

ventral surface as far as septum 5/6, where it becomes dorsal,

and lies over the pharynx ; it is surrounded by chloragogen cells,

or in the region of the pharynx by the pigment previously

mentioned. It bifurcates near the anterior end of the animal,

and the branches, turning ventrally, unite to form the ventral

vessel at the level of the first ventral setal bundles. There are

four pairs of lateral commissures in the pharyngeal regton, which

form a plexus; behind this, from segment vi. onwards, there is

a commissure on the anterior face of each septum,—in some

specimens, however, these were only seen as far back as segment

xvi. The body-wall is devoid of capillaries.

The first nephridium les in segment vii.

The cerebral ganglion is large and bilobed, deeply indented in

front and behind. ‘The ventral nerve-cord has an irregular

lobulated outline, ganglia not being clearly distinguishable.

Text-figure 2.

ce. 4)

ie fll

ty i}

FOR

ies A\-

u.defr 3 yy.

Genital region of Nais pectinata var. inequalis. Abr., atrium; ¢7.d., ejaculatory

duct; fem.f., female funnel; ov., evum ; ovis., Ovisac ; sep. 4/5, 5/6, and

6/7, the septa between segments iv. and v., y. and vi., and vi. and vii. ;

s.f, seminal funnel; sp.s., sperm-sac; spth., spermatheca; v.def., vas

deferens; y., yolk; g, male aperture. X ca. 120.

Text-fies. 2 and 3 are semidiagrammatic, and are compiled from several

successive vertical sections.

Genital Organs (text-fig. 2).—As usual in the Naidide, the

gonads appear first, and disappear entirely before the rest of the

genital apparatus has reached its full development.

The testes are a pair of ovoid bodies attached to the posterior

face of septum 4/5. The ovaries are similar in appearance in the

living specimen, but smaller; they arise from the posterior face

of septum 5/6.

The seminal vesicle is formed soon after the appearance of .the

gonads, as a backward bulging of septum 5/6, which later on,

SEXUAL PHASE IN INDIAN NAIDIDA. 461

when distended with the male products, may reach as far back as

seoment xii. The vas deferens, one on each side, is a short tube

with only a single bend; in diameter it is 15 pu, except where it

joins the atrium, where it is only 9; it has a uniform lining

of cubical epithelial cells. It passes vertically downwards from

the funnel on the posterior face of septum 5/6, and then after a

slight bend enters the atrium on its anterior aspect close to the

origin of the ejaculatory duct.

The neck of the male funnel lies in the mouth of the sperm-

sac (seminal vesicle). The funnels fill up the mouth of the sac,

and are directed upwards and backwards just within it; they

meet each other in the middle line above the gut by their inner

margins, and their outer surfaces are fused with the contiguous

part of the wall of the sac. They are cup-shaped, with everted

lips, and are lined by columnar ciliated cells with prominent oval

nuclei at the base. The greatest diameter of the funnel is

about 45 p.

The atria are ovoid chambers with their long axes vertical,

lying one on each side of the seminal vesicle in segment vi. Hach

is 80-90 » in height and 45-50» in breadth, and is lined by an

epithelium of cubical cells with indistinct outlines and con-

spicuous nuclei; outside the epithelium is a thin coat of circular

muscular fibres, outside which again the peritoneum is indicated

by a few scattered nuclei. The lumen may contain spermatozoa

or only a little coagulum. The ejaculatory duct is short,

about 30; its epithelium consists of closely packed columnar

cells with large peripherally situated nuclei; it has a fairly thick

investment of cireular muscular fibres. The duct may be in-

vaginated into the base of the atrial cavity; it opens to the

exterior in the depth of a short tubular depression of the ventral

body-wall, about 15 in length, which is narrower at the surface,

and broader above, where it receives the duct. There are no

‘‘ prostatic” cells in connection with vas deferens or atrium.

A thick band of muscle-fibres runs vertically upwards from the

ventral body-wall, lying internal to the atrium and supporting

the setal sac containing the penial sete.

The ovisac, formed by the backward bulging of septum 6/7,

surrounds the sperm-sae which lies within it. It may reach back

to segment xvi. ; it contains a large mass of yolk granules, which

stain faintly with eosine; and in its hinder part a number of

ova. The septa of the several segments behind the seventh

retain a transverse position between the body-wall and the ovi-

sac, fusing closely with the periphery of the latter. Large

blood-vessels are seen closely applied to the sperm-sac and inner

face of the ovisac.

The female funnels are attached to the anterior face of septum

6/7 near the ventral parietes ; the cells lining it are small, and

appear to be modified peritoneal cells containing little else than

nuclei. In ‘one specimen the funnels were seen to open on the

yentral surface at about the level of septum 6/7. They are seen

462 PROF, HARU RAM MEHRA ON THE

only in specimens which have reached full sexual maturity, and

considering the large size of the ova seem to be too minute to be

of any functional importance.

The spermathece occupy segment v., and their openings lie at

the anterior edge of the clitellum immediately behind septum 4/5.

The ampulla attains a maximum height of 105; its posterior

surface lies near the mouth of the sperm-sac, while in front 1t

may push forwards septum 4/5 so as to encroach on segment iv.

The ampullze when distended are ovoid, and meet and press on

each other in the middle line above the alimentary tube. The

epithelium is low and flat, except near the duct where the cells

are fairly high and cubical. There is a thin layer of circular

muscular fibres, and a few periteneal cells on the outside. The

spermathecal duct arises anteriorly instead of from the middle

of the base of the ampulla; it is slightly oblique in position,

cylindrical, about 45 in length; its lining consists of closely

packed columnar cells with nuclei peripheral, and a fairly thick

coat of circular muscular fibres surrounds this epithelium.

The clitellum covers more than half of segment v., and all vi.,

vil., and vill.; to -the naked eye it is opaque white. The cells

are four times as high as the ordinary surface epithelium, are

vacuolated, and when fully developed lose their distinetness of

outhne. The clitellum is absent from the regions of the body-

wall between the spermathecal pores and the male apertures.

The alimentary canal undergoes great degeneration in the

sexually mature worm. Though known in Dero limosa and

Hemonars laurentii, the phenomenon has not so far been observed

in the genus Vais. In advanced stages of maturity the mouth

becomes closed; the buceal cavity and anterior part of the

pharynx lose their lumen and become reduced in size ; the pharyn-

geal cells lose their distinctness of outline, and those of the

ventral wall are reduced in size, low and cubical. Behind the

pharynx the gut is comune! as a narrow band without a lumen

as far as segment xil.; the cells lose their regular epithelial

arrangement, and the solid cord is, seen in section, smaller than,

or sometimes about the same size as, the ventral nerve-cord ;

there may be small spaces here and there, filled with fluid; there

are large blood-vessels around it, in close contact with it.

Behind the sperm-sac there are small cavities in the solid cord of

disintegrating cells; but after segment xvii. the intestine, though

still narrower than in the normal worm, retains its proper form,

and is iined with columnar cells surrounded by chloragogen

cells; the lumen is either empty or contains some granular

matter. Although the gut is thus degenerating in the anterior

part of the body, the nephridia are normal, the blood-vessels are

larger than usual, and the same is the case with the cerebral

ganglion ; the specimens manifested the characteristic wriggling

movements, and were thus in no way pathological.

When such specimens were kept under observation for three

days, their anterior portions, containing the genital organs,

SEXUAL PHASE IN INDIAN NAIDIDA. 463

separated off as a sort of cocoon, while the hinder part of the

animal lived for some time, but was unable to regenerate and

ultimately died. It appears, then, that death ensues after the

full attainment of the sexual phase, and the cocoon is probably

the whole anterior region of the worm which has been separated

off.

BRANCHIODRILUS HORTENSIS Stephenson,

Many examples of this species were collected during the last

two weeks of October and the first two weeks of November, from

the mud at the bottom of a pond at Sikandra; and of these

nine were found to possess fully developed sexual organs (text-

Text-figure 3.

2 & AT ee r

Saad KS

= = a —

———$SSSS

ene ur | Sy

rage aS em at

Sp.S. sep.$/ femf£ Ss.

Genital region of Branchiodrilus hortensis. The atria lie one on each side of the

sperm-sac, and the vas deferens is seen running internal to the atrium of

its side; the funnel lies far behind in the sperm-sac. Pr., prostate; the

remaining letters as before. XX ca. 125.

The clitellum occupies segments v.—viil.; it is opaque white in

the living animal. It is about ‘05 mm. thick; the cells have a

coarse reticular structure, the meshes of the network being clear

spaces; the nuclei are indistinct. The clitellum is absent

between the spermathecal openings and on the ventral surface in

the anterior portion of segment v. It is formed after all the

other sexual organs have been developed.

‘The testes had disappeared in all the specimens examined.

The sperm-sac may reach as far back as segment xix, or Xx.,

in two cases to xxvi.; usually it extended to xviil.

The male funnels are within the sperm-sac, some distance

behind its mouth, and here they nearly fill up the available space.

Their lips are everted, and their outer margin is attached to the

wall of the sperm-sac, The width of the funnel is 90; its cells

are high and ciliated, with nuclei at their base. The cells and

cilia of the upper lip seem to be taller than those of the lower.

The vas deferens is 30 » in diameter; it is a fairly long tube,

about *2 mm, in length, and consists of two parts, a posterior

4.64 PROF. HARU RAM MEHRA ON THE,

longitudinal and an anterior ascending portion, ‘The longi-

tudinal portion passes forwards from the funuel, and is abbut

105» im length ; its last part lies over the upper wall of the

sperm-sac. The tube then bends upwards; the vertical ascending

portion, about 90 p in length, lies just bebind septum 5/6. Ina

few specimens the longitudinal portion formed a distinct curve

towards the ventral body-wall before rising to be continued into

the ascending portion. The cells lining the vas deferens are

about half as high as those of the funnel, have oval nuclei at the

base, and are without distinet cell-outlines; there is a thin

covering of muscle-fibres outside the epithelium. The tube

enters the atrium on its anterior face much above the middle ;

the ascending part of the duct as it opens into the atrium is

surrounded by a thick coat of muscle-fibres, continued onto it

from the muscular covering of the atrium ; the change from the

cubical cells of the duct to the columnar cells of the atrium

is sudden.

A bundle of muscle-fibres directed upwards from the ventral

-body-wall is attached to the ascending portion ; and a few fibres

connect it above to the dorsal body-v ral behind septum 5/6.

The atrium is a large pear-shaped body, taking up nearly the

whole length of its segment, 230-240 w in height and about

190 » in length antero-posteriorly ; in only one specimen were

spermatozoa seen in its interior. The epithelium is columnar,

the cells 18» by 15, with oval nuclei lying at the base ; outside

the epithelium is a thick coat of circular muscular fibres.

The ejaculatory duct, about 78 4 in length, opens at the top of

a tubular depression of the ventral body i yall ‘about 42 p in depth,

and is capable of being everted, when it projects slightly as a

short pseudo-penis. The epithelium of the duct consists of

columnar cells with elongated nuclei, and has a thick investment

of muscle-fibres.

The “prostate” consists of a large mass of pear-shaped cells

around the ejaculatory duct; the cells contain granular pro-

toplasm and a large oval or rounded nucleus near the base.

A few muscle- fibres surround and enter the mass of cells from

the ventral body-wall. There are a few blood-vessels around the

atrium and prostatic cells. Two bands of muscle-fibres, arising

from the setal sae, are attached to the atvium behind. The male

opening lies internal to and at the level of the ventral sete about

the middle of segment vi

The ovisac, fanned by the backward bulging of septum 6/7,

and, as usual, enveloping the sperm-sac, is fil Jed with a large

mass of spherical and elliptical yolk granules about 7-15 in

diameter. The ova are fairly large, and lie in two or three

masses.

The female funnel, about 45 in height, lies over. the lower

portion of septum 6/7 near the ventral Sparictes in segment vi

No female opening was seen.

The large spermathecz occupy the fifth seement.. The ampulla

SEXUAL PHASE IN INDIAN NAIDIDA. 465

is somewhat heart-shaped, or ovoid and notched below where the

duet arises. Its size varies somewhat ; in height it may be from

207 to 270, its length antero-posteriorly rather greater and its

width rather less. ‘The ampulle ave filled with spermatozoa, and

are so large as nearly to fill up the whole segment, the remaining

organs occupying only a small space below their contiguous inner

walls. The part of the ampulla anterior to the duct is somewhat

bulged downwards, and is lined with columnar cells ; the part of

the wall behind the duct is lined with cubical cells, which

gradually decrease in height as they pass upwards on the posteri 10r

wall; the rest of the ampulla i is lined by a very thin epithelium

of attenuated cells whose outline is quite indistinct. Outside

the epithelium there is a thin coat of circular muscular fibres—

the only part of the wall visible over a large portion, on account

of the thinness of the epithelium. The spermathecal duct leaves

the ampulla below, nearer its anterior wall, and is about 130-140 p

in height, including the depression of the body-wall where it

opens to the exterior. The duct is narrow above and below, but

somewhat swollen in the middle; its epithelium consists of

narrow columnar cells having elongated nuclei. There is a thick

covering of muscle-fibres outside the epithelium. The sper-

mathecal opening lies internal to and at the level of the ventral

set of the fifth segment, some distance behind septum 4/5.

The penial setee are the modified ventral sete of segment vi.

They are two or three in a bundle, somewhat hooked at the

distal end, which is not bifid. In length they are about 132

the shaft consists of a distal narrow portion about 36 u long, and

a proximal stouter part 96 » long; there is no distinct nodulus,

but the distal narrow portion is bent outwards and thus not in a

straight line with the proximal segment (text-fig. 1).

References to Literature.

1. Bepparp, F. E.—‘ A New Branchiate Oligochete (Br anchiura

sowerbyt).” Quart. Journ. Micr. Sci. vol. Exxi., 1892.

2. STEPHENSON, J.—“On Branchiura sowerbyt Beddard, and on

a new Species of Zimnodrilus with distinctive characters.”

Trans. Roy. Soc. Edin. vol. xlviii. pt. 2, 1912.

3. STEPHENSON, J.—‘‘On the Sexual Phase in certain of the

Naidide.” bid. vol. li. pt. 4, 1915

4, SrepuHEenson, J.—‘‘On Hemonais laurentii, sp. n., a represent-

ative of a little-known Genus of Naidide.” Jbid. vol. li.

pt. 4, 1915,

Vs i J, ‘a

f Vey ,

a iy

, cp

ON THE FLIGHT OF FLYING-FISHES,. 487

29. Observations on the Flight of Flying-Fishes.

By HE. H. Hanky, M.A., Sc.D., Agra, India*.

[Received August 4, 1920: Read October 19, 1920.]

(Text-figures 1 & 2.)

A point hitherto overlooked in the study of the flight of flying-

fishes is that the air is suitable for their flight to very different

degrees on different occasions. In this respect their flight

resembles that of soaring birds. This statement may be illus-

trated by the following examples.

1. Flight under unsuitable atmospheric conditions.

On the Ist June, 1920, at about a quarter of an hour after

sunset, the ship on which I was travelling across the Arabian

Sea was disturbing groups of small flying-fishes at the rate of one

or two groups per minute. The surface of the sea was either

glassy o1 disturbed by ripples too small to be easily visible.

Each fish, on emerging, jumped out of the water so far that,

while the body was supported on the outstretched ‘“ wings,” the

end of the tail was still immersed. This organ was thereupon

wagged vigorously from side to side, as is usual when starting,

thus forming a trail of ripples in the water. After proceeding

thus for the unusually long distance of four or five metres, the fish

raised its tail from the water and began to glide. The length of

the glide made by each fish was, at first, about a metre, Within

a few minutes a change was observed. The fishes of each group

disturbed by the ship made shorter and shorter glides, until at

length each fish fell into the water immediately it ceased to move

its tail.

One fish flapped its wings at starting, but made no better glide

than the others.

About half an hour previously, in sunshine, the ship had also

been disturbing small flying-fishes, which had flown for such

distances as are usual. Just before sunset the first symptoms of

lessened suitability of the air for their flight were observed. The

fishes began to show lateral instability. It may be noted that

both vultures and flying-fishes are more apt to show lateral

instability late in the afternoon than at other times of the day.

The stage of lateral instability shown by the flying-fishes soon

passed off and was replaced by one in which the course of each

fish, instead of being a horizontal straight line, was undulating.

Hach fish showed two or three alternations of gain and loss of

height before falling into the water. This condition soon

changed to that first described.

* Communicated by C. Tater Ruean, F.R.S., F.Z.S.

468 DR. E. H. HANKIN ON THE

In view of these gradual changes in the performances of the

fishes, there seems to be no room for doubt that they wished to

get away from the neighbourhood of the ship, by air and as

quickly as possible, throughout, and that the air was getting

progressively less and less fitted for their flight.

On the other hand, it has been observed by me on many

oceasions that the air remains suitable for the gliding flight of

flying-fishes after sunset if wind is present.

Flight wider fuily favourable atmospheric conditions.

During the same voyage, in sunshine and in the presence of a

hight ain flying-fishes were seen by me to fly at a uniform

height hoe the water till they were out of sight. This was

noted on two or three occasions. On a previous voyage I had

seen a flight till out of sight under cloud in a monsoon wind, A

binocular was used for these observations. Usually the longer

flights appear to be between 200 and 400 metres in length. Ina

flight of this kind the following phenomena may be noticed :—-

The fore wings (pectoral fins) are usually in the “ flat” position,

i. e. extended in the horizontal plane. Sometimes the wings are

slightly inclined upwards. In this case the outer part of the

wing a at a higher level than its base. This may be called the

SOND) 7 ’ position. Rarely the wings are inclined very shghtly

downwards. This may be deseribed as the ‘ down ” position.

This latter disposition, which I was only able to see distinctly on

my recent voyage, is probably that used for flight at highest

speed, as in slow-speed flight the wings are inclined upwards to a

strong degree.

Thus in 1 respect of its wing-disposition the flying-fish re esembles

the soaring vulture, for crn aries have their wings in the ‘“ up’

position for slow-speed flight and use the “ flat’ wing disposition

for flight at high speed. <A further resemblance is indicated by

the following very unexpected observation :—

Tf the flying- -fish is very carefully observed the extreme wing-

tips, for nearly an inch of their length, may be seen to be bent

up forming an angle of perhaps 45 degrees with the rest of the

wing. If “the sun is not far above the horizon, and if the fish is

travelling away from the observer in a poutuherly (presumably also

in a northerly) direction, then the wing-tip furthest from the sun

appears thicker than the other. This appearance has been seen

by me quite clearly and definitely in flying-fishes of two different

‘species. The appearance seems to be due to the upturned wing-

tip being bent round so that it has a negative angle of incidence

and hence, when the sun is low, the ahaclerasials of one wing-tip is

seen in shadow. When the sun gets higher the visibility of each

wing-tip is reinforced by shadow in this way. But, at best, the

upturned wing- -tip is by no means easy to see.

The bent-up wing-tips show no appearance of vibration. They

appear as if held in their position by a steady force, As I have

FLIGHT OF FLYING-FISHES. 469

stated elsewhere (‘ Animal Flight,’ [iffe and Sons, 1914), the wing-

tips of vultures in horizontal soaring flight show the terminal

quills bent up as though they were under the influence of a

steady force acting from below and behind.

As the fish is emerging from the water an interesting appear-

ance may sometimes be seen. The length of the trail (made by

lateral movement of the tail) is generally less than a metre and a

half. It may be only half a metre. At the end of the trail in

Text-figure 1.

Flying-fish starting, showing trail of ripples made by movement of tail.

Wings up and tail down as in slow-speed flight.

a few observations the fish appeared to make a sudden jump out

of the water, gaining height thereby to the extent of fowr or

five inches. Had this appearance of a jump been due to an extra

strong stroke by the tail, one would have expected to have seen

an extra large ripple in the water. Nothing of the kind was

observed. On other occasions, when the jump was not seen and,

I believe, did not occur, the trail was observed to get fainter

towards its end (text-fig. 1).

470 DR. E. H. HANKIN ON THE

3. Speed of flight of flying-jishes.

During my recent voyage a flying-fish was seen flying on a

course parallel with the ship and at the same apparent speed.

During eight seconds, timed with a stop-watch, it remained fixed

in position relatively to the ship. The latter was travelling

through the water at 154 knots or 7# metres per second. The

direction of the real wind was learnt by noting the movement of

the waves, that of the apparent wind by the position of the line

of smoke from the funnel. From these data it was calculated

that the velocity of the wind was 4 metres per second and that of

the flying-fish through the air was 10 metres per second. This

estimate involves the assumption that the wind-velocity was the

same near water-level as at the level of the top of the funnel.

A similar observation on another flying-fish gave an identical

result. Later in the day the wind decreased, and then flying-

fishes travelling on the same course as the ship were seen to move

distinctly faster than the latter.

These flying-fishes were all gliding with tails hanging down and

wings “up.” That is to say they were in slow-speed flight. In

‘Animal Flight’ reasons have been brought forward for believing

that, when in high-speed flight (wings flat and tail up on a level

with the body), flying-fishes may attain speeds of more than 20

metres per second.

Flapping sometimes occurs at starting, especially in the

presence of wind and in the colder months of the year. The

above-described observations of the position of the wing-tips, and

also the sharp-cut appearance of the wings in species whose

wings are opaque, definitely prove that flapping does not occur

when once the fish is well under way. Hence flapping cannot

be invoked as an explanation of the speed maintained during

flight.

Sometimes—when it may be supposed that the air is not fully

suitable for flight, or perhaps if the fish is exceptionally frightened

by the ship—during its flight the fish lowers its tail into the water

at intervals of about two seconds and wags it to and fro for a

fraction of a second with a resulting inerease of speed.

4. Comparison with the speeds attained by vultures

in horizontal soaring flight.

That the speed of flight of flying-fishes is similar to that of

soaring vultures when in horizontal flight is shown by the

following figures :—

During April 1920, I made several measurements, in Agra, of

the apparent speed of vultures with the help of a Souchier

telemetre by the method described by me in ‘ Animal Flight.’

Information as to the velocity of the wind in which the vultures

were gliding was obtained from measurements made with the help

of balloons sent up by Mr. J. H. Field (to whom I owe my thanks)

FLIGHT OF FLYING-FISHES. 471

from the Agra Aerological Laboratory. The balloons were sent

up at the time I was making my measurements. On one day

seven balloons were let off while I was observing. From data

thus obtained, it was possible to deduce the speed of the vultures

through the air.

Another set of measurements were made by me during the

first eight months of the year 1915. In these cases, however,

the balloons bad been liberated four or five hours before my ob-

servations were made. An error is thus introduced into the

calculations which, it may be noted, is likely to be less on days

of light wind than in stronger winds.

The results of these two sets of observations agree in indicating

that vultures soar at higher speeds in stronger winds. The mean

speeds caleulated from my data are as follows :—

Observations made in

April January to

1920. August 1915.

| |

' Velocities of wind,in which | Mean speed of vultures, in

the vultures were flying, | metres per second.

in metres per second.

| a | 116 11:7

0-3

4—6 | idee) 14]

7-9 15:0 17:2

10-20 20°0 Ue?

The above figures are based on 122 observations made in

April 1920 and 696 observations made in 1915.

Reasons have been adduced by me for believing that the

vultures on which the above measurements were made, were, as

they appeared to be, actually in horizontal flight and that their

speed, which they can maintain indefinitely, cannot be explained

by loss of height. An additional reason may here be mentioned

for this belief. It is that if a vulture glides, in soarable air, with

even a small loss of height, its speed will very greatly increase

beyond the above figures. For instance, in April last a measure-

ment was made of a speed of 42 metres per second in a vulture

flying nearly with the wind. The wind had a velocity of only

22 metres per second at the height at which the vulture was

flying. Hence the speed of the bird through the air must have

been about 402 metres per second. As this speed was exceptional

and as the bird was travelling in the direction of a place where

vultures frequently settle, it was assumed that it was gliding with

loss of height, though no loss of height was seen, and it was not

included in the ealculation of the mean speeds above given.

If vultures in soaring flight attain velocities of ten to twenty

472, DR. E. H. HANKIN ON THE

metres per second and if their velocities tend to increase as soon

as they begin to glide downwards, it is obvious that they must

possess some means of checking speed. Otherwise they would be

unable to land without accident. In‘ Animal Flight’ two inethods

of checking speed, one used for high-speed flight, the other used

for low-speed flight, have been deseribed. Two analogous methods,

for slow and fast speed respectively, employed by flying-fishes

have now to be described.

5. Method of checking speed in high-speed flight.

This method, which depends on changes of position of the hind

wings (pelvic fins), has only been followed by me in species whose

wings are coloured. It would be quite beyond my powers of

observation to see the adjustment in those species whose wings

are more or less transparent. Of the species that have yielded

most opportunities for these observations may be mentioned one

with black hind wings and another whose hind wings have a

chocolate-brown colour. The observations about to be described

are very difficult to make, and the following account is, to some

extent, compiled from glimpses of parts of the adjustment seen

on different occasions.

The flying-fish starts its flight with the hind wings directed

outwards and backwards and extended in the horizontal plane.

At some distance, perhaps 50 metres, from the end of its flight, it

may be seen suddenly to lower its hind wings (pelvic fins) so that

they point nearly vertically downwards. On my recent voyage,

when using a binocular of magnification 10, 1t appeared to me

that, when the hind wings are in the down position, their plane

is parallel to the direction of flight. In this position the leading

edge only of the hind wing would offer direct resistance to speed

ahead. But on a voyage in 1914, when I was using an Aitchison

binocular of the very unusual magnification 25, on two occasions

the hind wings, when in the down position, appeared to me to be

slightly rotated round their long axis so that their hind margins

approached each other. On one of these occasions, during part

of the flight, one hind wing was less visible than the other.

‘This was probably due to its having been seen end on. When

the fish changed its course so that it was travelling directly

away from me the two hind wings became equally visible. With

the hind wings thus disposed it is obvious that the air passing

between them must be,so to speak, entering a funnel, and therefore

much resistance would be offered to speed ahead. It is probable

that these two dispositions of the hind wings are used successively,

for, within about a second of the end of the flight, the two hind

wings are suddenly rotated through nearly a right angle round

their iong axes. he result of this maneuvre is that the plane

of the hind wines aequires a position at right angles to the

direction of flight. dn the first down position the hind wings are

not fully expanded. In the last position they are expanded to

FLIGHT OF FLYING-FISHES. 473

their greatest extent, thus offering the maximum resistance to

speed ahead. On one occasion the adjacent edges of the two

hind wings appeared to me to overlap. Almost or quite simul-

taneously with the full rotation of the hind wings, the tail is

lowered and the fore wings are placed in the up position. The

tail then touches the water, and the fish falls in with practically

no splash,

Text-figure 2.

A. Flying-fish in low-speed flight.

B. Hind wings advanced at end of flight.

In monsoon winds the fish may place its hind wings in the

down position and even rotate them almost from the commence-

ment of its flight. If this is done no high apparent speed is

attained even in cases in which the fish is travelling at right

angles to the wind.

Much practice in observing flying-fishes was needed by me

before it was possible to see these adjustments of the hind

wings. On my last voyage, in June 1920, I was able to see

clearly the full rotation of the hind wings in flying-fishes of three

different species.

6. Method of checking speed in slow-speed flight.

Tf the flying-fish is making a flight at slow speed, as happens

more frequently in the colder than in the hotter months of the

year, and as happens usually in the absence of wind, an entirely

different means of checking speed is employed and one which is

much more easy to observe.

During slow flight the fore wings are strongly inclined up and

the tail hangs down. The hind wings, as in high-speed flight,

Proc, Zoo, Soc,—1920, No. XXXII, on

474 ON THE FLIGHT OF FLYING-FISHES.

are directed outwards and backwards. Towards the end of the

flight the hind wings are somewhat suddenly advanced (text-fig. 2).

They still remain extended in the horizontal plane. The effect of

this movement is to bring the lift of the wings as a whole further

ferward in respect of the centre of gravity. ‘The fish consequently

rotates round its transverse axis. It is a rotation upwards,

i.e. the head goes up, the taildown. The angle of incidence of

the fore wings is consequently increased to such an extent that

their plane assumes a large angle with the direction of flight.

Consequently they present great resistance to speed ahead. ‘The

speed rapidly decreases and the fish drops. The tail is the first

part to be immersed, and the animal falls in with a very small

amount of splash.

7. Termination of flight without checking speed.

A species of flying-fish is known to me which is of small size

and which generally appears in groups of eight or ten individuals.

They do not, as a rule at least, fly either so far or apparently so

fast as individuals of other species that have come under my

notice. Their flight also differs in two other respects. First,

they usually fly in a straight line and do not follow the water-

surface as is generally the habit with other species. Secondly,

at the end of their flight they steer downwards by retiring the

wings through an angle of about forty-five degrees. They then

plunge head “for emost into the water without any visible attempt

to check their speed.

It is possible that this species 1s identical with Haocwtus evolans.

This is the name of a species, or of a group of closely allied

species, which differ from other flying-fishes in that their pelvic

fins are small and placed far forward. Hence these fins are

unfitted to function either for checking speed or for steering in

the vertical plane.

It is probable that the frequent shortness of the flights of

flying-fishes is often involuntary. My observations indicate that

flying-fishes sometimes behave as if they had made a mistaken

estimate of the suitability of the air for their flight. On one

occasion I noticed that they were starting with their wings

disposed for slow-speed flight, that they immediately changed over

to the high-speed disposition and then, at once, fell into the

water with a splash. On my recent voyage I noted, on one

occasion, that flying-fishes were starting with tails up (7. e. as in

high-speed flight). They then, either at once fell into the water

or lowered their tails and flew on. On other occasions when

groups of small individuals emerged they all at once fell in with

splashing or, in some cases, while the rest splashed in, one or two

got away and made flights of normal length,

ON THE LIFE-HISTORY OF THE LADY-BIRD BEETLE. A475

30. Observations on the Life-History, Biology, and Genetics

of the Lady-bird Beetle, Adalia bipunctata (Mulsant).

By Onzra A. Merrirr Hawkes, M.Sc. (Birm.),

B.Se. (Lond.) *.

[Received July 3, 1920: Read November 2, 1920.]

ConTENTS.

Page

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e. The Imagines .. : PR Cree Bare ce ees

ihe Tie Hnemies of the Lady-bitd ag ote wa tadeee: eS

g- Hibernation ......... ya agtid cee ee A

4. The Genetical Relations of fe Towetion Meco cc pando un tele

Ge STATE cic uc op NOt EER OC EEE DE OE REET ERCP RCeT ame E ME RrcCD Sc Cae Peels:

1. Lntroduction.

During three years of breeding pure and hybrid silkworms

(Philosamia), I have been increasingly impressed by the apparent

reasonableness of what would be regarded as fluctuating varieties ;

and by reasonableness, | mean a genetic relation between the

variations of the parents and those of the offspring. To under-

stand such variations, what appears to be needed is a large

number of exhaustive studies of normally occurring varieties in

animals untouched by Man. Insects present a large number of

variations, but little or nothing is known of their inheritance.

The Japanese have made extensive studies of silkworms (largely

domesticated), and in America, Roswell Johnson (13) has made a

wide study of Hippodamia, as a result of which he has been able

to make the following important statement {13 ):—‘“‘ Every variety

or noticeable variation of Hippodamia that has been tested is

inheritable in some degree.” Miriam Palmer (21, 21) has also

made successful studies of the American varieties of Adalia and’

other Coccinellids. Meissner (19, 20) and Schroder (23), in

Germany, have made a small number of breeding experiments

with some of the varieties of Adalia bipunctata.

Work of this kind would have the added advantage that in

some cases the sciences of genetics and classification would be

able to go hand in hand, instead of, as is too usual, being mere

bowing acquaintances.

Adalia is highly suitable for work of this character, as there

* Communicated by BE. Green, F.Z.S., F.E.S.

32"

A76 MRS. 0. A. MERRITT HAWKES ON THE

are many easily procurable varieties, which, as far as is known,

are not regional.

Palmer (21) worked for some time at Adalia, but she had only

a few varieties to deal with, as in America there occurs only the

“type” and three varieties, whilst in Hurope there are more than

twenty-six named varieties. It would be impossible to get as

quick results here as in America, as two generations per annum

are the most one can obtain in England under natural conditions,

whereas Palmer succeeded in obtaining four and five during the

long American summer. Our erratic climate also makes the

food-supply (aphides) more uncertain than under more stable

climatic conditions.

Coecinellids have been extensively studied in the United States,

as they are there recognised to be of considerable economic

importance; but in England there has been little careful study

of their habits, their distribution, or the genetical and possibly

regional relations of the varieties which are so numerous in some

of the species. Donisthorpe (6) has published the only complete

history of any species, Coccinella distincta. The following obser-

vations were made in the course of genetical experiments which

are still in progress.

The varieties of <Adalia have been determined chiefly by

Meissner and Schroder, who based their determination principally

upon the variations in the colour of the elytra, but neglected the

very considerable variations of the thorax. Schroder believes

that the varieties from pure red elytra at the one extreme to pure

black at the other are modifications of a scheme of seven spots

and a scutellar mark. This is an interesting matter of specu-

lation, but we do know that in Europe there are two principal

forms: (1) “type,” in which the elytra are a reddish brown

with an approximately round black spot in the middle of each

elytron; and (2) a black form with either two or three red spots

in each elytron: when two are present, an upper lateral and a

mid-dorsal, the variety is known as var. 4-maculata; when three

are present, an anal red spot being added to the above two, the

variety is named 6-pustulata. All other varieties, none of which

occur in large numbers, can probably be grouped about these two

main forms.

The Red varieties are generally acknowledged to be commoner

than the Black, though very few accurate statistics have been

recorded. Meissner found 35:2 per cent. Black in Summerfeld

and 42:3 in Potsdam. In Staffordshire and Warwickshire I

found 29 per cent. Black in 1918, 25 per cent. in 1919. In 1920

I found 2°75 per cent. Black at Crouch Hill, London, N., 42-7 per

cent. Black on hops at Bishop’s Frome, Herefordshire, 11-7 per

cent. at Enville, Staffordshire, and, very much to my surprise,

78°5 per cent. Black in che Edgbaston district of Birmingham.

Further observations will show whether localities are characterised

by certain varieties or whether the prevalent varieties vary from

time to time, If the varieties remain the same, will it mean that

LIFE-HISTORY OF THE LADY-BIRD BEETLE. AT7

there is little or no migration, that hibernation takes place

locally, and that the varieties breed true? Or, does the particular

environment allow only certain varieties to survive 2

Two, possibly four, factors have to be considered in a genetic

study of the Black and Red forms :—

1, Pattern: that is, the shape and distribution of the spots.

2. Colour: (a) black, which is the same in both “type” and

black varieties ;

(b) ved, which, contrary to accepted opinion, I believe

different in the Red and Black forms.

3. Size: there is great range in size in the Coccinellidee, which

Johnson believes is chiefly adaptational, but the mean size

of the varieties under consideration is different, and perhaps

constitutes a definite character.

4, There may be a physiological factor in the reaction of these

varieties to heat.

Microscopical examination by transmitted light shows that the

reddish colour in both varieties is due to a clear yellow, to which

brownish-red, semi-transparent particles are added in the Red

form and red-brown and carmine in the Black variety.

There is probably a physiological factor to account for the

seasonal appearances of the two forms, in England the Black

usually appearing later and disappearing earlier than the Red.

I have also observed that when the breeding-boxes were over-

heated, the Black could live at a higher temperature than

the Red,

The Food and Feeding of Adalia bipunctata.

There is considerable ditticulty in feeding Adalia with its food,

living aphis. The occurrence of aphis is very erratic, being much

affected by sudden climatic changes, such as cold, wind, and heavy

rains. At times they disappear inexplicably.

Adalia will eat many but not all species of aphis. During two

years breeding [ have found three species which are not accept-

able :—(1) Aphis rumicis, the black bean aphis which is found on

broad beans in early summer. The young of this species are

eaten to a small extent, but the large only under stress of circum-

stances. One may continually find a bean-row black with this

destructive aphid, but very few lady-birds upon them when, on

a bed of nettles a few yards away, can be found large naeEs

of Coccinellide—C. septempunctata, C. variabilis, A. “bipunctata,

Halyzia 14-guttata, I, 22-pwnctata, and H. 14-punctata,—even

when the aphids are not obviously numerous. (2) A bluish-green

aphis (Macrosiphum aconitum (Van der Goot)), which I have

found only on Monkshood. ‘This has never been eaten by Adalia.

Johnson says ‘‘ several of the highly-coloured aphis cannot be suc-

cessfully fed to Coceinellids.” (3) HTyalopterus prunti: this aphis

478 MRS. O. A. MERRITT HAWKES ON THE

has a grey-green mealy exudation, which fills the stomata of the

larvee and so kills them. When a Victoria plum was covered with

this aphis, no larvee or imagines were ever found among them,

and very few larvee of the Hover-fly, which is probably an even

more prolific eater of aphis than the Coccinellide.

The presence of large numbers of Coccinellide on the Stinging

Nettle may be due to the fact that these insects are not univer-

sally carnivorous. C. septempunctata certainly eats the abundant

pollen and possibly other products of this plant. Donisthorpe

found that C. distincta ate the honey supplied to ants, and I

have found C. sepiempunctata eat the exudation from bean plants

and Jerusalem artichokes in the autumn. During the summer

I have observed 4. bipwnctata apparently eating the juice of rasp-

berries and split plums,

The Adalia larve in the first two instars have to be fed with

minute aphis, as, although sufficiently pugnacious to attack large

aphis, the latter can easily shake them off.

Any species of aphis were used in 1918, but in 1919 practically

two species only —in early summer Aphis prauni Fab., which attacks

damson-trees; and for the remainder of the year Aphis fagi,

which lives only on the beech. Although the latter is slightly

woolly when full grown, it is in the earlier stages very acceptable

to bipunctata. The copper-beech is attacked more frequently

than the green beech, and on the former numbers of bipunctata

are to be found all the summer. The supply of beech aphis has

the advantage of being fairly constant, as the leaves and branches

ave so flat and so close together that it takes a great deal of cold

and rain to clean the aphis off the tree. The flat leaves, which

do not wither readily, are very suitable in boxes. The aphis of

the rose (Siphonora rose) and of the nettle (Aphis urticia) both

die quickly in boxes, presumably owing to the wilting of the thin

leaves of the food-plants.

In August 1920, after a variety of experiments, I found that

the aduits will eagerly eat pounded dates, upon which they can

live for months. The newly hatched larve cannot use this food

at all, but I succeeded in bringing one larva from the second

instar to the perfect insect on dates alone, the imago being of

average size. Most of the older larve kept alive on this diet for

some time, but did not grow much and ultimately died.

I have obtained the best results by feeding in closed boxes.

The boxes must be opened every day, not only to introduce fresh

food but also fresh air, which is of paramount importance to this

species. Great care has also to be taken to remove all mouldy

leaves. A mould (probably Cladosporium aphidis Jhiim) quickly

grows on the honey-dew produced by the aphis, especially that

found on beech and sycamore leaves. This mould fatally attacks

the larve. It first appears on the thorax, but its method of

producing death is undetermined.

The main difficulty of rearing is the cannibalistic habit of not

only A. bipwnetata, but also C. septempunctata, C. variabilis, and

H. \4-punctata. The larve eat eggs and one another in either

LIFE-HISYORY OF THE LADY-BIRD BEETLE. 479

the larval or pupal stages. This phenomenon is not due to con-

finement or lack of food, as it takes place in the presence of

abundant food and in normal natural conditions. One can easily

watch what appears to be a form of Coccinellid degeneracy among

the larvee which frequent red and black currant bushes. I have

seen adult bipunctata and septempunctata eat their own eggs in

boxes and on the trunks of trees, but never eat either larve or

pupe. This habit was also observed (6) in (. distincta in con-

finement. Palmer has, however, observed the imagines eating

larve when there was no other food in the cages. On one

oceasion I watched a bipunctata eat the abdomen of a small fly

caught in a spider’s web. After a period of experimental star-

vation, two Adalia bipunctata feasted upon a small weak relation.

Cannibalism has also been noticed by Palmer, Johnson, Burgess,

etc.

On hatching, the almost colourless larvee remain clinging to the

empty egg-shells, part (never all) of which forms their first meal.

They next fall upon one another. This same early habit of

cannibalism also occurs in Mpilachna (27), the vegetarian Lady-

bird. If by chance an egg lags behind the others, it never

hatches, for it is eaten by a larva which was born a little earlier.

The larve remain on the egg-mass until fully coloured, and then

suddenly and simultaneously disperse. Approximately a quarter

of the larvee are eaten before this dispersal.

During the first instar there is great mortality, partly owing

to lack of sufficiently small aphis and partly to cannibalism. The

young larvee do best in small boxes, as they seem incapable of

wandering far at this stage to find food, which, I believe, is

chiefly obtained by good luck and persistent wandering. As

early as possible the larvee were isolated in separate boxes, but L

was inclined to think cannibalism in the early stages a physio-

logically highly suitable diet, and, with this in view, I fed the

bipunctata with small larvee of other species. This practice was

not, however, always successful, as sometimes the eater became

the eaten. Partially successful attempts were made to feed the

larve on cooked and raw hen’s egg. Coccinellid eggs were

always eaten when put in the boxes, but this is not a suitable

proceeding in the early instars, as an odd egg may escape and

hateh. During 1918, 21 of 22 families were reduced to one, as

the full significance of cannibalism was not then recognised.

Jt is interesting to consider the intermittent competition

between larvee of the same batch of eggs. It is not the larva

which first starts in the race which necessarily first reaches the

winning-post. It may be the first to reach the end of the first

instar, but during the short qwuescent period which precedes

ecdysis, it becomes a defenceless prey to those who have lagged

behind. This special danger recurs at the end of each of the four

instars.

The method of attacking the pupa is always the same, the

larva beginning on the ventral side near the anal point of

attachment and eating its way upwards and forwards, half

480 MRS. O. A. MERRITT HAWKES ON THE

burying itself in the process. The head, the thorax, and the

tough dorsal skin are frequently left untouched.

There is, however, no special mode of attacking the aphis, and

certainly no painless method of dining. The bipunctata catches

hold of a passing aphis and at once proceeds to devour it in spite

of its struggles, which continue far into the meal. The legs, the

wings, the head, and part of the cuticle of abdomen and thorax

are usually left. The meal may take as long as five minutes.

The mother bipwnctata is not very wise in her choice of a posi-

tion for her eggs—many are laid on leaves, but also many on the

branches and trunks of trees. J have even found them on wooden

palings and on stones by the side of garden-paths. As the young

larvee cannot wander far in search of food, and as the parent is

lacking in the instinct to lay eggs where there would be an

adequate larder, the capacity for eating eggs and one another may

be of considerable biological value. Tinted knowing what we

do of aphis, one realises that 1t would indeed be a wise Lady-bird

that could guarantee aphis for its newly-hatched young. The

period of incubation varies from three to nine days, and in that

time the aphis might have disappeared. In May 1918 the

damson-trees were covered with aphis for three weeks ; without

any apparent change in the weather, the aphis began suddenly to

descend the tree-trunks in myriads, and at the end of two days

hardly an aphis could be found. I have found eggs laid on the

petals of a rose, but the petals fell before the eggs hatched.

The use of cannibalism was well demonstrated in July 1920.

An isolated gooseberry-bush had every shoot attacked by aphis,

upon which large numbers of bipwnctata larvee were feeding. At

the time when the larve were beginning to pupate, a few cold

rainy days caused the aphis to completely disappear. At once

the larvee began to attack each other and the pup, and, finally,

themselves pupated on an entirely cannibal diet. J saw no signs

of migration from this bush on account of shortage of aphis.

Cannibalism is probably of use in preserving the race during

periods of aphis famine, as the larve, unlike the adults, cannot

live long without food. Cannibalism is not of the same biological

use to the imago, as its power of flight gives it a large range,

and, when necessary, it can remain long periods without food.

I have starved Lady-birds for three weeks, and at the end some

of the males were capable of successful fertilization.

3. Notes on the Life-History.

a. Mating.

Mating takes place in captivity in the presence of plenty of

light, especially direct sunlight. ‘The response to light is very

rapid. Immediately a box containing a number of Lady-birds is

put im the sunlight, they begin to move quiekly, fly, and mate.

The same activity does not ie place if the box is warmed with-

out light. Fresh air, especially moving air, is also conducive to

mating.

A. bipunctata is both polyandrous and polygamous. When a

LIFE-UISTORY OF THE LADY-BIRD BEETLE, 481

male and female are isolated in a box, mating takes place at

intervals during a period of two to nine weeks. Copulation lasts

from half to three-quarters of an hour. It takes place when no

eggs are laid and when the egg-laying period of the female is

finished. Palmer states that the female lays fertile eggs three

weeks after fertilization. Mating takes place in Staffordshire

and Warwickshire during the whole summer, but the principal

season is May to June, with a subsidiary season trom the middle

to the end of August. Probably only a very few of the newly

emerged Lady-birds breed until the following year. The insects

have been found mating on sunny days in late ‘September, but at

that time no eggs were aid.

b. Lhe Eggs.

TABLE [.

Detailed Record of Heth es 33 eee i

Dates of Number of Eggs Incubation. , Larval life. | Pupation. | Number of

Oviposition. | and Colour. (Days.) | (Days.) | (Days.) Imagines.

(1) 22.7.19 | 16 (yellow) or |

(2) 22.7.19 24 (orange) } 2 | ae e f

(3) 23.7.19 | 8 (deep orange). 9 16 7 1

(Ny eer a I (on yh 6 | — = =

(@) Zerstrs Wy WC hat hs 6 | 17 13 3

(6) 27.7.19 3 (yellow base, 6 | 24 7 1

orange tip). |

(7), *28R 7019 21 (orange). if | 17 8 2

(8) 29.7. 19 26 (yellow hase, 5 | 22 9 1

orange tip).

(9) 1.8.19 25 (deep orange). — | = = =

(10) BBY Siow yi ear) re), 6 | 19 ss =

(GED Peon Sones aay 3 18 7 9

(12) 9.8.19 BCs mies 3 | 24 Uf i

| Total 185 Av. 5% | Av. 202 Av. 82 Total 24

Mating took place on 17.7.19 between two Red “ type” Adalias. In batches (1)

and (8) the larvee had light rings around all the abdominal and thoracic

tubercles: hence these larve appeared much lighter than the larvae of the

remaining batches, which only had light rings around the lower lateral

tubercles. The resulting imagines were all red, those from these two batches

being indistinguishable from the others.

The eggs are spindle-shaped and laid in batches of from three

to fifty. The outer shell is colourless and covered with numerous

protuberances. The colour of the egg is due to the yolk, which

is usually yellow but may be orange—a range of shades similar

to those of the yolk of the hen’s egg. Usually the whole batch of

eges laid on a single occasion 1s of the same uniform colour, but

they may vary (see Table I.). In several batches the eggs shaded

from yellow at one end to orange at the other. The eggs do not

change in colour until a few hours before the larvee are to

merge, when they become a greenish grey.

482 MRS. 0. A. MERRITT HAWKES ON THE

The average number of eggs laid was between 140 and 148, the

largest numbers laid were 167 in 1918, 189 in 1919, and 418

in 1920. ‘The largest number laid in twenty-four hours was 80.

The Black and Red forms are not differentiated by the colour,

size, or shape of the eggs or by the number of eggs laid.

Clausen (3) found 190 the average number laid in America.

The eggs are usually laid on the under side of leaves, but may

also be found in slight depressions on the trunk and branches ;

on the trunk they are usually on the north side or at the angle

where a branch originates. The bright-coloured eggs are very

conspicuous on the brown bark. On the lower three feet of the

trunk of a plum-tree badly attacked by aphis, I found within a

week 42 patches of Coccinellid eges-—A. bipwnctata, H. 14-guttata,

H. \4-punctata, C. l1-punctata, and C. variabilis. Evidently

there is a tendency to lay the eggs away from the light, but I

have watched a septempunctata lay her eggs on a stone in bright

sunlight. The usual period of incubation is three days, but it

may extend to nine. ‘he average laying period of the first two

years’ observations was 24 days. Incubation varied from 3 to 13

days, the average being 5 daysin 1918, 4 in 1919, and 77 in 1920.

Clausen (United States) finds the average incubation 5 days and

the average egg-laying 28°2 days.

ce. The Larve.

The larvee when about to pupate are of a blackish-brown colour.

The first upper lateral abdominal tubercle is always orange, and

an orange patch extends across the abdomen between the fourth

dorsal abdominal tubercles. This orange patch may be succeeded

by similar patches of decreasing size and colour-intensity in the

5th, 6th, and 7th abdominal segments. In many cases there may

be orange on the lower lateral tubercles of the fourth abdominal

segment. The lower lateral tubercles may be surrounded by @

cream-coloured ring (see note to Table I.). There is great varia-

tion in the depth of colour of the orange patches.

IT have made many unsuccessful attempts to determine the

differences in the larvee and pupz which are destined to become

respectively Red and Black imagines. In 1918 the average larval

period was 27 days, the minimum 22 and the maximum 27. In

1919 the average was 23, the minimum 16 and the maximum 39.

Clausen found the average 16:1 days for the United States. In

1920 the average larval period was 35 days, the minimum 23,

and the maximum 37.

d. The Pupe.

The larvee usually pupate on the under side of leaves, especially

where the edge is curled up by aphis or drought. Pup are

sometimes found on the upper side. In 1918 very few pupz were

found on the upper side of leaves, but in the early part of 1919,

LIFE-HISTORY OF HE LADY-BIRD BEETLE, 483

when there was a great deal of wet and cloudy weather, a much

larger proportion was found on the more exposed surface. During

the sunny part of July and August 1919 the pup were again

varely found on the upper side. I have found no special orien-

tation of bipunctata pupe, such as is so striking in Anatis ocellata,

which both on the trunk and needles of the pine normally faces

upwards.

The average length of pupation in 1919 was 13 days, the mini-

mum 9 and the maximum 21. In 1919 the average was 17, the

minimum 3 and the maximum 28. In 1920 the average was

20 days, the minimum 16, and the maximum 22. Clausen and

Girault (10a) both found the American average 6 days. The

very short period of three days was produced by placing the pupe

in a temperature of 101° F.

e. The Imagines.

On emergence the imagines vary from palest yellow to orange.

Air and light usually produce the normal colour in a couple of

days, but at times the original orange may persist for weeks. At

certain seasons when bipunctata are emerging the majority

caught are of an orange rather than the typical red colour.

This observation applies also to sepiempunctata and variabilis.

The variations of colour so obvious in some collections are not

all true variations, but many ave probably due to the develop-

ment being still incomplete when the specimens were taken.

Although the Lady-bird which emerges in August or Sep-

tember may live to the following June, a period of ten months,

its longevity during the summer is much shorter, the male

sometimes dying after a mating-period of three weeks, although

the female has lived two months after laying her eggs. Captivity

would no doubt greatly modify this character.

The combined duration of incubation, larval life, and pupation

varied from 28 to 56 days. The shortest period occurred during

the very hot weather subsequent to July 22,1919. Burgess (1)

gives the complete period as 28 days in America, where the

temperature would be very similar to that during this particular

period in 1919.

Only a small percentage, 8 per cent. to 17 per cent., of the eggs

laid became imagines. Palmer’s results were better, as she

obtained from 13 per cent. to 25 per cent. of imagines.

f. The Enemies of the Lady-bird.

The colour of the Coccinellids is supposed to be protective, and

it is stated that they are not eaten by Insectivores. The yellow

excretion produced by both larve and imagines is supposed to

protect them from predaceous enemies. ‘The latter contention

appears to be partly based upon the fact that the excretion is

disagreeable to Man both as regards taste and smell. Certainly

484 MRS. O. A. MERRITT HAWKES ON THE

the Lady-bird has some enemies besides parasitic flies. In this

country the larvee are eaten to some extent by Syrphid larvee, for

which veason it was imperative to examine carefully all leaves

put in the breeding-boxes, as if any of these voracious Syrphids

were inadvertently introduced, they soon made an end of the

Coccinellid larvee as well as the aphis. Spiders also eat these

larvee, as they have been found sucked dry in spider’s webs on

several occasions. In England I have observed Flycatchers and

Sparrows taking septempunctata and bipunctata, and also found

considerable numbers of the elytra of septempunctata in the

droppings of Sea-Gulls when that Coccinellid was abundant on

the coast. Elytra of bipunctata and septempunctata have been

seen in the pellets of the Brown Owl. In America, Flycatchers

and young Swallows are stated to eat Lady-birds. There also

all stages of the Coccinellide are eaten by Mantis religiosa and a

Wood-Bug, Podisus serieventris (1).

g. Hibernation.

The periodical appearance of vast numbers of various Cocei-

nellids has long been known. These appearances are of two types :

(1) increased numbers of the insects in various districts of the

country during the spring or summer, in places where there were

or had been large numbers of aphis; and (2) large numbers of

the insects at specified places, usually in the spring or autumn,

where there was or had been no food. In the latter case the

insects are in close juxtaposition, and may occur in such quantities

that the term “masses,” used by Johnson, most fitly describes

the case.

Septempunctata have occurred in masses by the seaside in

England in the autumn, and Fabre (10) has observed them on

Mount Ventoux in the Vaucluse Alps at a height of 4500 feet

in June and October. This is a condition similar to Hippodamia,

which is constantly found in large masses on the western

American Mountains, where it is known that they gather to

hibernate. ‘The same phenomenon possibly also occurs in the

Alps and other Kuropean mountains. The masses by the sea are

probably pre- and post-hibernation gatherings. Johnson suggests

that the yellow excretion may play some part in gathering these

numbers together. If this is so at the time of hibernation, it

may also be a factor in gathering Coccinellide together for

feeding purposes.

In Staffordshire I have found hundreds of septempuunctata

swarming over sand-hillocks by the roads and on the commons,

on warm days in April, at a time when they are beginning to

emerge, in places where there is no sign of aphis. J have never

found bipunctata in large quantities, although they do collect in

small groups under bark for the winter hibernation. Mr. Alfred

Priest reported to me the presence of large numbers of bipwie-

tate in the neighbourhood of a disused chimney-stack im

LIFE-HISTORY OF THE LADY-BIRD BEETLE 485

September 1919. Karly in 1920 he visited this place, and found

large numbers of the insects lying massed upon one another in

the cracks between the bricks where there was practically no

shelter. The Lady-birds were on all sides of the chimney.

In September 1919 large numbers of septempunctata were

reported from various parts of England, but in Staffordshire

there were at the same time large numbers of bipunctata and

C. 11-punctata, which, on account of their small size, were much

less obvious. ‘This seen ance was entirely acc ounted for by the

weather-conditions. Early in August there had been a period

of warm fine weather during which aphis and Coccinellid larvee

flourished : this was succeeded by a wet, cold period, which no

doubt retarded the pups; but when a second period of hot

weather supervened, the retarded pupz responded, as my breeding

experiments show they will do, and simultaneously, instead Pe

over a protracted period, the adults a ppeared.

A, The Genetical Relations of the Varieties.

Matings were made during the summers of 1918 and 1919

During the first year, only 22 of the 57 fertile matings produced

a complete generation, the family in each case being reduced to

one imago. In 1919, 23 complete generations were produced

from 35 matings.

Eleven of these successful matings were Red x Red, and with

the exception of one, Mating 32, produced only Red offspring,

among which there occurred small variations from their parents

and from “type.” The Red bred true, producing none of the

Black variety.

The exceptional mating (32) consisted of two Reds taken when

mating. The female laid one small batch of eggs (7), from which

two Black imagines were produced. Clearly the female was at

the end of her productive period, and had probably mated with a

Black before she was caught.

Six matings of Black x Black were made, five of which pro-

duced both Red and Black imagines, a phenomenon quite

different from the Red x Red matings; but as it was not possible

to guarantee that the females had not had partners previous to

the scheduled one, it cannot at present be deduced that the

genetic constitution of the Black as regards the purity of the

genes is different from that of the Red.

The sixth mating (24), which produced only Blacks, was as

follows :—<¢ var. 4-maculata x @ 4-maculata produced five

4-maculata and one 6-pustulata. The female had certainly had

no previous inate.

The five matings of Black x Black produced 8 Blacks and

4 Reds. It must be noted that these Reds were all variations

from “Type” and not exactly like the Red parent. The problem

therefore arises—are these variations heterozygous forms with ay

intermediate appearance ? Similar varieties have, however, been

486 MRS. O. A. MERRITT HAWKES ON THE

found in the pure Red families, where, moreover, the variations

of the offspring appear related to those of the parents; if, how-

ever, these variations are the heterozygous forms, one would

expect to find some Blacks appearing when these Reds are mated

together, but, so far, that has not been the case. A large

number of matings will have to be made, as the numerical pro-

portions in which the varieties oecur in nature indicates that

there are certain points of genetic stability, and that many of the

intermediate varieties, 1f such there are, must have a ‘definite

relation to certain parent types.

The condition in bipunctata may be similar to that observed

by Lutz (17) in Crioceris asparagt. He says: “Thus we have a

good case of a perfectly graded variation (from spots discrete to

spots joined) obeying Mendel’s law. Spots joined even lightly

and on one elytron only make a ‘‘pure” recessive individual.

One of these intermediate forms seems characteristic of the (DR)

heterozygous form.’

Six matings as follows were made between Black and Red :—

m, 20 ¢ Red x 9 6-pustulata produced one Red.

m. 21 ¢ Red x @ 6-pustulata produced one 4-maculata, one

6-pustulata, two Red,

m. 22 3 Red x 2 4-maculata produced five 6-pustulata, three

Red.

m. 23 ¢ Red x 2 6-pustulata produced four 4-maculata, five

6-pustulata, two Red.

m. 25 ¢ 4-maculata x 9 Red produced one 4-maculata.

m. 29 § Red x 2 6-pustulata produced one 4-maculata, two

6-pustulata, one Red.

There is here no sign of dominance, but there is a large pro-

portion of Blacks (20) to the Reds (9). This proportion is

surprising, as the Blacks are normally in the minority according

to data so far collected. If the larvee destined to become Black

imagines have the same phvsiological response to heat as the

Black imagines (see section 1), their large proportion might be

accounted for.

So far, only two matings (m. 57 in 1918 and m. 25 in 1919)

were made between a Black ¢ and a Red 9; both these produced

Black offspring only, so that not sufficient data is yet at hand

to judge if there is a sex factor in the inheritance.

There are usually two medium posterior whitish spots on the

pronotum of the Red form, These may be absent, or the two

may be large enough to fuse in the middle line. So far, no

regular method of cater itance of these spots has been determined.

Palmer (1911) found “the presence of the whitish basal markings

on the pronotum (of annectans and melanopleura) dominate over

its absence in coloradensis and humeralis.” The degree of the

development of these spots is not related to the degree of the

development of the black dorsal spot in any way ; these white

spots are absent in the Black form, the pronotum of the Black

LIFE-HISTORY OF TILE LADY-BIRD BEETLE, A487

and Red differing thus in degree of pigmentation. But those

Red forms, in which the black spot even stretches as a band across

the elytra, do not necessarily have a more pigmented pronotum

than in “type.” If an increase in the size of the black spot of

“type” means a step forward in a melanistic direction, one would

expect this tendency also to affect the pronotum.

C. variabilis.

C. variabilis has two forms of imagines, which in general

colour arrangement are similar to the two principal forms of

A. bipunctata. Vhere is a Black form with ten red-ochreous

spots and a Red (ochreous) form with fourteen black spots. The

difference in the shade of the red between the two forms is

much more marked than in Adalia. The red spots on the Black

frequently tend to carmine at the edges, but the red ground-

colour of the Red form would more accurately be described as

ochreous; it is never as deep a red as occurs in A. bipwnetata

(type). I have no information of the percentages in which these

types occur. The Black form of variabilis is the “ type,” but

whether this indicates that it is the more common is unknown.

The dimorphism in bipunctata and variabilis is not equivalent

to that of Zina (18), in which there is a spotted and a melanic

form, for in that species there is certainly only one colour-pattern,

as the spots can be sometimes seen in the melanie form by

holding the wings up to the light. MacCracken (18) found the

melanism recessive.

Two matings of variabilis were made: (1) mating 17, ¢ Black

x 2 Red produced 4 Red imagines ; and (2) mating 18, ¢ Red

x 2 Black produced 4 Black and two Reds. As in bipwnetata,

there is no sign of dominance.

A third mating (10) was made between a ¢ Red bipunctata

and a @ Black variabilis, the result of which was 7 Red variabilis

and 5 Black variabilis. The two mated at intervals from

May 29 to June 26, and during that period laid eggsas follows :—

15 on June 3, 16 on June 4, 24 on June 14, 6 on June 18, and

15 on June 26; a total of 76. As the results were all variabdilis,

one cannot but suppose that the female was satisfactorily ferti-

lised by a male of her own species before she was caught. ‘This,

however, involves the necessity that some of the spermatozoa

remained capable of fertilization for 29 days at least ; but Palmer

states, p. 235 (21): ‘“‘A female Adalia would not seem to be able

to lay fertile eggs for more than three weeks after being isolated

from a male.”

On one point these two species differ considerably, for it is the

Black form of bipunctata which is the more variable in the

number of spots, whilst in variabilis it is the Red form which

is the more variable. The recurrence of the same type of

variation in species so nearly related and which constantly occur

together, makes an understanding of the one imperative to a

satisfactory understanding of the other,

488 MRS. O. A. MERRITT HAWKES ON THE

5. Summary.

1. dAdalia bipunctata has so many yarieties that it is highly

suitable for the needed study of inheritance of normally occurring

variations.

2. Adalia can be mated in captivity with ease, but their canni-

balistic habit makes it very difficult to rear them satisfactorily

in large numbers.

3. The time passed in the various stages 1s variable, depending

not only on the food, but also on the temperature, to which all

stages, but especially the pupal, make a rapid response.

4. Little is known of either hibernation or migration, but

apparently both septempunctata and bipunctata tend to collect in

numbers in the spring and autumn, and may at times be found

hibernating, piled upon one another, similar to the ‘‘ masses” of

Hippodamia found in the western U.S.A.

5, There is no evidence of dominance in crosses between the

two main forms—the Red “type” and the Black (var. 4-maculata

and var. 6-pustulata).

6. Eleven matings of Red x Red produced only the Red type.

7. Black and Red forms also occur in C. variabilis, and, as in

bipunctata, neither is dominant.

8. The pronotum of the Black form is darker than that of the

Red, but there is no progressive blackening of the pronotum of

the Red coincidentally with the increase in the size of the black

spots of the elytra.

9. Although it is possible to make a series from pure red

elytra on the one hand through various patterns to pure black on

the other, this cannot be regarded as proof that the variations

are merely fluctuating. The percentages of the variations is, as

far as is known, approximately stable, and certain types are far

more common than others. These facts, combined with the

observation that the children tend to show the variations of

the parents, strongly suggest that there are probably certain

points of genetic stability, and that only analysis is needed to

show that there is some regular method of inheritance,

I have much pleasure in thanking Dr. A. D. Imms for naming

several of the aphis species and Mr. W. B. Grove for naming the

mould, Cladosporium aphidis, which he believes has never before

been recorded as British.

LITERATURE.

1. Bererss, A. F. 1903—Economic Notes on the Family

Coccinellide. U.S. Dept. Agric. Div. Ent., Bull. 40

(New Series), pp. 25-32.

2. Bureess, A. F., & Conims, C. W. 1911.—The Value of

Predaceous Beetles in destroying Insect Pests, U.S. Dept.

Agric. Year-Book, pp. 453-466, ;

10.

LIFE-HISTORY OF THE LADY-BIRD BEETLE. 489

. Unausen, GC. D. 1915.—A Comparative Study of a Series

of Aphis-leeding Coccinellide. Journ. Econ. Ent. vol. viii.

pp. 487-491.

. Cuyzer, Beta. 1886.—Note on Cannibalism of Coccinellide.

Revatani Lapok. vol. iii. p. 107.

. Curtis, JoHN. 1860.—-Farm Inseets. London: pp. 71-73.

DontsrHorPE, H, 1919.—Myrmecophilous Lady-bird, Cocci-

nella distincta Fald.: its life-history and association with

Ants. The Entom. Record, 1919, vol. xxxi. No. 12,

pp. 214-222, & 1920, vol. xxxii. No. 1, pp. 1-3.

. Dunntne. 1869.—Discussion on swarms of Coccinelle, Nov.

1869 by Mr. Dunning. Trans. Ent. Soc. London, p. xxv.

. Ewine, H. HE. 1914.—Some Coccinellid Statistics. Journ.

Keon, Ent. vol. vii. pp. 440-443.

. Ewine, H. KE. 1913.—Notes on Oregon Coccinellide. Journ.

Keon. Ent. vol. vi. pp. 404-407.

Fasre, Henri. 1897.—‘ Souvenirs Entomologiques.’ The

chapter entitled ‘“‘ Les Emigrantes.”

10a. Grrautt, A. A. 1914.—Pupal Stages of Adalia bipunctata.

11.

12.

13.

14.

15.

16.

Whe

18.

19;

20.

21.

Ent. News, vol. xxv. p. 155.

Hourrz, Martin. 1907.—Ueber Adalia bipunctata L. typ. und

deren Varietat sex-pustulata L. Ent. Woch., 24 Jahre.

pp. 181-182.

Jounson, Roswetnt. 1907.—Economic Notes on Aphids and

Coccinellids. Ent. News, vol. xviii. pp. 171-174.

Jounson, Roswentu. 1910.—Determinate Evolution in the

Colour-pattern of Lady-beetles. Carnegie Inst., No. 122

Jounson, Rosweri. 1912.—Heredity of Colour-Pattern in

Coecinnellid Beetles (abstract). Proc. 7th Int. Zool. Con-

gress, Cambridge, Mass., pp. 409-410.

Krause, A. H. 1904.—<Adalia bipunctata Varietaten aus

Nord-Thtiringen. Ent. Zeits. vol. xvi. p. 112.

Lurz, K. G. 1888.—Hibernation of the Two-spotted Lady-

bird. (Correspondence.) Insect Life, vol. i. p. 56.

Lutz, F. KE. 1908.—Notes on the Inheritance of Variations in

the Colour-pattern of Crioceris asparagi. ‘ Psyche,’ Boston,

vol. xv. pp. 50-52.

McCracken, I. 1906.—Inheritance of Dichromatism in Lina

and Gastroidea. Journ. Exp. Zool. vol. ii. pp. 331-336.

Meissner, Orro. 1912.—Statistik und Biologie von <Adalia

bipunctata. Ent. Zeits. vol. xxvi. pp. 82-85, 87-88, 94-

95.

Meissner, Orro. 1912.—Cannibalismus bei Coccinelliden.

Wien. Ent. Zeit. vol. xxvi. p. 322.

Paumer, Mirtam A. 1911.—Some Notes on Heredity in the

Coccinellid genus dAdalia (Mulsant). Ann. Ent. Soc.

America, vol. iv. pp. 283-302, 4 pls.

21a. Paumer, Miriam A. 1914.—Some Notes on the Life-

history of Lady-beetles. Ann. Ent. Soc. America, vol. vii.

pp. 213-237, 2 pls.

Proc. Zoou. Soc.—1920, No, XX XITI, 33

490

23.

25.

26.

27.

28.

ON THE LIFE-HISTORY OF THE LADY-BIRD BEETLE.

Rercuart, A. 1904.—Adalia bipunctata (L.) varietaten. Ent.

Jahrb. Leipzig, vol. xii. pp. 179-181, 1 Taf.

Scuréper, Cur. 1901 & 1902.—Die Variabilitat der Adaha

bipunctata, gleichzeitig ein Beitrag zur Descendez-Theorie.

Allg. Zeits. fiir Ent., Bd. vi. (1901), Nos. 23 & 24, pp. 355,

371; Bd. vii. (1902), pp. 5, 37, 65, mit Tafel 5 und 5

Textabbildungen.

. ScuroOpER, Cur. 1904.—Selektions und der Theorie der

Zeichungs-phylogenie bei den Lepidoptera. Allg. Zeits.

fiir Ent., Bd. ix. pp. 215, 249, 281.

Scur6pER, Cur. 1909.—Die Erscheinungen der Zeichnungs-

vererbung bei A. bépunctata und ihren ab. 6-pustulata u.

4-maculata. Zeits. wiss. Inseckten biol. Berlin, vol. iv.

pp. 182-134.

Sravrer, J. W. 1889.—Cannibalism with Lady-birds. Insect

Life, vol. i1. p. 55.

SmirH, J. B. 1892--General Notes: Hpilachna borealis.

Insect Life, vol. v. Nov., p. 98.

Smitu, J. B. 1893.—Carnivorous and Herbivoxous Insects.

Ent. News, vol. iv. p. 123, figs. 1 & 2.

Birmingham, November, 1920.

ON FILARIID WORMS FROM MAMMALS AND BIRDS. 49]

31. Filariid Worms from Mammals and Birds in the

Society’s Gardens, 1914-1915. By C L. Bounencrr,

M.A., D.Se., F.Z.8., Professor of Zoology, University

of the Panjab, Lahore.

[Received September 1, 1920: Read November 16, 1920. |

(Text-figures 1-12.)

INDEX.

Page

IsitiRoTahee OVI. sodpisnc qedhan deenee pn Rea cone won te paaAnticitie meracancrateee aC oul

UAT SUUCULONER Wa WINSte) eaaecsccteseesae cco eee AOS

Acanthocheilonema diacantha (Molin) ........................... 494

al pee GIRCCHIE (VOC) soscon cossssegooondannoaencoanany 2Ie

Dicheilonema horridum (Dies.) ...........0 ccc eee eee eeeeeece sevese 499

Diplotriena tricuspis (Fedtsch.)) ...........-............0005-.... 499

* diuce, sp. 0. : BRAGA A sachen tone eenaeenay DOL

. flabellata (v. Tans) sats ek ab aaueentha 0

J ROL PUTE CT ACH DATOS S FA Doty. Uaaa Anis se anna sebSFiwd acosaadarmnonocencicaseden e108:

In February 1914, I was asked by the Prosectorial Committee

to undertake the identification of the Nematode parasites

obtained at the Society’s Gardens, and material was forwarded

to me from the Prosectorium at regular intervals until the

beginning of 1916. During this period monthly reports, con-

taining provisional identifications, were sent to the Committee,

the material being laid aside for further investigation. My study

of the collection thus accumulated was interr upted by my depar-

ture for Mesopotamia on war service, and was only resumed three

years later at the end of 1919

The material described in the present memoir consists of the

representatives of the Family Filariide obtained from Mammals

and Birds in the Gardens. Altogether eight species were met

with, two of which are described as new. Of the known forms,

three have been previously observed on single occasions only anid

are species about which further information was desired; the

other three are worms commonly reported from Zoological

Gardens ; yet even of these our knowledge is by no means com-

plete, and I have therefore added a short account of two of them

to my descriptions of the less known species.

During recent years renewed attempts have been made | vv

various authors, especially Railliet and Henry, to break up the

old genus Paltorinn s.1., and have resulted in a much improved

knowledge of many of the commoner Filariids ; some of the older

generic names proposed by Viborg, Diesing, te have been revived

and several new genera established. In this paper, I have

attempted, wherever possible, to refer the worms described to

these new groupings,

33*

4992 PROF. CG. L. BOULENGER ON FILARIID

Genus FiuartaA Mueller, 1787.

FILARIA SUBCUTANEA v. Linstow, 1899, non Parona, 1894.

A number of specimens of this species were obtained from the

subcutaneous tissue of the Canadian Porcupine, Wrethizon

dorsatun.

This species, which is known only from the type-specimens

obtained from the same host in the Berlin Zoological Gardens,

Text-figure 1.

Seek

— eee

Ea:

VU_

‘

Filaria subcutanea v. Linst.

Lateral view of the anterior extremity of female. X 38. ~

i., intestine; 2.7., nerve-rine; c., cesophagus ; va., vagina; vw., vulva.

is of special interest, for, as pointed out by Hall (1916), there is

reason to believe that PF. subcutamea v. Linst. is a synonym of

Lf, martis Gmelin, the species generally regarded as the type-

species of the genus Milaria s. str.

My account of 7. swbcutanea will be found to differ in some

respects from that given by v. Linstow, yet I think there is little

WORMS FROM MAMMALS AND BIRDS. 493

doubt that we have both dealt with the same form. /. martis

is very imperfectly known, and until new material is described

the relationship between the two species must remain doubtful ;

there are certain characters, ¢.g., the position of the vulva, which

in our present state of knowledge it is impossible to reconcile.

Hall has pointed out that v. Linstow’s specific name is a

homonym of 7. swhewtanea Parona, 1894. I have followed his

example and refrained from renaming the species until its proper

relationship with Yilaria martis is ascertained.

My material was obtained from the same animal as the micro-

filaria recorded by Plimmer (1915) in his annual report on the

blood-parasites; Hall is therefore right in assuming that this

larval stage is to be referred to /. subcutanea v. Linst.

Text-figure 2.

Filaria subcutanea v. Linst.

Posterior extremity of male, lateral view. XX 225,

po., postanal papilla; pr., preanal papilla; sp., short spicule; spi., long spicule.

Specific diagnosis.—Filaria : Body filiform, more attenuated

posteriorly than anteriorly, both extremities rounded.

Cuticle appears smooth except under very high magnifications,

when a fine transverse striation is apparent.

Head-papille very small, four submedian papille present,

lateral papiilee not noted.

(Hsophagus simple, 7. ¢. not divided into anterior and posterior

regions, and short.

Male 48-56 mm. long, with a maximum thickness of about

0-45 mm. Césophagus 1:2—1:4 mm. in length.

The posterior end of the body is coiled in a loose spiral and

provided with cuticular ale at its extremity. Cloaca 0-07 mm.

from the posterior extremity. There seem to be nine pairs of

papille, of which four are preanal and five postanal. The papillee

494 PROF. C. L. BOULENGER ON FILARITD

of the last pair are small and flat; they no doubt correspond to

the similar pair at the posterior extremity of the female. The

eight anterior pairs of papille are elongated and decrease in size

from before backwards.

The two spicules are very unequal, measuring 0°38-0°42 mim.

and 0:13-0:14 mm. respectively. ‘The long spicule consists of an

anterior tubular region followed by a partly membranous posterior

region. The small spicule is conical and terminates 1n a point.

Female 160-175 mm. in length, with a maximum thickness of

about 0'7 mm. The esophagus measures 1°5-1'7 mm. in length.

Anus about 0°15 mm. from the posterior extremity, the latter

rounded and provided with a pair of small flat papille. At the

level of the anus the body has a thickness of 0°2 mm.

Vulva 1°9--2°4 mm. from the anterior extremity. ‘The slender

vagina has a forwardly directed loop which extends to the level

of the nerve-ring, 0-4 mm. from the oral end.

Ovoviviparous. Eggs thin-shelled, measuring 0°022 mm. x

0-015 mm.

Genus ACANTHOCHEILONEMA Cobbold, 1870.

ACANTHOCHEILONEMA DIACANTHA (Molin, 1858).

Filaria diacantha was originally described from worms obtained

in the abdominal cavity and lungs of various Rodents in Brazil.

The specimens from the Society’s Gardens ave from the Canadian

Poreupine, Vrethizon dorsatum, and were found in the peritoneal

cavities.

The original description is somewhat incomplete, and Hall, im

his monograph of the Nematode parasites of Rodents (1916), is

inclined to regard this form as a species of Seéaria ; the well pre-

served material before me makes it, however, quite clear that the

worm is to be referred to Cobbold’s genus Acanthocheilonema as

recently redefined by Railliet, Henry, and Langeron (1912).

Specific diagnosis.—Acanthocheilonema: Body slender, filiform,

diminishing in breadth at both extremities and much attenuated

posteriorly. Anterior extremity somewhat clavifovm, the head

being separated from the rest of the body by a slight, neck-like

constriction.

The cuticle appears longitudinally lined under a low power of

the microscope ; nigh magnifications, however, reveal a very fine

transverse striation.

Six head-papille are present, four submedian and two lateral.

The lateral papille are large and project in such a way that the

head appears almost square in dorsal or ventral view.

The esophagus consists of distinct anterior and _ posterior

regious ; the former is narrow and measures under 0-5 mm., the

posterior region is stouter and has a length of 2.3mm. The

nerve-ring surrounds the middle of the anterior part of the cso-

phagus.

WORMS FROM MAMMALS AND BIRDS. 495

The caudal region is long and slender, and its extremity in both

sexes is provided with a pair of conical appendages ; in front of

these on the ventral surface is a pair of small flat papillee.

Male 35-40 mm. long, its greatest breadth about 0°25 mm.

The head has a breadth of only 0°07 mm.

The posterior end of the body is coiled in a loose spiral.

oacal apertur 25-0°28 mm. from the extremity. a

Cloacal ture 0:25-0°28 f tl tremit Male

Text-figure 3.

Sp.

Acanthocheilonema diacantha (Molin).

A. Lateral view of anterior extremity of female. * 120. B. Dorsal view of

head. x 225. OC. Tail of female, lateral view. X 225.

a.@., witerior region of wsophagus; c.a., caudal appendage; ¢.p., caudal papilla ;

L.p., lateral head-papilla ; 2.7°., nerve-ving; p.c., posterior region of cesophagus

s.p., submedian head-papilla; v., vulva.

papille small, consisting of five pairs, four preanal and one post-

anal close behind the cloaca.

The unequal spicules measure 0°14 and 0-11 mm. in length

respectively ; the longer spicule is tubular in shape, with a some-

what ill-defined membranous posterior region, the smaller spicule

is in the form of a strong tube, bent in the middle.

Female 58-65 mm. in length, with a maximum breadth of

about 0°35 mm. Head 0:08 mm broad.

Auus about 0°2 mm. from the posterior extremity.

496 PROF. C. L. BOULENGER ON FILARIID

Vulva 0°65-0°9 mm. from the anterior end. The long vagina

runs almost straight backwards.

Ovoviviparous. Hggs thin-shelled, measuring 0:03 x 0:02 mm.

Text-figure 4.

Acanthocheilonema diacantha (Molin).

Posterior extremity of male, lateral view. X 225.

c.a., caudal appendage; cl., cloaca; ¢c.p., caudal papilla; po., post-anal papilla ;

pr., preanal papilla; sp., long spicule; spi., short spicule.

ACANTHOCHEILONEMA GRACILE (Rudolphi, 1819).

Filaria gracilis is a common parasite of Monkeys and has been

listed from more than twenty-five species, including both Old-

World and New-World forms. My material is derived from the

peritoneal cavities of the following species, all from the Society’s

Gardens: White-fronted Capuchin (Cebus hypolewcus), Smoky

Woolly Monkey (Lagothriz infumata), Moustache Marmoset

(Leontocebus mystas), and Squirrel Monkey (Sainiris sciwrea).

The systematic position of this worm is somewhat uncertain.

Diesing, in his famous ‘ Revision der Nematoden’ (1860), placed

it in his genus Dipetalonema, a genus which is not only ill-defined,

but contains also widely separate forms from Birds and Reptiles.

In many of its characters, e.g., position in the host, structure of

head-papillz, cesophagus, caudal appendages, spicules, ete., Milaria

gracilis closely resembles certain species of Acanthocheilonema, and

I have decided to include it, at least provisionally, in this genus.

Specific diagnosis.—Acanthocheilonema: Body filiform, very

elongated, much diminished in breadth posteriorly. Head

narrow, not separated from the body by a neck-like constriction.

Cuticle with very fine transverse striations, only visible under

high magnifications.

WORMS FROM MAMMALS AND BIRDS. 497

Text-figure 5.

A B

Acanthocheilonema gracile (Rud.).

Head: A. Dorsal view, B. Lateral view. XX 225.

L.p., lateral head-papilla ; m., mouth; s.y., submedian head-papilla:

Text-figure 6.

Acanthocheilonema gracile (ud.).

Posterior extremity of male, lateral view. X 225.

c.a., caudal appendage ; ¢.p., caudal papilla; c7., cloaca; po., postanal papilla;

pr., preanal papilla: sp., posterior part of long spicule; spi., short spicule.

498 PROF. C. L. BOULENGER ON FILARIID

Six head-papille, the lateral pair prominent giving the head

a truncated appearance when viewed from the dorsal or ventral

surface.

(Esophagus consisting of unequal anterior and_ posterior

regions.

Caudal region long and very slender, in both sexes provided

with a pair of conical appendages.

Male 96-110 mm. in length, with a maximum thickness of

about 0°27 mm. Head 0:09 mm broad.

Text-figure 7.

Acanthocheilonema gracile (Rud.).

A. Spicules of male. X 225. 1B. Caudal extremity of female. 225.

c.a., caudal appendage ; sp., long spicule; spi., short spicule.

The posterior end of the body is very slender and coiled in a

loose spiral. Cloaca 0°26-0°28 mm. from the extremity. Three

or four pairs of preanal papille and one pair of postanal papille

close to the cloaca. A pair of small conical appendages are

situated 0°02 mm. from the posterior extremity ; just anterior to

them are a pair of small flat papille.

Spicules very unequal; the long spicule is 1:23 mm. long, and

consists of an anterior tubular region, 0°33-0°36 mm. in length,

followed by a long thin terminal region; the short spicule

WORMS FROM MAMMALS AND BIRDS. 499

measures 0°27 1nm., it also has the shape of a tube, terminating

in a membranous, hooked extremity.

Female 160-250 mm. iong, with a greatest thickness of about

0°38 mm. Head 0-11 mm. broad.

The anterior region of the cesophagus measures 0°58 mm., the

posterior 2°5 min. in length.

Caudal region with a pair of elongated appendages, about

0-03 mm. from the end of the body. Anus 0°63 mm. from the

extremity. Z

Vulva 0°8-1:15 mm. from the oral end. The long,

vagina is directed posteriorly and has a Jength of 8 mm.

Ovoviviparous. Eggs small and _ thin-shelled, measuring

0:015-0:018 x 0-01 mm.

straight

Genus DicHEILONEMA Diesing, 1860.

DICHEILONEMA HORRIDUM (Diesing, 1851).

Examples of this well-known species were sent to me from the

Prosectorium on two occasions ; they were obtained from the body-

cavity of Rhea americana. It is a worm commonly reported from

Zoological Gardens.

Genus DreLtorriana Railliet et Henry, 1909.

The genus Diplotriena was established by Henry (in Henry

and Ozoux, 1909) to include a group of very closely allied species

of Filavia from the body-cavities of various birds, chiefly Passeri-

formes. They are easily recognised by the presence of a peculiar

apparatus in the form of a pair of chitinous * tridents” situated

in the front of the head, and by the structure of the male

spicules, one of which is straight, the other twisted.

Fifteen species have been referred to this genus; the majority

have, however, been so imperfectly described that their recog-

nition is very difficult. The material from the Society’s Gardens

includes three species which are to be placed in this genus; two

of these I have referred to established species, the third is

described as new. Diagnoses of the species of Diplotriena must

be regarded as merely provisional until a thorough revision of all

known forms has been made.

DreLorrieNxa rricuspis (Fedtsch., 1883).

Stossich’s monograph of Filariids (1897) shows that Milaria

tricuspis has been recorded from a large number of different

Birds; the measurements given in his specific diagnosis make it

clear, however, that a number of species have been confused

under this name.

The specimens which I have referred to D. tricuspis were

found in the body-cavities of the White Headed Starling,

500 PROF. C. L, BOULENGER ON FILARIID

Poliopsar leucocephalus ; in their measurements they agree fairly

well with the worms described by v. Linstow (1891) as belonging

to this species. My material consisted of females only. :

Specific diagnosis.—Diplotriena: Body long, filiform, tapering

close to the anterior and posterior extremities which are of

approximately the same thickness. Cuticle not transversely

ringed.

Text-figure 8.

Diplotriena tricuspis (Fedtsch.).

Lateral view of anterior extremity of female. > 120.

d.c., anterior region of cesophagus ; n7., nerve-ring ; p.c., posterior region of

cesophagus ; ¢7., trident; va., vagina; vu., vulva.

Six head-papille; the four submedian are very small and

inconspicuous. The cesophageal trident has a length of 0-15 mm. ;

its anterior stem is truncated.

(Esophagus consisting of a short anterior region and a very

long posterior region. The nerve-ring surrounds the anterior

cesophagus about 0-25 mm. from the anterior extremity.

Female 160-180 mm. long; the greatest breadth of the body is

about 6°6 mm.

The narrow anterior region of the cesophagus has a length of

WORMS FROM MAMMALS AND BIRDS. 5O1

0°35 mm., the wider posterior region 8°8 mm. The anus is sub-

terminal.

Vulva 0°55-0°6 mm. from the oral extremity. The stout,

muscular vagina runs straight backwards, and has a length of

25 mm.

Oviparous. Hges thick-shelled, 0:045-0-05 x 0:035 mm., con-

taining well-developed embryos.

Text-figure 9.

Diplotriena diuce, sp. n.

Lateral view of anterior extremity of female. x 120.

Lettering as in the preceding text-figure.

DIPLOTRIANA DIUCH, sp. n.

This worm was obtained from the body-cavity of the Diuca

Finch, Diuca grisea; the material included both male and female

specimens.

Specific diagnosis.—Diplotriena: Body comparatively short,

semi-transparent, tapering at both ends. The anterior extremity

is narrower than the posterior.

Cuticle thin, transversely ringed.

Cephalic extremity with six very inconspicuous, flattened

papille.

Kach cesophageal trident has a length of 0:13-0:14 mm.; its

anterior stem terminates in a fine point. CHsophagus 2:°5-4 mm.

long, narrow throughout, and without division into anterior and

posterior regions. The nerve-ring surrounds the cesophagus

0:2 mm. from the anterior extremity.

502 PROF. G. L. BOULENGER ON FILARITD

Male 12-20°5 mm. long, with a maximum breadth of about

0°75 mm.

Cloaca 0:06-0:07 mm. from the posterior extremity. Tail

broad and rounded. ‘There are six pairs of inconspicuous,

flattened papille ; four pairs are preanal, two postanal.

Spicules unequal; the long spicule is straight and has a length

of 0°72 mm., the short spicule is twisted and measures 0°45 mm.

Text-figure 10.

Diplotriena dince, sp. n.

Posterior extremity of male, ventral view. X 75.

7., intestine; po., postanal papilla; pr., preanal papilla; sp., long spicule ;

spi., short spicule.

Female 30-43 mm. in length; the body attains a thickness

of 0°'9 mm.

Anus about 0°71 mm. from the rounded extremity. The vulva

projects slightly from the ventral surface, 0°35 mm. from the

cephalic end. Vagina short, 0-9-1 mm. in length.

Oviparous. Eggs thick-shelled, measuring 0:045—0-05 x

0:03 mm.

DIPLOTRIZNA FLABELLATA (v. Linstow, 1888).

My material includes a single specimen of a female Diplotriana

from the body-cavity of the Red Bird of Paradise, Paradisea

rubra. 1 take it to be the same species as D). flabellata described

from Paradisea apoda by v. Linstow in the ‘ Challenger’ Reports

(1888).

Specific diagnosis.—Diplotriena : Body short, more attenuated

anteriorly than posteriorly.

WORMS FROM MAMMALS AND BIRDS. 508

Cuticle transversely ringed. Cisophagus not divided into two

regions, harrow anteriorly, increasing in width gradually behind

the nerve-ring, the latter 0°3 mm. from the oral end.

Trident 0°25 nm. long, its antevior stem truncated.

4 Text-figure 11.

Diplotriena flabellata (v. Linst.).

Lateral view of anterior extremity of female. X 120.

Lettering as in the preceding text-figures.

Female 44 mm. in length, with a maximum thickness of about

1 mm. Caudal extremity rounded, broader than the head.

Position of anus not ascertained.

Vulva prominent, 0°6 mm. from the anterior extremity; the

thick muscular vagina has a length of 2 mm.

Oviparous. Eggs thick-shelled, 0°04—0:045 mm. x 0:25 mm.

INCERT SEDIS.

FILARIA ARAMIDIS, sp. Nn.

A single specimen of this species was obtained from the leg-

muscles of the Cayenne Rail, Aramides cayennensis. In the

absence of the male it is impossible to place the form in its

504 PROF. C. L. BOULENGER ON FILARIID

proper systematic position; I have, therefore, retained it in the

genus Filaria s.1. In some respects it appears allied to certain

species known from the muscular system of Mammals, e. 9.

ilaria scapiceps Leidy (Hall, 1916).

Specific diagnosis—Filaria s.1.: Body comparatively short

and thick, tapering nearly equally at both extremities. .

Cuticle finely striated transversely.

Text-figure 12.

Filaria aramidis, sp.n.

A, Anterior extremity; B. Posterior extremity. Lateral view. » 75.

Lettering as in the preceding text-figures. ov., loop of ovary.

Head rounded, bearing four very small submedian papille ;

lateral papille not seen.

(Hsophagus narrow and simple, z.e., not divided into anterior

and posterior regions. Nerve-ring close to the anterior extremity.

Female 25 mm. in length, with a maximum breadth of about

lmm. Width of nead 0-2 mm. The esophagus has a length of

1-lmm. Posterior extremity rounded ; position of anus uncertain,

about 0:15 mm. from the end of the body,

WORMS FROM MAMMALS AND BIRDS. 505

Vulva 0°82 mm. from the anterior extremity; the slender

vagina is directed posteriorly and has a length of 2°4 mm.

Kees thin-shelled, measuring 0°03—0:035 x 0:02 mm.

BIBLIOGRAPHY.

Drestye, K. M. (1861).—“ Revision der Nematoden.” Sitz-

ungsber. d. Kais. Akad. Wiss. Wien, xlii. 1860, pp. 595-736.

Haut, M. C. (1916).—“ Nematode Parasites of Mammals of the

Orders Rodentia, Lagomorpha, and Hyracoidea.” Proc. U.S.

Nat. Mus. Washington, |. pp. 1-258.

Henry et Ozoux (1909).—“ La Filaire du Foudi.” Bull. Soc.

Path. Exot. Paris, 11. pp. 544-547,

Lrystow, O. von (1888).—‘‘ Report on the Entozoa collected by

H.M.S. ‘Challenger’ during the years 1873-6.” ‘Challenger’

Reports, xxii. pp. 1-18.

—— (1891).—‘ Veber Filaria tricuspis und die Blutfilarien der

Kraihen.” Arch. f. Naturg. Berlin, lvii. p. 292.

—— (1899).—** Nematoden aus der Berliner Zoologischen Samm-

lung.” Mitt. a.d. Zool. Samml. d. Mus. f. Naturk. Berlin,

1. (2), pp. 1-28.

Purmer, H. G. (1915).—‘ Report on the deaths which occurred

in the Zoological Gardens during 1914, together with a list

of the blood-parasites found during the year.” Proce. Zool.

Soc. London (1), pp. 123-130.

Ratiiuer, A., Henry, A., et Lanceron, M. (1912).—‘‘ Le genre

Acanthocheilonema Cobbold, et les Filaires péritonéales des

Carnivores.” Bull. Soc. Path. Exot. Paris, v. pp. 392-395.

Srosstco, M. (1897).—“ Filarie e Spiroptere. Lavoro Mono-

grafico.” Bull. Soc. adriat. di sci. nat. Trieste, xvii.

pp. 11-162

Proc. Zoou. Soc.—1920 No. XXXIV, 34

ON THE FAUNA OF THE AFRICAN LAKES, 507

32. The Fauna of the African Lakes: a Study in Compara-

tive Limnology with special reference to Tanganyika.

By Wiiitam A. Cunnineron, M.A., Ph.D., F.Z.8.

‘Received May 31, 1920: Read November 16, 1920.]

(Text-figures 1-2.)

InpDEx.

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1. HISTORICAL SURVEY.

Ever since the days of the great African explorers and their

quest of the sources of the Nile, there has existed a certain

fascination about the big lakes of Central Africa. While the

scientific world was at first concerned principally with the

geography of the heart of the continent, opportunity soon offered

of learning something of the organisms which live in the vast

inland seas, and it then became evident that matters of no little

biological meeeest were involved.

It is the object of this paper to deal in comparative fashion

with the animals of the principal lakes, and since Tanganyika*

holds a unique position as regards its fauna, a somewhat detailed

account of its zoological exploration seems fitting, as it will afford

both an introduction to a study of the lakes and an enumeration

of certain outstanding facts.

Little more than sixty years have passed away since Burton

and Speke, the first Huropeans to penetrate to Lake Tanganyika,

reached the shores of this big inland sea, Their expedition, after

encountering numberless hardships, arrived at the lake between

the Malagarasi River and Ujiji on the thirteenth of February,

1858. Although little was done in the way of exploring Tan-

ganyika, Burton described, with a surprising degree of accuracy,

various facts concerning it which he learned from the Arabs and

natives, while Speke, on his return, brought certain shells from

the lake shore, which came into the possession of the Br itish

Museum.

* Some years ago I called attention to the orthography of the name ‘T'anganyika,’

sdivoastitig the adoption of the spelling ‘Tanganika’ by English writers (69).

The arguments advanced in favour of the change have not lost their force, and still

seem worthy of consideration. As there is no indication that writers in this country

ave prepared to accept the modification, I adhere reluctantly to the old form.

34*

508 DR. W. A. CUNNINGTON ON THE

It was in the examination of these shells that the unusual

interest which still attaches to certain forms of life found in

Tanganyika had its origin. 8. P. Woodward, who reported on

the collection, was str mee by the fact that some ‘of the shells were

curiously marine in appearance. Of one of them, which he

named Lithoglyphus zonatus (now known as Spekia zonata), he

said (205, p. 348) that it so much resembled a Verita or Calyptrea

that it might have been taken for a sea-snell. Of another, he

remarks that ‘Its colour, solidity, and tuberculated ribs give

it much the appearance of a small marine whelk (Vassa).” At

the time, these opinions attracted little attention, although the

matter was destined to come into prominence at a later date.

Some twenty years later-—in 1878—there was established the

first settlement of Huropeans on the shores of Tanganyika, when

agents of the London Missionary Society settled at Ujiji in

August of that year. With praiseworthy public spirit, the

Directors of the Society had given instructions that the gathering

of scientific information should be considered among the duties

of this pioneer expedition, and thus it is that E. C. Hore (101),

a member of the party, was the first to attempt a systematic

collection of the forms of life occurring in the lake. While the

collections since made by trained naturalists have, of course,

added very largely to the knowledge of the fauna and flora, the

forms obtained by Hore embraced many different groups, and

afforded the first comprehensive glimpse of the organisms which

the lake contains.

This English Mission was very shortly followed by the estab-

lishment of a French Catholic Mission, and it should be noted

that these White Fathers of Algeria have also been instrumental

in gathering information, and in particular have made large

collections of shells which have been described in later years by

French conchologists.

At the end of 1879, Tanganyika was visited by Joseph

Thomson. Collections of ‘shells made by Hore and Thomson were

reported on by Hdgar Smith in 1880 and 1881 (159, 160, 161),

and he also had occasion to remark on the marine appearance

of some of them. This matter being brought to the front once

more, Thomson put forward the view (190, vol. ii. p. 85) that the

water of the lake had probably been saline until recent geological

times, when it became freshened through circulation and outflow

to the Congo (compare also 191, where he goes further, and

suggests hye origin of the lake from a former sea occupying the

Congo basin).

The discovery of meduse by Bohm (114), a German explorer

who crossed Tanganyika in 1883, only emphasised matters and

laid stress on the supposed marine character of the lake fauna,

since, with few exceptions, such organisms are confined to the

ocean. Von Wissman (204, p. 253) confirmed this discovery

when sailing on the lake in 1887, and a few years later, F. L. M.

Moir, the manager of a British trading company, undertook to

FAUNA OF THE AFRICAN LAKES. 509

obtain specimens of these medusze properly prepared for scientific

investigation. This valuable material, on its arrival in England

at the Sond of 1892, was placed for examination in the hands of

R. T. Giinther of Oxford.

It was Giinther who first put forward a detailed theory to

account for the presence in a central African lake of animals

with apparently marine affinities. In his report on the Tan-

ganyika medusa he suggested (95, p. 289) that if at some early

period the interior of Africa stood at a much lower level, with

the Atlantic Ocean extending over the Congo basin and the lake

existing as an arm of the sea, it would then be possible readily

to explain the presence in 1t both of the medusa and the remark-

able molluscan forms. As the lake became separated from the

sea, so the salinity of the water would tend to decrease if it were

supplied with inflowing rivers and found an outlet through which

to carry away the salt.

Since the problem presented by Tanganyika had now become

so interesting and definite, it seemed very desirable that the lake

fauna should be investigated upon the spot by some trained

zoologist. The matter was taken up by Ray Lankester, who,

with the help of others interested in the question, obtained the

necessary grants from the Royal Society and organised the First

Tanganyika Expedition. This was put in charge of J. E. S.

Moore of the Royal College of Science, who from that time

onwards contributed largely to the knowledge of the lakes of

Central Africa.

The expedition left England in the autumn of 1895, and Lake

Shirwa was visited as well as Nyasa, while Moore spent several

months on the shores of Tanganyika. As a result of this journey,

the original problem seemed far from being solved, for the expedi-

tion brought matters to light which only made the lake appear

more remarkable. Many strange and unique animal forms were

collected, certain of which were thought to show marine aflinities,

while the majority were new to science. The collection included

a number of types not unexpected in a tropical lake; only a

proportion of the fauna exhibiting the unusual features to which

reference has been made. Moreover, although the expedition had

visited Nyasa and Shirwa, in neither of these lakes had marine-

like forms been found resembling those of Tanganyika.

Through a study of the newly acquired material, especially the

anatomy of certain of the molluscs, Moore arrived at the conclu-

sion that the forms exhibiting a marine appearance could not be

compared directly with any marine types of the present day, but

rather possessed archaic and generalised characters which sug-

gested that they were the relics of a marine fauna of a previous

age. Further, he considered that some of the Tanganyika shells

offered a strong resemblance to certain marine Jurassic fossils, in

particular those of the Inferior Oolite of the Anglo-Norman

basin. Taking this as evidence of the period from which these

remarkable living forms had been derived, Moore propounded

510 DR. W. A. CUNNINGION ON THE

once more a theory of the former connection of Lake Tanganyika

with the sea, asserting that this connection existed in Jurassic

times (131).

The discoveries of the First Tanganyika Expedition, while

stimulating interest in the faunas of the African lakes, only

emphasised the need for further information. It was still to

be determined whether any of the other big lakes contained

organisms with a marine appearance, and it was very desirable

that more should be learned of the geology of the Tanganyika

basin, as having a direct bearing on the question of a former con-

nection with the sea. Ray Lankester, therefore, took the initiative

once more, formed a committee of scientific men interested in the

subject, and organised a second expedition. In view of the

geological and geographical work to be undertaken, a liberal

grant was made by the Royal Geographical Society, and Moore

was accompanied by Malcolm Fergusson as surveyor and geologist.

The Second Tanganyika Expedition (186) left England in 1899,

and after proceeding up lakes Nyasa and Tanganyika, visited

Kivu, Edward Nyanza*, and Albert Nyanza before returning to

the coast by way of the Victoria Nyanza.

An examination of the collections obtained in the latter lakes

only laid additional stress on the unique nature of Tanganyika,

for in none were found animals suggestive of the sea. Indeed,

the already long list of forms found only in Tanganyika was aug-

mented by the expedition, more particularly as regards the fish.

Certain of the little-known regions between Tanganyika and the

Albert Nyanza were mapped, and considerable attention was paid

to the study of the geology of this district as well as that of the

Tanganyika basin. Not long after the return of the expedition,

Moore published a volume entitled “The Tanganyika Problem”

(137), in which he dealt at some length with the question which

had so occupied his attention. A large amount of new informa-

tion was given in this work. It treated in a comprehensive

manner the physiography and geology of the part of Africa con-

cerned, and besides giving a complete review of the fauna of

Tanganyika as then known, discussed the origin of fresh-water

faunas in relation to the matter.

While a great deal of information bearing on the geology and

geography of the lake regions had been obtained, and extensive

collections had afforded a knowledge of the fauna of Tanganyika,

the flora of the lake had not been dealt with as it deserved. To

those who had been impressed with the marine characteristics of

certain animals living in the lake, it appeared quite conceivable

that when the flora came to be investigated, plants equally

remarkable and equally suggestive of marine aftinities would be

discovered. If, for example, specimens of the red Alge so charac-

* The lake, formerly known as the Albert Edward Nyauza, is always referred to in

this paper as Edward Nyanza or Lake Mdward. he change of name was approved

by the late King Hdward, and should be universally adopted in order to ayoid

coufusion with the Albert Nyanza or Lake Albert (ef. 55, p. 129).

FAUNA OF THE AFRICAN LAKES. yall

teristic of the ocean were to be found, this would afford very

valuable evidence concerning the ancient history of the lake.

It was with this end in view that a Third Tanganyika Expe-

dition (68) was organised by the Committee, with the conduct of

which I had the honour of being entrusted. One of the principal

objects of this expedition was therefore to make a careful collec-

tion of the water plants of Tanganyika, at the same time

collecting in Nyasa, with the idea of affording a comparison with

a more normal African fresh-water lake. My instructions also

provided ‘“ That other groups of organisms likely to throw light

on the Tanganyika Problem, and especially fishes, shall not be

neglected.” Including observations cn various points of interest,

the work before the expedition was thus sufficiently extensive,

although matters of geology and geography were outside the field

of enquiry.

I left England in the spring of 1904, proceeding via the

Zambezi and Shiré rivers to Lake Nyasa, where a few weeks were

spent in collecting. My stay on and around Tanganyika lasted

about eight months, which were fully occupied in making collec-

tions and observations as far as facilities offered. Returning to

the coast by way of the Victoria Nyanza, an opportunity was

afforded of obtaining representatives of the flora of that lake for

purposes of comparison with the plants collected in Nyasa and

Tanganyika.

It may at once be stated that the flora of Tanganyika fails to

exhibit quite the remarkable features which some had expected

it to show. As regards the higher aquatic plants, a comparison

of the species from Tanganyika with those from Nyasa and

Victoria Nyanza does not throw any light on the hypothesis of a

marine origin for Tanganyika. The fresh-water Alge of the

lake, however, have proved to be a strange and interesting

assembly, a considerable number being peculiar to Tanganyika

while several show undoubted marine aftinities. The lower forms

of vegetable life which occur thus tend to confirm the unique

biological nature of the lake.

On the zoological side, my collections have added not a little

to the number of forms known from the lakes, the results

appearing as a series of memoirs principally in the Proceedings

of the Zoological Society. For the fist time systematic tow-

nettings were made in Tanganyika on an extensive scale, and as

a result, detailed information is to hand about whole groups of

organisms, only the bare existence of which in the lake was

known before. Thus reports have now been published on the

smaller Crustacea (Copepoda and Ostracoda) as well as on the

Rotifera, while in addition the groups Branchiura and Hydrach-

nida are new records from the lake.

In completing this brief review of the zoological exploration of

Tanganyika, it is only necessary to refer to the Belgian expe-

dition to that lake and to Lake Mwero undertaken by the late

Louis Stappers. This expedition visited Tanganyika in 1912-

IL DR. W. A. CUNNINGTON ON THE

1913, and brought back valuable information concerning the

depth of the lake and the salinity of its water, as well as repre-

sentatives of its fauna. A large collection of fishes was found to

contain a number of new species. brief diagnoses of which have

been published (32, 33, 37), but beyond this, so far as I am aware,

no further particulars have been forthcoming, owing to the out-

break of the HKuropean war and the subsequent lamented death ~

of Stappers.

2. GENERAL INTRODUCTION.

This paper aims, in particular, at giving an account of the

animal forms now known to occur in Lake Tanganyika, but while

special attention has been directed to enumerating these, types

from other lakes have been noticed and discussed. The account

is therefore a comparative survey, and shows clearly the re-

markable nature of the fauna of Tanganyika to which reference

has frequently been made.

It must, of course, be understood that the systematic review

and the deductions drawn therefrom, are of necessity based upon

present knowledge, and may need modification in the light of

future discoveries.

In order not to exceed reasonable limits, the scope of the

paper has been restricted, and it does not deal with ail the big

lakes of Africa. At the same time, the largest and most impor-

tant lakes have been considered, and only those of lesser mterest

are outside the field of enquiry. Whenever a tabular form has

been adopted, columns for the lakes are uniformly arranged in

the following order:—Tanganyika, Victoria Nyanza, Nyasa,

Albert Nyanza, Edward Nyanza, and Kivu.

It is not possible to consider here at any length the geography

and physical features of the lakes concerned, but certain out-

standing particulars may be mentioned.

With the exception of Victoria Nyanza, all the lakes under

review occupy portions of the Great Rift Valley, as it has been

called, and are, therefore, of a very characteristic type. Without

entering into details, it may be briefly explained that Suess, in

discussing the geology of Central Africa, put forward the view

that the long deep valleys which run through the country in an

approximately north and south direction have been caused by

trough-faulting on an extended seale. Hach ‘‘ Graben ”—as he

terms these long narrow valleys, with their steep, almost preci-.

pitous sides—he believes to have been produced by the faulting

and sinking of a long narrow tract of country, and it is but

natural that a number of these characteristic depressions are

filled with water. Thus Tanganyika and the other typical

“Graben” lakes are situated in long trough-like valleys, bounded

for the most part by steeply rising sides, which may reach in

places a height of two or three thousand feet above the level of

the water.

FAUNA OF THE AFRICAN LAKES. 513

From the very nature of the troughs in which the lakes are

situated, the latter are likely to be deep, with shores quickly

shelving, and this is precisely what investigation reveals. Both

Nyasa and Tanganyika have now been pretty adequately sounded,

and the figures obtained are very remarkable for fresh-water

basins. Nyasa shows a maximum depth of about 786 metres

(430 fathoms), while ‘Tanganyika reaches no less than 1435

metres (about 784 fathoms)*. Tanganyika is thus the second

deepest lake in the world, Lake Baikal coming first, while Nyasa

takes fourth place, following the Caspian Sea. Where sufficient

soundings have been taken, the contours constructed show in a

number of instances the remarkably steep shores which were only

to be expected in such cases.

Victoria Nyanza, as already indicated, is a lake of a ver

different nature from those of the Great Rift Valley. Witha

length of some 200 miles and a breadth of 130 miles, it has

roughly the form of a parallelogram. While it has the largest

area of any of the African lakes, it occupies only a shallow depres-

sion, and instead of being bounded by steeply rising shores two

or three thousand feet in height, the surrounding hills reach a

maximum of 300 feet or so. The greatest depth is a mere

73 metres (40 fathoms).

In most other respects, the lakes included in this survey do not

differ greatly so far as physical conditions are concerned. ‘They

are all large sheets of water, where consequently almost oceanic

conditions prevail, while the three biggest lakes are subject to

storms of a violent description. In dealing with the evidence

afforded by the plants and animals inhabiting them, this fact

must not be overlooked, as physical surroundings may have

induced some of the features which are noted.

Although extending considerable distances both north and

south of the equator, these lakes all exist under practically the

same climatic conditions. They are tropical lakes, and differences

in fauna and flora are clearly not to be correlated with the very

slight climatic variations which may possibly occur.

The temperature of the water is naturally high. I took

sufficient temperature readings during my visit to Tanganyika to

afford some idea of the general conditions, although an exhaustive

survey was impossible. Only a few figures need be mentioned

here. The lowest reading I ever obtained in Tanganyika was

73°3° Fahy, and the highest 81-0°. Surface readings at different

periods of the year varied only between 74°7° and 81:0°, and

an average of 51 observations gave a mean of 78:0°. At con-

siderable depths the water temperature remains relatively

constant, for readings taken at a depth of about 139 metres

(76 fathoms)—the length of the sounding line—on various

* For further details concerning Nyasa, consult Moore (184, p. 7) and Rhoades

(148)-—concerning Tanganyika, see Stappers (174 & 175) and Marquardsen

(111).

514 DR. W. A. CUNNINGTON ON THE

occasions and at different places only vary between 74°8° and

741°. Stappers gives the temperature of the surface water as

oscillating between 25° and 28° C. ie 82°4° F.) (176). Mar-

quardsen, however, quotes exactly 25-0° C. (77° F.) as a mean

of 70 surface readings made by Jacobs. The latter was also

able to carry out deep water observations, to which Marquardsen

refers as follows :—‘* Unter 400 m. bis in die gréssten Tiefen

liegt dann eine fast homotherme Wasserschicht von 25°15° bis

23°12°” (73°67°-73°61° F.) (111, p. 97). A less extended series of

water temperature observations was made in Nyasa * and also in

Victoria Nyanza during the Third Tanganyika Expedition, but

the figures are sufficient to show that there are no inportant

differences from the results obtained in Tanganyika. It is

obvious that the amount of water lost by evaporation must be

very great in _ these cases.

As far as I am aware, there is relatively lttle known con-

cerning the Rate of the water in these Afiican lakes. They

are considered in general terms to be “fresh” +, but, except in

certain instances, no analyses have been made, and the nature

and quantity of the salts dissolved in the water are unknown.

The information which is to hand regarding ‘Tanganyika and

Kivu makes it clear that they differ from normal fresh waters,

and this difference appears to be reflected (as will be shown in

due course) in the nature of the organisms they contain. Tan-

ganyika water, while containing a relatively small amount of

calcium salts, is unusually vich in salts of magnesium (176).

Lake Kivu is an extreme case, where the salinity is very high.

In this instance, salts of calcium appear to have been aimost

entirely replaced by considerable amounts of sodium and

magnesium salts, the latter being quite excessive in quantity

(103).

It is possible that the water of Tanganyika has somewhat

changed in nature within recent years, for most of the early

explorers agree that the water was brackish, or at least ‘‘ pecu-

liar” in taste. If the rather vague statements are to be relied

upon, this alteration in salinity is probably to be associated with

the question of the outflow of the lake. A period during which

the lake possessed no outlet would result in an accumulation of

mee producing brackish water, while a subsequent period with

regular outflow would « Jiminish the quantity of saline matter

in solution. An examination of the available records suggests

that something of this kind has taken place, and it thus becomes

necessary to consider the whole subject of the outflow of Tan-

ganyika.

This is not the place to discuss in detail all the evidence

* Certain observations concerning Nyasa were made by Fiilleborn (83, p. 334

et seq.).

+ Grogan, referring to Lake Edward, says :—“ Although the water was salt, it

was drinkable”? (91, p. 191). In the same connection ‘Bright remarks :—* The

water of Albert is drinkable, but not good” (55, p, 188).

FAUNA OF THE AFRIGAN LAKES, Filta

which bears on the matter, but the conclusions arrived at may

be summarised as follows :—Since the area drained is not very

ereat, and the rainfall is relatively low, the amount of water

which finds its way into the lake is nearly balanced by the evapor-

ation (cf. 190, vol. il. p. 68 et seq.). It follows that under

average conditions there will be little change in the surface-

level and no occasion for an outlet for the surplus water. There

are reasons for believing that at some former period the water's

of Tanganyika did not flow out at all, and were consequently

more Sine than at present. As a result of the formation of

ereat voleanic cones north of Lake Kivu, that portion of the

Nile drainage system which once drained north into Edward

Nyanza became cut off, and its waters finally diverted south-

wards into Tanganyika (cf. 137, p. 89 e¢ seq.).

Slowly raised in level by this additional supply, the water

eventually found an outlet to the west, along the course of the

Lukuga, a former affluent, and thr ough a sandstone ridge

denuded in part by the action of a tributary of the Congo. The

lake having risen abnormally before the establishment of an

outlet, sank at first rapidly, remaining at a lower level, subject

to fluctuations with the time of year and the amount of rainfall,

A few consecutive seasons of unusual drought would suttice to

reduce the level of the lake to a point at which all outflow would

cease. It seems likely that under such circumstances a. barrier

might be formed in the bed of the effluent, due to the combined

effects of wind-blown sand and tropical swamp vegetation, and

varying in nature with the length of time taken to prodtce it.

If a tolerably firm dam was formed, a considerable rise in the

level of the lake would be necessary in order to develop enough

power to sweep away the obstruction. There is reason to believe

that the high water level which Cameron and Stanley observed,

and the subsequent re-establishment of an outflow, were phases in

a cycle of this nature. If a periodicity of this kind really exists,

it must of necessity be extremely variable in nature, and due t0

irregularities in the rainfall.

The important bearing of these inferences on the problem of

the fauna of Tanganyika it is hardly necessary to emphasise.

If the Nile has lost a not inconsiderable drainage area, which

has been added to that of Tanganyika, it seems very reasonable

to suppose that this accession of water produced, for the first

time, the overflow of the lake. If this supposition be correct,

Tanganyika must have been completely isolated and without an

outlet, until the formation of the great voleanic dam in geolo-

gically recent times.

The lists of animals from the various lakes given in the body

of this work are, as regards extent, partly dependent on the

amount of biological exploration which each lake has received.

That is to say, a ‘comparatively small fauna may be merely due

to less complete examination. Whether the Jakes considered

here can be regarded as having beeu at all equally investigated,

516 DR. W. A. CUNNINGION ON THE

it 1s not easy to pronounce, but I incline to the view that at the

present time the disproportion-—-if any—is not great*. It is

perhaps the smaller lakes which have been less fully explored,

and from them, therefore, new records might be expected,

although their smaller size does not offer the same occasion for

other organisms to come to light. The larger lakes, more

thoroughly, but still incompletely explored, may well from their

very vastness contain a number of forms which have never yet

been observed or collected. On the whole then, while recog-

nising that future investigations will probably add considerably

to the records from the individual lakes, there is reason to believe

that a comparative survey of the forms at present known will

afford a relatively true conception of the actual state of affairs.

It is not so easy as might at first sight appear, to give complete

and satisfactory lists of the animals known to inhabit the lakes

in question. I do not refer only to the difficulties of synonymy,

though these are by no means inconsiderable, particularly where

the Mollusca are concerned. ‘There is also the difficulty of

deciding which forms should be regarded as true aquatics, since

there are of necessity a number of types which are on the border

line. While in some groups the question does not arise, in

others 16 is clearly impossible to draw a hard and fast distinction

between aquatic and non-aquatic species. Thus in making a

decision in individual cases, as was necessary for the purposes of

this paper, I may have excluded forms whicn some think should

have been included and vice versd. For example, I have deliber-

ately excluded all reference to the Aves, although the lakes are

well stocked with water-fow], and similarly I omit the group

Insecta, notwithstanding the presence of water beetles and

insect larvee. Certain Oniscide were in the collections which I

brought from Nyasa and Tanganyika, the specimens having been

obtained from the lake shore. These truly terrestrial Isopods

are likewise omitted from the systematic portion of the paper, as

are also certain of the Oligochete worms. All these types, while

moisture-loving, are obviously not to be associated with individual

lakes, and their precise distribution cannot have the significance

which attaches to the distribution of strictly aquatic species.

There is finally the problem of deciding whether forms recorded

from the neighbourhood, but not actually from the waters of a

particular lake, are to be reckoned as belonging to that lake’s

fauna or not. This becomes, in a sense, a matter of special

importance when a species is described as endemicy?, Shoulda

species, for instance, be regarded as peculiar to a certain lake

* As concerns Nyasa, the collections of Fiulleborn do not come only from the lake

itself, but also from adjacent ponds, swamps, and rivers. Regarded in this light,

Nyasa has been more thoroughly investigated than the otber lakes, where records

from the surrounding neighbourhood are almost unknown.

+ Jt is necessary to explain that I use this word deliberately im the sense of

peculiar to a locality, z.e. not found elsewhere. While the word is often used with

this significance in works on geographical distribution, its meaning is a little

ambiguous, and it is desirable to make the matter quite clear.

FAUNA OF THE AFRICAN LAKES. SIG

when it was actually obtained from a neighbouring swamp ?

I have dealt with the difficulty in what seems the most common-

sense way, by definitely including all forms recorded from the

drainage area of a lake as belonging to it. Nevertheless it must

be pointed out that even this decision does not avoid the trouble

altogether, for it is not an easy matter to determine what con-

stitutes the drainage area of a lake in a country where the

boundaries of river basins are quite indefinite in the rainy

season. Again, the possibility is not excluded, that organisms

from adjoining fresh-waters may not find physical conditions

suitable for them in the big lake in their vicinity, in which case

they should not be included in its fauna. Evidence of the

existence of this very phenomenon is to be found as regards

certain plankton organisms which are absent from both 'Tangan-

yika and Kivu. Throughout this paper I have placed a § against

the names of all species which have not definitely been recorded

from the waters of the lake in question.

Subject to these difficulties in compiling exact lists, the

following approximate figures can be given :—Tanganyika contains

402 species of which no less than 293 are endemic, Nyasa with

a total of 361 and 86 endemics coming second. Victoria Nyanza

has a somewhat smaller total, viz. 289, but a larger number of

endemic forms (110). The three smaller lakes show a great

falling off in numbers, Albert Nyanza mustering only 67 species

of which 9 are endemic and Edward .Nyanza 54 with 11 endemie

species. Kivu, finally, only contains 23 types in all, 4 of which

are peculiar to the lake. Thus, as regards its fauna, Tanganyika

is distinguished from the other lakes not only by containing a

larger number of forms, but especially by displaying a remar ail

proportion of endemic species.

While it is not necessary to discuss here the origin of fresh-

water organisms, a few observations on one aspect of the question

are needed at this stage. By movements of the earth’s crust,

portions of the sea have, from time to time, been cut off with

their inhabitants, and by gradual freshening have been sub-

sequently converted into fresh-water lakes. During the latter

process, there is little doubt that certain of the organisms would

be unable to withstand the change and would perish. Inland

lakes which appear to be the modified remainders of isolated

portions of the ocean are sometimes spoken of as ‘relict lakes”

(Reliktenseen). It is, of course, precisely this view of the origin

of Tanganyika which was put forward by Giinther and afterwards

elaborated by Moore. On this hypothesis, the peculiar fauna

was originally that of a former sea, then that of a salt lake, and

finally that of a fresh-water lake.

The marine-like appearance of certain of the Gasteropod shells

of Tanganyika was expressed by the conchologist Bourguignat

by employing the term ‘“thalassoid” with reference to them—a

term which did not commit its author to any view as to origin

(42, p. 9). In his paper ‘‘On the Zoological Evidence for the

518 DR. W. A. CUNNINGTON ON THE

connection of Lake Tanganyika with the Sea,” Moore indicated

his belief in the origin of these forms as relicts from a former

ocean, by fae the expression “ halolimnic” (126, p. 453 :

eee also 188*). Of course from Moore’s point of view, a

number of animals from other groups—all those, in fact, which he

believed to be the remains of an old marine fauna—are similarly

to be included under the term halolimnic. It is necessary for

an understanding of what follows, that the significance of these

words should be explained at this point.

It remains to give a systematic account of what is known of

the animal groups represented in the lakes, after which an

attempt will be made to draw conclusions from the facts which

this detailed study reveals.

I wish to take this opportunity of acknowledging my indebted-

ness to the experts who have so willingly examined and reported

on the collections I made in Africa. In a number of cases, too,

they have given me valuable advice during the compiling of those

sections of this paper on which they are authorities, and for this

also [ express my thanks.

3. SYSTEMATIC ACCOUNT AND LISTS OF SPECIES.

In dealing in systematic fashion with the groups of animals

which are reviewed in detail, the latter are arranged approxi-

mately in order, beginning with higher and ending with lower

types. No attempt is made to conform to any particular scheme

of classification, the headings employed being merely such as are

suflicient to indicate the “contents of the different sections.

Where certain animal groups are conspicuously wanting, it should

be understood that they are either intentionally omitted, as not

truly aquatic forms (cf. p. 516), or are unrepresented in the lakes.

In each case a list is given (in tabular form where more than a

few types are concerned) of the species hitherto recorded from the

different lakes, and this is followed by a brief discussion of the

points of interest and of the significance of the records. The

complete discussion of the zoological evidence and the conclusions

to which it points is deferred “until the general survey which

follows the systematic account.

MAMMALIA,

The biggest, though by no means the most interesting in-

habitant of the lakes of Central Africa, is the hippopotamus,

which is widely distributed in the tropical parts of the continent.

The common species, ippopotamus amphibius, is probably to be

* YT do not agree with Moore’s subsequent statement that types which are

thalassoid cannot be halolimnic. It is true that the words are not synonymous,

bnt surely organisms may be marine-like in appearance (thalassoid) and at the same

time relict types (halolimnic),

FAUNA OF THE AFRICAN LAKES, 519

found in all the lakes under review *, but definite records are not

in every case available.

Tn addition to the hippopotamus, there are two species of otter

which occur over the greater part of Africa, and there is little

doubt that both of these frequent the waters of the big lakes.

A specimen of the larger form—now known as Aonya capensis—

from Lake Victoria is in the collection of the British Museum.

The smaller otter is known as Lutra maculicollis. Hore refers to

the occurrence of otters in Tanganyika (101, p. 150), and both

Grogan and Bockelmann speak of them in Kivu (91, p. 123:

16, p. 371), but actual records are wanting in most cases. It

may nevertheless be safely assumed that both species form a part

of the fauna of the lakes with which this paper is concerned.

As far as I am aware, these are the only Mammals 7 which are

recorded from the lakes.

CrROcODILIA.

The largest reptile and the one most in evidence in the African

lakes, is, of course, the crocodile, which is extremely widely

distributed in tropical Africa. Two species = have been recorded

as occurring in Lake Tanganyika.

1. Crocodilus niloticus.

This is the common African crocodile with the most extended

distribution, ranging from Palestine to Natal and known also

from Madagascar. It is found in Nyasa and Victoria Nyanza as

well as in ‘Tanganyika (see British Museum Catalogue (19) and

lists given by Tornier (192, 193, and 194)), but, as far as I can

ascertain, it has not been definitely recorded from the other big

lakes, though it very probably occurs in them all |).

2. Crocodilus cataphractus.

Recorded from Ujiji, eastern shore of Tanganyika, by Tornier

(194, p. 579). Essentially a West and Central African form,

extending from the Senegal to the Gaboon and nearly all over

the Congo.

OPHIDIA.

Four snakes adapted to an essentially aquatic life have been

found on the shores of Lake Tanganyika. In addition to these,

* Grogan states that the hippopotamus is not found in Kivu (91, p. 122), and

sockelmann and Hundeshagen refer to this also (16, p. 371: 108, p. 204).

7 Carl Peters, in his book “New Light on Dark Africa,” English translation,

London 1891, speaks on page 445 of having seen porpoises in the Victoria Nyanza.

There is to my mind little doubt that he was misled, either by the appearance of an

otter or even perhaps by one of the great fishes of the lake.

~ For specitic distinctions and synonymy consult Boulenger (19, p. 278 et seq.).

|| A statement is made by Schubotz (156, p. xix) that there are no crocodiles in

Take Edward, but this does not agree with the accounts given by other travellers.

Grogan (91, p. 122), Bockelmann (16, p. 371), aud IHundeshagen (103, p. 204).

however, speak of the absence of crocodiles from Nivu, ; ;

520 DR. W. A. CUNNINGTON ON THE

a snake allied to the cobras was taken swimming in the lake

during the Third Tanganyika Expedition. The five species *

may be enumerated as follows :—

1. Tropidonotus olivaceus.

This is a well-known and common water-snake which is

distributed over nearly the whole of Tropical Africa. It has also

been recorded from Victoria Nyanza (180, p. 265) and Edward

Nyanza (192, p. 67),and may very well occur in most of the other

big lakes.

2. Glypholycus bicolor.

Brought from Tanganyika by Hore, one of the earliest

collectors, and also recently obtained from the same locality. In

contrast to the preceding, this form has been found nowhere else.

The genus was established by Giinther (93, p. 629) to receive this

species, but a second species has since been found in Nyasaland‘.

3. Grayia ornata.

A Tropical African snake, distributed in the west from the

Cameroons to Angola and extending eastwards to the Uellé and

Ituri. Recently recorded from Albertville on the western shore

of Tanganyika.

4A. Grayia thollons.

Known from the Congo region, the Upper Nile, and the

western shore of the lake.

5. Boulengerina stormsi.

This snake, which belongs to the same group as the cobras, is

probably highly poisonous, but not strictly aquatic in character.

‘The species was described from a single specimen (a young one,

24 em. long) discovered by Storms on the western shore of

Tanganyika, A second specimen, which was obtained by my

expedition, was taken actually swimming in the lake. It is in

all probability adult and measures 125 cm. This form has not

been obtained elsewhere.

From an examination of this list it will be seen that more is

known of the snakes of ‘Tanganyika than of those of the other big

African lakes, from which indeed there are but two definite

records—to wit, those of Tvropidonotus olivacews in Victoria

* For particulars relatmg to these snakes, as also synonymy and means of

identification, consult Boulenger (84: 35).

+ The Nyasaland species of Glypholycus—G. whytii—is also a truly aquatic

snake, and may quite well occur m Nyasa, although it has not hitherto been reported

from that lake. :

FAUNA OF THE AFRICAN LAKES, 521

Nyanza and Edward Nyanza. Although Tanganyika has certainly

received a considerable amount of attention, this cireumstance

would not alone suffice to explain the relative richness of its

Ophidian fauna, which is the more remarkable in that two of the

species are endemic forms. It is hardly among the snakes that

types characteristic of a lake would be expected, but they never-

theless afford an interesting parallel to the state of affairs which

is so constant and striking in most of the groups of typically

aquatic animals.

CHELONIA.

Five species of aquatic tortoises come within the scope of this

survey *, of which four have been found in Tanganyika.

1. Sternotherus sinuatus.

A form well known from Tanganyika and obtained by Stuhl-

mann in the Victoria Nyanza (192, p. 4). It has more recently

been reported from Nyasa (193, p. 583) and from Lake Albert

(180, p. 200), while it is widely distributed in South Africa and

in Hast Africa south of the equator.

2. Sternotherus derbianus.

This species appears to be more especially West African, but

also occurs in Tanganyika.

3. Sternotherus nigricans.

A specimen of this tortoise from Usumbura, Lake Tanganyika,

is referred to by Sternfeld in his report on the reptiles of the

German Central Africa Expedition (180, p. 201). It is also

known from the region of Mozambique and from Madagascar.

4, Pelomedusa galeata.

This is likewise a widely distributed form, and is recorded by

Yornier (193, p. 583) both from Tanganyika and Victoria Nyanza.

It has again been reported from Victoria Nyanza by Sternfeld

(180, p. 201), and the British Museum also possesses a specimen

from that locality. The species does not seer to have been

mentioned hitherto from any of the other big lakes, although it

is well known from Southern and Tropical Africa and occurs

even as far north as the Sinaitic Peninsula.

5. Cycloderma frenatum.

y :

This well-known Zambezi type has been collected by Fiilleborn

* Particulars of these Chelonians will be found in the British Museum Catalogue

(19).

Proc. Zoou. Soc.—1920, No. XXXV. 315)

Fu) DR. W. A. CUNNINGTON ON THE

(193, p. 583 & 194, p. 580) in Lake Nyasa also. It is not known

from any other African lakes.

In this case again a larger number of forms is on record from

Tanganyika than from the other Central African lakes, but the

species are all widely-distributed, and their precise distribution

in the lakes is probably a matter of little significance. <A

curious piece of negative evidence of considerably more interest

is the fact that, with one exception, there are no records of

Trionychide from any of the lakes, although these forms are so

eminently aquatic and are known from other localities in Central

Africa. The exception to which I refer is Tornier’s allusion

.(193 & 194) to Cycloderma frenatuwm, a Zambezi species, in

Lake Nyasa.

BATRACHIA.

While there are many Batrachiaus which are widely distributed

in the African contiment, the number of truly aquatic forms—with

which alone this paper is concerned—is very small. So far as I

am aware, the only true aquatics found in the lakes of Africa

belong to the Anuran genus Xenopus, of which two species* need

to be enumerated,

1. Xenopus muelleri.

This form appears to be rather the commoner of the two. The

British Museum collection contains specimens from Lakes Tan-

ganyika and Nyasa, and according to Tornier (192, p. 162) this

species was collected by Stuhlmann at Bukoba on Victoria Nyanza.

Its occurrence does not seem to have been recorded from any of

the other big lakes, but as it is widely distributed in the Eastern

parts of Tropical Africs) it is quite likely that it may exist in

them too.

2. Xenopus levis.

Recorded by Tornier (192, p. 163) from Victoria Nyanza.

Subsequently recorded by Nieden (141, p. 186) from the same

lake and also from the western bank of the Rusisi River which

flows into Tanganyika—though not reported actually from the

latter lake itself. This form likewise has an extended distribution

in Central Africa, and may very well occur in the other lakes also.

Since both these species are so widely distributed, it is probable

that their exact distribution in the lakes is of little moment.

* For the distinctions between the two species of Xenopus consult Boulenger

(27, p. 249).

FAUNA OF THE AFRICAN LAKES. 523

PIscEs.

The fish of Tanganyika constitute no inconsiderable proportion

of the animal forms which are known from the lake, since there

are now recorded from its waters no less than 146 species. This

result is mainly due to the activities of the three British ex-

peditions and more recently the Belgian expedition under the

late Louis Stappers, though a number “of smaller collections have

added considerably to our “lsaae ledge.

There was a period during which the fish fauna of Tanganyika

had probably been more car efully i investigated than those of the

other big lakes, but at the present time this is no longer the case.

So far as Victoria Nyanza is concerned, this is due particularly

to the researches of the late W. G. Doggett, M. Alluaud.

K. Degen, and E. Bayon, while in Lake Nyasa the recent

collectors have been F. Fulleborn and E. L. Rhoades. It is

nevertheless very doubtful whether all the fish forms which in-

habit these vast inland seas have as yet been even approximately

discovered, for each extensive collection still adds new species,

often of very striking appearance. At the same time it will be

agreed that the investigations of many explorers, spread over

a fairly considerable period of years, must have furnished data on

which it is possible to rely, even though further knowledge may

occasion some revision in points of detail.

The fishes are better represented in the lakes than any other

group. Nyasa appears to form an exception, as in that case

the number of Rotifera even exceeds that of the fishes. This

is due to the inclusion of a large number of extra-lacustrine

records.

In the list which follows, 308 species of fish are enumerated,

and it is a point of considerable interest that only three of these

are recorded from outside the continent. They are Clarias

lazera, Tilapia nilotica, and Haplochronvis desfontainesti, all known

from Syria, with which, through the valleys of the Jordan and

Nile, there is believed to have been communication in former

geological times.

The outstanding features in the case of Tanganyika will be

seen to be firstly the large number of different forms found

within the limits of a single lake, and secondly the remarkably

large percentage of those forms which are not known to occur

elsewhere.

35*

DR. W. A. CUNNINGTION ON THE

Table of Distribution of Pisces *.

inna a2 Sinanies Tangan- Victoria Nyaa Albert Edward Reaue Other parts of

‘ j yika. Nyanza. * Nyanza. Nyanza. the world.

Family PoLyPrERiDZ.

Polypterus congicus ............ es a Congo R.

3 senegalus ......... 12 W. Africa, Nile.

Family LeprposirENIDz.

Protopterus zthiopicus ...... P P 18 Nile, Soudan.

Family MormMyRip2&.

Mormyrops deliciosus ......... Ie W. Africa, Zambezi.

Petrocephalus catostoma ...... Fels 1 Rovuma R.

ns decent ieane.ce: E

Marcusenius nigricans ......... abe 12 AN: East Africa.

a discorhynchus .. 1p cue 12 Zambezi.

4s petherici ......... P§ Nile.

Gnathonemus longibarbis E Victoria Nile.

Mormyrus kannume......... cae 12 we Nile.

es longirostris ......... 12 12 Bangweolo, Zambezi.

Family CLurEip®.

Pellonula miodon .............. iE

Stolothrissa tanganice......... Gen. E

Family CHARACINID®.

Hydrocyon forskalii ............ De 12 W. Africa, Nile.

be INTIGEMIUS aay oaoosn 48 Pp at Tropical Africa.

lestes dentex MEO os le W. Africa, Nile.

5, macrophthalmus...... Ie ay: P?|| Mwero, Congo, Gaboon.

See SACL OTA ek neces circ a i eet Malawa R.

eT SNES Oho t coreaseencdene es i W. Africa, Nile.

ty NOUS ONTUN. Gaede aed aacate ES ut Malawa R.

>> lmberi As er 12 Congo, Zambezi.

Fa My WAH ALUSz 6 or ceiibee Meech E

Pen od opleunsamerenstcaene E

Citharinus gibbosus ............ P

Congo R.

+ For further details relating to the Pisces, see in particular the comprehensive ‘ Catalogue

of the Fresh-water Fishes of Africa,’ by G. A. Boulenger (86), which is the authority on

which, in the main, I have relied in preparing the adjoining list. It is necessary to make it

quite clear that I follow Boulenger in regarding (1) the fishes obtained from that part of

the Victoria Nile which lies between Lake Victoria and the Murchison Falls as belonging

essentially to the lacustrine and not to the Nile fauna, and (2) the fish collected from the

Upper Shiré River (as opposed to the Lower Shiré) as constituents of the fauna of Lake

Nyasa. Dealing with the records in this sense, therefore, fishes have been described as

eudemic in Victoria Nyanza when they have been taken in the Victoria Nile as well as in

the lake itself, or even should they have been obtained only from the Victoria Nile. For

Nyasa and the Upper Shiré the same principle is adopted. In a few cases also, fishes taken

within the obvious drainage area of a lake ave included in the list, as it would clearly be

illogical to omit them. All the instances, however, in which records are not actually from

the lakes themselves, are specially indicated by a §.

* Throughout this paper, wherever a tabular form has been adopted, an HE is used to

indicate that the species referred to is endemic; P, that it is present in the lake named but

known elsewhere. Gen. E implies that the genus is endemic.

§ Not actually recorded from the lake itself. See footnote above.

- || This record needs confirmation.

FAUNA OF THE AFRICAN LAKES.

= eC Tangan- Victoria Albert Edward ,-. Other parts of

Name of Species. yika. Nyanza. Nyasa. Nyanza. Nyanza. Kiva the ead.

Family Cyprinip2.

Labeo horie ae Je Nile.

+ mesops ube K Upper Shiré R.

> Victorianus sheeneer ae E is Victoria Nile .

» eylindricus ..... lr aE. Ve Abyssinia to fambezi.

Discognathus johnstonii ...... Xe 12 K. Africa.

Varicorhinus stappersil ...... o}

tanganice ...... E

Barbus tropidolepis Ba Sasi E

>» lobogenys ws E

Bo, MISLVUENSISE Siccuiocnacedis igi K

> vradceliffii EK

> altianalis ath Pp Rusisi R.

5 tergussonli ...... ae ste E

» bayoni stee ES ae Victoria Nile.

pee ECU AONAIIILS menses -e. a E

5 eurystomus .. E

BPP NOAGESIN Gem vos/et tena: ee 10}

co. wei DIET AHOVEOUNSS Conseonsecee i

Me TNYASSH see sch sess dass E

Bee OMM SHOR tee tote. oe-. E

Sa UMUMNACWIAGUSE cee. ae ke ae: 12 AC Zambezi, Angola.

Pe TIAUIIMNINLED acne a Pat sree 1 i. Africa.

Soh MEPANTAMOSUS|...c0oten-- = J E. Africa, Natal,

» thikensis Le kan es IP 2s E. Africa. Angola.

PUES OUPILGI ee toe ccicerseasks 1p Ae Uellé R.

> minchini E Malawa R.

» tetraspilus iE

» sexradiatus ‘¢ H

PEP MUTATE oct suc sbeen stan ae y

» apleurogramma ...... 9

» imnocens ae - : K. Afinga.

Pe Team ODLEUVA) snes .)-> ee. 4}

Pete UNOS OWNA <2. ce csear i

» arcislonge E

nop. WOSRWCOanonee ae Peal Vile.

9 trispilopleura- Sete P? | Abyssinia.

» doggetti ey: E

» latukiensis 13)

magdalen E

Barilius microcephalus......... st 3} ... . Upper Shivé.

Pr IMTOO Gea cecasnenceneese 12 P

, tanganice i

PUCLOLep ISN yEe..c: E

Engraulicypris sardella as y Upper Shiré.

us argenteus ies EB

5 minutus ...... EH

Family SrnuRIpz.

Clatias anguillanis .....0.......: 5 Nile, Chad.

Fe EDO OLR eye ed sco ces Ee

SeeMMOSSAMUbIeUS: a. oss 12 se Ee Abyssinia, E. A‘rica.

PMOL AZELAN Ort: seccfaiesse oe iP 12 Syria, Nile, W. Africa.

pa GALSOMME eet eaeeelan'sioze 12 ... Uganda.

3 SUbMaALeINAbus) oo. ae 12 Cameroon, Ituri R,

», liocephalus eee 12 aoe Ubanghi R.

gf MURR, eos oc aman E

Werner ......... Bane As Pp Uganda.

Dinotopter us cunningtoni Gen. E

Eutropius niloticus ee i Ie : Nile, Senegal.

Schilbemaiysiuses seers, 1

Nile, Tropical Africa.

§ Not actually recorded from the lake itself.

|| This record needs confirmation.

——

526

DR. W. A. CUNNINGTON ON THE

ch hegt Tangan- Victoria ., Albert Edward 47,.. Other parts of

Name of Species. yika. Nyanza. “ts Nyanza. Nyanza. LON the world.

Bagrus bayad wages ae Nile, W. Africa,

ny. GHEINEUDIS 53 chon cho wne one i ia 12 E. Africa.

er OCMMAC HERE aye heyy eae 2 12 Nile.

vy OLOSCIMIE Cremer es caice eee H

rp NGAICHOMANINS cnoeceeecone ie K§ Upper Shiré R.

Chrysichthys sianenna......... E

Be GPPMUGL soonen sae E

. Cranchueseeeee }2 Congo.

5 stappersii ...... E

Me myriodon ...... 10)

i NAINCNS joe cenone E

5 brachynema 0)

Phyllonemus typus ............ Gen. E a

Amphilius platychir. .......... 1p 12 Re K. Africa.

Auchenoglanis occidentalis ... Ie 12 Nile, W. Africa.

Synodontis granulosus ......... HK

3 GIRO — suosovoo0e00 0)

Ee zambesensis ...... a6 i Zambezi, E. Africa.

a melanostictus ... P Zambezi, Bangweolo,

x multipunctatus ... E | Mwero.

e WCEOINES oo usccosann E Malawa R.

a afro-fischerl ...... E

ie WIGAN snocbonnsoao ae ana 12 Nile, W. Africa.

Malapterurus electricus ...... 12 12 Nile, Tropical Africa.

Family CyPRINODONTID&.

Fundulus teniopygus ......... ae 12 K. Africa.

Haplochilus dhonti ............ K§ Be Lukuga R.

3 FOOTIE — oeanoenc 12 12

3 johustoni ...... abe P§ Nyasaland, Zululand.

Lamprichthys tanganicanus.. Gen. E

Family SERRANIDZ. '

Watessnuiloncusmee ieee eee ae 2 Nile, W. Africa.

6 WOKGRONEINS — scensocoossanse E

> angustifrons E

Luciolates stappersii. ......... Gen. H

3 IDIGVAOT s5nd00000 coon00 EK

Family CrcHiip» +.

iil ajpiats hiirariceee eee eee 12 Zambezi.

np MOOSISENTMONGD, aoe oon oso oon zi PS§ an oe .. &. Africa, Natal.

COPA OLICAY eerste a 12 he see 12 12 12 Syria, Nile, W. Africa.

an COLE MOEN, Sob aco cee nus P§ PS§ KH. Africa.

sp WEXPRNDINS cos censs0500 ano EH ee aan Victoria Nile.

ee auromarginata ......... E

> Squamipinnis ......... me i ... Upper Shiré.

> Melanopleura ......... 12 12 12 W. Africa, Zambezi,

op WRURKO So: Sas odesooscoase 1p Pp ; { Zululand.

» adolphi-frederici ...... ee H

oy | OAUTOUOTEN Sesecrsotede vac 12 Zambezi.

a WHIDKENTISHIL,..scoces conten ses E

3 NOT Eras etc sos ee a)

§ Not actually recorded from the lake itself.

+ Quite recently, and since this section on the Pisces was completed, an important paper

dealing with the classification of the Cichlids has been published by Regan (145). Entering

on a revision of the fishes of this group, he deals in this introductory article with the

Tanganyika genera.

to revise my list in accordance with his views, and merely call attention to the paper in this

place.

Since he has not yet completed his investigations, I have not attempted

It is clear that in this tabulation I am compelled to conform to the standard

originally adopted, which cannot be brought into line with the suggestions put forth in

the above-mentioned preliminary essay.

FAUNA OF THE AFRICAN LAKES. 527

Tangan-

yika.

Albert Edward

Victoria Weisa

“~~ Nyanza. Nyanza.

Other parts of

Nyanza. ~/

Name of Species. Civ

: pecies Kivu, the world.

——

AEilapia PELVierl .:.........:<eeee

>» humilior

on pallid ates Sectteaeciso sc

>> pappenheimni

» Jacrimosa

54 nubila

+ macrops

Sep NALIN sete acs eee

55 DAVONaeseseceees- ae

5 MN ETAU ORNS \s pop ece eanloe sec

a SUM OUES ea scnraia dey tiles

En ab REED La a yee len

9 aurata UN Seton

sy Pleuwotenwtasesceses sc:

> johnstonii

ss) EebebLASWIONNG ee a2cc ene

Spo at abl loti a hacrcoeeeee ease

» lateristriga

a lethrinus

35 MOSUUALA aera

» . dardennil ....

+» macrophthalma

», brevis

x mornata S66 bo5 5008eE

» trematocephala ......

» boops Seaodeeneenee

em erandoculisme 4-4-2:

Petrochromis tanganice ......

3 andersonil

35 polyodon

53 MNVASSEES faeries

35 fasciolatus......

Cunningtonia longiventralis ..

Simochromis diagramma......

Tropheus moorii ...............

an annectens ............

Asprotilapia leptura ............

Lobochilotes Jabiatus

Docimodus johnstonii

Haplochromis livingstonii

2 WEMUSEUS <<. c.-ccs<

5 SCMUPOLAN) ease se

3 nuchisquamulatus

3 HERON! —o.%5 02-2.

s angustifrous ......

5 ASHMAN se area. be

5 MODEL Ula seeeeecos ioe

3 Stanleyieue ssaasce =.

5 percoides............

5 grauerl

Bs strigigena .........

z desfontainesii...

9 ENINDERDICO! aq agancecen

Paratilapial eesti essceecrcceo-

n parvidens

aS pfetteri ee

‘< PULE Rte aaa aes

3 modesta ............

55 thumbergii ;

3 WitiIEIEY ” Respecdanade

5 compressiceps

55 longirostris

= prognatha

lool

Gen. E

Gen.

Gen. E

eoesiies)

ferpesite>[ineiineie jae

te:

bit:

lesfesiie> mele

tan

esfeoleolacla- look

lesfepiic>]

mtd: oe

P? ||

rot:

Hoh HY:

§ Not actually recorded from the lake itself.

|| his record needs confirmation.

Victoria Nile.

Rusisi R.

Nyasaland.

Upper Shiré.

Victoria Nile.

Nile, HE. Africa.

Syria, Nile, Tunisia.

Victoria Nile.

Upper Shiré.

Zambezi, Bangweolo,

| Angola.

Victoria Nile.

== ~ ——

528

DR. W. A. CUNNINGTON ON THE

7 ne Tangan- Victoria } Albert Edward j--_ Other parts of

Name of Species. yika. Nyanza. Nyasa: Nyanza. Nyanza. ea the world.

Paratilapia serranus ............ 12 ase aah 12 Victoria Nile.

‘5 cuiarti {| A Victoria Nile.

‘ bayoni E Ee Victoria Nile.

a chilotes HS fe .. Victoria Nile.

5 pectoralis uae HS Victoria Nile.

os maculipinna ...... i

“ victoriana ......... E Victoria Nile.

fe SAM GI see ete K

a cinerea aaah H ae sprehies

Fa crassilabris ..... EH Victoria Nile.

a WHOGNO — scssssaarane KH vt Victoria Nile.

cy retrodens ......... E Victoria Nile.

5 polyodon ......... a0) 55 Victoria Nile.

35 nototeenia a K

33 Ghia HAD, soann5 005 1) Upper Shiré.

Bs rhoadesii ales K

sp chrysonota ......... E

55 intermedia .. ee H

a WORMIFAINE oon ac0 sooene 19)

op dewindti............ H)

2 lukugee nae E§ os Lukuga k.

ss THEOREM RAPEY soo nas eseo0n Hy

55 stenosoma H

re microlepis ......... EK

S leptosoma ......... E

<p nigripinnis......... KH

oA calliura ae EK

Pelmatochromis polylepis 13

Le riponianus .. 2 E§ he Victoria Nile.

ms microdon E

D obesus ...... ne H

55 ENGECTE) son ese a0)

eS SSM sooces P ie Ee Ty Victoria Nile.

sp flavipinnis .. HK :

; cavifrons Ete a0) hs , Victoria Nile.

33 frontosus ... E

Wks 6, macrops 1D)

Stl stappersil ... EK

3 pleurospilus H

rhodostigma 10)

Platyteniodus degeni : ) Gen. E

Champsochromis ‘ceruleus ... Gen. B ,

‘3 HopEreep® ie 1) be ... Upper Shiré.

eSOX . wit a9)

Bathybates eraueri Gen. H

es ferox meg eas K

5 IOP, sag goo55sa55 nnn E

a VAULENEDS soocspeoca- H

A VEICIENTUS sec ancaae E

minor . 2 auras 1)

Haplotaxodon microlepis... WAG Gen. H

Cyrtocara moorii Bee aa Gen, E

Ketodus descampsii ............. Gen. E

Hnantiopus melanogenys Gen. E

= ochrogenys ...... H

3 ? albini H

Stappersia singularis Gen. E

Xenotilapia sima sovosn G1@iMs 1B

5 ornatipinnis ast EK

Grammatotria lemairii......... Gen. E

Trematocara marginatum Gen. E .

i nigvifrons ...... K

- unimaculatum... aj

Gephyrochromis moorii ...... Gen, E

§ Not actually recorded from the lake itself.

FAUNA OF THE AFRICAN LAKES. 529

: cen a Tangan- Victoria y_. Albert Edward jy... Other parts of

Name of Species. yika, .. Nyanza. Nyasa. Nyanza. Nyanza. Kivu. the world.

Lamprologus brevianalis ...... 10)

“A tetracanthus ... y

cf marginatus... EK

io tong se. q)

a tretocephalus... K

cS INVA Aonendeor D

is multifasciatus .. E

* modestus ... .. E

Ry mondabu...... E

s ? steindachneri.. EK

ss Pboulengeri ... Ki

es elongatus ...... q

- pleurostigma ... E

os cunningtoni 4

z lemiarrnl) 222-22... E

55 callipterus ...... E

5 fasciatus sje. EK

. [Ore yign eae eee es E

“5 TOOL Re eee a E

5 compressiceps .. H

3 VES WNDU) ose naneG E

mn Callitunns;.e5..c-< E

- reticulatus ...... {)

is URUBREUREP! soon age q

Julidochromis ornatus......... Gen, EH

‘ Pocellatus...... 10)

Telmatochromis temporalis. Gen. E .

vittatus ... 1m;

Bayonia xenodonta seaindeyadaie Sica Gen. E§ vee oo - ... Victoria Nile.

Hemitilapia oxyrhynchus a oes E . i :

ss ayanie-sees ee aK E ie He on ... Wictoria Nile.

5 materfamilias ... ant u Bis ;

Corematodus shiranus ......... i fam Gen Eas vote ... Upper Shiré.

Eretmodus cyanostictus ...... Gen. E

Spathodus erythrodon ......... Gen. E

Perissodus microlepis ......... Gen. E

Chilotilapia rhoadesii ......... sh Gen. E

Schubotzia eduardiana......... See ve Gen. E

Xenochromis hecqui............ Gen. E

Plecodus paradoxus ............ Gen.

‘Family ANABANTID®. ‘

AMS DASHIMUTIOUN ee aed. e sees: : Ie ve ves ves reales

Family MasTaceEMBELID®.

Mastacembelus frenatus ...... E

33 cunningtoni.. K

% MAO OU eee sens 1)

- shivanus ...... ae sei E eb sh ... Upper Shiré.

+ victoriw ...... ar 0) te nee 6 ... Victoria Nile.

Pa ellipsifer ...... 13) :

3 mellandi ...... PS N. Rhodesia,

7 teenlatus ...... i

35 tanganice ... 10;

93 ophidium i

12 families ....... i 9 uf 6 4. 3

ULEAD EON «cocoon acon eupane 96E,29P. ° 2H,23P. 5H,20P. 14P. 12E,7P 6P

(55) (25) (25) (8)

308 species .................. 121E,25P. 60H,26P. 43H,20P. 17P. 6E,12P. 3E,10P.

(146) (86) (63) (18) (43)

§ Not actually recorded from the lake itself.

530 DR. W. A. CUNNINGTON ON THE

It is evident that a table which enumerates 308 species and

extends over several pages is far too unwieldy to convey a

correct impression of the outstanding features of distribution.

The principal table of distribution is therefore supplemented by

two tabular lists, which will serve as a summary and at the same

time emphasise the points which are most striking. The first

supplementary table supphes the number of species represented

in each lake classified under their respective families.

Number of Species represented in Lakes.

i : D ofu - navn

Nene ot —tanganyite, YOO Nyaca, ltt Elvan iy,

Polypteride ...... We tae ah iJ

Lepidosirenidex ... il 12 1 be i iL

Mormyride ...... 2 1 2K 32 4 Pp

‘Clupeidze ......... 2H

Characinide ...... 2H 3P 2H1P Ie 3P

Cyprinide ......... Qh Bie I, HIP NO 18 BI 1S eZ SE eee eRe os

SMUG Soocnnceanon LIL IB) 1 4H6P 1H 4P We a 12

Cyprinodontide... 2H1P 2P 1P

Serranidse ......... 4 a ee 1 P

ChENWClE oo, sso san ene 84H5P 40H 7P 31H 7P ATE BID Dy 7 1e

Anabantide ...... ae ide

Mastacembelide .. 7H1P 1H 1K

308 species ... 121E 25 P. 60H26P. 43E 20P. 17P. 6E12P. 3E 10P.

(146) (86) (63) (18) (13)

The other supplementary list simply gives the number of

endemic genera known from the lakes in question arranged

according to families.

Number of Endemic Genera in Lakes.

Name of Family. Tanganyika. Vhrotowne Nyasa. Swen

Nyanza. Nyanza.

Chupeideem nee 1

SHUUPBITIED seatoosesaakagaaeonne 2

Cyprinodontidee ......... il

SETAC) oouakone sscecanne il

Crchilidceere ee eee 21 2 5 ih

5 families ......... 26 2 5 1

It is clearly impossible to allude specifically to more than a

few of the points of interest which emerge from a study of the

—————-— Sl eee

FAUNA OF THE AFRICAN LAKES. 531

preceding tables. Many details of lesser importance will be

found discussed in the various papeis on the fresh-water fishes

of Africa mentioned in the bibliography or in the British Museum

Catalogue (36). Attention must, however, be drawn to the most

significant features, many of which ave now brought into special

prominence by means of the tabular method employed.

The point that first arrests attention is that Tanganyika con-

tains no fewer than 146 different forms of fish, which is in

itself a remarkable number to come from within the bounds of a

single lake. The fact becomes still more striking by a comparison

with Victoria Nyanza which contains 86 forms, eae Nyasa which

contains 63. Thus, Tanganyika has more than half as many

species again as Lake Victoria and over twice as many as Nyasa,

and this ean the fact that Victoria is a bigger lake than

Tanganyika and Nyasa not much inferior to it im size.

Jt must next be observed that of the 146 species known from

Tanganyika, no less than 121 are peculiar to that lake, while in

comparison 60 out of 86 are endemic in Victoria Nyanza and 43

out of 63 in Nyasa, That is to say, nearly 83 per cent. of the

fish of Tanganyika are found in that lake alone, while of the

species found in Victoria Nyanza, 69 per cent. are peculiar to

the lake and in Nyasa 67 per cent. Tanganyika, indeed, contains

more than double as many endemic species as Lake Victoria,

which shows the next largest total.

Although the number of endemic species in Tanganyika is

clearly remarkable, the number of endemic genera is still more

so. Of the 55 genera known to occur there, nearly half (26) are

found in that lake alone. A comparison with the two other

lakes, as instituted above, serves to emphasise the exceptional

nature of this fact. Two genera only out of 25 are endemic in

Victoria Nyanza and five out of 25 in Nyasa. The number of

genera recorded from Tanganyika forms a remarkably large

proportion of the total enumerated in the table of distribution.

A comparison of the figures shows that whereas this lake contains

146 species out of a grand total of 308, 7. e. 47 per cent., it con-

tains 55 genera out of 72—no less than 76 per cent.

Tt will now be well to review the list of fishes in rather more

detail, dealing in general terms with the various families and the

distribution of thei representatives in the lakes. ‘There is little

to comment on as far as the Polypteride and Lepidosirenide are

concerned. Species of Polypterus and Protopterus are fairly well

represented in the big lakes, but the details of their distribution

have no particular significance.—Of the Mormyride no species

have as yet been obtained from the smaller lakes, and the family

has but few forms in the three bigger ones. Only Victoria

Nyanza contains en:lemic species.—It is interesting to note that

the Clupeide are to be found in Tanganyika alone, where there is

an endemic species of the fresh-water genus Pellonaala, as well as

a representative of a closely allied endemic genus.—A few forms

of Characinide occur in the bigger lakes, of which Tanganyika

and Victoria each contain two types peculiar to themselves.

532, DR. W. A. CUNNINGTON ON THE

The family Cyprinide is better represented than the others.

hitherto considered, but this is mainly due to the number of

species of the very widely distributed genus Barbus, of which

about 250 forms have been already described from the fresh-

waters of Africa. There are types of this genus peculiar to each

of the lakes under review with the exception of Lake Albert, in

which, strange to say, no representative has yet been found.

The genus Varicorhinus, found in Asia and various parts of

Africa, occurs in the lakes only in Tanganyika, where there:

are two characteristic species. It is noteworthy of the genus

Lingraulicypris that each of the three big lakes appears to contain

a form peculiar to itself.

The family Siluride cannot be expected to afford evidence of

much value concerning geographical distribution, since many

of its members can remain for prolonged periods out of water

and even progress for some distance over land. There are there-

fore few points of significance to which attention need be drawn,

but it may be pointed out that even in this eroup. Tanganyika

5 ps gan)

exhibits two endemic genera. |*urther, it is curious to note that.

the genus Chrysichthys, while common in the Nile and Tropical

Africa, occurs in Tanganyika alone, in which six of the seven

species named are likewise endemic.

Only a few types of the family Cyprinodontide are represented

in the list, but among them is Lamprichthys, a genus peculiar to-

Tanganyika.—The genera ZLates and Lueciolates ave the only

members of the Serranide to be found in the big lakes, and of

the species which occur, only one—the well-known Lates niloticus

—appears outside the contines of Tanganyika. Thus in this.

group also there is a genus (Luciolates) endemic in that lake.

While the families hitherto considered show various noteworthy

features of distribution, it is the Cichlide whieh far surpasses.

them all, both in interest and in the number of forms represented

in the African lakes. Of all the species enumerated in the table-

of distribution, more than 70 per cent. are Cichlids, or to put it

another way, there are considerably more species of the family

Cichlidee than of all the other families taken together. In the

case of the genera, exactly half ave Cichlids, there being 36 out

of a total of 72. These are remarkable facts, but they are not

merely an expression of the common occurrence of this group of

fishes on the continent, as an examination of the list will show.

It is quite clear that there are groups of forms which are peculiar

to these big lakes, and that the three biggest contain more

than the smaller ones, while a culminating point is reached in

Tanganyika with a succession of endemic genera.

Like Albert is alone in containing no endemie species, but the

numbers grow from 2 and 3 endemic forms in Lakes Kivu

and Edward respectively, to 31 in Nyasa, 40 in Victoria Nyanza,

and no less than 84 in Tanganyika. Nor is this all, for in the

number of endemic Cichlid genera the remarkable nature of

the fauna of Tanganyika is particularly exemplified. In the

FAUNA OF THE AFRIJAN LAKES, 533

latter lake 21 peculiar genera occur, as compared with 2 in

Lake Victoria, 5 mm Nyasa, and a single genus in Lake Edward.

As coneerns Tanganyika, the cas? may be stated even more

forcibly by pointing out that 41 genera in all are recognised

from the whole continent, of which 21 belong exclusively to this

lake: in other words more than half the genera of Cichlide

known in Africa are confined within the limits of Tanganyika.

ft may be worth while to give a few more comparative figures

in the case of the three bigger lnkes around which interest

principally centres. In Nyasa 38 Cichlid fishes are found out

of a total fish fauna of 63, which is 60 per cent. of the rea

in Victoria Nyanza 47 out of 86, which is some 54 per cent. ;

‘Tanganyika 89 out of 146, or nearly 61 per cent. of the eo,

With this remarkable total of 89 Cichlid fishes, Tanganyika, in

fact, establishes another record, possessing the richest Cichlid

fauna in the world. Next note how extremely large a proportion

of the endemic species are Cichlide. In Nyasa there occur 31

endemic Cichlid species out of a total of 43 endemic species in

the lake, z.e. 72 per cent.; in Victoria Nyanza 40 out of 60, or

66 per cent.; in Tanganyika 84 out of 121, or 69 per cent. The

proportion of endemic genera belonging to this family is even

more extraordinary, for all the, genera which are endemic in

Nyasa and in Lake Victoria are Cichlidie (the same is true of

the single endemic genus found in Edward Nyanza), and in

Tanganyika there are 21 out of 26 endemic geneva, or 80 per

cent. Lastly, the number of endemic Cichlid forms as compared

witli the total number of Cichlid forms known from the lakes in

question is a basis for further striking figures. Thus in Nyasa,

out of 38 species of Cichlid, 31 are endemic, or 81 per cent.; in

Victoria Nyanza, out of 47 species, 40 are endemic, or 85 per

cent.; in Tanganyika, out of 89 species, 84 are endemic, or over

94 per cent.

From a consideration of all these figures it becomes very plain

that the large number of unique forms characteristic of Tan-

ganyika is especially due to a great development of the fishes of

this group, and that the same is true, though in lesser degree, for

Nyasa and Victoria Nyanza.

The review of so largely represented a ane must of necessity

be brief. The very characteristic genus Vilapia extends to

AQ species in the list, but though some are peculiar to certain

lakes, there are no very striking features of distribution to be

noted. The genus Petrochromis appears to be mainly typical

of Tanganyika, for though an endemic form occurs in Nyasa and

there is a doubtful record from Lake Albert, it is not found in

Victoria Nyanza or any of the other lakes considered. The case

of Haplochromis is interesting as being, on the contrary, a genus

with a single widely distributed species recorded from 'Tan-

ganyika, while it is better represented in all the other lakes

under review. Perhaps a more important—though unexpected—

feature is the existence of H. angustifrons and H, graueri in both

534 DR. W. A. CUNNINGTON ON THE

Lakes Edward and Kivu and nowhere else. This is a matter of

some interest, since there is evidence for believing that these two

lakes, between which there is now no connection whatever, were

in relatively recent times in communication with one another

(cf. Moore, 187, p. 89). The genus Paratilapia is another repre-

sented in the hst by a large number of species, these being

associated in groups which in most cases are peculiar to one of

the three bigger lakes. Pelmatochromis, which is widely dis-

tributed in West Africa, occurs in Tanganyika and in Victoria

Nyanza, but otherwise only in a single instance (Lake Edward),

in the lakes with which this paper is concerned.

There is no occasion to refer individually to the numerous

Cichlid genera which are endemic in the African lakes, the

important point bemg that so many of them are found in Tan-

ganyika. Attention may, however, be called to Champsochromis

from Nyasa with three species, and Lathybates from Tanganyika

with six, as these exhibit the greatest number of species among

the different endemic genera. The genus ZLamproloyus is otf

considerable interest and importance, being represented by a

total of 27 species from the Congo and Lake Tanganyika. It is

nevertheless essentially characteristic of the latter, for 24 endemic

species are recorded from the lake, while the remaining three

species are peculiar to various parts of the Congo river system.

The distribution of the three forms of Hemutilapia is quite

different from anything else met with in the list, since the genus

is only known from Lakes Nyasa and Victoria. A single endemic

species occurs in the former lake and two endemic species in the

latter.

Since the Cichlids are so pre-eminent among the fishes of

Tanganyika, and indeed constitute a most striking feature of the

lake fauna as a whole, it becomes necessary to refer to certain

conclusions which have been drawn from a study of the anatomy

of these particular forms. From various considerations, such as

the number of rays, the multiple lateral lines, and the number of

vertebra, Boulenger (20, p. 2: 25, p. 393) at one time regarded

certain Tanganyika genera as showing features of generalisation.

At a later date, however, after a further investigation into the

anatomical evidence bearing on the phylogeny of the group,

he arrived (26, p. 422: 38, p. 43) at a different conclusion,

and considers the Tanganyika genera and species as essentially

specialised. This is an important pronouncement, since it brings

the case of these fishes into line with most of the other animal

groups under discussion, in which certain marks of specialisation

are found to be characteristic of the endemic Tanganyika forms.

Passing now to the family Anabantide, a solitary representative

(Anabas mufiei) has been obtained from Victoria Nyanza. It is

a Nilotic type, and its existence here calls for no particular com-

ment.—The last group in the list, that of the Mastacembelide, is

represented in Africa only by the genus MJastacembelus itself.

Many species are known fiom the fresh-waters of the continent,

FAUNA OF 'THE AFRICAN LAKES. 535

but examples are found only in the three biggest of the lakes

included in this survey. It is interesting to note that while

there are single forms peculiar to Nyasa and Victoria Nyanza

respectively, there are no fewer than seven endemic forms in

Tanganyika. Here there is again exemplified in very typical

fashion the striking nature of the Tanganyika fauna.

In his work “The Tanganyika Problem” (187. p. 146) Moore

has pointed out that, on the whole, the number of different

animal forms present in any particular African lake is definitely

proportional to its size. He expressly omits, in making this

statement, the so-called “halolimnic’”’ forms which he distin-

guishes in Tanganyika, and deals only with what he calls the

ordinary fresh-water fauna. Regarding as inadmissible the

exclusion of certain forms as “ halolimnic,” Tanganyika will be.

seen to far outstrip the other lakes in the number of types.

represented, but with that exception, the figures undoubtedly

do vary in proportion to the size. So extensive a list as that of

the fishes affords an excellent opportunity for testing the general

accuracy of this statement, and a comparison may be instituted

not only as regards number of species, but as regards number of

genera and even families. The results ave not dissimilar in the

three cases, as the adjoining table will show. With the ex-

ception of Tanganyika, the lakes are arranged in order of size,

te!

from the largest to the smallest of those under review.

Tangan- Victoria Neder Albert Edward Ker

yika. Nyauza. yasa- Nyanza. Nyanza. cls

Number of Species ......... 146 86 63 17 18 13

a Generaye was 55 25 25 14. 8 6

3 Families......... 11 9 7 6 4, 3

Having thus emphasised from many points of view the re-

markable nature of the fish fauna of Tanganyika, an examination

into its significance might well follow. As however the matter

merits a detailed discussion, and is moreover by no means con-

cerned with the distribution of the fishes only, this must be

deferred. In this place it is enough to indicate that the evi-

dence points to a period of complete isol

period which was long enough for the inhabitants of the lake to.

assume the characters of species, and even genera, distinct from

those of the neighbouring parts of the continent. This is indeed

fully borne out by a comparison of the fish fauna of Tanganyika

with that of the Congo, to the drainage area of which the lake

now belongs. The striking dissimilarity of the fishes of Tan-

ganyika and the river into which it drains may be illustrated

by pointing out that of 25 species non-endemie in the lake, only

536 DR. W. A. CUNNINGTON ON THE

14 are known also from the Congo system, and that of these

about one-third are of very wide distribution, by no means

characteristic of the Congo,

A few lines may now be devoted to a general survey of the

fishes of the other big lakes, for the purpose of pointing out any

characteristic features and affording a ground of comparison with

Tanganyika. It has already been seen that Victoria Nyanza,

while containing representatives of nearly.as many families as

Tanganyika, contains a much smaller number of species. Very

few endemic genera are found in the lake, but a considerable

number of endemic species occur, especially among the Cichlide,

Tt may also be noted that Lake Victoria shows a larger number

of Mormyrids (including two endemic types) than does Tan-

ganyika. Again, as regards the Cyprinide, this lake is richer in

forms, with 11 endemic species mostly belonging to the genus

Barbus, while the Anabantide is represented here alone.

Speaking generally it may be said, that although now one of

the sources of the Nile, only a small proportion of its fish are

found also in that river, so that there are good reasons for sup-

posing that Victoria Nyanza was also isolated for a certain

period, and only included in the Nile basin in times, geologically

speaking, recent. Meanwhile it is clear that the modifications

of form which exist are not comparable in importance to those

occurring in Tanganyika, but go to prove that the lake has a

peculiar fish fauna, not closely related to that of either Nile or

Congo. ;

Nyasa, having a lesser area, shows something of that decrease

in number of types which is related to the smaller size of the lake.

Nevertheless it still contains more Cyprinide and Mormyride

than Tanganyika, and it is relatively rich in Cichlid fishes. Of

the latter there are indeed five endemic genera, yet out of a total

of 38 species, 27 belong to the two genera Vilapia and Para-

tilapia, which are of very wide distribution. Thus, although

Nyasa possesses a fish fauna which is in part peculiar and

characteristic, it has a far less striking assemblage of forms

than Tanganyika, or even than Victoria Nyanza. While it is

probable that a period of isolation favoured the production of

species peculiar to Nyasa, it 1s nevertheless clear that its fishes

belong essentially to the Zambezi river system, and do not differ

from those of that river to the same extent as do the fish of

‘Tanganyika, and to a less degree of Victoria Nyanza, from those

vespectively of the Congo and the Nile.

Speaking of the fishes, it may therefore be said that each of the

three largest lakes contains a group of forms peculiar to and

characteristic of itself. These forms differ from the fish of the

rivers which drain the lakes, but not to the same degree in each

instance. Nyasa exhibits a number of types which are not found

in the Zambezi, and Victoria Nyanza a larger and more diver-

gent series of forms which are not found in the Nile; but in

these eases the differences are for the most part only specific.

FAUNA OF THE AFRICAN LAKES. ¥ 537

Tanganyika, however, is the extreme case, where not merely

specific but generic distinctions occur, and a remarkable fish fauna

exists which has very little in common with the fishes of the

Congo, and which, moreover, has no parallel elsewhere.

Albert Nyanza is the only lake of those under review which

does not contain a single endemic species of fish. The family

Siluride is best represented, with 7 different forms, belonging to

6 different genera, bub these are without exception well-known

Nilotic species. Very few Cichlid fishes are found in the lake,

the most interesting being Petrochromis andersonii, a Tanganyika

type, the existence of which in Lake Albert needs confirmation.

With this exception, and with the exception of Alestes macro-

phthalmus, a Congo-Tanganyika type, the occurrence of which in

the lake is also rather doubtful, the whole of the fishes belong

essentially to the Nile system, of which the lake is thus shown to

be an integral part.

Lake Edward, which lies further south, bat is similarly in

direct connection with the Nile, is nevertheless not without types

peculiar to itself. Two endemic speces of Barbus have been

described, and one of Claritas. The Cichlide are well represented

here with 12 forms, 3 of which, including the genus Schubotzia,

are endemic. While the greater number of the fishes are either

Nilotic types or are peculiar to the lake, there is a series of

6 Uichlids which are common to Lakes Edward and Victoria and

are not found elsewhere. This is a rather unexpected state of

affairs, and it is by no means easy to account for the association,

since the lakes do not communicate in any way. It may be noted

that in Lake Edward the total number of genera is reduced to

eight and the number of families to four.

Dealing lastly with Lake Kivu, the smallest lake is seen to

contain the smallest number of different species of fish. Yet

even here an endemic species of Barbus and two endemic Cichlids

occur. Apart from these, the fish fauna gives indications of

being considerably mixed. ‘Three species belong clearly to the

Nile group, while reference has already been made to the two

forms of Haplochromis found in Lakes Kivu and Edward only.

On the other hand, two fishes are found in Kivu and Tanganyika

only: a third occurs in Vanganyika and the Rusisi River which

drains Lake Kivu, as well asin the lake itself. The two remaining

types have a wider distribution, extending into West Africa.

In works dealing with Geographical Distribution it has- long

been recognised that undoubted affinities exist between the

animals of Southern Asia—and of the Indian Peninsula in par-

ticular—and of East and Central Africa. While evidence of this

is more marked in some groups than in others, clear testimony

as to the general truth of the assertion is to be gained by a study

of certain families of fishes. Of the families under discussion,

it is pre-eminently the Cyprinide and Mastacembelidee which

exhibit this affinity in a striking degree. Among the Cyprinide,

various genera are common to India and Africa, and a close

Proc. Zoot. Soc.—1920, No. XXXVI. a6

538 DR. W. A. CUNNINGTON ON THE

relationship is sometimes recognisable even between species. The

genera found in the African lakes which best exemplify this

relationship, are Labeo, Discognathus, and Barbus (of the group

allied to Barbus bynni). Mastacembelus itself is the only genus

of the Mastacembelidze which occurs in Africa, but it is also well

represented in India and the Malay Archipelago. This case is

specially interesting, since the genus was at one time considered

quite characteristic of the Indian region, yet at the present day

an even larger number of forms is known from the continent of

Africa.

Summary.—The lakes of Central Africa contain a large and

interesting assemblage of fishes. The bigger lakes as a rule possess

more forms than the smaller, but while Nyasa and Victoria

Nyanza each possess an extensive fish fauna, Tanganyika can

show a far larger number of types. In Nyasa and Lake Victoria

a considerable percentage of these types are endemic, but this is

more strikingly the case in Tanganyika, where indeed not merely

endemic species, but a very exceptional number of endemic genera

are found. Of the 12 families which are represented, interest

centres in particular on the Cichlide, which are very promi-

nent in all tie lakes except Albert Nyanza. A large proportion

of the Cichlid species are endemic in the lakes where they occur,

the extreme case being that of Tanganyika, whivh, with 84

endemic species and 21 endemic genera, possesses the richest

Cichlid fauna in the world. There are reasons for believing that

these remarkable endemic Cichlidee show signs of specialisation.

An explanation of the unique fish fauna of Tanganyika is

probably to be found in a period of complete isolation of that

lake: it is possible that isolation for a lesser period has produced

an effect in the cases cf Victoria Nyanza and Nyasa. The fishes

of the families Cyprinide and Mastacembelide in particular

exhibit indications of Afro-Indian affinities such as are displayed

by other groups of organisms.

PoLyzoa,

The representatives of this group, as far as Lake Tanganyika

is concerned, are of particular interest. This is due in the first

place to the fact that while examples of the Phylactolemata were

quite to be expected, the latter are accompanied in Tanganyika

by two species of Gymnolemata, the members of which are, with

few exceptions marine. It is also true that the gymnolematous

form which was first discovered by Moore (187, p. 295) has a ve-

markable resemblance to the marine genus Arachnidium. With

the exception of this type, which Moore named Arachnoidea™,

the species from ‘Tanganyika were all obtained for the first time

by the author during the Third Tanganyika Expedition.

* T follow Annandale (7, p. 198) and Harmer (99, p. 50) in adopting Arach-

noidea as the correct spelling of this generic name, which Moore originally wrote

Arachnoidia.

FAUNA OF THE AFRICAN LAKES, 539

Table of Distribution of Polyzou *.

: : Tangan- Victoria Albert Edward Other parts of

Name of Species. a ie : ae eae

I yika. Nyanza. Nyanza. Nyanza. the world.

Order GyMNOL@MATA.

Victorelia symbiotica ..... eee he LET, Birket Qarun ?

Avachnoidea ray-lankesteri... EH

Order PHYLACTOLmMATA.

Fredericella cunningtoni ...... KH

Plumatella NG OGVCST i poenuo succes 1B Je P 1B) Cosmopolitan.

‘ emarginata ...... at \P 2 re Cosmopolitan,

A (Afrindella)

tanganyike ... P Mt a ie India.

GISpecles ee eReeree ecen ic Lia) ie 2P 2P 2P

The above table illustrates an interesting feature of distri-

bution, paralleled in the case of other organisms, where one or

more than one almost cosmopolitan species occurs in Tanganyika,

associated with a larger number of endemic forms. Plwmatella

repens, a form with a most extensive range, is represented in

Tanganyika as well as in Lakes Victoria, Albert, and Edward,

while the closely related P. emarginata is found in the latter

three lakes, but has not been recorded from Tanganyika. The

case of P.(Afrindella) tanganyike is of considerably more interest,

for it was originally described as P. tanganyike by Rousselet

(149, p. 252) from material collected by the Third Tanganyika

Expedition. it was at that time believed to be peculiar to Tan-

ganyika, but Annandale (8, p. 225) subsequently referred to this

identical species, specimens which he obtained from Igatpuri

Lake in the Western Ghats, Bombay Presidency *. Still more

recently (8, p. 140) the same author established the subgenus

Afrindella to receive the species, indicating in the name the

striking feature of its distribution. The absolute identity of

forms from Tanganyika and from India is a most remarkable

instance of those Afro-Indian affinities to which reference has

already been made £.

The remaining phylactolematous form, Mredericella cunning-

toni, has sufficiently marked peculiarities to distinguish it from

the well-known /. swltana, which has an almost world-wide

distribution. The former is confined to Tanganyika as far as is

at present known.

Among the great lakes of Central Africa, Tanganyika alone

has been found to contain representatives of the Gymnolemata.

+ For descriptions of most of these species and remarks on distribution, see the

report by Rousselet on the Polyzoa of the Third Tanganyika Expedition (149).

* These specimens he at first named P. bombayensis (4, p. 169)—a species after-

wards discarded.

{ A table of African and Indian Polyzoa is given by Annandale in his paper on

the resemblances between African and Indian fresh-water faunas (10, p. 582).

540 DR. W. A. CUNNINGYON ON THE

The well-known genus Victorella, essentially a brackish and fresh-

water genus, 1S represented by the form V7. s ymbiotica, a species

found associated with and growing through the sponge Spongila

tanganyike, During an expedition more recently undertaken to

the Birket Qarun in Lower Egypt, on which I was accompanied

by C. L. Boulenger, further specimens of a gymnolematous

Polyzoon were collected, which were submitted to Rousselet for

examination. No definite report on these has been received, but

some of this material has evidently been transmitted by Rousselet

to other writers on the Polyzoa, for Annandale (7, p. 197) records

his belief from an examination of the specimens, that the Qarun

species is identical with Victorella symbiotica from Tanganyika.

Braem, on the other hand (53, p. 33), whilst affirming that the

forms V. continentalis, V. bengalensis, and that from the Qarun

have close affinities with the EHuropean V. pavida, asserts that

‘“‘Rousselets V. symbiotica aus dem Tanganyika .... durch

einen abweichenden Bau des Darms ausserhalb dieses Kreises

steht ” (Joc. cit. p. 34).

Be this as it may, there is a further matter concerning this

genus which demands consideration. The view generally held by

specialists on this group has been that Victorella only recently

migrated from the sea into brackish and fresh-water. Braem,

however (53, p. 34), in the light of the Qarun specimens and ibe

species he describes from an inland lake in Turkestan, declares

his belief that on the contrary the genus is one which primitively

adapted itself to fresh-water conditions, and that the specimens

in question are “relict” forms. He considers that the existence

of the genus in Tanganyika further strengthens his supposition.

Since Tanganyika has been claimed by Moore as an undoubted

‘“‘ Reliktensee,” the true nature of this Polyzoon genus be-

comes a matter of some importance. Further light is fortunately

shed on this problem by the recent discovery by Harmer (99,

p- 45), in the Siboga material from the Dutch East Indies, of a

truly marine species of Victorella. The usual view would thus

seem to be definitely supported, and the existence of V. sym-

biotica in Tanganyika, while of considerable interest, would not

tend to prove the occurrence in the lake of an ancient marine

fauna. :

The second gymnolematous type—<Arachnoidea ray-lankestert

—is certainly the most remarkable form from the African fresh-

waters. Despite recent discoveries, it remains one of the few

instances known of a fresh-water incrusting gymnolematous

Polyzoon, and is of peculiar interest accordingly *. In con-

sequence of its undoubted resemblance to the marine genus

Arachnidium, to which reference has already been made, Moore

(187, pp. 330, 332) regarded it as an important member of that

group of primitive marine forms which, he contended, still exists

* Another genus is Hislopia.

FAUNA OF TUE AFRICAN LAKES. 541

in Tanganyika. Quite a different conception now suggests itself,

in view of the recent description by Harmer (99, p. 50) of another

species belonging to this genus collected by the Siboga Expedition

in the Straits of Makassar. No one would have ventured to predict

that Moore’s genus would be found to have a representative still

living in the sea, but the discovery shows that, as in the case of

Victorella, the Tanganyika species has affinities with a present-

day marine type and not problematical affinities with marine

Polyzoa of a past era. It may be pointed out that a further

instance of Afro-Indian associations is afforded by the finding of

this species of Arachnoidea in East Indian seas.

A piece of negative evidence of some interest, but to which

perhaps no great importance need be attached, is the apparent

absence of Polyzoa from Lake Nyasa. It is admittedly a fact

that these organisms are small and inconspicuous, and little

likely to attract the attention of any but skilled naturalists ;

moreover, their distribution in the African lakes has in some

cases been merely deduced from a discovery of their sessile stato-

blasts. On the other hand, Nyasa has received a considerable

amount of attention from a trained observer in the person of

Prof, Filleborn, and it is certainly strange that (so far as I am

aware) he collected no representatives of this group from that

lake.

Polyzoa are also unknown in Kivu, where Schubotz states he

was unable to discover them (156, p. xiv). This is less surprising

on account of the unusual salinity of the lake.

Mo.uusea.

It is a task of exceptional difficulty to give an accurate account

of the mollusecan fauna of the big lakes. This is in a large

measure due to the work of the late J. R. Bourguignat, who

described a very large number of types, a considerable but varying

proportion of which have not been accepted by other authorities.

From Tanganyika alone, Bourguignat described no less than 242

species *, and to this total must be added a number of species for

which other writers are responsible, yet in the opinion of some of

the principal specialists the number of different molluscan forms

living in that lake is very much smaller than the enormous total

which has been credited to it.

The great interest attaching to certain Tanganyika molluscs

owing to their remarkable marine-like appearance has already

been referred to, and on account of this, it is the more to be

regretted that both genera and species have been unduly multi-

plied and that a generally accepted list of forms from this lake is

not available. In writing a paper such as this, it is necessary to

compile a definite list, and in doing so, I have followed more

* Speaking generally it may be said that Bourguignat and his school, of which

Locard was the principal exponent, elevated species into groups, and varieties into

species,

549, DR. W. A. CUNNINGTON ON THE

particularly the account given by Edgar Smith (170), who had an

unusual opportunity for “the sy stematic study of African fresh-

water shells.

To a lesser degree, the “species-making ” of Bourguignat has

affected the lists ‘of mollusca from the other big lakes of Africa,

and in making a selection of forms to be retained and forms to

be rejected, I have no doubt acted arbitrarily. ‘There is no

need to enlarge upon this point, but while all the lists have

been carefully considered with a view to the elimination of

synonyms, they must be taken merely as the writer’s expression

of opinion. It is only necessary to emphasise that while finality

in such a matter is always impossible, it is especially so in the case

of the Mollusca.

Although the number of genera and species to be included in

the fauna of Tanganyika may be a matter of opinion, there are

certain striking facts which remain undisputed. Firstly, it is

only among the Gasteropods that species described as thalassoid

are to be met with; that is to say, all the Lamellibranchs are non-

thalassoid. Secondly, while there are a number of non-thalassoid

Gasteropods known to live in the lake, there are more than twice

as many which are considered to show a thalassoid aspect.

The arrangement of the families and genera which has been

adopted is, in the main, that given by Pelseneer (Treatise on

Zoology, Ed. by Lankester, Part V. Mollusca).

Gasteropoda.

It has more than once been emphasised that the Gasteropoda are

among the most remarkable of the peculiar animal forms which

inhabit Tanganyika. In the adjoining table of distribution, a list

is given for each of the lakes under review, of those species which

appear to be admissible. This list reaches a total of 133 in all,

and it is evidence of the unique position which Tanganyika holds,

that 84 of these forms are recorded from that lake, 76 of them

being unknown elsewhere. Of these endemic ‘Tanganyikan

species, the great majority were described by Bourguignat and

others as thalassoid and alternatively by Moore as halolimnic

(cf. p. 517). It has been thought desirable, in view of the special

interest attaching to these types, to mark with a * all the genera

which have been so designated. Perhaps it is well to explain that

none of the species belonging to genera which occur elsewhere

have been described as thalassoid by the conchologists, though

this does not mean that all the endemic genera are necessarily

thalassoid in aspect, there being a notable exception in the case

of the genus Veothauma,

FAUNA OF THE AFRICAN LAKES. 543

Table of Distribution of Gasteropoda *.

Name of Species.

Family VIvIPARID2.

Viviparus constrictus

- costulatus

. foal

a meta

5 rubicundus

unicolor

? brineatianus

Neothauma tanganyicense ...

Cleopatra emini

guillemeti

jouberti

”

bE]

”

PUUOU Ms aces scusce

PEISUIGAAN ese scans

ABN dOuxIa SITU Ieee 2-25 =e

Family AMPULLARIID.

Ampullaria bridouxi

letourneuxi

se ovata

speciosa

3 stuhlmanni

Lanistes ellipticus

jouberti

nyassanus

olivaceus

ovum

schweinfurthi

3 sinistrorsus

*Leroya bourguignati

eS)

Family HypRopiipm.

*Spekia zonata sae

*Tanganyicia rufofilosa

*Rumella neritinoides

*Stanleya giraudi

5 rotundata

3”

PUEDUECUST seenen esses

Smt Mina... see

Ordon ee es

VICKERY anger PoceR Bae

TYAN as. Secs

*Limnotrochus thomsoni ......

Ohi tras karkdt eee sciee scases

Bithynia alberti

a humerosa ......

multisulcata

4 Stamleyi'his 3 ace:

walleri

*Syrnolopsis carinifera

43 lacustris

Tangan- Victoria By. Albert Edward j1,.. Other parts of

yika, Nyanza. BES Nyanza. Nyanza. i the world.

~ E

1p ie 1p eh R. Nile.

ee P 1p P P Egypt, E. Africa.

Gen. E

12 P E. Africa.

one 12 E. Africa, Abys-

E [sinia.

Gen. E

nee 1p E. Africa ?

12 K. Africa.

1p P 1 ae 1p Egypt. E. Africa.

Je 1B E. Africa.

us 12 FB. Africa.

he 12 EK. Africa.

1p Nyasaland.

1p E. Africa.

12 E. Africa.

ace Ve E. Africa.

P E. Africa.

Gen. E

Gen. Ei

Gen. E

= sé te ye Pe

eg 12 P 12

Gen. E

+ Additional particulars and bibliographies will be found in von Martens’ account

ot East African forms (116) and Smith’s review of the Mollusca of Tanganyika

(170).

There have been but few additions since the dates of these )apers,

* Genera described as thalassoid,

D44

DR. W. A. CUNNINGTON ON THE

Name of Species.

Tangan- Victoria

yika. Nyanza.

Nyasa.

Albert Hdward

Nyanza. Nyanza.

Kavu

Other parts of

the world.

Family TraRip2z.

Tiara || admirabilis .........+5 E

= arcuatula

x liricincta

ts nodicincta

i nyassana

2 pergracilis

3 polymorpha ..

eS pupiformis

‘ simonsi ........-- Reicha ee

5 tanganyicensis ...... K

i tuberculata ............ 12 1?

turritispira se

*Giraudia foai . sccocn (GxEio JB

eae andidieriana ...... K

i horei panes We H

i lavigeriana ......... K

53 FADIAOWE o09 spaneaqes 50400 E

5 JORENCINAD, ooo gosace onc H

i quintana suk E

tanganyicensis ...... HK

«Lechaptoisia ‘ponsonbyi inet Gen. E

*Burtonilla terebriformis ...... Gen. E

*Baizea giraudi eee Genny

Wy leucoraphere eee E

*Anceya admirabilis ............ Gen. E

rp SRMEENWGN sonnaa sooson-coneas a0)

a rufocincta oo... E

*Bythoceras iridescens ......... Gen. E

55 TRDNTOVP aoaonaataccaece EH

*Paramelania pridouxi ......... Gen. E

un crassigranulata .. E

damnit saeeus K

#Joubortia baizeana.........-.- Gen. E

a spinulosa ............ E

stanleyana............ 10)

*Lavigeria callistay.).. ce) Gen. BI

3 GOPENH con congao ane aoe EK

a GOROBENE) goo ssn nones KE

fe diademata ......... E

a PABINGWS .osonesscavsd0 E

FOUN! 55 osbsso don c00 E

. }OEELONTTNEY, Soo aoa goonen EH

5 ruellaniana ......... E

*Randabelia catoxia ............ Gen. E

a Ihiamniyanaye essen K

*Edgaria bourguignati ......... Gen. E

oy crassilabris ...........- E

» lechaptoisi H

» nassa Be eS" HK

% paucicostata neon eee aD)

» reymondi H

a GHAAWAAG 355 45000008 EK

5 tiarella E

VATIAD IIS eee eaen cee a0)

*Hirthia elobosa.. Bes Gen. Ei

a hittorina~ Bae e oun H

Family TrPHoBiip™,

HINT) OVE) WNOVWE 53.5.snp ann op soan con Gen. E

*Bathanalia howesi ............ Gen. E

fesilesileaesyte>|imels

lesjiac)

P Pp Pp

Angoni Land.

Tndia, Malay Pen.

* Genera described as thalassoid,

|| Olim Ielania,

FAUNA OF THE AFRICAN LAKES. 545

Tangan- Victoria

Albert Edward aa Other parts of

yika. Nyanza. ’

Name of Species.

Name of Species Nyanza. Nyanza. the world.

Nyasa.

Family Limnmipz.

Limnea natalensis ............ 12

S| CIYANSe teeeeaneeec

3) cundussume sees. Efe

Msi oraicoullosiaessseses ess K

pe TOUSK AYR ceer eee se A 1p Xs a age ... Egypt, Natal,

sy LM ASSANAN eee tee ae He K (Angola.

* randabelue sess 1)

OUP stticasapa meanest a 2 Bis &. de ... HH, Africa.

5, succineoides ............ Be ae 1)

PU ebLANSVersaliseecenesee- ae HE

Ay LODIEXONEN soe pmnangospdhoose &e 0

Physopsis africana ee a ide 12 ae sh ... KH, Africa, Natal.

FA OVOIGER ee ceenen ce nO 12 ax be she ... 4, Africa,

5 fanganyice ......... EK

1 Se ve) ee Navale

P ... HH. Africa.

lasHies}iae]

Family PLANORBIDZ.

Planorbis adowensis ............

is alexandrina .........

3 ENOANBS) cco cbhconesoune

= bridouxiana .........

95 choanomphalus_...

ss crawfordi ............

53 eibbonsi “sae

os J ial Bese censtacee at

Pr lavigerianus .........

= INOMCE Libee eee

3 Stamll eye easeaces tera.

i Sudamicus) -....2s..-:-

5 VWACUOMED Sadoepnenadants ae y

a wis ie ... Abyssinia.

12 as ee ... Egypt.

i 62 _.. Cape Colony.

1p fi eEeAunica:

soliac)iae)

bd: ee:

P oe ReeNule?

Family ANCYLIDm.

Ancylus stuhlmanni_......... an 1D)

> tanganyicensis ...... E

Shomulliesimpeee eee ae ene Lenses C2", (le) 5 1eh 6P. 2P.

ee ee

SIS; CTC), cadcaoasconecesbenuene PU ONL Be Tk Wee 9P. Gk: G12, 2P.

(35)

MSSISPECIES) 0.0 6. ct. ae d..e.2 COLL. 17 1OR 14 Ps 4b ORs ih SPs 2p

(84) (28) (24) (13) (10)

A list such as this, with 133 species, requires a somewhat

detailed analysis to bring out the points of most interest and

importance, although, indeed, the tabular form reveals the excep-

tional nature of the Gasteropod fauna of Tanganyika. In the

first place it will be noticed that no less than 84 species are

recorded from Tanganyika, which is clearly an exceptional number.

That this is so, is shown by a comparison with the known fauna

of the two lakes which come nearest in size, for Victoria Nyanza

can muster but 28 forms and Nyasa only 24. Thus Tanganyika

contains three times as many Gasteropods as Lake Victoria and

more than three times as many as Nyasa.

Secondly, be it observed, that of the 84 Tanganyika species, no

fewer than 76 are endemic, while correspondingly only 11 out of

28 are peculiar to Victoria Nyanza and 10 out of 24 to Nyasa.

546 DR. W. A. CUNNINGTON ON THE

Of the smaller lakes, Albert Nyanza has 4 endemies out of a total

of 13, Edward Nyanza 1 out of 10, while Kivu only contains two

species, neither of which is dene In other words, more than

90 per cent. of the forms in Tanganyika are only known from that

lake, while Victoria Nyanza has 39 per cent. of endemic species

and Nyasa some 41 per cent.

In the next place it must be emphasised once more that the

greater number of the endemic species of Tanganyika are types

which have been described as thalassoid—58 out of 76 belonging

to that category. Thus there are more than three times as many

thalassoid as non-thalassoid endemie species, actually some 76 per

cent. having this characteristic appearance. Even when the non-

endemic forms are added to swell the total, the shells having this

marine aspect outnumber the normal series by more than two to

one, there being respectively 58 and 26 species.

The figures which refer to the genera are even more arresting

than those which have been dealt with. Tanganyika alone con-

tains one or more representatives of each of the 35 genera named

in the table. The number of genera represented in Lake Victoria

sinks to 11, but it is significant that this 1s due to the absence

from that lake of the large total of 24 genera which are peculiar

to Tanganyika. Of this total of 24, 23 are regarded as thalassoid,

the single exception being the genus Weothawma, as already

mentioned, Apart from these endemic genera, both Tanganyika

and Victoria contain species belonging to the same 11 genera—

those of the ‘normal African fresh-water fauna” to adopt

Moore’s term. It is interesting to observe that while 28 species

of these less specialised genera (including 11 endemics) occur in

Victoria Nyanza, 25 (including 17 endemics) are found in Tan-

ganyika. Lake Nyasa with 9 of these ordinary fresh-water

genera comes next, and the other lakes follow with still smaller

numbers. No endemic genera are to be observed outside Tangan-

yika. It is thus clear that over and above the representatives of

certain well-known fresh-water genera, there is, in Tanganyika,

a whole series of unique Gasteropods which are not represented

elsewhere.

Tt may not be unprofitable to institute a comparison with the

group of the fishes, in which alone so large a number of endemic

genera is known. Tanganyika contains 25 endemic genera of

fishes, as compared with 24 endemic genera of Gasteropods! but

whereas with the fishes there are in addition 29 non-endemic

genera represented, there are only 11 non-endemic genera of

Gasteropods, Again, among the fishes there are a fou endemic

genera found in the remaining lakes, while this is not the case

with the Gasteropods. The comparison serves to show that while

the actual numbers both of genera and species are less in the

group now under discussion than in that of the fishes, it affords

quite as conspicuous an instance of the peculiar Chavacher: of the

Tanganyika fauna.

The table of distribution already furnished now calls for more

FAUNA OF THE AFRICAN LAKES. HAT

detailed examination. Among the Viviparide, Viviparus itself

is represented in Tanganyika only by two (one doubtful) endemic

forms. In Lake Victoria there is a series of 5 forms, 3 of which

are peculiar to its waters. It is noteworthy that the widely dis-

tributed species V. rubtcundus and V. wnicolor have not been

obtained from Tanganyika. The genera Veothaumaand Lridouaia

are of more interest, since they are entirely confined to Tan-

ganyika. The latter genus has been regarded as exhibiting a

thalassoid facies. Each i is represented by only a single species.

The family Ampullariidz contains representatives “of the well-

known genera <Ampullaria and Lanistes in addition to the

thalassoid genus Leroya from Tanganyika. Only Victoria

Nyanza and Tanganyika possess endemic forms of the two first-

mentioned genera, the remaining species—especially Ampullaria

ovata—having a wider distribution. Lake Victoria will be seen

to have five species of dAmpullaria, Nyasa a similar number of

Lanistes. The latter genus has not been recorded from the

smailer lakes.

A larger number of genera, several of great interest, are asso-

ciated in the family Hydrobiude. Bithynia* alone among the

genera enumerated is found outside the confines of Tanganyika,

but the species on record from the lakes have not been found in

other parts of the continent. Of the thalassoid types, Stanleya

is represented by 3 species and Syrnolopsis by 2. Chytra and

Limnotrochus, as the name of the Jatter indicates, are forms in

which the shell is suggestive of the marine genus T’rochus.

The genera belonging to the family Tiaride (formerly Melaniidze)

form by far the largest assemblage in the list under consideration.

Thirteen genera are repr seerhed. but here again only one of them,

the type-genus Ziara itself (perhaps better known as MJelania) is

of wide distribution, the remainder being thalassoid forms found

only in Tanganyika. The latter lake contains three species of

Tiara, two ot which are endemic; Albert Nyanza contains one

endemic as well as one non-endemie species, while Nyasa contains

a remarkable series of nine different types, seven of which are

endemic. It is interesting to note that Tiara tuberculata, which

has a wide range in Africa and extends into India and the Malay

Archipelago, is the only one of the 133 species enumerated which

occurs in all the lakes included in this survey. There is little

need for comment on the remaining genera in this family. They

contain the greater number of those types of marine aspect which

are so remarkable a feature of the Gasteropod fauna of Tangan-

yika. Certain of the genera are, however, represented by quite

a number of species, notably Hdgaria by 9 species and Giraudia

and Lavigeria each by 8.

Under the family name of Tiphobiide have been associated the

* A species of Bithynia is stated by Moore (187, p. 129) to occur in Lake Kivu.

I believe no specimens have ever reached this country, and in view of the negative

evidence which the visit of the German expedition to the lake affords, I regard the

record as of doubtful yalue and have intentionally omitted it.

548 DR. W. A. CUNNINGTON ON THE

genera Tiphobia and Bathanaliw, which are among the most

striking of the thalassoid fornis from Tanganyika. At present

this remains the only example of an endemic family characteristic

of that lake.

The representatives of the three remaining families are all

well-known fresh-water genera, and do not offer much of interest

on which to comment. Of the Limneide, specimens are on

record only front the three large lakes and Lake Edward, which

is rather surprising as the group has a wide distribution in

Tropical Africa. Limneea natalensis is the most widely distributed

species, being known from Tanganyika, Victoria, and Nyasa, as

wellas from Natal. The families Planorbide and Ancylide are

each represented by the type-genus only. Thirteen species of

Planorbis are enumerated, of which Tanganyika contains the

largest series, viz. 7 forms, 4 of which are endemic. In Nyasa,

it is strange to note, only a single widely distributed species

has been found. The form of commonest distribution in the

lakes is P. swdanicus, which occurs in four of them as well as in

the River Nile. Turning to the Ancylide, it will be seen that

an endemic species of Ancylus is on record both in Tanganyika

and in Lake Victoria. No specimens have been observed in the

other lakes, but they may have been overlooked owing to their

small size.

It is not unreasonable to refer here to the suggestion already

cited in the section dealing with the fishes (of. p. 535) that the

number of different animal types living in a lake is directly

proportional to its size. The figures for the Gasteropoda are

sufficiently large to afford some evidence for this contention, and

they are accordingly given in the following table, which records

the number of families and genera, as well as species :—

Tangan- Victoria : Albert Edward }

yika. Nyauza. ite Nyanza. Nyanza. Lom

Number of Species ......... 84, 28 24, 13 10 2

a COND ssosscanc SB 11 9 6 6 2

. Families ......... 8 7 6 5 6 2

Tt will be seen that Tanganyika, with its admittedly unique

fauna, is in a category by itself, but the remaining lakes, which

are arranged in order of size, form a series with decreasing totals,

and thus support the general truth of the assertion.

The distribution of the Gasteropoda in the lakes may be sum-

marised in a few sentences. ‘Tanganyika, containing 84 species,

of which 76 are endemic, is altogether exceptional and is prin-

cipally characterised by a remarkable series of marine-like or

thalassoid forms. While the latter are not confined to one

FAUNA OF THE AFRICAN LAKES, 549

family, they belong mainly to that of the Tiavide. Tanganyika

is the only lake of those under review which contains represen-

tatives of all the genera enumerated in the table of distribution.

— Victoria Nyanza, with a far smaller number of individual forms

(28), can nevertheless show quite a comprehensive series of the

normal fresh-water types, each of the genera being represented.

The thalassoid genera and species are conspicuously absent.—

Nyasa, with only slightly lower figures (24), does not differ very

markedly from Lake Victoria. The genera Cleopatra and Ancylus

are unrepresented, but the lake is rich in species of ZLanistes and

Tiara, especially the latter, of which 9 forms (7 endemic) are

known.—A!bert Nyanza and Edward Nyanza exhibit a progressive

reduction in the number of species which they contain, but

otherwise possess no outstanding features.— Lake Kivu, finally, is

the extreme case, with a very reduced Gasteropod fauna. Of the

two species which are known from its waters, Ziara tuberculata

is the more widely distributed, it being in fact the only form

enumerated which has been found outside the continent of Africa.

It was in the first degree the Gasteropoda. which constituted

Moore’s argument for a halolimnic (¢.e. relict) fauna living in

Tanganyika side by side with types which are common in tropical

fresh-waters. It is thus necessary to consider whether the balance

of evidence is still in favour of the views regarding these Mollusca

which Moore originally expressed. He held that the peculiar

forms in question were essentially primitive types, and after com-

parisons based on their anatomy he stated :—*‘ It is difficult, or,

I may say, impossible, to view these extraordinary molluscs as

either the forerunners or the derivatives of the fresh-water

molluses which we find in the lakes and rivers all over the world

to-day. . They are, however, readily intelligible if we regard them

as the forerunners of several marine groups, such as the Strom-

bidee, the Naticas, and the early Ciriths, to which I have referred ”

(135, p. 466).

These statements did not go by any means unchallenged, nor

did the astonishing suggestion of a resemblance between some

species and certain Jurassic fossil shells. Hdgar Smith, in his

important paper on the Mollusca of Lake Tanganyika, pointed

out that Moore’s conclusions were drawn from a study of only 9

out of 23 so-called halolimnic genera, adding that it became

‘““mere conjecture ” to suppose a relationship with marine forms

on account of the appearance of the shell (170, p. 78). As

an expert conchologist, Smith dealt unfavourably too with the

supposed resemblances to Jurassic fossils. The matter was

approached from the geologist’s standpoint by Hudleston, who

had made a speciality of Jurassic Mollusca (102). In a com-

prehensive paper on the origin of the halolimnic fauna of

Tanganyika, he failed to find evidence of a satisfactory character

in favour of Moore’s views. In an appendix, Hudleston also

dealt seriatim with the comparisons between living and Jurassic

Gasteropods, only to reject likewise Moore’s conclusions. In a

550 DR. W. A. CUNNINGTON ON THE

discussion on this topic at the British Association Meeting in

1906 (188), Moore expressed less extreme views, while Pelseneer,

speaking with a wider knowledge of the anatomical characters

of these molluscs, asserted positively that there was no special

resemblance between them and marine types and that they were

no more primitive than other well-known fresh-water genera

(144). Pelseneer added that the halolimnic Gasteropods were all

to be regarded as belonging to the Melaniide (Tiaride) or families

nearly akin, in which case they belong to an essentially fresh-

water group. If this be indeed a fact, Moore’s whole hypothesis,

as far as it rested on these molluscan forms, falls to the ground.

It no longer becomes necessary to speculate as to how and when

Tanganyika received animal types from an ancient sea, it is suffi-

cient to invoke prolonged isolation to account for endemic genera

and species—and this is precisely what has been done in other

groups.

It is true that no satisfactory explanation of the marine aspect

of these Gasteropod shells is forthcoming, although certain

guesses may he hazarded. Without attributing it to a direct

community of descent with certain salt-water types, it might be

regarded as simply due to convergence. It might be urged that

the resemblance is purely accidental, or that it results from the

quasi-oceanic conditions which prevail in Tanganyika, It might

be due to the magnesium salts in the water, or, perhaps, to the

greater degree of salinity which is believed to have formerly

existed. Beyond such suggestions it is impossible to go, but it

must not be overlooked that a number of fresh-water shells from

other parts of the world exhibit the same characteristically marine

aspect, though there may be no other case with so extensive a

series of forms. Indeed, Bourguignat and others have not hesi-

tated to claim a thalassoid aspect for certain Melaniide from

Nyasa and even from the River Congo (cf. 48, p.40: 87, p.564).

A discussion as to the source from which the striking group of

thalassoid Gasteropods has been derived might follow here, but is

deferred for consideration at a later stage. It may, however, be

ointed out that forms possibly akin to those in Tanganyika

which have aroused so much interest, formerly occurred in the

Balkan Peninsula (cf. Brusina, 58 : 59), from which neighbourhood

distribution through the valley of the Jordan and the Great Rift

Valley may have been effected.

Lamellibranchiata.

The Lamellibranchs of the African lakes fall far short of the

Gasteropods in point of general interest, since there do not exist

in Tanganyika or elsewhere any of those types which have been

deseribed as thalassoid. The species belong, in consequence, with

but few exceptions, to well-known fresh-water genera such as

Corbicula, Unio, and Mutela. ‘The list of species admitted reaches

a much smaller total (53) than in the case of the Gasterepoda,

FAUNA OF THE AFRICAN LAKES. il

and it is quite surprising to find that in this instance ‘langanyika

does not take the first place with the largest number of different

forms. This position is held by Victoria Nyanza with 18 species,

but Tanganyika follows very closely with 17, and maintains its

reputation to some extent by exhibiting the highest number of

endemic species and even certain endemic genera. In the

following table no column is provided for Kivu, since there is

only a doubtful record of Lamellibranchs from that lake *.

Table Pee Distribution go Lamellibranchiata *.

Tangan- Victoria Albert Edward Other parts of

Name of Species. yika. Nyanza. Nyasa Nyanza. Nyanza. the world.

Family CyYRENIDZ.

Corbicula astartina ......... ee s 12 Ae af Zambezi.

i cunningtoni...... vs E

35 TRAGHENE | cocarredguue 12 Je) ie Pp E. Africa, Nile.

Family SPHEZRIIDZ.

Spherium nyanze ......... Bes lz es 12 18 FE. Africa.

3 stuhlmanni...... ee E

VichOTle ......-.. Ae 1)

Eupera parasitica ............ ae P et 5c it Nile, Abyssinia.

Family Unronip.

Unio acuminatus ............ ae a ae 1

op ESCAADDEKOUS coocsbnoe scr tn ae on iP sm Nile.

Pe DAK ELI y Sxauece tors eis ie Si nie E

Mme bore lies. .s teas ceca. sep ate E

re) RUSROTN Coc oee ne ienbaccoon H

pee carlltadtieseeeste ees 50 P fe Nile.

> calathus ae K

net CULO Pgs Se aA ten Berohes ar E

EPSP OERTALGIN Vee teceeenesr = E

epsrandidiertyscs see eecea- tee a0)

Pee lianitbecceuniee ase tes ao i

bp JO utapecee nb cedrceeneee E,

>, lypsiprymuus ......... K

” kirki sStendoopceo seo oncaog ig Nyasaland.

5, lechaptoisi E

>, Hederi oe de K. Africa.

» lourdeli E

+, monceti E

5» mossambicensis ...... aes 12 Zambezi, E. Africa.

ry TOUCOMO okdseenosoe Se E

Py ueesimus) ee aoe Eee iM Ene et E

P mDyASsacnsisnnes) a sth IY ‘if te Angoni Land.

RP ROS GLANS yee centn ates. 1D

a Ue anise sen tees she EH

S stuhlmannii a Be we Ree i

* Speaking of the fauna of Kivu, Moore refers to “one or two species of fresh-

water bivalves, closely allied to the Unios found generally in the African lakes”

(137, p. 129). Against this assertion is to be set the statement of the naturalist

at the head of the German Central Africa Expedition, “ Ebenso scheinen lebende

Lamellibranchier zu fehlen” (156, p. xiv).

+ The principal sources which may be consulted for information on the Lamelli-

branchs are Smith (170) and von Martens (116).

5d2 DR. W. A. CUNNINGTON ON THE

Tangan- Victoria Albert Edward Other parts of

Name of Species yika. Nyanza. Nyasa. Nyanza. Nyanza. the world.

Unio tanganyicensis ......... E .

,, teretiusculus ......... ote Ne is 12 se R. Nile.

Fy UINTONEOVNL eclancnop asabee E

Burtonia bourguignati...... Gen. H

5 tanganyicensis ... K

Brazzwea anceyl .............. Gen. EH

Family MureLipa.

WKH, AIEWEW. sasdsc oor oodeen ene AM 5H a0)

» bourguignati .. ... ee IP a 83 ms Ki. Africa.

Sy. GEROUNER), | ocongdsassandoo Pp ae ee a se N. Africa.

Pe enilobicases torre rere: Hee ue x. Pp IP Heypt.

», soleniformis ......... 10)

» subdiaphana......... vols E

Pleiodon spekei ...........-..- y

Spabthakanceyiueessceres-seer sss sais E

ye ulblkoy Gl nlmneeeesemaseoanas ems staan 12 aot ae as K. Africa.

By ab auckattreninec steer a-tecr one ein E

» mhyassaémsis ......... oe an E

gota ez ai acernceerr cote ay E

Moncetia anceyi_ ............ Gen. E

Family AXTHERIID#.

AMtheria elliptica ...........- 12 12 ya - abe Tropical Africa.

BS eMMITES cocgeaacoasoor 4 5 3 4: 4,

TH exeniitey “Gonobaseaceesan 3H, 5P. Uf 12s AP, 4P. 4 P.

(8)

53 species .............. J4H,3P. 12H 6P, Tier, 2H 6P. 2H 3P.

Gi. SCiieds) a 1), oC)

The table of distribution in this case displays no outstanding

features such as are to be seen in other groups. The not incon-

siderable total of 53 species is reached, but the details of

distribution have, in most cases, little significance. Firstly, it

may be noted that the vast majority of the forms (there are only

4 exceptions) are on record merely from a single lake ; secondly,

it is interesting to find that no species in the whole list occurs

outside the African continent.

Victoria Nyanza, as already stated, exhibits the largest number

of types, viz. 18, but is closely followed by Tanganyika with 7

and by Nyassa with 13. Lakes Albert and Edward follow in the

usual order with 8 and 5 species respectively. A fact which

emerges from an examinaticn of these figures, is that the two

lakes with the highest totals only contain about one-third of the

number of species enumerated (Lake Victoria 35°9 per cent.,

Tanganyika 32 per cent.). Contrastec with this, the Gasteropoda

of Tanganyika constitute over 63 per cent. of the total number

of Gasteropods enumerated, while in other groups the corres-

ponding figures for Tanganyika reach 80 per cent. and even

90 per cent. (Branchiuia).

FAUNA OF THE AFRICAN LAKES. 553

Tanganyika with 14 endemics out of 17 leads the way as far

as endemic Lamellibranchs are concerned (82 per cent.), Lake

Victoria coming near with 12 out of 18 (66 per cent.), while

Nyasa has 7 out of 13 (54 per cent.). Even the smaller lakes

have each two endemic species. It will be observed that the

proportion of peculiar forms is very high in this group also;

indeed, in some instances it is higher than in the case of the

Gasteropods. Hndemic genera are known only from Tanganyika,

three being retained in the list out of a larger number distin-

guished by Bourguignat. The genus Moncetia of Bourguignat,

inserted in the table as endemic, though not definitely rejected,

is nevertheless considered by Smith as only doubtfully separable

from Spatha (170, p. 101).

A brief survey of the list of species will suftice, Corbicula

radiata, a representative of the Cyrenide, is the only form which

is on record from all the lakes concerned. It is a widely distri-

buted African type, being known from other parts of East Africa

and from the valley of the Nile——The family Spheeriide is

represented by the genera Spherium and Hupera. It is some-

what strange that the four species enumerated are all found in

Lake Victoria, but not in Tanganyika or Nyasa. Sphervwm

nyanze is known from Victoria, Albert and Kdward Nyanaas, as

well as from other parts of the continent, but it is associated in

Victoria with two additional species which are peculiar to that

lake. Hupera parasitica, which is a Nilotic and N. African form,

is at present only recorded from Victoria Nyanza.—In the family

Unionide there are associated with the extensive genus Unio

itself only the two closely related genera Burtonia and Brazzea,

which are confined to Tanganyika. No fewer than 29 different

species of Unio are enumerated, not one of which is on record

from more than a single lake. ‘[anganyika exhibits 8 and Lake

Victoria 7 endemic species. Of 7 types in Nyasa 3 are endemic ;

of 5in Lake Albert 2 are endemic. Lake Edward, lastly,

contains 2 forms, both of which are endemic.—In addition to

three genera of wider distribution, the Mutelide contains the

doubtful genus Moncetia, to which reference has already been

made—a genus described as peculiar to Tanganyika. J/utela

nilotica is the only species in the family recorded from more than

one lake, it being found in both Albert and Edward Nyanza as

well as in Egypt. Each of the bigger lakes exhibits a single

endemic form of Jutela. he genus Pleiodon is only represented

by P. spekei, confined to Tanganyika—it is one of the species

named by Woodward from Speke’s original collection. Spatha is

unrepresented in Tanganyika and the smaller lakes, but a series

of three endemic forms is known in Nyasa.—Lastly, Wtheria

elliptica, sole representative of the Aitheriide, occurs in 'Tangan-

yika and Victoria Nyanza, but has not been recorded from the

other lakes, although it is widely distributed in Tropical Africa.

While a considerably smaller number of forms is involved here

than was the case with the Gasteropods, there is yet an indication

Proc. Zoou. Soc.—1920, No. XXX VII. 30

554 DR. W. A. CUNNINGTON ON THE

that fewer species occur in the smaller than in the larger lakes.

With the exception of Tanganyika, the lakes are annie in

order of size, and the figures for the species

Tanganyika 17, Victoria 18, Nyasa 13, Albert 8, oa aware 5.

Regarding Tanganyika as a lake apart, the remaining figures fit

in satisfactorily with this suggestion.

The features which are characteristic of the different lakes may

be summed up in a few words. ‘Tanganyika alone contains

endemic Lamellibranch genera. Spheriuwm, Hupera, and Spatha

are unrepresented, but a number of endemic species of Unio are

known.— Victoria Nyanza, with the largest total of species,

displays, notwithstanding, little of interest. Most of the genera

are represented, except those peculiar to Tanganyika, and a series

of forms belonging to Spheriwm and Unio are to be observed.—

Of Nyasa there is likewise little to record. The lake only

contains species from the four well-known genera Corbicula,

Unio, Mutela, and Spatha, 3 endemic types of the last-mentioned

being an outstanding feature.—Both Albert and Edward Nyanza

are similar in type to Nyasa, but contain a representative of

Spherium and not Spatha.

From this survey of the Lamellibranchs, the following points

emerge :—No thalassoid types occur in Tanganyika, but that lake

contains a number of endemic species, as well as three endemic

genera. All the lakes show a high percentage of endemic forms,

but in most cases these are merely species belonging to widely

distributed genera. While Tanganyika does not exhibit in this

instance so remarkable a series of unique genera and species as

in the case of the Gasteropods and other groups, it retains never-

theless a distinctness from the remaining lakes quite in keeping

with its general character.

MaAcruRA.

The only Macrurous Crustacea which appear to occur in the

big African lakes are the prawns, these being quite common

types in the fresh-waters of the tropies. An examination of the

forms now known to exist in the lakes of Africa reveals, how-

ever, many points of interest. In the first place, no fewer than

twelve species have been discovered in Tanganyika, of which ten

were obtained for the first time by the Third Tanganyika

Expedition. ‘This is in itself a large number of different species

to be found within the limits of a single lake, but the interest is

greatly increased when it is realised that all these forms occur

in Tanganyika alone. This is the only well represented group in

which such is the case. Again, with the exception of a single

species of the well-known fresh-water genus Palemon, all the

genera are equally to be regarded as endemic.

FAUNA OF THE AFRICAN LAKES. 555

Table of Distribution of Macrura *.

Tangan- Victoria Albert Edward Other parts of

es Baas

Waiatesot = pebses? yika. a anza. Nyass. Ne Nyanza. the world.

Family PALHMONIDZ.

Paleemon moorei .............:-

Family Ary1pz.

Caridina nilotica

var. eracilipes Ripe re hates ra 2! ip P * P ; Asia, Australia.

Limnocaridina retiarius ...... Gen. EH

ss parvula...... a0)

ie tanganyike ... iH

a SIMMS | tees ss K

‘ Tatipesmcgess cs. i

rs socius ......... E

spinipes ...... 0)

Limnocaridella alberti ......... 4¥ ui}e ce Gen, E

Caridella cunningtoni ......... Gen, E

sy minuta epee E

Atyella brevirostris ............ Gen. E

>) M longinostrist.:. 222-24. 0)

6 genera 3H, 1P. 1) TP. ide 1K, 1P. WP

TM SHNGOS coon ssescessessesseas | pal qI0e id TED eee TUB e i} 12

The adjoining table shows at a glance the most striking

feature of distribution, namely that while Caridina nilotica var.

gracilipes = occurs in nearly all the big lakes of Africa, it is not

found in Tanganyika, but is replaced there by 12 endemic species

belonging for the most part to endemic genera. Lake Albert

-also contains an interesting endemic form (obtained by the

German Central Africa Expedition and described by Lenz

(109, p. 132) and Bouvier (52, p. 575)) in addition to the above-

mentioned widely distributed Caridina, while Kivu is the only

lake of those at present under review in which prawns have not

hitherto been observed.

‘The first species of prawn ever taken in the African lakes was

+ A detailed account of the Macrura of the Third Tanganyika Expedition is given

by Calman (61), and forms the principal source of information on this group.

* The species of prawn collected by Schubotz at Kassenje on Lake Albert was

described by Lenz (109, p. 180) as Caridina longirostris Milne-Edwards. It is,

however, commonly agreed that longirostris is identical with the earlier nilotica,

so that, apart from the varietal name, this is the same form that occurs so widely

distributed in Africa. Since the type described as Caridina nilotica var. gracilipes

is certainly rather variable, it seems probable that the Lake Albert specimens are not

sufficiently distinct to be recorded under a different name, and this is the view which

I have taken in compiling the table of distribution above. For a discussion of these

questions of synonymy consult the paper of Calman (61, p. 189 e¢ seq.) and the

subsequent paper of de Man (110), which arrives at somewhat different conclusions.

+ I adopt the varietal identifications of Hilgendorf (100), Calman (61), and Lenz

(109), bat it is only right to point out that de Man (110) takes a different view.

He appears to consider that the form occurring in Lake Victoria is to be referred to

the typical Caridina nilotica itself, and the form trom Nyasa to his newly estab-

lished variety natalensis, The type Caridina nilotica var. gracilipes he records

only from the islands of Celebes and Salayer in the Malay Archipelago.

37%

556 DR. W. A. CUNNINGTON ON THE

obtained in Victoria Nyanza by Stuhlmann in 1890 (100, p. 36).

It was re-taken in the lake by Neumann, and more recently by

Alluaud, by myself, and by Degen. From Nyasa a species of

prawn was brought for the first time by my expedition. These

species prove to be one and the same form—the Caridina nilotica

var. gracilipes, to which reference has just been made. During

the German Central Africa Expedition of 1907-08 this prawn

was collected by Schubotz in both Lakes Albert * and Edward

(109, p. 130). It is thus the only common species in the African

lakes, while it has in fact a yet wider distribution, ranging in

Africa from Natal to the Nile and extending into Asia and

Australia. The eastward range of this type has indeed an

additional interest on account of the well-marked resemblance

noticeable in other groups between the Hast African and Indian

faunas.

Only the first of the twelve species enumerated from Tangan-

yika can be compared at all closely with forms which are known

from other parts of the world, and it has no very pronounced

affinities. ‘The remaining eleven species belong to the group of

the Atyide, and are not so nearly connected with types hitherto

known. Moreover, in common with Limnocaridella alberti, they

differ from all the other species of the family in having a smaller

number of branchie +, which is a feature undoubtedly due to

specialisation.

To sum up then, there occurs in most of the great lakes of

Africa only a single species of prawn having a very wide

distribution. In Lake Albert this is associated with an endemic

form, and in Tanganyika it 1s replaced by twelve other endemic

forms, the majority of which are among the most highly special-

ised members of the family to which they belong. Under thé

circumstances, it is impossible to resist the suggestion that there

is exhibited here something very similar to what was seen in the

case of the fishes, and particularly the Cichlide. If the extra-

ordinary variety of form, and high degree of specialisation, which

is characteristic of the Cichlids of Tanganyika, may have been

due to prolonged isolation and comparative freedom from com-

petition, it at least seems not improbable that the remarkable

Macruran fauna of the lake owes its origin to the same cause {.

BRACHYURA.

There are five different species of crabs now known from

Tanganyika, of which one has been left unnamed for the present.

In a manner closely corresponding with the case of the prawns,

* See footnote above.

+ A. reduced branchial formula is also characteristic of the remarkable West

Indian form for which Bouvier has established the genus Micratya (51, p. 181),

formerly Calmania (50, p. 334). Bouvier regards this as allied to the Tanganyikan

genera, but this view is not accepted by Calinan (62, p. 796).

+ An important paper dealing with the origin of the peculiar prawn fauna of

Tanganyika was communicated by Bouvier to the International Zoological Congress

at Monaco (52). Consult also Bouvier’s paper on the classification of the Atyidee

(51).

FAUNA OF THE AFRICAN LAKES. Otay

while two species belong to a widely distributed and common

fresh-water genus (Potamonautes), the remaining three, though

members of the typical fresh-water group (Potamonidee), con-

stitute a remarkable genus, which occurs only in Tanganyika.

All the forms from the other lakes under review belong to well-

known genera of the same family. No Brachyarans have as yet

been reported from Albert Nyanza.

Table of Distribution of Brachyura*.

= ——.- a

ers Tangan- Victoria , Edward j,,. Other ;

Name of Species. yika. Nvanza) Nyasa Ri pante Kivu. fs ite:

Family PoTAMONID#.

Potamon (Potamonautes)

inflatus ... + se IP ha ... Natal.

s », orbitospinus iis Se E

. », platynotus.. iE

- eeSD ete sects: =e R

rs oan SIDS 5 Mgreee aoe Py

Potamon (Geothelphusa) _

berardi aes ct eee oes ae 1p Egypt, Abyssinia.

55 5 EET cncicoe 200 Je - P PK. Africa, Abyssinia.

Potamon (Acanthothelphusa)

= TUPD ATONE Leos goonbanseonce Aon! zee P a od ... Egypt, Abyssinia.

Platythelphusa armata......... Gen. E

re maculata... K

i conculcata ... E

4 genera and subgenera... 1H, 1P. a Nee ible SEs iL Te:

1, GOCE erp ccotepenesd aan con, MUON whN ze N22, INDE ey DP Dike

The table of distribution makes it clear that while each of the

lakes in the list exhibits one or more representatives of the very

well-known genus Potamon with its sub-genera, these types are

associated in Tanganyika with three species of an interesting

endemic genus. Both Nyasa and Tanganyika possess species of

Potamonautes which are peculiar to themselves, but it is in

Tanganyika alone among the big lakes that an endemic genus is

found. All the species enumerated appear to be confined to the

continent of Africa.

Apart from the case of Tanganyika, it will be noted that (with

the exception of Potamon (Potamonautes) orbitospinus from

Nyasa), the forms from the different lakes are by no means con-

fined to. them, and indeed are often of wide distribution. This

whole series of types—types such as are known from all the

tropical fresh-waters of the Old World—may be considered as

the normal African group, and calls for little further remark.

In the paper already cited (70, p. 263) the present writer

expressed a conviction that the number of African species has

been unduly multiplied, and the unnamed forms included in the

+ For further details concerning most of these forms, consult the Report on the

Brachyurous Crustacea of the Third Tanganyika Expedition (70).

558 DR. W. A. CUNNINGTON ON THE

table are to be regarded as specimens which, in the existing state

of our knowledge, it is impossible to identify. It is nevertheless

clear that the precise distribution in the lakes of these repre-

sentatives of the subgenera of Potamon is of little, if any,

significance.

In this connection it should perhaps be explained that the

evounds of identification of the river-crabs are unsatisfactory on

the whole, external characters of a compavatively trivial and

fluctuating nature forming the basis of distinction. Systematists

are thus on less sure ground than in the case, for instance, of the

prawns, where a matter like the branchial formula affords more

satisfactory evidence of affinity.

Tanganyika, with no less than three species of the endemic

genus Platythelphusa, is evidently a case sui generis. There are

certain features in the anatomy of this unique genus which

suggest that it-is of a somewhat primitive and unmodified

character, but at the same time nothing to indicate that it is

more definitely marine than the other members of the family to

which it belongs. Further, it is only to be regarded as unspecial-

ised in comparison with allied forms which have adopted a semi-

terrestrial mode of life, so that it affords in reality no support

for Moore’s view that Tanganyika is the altered remains of an

ancient sea. The genus with its three distinct species is rather

to be looked upon as one more example of variation and

divergence, brought about, in all probability, by prolonged

isolation. Finally, it may be noted that this lake, with five

species, contains a larger number of different crabs than any of

the others under consideration.

Those groups of smaller Crustacea which are often associated

under the heading Entomostraca, are (with the exception of the

marine Cirripedia) well represented in the African lakes. In-

formation concerning them is, however, for the most part, of

fairly recent date, since such organisms would be overlooked

by any but trained biologists, and the use of the tow-net in

Central Africa has even yet yielded results which are but

fragmentary and incomplete. All the earlier records are due

to the zeal and energy of Stuhlmann, and concern in particular

Lakes Victoria and Edward. In 1898-1900, extensive collections

were made in Nyasa and its neighbourhood by Filleborn, and

this material, together with a supply from Victoria Nyanza

collected by Bor gert, forms the basis of a comprehensive treatise

by Daday (76), in which the whole of the microfauna is dealt

with. The collections made by the writer during the Third

‘Tanganyika Expedition have afforded information for the first

time concerning the Entomostraca of Tanganyika, as well as

providing additional records for Nyasa and Lake Victoria.

Finally, the work of Schubotz during the German Central Africa

Expedition has furnished further particulars relating to Lakes

Kivu, Edward, and Albert.

It may here be pointed out that these lower Crustacea, with

FAUNA OF THE AFRICAN LAKES. 559

the exceptional opportunities they are known to possess for

obtaining world-wide distribution, cannot afford evidence of the

same value as the higher forms. When—as is particularly

the case among the Cladocera—even specific forms of cosmo-

politan range reappear in the great lakes, it becomes clear that

the precise geographical distribution of such species is a matter

of no great significance. At the same time, the possibility is not

excluded that the lakes may differ in their suitability to harbour

certain types, while it is highly probable that new types can and

will develope in some cases, constituting species or even genera

of an endemic nature. Thus a study of the distribution of these

forms in the lakes of Central Africa is nevertheless not devoid of

interest, and may indeed furnish testimony of some importance.

HUCOPEPODA.

The Hucopepoda so far observed in the lakes with which this

paper is concerned reach the not inconsiderable total of 54

species. Of these, more than half (31) have been found in

‘Tanganyika, to which lake a large proportion exclusively belong.

There is every reason to believe that further investigation will

bring to light additional forms, particularly from Victoria

Nyanza and the smaller lakes, which, in this respect, have

received less attention than Tanganyika and Nyasa.

Table of Distribution of Hucopepoda *.

Other parts of

on Tangan- Victoria ... Albert Edward 4,-

Name/of Species. yika. Nyanza. See Nyanza. Nyanza. Eve the world.

Family CentTROPAGID®.

Diaptomus galeboides{ ... i Pp “eb Je

* TONetHNE! coneesoee Be: oe K

A africanus ...... ae: a 12 oe ae =. | Ey Adrica:

53 stuhlmanni ... Bic 0)

3 simplex ......... K

FF cunningtoni ... he wits E

$3 kreepelini ...... P EK. Africa.

Family Harpacticip®.

Canthocamptus schréder1 Ee E

Attheyella decorata ......... IPS Asia, S. America.

- erandidieri ...... P§ S. America, New Guinea.

Dactylopus jugurtha ...... be 12 Asia, New Guinea.

Schizopera inopinata ...... KE

55 Validione esc K

3 consimilis ...... iz 12

3 WNW atareeee aD)

55 minuticornis ... KE

+ Detailed information concerning a large proportion of these species is given by

Sars (151). The other sources of most importance are Daday (76, p. 106) and

Mrazek (139). For the Lernzid consult Cunnington (73).

t I follow Sars (151, p. 34) in assuming that this form, which is one of the

commonest and most characteristic species of Lake Victoria, is not identical with

the Egyptian D. galebi Barrois, as supposed by Mraézek and Daday.

§ Not actually recorded from the lake itself, but from within its drainage area,

560 DR. W. A. CUNNINGTON ON THE

Bae Tangan- Victoria , Albert Edward y,.. Other parts of

HNJaHEH OM epee yika. Nyanza. ~Y*S** Nyanza. Nyanza. oN mi the world.

Schizopera spinulosa ...... 13)

ys fimbriata ..... wat a ree a0)

are SCallavis) Seaeeuse E

Tlyophilus perplexus......... KH

Family CycLopip™.

Cyclops aspericornis ......... ie los PS eA ie ... Hast Indies.

> Jeuckarti -.. P P P 12 je P Cosmopolitan.

>» emini Beas Sete ie 1 P a ae .. KE. Africa.

pe eneglectuchcmemerre: P 12 Pp P IP P Senegal, Sumatra.

pnbenellus: vtaeenremsce E

5 INLCEES cocoscondaonoc as P ee A of ... Cosmopolitan.

np ETSTMBANBIS so.90> oon i

SAV ATICATIS greyeeepete 12 12 P ae .... Hurope, New Zealand.

VE Mm bLCOlON era wacern ce se IPS a ue ... E. Africa, Europe.

sal) MERIOUUUSI eae Mee aan: E

» cunningtonl ...... H

ee pachycomluss se 10)

f) serrulatus-p .-.- at PS P§ i P ... Cosmopolitan.

» semiserratus ...... K

by IBSNEOTENPERO). oo gascve 13)

fy WDAOPBIETS 350 coo nen 000 bee PPM RR A . ... Cosmopolitan.

ee ancushusmeees see JP Pp

» stuhlmanni......... oor

2 ranispimus\eceeeeee EK

5p ESHIIONGI@S cca sooo 000 1 12 Ie

Ee UACA UNIS eee E§

>» Ciliatus ae i

eo GUTUSE se akraccececree Be i

fy DEREMNOS codooaobocos 1 Ip Be as .. EH. Africa, Europe.

» Phaleratus ......... tke PS§ P§ er si ... Cosmopolitan.

> compactus ......... P§ : Central Africa,

Pe olicanthnusyerseerre E

Family ErGasinipH.

Ergasilus kandti ............ Er ih are 0)

e SDE ee eae bye 12 IP

Ergasiloides megacheir... iH

5 macrodactylus K

% brevimanus ... 12 1 P

Family LERN#ZID2.

Lernzocera diceracephala .. x’

Ps haplocephala ... P eb bi ae a ... Nile, Congo.

10 genera .....0ee cee 6 P. B12, Hil 3 P. le, 12,

54) species .........006-. “7, 22E,9P. 3E,10P. 46,19P: 1H,6P. 3 1 uke

; (31) (13) (23) (7)

§ Not actually recorded from the lake itself, but from within its drainage area.

* As explained by Sars (151, p. 51 & 74, p. 82), this species has been wrongly

identified with C. oithonoides G. O. Sars or with C. hyalinus Rehberg by other writers

on African Copepoda. I adopt the new name introduced by Sars.

+ A number of the forms which succeed this in the list belong to the group for

which C. serrulatus Fischer is the type. While the species C. serrulatus is recorded

by Mrazek (189, p.3) for Victoria Nyanza, by Daday (76, p. 108) for Nyasa, and

by van Douwe (80. p. 492) for Edward Nyanza, it is not unlikely that the specimens

in question might rather belong to one of the species in the “‘ serrwlatus” eroup as

described from the lakes by Sars, than to C. serrulatus, s. str. At present it is

impossible to settle this point, so I take the course least open to objection and insert

the records of Mrdzek, Daday, and yan Douwe as they stand,

FAUNA OF THE AFRICAN LAKES. 561

An examination of this long list of forms reveals in the first

place the large proportion of the recorded species which has been

observed in Tanganyika. ‘The latter lake contains 31 out of a

total of 54 different forms, Nyasa contains 23, Victoria Nyanza

13, and the remaining lakes still smaller numbers. The contrast

between the lakes is greater than is disclosed by these figures, for

out of 31 species known from Tanganyika, 22 are peculiar to that

lake, and in comparison only 4 out of 23 for Nyasa and 3 out

of 13, for Lake Victoria. In the three smaller lakes there is only

a single recorded species which is endemic, and that occurs in

Albert Nyanza. Expressed in percentage form, th’s means that

nearly 71 per cent. of the Copepods of Tanganyika are found in

that lake alone, while of the species found in Nyasa only some

17 per cent. are peculiar to the lake, in Victoria 23 per cent., and

in Albert 14 per cent. Excluding Tanganyika, it will be observed

that only 8 of the forms recorded are endemic, so that with 22

endemic species, Tanganyika has nearly three times as many

endemic Eucopepoda as the other lakes can muster between them.

Turning now to the distribution of families and genera in the

lakes, the genus Diaptomus, sole representative of the Centro-

pagidee, calls for little comment. A single species of this genus

occurs in Tanganyika, to which it is peculiar: it appears to be

a most characteristic plankton form in that lake. Lakes Victoria

and Nyasa each contain endemic species as well as types of wider

distribution, so that the three big lakes may each be said to have

their own particular forms. None of the species of Diaptomus

in this list have been observed outside the continent of Africa.

The representatives of the Harpacticide are of far greater

interest. An endemic species of Canthocamptus has been re-

corded from Victoria Nyanza, and widely distributed species of

Attheyella and Dactylopus from Nyasa. It is on the genera

Schizopera and Ilyophilus, at one time regarded by Sars as strictly

marine, that interest really centres. The genus Schizopera is

represented in the collections made by the writer by no less than

8 species, of which 6 are peculiar to Tanganyika and one to

Nyasa, while one occurs in both Lakes Tanganyika and Victoria.

The genus was established by Sars for the reception of a species

found in a brackish-water lagoon on the Chatham Islands, while

subsequently species have been obtained from the Caspian and

from Egypt. It is obvious that such a genus can hardly be

regarded as strictly marine. ‘Three questions suggest themselves

in this connection. How does the genus come to be represented

in the heart of Africa; how is it that brackish-water types are

living there in perfectly fresh-water ; and how is it that so many

different species are recognisable in Tanganyika ? Sars points out

(151, p. 69) that the most obvious explanation would be to regard

these as genuine “relict” forms, in which case the Tanganyikan

species would be important memhers of Moore’s so-called “ halo-

limnic” series. He rejects on general grounds the view advanced

by Moore, and indicates transport by the aid of migratory

aquatic birds -as sufficient explanation for this remarkable

562 DR. W. A. CUNNINGTON ON THE

discontinuous distribution. He asserts the probability that

forms transported from brackish water would not be unable to

survive in fresh water, and finally maintains that the several

species occurring in Tanganyika have been produced by diver-

gence during a period of isolation—a view entirely in accord with

that already expressed in the case of other animal groups.—The

genus //lyophilus is represented in Tanganyika by a single species

which is endemic. The type species of this genus was described

from the Baltic and has also been obtained near Christiania, so

that the remarks concerning transport and establishment in

fresh water are as applicable here as in the case of Schizopera.

No fewer than 27 species, all belonging to the genus Cyclops, are

enumerated under the family Cyclopide. It will be noted that

Tanganyika again leads the way with 17 species, while Nyasa

contains 13 and Lake Victoria 8: moreover, 11 of the Tangan-

yikan species are peculiar to that lake, while the other two lakes

each contain only a single endemic form. ‘The three smaller

lakes possess fewer representatives, viz.:—Lake Albert 5, Lake

Edward 3, and Lake Kivu only 2. All the latter are of fairly

wide—some of cosmopolitan—distribution.

The family Ergasilidee, which may be regarded as intermediate

between the free-living and the highly degenerate parasitic

Copepods, is represented by 5 species belonging to two genera.

An endemic form of the genus Hrgasilus has been recorded from

Albert Nyanza, and the genus has also been recognised both in

Nyasa and Victoria. In Tanganyika it would seem to be

replaced by the genus Hrgasiloides, described from material

obtained by my expedition. ‘Two endemic species are on record

from Tanganyika, while a third is found in that lake and also

in Nyasa.

The Lerneide with two forms of Lerncocera concludes the

list. The two species were obtained by the Third Tanganyika

Expedition from fishes in that lake, and while one proves to be

endemic, the second was found to be identical with specimens in

the British Museum collection taken on Nile fish.

Reference may again be made here to the statement that the

number of different animal forms existing in the lakes varies in

proportion to the size of the latter (cf. pp. 535, 548). It is only

when many forms are enumerated that such a comparison is

justified, but in this case it may fairly be undertaken. Apart

from Tanganyika, the lakes are arranged in the table in descend-

ing order as regards size, and the total figures are as follows :—

Tanganyika 31 (species), Victoria 13, Nyasa 23, Albert 7,

Edward 3, and Kivu 2. It is very clear that Tanganyika is

quite unique, but apart from that lake, beginning with Victoria

Nyanza, the Copepods exhibit a regularly decreasing total with

the exception of Nyasa, where the higher figure is probably due

to the more thorough exploration of its water's.

The distribution of Copepoda in the individual lakes may be

summarised as follows :—Tanganyika, with 31 species, 22 of which

are endemic, is clearly in a category by itself. Containing a

FAUNA OF THE AFRICAN LAKES. 563

single species of Diaptomus which is peculiar to it, the lake

is chiefly characterised by a number of remarkable forms belong-

ing to the genera Schizopera and Ilyophilus, nearly the whole of

which are not known elsewhere. Even of the cosmopolitan genus

Cyclops, Tanganyika exhibits 11 endemic types, as well as others

of wider distribution, while of the three species of Hrgasiloides,

two are found here alone. ‘Two forms of the parasitic Lerneocera

oceur in the lake, one of which is likewise endemic.

The Copepoda of Lake Victoria are referred not only to fewer

types, but form a much less interesting assembly. They comprise

a single endemic species in each of the genera Diaptomus, Can-

thocamptus, and Cyclops, but otherwise are forms of wider, often

of world-wide distribution.

A larger number of species is known from Nyasa, but out of

23, only 4 are endemic. Of the latter, two are species of

Diaptomus, while there is a single endemic form of the striking

genus Schizopera and one of Cyclops. Most of the remaining

species are widely distributed, a considerable proportion being

well-known forms of Cyclops. Nyasa contains a species of

Ergasilus as well as Lrgasiloides brevimanus which is otherwise

confined to Tanganyika. There is no reason to suppose that

Nyasa will be found to contain so many more Copepoda than

Victoria Nyanza when the latter lake has been equally well

investigated.

Seven species have been observed in Lake Albert, of which

merely a single form (Zrgasilus kandti) is endemic. Apart from

this and from a species of Diaptomus found also in Lake Victoria,

the lake contains only representatives of the genus Cyclops, which

are of fairly extended, and in some cases of the widest possible

range.

Lake Edward with 3 species, and Kivu with 2, come last in

the series, and it should be noted that not only do they contain

very few types of Copepoda, but that these types have little

significance, being in each case widely distributed species of

Cyclops.

It is thus apparent that with increase in size the lakes exhibit

an increase in the number of forms which they contain; that

only in the larger lakes do endemic species appear; and, finally,

that Tanganyika possesses a large number of unique types which

may well have been produced during a period of prolonged

isolation.

BRANCHIURA.

This small group of animals, which are parasitic for the most

part on fish, was for long but poorly represented in the collections

from the great lakes of Africa. The collections made by Stuhl-

mann in Victoria Nyanza (184, p. 154) and Edward Nyanza

(187, p. 47 & 188, p. 37) and by Fiilleborn in Nyasa (187, p. 47

& 188, pp. 37 & 44), resulted in a total of three species belonging

to as many genera. To these may now be added no fewer than

seven new species of Argulws which were obtained for the first

time by the Third Tanganyika Expendition (72), while there are

(57 ]

564 DR. W. A. CUNNINGTON ON THE

further records of distribution now available which make our

knowledge more complete. It is safe to predict that future

investigation will add to the localities in Africa from which these

parasitic forms are known, even though it may not materially

increase the number of recognised species.

Table of Distribution of Branchiura *.

Tangan- Victoria Albert Edward Other parts of

Name of Species. Nyasa.

yika. Nyanza. Nyanza. Nyanza. the world.

IDONOVOS WATETET coo bonesoen P Pp 12) af oe Nile.

Argulus incisus eeaeee KH

>» Yubropunctatus ... K

> personatus ......... K

> exiguus Sante K

5) angusticeps......... EK

» Striatus bt K

ae eubescens) ee EK -

op UAOANNUTG <oseon season P Ie 12 P PS Nile.

Chonopeltis inermis ......... a4 za Gen. E

MQ) SONOS sso scededcesace 18s BIL. |r. WE AIP dR iL IP),

The outstanding feature revealed in the above list is that while

two species occur widely distributed in Africa, these are asso-

ciated in Nyasa with a single endemic form, but in Tanganyika

with no less than seven. Thus out of ten known African species,

nine are found in Tanganyika, which is therefore seen to possess

not only a larger number of characteristic species, but.a much

richer Branchiuran fauna than the other great lakes with which

it may reasonably be compared. All the species enumerated are

confined to the African continent so far as is at present known.

Dolops ranarum is the only African representative of a genus

which is otherwise exclusively American. The genus Chonopeltis

was established from a single female specimen from Lake Nyasa,

while Argulus alone has a distribution which is world-wide.

OsTRACODA.

It seems probable that the Ostracod-fauna of the great lakes is

less well represented in the collections hitherto obtained than is

the case with any of the other groups of Entomostraca. This

is due to the fact that nearly all fresh-water Ostracods are

bottom-living rather than planktonic forms, and require other

methods than surface tow-netting for their capture, while the

bulk of the material at present examined was collected by the

tow-net. ‘This implies, of course, that many additional types

will probably be found when the lakes are more adequately

explored. Nevertheless, 22 species are on record for Tanganyika

(of which 20 ave peculiar to that lake), and 17 and 7 respectively

for Nyasa and Victoria Nyanza. No Ostracoda have so far been

recorded from Lakes Edward and Kivu,

+ For further particulars, bibliography, etc., consult Cunnington (72).

FAUNA OF THE AFRICAN LAKES.

ad Tangan- Victoria Ber Albert Other parts of

Name of Species. yika. Nyanza. Nyasa. Nyanza. the Hasile

Family Cyprip.

Cypria castanea............... ee Ex 1z na Natal.

Bo lenticulanisimmesne sc. te P§ ee Madagascar.

Paracypria declivis ......... iE

. complanata ... HK

5 conoidea ........ K

es ObGusteeente ees 1 ae Te P

3 denexauny nse. EK

F subangulata ... aD)

5. reniformis ...... K

os LemeTnlG) seuboo ees 13)

Be OpAacatee secs tet E

5 claviformis... K

e HexiTOSAaee=enene K

x CUIGAL jecetncenuee E

Iliocypris propingua......... a 13)

& Bi fae, ei ece ed: E

Heterocypris dubia ......... Be 1)

Hucypris flabella ............ = y

Fs IMELMOIS sso oko BX Ete P§ Natal.

& JatiGanidaepen sce E§

Stenocypris sinuata ......... oe i P§ Madagascar.

% malcolmsoni ... ot P tf Asia, Australia.

3 marginata... ... BR EK §

- perarmata... ... 12 je aes Natal.

45 fiilleborni ...... K§

- fasciculata... HS§

¥ Stagnalis ...... K be

Cypridella fossulata ......... Se P§ Tohlimandjaro.

Cypridopsis obliquata ...... Ki

as sinuata ........- H

5 cunningtoni ... Et E

* POSE, | a occ ae E

5 monodonta ... {|

i Selratal “ic... o}

* bidentata ...... i

3 tumidula ...... a0)

c: congenera ...... 0)

i miler eet..eees aa EK

Potamocypris fiilleborni ... sis eit H

-: viduella ...... for ee Pp : see Cape Colony.

Zonocypris costata ......... Se 1g df H. Africa, Madagascar.

5 ee VISM ene. - EK

Family CytHERIDa.

Limnicythere obtusata...... ae i

. michaelseni.. oe ee P§ Se Lake Rukwa.

I cen eraieny. oacaseraet: 4P, Guke 9 P. Te

ADS snecies eee ec OLye ee) 5 B2PS) SBOP: 1P.

(22) (7) (17)

+ This account is based in the main on the report published by Sars on the

Ostracoda collected by the Third Tanganyika Expedition (152).

§ Not actually recorded from the lake itself, but from within its drainage area.

566 DR. W. A. CUNNINGLON ON THE

The existence of a large number of Ostracods which are

peculiar to Tanganyika is the principal feature displayed by the

list of species. Of the total number of forms on record, it will be

noted that Nyasa with 17 species does not fall far short of Tan-

ganyika with 22, while Victoria Nyanza is a long way behind

with only 7. This is probably due—at all events in part-—-to the

fact that Nyasa and its drainage area have been more thoroughly

investigated in this direction than Lake Victoria. It may further

be remarked that less than half the records from Nyasa are from

the actual waters of the lake itself, the remainder coming from

ponds and swamps etc. in the vicinity. Corresponding collec-

tions from the surrounding ponds, swamps, and rivers are wanting

in the case of both Victoria and Tanganyika; imdeed, the whole

of the material collected in the three biggest lakes by the author’s

expedition came only from the lakes themselves. Additional

records are therefore to be expected, particularly from Lakes

Victoria and Tanganyika whenever a more complete investigation

of the Entomostraca becomes possible.

While Nyasa is shown to contain nearly as many different

forms as Tanganyika, it will be seen that in the nature of these

forms the lakes differ profoundly. More than half the species

from Nyasa have also been obtained elsewhere, but of the 22

species from Tanganyika only two have so far been observed out-

side the confines of that lake, or to put it in another manner,

AT per cent. of the Nyasa forms are endemic in the lake, while

nearly 91 per cent. are endemic in the case of Tanganyika.

Another feature of distribution which this table brings into

prominence is the association of certain groups of forms with

certain lakes. Only in three cases have species been recorded

from more than one lake—Stenocypris perarmata from Tanganyika

and Nyasa, Zonocypris costata from Victoria and Nyasa, and

Paracypria obtusa from Tanganyika and Albert. In every other

instance the species from the lakes differ from one another, and

appear to be either peculiar to or at least characteristic of the

lakes in which they are found. Without referring to the species

seriatim, it may be pointed out that the genera Stenocypris,

Potamocypris, and Cypria seem to be associated in particular with

Nyasa, and Cypripodsis with Tanganyika, while Paracypria, with

the exception of a single species, occurs only in Tanganyika

among the lakes.

Apart from the forms peculiar to Tanganyika (which number

20 out of a total of 44in the list of Ostracoda), it is to be observed

that 8 species are peculiar to Nyasa and 5 to Victoria Nyanza,

or 13 in all. It will thus be seen that when Tanganyika, is

excluded, more endemic forms are known from the remaining

lakes than is the case among the Copepoda. It is also singular

to note how restricted is the distribution of even the non-

endemic forms recorded, for only in a single case (that of Steno-

cypris maleolmsoni which is known from Asia and Australia) has

a species been obtained outside the continent of Afriea with

Madagascar.

To the genera Paracypria and Cypridopsis, which are so largely

FAUNA OF THE AFRICAN LAKES. 567

represented in Tanganyika, it is necessary to call special attention

on account of their remarkable nature. Of the former genus, no

fewer than twelve different species have been described from that

lake, eleven of which are peculiar to it, and they all, as Sars points

out (152, p. 756), exhibit striking features of specialisation.

It is also : at—that the

type species of Paracypria was obtained from the same brackish

lagoon on the Chatham Islands as was the type of the Copepod

genus Schizopera, which is likewise very characteristic of Tan-

ganyika among the lakes. The case of Cypridopsis is perhaps

more significant still, since of the ten species obtained by the

Third Tanganyika Expedition two only display all the features

-characteristic of the genus, and these are the two forms taken

respectively in Lakes Nyasa and Victoria. The other eight types,

which are confined to Tanganyika, exhibit certain fhe: eences

which suggest that they might more suitably be removed to a new

genus which would then be endemic in the lake (152, p. 757).

It is quite clear that in the case of this group also, Tanganyika

contains an unusually large number of forms, an exceptional

propertion of which are peculiar to the lake, and that of these

peculiar forms many show undoubted signs of specialisation.

CLADOCERA.

The outstanding fact concerning the Cladocera is the absence

of any representatives from the plankton of Tanganyika, and

also apparently of Kivu, which is in direct communication with

it. Thus the survey of this group of organisms resolves itself

into a comparison of the records from the remaining lakes, and

this, it 1s not surprising to find, lacks the interest which the

inclusion of species from Tanganyika usually affords.

Table of Distribution of Cladocera *.

ine i? Senta! Victoria yy ,., Albert Edward Other parts of

Nyanza. J Nyanza. Nyanza. : the world.

Family Srprpz.

Rarasidayszalayie a,c: anee. ns Ve ee ues Asia.

Diaphanosoma excisum ... 12 12 oY 1p i. Africa, Australia.

Family DarpHnipz.

Daphnia kirimensis ......... ae a £05 K

PP longispina ......... 12 Ps af a Cosmopolitan.

3 hiyalhindiyseesee secs P ae aoe ace Cosmopolitan.

% lumholtzi ......... le Ie Ae as Palestine, Australia.

- monacha ......... Bc a E

Hyalodaphnia barbata ...... li as a abe i, Africa.

Ceriodaphnia bicuspidata... iP Fide a 1g

33 cornuta ...... 12 P§ ~ ai Asia, Australia, S. America.

5 Qubiagey.. P a3 ts ue Sumatra.

3 rareeW OU kas 1p P§ ue le Almost cosmopolitan.

+ For further particulars consult especially Daday (76, p. 120). Additional

records for Lake Victoria are given by Delachaux (7'7) and for the smaller lakes by

Brehm (54).

§ Not actually recorded from the lake itself, but from within its drainage area.

568 DR. W. A. CUNNINGION ON THE

- Victoria Albert Edward Other parts of

Name of Species. Nyanza. Nyasa Nyanza. Nyanza. the aaah

Simocephalus vetulus ...... 1p ay a Cosmopolitan.

Moinodaphnia macleayi ... nist P§ =f o, Almost cosmopolitan.

WiC mHe) GIB, 45 cqocneenmnone= P 12 KE. Africa, Heypt.

oy WAMORUNAD, p00 s30sb oon: IP te a me Zanzibar.

Family Bosminip2.

Bosmina longirostris ...... 12 -P Cosmopolitan.

Bosminopsis deitersi ...... 2 Central Africa, 8. America.

Bosminella anisitsi ......... IP S. America.

Family MAcRoTHRICID#.

Macrothrix laticornis ...... 1p P§ Cosmopolitan.

Z hirsuticornis ... = P§ Algiers, Bevoes:

A cheyreuxl ....., 1p P Africa, S . America.

Ilyocryptus longiremis iP P§ Chemenelien,

Family CHyDORID®.

Camptocercus australis... P bis ore ee Australia.

Alonopsis singalensis ...... oe 2 es oe Ceylon, S. America.

Huryalona orientalis .. Bes. A zis P ‘a a3 Ceylon, S. America.

Pseudalona longirostris a P§ ss a3) S. America, New Guinea.

INIOMAY BUTANE 56 303990050 209 900 Ie Cosmopolitan.

“hy GUSUENEY) sboudsnooudobos PS§ si se se Kurope.

ry) GROWING, poocnddnenodnse PS§ 3 a8 ty, K. Africa, Europe.

Pee pulchel | aiereeeeercecerrt a Ie aor nat Australia, 8. America.

» quadrangularis ...... me P§ 00 = Europe.

rectangula Heh eee 1 IP ih 1 Cosmopolitan.

Rhy nchotalona rostrata... PS§ ae RS Sch Kurope.

Nonelilanexcis aieyeerer eens a: IPS a a Kurope.

3 elopulosa) eee: as P Me iy Ceylon, S. America.

ie RETRO Te aobaonancaasy bad as 12 iA seh Australia, S. America.

es punctata.:..-...0--. iP, st a cf Ki. Africa, S. America.

i diaphana ......... P fe Es Bb Australia, S. America.

Pleuroxus similis ............ P§ a e S. America,

is RYFPNAEDIS 65 aoc co0.000 fl PS§ rs ek Europe, N. America.

or ASS IMUIIS Neeeeee eee P§ se ee: a Natal.

Chyderus barroisi............ 12 Wa a ee Asia, S. America.

x spheericus ...... Pp P on oe Cosmopolitan.

- ventricosus ...... P§ P§ a8 ee Ceylon, S. America.

be leonardi ......... 12 ae ie cal Australia.

a [DIGREE cocoa coo o0nve. PS§ oe Ea et Europe.

clobosus eee a P§ es he Hurope.

Daday Al TMEKGROOS — cov noascosee ase PRS oe ee. Asia, S. America.

Monospilus dispar ......... W ac ae fe: Hurope, N. America.

Wh GHNINTBY soobsee soos 39. 060 1G IE 20 P. ues SIP,

51 Species Paeetecce SLE 30 P. 1E. ee eee

§ Not actually recorded from the lake itself, but from within its drainage area.

The list of Cladocera from the big lakes of Africa differs from

all the corresponding lists in the absence of a column for Tan-

ganyika. While the group 1s entirely unrepresented in Lake

‘Tang anyika itself, various Cladocera occur in the surface tow-

nettings which I made in the broad mouth of the Lofu River,

which enters the south-western corner of the lake. I had intended

myself to report on the Cladocera collected by the Third Tan-

ganyika Expedition, but I have not yet been able to do so, and

the specimens from the Lofu remain unexamined. During his

FAUNA OF THE AFRICAN LAKES. 569

investigations on the Copepoda and Ostracoda of the expedition,

G. O. Sars, however, kindly isolated and identified certain

Cladocera from Victoria Nyanza and Nyasa, and these forms are

included in my list of species without further comment, although

they constitute in some cases new records of distribution.

The reason for the absence of Cladocera from Tanganyika

remains a matter for speculation, but the most likely explanation

would seem to be that the water of the lake either contains

certain salts which ave fatal to sach organisms, or is perhaps of

too saline a nature to permit them to survive. Colour is lent to

this view by the already-mentioned occurrence of Cladocera

in the river Lofu and also by the marked difference in nature of

the Rotatorian fauna of the lake proper and the river mouth.

Rousselet (150, p. 794) reports that in a small quantity of

material from the river he distinguished 23 different species

of Rotifera, while in a large series of tow-nettings from different

parts of Tanganyika he was only able to find 10 species, of which

but one was common to Jake and river. It thus seems highly

probable that though the salinity of the lake water has not made

it impossible for certain Rotifers to live in Tanganyika, it is at

least the factor which has determined the striking difference

between lake and viver in this respect, Further, it may be

pointed out that a somewhat similar state of affairs has been

observed in the case of the phytoplankton. G.S. West in his

report on the fresh-water Alge of my expedition states (200,

p- 96) :—‘“‘ OF the thirty species observed in this river-plankton,

only one occurred in the plankton of the lake itself.” This

evidence is very suggestive, and appears to confirm the view put

forward, as it is well known that the distribution of these delicate

plant organisms is profoundly influenced by the nature of the

water.

This is not perhaps the place to discuss at all fully what is

known of the nature of the lake water, but the principal features

may be indicated as follows. The total amount of dissolved

saline matter is rather higher than in the case of Thames water,

but is not excessive. The amount of calcium salts is, however,

low, while salts of magnesium are present in much larger

quantities than is characteristic of fresh waters. It 1s an obvious

suggestion that the presence of this excess of magnesium salts—

which are characteristically bitter to the taste—while not ren-

dering the water unfit to drink, might be sufficient to affect these

organisms and prevent them from obtaining a footing in the lake.

It becomes necessary in this connection to refer to the nature

of the water of Lake Kivu, which is in direct communication with

Tanganyika. Here the quantity of saline matter is excessive,

and indeed most remarkable for a relatively large lake, From an

analysis by Hundeshagen (103) it appears that while the water

is almost entirely free of calcium salts, it contains a very unusual

amount of salts of sodium and magnesium—particularly salts of

magnesium, of which there are_more than twice the amount

present in Tanganyika.

The plankton of Kivu, among other lakes, was collected by the

Proc. Zoon. Soc.— 1920, No. XX X VITI. 38

570 DR. W. A. CUNNINGLON ON THE

German Central Africa Hxpedition of 1907-08, and Brehm,

reporting on the Cladocera obtained by the expedition (54, p. 168),

refers to the complete absence of these forms from Kivu, and

expresses the opinion that the nature of the water may explain

the fact. It seems, nevertheless, that a very occasional specimen

may be found—presumabl, y in the littoral zone,—for in the collec-

tion examined by Brehm was a tube labelled ‘‘ Surface, Lake

Kivu—one specimen Alona.” his specimen he failed to find,

but there is httle doubt that such individuals must be carried

into the lake by streams entering it. Indeed, the conditions are

probably paralleled in Tanganyika, for not only does the existence

of Cladocera in the Lofu River suggest this, but Sars (151, p. 67)

actually observed an isolated specimen of Moina during his

examination of plankton from the lake. Both these records I

have purposely omitted from my list.

It only remains to suggest that since the river Rusisi, which

is the outlet of Kivu, brings a considerable volume of water into

Tanganyika, it is likely that Kivu is the principal source of the

magnesium salts which appear to have an influence on the fauna

of both the lakes in question.

It would be interesting, and should not prove difficult, to

determine by experiment whether Cladocera are susceptible to

the presence of small quantities of magnesium salts in the water,

and, if so, whether they are affected by amounts too small to have

any adverse influence on ether Kntomostraca*. Indeed, a whole

field of experimental research is indicated, which might go some

way towards determining whether the very special nature of

the fauna of Tanganyika may not in some degree be related to

the unusual nature and quantity of the salts dissolved in the

water.

Turning from these matters to a survey of the list of Cladocera,

it will be noted that 51 species in all have been enumerated.

Of these, 31 species are recorded from Lake Victoria, 30 from

Nyasa, 6 from Edward Nyanza, and only one from Albert Nyanza.

It is interesting, though perhaps not surprising, to observe that

an overwhelming majority of the forms were already known from

other parts of the world, only 2 species out of 51 being described

as endemic, to wit Duphaad monacha from Lake Albert and

D. kirimensis from Lake Edward. A number of the forms are

truly cosmopolitan and many more are known from two or three

continents, while only four are restricted to other parts of Africa,

in addition to the lakes. On comparison with the particulars

given for the Copepoda and Ostracoda, it becomes clear that while

the former group holds an intermediate position, the Ostracoda

are the most lmited in range and the Cladocera. the most

extended. Of 44 species of Ostracoda enumerated, 33 are described

as endemic in one or other of the lakes: out of 54 species of

CULE 30 are endemic: while out of 51 species of Cladocera

only 2 are endemic. It can hardly be doubted that this is due to

* A brief reference is made by Gurney to his observations of the injurious effect

of magnesium salts on the growth of fresh-water Daphnia (97, p. 469).

EEO Oe eee a et ie

FAUNA OF THE AFRICAN LAKES. 571

the exceptional facilities for distribution afforded by the thick-

shelled “ resting” eggs of the Cladocera.

A detailed survey of the distribution of the genera in the lakes

does not seem called for, but certain features may be pointed out.

All the well represented and more important genera appear to

occur in two or more of the lakes, while it is the less outstanding

forms which are only known at present from one of the lakes in

question. Chydorus and Alona are the best represented genera,

each with 6 species, Daphnia and Alonella following closely with

5 species each. From Albert Nyanza, as already mentioned,

there is only the record by Brehm (54, p. 169) of a single species

of Daphnia. I have, however, in my See ae a small quantity

of plankton from the lake collected in July 1907 by my friend

R. 'T. Leiper (cf. 74), and though I have not yet found an oppor-

tunity of properly “eee the Cladocera which it contains, I can

at least indicate the occurrence of the genera Diaphanosoma,

Moina, Canyptocercus, Alona, and Chydorus. Waving made no

specific determinations, these new records are not included in my

list of forms, but attention is merely called to them in this place.

The addition of these genera to its ‘fauna brings Lake Albert far

more into line with Lake Edward and the larger lakes.

To sum up—Lakes Tanganyika and Kivu quite unexpectedly

contain no Cladocera. This is very probably due to the salinity

of the water, and perhaps in particular to the excess of magne-

sium salts in those Jakes. Apart from this, the distribution of

species in the lakes affords little of interest, many of the forms

being cosmopolitan and only an insignificant number endemic.

BRANCHIOPODA.

Although representatives of this group have been found in

various parts of the African continent, there is, as far as I am

aware, only a single record which concerns the lakes under

review. A species belonging to the family Estheride, viz.

Cyclesiheria hislopi, was collected by Fulleborn in the neigh bow

hood of Nyasa, though not actually in the lake itself (76, p. 159).

This form, originally described as /stheria, but subsequently

separated from that genus, is of wide distribution in the tropics,

and its occurrence in the fauna of Lake Nyasa has no particular

significance.

PENTASTOMIDA.

These parasitic Arachnids have not, I believe, been hitherto

recorded from any African fishes. 1 obtained a number of speci-

mens of Porocephalus from Tanganyika fish, encysted forms from

the body-cavity and others from the int eee The materiai has

been submitted for expert examination, but no detailed report is

yet available. Since the specimens came from very distinct

hosts—a Siluroid in the one case and a large Lates (Serranid) in

the other,—it may be they will prove to belong to different

species. It is open to question whether such foun should be

included in a description of lake faunas.

3e*

572 DR. W. A. CUNNINGION ON THE

HYDRACHNIDA.

It is probable that the Hydiachnida of the great African lakes

are but imperfectly known to science, although 31 species in all

have been enumerated. Comparatively small and inconspicuous

forms as these mites are, there seems little doubt that a good

many more species will come to light in the future. Following

on the earliest collections made by Stuhlmann, come those of

Filleborn in and around Nyasa and of Borgert in Victoria

Nyanza. Specimens from both those lakes and also from Tan-

ganyika were obtained by the expedition which I conducted

in 1904-05. It seems likely that Lake Nyasa, from which

the largest number of types (14) is on record, has been better

explored in this direction than the remaining lakes. No

Hydrachnids have hitherto been collected in either Albert

Nyanza or Kivu.

Table of Distribution of Hydrachnida *.

: Bie ete Tangan- Victoria Aas Hdward Other parts of

Name of Species, yika. Nyanza. Nyasa. Nyanza. the world.

Family HypRopHANTID®.

Georgella incerta ...........0.-. Bee ae P§ be Quilimane, Noss1-Bé.

Diplodontus despiciens ...... P ae Bee wee Cosmopolitan.

Family Hye@roparip2.

Limnesia armata ............... 1D)

xs campanulata ...... rs K

Anisitsiella africana ............ ae a K§

Hygrobates edentipalpis ...... H

Oxus stuhlmanni ............... 0)

Unionicula borgerti ............ a0)

5 Crassipes ......... 1p a as Hurope.

25 digitatus ......... au 1p ee Nossi-Bé.

Bi taleifer cory Ba E

i TEMIRAMIS) Soy soe eo 12 ie ee ee Europe.

- IKPRGRWS soosasoooce us Had sae E

is cunningtoni ...... ne ee 10)

Encentridophorus borgerti ... aps EH

koenikei ... sas a H§

spinifer ... rie? 1P ie cay Zanzibar.

Neumania paucipora ......... 5 12 a2 ae Hast Africa.

55 STN ENING 6 ud eco sce ae as P§ ie Soudan, Quilimane.

5 spimipes ............ ae P aM He Europe.

= palptlNOSa) eeeeeree eee aD)

Koenikea tessellata ............ wae He _&-£ :

ionaycaliciteta nese eee a ace 12 ane Madagascar.

oj OWMUUNCING) 5. casnanoce coon eg 12 e: Ags Hurope.

Forelia liliacea Be aactinn 12 Hurope.

Mideopsis minuta ............... 0)

Arrhenurus plenipalpis ...... is a 12 oe Madagascar, Quilimane.

5 sarcimabtus ......... aa a P§ aus Madagascar.

BS HESTON) spoonvece a ae HS§

a WINS“ no5 ocnoce ey of PS§ ae Nossi-Bé.

% voeltzkowi ...... a Bf P§ ae Madagascar, E. Africa.

TAroenera hon goo cc eeeeet Ones Gee 8P. Wet

GIL NACHOS ysncooennbepascnsen ily BIE 6H,6P. 54H, 9P. 1E.

©) 1 hae) (14)

+ For additional information refer to Daday (76, p. 237) and Soar (172).

§ Not actually recorded from the lake itself, but from within its drainage area.

FAUNA OF THE AFRICAN LAKES. 573

The distribution in the lakes of the forms at present recorded

is made clear by the adjoining list of species, and it will at once

be observed that the exceptional position held by Tanganyika in

the case of other animal groups is not in evidence here. While

the possibility is not excluded that further investigation may

bring to light an unusual number of remarkable species in the

lake, it is not in this group of animals that such a discovery is

very probable. Nyasa with 14 different forms is closely followed

by Lake Victoria with 12. Only 6 species are recorded from

Tanganyika and one from Lake Edward. The number of species

which are endemic affords no striking contrasts, 5 out of 14

being peculiar to Nyasa, and half the total number in each

case being peculiar to Victoria and Tanganyika. The only species

known from Edward Nyanza is endemic. There are no endemic

genera on record.

A point of more interest is the fact that of 31 species men-

tioned in the table, only two have been observed in more than

one lake, viz. Unionicula figuralis in Tanganyika and Victoria

and Hneentridophorus spinifer in Victoria and Nyasa. In every

other case the species from the Jakes differ from one another,

and have been recorded only from a single lake, where, in some

instances, they are endemic. Although this is so, there is not

much evidence of that association of certain groups of forms with

certain lakes, which is elsewhere a noticeable feature. It may,

however, be pointed out that both species of Limnesia are re-

corded only from Lake Victoria, while all the species of Arrhenurus

come only from Nyasa. Other genera well represented are

Unioncula and Newmania, but they both have an extended

range and have representatives in each of the three bigger lakes.

While 15 species in all are described as endemic, 10 of the

remainder are known from other parts of Africa (including

Madagascar) and 6 possess an even wider distribution. Under

these circumstances there seems every probability that future

exploration will prove many of the types to occur more generally

in the lakes, though on the whole it may be predicted that in

any case their precise distribution is little likely to afford evidence

of a striking character.

TARDIGRADA.

It is not surprising that inconspicuous organisms belonging to

this group have been seldom observed in the fresh-waters of

Africa. Apart from a record of the existence of these forms in

the Azores, there is, I believe, only the account given by Daday

(76, p. 236) of specimens which Fiilleborn collected in and around

Lake Nyasa. The specimens in this collection all belong to the

species Macrobiotus tetronyx, which Daday established to receive

them. While this form must be regarded as endemic in Nyasa,

the genus Macrobiotus is known to have a cosmopolitan distri-

bution, so that the existence of this particular species in the lake

is doubtless a matter of little import.

OLIGOCH ETA.

it is necessary, in compiling an account of the Oligochete

worms from the lakes, to distinguish as far as possible between

574 DR. W. A. CUNNINGION ON THE

the purely aquatic and the purely terrestrial forms. Yet this is

not a simple task, for not only are there a number of types which

he on the border-line, but even the true earthworms in the

tropics are semiaquatic in habit, living in very wet places not far

from water. for the purposes of this paper, the border-line

species as well as the true aquatics have been retained, those

forms being excluded which are generally recognised as earth-

worms. It follows from what has been said, that even worms

with specific names such as tanganyike have been omitted fiom

the list when the name has been applied to a terrestrial species.

Aquatic Gligochta ave recorded from five out of the six

lakes under review, but there is reason to suppose that a good

many more records will be forthecming. The list which follows

gives the impression of being incomplete, yet it incorporates, I

elie, all the available ntion mation. Since a number of these

forms are small and inconspicuous, it is not unlikely that they

have hitherto escaped notice. No species have been reported from

Kdward Nyanza. In a few cases, records have been inserted

without specific determination. While these are unsatisfactory,

they still have a certain value, as proving the occurrence of a

particular genus.

Table of Distribution of Bos ike i

Other parts of

Be Tangan- Victoria 5 Albert eee

Nameb ef Bmevien: yika. Nyanza. Wyasn Nyanza. = the world.

Family NatpomorPnHa.

INEM HTT, SO as doovecncoseessosen i P

INaissp. he puede tie Aiea! ie is iP

Dero sp. . eae on & P

Anlophorus stuhlmanni ...... sis iH

Pristina longiseta ............... Be ae P a8 ... Hurope, Ameriea.

Family ALLUROIDID.

Alluroides tanganyike ...... P es Me ey .. Rhodesia.

Family Mre@ascoLEcipm.

Sub-family Ocnerodriline.

Ocnerodrilus ivencua)

CUNARD Sas cessesecceoo0e 0)

Pygmeodrilus affinis ......... ae bh

y bipunctatus ... ae ae #3 1 P i. Africa.

rs bukobensis ... eee 19)

ie grawertl ...... 1)

es kiwuensis ...... she “ie Bes st E

Family GiossoscoLneipa.

Subfamily Criodriline.

ANIOTE, CHMIAIIE GG as soceocnaebanon’ sei i

» stuhlmanni «0.0... ne 1p af 1p .. KE. Africa.

9 ENGL a weston en scene 3 P. 4P. 3 12h BAP. 1)

WANS! Go cudvesagsecsosoons lo, ILIeS) 418, oP. 8) ie) oP. LE PP!

(3) (6) (2)

; + The records of these forms from the African lakes are somewhat scattered in the

literature of the subject. Refer especially to Beddard (12: 18) and to Michaelsen

(120: 121), who gives a bibliography.

FAUNA OF THE AFRICAN LAKES. 575

An examination of the table of distribution suggests in the

first instance that there are certain gaps in the list which future

investigation will surely fill. It is strange, for example, that no

representatives of the Naidomor pha have been found in Tangan-

yika or in the smaller lakes, for it does not seem likely that

they are entirely wanting. Secondly, it is cleav that very few of

the forms enumerated have been observed in more than one

lake, half of them being classed as endemic. Out of a total of

14 species, the highest number recorded from a single lake is

6 (Victoria Nyanza).

There is no indication in this case of a marked pre-eminence

for Tanganyika either in the number or peculiarity of its forms.

Lake Victoria leads the way with 6 species, 4 of which are

endemic, Tanganyika coming second, with exactly half that

number (3 in all, 2 endemic). Lake Kivu also contains an

endemic form. The species are distributed among a. large

number of genera (nine), only two of which—Pygmcodrilus and

Alma—are represented by more than a single type. Lake

Victoria again heads the list with representatives of four genera,

Tanganyika and Nyasa coming next, each with three. No

endemic genera occur.

Tt would serve no good purpose to review in detail the distri-

bution of the different genera and species, but reference may he

briefly made to the nature of the Oligochete fauna of each

lake. Tanganyika alone contains a member of the Alluroidide.

This worm—Alluroides tanganyike—was first described from

Tanganyika, but has since been obtained from Rhodesia in the

neighbourhood of the Victoria Falls. Two endemic species

of Ocnerodriline complete the list: there are no Naids or

Criodrilines known.—-In Victoria Nyanza the types are more

uniformly distributed among the families concerned. Of four

endemic species, two belong to the genus Pygmcodrilus and

one each to 4dulophorus and Alma—From Nyasa only three

Naid genera of wide range are on record. Pristina longiseta,

which I collected in that lake, is the only worm in the list which

occurs outside the continent of Africa, being known both from

Europe and America. It is strange that other aquatic Oligo-

cheetes have not been forthcoming from this lake as a result of

its careful exploration by Fiilleborn.—Lakes Albert and Kivu

each contain but two species. Two forms of Pygmcodrilus

(one endemic) come from Kivu, with one Pygmeodrilus and

an Alma from Albert. Neo Naids have so far been observed.

These lakes, together with Lake Edward, have recently been

visited by the German Central Africa Expedition, and it is.

therefore, the more surprising that no Oligochete worms have

been found in Lake Edward.

HIRvUDINEA.

There is but little known of the leeches which inhabit the

great lakes of Central Africa: indeed, there appears to he only

a single published record concerning them, which is that of

576 DR. W. A. CUNNINGTON ON THE

Blanchard (15) deseribing forms collected by Stuhlmann. It is

in a measure surprising that these creatures should have escaped

attention, since they are neither minute nor inconspicuous.

During my expedition to the great lakes I was able to collect a

considerable number of specimens, the majority of which were

obtained in ‘Tanganyika. Some examples, however, were found

in Victoria Nyanza (from which a representative had already

been recorded by Blanchard), and also in Nyasa, where they

constitute a new record. With this opportunity for gaining a

fuller knowledge of these forms, it is the more to be regretted

that no report has yet been received from the expert to whom the

collection was submitted for examination.

The leech which is already known from Lake Victoria was

termed Hirudo hildebrandti by Blanchard (15, p. 5). It is not

confined to the lake, specimens having been collected also in

Mombasa. In the same paper a new genus and species belonging

= Oo

to the family Herpobdellide is described under the name of Salifa

perspicax. This form was collected by Stuhlmann in Kiriwia,

which is described as north-west of Edward Nyanza. I have not

been able to identify this locality, but it appears doubtful whether

this species should be included in the fauna of this or any of the

lakes.

It is hardly safe to foreshadow the results of a proper exa-

mination of the specimens collected by the Third Tanganyika

Expedition. It may nevertheless be stated that they will in all

probability prove to belong to some five or six different species,

of which the greater number come from Tanganyika. This is only

in accordance with what has already been observed in so many

animal groups. The leeches were found for the most part in

shallow water, under stones or on shells, but some were dredged

in about 10 fathoms.

NEMATODA.

There is hardly any information available concerning the

Nematodes of the great lakes. In the extensive plankton

material collected by Fiilleborn in and around Nyasa, Daday

detected certain free-living forms, which he identified and de-

scribed. Reporting on Sonlenamca s collections, Collin records a

single specimen of a free-living Nematode (which he leaves

undetermined) from Victoria Nyanza, and also examples of

well-known parasitic type—Ascaris spiculigera—obtained from a

pelican on Lake Edward (64, p. 10).

During my expedition to the three big lakes, I was able to

procure a considerable number of parasitic Nematodes from the

fish which passed through my hands. Whether such forms can

be legitimately regarded as part of the lake fauna may be ques-

tioned, but as they are associated in this case with genuine

inhabitants of the lakes, Iam disposed to include them. Un-

fortunately these specimens have not been reported upon, so that

little can be added to this bare record of their existence. The

majority of them are from the body-cavity or gut of Tan ganyik

FAUNA OF THE AFRICAN LAKES, 577

fishes, but some were obtained from a large Siluroid taken in

Victoria Nyanza. In all, these parasites were collected from

8 different species of fish, belonging to 8 genera and 6 families.

Since these fishes are, with a single exception, endemic forms,

it is likely that their parasitic Nematodes will prove interesting

and perhaps new to science.

It does not seem logical to regard the Ascaris from the Edward

Nyanza pelican as belonging to the fauna of that lake, and as the

specimens I collected have not been identified or described, the

only records available for the purposes of this paper are those of

the free-living types from Nyasa. The species *, which all belong

to the family Anguillulidee, may be enumerated as follows :—

1. Monhystera filleborni.

Described as new from a single specimen in the surface

plankton.

2. Monhystera vulgaris.

A species known previously only from Europe. Obtained near

the mouth of a river which enters the lake.

3. Monhystera similis.

Likewise recorded from Europe and found in the neighbour-

hood of Nyasa.

4. Trilobus graciloides.

This species has been described as new by Daday. It comes

from the vicinity of Nyasa, but has also been taken on the

swampy shores of Lake Rukwa.

5. Mononchus tenuis.

Under this name Daday described a single male specimen

which he found in a sample of mud brought from the bottom of

Nyasa at a depth of 200 metres.

6. Monochus macrostoma.

Collected in a pool near the Jake. Also known from Europe.

7. Dorylaimus macrolaimus.

A European species which appears to be common in the waters

around Nyasa, but has not been observed in the lake itself.

Out of a total of 7 species, 2 may be regarded as peculiar to

the Jake fauna, while one of the remaining forms is confined to

Kast Africa. The others are known also in Europe.

Thus there is no indication that the free-living Nematoda. of

Nyasa are in any way remarkable or specialised, and information

is not available concerning either free-living or parasitic types

from any of the other lakes.

* Particulars of these will be found in Daday’s monograph (76, p. 44).

578 DR. W. A. CUNNINGTON ON THE

Rortrera.

The Rotifera of the African lakes are still very imperfectly

known, and the table of distribution which follows probably

gives quite an inadequate idea of the fauna. No information is

available for Lakes Edward and Kivu, and very little for Lake

Albert, Tanganyika, Victoria Nyanza, and Nyasa have been

better, though very unequally investigated. The latter has been

fairly well explored in this direction by the efforts of Fulleborn,

who obtained numerous examples of the lake plankton and, in

addition, suitable material from neighbouring ponds, swamps,

and rivers, ‘Tow-nettings from all three lakes were procured by

my expedition, the largest series coming from Tanganyika, but

with the single exception of samples taken in the broad mouth

of the Lofu River, which enters Tanganyika, I did not collect

outside the confines of the lakes themselves. Thus, while the

truly lacustrine types from Nyasa and Tanganyika may be fitly

compared, the Nyasa total is enormously swollen by species from

the adjoining neighbourhood, whereas the region around Tan-

ganyika remains almost entirely unexplored. In addition to my

small collections from Victoria Nyanza, there have been those of

Stuhlnann, Borgert, and Alluaud, but the Rotifera of the rivers

etc., within its drainage area are likewise practically unknown.

Table of Distribution of Rotifera*.

aah Tangan- Victoria : Albert Other parts of

Name of Species. yika. Nyanza. Nyasa. Nyanza. the PAA

Order Prorma.

Notommata copeus ............ yas P§ : ... Europe, Asia, N. America.

pachyura ......... P§ Kurope, N. America.

TAD OWGIG Aes tat ag sos oe P§ Cosmopolitan.

Pleurotrocha daphnicola ...... P§ ae Kurope, N. America.

Cephalodella forficula ......... PS Cosmopolitan.

Diaschiza forficata ............ P§ Kurope, America.

35 IDM genoa pena PS Cosmopolitan.

Bf TREAPHOUE ono 99nsaecon00e PS Europe.

Monommata orbis ............ P§ Kurope, N. America.

Dicranophorus auritus ...... P§ ... Europe, Asia, N. America.

forcipatus P§ ... Europe, Asia, N. America.

Encentrum caudatum ......... PS Europe, N. America.

Epiphanes oblonga ie ce die ES

ks clavulatus ......... #4. P fee Kurope.

” lotos P§ Asia.

Pa brachionus

spinosus ......... Ae 3 P§ Europe, S. America.

a Pmacrourus ...... a 12 1 Asia Minor.

+ The principal sources from which this table has been compiled are Rousselet

(150), Collin (64), and Daday (76, p. 59), where further particulars will he found,

Tt is weli to explain that the names under which the species are enumerated are

mostly those adopted by Harring in his “Synopsis of the Rotatoria’”, Washington,

1913.

§ Not actually recorded from the lake itself, but from within its drainage area,

Name of Species.

Brachionus angularis

39 bP)

caudatus ...

5 calyciflorus

bP) 5°

GLOGS) sen oe posoae

s capsuliflorus ......

bidentatus ......

TENGEN WS) segeneseones

forficula

mirabilis

JOELIDIIG IS s Goegaopsaoes

quadridentatus ...

i urceus

Platyias quadricornis

” bb)

brevispmus

Schizocerea diversicornis

homoceros) = se

Keratella cochlearis ....

PA quadrata

- 5 valga ......

Notholea striata labis .........

Anuropsis fissa nore

Mytilina compressa ............

9 MUCroOnaial eae

Py, aA spinigera

58 venbrallisiny. sees tees

oe 3 brevispina.

= ee

Huchlanis dilatata :

se OHO) cocosocacsooase

Bs triquetra® ..........

3 7 hyalina ..

Dipleuchlanis propatula ......

», leontina Ropes

gem OMAN Aste:

> ungulata

Monostyla bulla

53 lunaris

fe hamata

Lepadella ovalis

ss acuminata .........

ip cristata 8 20.

= patellaysa34) -3f.5.0.!

Colurelllajcolunay see

A adriatica

3 bicuspidata

a deflexa

53 DHOUENBD ds scone

Trichotria pocillum ............

Scaridium longicaudum ......

Redalitajmainal % cfs ae-ayeeass.

Filima longiseta

Tetramastix opoliensis .........

Trichocerca bicristata .........

“6 GUBRIA ik as sc

FAUNA OF THE AFRICAN LAKES.

Tangan- — Victoria

yika. Nyanza.

Pes JP

1p P

PS P

se 1p

PS§ Ie

P§

oes P

P 12

P§ 12)

P§

P a

P§ P§

ies

PS ee

IP 1p

PS§ P

P§

Nyasa.

Mtr

MMM ror hor ho roo:

Loe)

26) DIT IATL?

PDD PLIVLILLIL LIAL LIL

eiigehine incline hae iaeiineigciige)igelineigeiae)iae)

Albert

Nyanza.

579

Other parts of

the world.

Cosmopolitan.

Rhodesia, N. America.

Cosmopolitan.

Europe, N. America.

Cosmopolitan.

Asia, Europe.

Cosmopolitan.

Europe.

New Guinea, $8. America.

Cosmopolitan.

)

Asia, S. America.

Kurope.

Cosmopolitan.

bE]

Europe, N. America.

Cosmopolitan.

Europe, Asia.

Cosmopolitan.

bb)

Europe, N. America, Aastralia.

Cosmopolitan.

Europe.

Europe, N. America, Asia.

Europe, N. America.

Europe.

Cosmopolitan.

Asia, America.

Europe, N. America, Asia.

Cosmopolitan.

Europe, America.

Cosmopolitan.

Europe, N. America.

Cosmopolitan,

Europe, N. America, Australia.

Europe, Asia.

Cosmopolitan.

Europe, America.

Kurope.

Cosmopolitan.

Europe, Rhodesia.

Europe, Asia, N. America.

Cosmopolitian.

§ Not actually recorded from the lake itself, but from within its drainage area.

580

DR. W. A. CUNNINGTON ON THE

: : Tangan- Victoria Albert Other parts of

Name of Species. vik a Nivenie Nyasa. Wyanza. vemcenide

Trichocerca elongata ......... ab PS§ Cosmopolitan.

5 NEITANS sadopdlanebacees P§ ae Europe, N. America.

5 longiseta ...... 1S Cosmopolitan.

s TEKH, “Gooocapoa tas PS he Europe, N. America.

- rattus sochpptes Re P Cosmopolitan.

a SClplOR a reeeeeeeer es ie P§ 5s

s Stylataiescsren 1P 12 Europe, S$. America.

Diurella stylata 12 Europe, N. America.

» tenuior les Cosmopolitan.

ae tienis tendered he PS .

Syncheta oblonga............... P§ P§ Europe, N. America, Australia.

at pectinata <2. ......... re P Cosmopolitan.

THREAD os coo sos adones ae P§ is

IPO ARADO, DOEABY 5.6580 co9s0e090 12 P Bi

Sphyrias lofuana ......... ..... P§ a N. America.

Asplanchna brightwellii ...... a 12 1? Europe, N. America, Australia.

Be intermedia ...... iP ay Europe, N. America.

Asplanchnopus multiceps P§ Cosmopolitan.

Testudinella patina ............ ee P§ i

a trilobaitall seee. oe P§ Australia.

Order KH1IzoTA.

Floscularia ringens ............ P§ Cosmopolitan.

Limnias melicerta : Uae P§ 53

IP ADR, WANOGOA, oosscccx000050: P§ ae N. America.

Sinantherina spimosa ......... sau PS§ Cosmopolitan.

Conochilus hippocrepis_...... P§ a

Collotheca ornata ............... 12 Europe, N. America, Australia.

Order BDELLOIDA.

IPiniloyabing, GuRBw) .2.co5qn9 000002 sos P§ Cosmopolitan.

# CHOI eeeeer ee Le E§

MOSCOLAlg Nee er ee he P : Cosmopolitan.

Rotifer macroceros Bite 1? -

a4) NOTBUPDIS -ntys ons anode eco a P § 3

Pe avloarisiye .-etessecrss. PS§ P§ as

5p MOTIFS <6 ocenv a0 008 a P§ cn

42 venera 16P. ig}12 37 P. 4P,

105 species and subspecies... 1H,28P. 1H,24P. 1K,84P. 4P.

(29) (25) (85)

§ Not actually recorded from the lake itself, but from within its drainage area.

Attention has already been called to the fact that the Rotifera

of Nyasa are in all probability better known than those of the

other lakes.

Of 105 forms enumerated, 85 are reported from

that lake, while only 29 are given for Tanganyika, 25 for Victoria

Nyanza, and 4 for Albert Nyanza.

Examination of the Nyasa

records shows that a much smaller total has been observed in the

lake proper, and indeed it is the latter figure, compared with cor-

responding figures.from the other lakes, which affords a truer basis

of comparison than those given above, since the forms collected

from the surrounding neighbourhood are almost unknown except

for Nyasa.

Disregarding Albert Nyanza—from which informa-

tion is very meagre,-—the totals of those Rotifers taken within

the limits of the lakes themselves are found to be:—Nyasa 22,

FAUNA OF THE AFRICAN LAKES. 581

Victoria Nyanza 21, and ‘Tanganyika only 8. Clearly there is

little disproportion between Lakes Victoria and Nyasa, but Tan-

ganyika shows a marked reduction in number. It is probable

that the low figure for Vanganyika proper has some relation to

the nature of the lake water, which appears to be somewhat un-

suited to these organisms. Rousselet points out in his report on

my collections (150, p. 794) that there is a striking difference

between the scanty Rotiferan fauna of the lake and the far

richer fauna which he observed in a small quantity of material

from the Lofu River. Tanganyika water contains an unusual

amount of magnesium salts, and though very little is really

known concerning the influence of such salts on fresh-water

organisms, this seems likely to be the cause of the dissimilarity

between lake and river in this respect. he matter has already

been discussed in some detail in connection with the complete

absence of Cladocera from the lake (cf. p. 569).

On examining the list of Rotifera more closely, it will be

observed that with three exceptions, the forms are all known

from other parts of the world, many of them, in fact, being

cosmopolitan in their distribution. A single endemic species is

enumerated from each of the three bigger lakes, but apart from

Lecane lofuana described from the Lofu River, these are of only

doubtful value. The facilities for dispersal which the Rotifers

possess are well known, and quite account for this wide distri-

bution. In the case of these organisms, therefore, no deductions

of any value can be made from the presence or absence of a

species in a particular lake, and the interest attaching to a com-

parative table of distribution is, in consequence, small.

Nor does a study of the actual genera and species occurring

disclose feattres of much significance, although one or two com-

ments may be offered. In all, 42 genera are enumerated, of

which again the largest number is found in Nyasa. ‘Those best

represented are Brachionus with 12 species, or well-marked

varieties, 7'richocerca with 9 and Lecane with 6, while nearly all

the more important genera are known from at least two of the

lakes. Certain cosmopolitan species have been identified from

all the four lakes under consideration. There is little to note in

the way of unexpected forms or of types unaccountably absent.

A subspecies of Brachionus—B. capsuiiflorus bidentatus,-—which

was isolated from the Albert Nyanza material, is apparently very

rare, having been observed only i in Calcutta andl more recently in

Bulgaria. Furthermore, Rousselet has pointed out that the

genus 1splanchna is recorded from the lakes, but has not been

obtained in South Africa.—In conclusion, it will be observed that

asin the case of the Cladocera — which are also of extended distri-

bution,—there is no indication of an exceptional fauna peculiar

to Tanganyika, or indeed to any of the lakes.

GASTROTRICHA.

Representatives of this group may eventually prove to be

widely distributed in the African lakes, but at present there is

584 DR. W. A. CUNNINGTON ON THE

little information at hand concerning them. An account of

cartain forms collected by Stuhlmann at Bukoba, Lake Victoria,

is given by Collin (64), while other species which Fiilleborn

obtained in the neighbourhood of Nyasa have been identified by

Daday (76, p. 56). No Gastrotricha were observed by me during

my expedition to the great lakes. The species* may be enume-

rated as follows :—-

1. Lepidoderma squamatum.

A species well known in HKurope and occurring also in North

America. A single specimen was collected in a pool near the

shores of Nyasa.

2. Lepidoderma hystrix.

Under this name Daday deseribed a form obtained from the

sane locality as the above. It has not been observed elsewhere.

3. Ichihydium macrurum.

This type was described as new by Collin, but the species rests

upon a diagrammatic figure and a very incomplete description.

It was found at Bukoba, a station on the shores of Victoria

Nyanza, and has not been re-discovered, so that it may be looked

upon as a species of rather doubtful value.

4. Chetonotus formosus.

This form, like the two species of Lepidoderma, has been

identified from the vicinity of Lake Nyasa. Previous to this

discovery it was only known from North America.

5. Cheetonotus pusillus.

A single specimen of this type, hitherto only recorded from

Paraguay, was isolated by Daday from material collected in the

Mbasi River close to its entrance into Nyasa.

Two species of Chetonotus are, according to Collin, among the

Gastrotricha which Stuhlmann recorded from Lake Victoria, but

in neither case has it been possible to identify them further.

Brief notes made on the spot, accompanied in one case by a sketch,

form all the information available. It is doubtful whether either

of these types was observed in water from the lake itself.

6. Gossea pauciseta.

Another species previously known only from Paraguay.

Several specimens of this were obtained from a pond in the

Nyasa district.

On a survey of this list the following facts appear. Five

species are on record from Nyasa, one of which is peculiar to that

* Hor further particulars of these forms, consult Collin (64, p. 9) and Daday

(76, p. 56).

FAUNA OF THE AFRICAN, LAKES. 583

lake. From Victoria Nyanza three different forms have been

reported, of which one was described as new and has not been

found elsewhere. The two remaining types were not specifically

identified, and indeed all three species from this luke rest on

a somewhat insecure basis. None of the recorded species occur

in both Victoria and Nyasa, and it appears probable that none

were actually observed in the waters of the lakes themselves.

No Gastrotricha are known at present from any of the other

African lakes.

TURBELLARIA.

There can be little doubt that Turbellarians are relatively

uncommon in the lakes as well as in other parts of Central

Africa. Stuhlmann, a careful observer, who had opportunities

for collecting in many parts of the country, remarks on the

scarcity of these organisms, and in particular on the scarcity of

the fresh-water Dendroccels, which are usually common in ponds

and streams (181, pp. 1262 & 1268: 182, p. 652: 185, p. 349).

During my expedition to the great lakes, I only observed ‘Turbel-

larians in Tanganyika. They were found on the under side of

stones in shallow water, and all proved to belong to a single

species of the well-known genus Planaria. Prior to the

description of this Tanganyikan form by Laidlaw (107), the only

work dealing with species from the lakes was that of Bohinig (17),

who reported on the collections made by Stuhlmann. Untortu-

nately, the accounts in this paper are based almost entirely on

sketches and notes made on the spot, which often lack details of

importance for systematic determination. Although the records

must thus be regarded as of doubtful value, I msert them here

without further comment. The following species * come within

the scope of this survey :—

1. Planaria tanganyike.

This is the type from Tanganyika to which reference has been

made. It is the only species known from that lake and has not

been obtained elsewhere.

2. Stenosiona leucops.

A form known in Europe and also in North America. On the

evidence of drawings, recorded from the neighbourhood of

Bukoba, on Victoria Nyanza.

3. Stenostoma stuhlmanini.

Described as new by Béhmig from Stuhlmann’s notes and

sketches. Observed at Bukoba, Lake Victoria and unknown

elsewhere.

4, Stenostoma gilvum.

Another species based only on a drawing and a few notes and

* Vor details, consult the above-mentioned papers: Bohmig (17) and Laidlaw

(107).

584 DR. W. A. CUNNINGION ON THE

recorded from the same locality as 8S. stuhlinanni. Nothing

further is known of this form.

Examples of two species of Séenostoma from Bukoba were

among the material examined by Béhmig, but their unfavourable

state of preservation did not permit of nearer identification.

5. Gyrator hermuphroditus.

A pelagic Turbellarian collected in Victoria Nyanza is identified

with this species by Bohmig. It is well known in Europe.

6. Vortex quadridens.

This type is established merely on the evidence of ketalvas and

notes by Stuhlmann. It is admitted that certain important

features of the genital apparatus are unknown. Obtained from

stagnant water at Bukoba,

From the above it will be observed that Turbellaria are only

recorded at present from Lakes Victoria and Tanganyika. From

Victoria Nyanza and its neighbourhood five forms have been

specifically identified, three of them being peculiar to the lake.

There are, in addition, two types which are unnamed. ‘These

records of Bohmig, however, need confirmation, as they are based

on very meagre evidence. From Tanganyika a single endemic

species 1s known, which belongs to the widely distributed genus

Planaria *, No doubt farther representatives of this group will

eventually be found in the great lakes, but they appear to be less

common than might have been expected. There is no suggestion

of a striking Tarbellarian fauna in Tanganyika and little indica-

tion that such will be discovered in the future.

TREMATODA.

Practically nothing is known of the distribution of these

exclusively parasitic animals in the lakes of Africa. Following

the argument advanced in the section dealing with the Nematoda,

it seems only logical to include such forms in a lake fauna. In

the case of these organisms they may be obtained in the free-

swimming larval stage, or infesting an intermediate Molluscan

host or in their final vertebrate host. As far as I am aware, the

description given by Daday (76, p. 39) of two Cerearia larvee from

the neighbourhood of Nyasa is the only account which concerns

any of the lakes. Both were found in material from ponds near

Nyasa and were described as new larve, though it seems doubtful

whether such determinations have much systematic value. They

have received the names of ‘ Cercaria” schizocerca and “ Cercaria

hoplophora 76, p. 288).

* Tt seems clear that. Stuhlmann’s statement that Planarians cannot survive

temperatures of over 25° C. (185, p. 349) is not universally true. The specimens I

collected in Tanzanyika were taken in quite shallow water, where the temperature

tends to be highest, vet my thermometer readings for the surface of the lake showed

a higher average than 25°,

FAUNA OF THE AFRICAN LAKES, 585

In the course of my expedition I collected in Tanganyika a few

Trematodes which are parasitic on fish. They constitute a new

record for this lake, but have not yet been examined and

deseribed. The specimens were taken from large Siluroids, in

one case from the gill-arches, in another from the gut.

CESTODA.

As far as I have been able to ascertain, no records have been

published of tape-worms from the lakes included in this survey,

although Daday described two new larvee which he observed in

Kast African Copepods. Different forms of tape-worm proved

common in the gut of fishes examined for parasites by my

expedition, and I succeeded in getting a considerable number of

specimens. These all came from Tanganyika, where I had better

opportunity than elsewhere to seek such organisms, but it is

clear that systematic examination would not only bring to light

enteric parasites from the fishes of other lakes, but would result

in a far richer series from Tanganyika. My material has only

received a preliminary examination, so that little information

can yet be given as to the nature of the Tanganyika forms. A

species which occurred abundantly in an endemic Siluroid proves

to be one of the unsegmented Cestodes, and has been referred to

the genus Caryophylleus. It has not been more fully identified.

Almost all the remaining specimens come likewise from endemic

fishes belonging to different genera of Cichlids and Siluroids.

Thus it is quite probable that other species of tape-worm—-perhaps

new forms—are represented in the collection. It is to be hoped

that the investigation will shortly be completed.

CQ@LENTERATA.

The only representatives of this group at present recorded are

the common fresh-water Hydra and the medusa which so stimu-

lated interest in the fauna of Tanganyika. Stuhlmann obtained

specimens of Hydra from Victoria Nyanza which resemble the

common fH, fusca, though Weltner, reporting on this material,

would not venture to identify the species in the absence of eggs

(199, p. 2). Hydra has never been discovered in any of the other

lakes so far as J am aware, which is perhaps rather strange.

The Tanganyika medusa was described by R. T. Giinther under

the name of Limnocnida tanganice * (94), and being peculiar to

the lake, was, of course, regarded by Moore as one of the most

striking halolimnic or relict forms. The significance of its

occurrence in this lake in the heart of Africa, which communicates

with the sea only by some thousand miles of river, broken by falls

and rapids, was obvious, especially since the number of fresh-

water medusx then known was very small. Discoveries made since

Moore’s expeditions, however, have put a very different complexion

* The specific name is that of Bohm, the discoverer of the medusa, who wrote it

“tanganjice.” I follow Giinther (96, p. 651) and most subsequent writers in

adopting tanganice as a more rational spelling.

Proc, Zoou, Soc,—1920, No, XX XIX, 39

86 DR. W. A. CUNNINGTON ON THE

on the matter. Fresh-water jelly-fishes of other tiypes have been

found in the Yang-tse-Kiang, in a lake in Egypt and in the

Caspian, Moreover, the distribution of the genus Limnocnida

itself has proved wider to an unforeseen extent. In August 1903

a medusa was discovered in the great Kavirondo Gulf of Victoria

Nyanza and another in a fresh-water lagoon in the Niger delta.

The specimens from Lake Victoria were submitted to Giinther,

who pronounced them indistinguishable from the Tanganyika

species, though he subsequently modified his view, and described

them as ZL. tanganicew var. victorie (96, p. 651). The Niger

medusee were reported on by H. T. Browne, who considered them

undoubted specimens of L. tangunice (56: 57). In 1908, medusee

were found in a tributary of the Zambezi. ‘These were regarded

by C. L. Boulenger as specifically different from the Tanganyika

form, and were named by him Zimnocnida rhodesiw (18, p. 429).

The latter species has still more recently (1913) been obtaimed

from another part of Rhodesia, where it occurred in a tributary

of the Limpopo River (11). Remarkable as it was to find this

genus so widely distributed in Africa, the discovery of a species

in India in 1911 was more remarkable still (6). This type,

which comes from streams in the Western Ghats, differs slightly

from the African species, and was termed LZ, indica by Annan-

dale (9).

The bearing which these discoveries have on the problem of the

origin of the Tangany ika fauna, needs little pointing out. There

ean be no hesitation in deriving meduse from the ocean, and

when Tanganyika was believed to be the only locality in the

whole continent in which such organisms occurred, the lake was

naturally regarded as unique, and the medusze were looked upon

as strong evidence in favour of a marine origin for its fauna. In

the light of present knowledge, this evidence becomes greatly

weakened. If ZLimnocnida is to be regarded as a relict form, it

has now attained so wide a distribution that its presence no

longer singles out Tanganyika as a probable ‘ Reliktensee.”

Moreover, it must be noted that the new records are none of

them from the basin of the Congo, where the medusa should

be forthcoming in accordance with Moore’s view of the origin

of the lake.—The finding of a species of Limnocnida in India

affords another example of those Afro-Indian affinities to which

reference has more than once been made.

Tt will be more satisfactory to postpone a consideration of the

sources from which the medusa may have been derived, until the

general discussion—which will follow this systematic account—of

the zoological and of other evidence.

PoRIFERA.

Representatives of this group are known at present only from

the three largest of the six lakes under consideration. Since

however, fresh-water sponges are usually inconspicuous objects, it

ds very probable that sooner or later specimens will be found

FAUNA OF THE AFRICAN LAKES. 587

in the other lakes*, although Schubotz states that he completely

failed to find any in Kivu (156, p. xiv). The species from

Victoria Nyanza and Nyasa are quite ordinary forms, but those

known from Tanganyika exhibit divergences of at least specific

rank from the types cclleeted in any other part of the continent.

Of the eight species enumerated in the adjoining table, seven

helong to the cosmopolitan fresh-water genus Spongilld, the

eighth—about which doubts have been raised —being referred

provisionally to the genus Potamolepis.

Table ee Distr abaition of Por Ls:

Tangan- Victoria Ninn Other parts of

Name of Species. :

I yika. Nyanza. the world.

Family SPonGILLIp sm.

Spongilla biseriata ...... ve is P Egypt.

e (gohan. PS act ee ee Aun Gar

CAnveLl sesso a Ve .. Hungary, India, Java.

BS cunningtoni ... EK

a MOOLENN enh ee.. 1)

_ TANIGIER,, Gaede cae P§ White Nile.

tanganyikee ... 1D)

Potamolepis weltneri ... 13)

SIspeciesweeeesceae sn bin, es ie, ibe

§ Not actually recorded from the lake itself, but from within its drainage area.

The distinctness of the fauna of Tanganyika is once more

exemplified in the case of the sponges, and is shown graphically by

the above list. While Victoria Nyanza and Nyasa each contain a

single species of somewhat extended distribution, Tanganyika

contains six other types, four of them (those occurring in the

waters of the lake itself) being endemic.

The forms from Lake Victoria and Nyasa, both of which were

obtained for the first time by my expedition, do not call for much

comment. Spongilla cartert, now known from Victoria Nyanza, is

the only sponge from the lakes to occur outside the continent of

Africa. While it ocew's in Kurope, its occurrence also in India and

Java is noteworthy in view of the affinities of other African and

Indian types to which reference has already been made. Five

species of Spongilla have been recognised from the Tanganyika

area, three of them—all endemic—from the lake proper, and the

remainder from the Ugalla River, a tributary of the Malagarasi,

which in its turn enters the lake. The forms from the Ugalla

River, S. béhmii and SN. nitens, occur also in other parts of Africa.

Following the usual procedure, from which it is perhaps illogical

to depart, these species are inserted with an explanatory footnote

in the Tanganyika column, but it may be pointed out that the

* Sponge spicules were observed by West in tow-nettings made in Albert Nyanza

(201). It is probable that these occurred in the sample obtained from near the

bottom in about 30 feet of water.

+ The principal source of information on this group has been Kirkpatrick (105).

where references to other authors are given.

588 DR. W. A. CUNNINGTON ON THE

district from which they come is some 200 kilometres or more

east of the lake, and that they show East African rather than

Tanganyikan affinities.

The last of the forms enumerated—Potamolepis welineri—was

established by Moore (187, p. 323), and regarded by him as peculiar

to Tanganyika. Moore explains that a small specimen of this

sponge was dredged in the lake from great depths during his

second expedition *. It has never been adequately examined and

described, but Weltner expressed the opinion that it was a

distinct species, with a framework very similar to that of Spon-

gilla béhmii. It was accordingly given the specific name welinert

and placed provisionally in the genus Potamolepis. Kirkpatrick,

who has also examined some of the material, expresses the view

that these two species may possibly be synonymous (105, p. 222).

It is perhaps well to leave the matter sub judice, particularly as no

further supply of material has been forthcoming to aid in deciding

this point. Moore’s comparison of the spicules of this sponge

with those of the genus Reniera (which he misquotes “ Renieria”)

does not strengthen his case. Instead of being “the old fossil

genus ” (loc. cit. pp. 331: 354) it would be more accurately de-

scribed as the recent marine and brackish-water genus—although

fossil representatives also exist in the tertiary strata. In con-

clusion it may be stated that despite the suggested resemblance of

the spicules of Potamolepis weltneri to those of Reniera, the

sponge fauna of Tanganyika consists essentially of fresh-water

and not marine types, although the forms inhabiting the lake are

peculiar to its waters.

PROTOZOA. :

It is to be regretted that the Protozoa of Tanganyika have not

yet been investigated. There are, it is true, a few species to

record, but these are principally forms belonging to the Phyto-

flagellata, which are enumerated in the report on the Algee of my

expedition (200). This is the more unfortunate, since a good deal

of information on this group is forthcoming respecting Victoria

Nyanza and Nyasa, although much is probably unknown. There

is no reason to suppose that such a marked disproportion really

exists between the Protozoan fauna of Tanganyika and those of

the two other big lakes; indeed, the evidence afforded by many

groups of animals suggests that Tanganyika is likely to lead the

way with a larger and more unusual assemblage of species. An

examination by a protozoologist of the extensive series of tow-

nettings and other suitable material which I brought from the

lake should go some way towards determining this point, and it

is to be hoped that this will be carried out in the near future,

From Albert Nyanza there are a few forms recorded among the

* It may be mentioned in this connection that the species obtained by Moore’s

first expedition were, through some misunderstanding, said by Evans to have come

from a depth of 350 fathoms (82, p. 471). Moore himself states (127, p. 405) that

on this expedition 1200 feet (200 fathoms) was the greatest depth at which be was

able to dredge. ‘hese sponges are not in reality deep-water forms, for, during the

third expedition, I found them living in quite shallow water at the lake margin

FAUNA OF THE AFRICAN LAKES. 589

Phytoplankton, and from Edward Nyanza a single species of

Rhizopod, but it is obvious that these lakes too are quite

unexplored as yet. Nothing is known of the Protozoa of Kivu.

Table of Distribution of Protozoa 7.

N £ Specie Tangan- Victoria Nice Albert Edward Other parts of

AMG ORNS PECIes: yika. Nyanza. ~“9*5* 9 Nyanza. Nyanza. the world.

Class RHIZOPODA.

Order Loposa. :

Awnceb a proveuseeeaeres eee ee P i a iat Cosmopolitan.

AGRLlAlapIcdiac.ssccetcseeeccs eet Ve BAG wee ae Ki. Africa.

ay hy Gentatare caoeterteesc cs. ae “es PS ue ty Cosmopolitan. ‘

ReUP AISCOLMES ae ene ek ony P§ 12 “ A. Hurope, N.America, Australia.

Foie ye] CAUMUICIIO meee pene co ataok oe E i

Si ASCUILIC Ree nennen ape ane me 12 P§ Bs if Huvope, a America.

“7 > DB} Nos . ite n

UP SeVILULOAIS ee sete eas ste ie I Sas nee Cosmopohi an,

? Hyalosphenia papilio ......... sae P +S ie nd Europe, N. America.

Quadrula symmetrica ......... # 12 PS e Se Europe, N America.

Ditiueiaacuminataysesss-....0 | cos ae P§ ae Re Cosmopolitan. :

3 Constrictan..,.css.0 Sn oe PS A vehi Kurope, N. America.

- (ONOIND) — Gen ocasecseanas ant Ae IE nts = Europe, N. America.

is lobulossis.esscen a iE te ioe fee Cosmopolitan.

ps Mimme tre aenseeeseeeeee a 1p) 283 a fe Kurope. _

5 lobostoma ............ et a P§ 68 a Cosmopolitan.

DyLOMINS Nae eee eee P Ve tee aa Cosmopolitan. _ .

sa TUNGCOIENIEY Bo ohue accuse i in 12 1% 38 Europe, N.America, Australia,

Centropyxis aculeata ......... 12 IP a me Cosmopolitan. :

Webelaicanimnatalseeres deca s 12 Europe, N. America,

Pie CONMATISE eases ihe eee: Fs 1p P§ te one Europe, N. America, Asia.

og, LRAGXD AVI ODOT Boehesponsae Hee R ee ne a Europe, N. America.

Lecquereusia spiralis ......... P§ Ps Cosmopolitan.

Kuglypha alveolata ............ P§ ate Cosmopolitan. _

- COIVENEN Sanqnesseoncuee 6 : P§ Kurope, N. America, Australia.

Trinema enchelys ............... 0. P PS§ Cosmopolitan.

Cyphoderia ampulla ............ 1 PS§ Europe, N. America.

Order HELIOZOA.

Actinospheerium eichhornii... ... re P§ fee Af. Cosmopolitan.

Raphidiophrys elegans ...... if P a ae ie Cosmopolitan.

Clathrulina stuhlmanni ...... a iP ae Mee a East Africa.

Class MASTIGOPHORA.

Order EUFLAGELLATA.

Ruclenaacuse ere. te je § Cosmopolitan.

ae SPIO LAMENT eee tad a P§ Cosmopolitan.

aes 4 :

#3 VADISI 3 eas, pi 12 }e Cosmopolitan.

Colactumicalwuomiieens.. o-oo: 12 Europe. -

Trachelomonas annulata ...... P§ S. America.

x6 hispida .. ... PS Cosmopoliton.

5 OUTS ae asecere ES ‘ :

os volvocina PS§ Cosmopolitan.

Lepocinelis ovum ............... PS§ Burope, Australia. _

Phacus longicaudus ............ P§ oe ase Europe, Asia, America.

3 a :

5, pleuronectes ............ : ae PS§ ye ao Cosmopolitan. ‘

Pandorina morum ............ a 12 \e Kurope, Asia, America.

+ The majority of these records will be found in the accounts given of East

African forms by Schaudinn (154) and Daday (75: 76, p. 6), where further

particulars are given.

§ Not actually recorded from the lake itself, but from within its drainage area.

590

DR. W. A. CUNNINGTON ON THE

5 : Tangan- Victoria a ad Albert Edward Other parts of

Name of Species: yika. Nyanza. Nyeee: Nyanza. Nyanaza. the world.

Kudorina elegans P a a Europe, Asia.

Volvox africanus Ly ae ne E. Africa.

> aureus P§ Europe.

> globator PS Europe, Asia, America.

Order DINOFLAGELLATA.

Ceratium brachyceros ......... H

es hirundinella ......... 12 see f an Cosmopolitan.

ie macroceros Ags P Kurope, Asia.

Peridinium africanum ......... IP Je

oh berolinense......... 12 a6 Re: Europe.

a imconspicuum “a P ve Cosmopolitan.

5 palatinum ......... P§ a Pa Europe.

5 quadridens.. P a Re Europe, Asia, 5. America.

tabulatum ......... $5 1 Cosmopolitan.

Per idiniopsis cunningtonil ... 0}

Glenodinium pulvisculus Ip 1 1p Europe, Australia.

Class CILIATA.

Order CILIATA VERA.

Enchelyodon farctus Pp ee Be Europe, N. America.

Coleps hirtus .. P§ ue Sa Cosmopolitan.

Loxophyllum meleagris | ida IP oe ae Europe, N. America.

Trachelius ovum 12 kee se Cosmopolitan.

Dileptus anser ee ey re &, Europe, America, Australia.

Nassula sp. ct 12

Colpidium sp. ae ee P

Parameecium aurelia : ts eas Cosmopolitan.

Spirostomum ambiguum ...... By P te Cosmopolitan.

Condylostomarspsueeee ee

Stentor roeselit 1.0.0.0... We age a see Europe, America.

Tintinnopsis ovalis 1p Europe.

Uronychia paupera Ae HS§

Trichodina sp. iota eta EY

Worticellanlunagisis ssn Be P§ Kurope, N.America, Australia.

Ms microstoma ......... P 1P Cosmopolitan.

f MOOD, so cseen0 Pp Ps Europe, America, Australia.

Zoothamnium arbuscula ...... P§ Europe, America.

CENT) oscana ony Ds PS Kurope.

EKpisty lis anastatica IP PS§ Cosmopolitan.

53 brevipes ......... 12 Europe, S America.

i plicatilis ue P§ Europe, N. America.

ys WALT, oc nenesos 1p P§ Kurope.

Opercularia nutans 1p i Europe, America, Australia.

Cothurnia crystallina ......... P IPS Cosmopolitan.

60 incisa... 10)

ie lobata a0)

Order SucTORTIA.

Podophyra sp. Portree ii

Tokophyra cy clopum P§ Kurope, N. America,

Acineta symbiotica 1 H. Atrica.

> tuberosa Ps Europe, Asia.

AS ISenera sco ae rerqaecn ae oles 5) 122, 34 P. 4P. oe

Si/ DAC, sosnnccsoonvenceascon WSOP, AVBL BID, BIBLE. 4 P. LIP,

(7) (87) (60)

§ Not actually recorded from the lake itself, but from within its drainage area.

FAUNA OF THE AFRICAN LAKES. 59]

The most obvious comment on the table of distribution is the

great Inequality in the totals recorded from the biggest lakes—an

inequality which is obviously due in this instance to unequal

investigation. Of the 87 species enumerated, 60 occur in Nyasa,

37 in Victoria Nyanza, and only 7 in Tanganyika. While it is

impossible to predict what figures will be forthcoming when the

Protozoa of the lakes are better known, the figures at present

available are merely an index of this disproportionate investiga-

tion. The records from Nyasa are the result of the extensive col-

lections made by Fiilleborn, which were reported on by Daday (76).

It will be noticed that more than half the species identified were

not obtained in the lake itself, but came from river mouths,

pools, and swamps in the vicinity. Victoria Nyanza has been less

adequately explored in this direction, smaller collections being

made by Stuhlmann and more recently by Borgert and others.

In his book on the Tanganyika Problem, Moore devotes a few

paragraphs to the Protozoa of the lake (187, p. 323). The two

forms which he mentions by name—a Condylostoma anda large

Infusorian which he refers with some hesitation to the genus

Colpidiam— have been inserted in the table of distribution. He

describes the latter organism as the cause of the yellow clouds

which occur on the surface at times and make the water appear

“as if tinged with a fine golden dust.” This effect was also

observed by Livingstone, who thought the yellow scum to be of

vegetable origin. [I have repeatedly observed the phenomenon

myself, not only on Tanganyika, but on Nyasa, and without

denying other possibilities, I can confidently assert that it is

usually due to limnetic Alge. In conclusion, Moore remarks that

he found some twenty types of Protozoa belonging to groups

common in tropical fresh-waters.

A consideration of the list of species shows that in the

great majority of cases the forms are widely distributed if not

cosmopolitan in range. There are very few endemic species

enumerated—four in Lake Victoria, two in Nyasa, and one in

Tanganyika—and most of these are of little interest. Peridini-

opsis cunningtonti trom Tanganyika, and Uronychia paupera from

Nyasa, are perhaps of more interest, since in each case they are

only the second described species ‘of the genus. Forty-eight

genera ave mentioned in the table of distribution, of which the.

greatest number occur in Nyasa. This is a large proportion of

genera to species, but is accounted for by the number of genera

represented by only a single species. None of the genera are

endemic.

No good purpose would be served by commenting in detail on

the genera and species in the list, but one or two further remarks

on their distribution may be offered. As far as the Rhizopoda

are concerned, the lists from Victoria Nyanza aad Nyasa may be

reasonably compared, and they exhibita close degree of similarity.

The well-known genera Arcella and Difflugia are each represented

by a number of species. It is in the remaining groups that there

592 DR. W. A. CUNNINGTON ON THE

has been inequality of investigation, so that the Euflagellata, for

example, are only represented in Lake Victoria by a couple of

forms as contrasted with a large number from Nyasa. Volvow

africanus is an interesting species which was established to

recelve specimens brought by Leiper from Albert Nyanza. It

can no longer be regarded as endemic, since it has been observed

in another part of Hast Africa. he Dinoflagellates are recorded

from four lakes, and a comparison may thus be instituted. Lake

Victoria contains representatives of the genera Ceratium and

Glenodinium*. The former genus is wanting in Tanganyika,

where, however, Peridiniwm and Peridiniopsis replace it. In

Nyasa, Ceratium and Peridiniwm occur; in Lake Albert, Peri-

dintum and Glenodinium. It is particularly among the ciliated

forms that a number of genera occur represented only by a single

species. ‘The 7ichodina mentioned, which has not been specific-

ally identified, is parasitic upon the Tanganyika jelly-fish. Since

the latter, or a variety of it, is known from Victoria Nyanza, it

may well be that a 7richodina accompanies the medusa there

also, but I have no information on the subject.

In conclusion, it only remains to point out that the Protozoa,

as at present known, give no indication of an exceptional fauna

peculiar to any lake. At the same time it is precisely in

Tanganyika, if anywhere, that such might be expected, and that

lake remains virtually unexplored in this direction.

4, GENERAL DISCUSSION AND CONCLUSIONS.

Having completed the systematic review of the animals at

present known to occur in the lakes, the points of interest

concerning their distribution may now be fittingly discussed.

Far the most noticeable feature is that which has so repeatedly

shown itself, namely the unique nature of the fauna of ‘Tan-

ganyika. With a recorded total of over 400 different animal

types, the Jake is clearly exceptional; moreover, no details of

certain groups which are known to occur are yet available for

incorporation. As this treatise is concerned so largely with

Tanganyika, it may not be out of place to illustrate here in a

graphic manner, the growth of knowledge concerning its fauna.

The curve which follows may be regarded as approximately

correct. It is clear that the labour of ascertaining the date of

discovery or description of every type—even if possible—would be

out of proportion to the value of such a record. The largest

addition to the total (159 species in all) was made by the writer’s

expedition, which obtained, moreover, practically all the infor-

mation on the flora of the lake.

* Virieux states that in the plankton which he examined he observed a specimen

of Peridiniwn, but was unable to identify the species (197, p. 6).

FAUNA OF THE AFRICAN LAKES.

Text-figure 1.

D983

Curve showing growth of knowledge of Tanganyika fauna.

500

400 ——

Re)

S 3

+ Ke)

300 — - = i a

S <

2 D

200 ae ne :; S a

s &

100 ree

S Ry

s Ss

S S

0 cs

1850 1860 1870 1880 1890 1900 191d 1920

The relative peculiarity of each lake fauna, as far as totals only

are concerned, can be best illustrated by giving the figures in

parallel columns with the marks E or P against them.

Tanean-

yika.

57H, 111P.

(168)

Genera |

Species 293 H, 109 P.

(402)

Victoria

Nyanza.

2K, 137P.

(139)

110#, 179°.

(289)

Albert

Nyasa. Nyanza.

6H, 172P.|1, 47P. |

(178)

86H, 275 P.| 9H, 58P.

(361) | (67)

(48)

Edward

| Nema Kivu.

eau, 2 129 a 2

| (88)

| (64) (23)

An examination of these totals at once reveals the chief points

in which the fauna of Tanganyika is distinct.

the lake contains a more extensive series of forms (402) than any

other.

In the first place,

In the second place, those types greatly predominate

which are unknown elsewhere (293 out of 402).

Both these

points, viz. total number of species and corresponding number

of endemics, are shown graphically for each lake in the following

? t=) ro)

table :—

11H, 48 P. 41, 19 P.

594 DR. W. A. GUNNINGTON ON THE

Text-figure 2.

Table to illustrate richness of fauna and proportion of

endemic species to the whole.

450 a

400

350

300 i lind

250 ee Hl jue ca

150 oe ; —_——

100 E a

VIC TORIA ALBERT EDWARO KIVU

TWIG SES NYANZA NYANZA NYANZA

The height of each column represents the total number of species

2 : : I ,

and the shaded portion the number of endemic forms.

FAUNA OF THE AFRICAN LAKES. 595

It will be observed that the total figure for Nyasa, though

some way behind that of Tanganyika, is still very large. Since

in many cases—e. g. fishes, molluscs, prawns—Tanganyika has a

much richer fauna, such a result is perhaps a little surprising.

It is worth while to recount the facts which explain this. Firstly,

the group of the Cladocera is entirely absent from Tanganyika,

but well represented in Nyasa and elsewhere. ‘This is probably

due to the nature of the salts dissolved in the water, which

appears also to have a restrictive influence on the Rotifera of the

lake. Secondly, there are one or two groups of animals—notably

the Protozoa and free-living Nematoda—almost uninvestigated as

far as Tanganyika is concerned, although tolerably well known

for Nyasa. Lastly, the collections made by Fulleborn extended

to the waters surrounding Nyasa, while the rivers, ponds, and

swamps in the neighbourhood of Tanganyika remain unexplored.

This has greatly increased the records for Nyasa, particularly

in the groups Rotifera, Protozoa, and Entomostraca.

In the matter of endemic types, however, Tanganyika leaves

Nyasa (and the other lakes) far behind, exhibiting an astonishing

series of forms for which it is difficult to find a parallel.

Hxpressing in percentages the figures already furnished, nearly

73 per cent. of the species in Tanganyika are peculiar to the lake,

whilst Victoria Nyanza comes a poor second with 38 per cent.,

and Nyasa still further behind with some 24 per cent. The

smaller lakes have not only smaller totals, but have a reduced

proportion of endemic forms, namely 20 per cent. for Lake

Kdward, 17 per cent. for Kivu, and only 13 per cent. for Lake

Albert. Nor is this all, for the number of endemic genera which

Tanganyika contains places it in a category by itself. No fewer

than 57 out of 168 are regarded as peculiar to the lake, or rather

more than one-third of the total. As against this, Nyasa has

6 endemic genera out of 178, and Victoria Nyanza 2 out of 139,

while Lakes Albert and Edward each possess but a single endemic

genus. Tanganyika alone among these lakes has a family which

ean be regarded as endemic—that of the Tiphobiide (Gasteropoda).

It is thus clear that Tanganyika exhibits by far the most

striking series of endemic animals of any of the lakes under con-

sideration—indeed, it must be recognised as one of the most

remarkable lakes in the world. There are only two lakes, as far

as I know, which merit comparison with Tanganyika in this

direction, viz. the Caspian Sea and Lake Baikal. Unfortunately

I have been quite unable to obtain trustworthy figures of recent

date with which to compare the particulars now available for the

African Jake. It seems probable, however, that Baikal even

surpasses Tanganyika in the number of animal forms peculiar

to its waters*. It appears that while relatively few groups are

* Consult the series of monographs dealing with the results of the most recent

scientific expedition—Wissenschaftliche Ergebnisse einer Zoologischen Expedition

nach dem Baikal-See unter Leitung des Professors Alexis Korotneff in den Jahren

1900-1902. Lieferungen 1-5. Kiew und Berlin 1905-1912. It is to be regretted

that this work remains uncompleted, doubtless on account of the European war.

596 DR. W. A. CUNNINGLON ON THE

represented, there is often a great richness of species within the

groups. The fishes are far fewer than those of Tanganyika and

only half of them are endemic, while the Mollusca agree pretty

closely in the number of endemic types. On the other hand, there

are most extensive series of Oligochetes and Turbellarians, andan

extraordinary wealth of Gammarids, the species being in each case

nearly all endemic. In the case of the Caspian, I am able to

quote figures, according to which some 64 per cent. of the animal

forms are found nowhere else in the world (155, p. 34). If this

statement can be relied upon, the Caspian Sea, while sufliciently

remarkable, is less so than Tanganyika, which has nearly 73 per

cent. of endemic types.

Returning to a consideration of the fauna of Tanganyika,

other features revealed in the Systematic Account may be

summarised in a few sentences. Those groups which are most

conspicuous in possessing endemic genera and species are the

Pisces, Mollusca (especially Gasteropoda), and Macrura, with the

Brachyura following closely. The Copepoda and Ostracoda are

well represented by endemic species (but not genera), with the

Porifera and Polyzoa showing smaller numbers. While other

groups with few endemic species, appear, by contrast, devoid of

significance, there are only five of ali those represented in the

lake—Mammalia, Crocodilia, Chelonia, Batvachia, and Coelen-

terata—which do not contain endemic types. It may be added

that certain endemic forms are held to exhibit a marine aspect

and have been termed thalassoid (halolimnie according to Moore).

Such are many of the Gasteropod molluses and perhaps a Polyzoon.

In the same category comes the medusa, which, of course, is not

confined to Tanganyika.

The exceptional character of the Tanganyika fauna having

been sufficiently emphasised, an explanation of this marked

peculiarity must be sought. In other words, a general con-

sideration of what Moore called the Tanganyika “ problem” must

be undertaken. In order to appreciate the actual value of the

purely biological evidence, it is necessary clearly tc understand

the relations whieh exist between marine and fresh-water

organisms. ‘The essential points may therefore be stated as

briefly as possible *.

In the first place, while certain organisms are characteristic of

the sea and others of fresh water, the distinctions which exist

between marine and fresh-water forms are neither very great nor

very definite. Secondly, it may be emphasised that the barriers

which tend to prevent a change of medium are not wholly insur-

mountable. At the same time fresh-water types are usually

recognised in consequence of certain structural peculiarities

directly due to their mode of life. Such features should be

excluded as far as possible when deciding the systematic position

of an organism, for it is only thus that a true idea of its inter-

% These matters are discussed at greater length in several recent papers. Consult

Sollas (173), Cunnington (71), Gurney (97).

FAUNA OF THE AFRICAN LAKES. 597

velationships—which are quite independent of habitat—can be

obtained.

The undoubted affinities existing between marine and fresh-

water organisms are the direct result of a community of descent,

for there is no escape from the conclusion that life had its origin

in the ocean. Thus the forms now found in fresh-waters must

have attained their present distribution in one of three ways :—

(1) by a direct, active or passive migration from the sea; (2) by

becoming terrestrial or swamp-loving in nature, and secondarily

adapting themselves to life in fresh water ; (3) as a result of the

isolation and subsequent freshening of some portion of the sea,

due to movements of the earth's crust. Without speculating as

to which of these methods has played the most important part, it

may be pointed out that the salinity of the ocean has not been

constant throughout the ages, but is doubtless greater now than

in past geological times. Since certain types are known to have

recently migrated from the sea, it is not hard to believe that many

forms may have achieved the change during former epochs when

the obstacles to be surmounted were somewhat less. It is hardly

necessary to repeat that the view advocated by Moore assumes

that the remarkable organisms found in Tanganyika have attained

their present distribution by the third means, and have been

modified from marine types in a basin cut off from the sea.

Since the flora of a lake perforce exists under the same con-

ditions as the fauna, it will be well, before proceeding, to make

further reference to the plants of Tanganyika. The higher plants

show no outstanding peculiarities. There are certainly 8 species

of true aquatics which have been collected in Tanganyika alone

among the lakes, but all these are well-known African—-or even

cosmopolitan- -forms (ef Rendle, 147). On the other hand, the

Alge of Tanganyika differ markedly from those found in the

other big lakes, a number being endemic, while a few are usually

marine or brackish in habit. In all, some 21 species and 5

varieties are described as peculiar to Tanganyika. It is, however,

the phytoplankton of the lakes which affords the most interesting

comparisons. The plankton of Tanganyika is much richer in

species than that of either Nyasa or Victoria Nyanza. Out of a

total of 85 species, more than 70 per cent. do not occur in the

other two lakes, so that in this instance once more the features

characteristic of Tanganyika are exhibited. The presence in the

lake of brackish-water and quasi-marine Algee may perhaps be

accounted for by a period of growing salinity prior to the estab-

lishment of an outlet. (For further particulars consult the

detailed Report on the Fresh-water Algze of the Third Tanganyika

Expedition, 200).

It is natural that a good deal of attention has been attracted

to Tanganyika by the singular nature of its fauna, and it is

not surprising that various suggestions have been offered in

explanation of the facts. While it will be necessary to recount

the several views which have been advanced, it is appropriate to

598 DR. W. A. CUNNINGTON ON THE

deal in the first instance with Moore’s hypothesis that Tanganyika

represents an old Jurassic sea. This view, first put forward in a

paper published in 1898 (131), was subsequently developed and

the evidence detaiied at considerable length in “ The Tanganyika

Problem,’’ 1903, (187). Being thus a widely known theory, it is

only necessary to inquire how it accords with the facts which have

since come to light. From the botanical and geological, as well

as from the zoological side, more information is available than

when this hypothesis was propounded, and it may at once be

said that recent discoveries do not favour the theory. As the

zoological evidence has been examined and discussed in the body

of this paper, it is only needful to summarise the conclusions.

Moore’s view rests in the main on his comparison of certain

marine fossil shells of the Jurassic period, with those of Gastero-

pods living in the lake at the present day—coupled with his

deductions as to the anatomy and relationships of the latter forms.

Tt has already been explained (p. 549) that, in the opinion of

leading experts, neither of these claims can be substantiated. If

the counparison with Jurassic fossils is held to be inadmissible,

the period of the suppesed connection with the ocean remains in

doubt, but while Moore subsequently declared that he attached

no great weight to this comparison (138, p. 602), he still adhered

to his view that the so-called halolimnic animals were truly

marine or relict forms. As faras the molluscs are concerned, the

opinion of Pelseneer is in direct conflict with this view, for he

regards the halolimnic Gasteropods as emphatically fresh-water

types.

In this connection it may well be asked—-why are there no

thalassoid Lamellibranchs in Tanganyika? There seems no

obvious explanation of the fact, yet surely some members of this

group would also exhibit a thalassoid appearance had they been

relict forms from the ocean. This itself is evidence which tells

against the view of a relict origin for the fauna.

Nor do the other members of Moore’s ¢ group of halolimnic

animals definitely support his contention. The medusa, once so

important, and admittedly a marine type, has lost most of its

significance. Not only are fresh-water meduse known to occur

more widely than was formerly supposed, but the Tanganyika

species has been found in the Niger and in Victoria Nyanza.

The Decapod Crustacea—prawns and crabs—regarded in ‘ The

Tanganyika Problem” as constituents of the halolimnic group,

have been shown to belong to typically fresh-water families.

Much stress was formerly la, id upon the incrusting gymnolematous

Polyzoon Arachnoidea, but recent discoveries have modified its

importance. Since the genus is now actually living in Asiatic

seas, it can hardly be regarded as an ancient marine type

persisting only in Tanganyika, but must rather be looked upon as

a recent importation (cf. the analogous case of Victorella, p. 540).

Finally the sponges belong to the. family Spongillide, a charac-

teristically fresh-water assemblage. Thus, of the succession of

FAUNA OF THE AFRICAN LAKES. 599

animals from different groups which constituted Moore’s halo-

limnie series, none are accepted as peculiarly marine save the

Polyzoon Ar achnoidea and the medusa.

But more than this. The endemic animal forms have been

described almost without exception as specialised and not primi-

tive types. It is true that Moore regarded the remarkable

Gasteropods as essentially primitive in nature, but this view is

not shared by other writers (p. 550). If the halolimnic animals

are indeed relict forms, they must have been cut off at some

remote era—though it need not have been the Jurassic period—

and ought in consequence to exhibit primitive rather than

specialised characteristics.

This review of the zoological evidence makes it clear that on

such grounds it is impossible to justify the contention that

Tanganyika was connected with the sea in Jurassic times, or

indeed that a connection with the sea ever existed. It is there-

fore necessary to inquire what light may be thrown on the subject

by the evidence of geology.

In his book, Moore maintained that resting on the Archean

granites, gneisses, and schists which appear to constitute the

basement rocks of the continent, three types of sedimentary rock

are to be recognised. ‘The lowest of these consists of beds of

sandstone and shale, which are not only well developed in the

neighbourhood of the great lakes, but appear to extend over vast

areas of the African interior, including a large part of the Congo

basin. Then follow the beds discovered by Drummond _ north-

west of Nyasa, and covering these in turn, white shelly deposits

(Pleistocene) laid down by the lakes themselves. Drummond’s

beds being regarded as Triassic in age and probably estuarine,

fo)

Moore considered the great beds of sandstone and conglomerate

as evidence of an extensive ocean which at some still earlier

period covered a great part of the lake regions of Central Africa

(137, p. 65 et seq.).

It is particularly to this last point that exception is taken by

other writers on the geology of these regions, Some regard Drum-

mond’s beds and the great sandstone series as of like age, and

on the evidence of the fossils occurring in the former, regard the

whole as beds of the lower Karoo (Trias)—or at least as a forma-

tion of corresponding age, deposited under similar conditions. A

very recent writer on the stratigraphy of this part of the continent

(Behrend) speaks of the unfossiliferous conglomerates, quartzites,

and sandstones which are particularly well displayed in the

neighbourhood of the great lakes, as the “ Tanganyika System ”

(14, p. 52). These beds he distinguishes as of different age from

similar rocks occurring near Nyasa and in parts of the Congo

basin, assigning them “to an earlier perilod—Devonian or even

prior to that (14, p. 73 and Taf. iii.). While it may be that the

relative age of these different strata is by no means conclusively

fixed, these recent investigations show that Moore’s lowest

series—the “ Old African sandstones” as he calls them—really

600 DR. W. A. CUNNINGTON ON THE

comprises two or more formations. In any case all the evidence

goes to prove that these sandstones and conglomerates, whatever

their age, were laid down under continental, i.e. fresh-water and

terrestrial conditions, and thus afford no support for the view

that the ocean formerly extended over these large tracts in the

heart of Africa.

There is yet another geological objection to the view that

Tanganyika contains relict domme from an ancient sea, and that

is, that the depression itself would not appear to date back to

the remote times required by Moores hypothesis. There is every

reason to believe that the extensive faulting which produced the

Great Rift Valley took place in Middle “Tertiary times, and if

this be accepted, the basin of Tanganyika was actually not

in existence at the time when the Jurassic theory supposes it to

have received its marine fauna *.

Clearly the geological evidence does net favour a marine

“relict” origin Bow the peculiar fauna of Tanganyika, and it has

been shown that the testimony of zoology is against it. In order

to be convincing, a theory must not run counter to the findings

of either branch of science.

Since it does not seem possible to accept the hypothesis put

forward by Moore, alternative suggestions have now to be con-

sidered. In the first place, it 1s important to point out that

shells of the thalassoid Tanganyika genus Paramelania had been

compared by White (202: 203) and Tausch (186) with those of

the fresh-water Cretaceous genus Pyrgulifera some time before

Moore drew his comparisons with marine Jurassic shells. It has

been held by conchologists that the resemblance in this case is

every whit as close as between any of the forms compared by

Moore. This may constitute slender evidence on which to

theorise, but it is significant that the beds from which the fossil

type comes are not only more recent, but are fresh-water in

character and not marine. Thus, if any value attaches to the

evidence, it would suggest that the unusual molluscan genera

should be regarded as the little modified representatives of a late

secondary fresh-water assemblage rather than those of a much

earlier marine one. This view, which implies that the thalassoid

Gasteropods are relics of an ancient lake fauna preserved in this

basin, obviously did not find acceptance by Moore. He urged

against it the pertinent fact that in such a case, similar types,

living or fossil, ought to be found in other areas, and yet they are

conspicuously absent (137, p. 335). While agreeing that this

constitutes a serious objection, the same objection, to my mind,

may be raised with equal force against the marine Jurassic

hypothesis.

Reference has already been made to the fact that certain fossil

Gasteropods from the Balkan Peninsula exhibit a considerable

resemblance to some of the thalassoid types from Tanganyika

* Certain geological experts, indeed, regard the Tanganyika basin as more recently

produced than other parts of the Rift Valley system.

FAUNA OF 'HE AFRICAN LAKES, 601

(p. 550). The forms in question, described and figured by

Brusina (58:59), come from fresh-water Pliocene beds in Dal-

matia, Croatia, and Slavonia, From this it might be argued that

the lakes of the Mediterranean region which existed in Tertiary

times were the source from which the Tanganyika Gasteropods

have been derived. It is not unreasonable to suggest that com-

munication was possible between these lakes and the region of

Tanganyika by way of the valley of the Nile and the Great Rift

Valley. While less objection can be taken to this view than to

Moore’s, or even to the suggestion of a Cretaceous origin for the

molluses, there are difficulties in accepting it as a complete

solution of the problem. ‘To confirm this theory, either fossil

forms of like nature should be forthcoming in some intermediate

region, or (aS a communication to the nortli of Tanganyika is

assumed) living types should be found in Lakes Kivu, Edward, or

Albert. Since neither are known to occur, the case for ne

source of origin is unsupported.

Since the shells of the thalassoid molluscs have been held to

resemble (1) marine Jurassic types of the Anglo-Norman basin,

(2) in one instance a widely distributed fresh- or brackish-water

genus from the Upper Cretaceous of Hungary and North

America, (3) a series of fresh-water fossils from beds of Newer

Tertiary age in Jugo-Slavia, the evidence afforded is so contra-

dictory as to offer little guidance in determining the origin of the

fauna of Tanganyika.

It is a very natural suggestion that the thalassoid appearance

of the Tanganyika Gasteropods is directly due to the size, depth,

and quasi-oceanic conditions prevailing in such a lake*. That is

to say, that a marine aspect has been produced in certain

members of the ordinary African fresh-water series as the result

of convergence. To those who have visited these lakes and

realised their vastness this seems plausible, but the difficulty has

then to be faced, that similar forms are not forthcoming in Nyasa

and Victoria Nyanza.

An hypothesis in which this view is introduced, but which has

other novel features, was brought before the International

Congress of Zoology in 1913 by Germain ( (87). He holds that

previous writers have been wrong in considering Tanganyika by

itself, and urges that a clearer under standing of the facts becomes

possible on taking into account the organisms which inhabit

neighbouring lakes and rivers. Dealing with the Prosobranch

Gasteropods, which exhibit par eacellence a thalassoid facies,

Germain asserts that while Tanganyika contains a much larger

series of such forms than any other lake, it is not the sole locality

in which they may be found. He considers that Lakes Mwero

and Nyasa, as well as the Upper and Middle Congo, contain a

number of Prosobranchs (chiefly Melantide) the marine aspect

of which it is impossible to deny. Proceeding to discuss the

* A corresponding suggestion has been made to explain the marine appearance of

certain Crustacea and other organisms in Lake Baikal.

Proc. Zoou, Soc.— 1920, No. XL. 40

602 DR. W. A. CUNNINGTON ON THE

geological aspect of the problem, he states that very extensive

lacustrine deposits are actually known in Central Africa, reaching

from the Congo basin to Tanganyika and from that lake to

Victoria Nyanza on the one hand and Nyasa on the other. From

such considerations he supposes that there formerly existed in

east-central Africa a vast lake basin which united the Middle

and Upper Congo with Nyasa and Tanganyika and_ probably

Victoria Nyanza. Living in this region was a uniform fresh-

water fauna specially rich in Prosobranchs. After the formation

of the Great Rift Valley the lakes were isolated in their present

basins, where modifications of the original fauna resulted from

the new environment. Lake Tanganyika, possessing most nearly

the characteristics of the ocean, became inhabited by molluses

which have assumed (by a phenomenom of convergence) a marine

aspect in the highest degree*

With this conception I do not find myself wholly in accord.

Without expressing an opinion on the nature of the Gasteropods

of Nyasa and the Upper Congo region, there are two serious

objections to its acceptance. In the first place, there is in-

sufficient geological evidence for a lake basin so large in extent—

the deposits in this area being probably terrestrial and fluviatile.

In the second place, there is no reason why the conditions in

Nyasa and Victoria Nyanza, which closely resemble those of

Tanganyika, should not have produced an equally striking series

of thalassoid Gasteropods in those lakes, and yet this is not the

case. At the same time I am quite prepared to agree that

the marine aspect of the molluscs is probably due to convergence.

Another obvious suggestion is that the salinity of the water

has been a determining factor in producing mavine-like forms.

Here it is much less easy to come to a decision, for the question

of increased salinity is of course directly associated with that of

a period of isolation. But prolonged isolation itself, with the

opportunity it affords of development free from competition with

the outside world, is obviously a cause predisposing to the pro-

duction of new characters. Where isolation and a gradual

increase in salinity have coexisted, it is difficult to recognise

which factor is responsible for a particular result. Experimental

evidence is not wanting to show that certain salts, even in

minute quantities, exert a profound influence on aquatic organ-

isms, but it does not follow that a marime aspect would be

produced as a result. West, indeed, goes so far as to assert

that the Algz of Tanganyika which exhibit marine affinities may

well have been produced by a gradual increase in the Salinity of

the lake during an extended period of time (200, p. 191). Here,

it is true, the two factors are inextricably associated, but if the

suggestion is not unreasonable for the Algz, the surmise may be

* Since this account has been in the press, a still more recent paper by Germain

has reached my hands—‘‘ Histoire Océanographique des Lacs de l’Afrique Orientale.”

Bull. Inst. Océanoer. Monaco, No. 369, 1920. In it, he adds little which is new,

merely re- affirming the opinions expressed i in his earlier ar ticle,

FAUNA OF THE AFRICAN LAKES. 603

hazarded that the thalassoid Gasteropod shells owe their nature

to the same cause. Be this as it may, there are additiona] com-

plications affecting the salinity of Tanganyika.

It has been shown that Tanganyika had probably no outlet

until a portion of the Nile basin became cut off and Kivu drained

south into the lake (p. 515). It has also been pointed out that

since Kivu water contains an excessive amount of magnesium

salts, that lake is probably the source from which the high per-

centage in Tanganyika has been derived (p. 570). If these

probabilities be accepted, certain conclusions as to salinity follow.

During the first period the salinity may well have been consider-

able, though there is no evidence as to its nature. The lake

subsequently freshened, but eventually its waters became rich in

salts of magnesium. Thus any effect which the saline nature of

the water may have exerted on the organisms of the lake may

have been due to either of these conditions, or to a combination

of both. As already suggested, there may even exist an irregu-

larity in the outflow of the lake, due to the forming and breaking

of dams in the bed of the Lukuga River (p. 515). If this be the

ease, the salinity of Tanganyika has not only changed consider-

ably in the past, but may still be changing materially from time

to time.

The view that Tanganyika owes its remarkable organisms—not

merely the thalassoid forms—to a long-protracted period of

isolation, has been advocated by several writers, and remains,

on the whole, the most likely suggestion put forward, The

' possible effect of an increased salinity, which isolation would

involve, must of necessity be coupled with this, but it is not

regarded as the prime factor. This view has the positive

advantage that it does not run counter to geological conceptions,

but fits in with what is believed to be the past history of the lake.

Testimony in favour of it is afforded by the very remarkable

nature of the Cichlid fishes which Tanganyika contains. This

group has long been known to show a. peculiar facility for

colonising isolated and often saline waters, though the agency by

which this is effected is not understood. What then more likel

than that the Cichlids were among the earliest inhabitants of the

Jake, where, without having to compete with other types of fish,

they multiplied unchecked and became differentiated into new

genera and species (cf. 26, p. 423).

It is hardly necessary to point out that this isolation

hypothesis does not assume that Tanganyika was stocked from

any exceptional source. That is to say, the lake did not receive

its fauna from an ancient sea, but in the same manner as the

neighbouring fresh-waters, the original similarity of its fauna to

those of the other lakes being secondarily lost by marked

divergences of form consequent upon prolonged isolation. The

marine aspect of certain Gasteropods would thus be regarded as

merely due to convergence. Viewed in this light, the case

of Tanganyika is closely analogous to that of oceanic islands,

40*

604 DR. W. A. CUNNINGTON ON THE

which, as isolated areas of land, are well known to possess faunas

and floras largely peculiar to themselves,

The last-mentioned hypothesis, even if satisfactory in the main,

makes no attempt to account for the presence of the medusa in

Tanganyika, and accordingly a few sentences are needed on this

aspect of the question.

In dealing with the distribution of African fresh-water fishes,

Boulenger has discussed the problem of Tanganyika, and states

that he cannot admit Moore’s contentions (26, p. 422). He refers

to the inconclusive evidence afforded by the so-called halolimnie

Gasteropods, and clearly regards the medusa (now known from

other parts of the continent) as the only organism in the lake for

which it is necessary to account in any special manner. He

points out that paleontological evidence exists of a Middle

Eocene (Lutetian) sea which extended over a large area in

Northern Africa (vide also Hudleston 102, p. 352), and suggests

that this would afford a rational explanation of the present

distribution of the medusa in Africa.

With this view Gravier is not in agreement (90, p. 221). He

gives it as his opinion that the medusa may well have migrated

from the sea at a recent epoch, especially should it possess a

hydroid stage, as is held likely by Browne (cf. 56, p. 306). Its

present distribution in Africa he explains by reference to the

possibilities of intercommunication between the river systems.

Having dealt at considerable length—as becomes its import-

ance --with the fauna of Tanganyika and the views put forward

to account for its very unusual character, 14 is now possible to

proceed to a brief study of the remaining lake faunas. While

there are smaller totals and fewer peculiar types, the bigger lakes

at least are not devoid of interest.

Victoria Nyanza, with 38 per cent. of endemic species, clearly

has characteristics of its own, these being more prominent in

some groups than in others. By far the most conspicuous group

is the Pisces, containing, as in the case of Tanganyika, the largest

series of forms. It comprises also over half the total number of

endemic species and the only two endemic genera. 'The Mollusca

again are noteworthy, but in this case the Lamellibranchs are

more striking than the Gasteropods, exhibiting a larger pro-

portion of endemic types. Of the few Ostracods recorded from

the lake, 5 out of 7 are described as peculiar, and the Oligochete

worms are represented by 6 species, 4 of which are endemic.

Victoria is the only lake besides Tanganyika which contains the

medusa, though this should perhaps be regarded as subspecifically

distinct. It is only in this lake that the common Hydra is

known to occur. Many groups are wholly without endemic

representatives.

Generally speaking, Lake Nyasa exhibits very similar features,

but with rather fewer peculiar forms. Fishes constitute half the

total number of endemics, and 5 endemic genera out of 6.

A considerable number of molluscs are known, nearly half being

er ee

FAUNA OF THE AFRICAN LAKES. 605

peculiar to the lake. Nyasa contains an endemic genus belong-

ing to the Argulide, and the Ostracoda are well. represented,

8 species being endemic out of 17. It is strange that no aquatic

snakes and no Polyzoa have yet been recorded ; on the other hand,

Nvasa alone of the lakes under review contains aquatic tortoises

referred to the Trionychidee.

The three smaller lakes contain representatives of fewer animal

groups, but it is possible that this is merely due to less syste-

matic investigation.—Albert Nyanza displays only i3 per cent.

of endemic forms, which are mostly Mollusca, while it has no

endemic fish. The genus Limmnocaridella (prawn) is the only

genus peculiar to the lake.—Hdward Nyanza contains a more

interesting series of fish, with one genus and six species endemic.

Only 3 molluscs are peculiar out of a total of 15, and there is

little else which calls for comment.—In Kivu, representatives

of only 6 groups of animals are at present lmown. to occur.

While future exploration may bring other forms to light, the

poverty of its fauna is probably connected with the exceptional

salinity of the water. Out of a total of 23 species, there are

13 fishes, 3 being endemic, while the only other endemic type

is an Oligochete worm. The apparent absence of the hippo-

potamus and the crocodile is a point of some interest. Similarly

no Lamellibranchs appear to occur, although two forms of

Gasteropod are known.

It is obvious that none of these lakes exhibit such striking

forms as Tanganyika, and that while endemic types are not

wanting, these are fewer in number and for the most part only

specific in character. Apart from the presence of the medusa in

Lake Victoria, there is nothing to suggest a special connection

with the sea. In broad terms it may be said that the lakes

contain the ordinary fresh-water fauna of Africa modified in

varying degree in each case. Where such modification is con-

siderable, as in Victoria Nyanza and less markedly Nyasa, it may

well have been caused by a period of isolation proportional to the

relative peculiarity of the fauna. It is more especially the fish-

fauna of these lakes which is rich in endemic species and shows

certain endemic genera, and from this evidence it would seem

that Lake Victoria remained isolated for a longer period than

Nyasa (ef. p. 536). If isolation be accepted as accounting for the

remarkable fauna of Tanganyika, it is clear that a still longer

period must have been necessary in that case to produce such

notable results.

There are certain other matters concerning the distribution of

animals in the African lakes which are brought out by this com-

parative survey. While representatives occur of most of those

groups which may be expected in tropical fresh-waters, there are

some interesting exceptions. It comes as a surprise to a Euro-

pean naturalist to find no fresh-water Isopods such as Asellus, or

Amphipods such as Gammarus, yet these familiar forms appear

to be conspicuously absent from the tropical parts of Africa,

606 DR. W. A. CUNNINGYON ON THE

though the genus Gammarus is recorded from the north and

south of the continent. Why such types are wanting it would

be idle to speculate, but the fact is also emphasised by Stuhlmann

in more than one place (181, p. 1268: 182, p. 652). It is strange

too, that among the intestinal parasites OF the fresh-water fish

there appear to be no species of Hchinorhynchus, although they

are common in the fish of European rivers. Daday, it is true,

has described a larval form from a species of Ostracod taken in a

small Hast African lake (76, p. 55). Thus, while Asel/ws and

Gammarus are commonly the intermediate hosts, it is evident

that the absence of these genera does not form a complete

barrier to the distribution of Lehinorhynchus, and Daday con-

siders that it will yet prove common in Africa. At present,

however, the adult ferm is entirely unknown.

Concerning the numerical distribution of species in the lakes,

there is one point which calls for further notice. From a study

of the lists of forms found in each lake, Moore believed that a

definite relation existed between numbers and size. After

reviewing the facts then at his disposal, he writes :—‘ It is thus

obvious that from some cause or other the number of specific

forms in an African lake is roughly proportional to the size of the

lake itself” (137, p. 146). This does not mean that the smaller

lakes are less well stocked with animals, but simply that the

number of species they contain is less. The matter has already

been referred to in the systematic section, and it has moreover

been shown that the principle appears capable of extension to the

number of genera and even families (pp. 535, 548). With the

total figures for the six lakes now available, it is possible to test

the correctness of this conception on a more extended basis.

The totals for species and genera are therefore given in tabular

form, with the lakes (apart from Tanganyika) arranged in order

of size.

Tangan- Victoria Albert Edward

yika. Nyanza. Where Nyanza. Nyanza. Kivu.

Number of Species ......... 402 289 361 67 54 23

i Generate tee, 168 139 178 48 35 13

Tanganyika, which heads the list, 1s in every sense to be

regarded as a case apart, but the figures for the remaining lakes

should accord with this law-—if such it be. A descending series

is seen to exist: Nyasa it 1s true constitutes an exception, but,

as already explained, its totals have been artificially swollen in

certain directions ‘p. 595). Nyasa conforms to the rule in the

case of the Pisces and Mollusca—it is the addition of many types

of Rotifera and Protozoa which chiefly accounts for the large.

total figure for the lake.

FAUNA OF THE AFRICAN LAKES. 607

A very similar result is arrived at on comparing the number of

groups represented in the different lakes. From Victoria Nyanza

downwards the decrease in size is accompanied by a decrease in

the number of groups present. The figures (including Tangan-

yika for comparison) are found to be as follows : Tanganyika

26 (groups), Victoria 25, Nyasa 24, Albert 15, Edward 14, ‘and

Kivu 6. It is possible, however, that the low totals for the

smaller lakes are, in part, a result ia less complete investigation.

Knough has been said to show that Moore’s contention is

substantially correct as far as these African lakes are concerned.

It would be interesting to discover whether a similar relation

between size and number of specific forms can be made out for

other groups of lakes or even if it is a principle of general appli-

cation. Moore makes a comparison, which would seem to be

justified, between this phenomenon and that exhibited by the

flora of oceanic islands, where the smaller the island,—although

it may be as thickly covered with vegetation as any other area,—

the fewer the species of plants which inhabit it.

The last matter to be considered is the undoubted affinity

which exists between certain African and Indian fresh-water

types—an aflinity which has been noticed already, when re-

viewing the groups in which it is most pronounced, ‘This inter-

relationship i is exhibited in many groups of animals, and extends

not merely to forms from the Indian Peninsula itself, but from

the whole of that part of Asia, including the Malay Archipelago.

An interesting account of these affinities is given by Annandale

in a paper entitled “The African Klement in the Freshwater

Fauna of British India” (10). So far as the present treatise is

concerned, consideration is limited to those animals which are

known to occur in one or other of the big African lakes. Briefly

enumerating the cases, the Cyprinide and Mastacembelidze among

the fishes indicate this affinity in a marked degree. The Polyzoa

afford striking evidence, since the genus Arachnoidea is known

from Tanganyika and East indienne seas, while in the case of

Plumatella (Afrindella) tanganyike the actual species has been

found in an Indian lake as well as in Tanganyika. Caridina

nilotica with its varietal forms occurs in several of the African

lakes, while it is widely distributed in Indian and Malay regions

and extends still further east into Australia. Among the

Ceelenterata a medusa has now been found in India which is

generically identical with that from Tanganyika and Victoria

Nyanza, while finally among the sponges Spongilla carteri from

Lake Victoria is known in India and Jaya.

This is not the place to discuss the geological evidence for a

former land-connection between these areas, but the views

commonly held can be stated in a few words. During the

Carboniferous period, and persisting subsequently through the

Permian and Triassic, there appears to have existed a vast

tropical continent which extended from Brazil to Australia,

embracing of course Africa and India. This continent is known

608 DR. W. A. CUNNINGTON ON THE

as Gondwanaland. In the ensuing period—the Jurassic—Gond-

wanaland began to break up, but there is some evidence that in

late Cretaceous or even early Tertiary times a land-bridge still

existed between Hast Africa and the Indo-Malayan region, by

way of the Seychelles and Maldives. he geological record

is thus quite in keeping with the facts of distribution to which

reference has been made.

In bringing this study to a conclusion, it must be admitted

that im many directions information is very limited. There is no

doubt that the discovery of additional species is to be expected

whenever a re-examination of any of the lakes occurs, but there

are other points of considerable interest on which knowledge i 18

much to be desired.

Despite the investigations of Moore in Nya: asa and Tanganyika,

very little is really (nowt concerning the deeper regions of any

of the lakes, and the existence of a distinct abyssal fauna is

uncertain. In a paper dealing with the distribution of the

molluses, Moore speaks of obtaining certain thalassoid forms in

Tanganyika at a depth of 800-1200 feet (244-366 metres)

(129, p. 171). He regarded these particular Gasteropods as a

deep-water assemblage, but a more thorough examination may

well reveal other animals which permanently inhabit the bottom

muds. There is little doubt from an analogy with other deep

lakes that the deeper waters of these African lakes are almost, if

not quite, devoid of life. At the same time, further investigation

may indicate a definite association of abyssal forms not only in

Tanganyika, but in the other lakes under review.— Associated

likewise with the distribution of organisms within the limits of

the lakes are questions such as the vertical distribution of

plankton forms, as ascertained by tow-nettings. During the

Third Tanganyika Expedition my operations were practically

confined to surface tow-nettings, but special tow-nets worked

from suitable craft would afford the necessary information.—-

Again, the seasonal variations of plankton organisms are almost

unknown, although [ was able to detect a marked decrease in the

quantity of material during the rainy season. Detailed know-

ledge of this kind can only be gained by the aid of large collections

extending over many months.—The distribution of local forms or

varieties within the limits of a single lake was discussed by

Moore in his book. He considered that certain well-marked

varieties or even species were confined to particular areas in the

greater African lakes (187, p. 149). My own observations lead

to a different conclusion, and the matter undoubtedly merits

further investigation.

Lastly, there are some outstanding physical matters which are

of importance on account of their relation to biological pheno-

mena. Very little is known concerning the chemical impurities

of the water in these lakes, and even the depth and general

nature of the basin is imperfectly known save for Tanganyika,

Victoria and Nyasa. Further knowledge is likewise desirable as

FAUNA OF THE AFRICAN LAKES. 609

to water temperatures, seiches, and the possible occurrence of

temperature seiches.

It is clear, nevertheless, that from the facts already established,

a reasonably tirue conception may be formed both as to the nature

of the lakes themselves and that of the organisms which they

contain. It has been the aim of this work to supply a connected

account of these facts, based on the most recent particulars.

Considering the difficulties which beset the investigator im a

tropical climate far from civilisation, the amount of information

available is not discreditable to those concerned.

5. SUMMARY.

The special interest attaching to this comparative study of

African lakes is due to the remarkable nature of the fauna of

Tanganyika. That lake was discovered in 1858 by Burton and

Speke, the latter bringing back with him shells considered to have

a distinctly marine appearance. Subsequent collections emphasised

this point, and interest was further increased by the discovery of

a medusa by Bobm. A scientific expedition to investigate the

fauna was despatched in 1895 in charge of J. E.S. Moore. The

rich and unusual nature of the fauna then collected led him to

formulate the hypothesis that Tanganyika represents an old

Jurassic sea. . In order to test the validity of this hypothesis,

a second expedition, on which Moore was accompanied by Fer-

gusson, left England in 1899. The result was held by Moore to

justify the theory, and he embodied his conelusions in a work

entitled ‘‘ The Tanganyika Problem,” published in 1903. As the

aquatic flora had not been taken into account, a third expedition

was despatched to rectify that omission and make a further

collection of animals. This left in 1904 in charge of the writer,

returning in 1905. More recently still, in 1912- 13, the Belgian

expedition of Louis Stappers visited the lake and obtained

additional information.

The scope of this paper includes, besides Tanganyika, the five

adjacent lakes of most interest, viz. :—Victoria INiyamcaly Nyasa,

Albert Nyanza, Edward Nyanza, and Kivu. All these, with the

exception of Lake Victoria, occupy portions of the Great Rift

Valley, which has probably been formed by trough-faulting on a

stupendous scale. They lie in long narrow depressions bounded

by escarpments rising to a height of two or three thousand feet

above the level of the water. Nyasa and Tanganyika are very

deep, the former reaching to over 780 metres and the latter to no

less than 1435 metres. Victoria Nyanza has the largest area, but

occupies only a shallow basin bounded by low hills.

In all the lakes, but especially the largest, conditions are almost

oceanic. Climatic differences are negligible, but water tempera-

tures are uniformly high, showing an average of about 26°C.

Analyses of the water have been made in very few instances.

The water of Tanganyika, while fresh, is unusually rich in salts

610 DR. W. A. CUNNINGYON ON THE

of magnesium, and that of Kivu contains excessive quantities of

the latter as well as sodium salts. It is likely that the salinity

of Tanganyika was greater formerly, and may still be subject to

variation. Evidence exists of a considerable rise and fall in the

level of the lake, yet it seems probable that rainfall and evapora-

tion are very nearly balanced. There is reason to believe that

Tanganyika had no outlet until it received an additional water

supply from the Kivu basin, which was cut off from the Nile, and

added to the drainage area: of the big lake by the formation of a

voleanic dam in recent geological times. ‘Tanganyika would thus

have been completely isolated and its waters more saline until an

outflow was established. The present etHuent appears to have been

formed as an affluent, its bed being finally captured by a tributary

of the Congo. A periodic rise and fall in the lake level may

be caused by a temporary damming of the bed of the effluent,

indirectly due to irregularities in the rainfall.

As regards fauna, it is probable that at the present time all

the six lakes have received fairly equal investigation. Only

strictly aquatic animals are considered in this paper, but forms

obtained from the neighbourhood of a lake are included in the

totals. ‘Tanganyika exhibits by far the most remarkable features,

containing some 402 species of which 293 (nearly 73 per cent.)

are endemic. Nyasa has a total second in point of size, but this

has been artificially swollen by extra-lacustrine records which

are wanting for other lakes. It has only 24 per cent. of endemic

forms, while Victoria Nyanza with a smaller total has a larger

percentage of endemics, namely 38 per cent. ‘lhe three smaller

lakes show a great reduction both in number of types and number

of endemics. Tanganyika is further distinguished in that 57 out

of 168 genera are peculiar to its waters, whereas Nyasa can only

muster 6 endemic genera and the other lakes fewer still.

The fishes of Tanganyika are of outstanding interest, comprising

146 species, of which 121 are endemic. The most notable feature

is the number and high degree of specialisation of the Cichlide,

which with 27 genera (21 endemic) and 89 species (84 endemic)

is the richest Cichlid fauna in the world. A species of incrusting

gymnolematous Polyzoon occurs, such forms being mostly marine.

There is a large molluscan fauna, and of the Gasteropods more

than two-thirds exhibit a marine-like appearance. These are

known as the thalassoid or halolimnie group and are without

exception endemic. There are no thalassoid Lamellibranchs.

Twelve species of prawns are known, typically fresh-water in

character, but specialised and all peculiar to the lake. There is

an endemic genus of crabs, with 3 species. The Eucopepoda,

Branchiura, and Ostracoda are well represented, each showing a

large proportion of endemic species. The Cladocera are con-

spicuously absent from the lake and the Rotifera are relatively

few in number. This may be related to the salinity of the water.

The medusa originally described from Tanganyika has now been

— ees

FAUNA OF THE AFRICAN LAKES. 611

discovered in Victoria Nyanza and the Niger. There are four

endemic species of fresh-water sponges. Only 5 groups of animals

contain no endemic types.

Tanganyika is one of the most remarkable lakes in the world,

the only cases comparable being the Caspian Sea and Lake Baikal.

Recent figures are difficult to ascertain, but while Baikal may

even surpass Tanganyika in the number of unique animal forms,

it appears that the Caspian is less striking.

There is reason to believe that the relations between marine

and fresh-water organisms are intimate and due to a community

of descent. The barriers which prevent a change of medium are

not insurmountable. Organisms originated in the ocean, and

have attained their distribution in fresh water in various ways.

Moore regarded many of the Tanganyika types as relicts from a

former ocean. 3

The aquatic plants of Tanganyika are of less interest than the

animals. The higher plants show no peculiarities, but the Algze

differ from those of the remaining lakes. A number of species

ave endemic and others are usually marine or brackish in habit.

The phytoplankton is rich in species, and more than 70 per cent.

of the forms do not occur in Nyasa or Victoria.

Recent discoveries do not favour Moore’s hypothesis of a

marine Jurassic origin for Tanganyika. Neither his comparison

of shells from the lake with marine fossil shells, nor his views on

the primitive nature of the halolimnic Gasteropods, have been

accepted by leading experts. No members of his halolimnic group,

save the Polyzoon and the medusa, can be regarded as peculiarly

marine. The Polyzoon is allied to a species still living in Indian

seas and the medusa is known from other parts of Africa. The

endemic animal types are held to be specialised rather than

primitive in nature. Geological evidence is not more favourable.

The extensive beds of sandstone and conglomerate which occur

in the lake regions were probably formed under fresh-water

and terrestrial conditions. ‘hey are considered by some to be

of Triassic age, but may belong to a much earlier period, 7. e.

Devonian. Thus there is no support for the view that the ocean

at one time extended over the Congo basin. Further, there is

much to show that the trough in which Tanganyika les was not

formed until Middle Tertiary times.

A comparison has been made between thalassoid shells from

Tanganyika and a fresh-water Cretaceous genus on the one hand

and fresh-water Pliocene shells on the other. Since the thalassoid

shells have been held to resembie types from such different sources,

they offer little indication as to the origin of the lake fauna.—The

quasi-oceanic conditions in Tanganyika may have produced an

effect on the organisms it contains. Germain asserts that

Gasteropods of marine aspect occur in other regions besides

Tanganyika, and are derived from the fresh-water types of a

former vast lake basin. On the present lakes becoming isolated,

612 DR. W. A. CUNNINGTON ON THE

the conditions in Tanganyika produced a more striking series of

such forms than elsewhere. This view is not regarded as accept-

able.—It has been suggested that the salinity of the Tanganyika

water has produced marine-like forms, but further evidence 1s

needed.—The view that Tanganyika owes its remarkable organ-

isms to a prolonged period of isolation is regarded as the most

likely suggestion. It does not run counter to geological

conceptions.

This theory does not account for the medusa. Boulenger

suggests that it may have survived from an Kocene sea in

Northern Africa. Gravier considers, on the contrary, that it

may be a recent migrant from the ocean.

The faunas of the remaining lakes are of less interest. Victoria

Nyanza is next in importance to Tanganyika. The most con-

Spicuous group is the fishes, with a large proportion of endemic

species. The Mollusca also are noteworthy. The fauna of Nyasa

is similar in character, but with fewer peculiar types. There are,

however, 5 endemic genera of fishes. The smaller lakes contain

representatives of fewer animal groups as well as fewer species.

Kivu is the extreme case, with only 23 species, of which 4 are

endemic. ‘The poverty of its fauna may be associated with its

exceptional salinity. Albert Nyanza displays the smallest pro-

portion of endemic types, viz. 13 per cent. It is suggested that

periods of isolation would account for the peculiarities of Victoria

Nyanza and Nyasa.

Certain animal types are unexpectedly absent from the African

lakes. Such are the fresh-water Crustacea d sells and Gammarus

and the fish parasite Eehinorhynchus. No explanation of this is

forthcoming.

The number of specific forms in these lakes appears to be

proportional to the size. ‘This would seem to hold good also for

the number of genera and families and even for the number of

groups represented. It is possible that this principle is of

general application. It is thought to be analogous to the phe-

nomenon exhibited by the flora of oceanic islands.

The affinity between African and Indian fresh-water types is

recognisable in several instances among the inhabitants of the

lakes. This affinity is explained by the former existence of a

continent which embraced these countries in the Carboniferous

and subsequent periods.

Little is known of the deeper regions of any of the lakes. It

remains to be discovered whether associations of abyssal animals

exist in them. Neither the vertical distribution of plankton

forms, nor their seasonal variations have yet been studied.

Further information is needed on the salts dissolved in the water,

the depth and nature of the lake basins, water temperatures, ete.

It is nevertheless possible from established facts to form a true

conception of the nature of the lakes and their organisms.

FAUNA OF THE AFRICAN LAKES. 613

BIBLIOGRAPHY.

[A number of references which are not cited in the body of this paper are included

in the Bibliography in order to add to its general value. All the more important

works on Tanganyika and its fauna are included, as well as the principal sources of

information on the fauna of the other lakes. |

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2. AnceEy, C. F. —“Résultat des recherches malacologiques de Mer.

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“ Recherches Bathymétriques sur les lacs Moévo et

Tanganika.” Ann. Biol. lacustre, T. vii. 1914,

p. 83.

“Composition chimique de V’eau de surface des lacs

Moéro et ‘Tanganika.” Renseignements de

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—“ Zoological Results of the Third Tanganyika Ex-

pedition, 1904-1905.—Report on the Isopoda

Terrestria.”” Proc. Zool. Sec. 1908, p. 554.

—“Ueber einige neue Fischarten aus dem T'ancan-

to} t=}

yikasee.” Anz. Akad. Wiss. Wien, 1909, p. 399.

“ Beitraige zur Kenntnis der Fischfauna des Tangan-

yikasees und des Kongogebietes.” Sitzb. Akad.

Wiss. Wien, Bd. exx. Abth. i. 1911, p. 1171.

—“Reptilia.” Wiss. Ergebnisse d. Deutsch. Zentyal-

Afrika-Exp. 1907-1908, Bd. iv. 1913, p. 197.

—“Vorlaufiger Bericht tiber eine mit Unterstititzung

der Koéniglichen Akademie der Wissenschaften

unternommene Reise nach Ost-Africa, zur Unter-

suchung der Stisswasserfauna.” Sitzb. Akad.

Wiss. Berlin, 1888, p. 1255.

“Zweiter Bericht tiber eine mit Unterstiitzung der

Kéniglichen Akademie der Wissenschaften nach

Ost-Afvica unternommene Reise.” Sitzb. Akad.

Wiss. Berlin, 1889, p. 645.

“Beitrage zur Fauna centralafrikanischer Seen.—

I. Stidereek des Victoria-Niansa.” Zool. Jahrb.

Syst. Bd. v. 1891, p. 924.

“Zur Kenntniss der Fauna central-afrikanischer

Seen.—I]. Ueber eine neue Art der Arguliden-

Gattung Gyropeltis.” Zool. Jahrb. Syst. Bd. v.

1891, p. 152.

“Mit Emin Pascha ins Herz von Afrika.” Berlin,

1894,

622

186.

187.

188.

189.

190.

191.

192.

193.

194.

195.

196.

197.

198.

199.

200.

201.

202.

203.

204.

205.

ON THE FAUNA OF THE AFRICAN LAKES.

Tauscu, L.

THIELE, J.

Tuomson, J.

Tornier, G.

ULMER, G.

Vavra, V.

VIRIEUX, J.

WELTNER, W.

West, G.S.

Wuits, C. A.

Wissmann, H. von.

Woopwarp, S. P.

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und deren fossile Verwandte.” Sitzb. Akad.

Wiss. Wien, Math-Naturw. Classe, Bd. xe.

Abth. i. 1884, p. 56.

—Diagnosen neuer Arguliden-Arten.” Zool. Anz.

Bd. xxiii. 1900, p. 46.

“ Beitrage zur Morphologie der Arguliden.” Mitt.

Zool. Mus. Berlin, Bd. ii. Heft. 4, 1904.

“Mollusken der Deutschen Zentralafrika- Expedition.”

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1907-1908, Ba. iii. 1912, p. 175.

— To the Central African Lakes and back.” Vols. 1.

& 1i. London, 1881.

“On the Geographical Evolution of the Tanganyika

Basin.” Report British Association, 1882,

p- 622.

.—“Die Kriechthiere Dentsch-Ost-Afrikas.” Deutsch-

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Ch. Alluaud et R. Jeannel en Afrique orientale.

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ON A NEW MOSQUITO. 623

33. Descriptions of the Adult, Larval, and Pupal Stages of

a New Mosquito from Lord Howe Island, S. Pacific.

By Henry F. Carrer, Liverpool School of Tropical

Medicine *.

{ Received September 25, 1920: Read November 16, 1920. }

(Text-figures 1-3.)

The material from which the following descriptions were

drawn up was collected by Mr. R. Douglas Laurie, M.A., of the

Zoological Department, University College, Aberystwyth, in Lord

Howe Island during the summer of 1914. The adult mosquitoes,

a male and a female, were reared by him from larve and pupe

found in a hole containing water in a fallen tree. The second

species, to which mere reference is made, was taken on the wing

in a dwelling-house at night.

OCHLEROTATUS LAUREI, sp. n.

A medium-sized, reddish-brown, mosquito with golden-coloured

lines on the thorax, and the dorsal surface of the abdomen

uniformly dark brown. Proboscis and legs blackish, some or

all of the tarsal segments with small pale bands. Male palpi

relatively short, scarcely more than half the length of the

proboscis.

Head: Integument blackish, the occipital region clothed mainly

with narrow, curved, and upright forked scales, but with small

areas of creamy-white flat scales laterally; curved scales black

and golden, the latter forming a median line, which broadens

posteriorly, and a border (composed of a single row of scales)

round the eyes; upright scales long and thin, black anteriorly

and medially, dark yellowish-brown posteriorly and laterally.

Hyes black. Antenne dark brown, each with the basal segment

and base of the second segment yellowish-brown. Proboscis

straight, long and slender, black; labellee small, pointed, tes-

taceous. Palpi black; in the female very short, each with two

narrow white rings and a white apex ; in the male slender, about

half the length of the proboscis, each with a narrow white band

at the bases of the four segments, terminal segment about two-

thirds the length of the penultimate, no distinct hair-tufts

present. Clypeus black, pruinose. Vhorax : Integument reddish-

brown, clothed with dark brown and golden, narrow, curved scales ;

the golden scales are arranged in a definite manner as follows :—

a narrow border round the anterior edge, two small more or less

circular shoulder-spots, a median line broad anteriorly and

gradually tapering to a point near the middle of the scutum, two

* Communicated by R. D. Laurie, ¥.Z,S,

624 MR. H. F. CARTER ON THE ADULT, LARVAL,

narrow dorso-lateral lines extending parallel from the scutellum

to a spot slightly beyond the centre of the scutum, then becoming

somewhat broader and curving outwards to meet the edge of the

scutum about the anterior third, two very short admedian lines

immediately before the bare space in front of the scutellum, and

lastly, a few scattered scales laterally over the roots of the wings.

Prothoracic lobes dark brown, conical, widely separated dorsally,

with black bristles anteriorly, and intermingled, curved, golden,

and creamy flat scales centrally and posteriorly. Scutelluimn

lighter in colour than the scutum, with sparsely arranged golden,

curved scales; border bristles black. Pleure dark brown with

patches cf creamy-white flat scales. Abdomen: Integument dark

brown. Dorsal surface in the female covered with dark, almost

black, flat scales; in the male similar, but with traces of pale

lateral basal spots (each formed of a few cream-coloured scales)

on the 5th, 6th, and 7th segments; ventral surface in both sexes

dark brown, with a pair of large ereamy-white lateral basal spots

on each segment. ‘Terminal segment of female paler brown, with

moderately long cerci. Wings: Veins rather densely clothed with

dark brown scales, the lateral scales somewhat broadly linear.

Fork cells with their bases almost level, the first cell long and

narrow, about three times as long as the petiole, the second wider,

about twice as long as the petiole; posterior cross-vein about one

and one-half times its own length distant from the median cross-

vein. Legs: Dark brown, blackish. Foreand middle femora each

with a narrow line of cream-coloured scales on the ventral surface,

basal half of each hind femur broadly cream-coloured, apices of

femora with small, yellowish spots. Tibie with inconspicuous

yellowish knee-spots. First and second tarsal segments of the

fore and middle legs with incomplete narrow, basal, whitish

bands, tarsal segments of hind legs with basal, white bands.

Ungues of female all equal and uniserrate; of fore and middle

legs of male unequal and uniserrate, of hind legs of male small,

equal, and uniserrate.

Length: Female approximately 5:5 mm., male 50 mm.; length

of wings of female 5-2 mm., length of wing of male 4°5 mm.

Male Hypopygium (text-fig. 1).—Side-pieces long and narrow—

from three to four times as long as the width at the extreme

base,—with prominent basal and apical lobes. Dorsal surface of

side-piece moderately hairy, the hairs relatively short towards the

base, longer apically and laterally; ventral surface with numerous,

long, curved hairs; outer margins with a few scales. Basal lobes

subquadrate, bearing numerous strong, short hairs; apical lobes

large, extending backwards to the basal lobes, with densely

arranged, short, inwardly projecting hairs. Terminal clasp-seg-

ment slender, comparatively weak, with a long terminal spine

and two short delicate hairs slightly before the apex. Tenth

sternites (harpes) slender, strongly chitinised, slightly recurved at

the tips ; connecting membrane with three small spines on each

AND PUPAL STAGES OF A NEW MOSQUITO. 62.5

side towards the apex. Claspettes (harpagones) large, rather

more than half the length of the side-pieces, each with a conspi-

cuous, attenuated, membranous foot-like appendage. Aldaagus

(unci) separate distally, without serrations or teeth.

Text-figure 1.

Male hypopygium of Ochlerotatus laurei, sp.n. X 120.

(Hairs of portion of right side-piece omitted.)

Larva (text-fig. 2)—Relatively stout, about 7 mm. in length.

Head rounded, slightly narrowed before the eyes; antennz cylin-

drical, sparsely covered with minute spines and bearing four or

five longer spines at the apex; antennal tuft small, centrally

situated, composed of four simple hairs arising from a common

stem. Dorsal hairs not feathered, compound, the anterior pair

each composed of four hairs, the small median of four and the

posterior of five or six; ante-antennal tuft formed of four to six

hairs. Mental plate triangular in shape, armed with from 12-14

teeth on each side; the terminal tooth longer and broader than

the rest, the apical lateral teeth elongate and somewhat crowded

626 MR. H. F. CARTER ON THE ADULT, LARVAL,

Text-figure 2.

\\ wes

\ NN WSs

Head and anal extremity of larva of Ochlerotatus lawrei, sp. n.

AND PUPAL STAGES OF A NEW MOSQUITO. 627

together, the posterior lateral teeth broader (almost triangular

near the base) and more widely separate, the last tooth small

and remote. Thorax wider than long, hairs moderate, feathered,

the stouter, multiple ones each consisting of from 8-10 hairs.

Abdomen somewhat stout. Eighth abdominal segment as shown

in text-fig. 2, the lateral comb composed of about fifty shghtly

chitinised scales arranged to form a triangular area; each comb-

scale is somewhat elongate and is armed apically with a fringe of

spinules. Air-tube about twice as long as broad: pecten com-

posed of some 16 evenly spaced teeth—each pointed and with

one or two lateral projections,—the most distal tooth situated

about half-way along the tube. Hair-tuft of the air-tube arising

immediately beyond the last spine of the pecten and composed of

four or five feathered hairs. Anal segment as long as its greatest

width, the chitinous plate rather small, not completely encircling

the segment; upper pair of papille large, about twice the length

of the anal segment, lower pair smaller, scarcely exceeding the

length of the segment. Dorsal hairs four in number, the upper

pair compound composed of 3-4 hairs each, the lower pair longer

and apparently single; ventral brush well developed, composed

of 14 or 15 compound hairs.

Text-figure 3.

a. Air-trumpet (X 80), and b. Paddle (x 45) of pupa of Ochlerotatus

laurei, sp. Nn.

Pupa: Anterior region (cephalo-thoracic mass) elongate, sub-

pyriform, compressed ventrally and posteriorly ; respiratory

trumpets (text-fig. 3a) conical, their apices obliquely truncate.

Abdomen relatively stout with weak hairs; dorsal, admedian,

pair of multiple hairs of the first segment each composed of about

ten short, branched, hairs; second segment with a pair of smaller

and less complex multiple, admedian, lateral hairs, and also a

pair of longer, single or double, dorso-hairs. Remaining segments

with similar dorso-lateral hairs, the seventh and eighth segments,

628 ON A NEW MOSQUITO.

in addition, with a short compound hair (that on the eighth

segment formed of 10-12 simple hairs) arising from each of the

dorso-antero-lateral angles. Paddles (text-fig. 36) large, ovoid,

each with a short terminal hair.

Mr. Laurie supplied the following data regarding this mos-

quito :—‘* Mosquito larve and pupze from a hollow in a fallen

tree-trunk. Sea-level. The hollow was about 12 ins. x4 ins.

and 4 ins. deep, and contained water. The larve and pupzx were

removed and kept in captivity (7. 14), and two adults emerged

(a male on 18.7.14 and a female on 21.7.14) before we left the

island.”

This species, which I have much pleasure in dedicating to its

collector, is a member of the genus Ochlerotatus in the most

restricted sense; it resembles O. eatoni Kdw. from the Madeira

Is., in regard to the length of the male palpi. In general facies

it somewhat resembles O. japonicus Theo., and apparently is

related to the Asiatic group of species formed by the last-named

mosquito, O. macfarlanei Kdw., O. pulcriventer Giles, O. oreophilus

Kdw., etc.

Five examples of a second and probably undescribed species of

Ochlerotatus were also captured by Mr. Laurie in a dwelling-house

on the island. This species is dull brown in colour, and, with

the exception of yellowish lateral abdominal spots, is devoid of

ornamentation. Unfortunately, the specimens obtained were all

females, and therefore until more material is available it is

advisable to refrain from attempting an exact diagnosis.

References.

Epwarps, F, W.—‘ Hight new Mosquitoes in the British Museum

Collection.” Bull. Ent. Res. vi. p. 357 (1915-

LONG):

35 as ‘* Notes on Culicidee, with descriptions of new

Species.” Bull. Ent. Res. vi. p. 201 (1916—-

US)

Liverpool, July 1917.

THE LIFE-HISTORY OF THE YELLOW DUNG-FLY. 629

34. The Jife-History and Habits of the Yellow Dung-Fly

(Scatophaga stercoraria) ; a possible Blow-Fly Check.

By G. 8. Correrett. With a Preface by Prof. Max-

WELL LEFROY, F.Z.S.*

(Text-figures 1-14.)

[Received October 19, 1920: Read October 19, 1920.}

PREFACE.

This paper is an account of work undertaken to complete an

inquiry into Blow-fly control, of which a partial account has

already been published by Mr. A. M. Altson in the Proceedings of

the Zoological Society, 1920, p. 195. Of the three important

controls of Blow-fly in England, Mr. Altson has already dealt with

the two common parasites: the present paper deals with what we

believe to be the most important direct enemy of the adult fly,

a check which appears to be very effective in this country. The

Yellow Dung-fly first showed itself in our work at the Zoological

Society in 1915 in connection with methods of trapping flies: it

came in numbers, persistently eating the adult Blow-flies, and

seriously interfered with experiments out of doors. My observa-

tions since show that while the fly preys on a large variety of

Diptera, it specially attacks Calliphora and Musca. It is a

constant and general feeder on the common species of Blow-fly in

England throughout the season.

The author of this paper undertook the investigation, and

submitted this paper as a thesis for the Diploma of the Imperial

College: he also investigated the best means of transporting this

species to countries where Blow-fly is a serious pest to sheep, in

the hope that it might be possible to utilise it as a check on

Blow-fly. This has not been possible as yet, but the species

seems to have much value in this connection, and its habits as a

maggot and an adult are so harmless that it 1s to be hoped it

will eventually be made use of —H. M. Lerroy.

Introduction.

The study of the life-history and habits of the Yellow Dung-

fly was first commenced in October 1919, at a time when the

Blow-fly problem was receiving a great deal of attention and

the discovery of an etticient control was sought for.

In conjunction with other methods of control then under

investigation, that of the Yellow Dung-fly, as a predator on the

Blow-fly, was also studied.

The adult fly was identified for me by Mr. Edwards, of the

Natural History Museum, South Kensington, as Scatophaga

* Communicated by Prof. Maxwett Lrerroy, F.ZS.

630 MR. G. S. COfTERELL ON THE LIFE-HISTORY

stercoraria L., belonging to the family Cordyluride of the

Acalyptrate Diptera.

A description of the species is given in a monograph by

Becker, which is at present the standard work on the classification

of the family Cordyluridee.

Both sexes vary from very large and robust flies to small

varieties. The largest measures 15 mm. in length and the

smallest 8 mm. The span of the wings is over double the length.

The head is globular; the eyes oval, brown, and separated in

both sexes by an area equal to half the width of the head. The

frontal stripe is rich yellowish brown in the male, dull yellow in

the female. The frontal margins of the orbits, cheeks, and face

are yellow. The facial bristles are strong; antenna black, and

the arista bare except for the upper third, which is feathered.

The thorax of the male is marked with longitudinal stripes on

the dorsal surface. There is a bunch of light yellow hairs under

each wing. The thcerax of the female is coloured darker, the

stripes more marked, and the hairs below the wings are absent.

Both sexes have more than two rows of acrostical bristles, and

well-marked scutellar bristles.

The wings are slightly yellow; anterior cross-veins very

distinct, with a smoky coloration round them.

The femora are covered with long yellow hair in the male,

especially the anterior pair; other parts yellow but not hairy.

In the female the femora are dark and clothed with a few dark

hairs.

The abdomen of the male is yellow and very hairy. In the

female yellow-green and not hairy. The male abdomen is

cylindrical, terminating bluntly. There is a dark area on the

ventral surface of the fifth segment marking the entrance to the

male genital atrium. In the female it is short, broad at the base

and conical, becoming more or less oval when gravid.

The female is smaller than the male and darker. owing to the

absence of the yellow hairs. The legs of both sexes are bristly

and the pulvilli well developed.

The sexes are distinguished by the size, difference in colour,

shape of the abdomen, and the black area on the ventral surface of

the abdomen of the male.

There are nineteen British species of the genus, but I have

never observed any other breeding in dung.

S. merdaria is the nearest related species, but is distinguished

by both sexes being duty green in colour and by there being only

two rows of dorso-central bristles on the thorax.

S. seybaiaria is distinguished by the colour of the third antennal

joint, which is reddish brown.

Scatophaga stercoraria is widely distributed. It occurs as far

north as Nova Zembla and Siberia, and as far south as North

and South Africa and the Canary Islands. It is common

throughout Europe and is found in Asia Minor. I have seen no

mention of it occurring in India or Austyalia.

It is evidently both'a temperate and sub-tropical species.

AND HABITS OF THE YELLOW DUNG-FLY. 631

Adult Feeding.

Both sexes of the adult fly ave predaceous on other Diptera.

The prey is never caught on the wing, but usually at the moment

of settling. The captured fly is grasped firmly by the middle and

hind legs, the bristles of the legs helping to make a firmer grip.

The prey is caught in such a fashion as to bring both flies in an

upright position and the heads one above the other. In this

position the wings of the victim are useless, its legs being the

only active part. The proboscis is then pressed against the neck

and a puncture made through both sides. Attacked flies may all

Text-figure 1.

Scatophaga stercoraria, g, attacking Musca domestica.

be seen to have the neck stretched and a distinct hole right

through it. In this way the nerve-cord is either cut or damaged,

causing a partial paralysis. The contents of the thorax are then

sucked out, at the junction of the neck, as far as the proboscis

will reach. A large amount of saliva is secreted during these

operations, probably acting as a solvent. The head is then turned

round by means of the front legs, bringing the oral margin upper-

most. Another puncture is made inside the oral margin, and the

contents of the head, including the eye-pigment, are also sucked

out. Access to the other parts of the thorax is obtained through

632 MR. G. 8S. COLTERELL ON THE LIFE-HISTORY

the thinly-chitinised membranes between the coxe and thorax.

The fly is then turned completely round, bringing the ventral

surface of the abdomen uppermost and the head in a posterior

position. The contents of the abdomen are then sucked out

through punctures between the sternites.

This order of feeding is invariably carried out, but, when food

is abundant, the parts are only partially sucked out, the abdomen

often not being touched.

Attraction for more food from one victim apparently ceases

when struggling has ceased.

Both sexes are very strong in flight and are capable of flying

short distances, grasping flies as large as themselves.

Adult Mouth-parts.

On account of its predaceous habits the mouth-parts are

modified accordingly.

Externally the proboscis is of the Muscid type, ¢.¢. the ordinary

labella consisting of the fused inner lobes of the labium, the

pseudo-trachee, the labrum-epipharynx, hypopharynx, and maxil-

lary palpi. These structures are identical with those found in

Musca domestica or Calliphora erythrocephala, in which the pro-

boseis is adapted for licking. The only modifications due to its

predaceous habit are found in the internal chitinised structures

of the labellum, or oral disc, and haustellum.

‘The structure of the rostrum, or proximal portion of the

proboscis, is identical with that of C. erythrocephala as described

by Lowne. The main chitinised structure is the fulcrum (text-

fig. 2,a) enclosing the pharynx. This has been compared by

Kraepelin to a Spanish stirrup-iron with a double foot-plate, the

foot-plate being posterior and the toe at the lower end. The

whole structure of the rostrum resembles that of a truncated

cone with the apex downwards.

The haustellum or arm of the proboscis is cylindrical in shape.

Proximally it is attached to the lower end of the rostrum, and

distally to the oral disc. It is enclosed on its lateral and

ventral sides by a convex sclerite, the theca (text-fig. 2,7). This

articulates proximally with the fulcrum and distally with the

furca (text-fig. 2,9), a tri-radiate sclerite forming the chief

internal skeletal structure of the oral disc.

The labrum-epipharynx (text-fig. 2,d@) and hypopharynx (text-

fig. 2,e), enclosing the salivary duct, lie over the dorsal portion of

the haustellum as in the House-fly.

The oral dise consists of two lobes united posteriorly by a ball-

and-socket joint, each grooved on its oral surface by a number of

pseudotrachez (text-fig. 2,4). The oral aperture is situated

between the lobes at their point of junction. A space below the

mouth is kept open by a pair of sclerites, the discal sclerites

(text-fig. 2,4), deeply embedded in the oral disc. These sclerites

AND HABITS OF THE YELLOW DUNG-FLY. 633

are united posteriorly, forming a U-shaped structure. This space

is the oral pit, and the common trunks of the pseudotracheze

open into this.

Up to this point the structure, except for minor details, is

identical with that of the Blow-fly or House-fly. In addition,

however, the haustellum is strengthened by a pair of long,

chitinised rods (text-fig. 2, /), articulating proximally with the

fulerum and distally with the discal sclerites. These occur also

in the House-fly and Blow-fly, but are only very slender rods.

Text-figure 2.

Mouth-parts of adult:—a, fulcrum; d, labrum-epipharynx; e, hypopharynx;

Ff, rods; g, furca; h, discal sclerite; i, teeth; 7, theca; &, pseudotrachez.

They play an important part in the articulation of the oral

dise. In this case the oral disc is extremely mobile and used for

vasping as well as sucking, therefore the rods are more strongly

developed. These reds are called paraphyses by Lowne in his

description of the Blow-fly.

The paraphyses articulate distally with the discal sclerite : thus,

there are two articulation points between the haustellum and

oral dise—the first between the theca and furea, and the second

between the paraphyses and the discal sclerite.

To each arm of the discal sclerite a set of teeth is attached

Proc. Zoor, Soc,—1920, No, X LIT, 42

634 MR. G. 8. COTTERELL ON THE LIFE-HISTORY

(text-fig. 2,7). Hach set consists of five teeth, the middle one

being the most prominent. They are placed on each side of the

oral aperture between the two lobes of the oral disc. They pro-

bably correspond to the teeth in the House-fly, forming the lateval

edges of the gutters or continuations of the common trunks of

the united pseudotrache described by Graham-Smith. There

are three rows on each side of the oral pit in the House-fly, but

only one pair on each in Scatophaga. They function to a very

small extent for rasping in the House-fly, and in Scatophaga they

exist essentially for rasping. If this point is correct, the mouth-

parts are identical with those of the sucking Muscids, except for

certain modifications of the chitinised structures, particularly of

the paraphyses and teeth, in accordance with its predaceous habits.

The method of sucking is similar to that of the Blow-fly or

House-fly. The liquid food is sucked up through the pseudo-

trachez into the oral pit and so into the mouth.

By the alternate upward and downward movement of the

paraphyses the oral dise is worked in a backward and forward

direction, and, consequently, a rasping is caused by the teeth on

any surface they are in contact with. In this manner the

thinly-chitinised parts of other Diptera are easily punctured.

The internal tissues are broken down by further rasping, and

apparently partially dissolved by the large amount of saliva

secreted. The liquid food is then sucked up through the pseudo-

trachee.

The theca serves not only as an external skeleton to the

haustellum, but also protects it from whatever damage it might

incur from coming in contact with the jagged edges of chitin

when inserted in a hole in its victim.

Breeding Media.

The breeding media consist entirely of excrement providing a

sufficient consistency for the larva to complete its life without it

becoming dry. This includes human excrement, poultry, sheep

and cattle excrement, and horse excrement. The last-mentioned

is very rarely used for oviposition. Sheep and cattle excrement

is preferred chiefly on account of its viscosity and the amount

obtainable in pastures. Cattle excrement was used in the

breeding-jars in the laboratory.

(At the Royal Naval Cordite Factory at Holton Heath a large

mass of sludge accumulated from septic sewage tanks: this sludge

was in the open, and was about the consistency of fresh cow-dung;

it contained an enormous number of this fly in all stages of

development, and formed a very suitable breeding medium.

I believed that the marked absence of House-flies and Blow-flies

at Holton Heath was in part due to this, but I had no direct

evidence.—H. M. L.)

AND HABITS OF 'HE YELLOW DUNG-FLY. 635

Female Genitalia and Method of Oviposition.

As seen externally, the apex of the abdomen of the female ends

in two setiferous lobes (text-fig. 3, a), between which the anus

opens. Below this again is the retractile ovipositor (text-fig. 3, 5).

This consists of a pair of chitinised blades attached, proximally,

to 1 membranous tube. The whole of this can be retracted within

theabdomen. The blades are pointed and, when placed together,

form a groove. The common oviduct, which is very large, opens

into it.

When oviposition is about to take place, the abdomen is

stretched out horizontally with the ovipositor extruded. In this

position an egg is passed down into the groove formed by the

juxtaposition of the two blades of the ovipositor. Ovipositor

and egg are then pressed down into the dung obliquely. The

blades are pulled apart and the egg released.

Text-figure 3.

Abdomen of female S. stercoraria, with genitalia extended.

The eggs are more or less scattered over the surface of the

dung, but ten or fifteen or more may be deposited in one small

area, particularly where there is a crack or a crevice where the

softer parts of the dung are exposed.

Fresh dung is preferred by the females for oviposition, but

occasionally eggs are laid in dung a week or more old. In this

case the crust on tke surface of the dung is too hard for the evi-

positor to be inserted deeply, and eggs are often seen only half

buried. This does not prevent the egg from hatching, but the

larva usually has difficulty in getting through the hardened crust

of the dung.

Oviposition occasionally takes place with the male in siéw, in

which case the male removes his abdomen from that of the

female, and, instead of clasping her with his two front pairs of

legs, drops back clasping her only with the front pair. The

female then can manipulate her ahdomen freely whilst the male

follows her about.

When a female is gravid, the abdomen is so distended as to

make her practically incapable of flight. The abdomen is re-

duced to nearly a fourth its size when oviposition has taken

place,

42"

636 MR. G. S. COTTERELL ON THE LIFE-HISTORY

The Lgq.

The egg is creamy-white when first laid, becoming darker as

the incubation period increases. It measures from 2 to 3 mm.

in length and is slightly curved.

It is especially characteristic in having two wing-like exten-

sions of the chorion at its anterior end (text-figs. 4 & 5, a).

These are covered with extremely short sets on their under

surfaces, and serve as a support in the dung. The egg is laid

obliquely at an angle of from 15° to 25° with the surface of the

dung, so that the two extensions lie flat on the surface. The

micropyle (text-figs. 4 & 5, c) is situated between the ex-

tensions, and is drawn out into a short crest (text-figs. 4 & 5, b)

at the anterior end between the two extensions. Its surface is

finely sculptured. The two extensions and the micropyle are all

Text-figure 4, Text-figure 5.

Text-fig. 4 Hee of S. stercoraria, showing supporting wings.

Text-fig. 5.—Hatched egy of S. stercoraria.

exposed on the surface. The wall at the base of the egg (text-

fig. 5, e) is thickened, giving greater strength, as this is the part

of the egg which first comes into contact with the dung when

oviposition is taking place. The egg is broken by means of a

transverse split behind the micropyle (text-fig. 5).

Tn summer the whole surface of a deposit of cow-dung may

be seen covered with the wing-like extensions of the eggs. The

incubation period varies from one to two days according to tem-

perature. This period is much more constant than other periods

in the life-history, as temperature is the only governing factor.

The newly-hatched larva breaks the egg behind the micropyle,

erawls out on to the surface of the dung, and seeks a convenient

erack by which to gain access to the softer parts of the dung.

This may take some considerable time if the dung has been de-

posited for some time, and consequently become dry on the surface,

AND HABITS OF THE YELLOW DUNG-FLY. 637

A number of newly hatched larve perish through not getting in

before they starve.

The Larva.—Wirst Instar.

The larva immediately after hatching measures from 2 to

3mm. in length. The body is cylindrical, composed of twelve

seoments, and generally tapers to a point at the anterior end.

The posterior end is truncate, forming the anal plate. In this

stage the larva is metapneustic, the two spiracles being placed

side by side on the anal plate. Each consists of a chitinised ring,

situated on a pair of short projections from the anal plate, en-

closing two slit-like apertures. ‘The anal plate is bordered by a

number of tubercles, the position of which will be described in

a later instar.

The first four segments are devoid of spines. Very delicate

spines occur on the anterior border of each segment from the

fifth posteriorly. The area covered on each segment increases

proceeding posteriorly, the whole surface of the last two segments

being uniformly covered.

Text-figure 6.

Larva of S. stercoraria. 1st instar.

The first segment is divided dorso-ventrally by a cleft forming

the two oral lobes (text-fig. 6,2). Hach lobe carries a pair of

sense tubercles (text-fig. 6,7). In addition, the ventral surface

of the first segment is provided with a transverse row of strong

backwardly-curved spines (text-fig. 6, ¢). These are locomotory

in function. Keilin mentions them as being present in the

first instar larva of Musca assimilis. Below the spines there are

two channels running more or less parallel with each other. They

originate at the mouth, and run out laterally over the ventral

surface of the first segment. They are fringed with hair, and

direct the liquid food towards the mouth (text-fig. 6, /).

638 MR. G. S. COTTERELL ON THE LIFE-HISTORY

The chitinised mouth-parts are slender. Commencing pos-

teriorly there are a pair of U-shaped sclerites (text-fig. 6, @) with

the arms directed backwards. The anterior end of each is ex-

tended (text-tig. 6, 6) for articulation with a large median sclerite

(text-fig. 6, c). The posterior sclerites, which correspond to the

lateral pharyngeal sclerites of the full-grown larva, are united

dorso-laterally by a chitinised band (text-fig. 6, ). A pair of

sclerites are also found embedded in the first segment (text-

fig. 6, d). These probably correspond to the buccal sclerites or

hooks of the full-grown larva. They articulate at their bases

with the median sclerite and give rigidity to the oral lobes.

The hooks of the second instar appear behind those of the first

a few hours before ecdysis, and become functional as locomotory

organs (text-fig. 7).

Text-figure 7.

Larva of S. séercoraria. 1st and 2nd instars.

The anterior and posterior spiracles and the remainder of the

mouth-parts of the second instar do not appear until just before

the moult. An entirely new set of mouth-parts can be seen in a

preparation made an hour or so before the moult.

Kedysis takes place on the first or second day after hatching,

the instar lasting from one to two days.

During this instar the larva is very active. In the jars used in

the laboratory, occasionally the larvee hatched out into too liquid

dung to allow of a sufficient supply of air below the surface.

They were observed to be hanging on to the surface-film with

their posterior spiracles exposed, disappearing on being disturbed,

reappearing later in the same place. On one occasion the dung

remained sufficiently moist to necessitate this during the first two

instars. It was only when the third instar was reached that a

sufficient air-supply was obtainable below the surface. Probably

under natural conditions, where the dung is exposed to wind and

AND HABITS OP THE YELLOW DUNG-FLY. 639

other drying factors, it would dry up to a sufticient consistency

for the lanes to commence living a normal existence buried in the

dung itself.

The Larva.—NSecond Instar.

After the first ecdysis the larva increased considerably in size.

It measures from four to six millimetres in length. The general

shape of the body and the number of secments | remain constant

throughout larval life. It is now amphipneustic, being provided

with an anterior pair of spiracles (text-fig. 8, 7) in addition to

the posterior pair. The anterior spiracles are situated at the

junction of the second and third segments, and will be described

in detail in the following instar. The posterior pair are identical

with those described in the preceding instar.

Text-figure 8.

Larva of S. stercoraria. 2nd instar.

The locomotory spines on the first segment in the previous

instar have now disappeared. Spines similar to those of the first

instar but proportionately larger occur on each segment behind

the fourth. They are concentrated on the anterior border of the

fifth, sixth, and seventh segments, but the area covered increases

posteriorly. The last few segments are uniformly covered.

The first segment is similar to that described previously, but

the locomotory spines are absent. Also the two parallel channels

converging into the mouth are now replaced by a number of

channels radiating from the mouth over the ventral surface of the

oral lobes (text- fig. 8 sige

The chitinised mouth- -parts have no similarity witn those

640 MR. G. 8. COTTERELL ON THE LIFE-HISTORY

described previously, except for the structure of the lateral

pharyngeal plates (text-fig. 8, a@). These aresimilar in structure,

but are more strongly chitinised and longer in proportion to their

breadth. They are united dorsally by a chitinised band, and ven-

trally by a thinly-chitinised membrane forming the floor of the

pharynx (text-fig. 8, 7). The median piece is replaced by a pair

of chitinised rods, the intermediary sclerites (text-fig. 8, 6), which

are united transversely by a bar of chitin. Dorsal to each inter-

mediary sclerite and lying close up to each there is a slender

chitinised rod.

The buccal sclerites are elongate and spoon-shaped (text-

fig. 8,d & text-fig. 9). The exterior lateral edge of each bears four

teeth. Ventrally, and at the base of the hooks, there are a pair of

short chitinised rods (text-fig. 8, m) which support the oral

opening.

The buccal sclerites of the third instar appear dorsally to

those of the second instar two days before ecdysis, becoming fully

chitinised and functional one day before ecdysis. The remainder

of the mouth-parts do not become fully chitinised until a few

hours before ecdysis. A complete second set of mouth-parts was

never definitely observed at this moult, but as all the mouth-parts

of the first instar are thrown off, the same may be taken for

granted to occur here. The new anterior and posterior spiracles

can also be seen underlying those of the present instar.

The second ecdysis takes place on the third and fourth day of

larval life, this period lasting from 36 hours to 8 days.

The Larva.—Third Instar.

In the third and final larval instar the larva increases greatly

in size, measuring, when full-grown and fully-extended, from one

to one and a half centimetres in length. The general external

shape is similar to that described in the preceding instars.

The anal plate (text-fig. 11) is surrounded by twelve tubercles.

The two largest are situated further forward than the rest on the

ventral surface of the last segment immediately behind the anus.

The remainder are situated on the border of the anal plate, eight

laterally (four on each side) and two dorsally. In addition, there

are three situated on the anal plate itself below the spiracles.

Spines (text-fig. 12) oceur on each segment from the fifth seg-

ment posteriorly, concentrated on the anterior border. The last

few segments are uniformly covered. <A belt of spines also occurs

round the middle of the fourth segment. The anterior borders

of the four anterior segments bear a number of small projecting

plates (text-fig. 13) arranged ina number of concentric rings round

the segment. Each plate lies against the side of the body, but

is hinged anteriorly and can be pushed away from the body.

These help to give the first segments a grip when pushed into

the dung, but they can also be pulled in against the side -of the

body if the larva wishes to withdraw its anterior segments. They

AND HABITS OF THE YELLOW DUNG-FLY. 641

are to be contrasted with the spines covering the remainder of

the body, which also help to give the segments a grip on the sides

of the larval burrow, but, being fixed, prevent a backward move-

ment. If the anterior segments were covered with these, the

Text-figure 9.

Larva of S. stercoraria. 3rd instar.

Text-figure 10.

Posterior spiracles. 3rd instar.

larva, once having pushed its head in one direction, would be

obliged to continue whether it wanted to or not. The hinged

plates give it a chance to make a second investigation in another

direction provided that it has not gone too far.

642 MR. G. S. COLTERELL ON THE LIFE-HISTORY

- The first segment is divided into two oral lobes with radiating

food-channels on their ventral surfaces as in the preceding instar.

The chitinised mouth-parts are similar in general structure to

those of the second instar, but are proportionately stronger. The

lateral pharyngeal sclerites are longer and more deeply cleft

(text-fig. 9, a). The intermediary sclerites are much thickened

and shortened. The buccal sclerites (text-fig. 9,@) have no

similarity with those of the second instar. They have lost the

spoon-shaped structure, and are now a pair of stout ventrally

eurved hooks. They are pointed anteriorly and thickened where

they articulate with the intermediary sclerites. They project

externally over the oral opening,

The anterior spiracles (text-fig. 9) are situated laterally at the

junction of the second and third segments. They consist of a

short chitinised trunk, projecting forwards and externally from

the junction of the segments. The trunk divides into two lobes,

each lobe bearing usually eight papille, making sixteen in all.

Each papilla is pierced by a small lumen for the ingress of air.

The number of papillee varies from sixteen to eighteen in different

larvee.

The posterior spiracles (text-fig. 10) are situated side by side in

the middle of the anal plate. Hach consists of a chitinised ring,

situated on a short projection, enclosing three slit-like apertures.

The apertures are bordered by inwardly projecting filaments

serving as a sieve for the incoming air. These spiracles are the

most necessary to the larva, being those it keeps above the surface-

film when living in very liquid dung. The respective anterior

and posterior spiracles are connected by two lateral longitudinal

tracheal trunks. The lateral trunks are connected, soon after

their origins from the posterior spiracles, by a transverse trunk.

In addition the longitudinal trunks give off small branches in

each segment.

The anus is situated on the triangular plate on the ventral

surface of the last segment. This plate is devoid of spines, and

appears to be glandular in structure.

The third instar period takes from 6 to 9 days, pupation

taking place on the ninth to twelth day of larval life.

The larva is active only during the first two days of this instar,

after which it seeks the drier parts of the dung or the soil,

preferably the latter. During the inactive period it changes

from a more or less transparent appearance to an opaque one,

due to the great develepment of the fat-body.

Great dithculty was experienced in determining whether the

whole of the chitinised mouth-parts were thrown off at ecdysis.

Owing to the thin epidermis and the impossibility of finding cast

skins in the dung, a set of old mouth-parts was never found.

Theoretically the complete set should be thrown off, and recently

I had the good fortune to mount a preparation within an hour

or so of the first moult showing a complete second set outside

the old set. Up to this time I had proof that the buccal and

AND HABITS OF THE YELLOW DUNG-FLY. 643

Text-figure 11.

Schematic drawing of anal plate.

Text-figure 12.

Spines of fitth and later segments.

Text-figure 13.

Modified spines of anterior four segments.

644 MR. G. 8S. COTTERELL ON THE LIFE-HISTORY

intermediary sclerites were thrown off, but it seemed impossible

that the larva could be capable of throwing off the large lateral

pharyngeal plates. No conclusive proof was obtained of the

complete throwing-off of the second instar mouth-parts at the

second moult, but this may be taken for granted as occurring as

well as at the first moult.

The total larval life takes from 9 to 12 days. The minimum

time observed was 8 days and the maximum 14 days. Eleven

days is the usual length of time under laboratory conditions.

The total larval life and the ultimate size of the larva, and

therefore the puparium, are controlled to a very large extent by

the temperature and condition of the breeding medium. Lack of

moisture is a very important factor: This explains the variety

in size of the adults.

The Puparium.

At pupation the two anterior segments of the larva are with-

drawn, bringing the anterior spiracles in a forward position. In

this position the larval skin slowly changes to the brick-red

colour characteristic of the puparium (text-fig. 14). The colour

changes to black as the pupal period advances. It is more or

less cigar-shaped; the anterior end is slightly flattened dorso-

ventrally. The anterior spiracles can be seen as two dark

projections at the anterior end. It measures from 6 mm. to over’

a centimetre in length. The size varies according to the con-

ditions governing the life of the larva, mentioned earlier.

From larve bred in the laboratory a large percentage pupated

in the hardened upper crust of the dung. This was probably due

to the soil being too moist in the jars, as there was no outlet

to allow the excess moisture to get away. Under ordinary

conditions the soil is preferred, as in field observations very few

puparia were observed in dung.

The pupal period takes from 10 to 17 days. The minimum

time observed was 6 days and the maximum 18 days.

The following data give the comparative larval and pupal

periods for a number of batches bred :—

Oviposition. Keg. i: instar | II. instar. | IIT. instar.| Pupa. Total. |Hmergence.

' Jan. 5th. | 48-54 hrs.| 36 hrs. | 36-60 hrs.| 6-7 days. 6 days. 17-19 days.|Jan. 22-24.

Feb. 17th. | 20-24 hrs.| 36 hrs. | 60 hrs. 6 days. |13-14 days.|24-25 days.|Mar. 12-13.

Mar. Ist. | 36-42 hrs.| 36 hrs. | 36hrs. | 6-7 days. | 9-10 days. 19-21 days. Mar. 20-22.

Mar. 18th. 48 hrs. | 36 hrs. | 36 hrs. 9 days. |16-17 days. 30-31 days.|Apr. 17-18.

May 7th. 48 hrs. | 36 hrs. | 48 hrs. | 7-8 days. 16 days. 28-30 days. June 4-6.

AND HABITS OF THE YELLOW DUNG-FLY. 645

The minimum time from adult to adult observed was 17 days

and the maximum 31 days. The average time from the table

given above is 24 days, but this time, as stated earlier, is governed

by temperature and the consistency of breeding media.

In one batch of larve bred in dry dung the larve were

observed to have collected in one spot where the only moisture

was. These did not reach the third instar until after a week of

larval life. They were transferred to fresh dung later, so that

the total larval life was not observed. Under similar conditions

the larval life may take as much as three times as long as the

maximum observed.

Text-figure 14.

Puparium.

Sexual maturity is not reached until after 21 days of adult life.

During this time the males have no attraction towards the

females, and, owing to the male being the more robust, the latter

are liable to be attacked unless a sufficient supply of food is

maintained. .

Copulation takes place about two days before oviposition. The

life of the male is considerably longer than that of the female.

No definite results were obtained on the life of the female. No

females were observed to lay more than 120 eggs. It is pro-

bable that one female is capable of laying from 100 to 150

eggs and then dies. Eggs are laid in one batch of from 40 to 80

and afterwards, 10 or 20 at a time at intervals.

The total life-cycle from egg to egg takes from six to seven

weeks.

646 MR. G. S. COTTERELL ON THE LIFE-HISTORY

Breeding takes place regularly from April to October, and

there are probably five broods a year.

No definite results were obtained on hibernation, but evidently

a large number hibernate as adults. A number of pupz were

exposed to cold weather during December, but the adults all

emerged without exception. This implies that hibernation does

not take place in the pupal stage, but it may be that the process

of histogenesis had proceeded beyond a certain point before being

placed in the cold, and the breaking down and building up of

tissues in the pupz were carried on. A large amount of adults

are seen about on warm sunny days in winter, but 90 per cent,

are males. I think that probably the majority hibernate as

adults, but that only those females survive that failed to ovi-

posit before the cold weather set in, This would account for the

difference in proportion of the sexes. Probably a few survive

the winter as puparia and the larger proportion as adults.

Investigations extended over another winter would prove this

point. Graham-Smith remarks that S. stercoraria hibernates as

a resting larva or pupa in the soil, a few surviving the winter

as adults.

A number of flies were placed in a glass-house heated by two

electric radiators. During one night the temperature rose to

over 90° F., with the result that they were all found dead the

following morning. It appears from this that the fly is not

capable of withstanding high temperatures. The fly is, however,

a subtropical one as well as a temperate one, and should there-

fore be able to stand this temperature. The sudden change from

a mild temperature to a hot one and the lack of ventilation were

probably the cause. It must also be remarked that the flies

were of a very large variety. ‘The largest and smallest varieties

are not so resistant to unfavourable conditions as the average

sized varieties.

The food of the adults is very varied, but confined to other

Diptera. The small Borborid fly (Borborus equinus) appears to

be the chief article of diet in the field, chiefly as it breeds

abundantly in horse excrement and as it passes the winter as an

adult. Larger flies, however, are preyed upon, such as Calliphora,

Lucilia, M. domestica, etc. Probably all species of Diptera are

preyed upon, with the exception of those of fast flight, such as

Syrphide and Stratiomyide. I observed, on one occasion, a

large male attempt to capture a small dung-beetle (Geotrupes)

as the latter was settling. At the Zoological Gardens in 1915

Professor Lefroy’s experiments with fly-traps were interfered

with by the abundance of the adult S. stercoraria that fed on the

trapped flies, chiefly blow-flies of the genus Calliphora.

In summer the adults may be seen on plants and flowers far

away from pastures waiting for other flies to settle in the vicinity.

M. domestica was used as food almost entirely in the laboratory,

each fly sucking out as many as a dozen a day.

AND HABITS OF THE YELLOW DUNG-FLY. 647

Natural Hnemies.

Natural enemies were not observed. he Scarabeid and

Staphylinid beetles and their larvee, which breed in dung, do not

appear to attack the larvee of Scatophaga.

One Ichneumonid parasite was bred out from pup, but, having

failed to reach the surface of the soil, was in too bad a state of

preservation to be identified.

A large number of birds are mentioned by Newstead as feeding

on Searabeid beetles and Muscid larvee breeding in dung, so that

they very probably are a considerable check on the spread of

Scatophaga.

A number of larve are probably destroyed by natural con-

ditions such as the too rapid drying up of the dung or owing to

the dung being trodden on and spread about by cattle.

Bibliography.

1. Becker, T.—Dipterologische Studien. I. Scatomyzide.

Berlin. Ent. Zeit. 1894, pp. 77-190.

2. Becker & Brzz1.—Katalog der Paliarktischen Dipteren.

3. GRAHAM-SmItTH, G. 8.—-Flies in Relation to Disease: non-

blood-sucking flies. Cambridge (Public Health Series),

1913. (Pp. xiv+292.) 24 pls.

. —— Observations on the Habits and Parasites of Common

Flies. Parasitology, viii. (1915-16), pp. 441-547.

. Hewrrr, C. G.—The House-Fly.—Biological Series, No. 1.

Manchester University Press.

—— Journal of Economic Biology, 1907.

Canadian Entomologist, xlvi. (1914), p. 2.

Keinin, D.—Recherches sur les Anthomyides 4 Larves

Carnivores. Parasitology, ix. (1916-17).

. Lowne, B. T.—The Anatomy, Physiology, Morphology, and

Development of the Blow-fly. Vols. I. & II.

10. Mentor, J. E. M.—Observations on the Habits of certain

Flies. Annals of Applied Biology, vol. vi. (Sept. 1919).

11. Newstreap, R.—The Food of some British Birds. Journ. of

the Board of Agriculture, vol. xv. No. 9 (Dec. 1908).

12. Parron & Crace.—Text-book of Medical Entomology.

13. Pouxtron, E. B.—Predaceous Insects and their Prey. Pro-

ceedings of the Entomological Society, 1906.

14. Witurston, 8. W.—Manual of North American Diptera,

(1908.) James F. Hathaway, New Haven,

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ON TILE RESPIRATORY MOVEMENTS OF NECTURUS, 649

35. Remarks onthe Respiratory Movements of Neeturus and

Cryptobranchus. By A. Wiuuny, M.A., D.Se., F.RS.,

BLS!

[Received Octoher 25, 1920: Read November 2, 1920.]

In a former paper certain facts were published showing that,

me the conditions of observation, Vecturus, the “ Mud Puppy”

r “ Gilled Salamander,” is normally a water-breather, not an

air-breather * Shortly afterwards I had the opportunity (May

24 and 25, 1918) of observing Vectwrus under the conditions

presented to it in the New York Aquarium, and of comparing its

behaviour with that of the ‘“hellbender,” Cryptobr whee

| =IMenopome |.

Between 11 4M.and noon on May 24, after watching for ten

minutes, [ saw one Cryptobranchus yawn slowly to the full

extent of its gape under water at the bottom of the deep

aquarium. Two others afterwards went thr ough the same

performance, 7. e. yawning under water. There were more than

a dozen individuals, all resting at the bottom of the aquarium

during my forty-minute vigil. One of them gave two rather

copious emissions of air from the month, and a bubble was seen to

vise from the right gill-pore. None visited the surface. While

at the bottom they have a way of swaying gently from side to

side, whereby the longitudinal lateral cutaneous flap waves up

and down.

At the same hour on the following day I found the hellbenders

actively engaged in ascending to the surface, remaining for some

minutes just below the surface suspended in the water with

arched back, retaining that attitude whilst sinking to the bottom,

or else deliberately swimming to the bottom. They protruded

their muzzles: above the surface without opening the mouth, and

in one case a deep inspiration through the nostrils was effected by

the dilatation of the hyobranchial apparatus, The air thus taken

in may be expelled quickly from the mouth in a continuous

stream of bubbles, perhaps indicatine that it had been used for

oropharyngeal respiration.

Giant Salamanders (C. maximus), in a neighbouring tank,

behaved like the hellbenders. On May 24 they were Ponti: at

the bottom of the deep aquarium ; one of them emitted some air

through the mouth; none visited the surface. But on the

following day they were restless. ascending to the surface. T saw

one of them, after pushing its muzzle above the surface, take a

slow and deep inspiration through the nostrils by the dilatation,

to full capacity, of the byobranchial apparatus. The latter

* A. Willey: “ Branchioderma and Branchiotrema.” Trans. Roy. Soc, Canada,

3rd ser. vol. xii. sect. iv. (May 1918), pp. 95-104. Ottawa, 1919.

Proc. Zoou. Soc.—1920, No. XLITI. 43

650 DR. A. WILLEY ON THE RESPIRATORY

bulges conspicuously downwards and backwards so that the

greatest convexity lies behind, at the base of the throat.

Presumably, in this act of dilation, the glottis is opened; and, in

fact, Cryptobranchus seems to show some increase of girth after

inhalation.

Inhalation through the nostrils was observed by G. B. Wilder

in a Japanese Cryptobranchus, 25 feet long, which he kept in

shallow water sufficient to cover it *. The downward protrusion

of the floor of the mouth in narial inspiration, the arching of the

back, and the swaying movement at the bottom have been

described by A. M. Reeser, who adds that the hellbender will

survive more than a week’s exposure out of water without

suffering any permanent diminution of vitality. The longest time

that any individual was observed by him to remain under water

was 43 minutes. B. G. Smith2 confirmed the observation that

the arching of the back is a sign that air has entered the lungs.

No writer seems to have reported the extracrdinary yawning

under water to which I have alluded. .

Another equally deep tank held at least a dozen examples of

Necturus. On both occasions they remained undisturbed at the

bottom of the aquarium without betraying any signs of unusual

activity such as would attend recent or approaching visits to the

surface. Some of them had lived for three years in the aquarium ;

their bodies were sleek, firmly rounded, and fully extended.

There is a point concerning the vascular system which requires

slight emendation. All the blood that leaves the heart of

Necturus passes through the afferent branchial arteries to the

external gills, J. E. V. Boas (Morph. Jahrb. vii. 1882) had

described an external carotid artery arising from the first afferent

artery on each side, between the heart and the base of the first

gill. I have satisfied myself by injection that this artery does

not exist in the position claimed for it by Boas. It arises from

the first efferent artery as described and figured by W.S. Miller §.

Unfortunately, two papers by H. L. Bruner had escaped my

notice earlier ||. In these regular oscillations cf the floor of the

mouth in aquatic respiration are attributed to WVectwrus, and,

under certain experimental conditions, of which temperature is

one of the factors, pulmonary respiration is stated to oceur at in-

tervals. Bruner’s analysis of the respiratory movements observed

* Burt G. Wilder: “On the habits of Cryptobranchus.” Amer. Nat. xvi. pp. 816-

817, 1882.

+ Albert M. Reese: “The habits of the Giant Salamander.” Pop. Science

Monthly, vol. Ixii. pp. 526-531. “New York, 1903.

t Bertram G. Smith: ‘ The life-history and habits of Cryptobranchus allegheni-

ensis.’ Biol. Bull. xiu. pp. 5-39, 1907.

§ W.S. Miller: “ The vascular system of Necturus maculatus.” Contributions

from the Anatomical Laboratory of the University of Wisconsin, Bull. Univ.

Wisconsin, No. 33, Science Series, vol. 11. No.3, pp. 211-226, pls. ix.-xi. Madison,

Wisconsin, 1900.

| H. L. Bruner: 1. ‘The Mechanism of Pulmonary Respiration in Amphibians

with Gill-Clefts.”” Morph. Jahrb. xlviii. pp. 63-82, 1914. 2. “ Jacobson’s Organ and

the Respiratory Mechanism of Amphibians.” bid. pp. 157-165.

MOVEMENTS OF NECTURUS AND CRYPYOBRANCHUS. 65]

by him in Vectwrus does not include the gross flapping of the ex-

ternal gills. ‘There is, therefore, an apparent textual discrepancy

between his account and mine, which will doubtless be cleared by

further independent observations. Bruner fortifies his statements

concerning the “ bueco-pharyngeal mechanism” by a careful

description of the choanal valve of wVectuwrus; and he seems to

assign a preponderating rédle to the gill-clefts in the branchial

respiration of this genus. It is not altogether inconceivable that

in different parts of its climatic range, as well as under diverse

laboratory conditions, the several components of the respiratory

tract may vary in the relative frequency of their turns. The

behaviour of Cryptobranchus informs us that not every yawn is

an act of breathing. No contrast could be more realistic than

that between the restless, air-craving Cryptobranchus and the

listless, gill-waving Vecturws when sieved at the right biological

moment in the splendidly appointed tanks of the New York

Aquarium.

McGill University, Montreal,

October 10, 1920.

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THE SECREVARY ON ADDITTONS TO THE MENAGERIE. 653

EXHIBITIONS AND NOTICES.

October 19th, 1920.

Prof. EK. W. MacBrips, F.R.S., F.Z.8., Vice-President,

in the Chair.

The Secrerary read the following Report on the Additions

to the Society’s Menagerie during the months of June, July,

August, and September, 1920.

JUNE.

The registered additions to the Society’s Menagerie during the

month of June were 495 in number. Of these 267 were acquired

by presentation, 32 were deposited, 155 were purchased, 12 were

received in exchange, and 29 were born in the Menagerie.

The following may be specially mentioned :—

Manmalia.

1 American Bison (Bison americanus), 3, born in the Mena-

gerie on June Ist.

1 Duke of Bedford’s Deer (Cervus wanthopygius), 9 , from Man-

churia, deposited by H.G. The Duke of Bedford, K.G., Pres. Z.8.,

on June 2nd.

1 Hippopotamus (/Zippopotamus amphibius), 2, trom the

Upper Nile, purchased on June 9th.

1 Oak-Dormouse (Dyromys nitidula), from Kedos, Anatolia,

presented by Capt. G. Noel Rogers on June dth. New to

the Collection.

Aves.

1 Grey-cheeked Thrush (Hylocichla alicic), from N. America ;

1 Red-eyed Vireo (Vireo olivacea), from N. America; 2 Cali-

fornian Ground-Finches (Pipilo crissalis), from N. America ;

2 Texas Colins (Colinus texwnus), from Texas; 4 Mexican Gold-

finches (Chrysomitris meaiconus), from Mexico; 4 Pine-Siskins

(Spinus pinws), from N. America; 2 Rufous-winged Tyrants

(Pitangus rufipennis), from Colombia; 1 San-blas Jay (Cissolopha

sanblasiana), from Mexico; 2 Inca Doves (Scardafella inca), from

Central America; 1 Venezuelan Motmot (Jomotus venezuele),

from Venezuela; 2 Red-winged Doves (Chamepelia rufipennis),

from Central America; 2 Mexican Ground-Doves (Chamepelia

pallescens), from Mexico; 1 Northern Guan (Ortalis maceallc),

trom 8. Texas; 1 Western Gull (Larus occidentalis), from Pacific

Coast of N. America; 1 Pacitic Gull (Gabianus pacificus). All

presented by the Zoological Society of New York on July 9th,

and all new to the Collection.

654 HE SECRETARY ON ADDITIONS TO THE MENAGERIE.

3 Burchell’s Glossy Starlings (Lamprocolius australis), from

S. Afvica, received in exchange on June 17th. New to the

Collection.

Reptilia.

1 Night Tree-Snake (Dipsadomorphus dendrophilus), from

Malay Peninsula, purchased June 4th. New to the Collection.

JULY.

The registered additions to the Society’s Menagerie during the

month of July were 413 in number. Of these 71 were acquired

by presentation, 24 were deposited, 231 were purchased, and 87

were born in the Menagerie.

The following may be specially mentioned :—

Mammalia.

2 Tigers (Felis tigris), from India, presented by The Maharajah

of Ratlam on July 16th.

1 Indian Elephant (Hlephas maximus), from India, presented

by The Maharajah of Uooch Behar, K.C.S.1.

Aves.

2 Yellow-collared Macaws (Ara auricapilla), from HKastern

Bolivia, presented by Walter Goodfellow, F.Z.S., on July 23rd.

New to the Collection.

5 Northern Boatbills (Caneroma zeledoni), from Mexico, pur-

chased on July 9th.

Reptilia, Batrachia, Pisces.

2 Indian Cobras (WVaia tripudians), 1 Hamadryad (Vaia bun-

garus), 2 Russell’s Vipers (Vipera russelli), and 1 Banded Krait

(Bungarus fasciatus), from India; purchased on July 16th.

AUGUST.

The registered additions to the Society's Menagerie during

the month of August were 144 in number. Of these 37 were

acquired by presentation, 73 were deposited, 24 were purchased,

8 were received in exchange, and 2 were born in the Menagerie.

The following may be specially mentioned :—

2 Black-and-White Guerezas (Colobus abyssinicus), from the

Lotuka District, Mongalla Province, 8. Soudan, presented by

Capt. W. R. Barker on August Ist.

2 Leopards (Felis pardus) and 1 Baboon (Papio anubis), from

Southern Nigeria, presented by A. E. F. Murray on August 19th.

2 Boddaert’s Squirrels (Callosciuwrus notatus), from Java,

purchased on August 12th.

THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 655

2 Collared Sunbirds (Anthothreptes collaris), new to the Collec-

tion, from Natal, presented by Harold Millar on August 2nd.

2 Brazilian Cariamas (Cariama cristata), from Morro Velho,

Brazil, presented by George Chalmers, C.M.Z.S.

2 Daudin’s Vipers (Vipera lebetina), new to the Collection, from

Algeria, deposited on August 19th,

SEPTEMBER.

The registered additions to the Society’s Menagerie during

the month of September were 158 in number. Of these 105

were acquired by presentation, 28 were deposited, 9 were pur-

chased, 4 were received in exchange, and 12 were born in the

Menagerie.

The following may be specially mentioned :—

3 Polar Bears (Thalarctos maritimus), ¢ 2 2, from Greenland,

purchased on September Ist.

1 Bosman’s Potto (Perodicticus potto), from Ashanti, presented

by Mrs. Beddington on September 22nd.

1 Kaffir Mongoose (J/ongos caffer), from Ikivun, 8. Nigeria,

new to the Collection, presented by RK. H. Lapage, Esq.

1 Banded Squirrel (Callosciurus vittatus), from N. Borneo, new

to the Collection, deposited on September 231d.

1 Green-headed Amazon (Chrysotis virenticeps), from Central

America, received in exchange on September 30th. New to the

Collection.

Dr. P. CHatmers Mircuent, F.R.S., F.Z.S., exhibited a

Double-tailed Lizard, and illustrated his remarks with a series of

radiographs and lantern-slides.

November 2nd, 1920.

Sir Sipney F. Harmer, K.B.E., F.R.S., Vice President,

in the Chair.

Mr. R. I. Pocock, F.R.S., F.Z.S., exhibited, on behalf of Mr.

H. Gerrard, the skin of the groin of an example of Z'ragelaphus

buaxtoni, proving the presence of inguinal glands in that Antelope.

Mr. E. G. Bouuencer, F.Z.S., exhibited, and made remarks

upon, living specimens of Vecturus, presented to the Society by

Dr. AD Walley, F.R.S., F.Z.8.

656 THE SECRELARY ON ADDITIONS TO THE MENAGERIE.

Mr. T. A. Barns, F.Z.8., gave an account of his recent

expedition through the forests of Africa in search of Gorilla and

Okapi, illustrating his remarks with a fine series of cinematograph

films,

November i6th, 1920.

Prof. J. P. Hinz, D.S8e., F.R.S., Vice-President,

in the Chair.

The Srcrerary read the following Report on the Additions

to the Society’s Menagerie during the month of October, 1920 ;—

The registered additions to the Society's Menagerie during

the month of October were 273 in number. Of these 80 were

acquired by presentation, 37 were deposited, 151 were purchased,

1 was received in exchange, and 4 were born in the Menagerie.

The following may be specially mentioned :—

1 Maned Wolf (Canis jubatus) and a Crab-eating Dog (Canis

thous), from Morro Velho, Brazil; presented by Mr. George

Chalmers, C.M.Z.8., on October 18th.

2 Bay Duikers (Cephalophus dorsalis) and 1 Maxwell’s Duiker

(C. macwelli), from Vogoland ; presented by Dr. Malcolm B. Hay

on October 7th.

A collection of mammals and birds from the East, purchased

on October 13th, containing 2 De Brun’s Wallabies (dZacropus

brunit), from Aru; 1 Ursine Tree-Kangaroo (Dendrolagus

ursinus), from Northern New Guinea, and the following birds

which are new to the Collection: 1 Sula Island Calornis (Lam-

procorax sulaensis) ; 1 Black-headed Butcher-Crow (Cracticus

cassicus), from Aru; 1 Rosenberg’s Lorikeet (Vrichoglossus

rosenbergr), from Mysore; 1 Mueller’s Imperial Fruit-Pigeon

(Carpophagau muelleri), from Aru; 1 Pinon Imperial Fruit-Pigeon

(Carpophaga pinon) and 2 Finsech’s Imperial Fruit-Pigeons

(Carpophaga finschi), from New Ireland.

Mr. J. T. Cunninacuam, M.A., F.Z.8., exhibited, and made

remarks upon, a specimen of the Leech 7’rocheta, recently found

in the Society’s Gardens.

Mr. F, Martin Duncan, F.Z.8., exhibited, and made remarks

upon, @ series of cimematograph films, which he had taken of

animals in the Society's Gardens,

No. 208.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

October 19th, 1920.

Prof. KE. W. MacBrivz, F.R.S., F.Z.S., Vice-President,

in the Chair,

The Secrerary read a Report on the Additions to the Society’s

Menagerie during the months of June, July, August, and

September, 1920. |

Dr. P. CHaumers Mircuet, F.R.S., exhibited and made

remarks upon a Double-tailed Lizard.

Dr. E. H. Hankin, M.A., communicated his paper ‘‘ Observa-

tions on the Flight of Flying- Fishes.”

In the absence of Dr. W. N. F. Woopianp, a réswmé of his

paper “On some Results of ligaturing the Anterior Abdominal

Vein in the Indian Toad (Lufo stomaticus)” was given by Sir

S. F. Harmer, F.R.S.

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent's Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications ; but it may be obtained on the

day of publication at the price of Sixpence, or, if desired, sent post-free for

the sum of Siw Shillings per annum, payable in advance,

Prof. MAxwE.Lu hanes, F.Z.8., communicated a paper by

Mr. G. Corrprent on “The Life-History and Habits of the

Yellow Dung-Fly: a Blow-Fly Check.”

The next Meeting of the Society for Scientific Business will

be held on Tuesday, November 2nd, 1920, at 5.30 p.m., when the

following communications will be made : —

aE nop Ses eee

Tans oe vations. on 1 the Structure anal Life-History of the

Common Nematode of the Dogfish (Scylliwm canicula).

Mrs. O. A. Merrirr HAWKES, M.Sce., B. _B.Se.

Olbsemarians on the Life- isto, Biology, and Genetics of

the Lady-bird Beetle, Adalia bipunctata Mulsant.

Haru Ram Menra, M.Sc.

On the Sexual Phase in certain Indian Naididz (Oligocheta).

KH. A. BARNES.

In the Land of the Gorilla and Okapi.

The following Papers have been received :—

Wo. A. CUNNINGTON, M.A., Ph.D. B.ZS

The Fauna of the cas oe a ey: in Comparative

Limnology, with special reference to Tanganyika.

Henry F. CARTER.

Description of the Adult, Larval, and Pupal Stages of a new

Mosquito from Lord Howe Island, ‘South Pacific.

C. F. SONNTAG, M.D , Ch. [Big Ay di

The Tee ae oe a the Tongues of the Mam-

malia.—lII. Fam. Simiide.

C. L. Boutencsr, M.A., D.Sc., F.Z.S.

Filariid Worms from Mammals and Birds in the Society’s

Gardens, 1914-1915.

Aveusta ARNBACK Crristiz- LINDE.

a ee REE EER UO

On the Reproductive Organs of the Ascidian Aiikenthalia

boreulis Gottschaldt.

EK. Horxtinson, D.S.O., M.A., M.B., F.Z.S.

Records of Birds which have bred in Captivity.

S. Monckton Corman, M.D., F.R.S.

Note on the Capture of a rare Parasitic Fly, Hammomyia

(Hylephila) unilineata Zett, on Primrose Hill.

EK. DuKINFIELD Jonus, F.Z.S., F.W.S.

Descriptions of New Moths from South-east Brazil.

D. M.S. Watson.

The Basis of Classification of the Theriodontia.

The Publication Committee desire to call the attention of

those who propose to offer Papers to the Society, to the great

inerease in the cost of paper and printing. This will render it

necessary for the present that papers should be condensed, and

be limited so far as possible to the description of new results.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Society or Lonpon,

Recent’s Park, Lonpon, N.W. 8,

October 26th, 1920.

No. 209.

ABSTRACT OF THE PROCHEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.’

November 2nd, 1920.

Sir Sipney F. Harmer, K.B.E., F.R.S., Vice-President,

in the Chair.

In the absence of Dr. A. Wiuuey, F.R.S., F.Z.8., a réswmé of

his communication, “‘ A Note on the Respiratory Movements of

Neciurus and Cryptobranchus,” was given by Mr. E.G. BouLENGER,

F.Z.8., who also exhibited living specimens of Vecturus, presented

to the Society by Dr. Willey.

In the absence of the Author, Mr. J. H. Luoyp, M.S8c., F.Z.S.,

his paper on “Some Observations on the Structure and Life-

History of the Common Nematode of the Dogfish (Seyllawm

canicula)” was taken as read.

In the absence of Mrs. O. A. Merritt Hawkes, M.Sc., B.Sc.,

a résumé of her paper, ‘ Observations on the Life-History,

Biology, and Genetics of the Lady-bird Beetle, Adalia bipunctata

Mulsant,” was given by Mr. F. Martin Duncan, F.Z.8.

Mr. R. I. Pocock, F.R.S., F.Z.S., exhibited, on behalf of Mr.

KE. Gerrarp, the skin of the groin of an example of Z'ragelaphus

buxtont, proving the presence of inguinal glands in that Antelope.

In the absence of the Author, Prof. Haru Ram Mrnra, M.Sc.,

his paper, “‘On the Sexual Phase in certain Indian Naidide

(Oligocheta),” was taken as read.

= = ~

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent’s Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications ; but it may be obtained on the

day of publication at the price of Siapence, or, if desired, sent post-free for

the sum of Six Shillings per annum, payable in advance,

Mr. T. A. Barns, F.Z.S., gave an account of his recent ex-

pedition through phe Romeats a Africa in search of Gorilla and

Okapi, illustrating his remarks with a fine series of cinemato-

graph films.

The next Meeting of the Society for Scientific Business will be

held on Tuesday, November 16th, 1920, at 5.30 p.m., when the

following communications will be made :—

The SECRETARY.

Report on the Additions to the Society’s Menagerie during

the month of October.

Wo. r A . Cunnineron, M.A., . Ph.D., he ZS.

The ean of ane Gea Lakes; a Study in Coropemttine

Limnology, with special reference to Tanganyika.

Henry FE. CARTER.

Descriptions of the Adult, Larval, and Pupal Stages of a new

Mosquito from Lord Howe Island, South Pacific.

C._L. Bourmene, M.A. D.Se.. 1.2.8.

Fil stad Aivoene =e ieee and Birds in the Society’s

Gardens, 1914-1915.

The following Papers have been received :—

C. F. Sonntag, M.D., Ch.B., F.Z.S.

The Comparative Anatomy of the Tongues of the Mam-

malia.—IT. Family Simuiidee.

(oye)

Augusta ARNBACK CHRIsTIE- LINDE.

On the Reproductive Organs of the Ascidian Avikenthalia

borealis Gottschaldt.

S. Moncxron CorEMan, ETE . F. R. 5.

Note on the Capture of @ rare Parasitic Fly, Hammomyia

(Zylephila) unilineata Zett, on Primrose Hill.

E. DuxkINFIELD Jonts, F.Z.S8., F.E.S.

Descriptions of New Moths from South-East Brazil.

D. M.S. Watson, F.Z.S.

The Basis of Classification of the Theriodontia.

B. P. UvARoveE.

= eee

The Geographical Distribution of Orthopterous Insects in

the Caucasus and in Western Asia.

Miss Joan B. Proctnur, F.Z.S8.

On the Variation of the Secapula in the Aglossa and the

Arcifera.

EK. G. BoutencsEr, F.Z.8.

Experiments on Colour-changes of the Spotted Salamander

(Salamandra maculosa), conducted in the Society’s Gardens.

The Publication Committee desire to call the attention of

those who propose to offer Papers to the Society, to the great

increase in the cost of paper and printing. This will render it

necessary for the present that papers should be condensed and

be limited so far as possible to the description of new results.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Society oF Lonpon,

Recent’s Parx, Lonpon, N.W.8.

November 9th, 1920.

No. 210.

ABSTRACT OF THE PROCEEDINGS

ZOOLOGICAL SOCIETY OF LONDON.*

November 16th, 1920.

Prof. J. P. Hitt, D,Sc., F.R.S., Vice-President,

in the Chair.

The Secretary read a Report on the Additions to the Society’s

Menagerie during the month of October 1920,

Mr. F. Martin Duncan, F.R.M.S., F.Z.S., exhibited and made

remarks upon a series of cinematograph films, which he had taken

of Animals in the Society’s Gardens.

Mr. J. T. Cunntnenam, M.A., F.Z.S., exhibited and made

remarks upon a specimen of the Leech Zrocheta, recently found

in the Society’s Gardens.

Dr. W. A. Cunnineron, M.A., F.Z.S., gave an account of his

paper on the “ Fauna of the African Lakes, with special reference

to Tanganyika.” After referring to certain physical and geo-

logical features which have a bearing on the subject, the nature

of the various animal forms inhabiting the lakes was dealt with.

Tanganyika was shown to havea very distinctive fauna, in that :—

(1) it includes many more different types than any of the other

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent's Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications ; but it may be obtained on the

day of publication at the price of Sixpence, or, if desired, sent post-tree for

the sum of Six Shillings per annum, payable in advance.

lakes, (2) an extremely large proportion of them are not found

elsewhere, (3) certain forms (notably Gasteropoda) are considered

to have a marine-like appearance. The view previously put

forward which regarded the lake as the remains of an old Jurassic

sea was considered untenable, since many of the types thought to

be marine and primitive belong to essentially fresh-water groups

snd show signs of specialisation. The Jurassic hypothesis proves

likewise incompatible with recent geological evidence. After

discussing various other theories, it was suggested that Tan-

ganyika probably owes its remarkable organisms to a prolonged

period of isolation—coupled, perhaps, with the effect of an

increased salinity which isolation would involve.

Tn the absence of the Authors, Mr. Henry F. Carrsr’s paper

cn ** Descriptions of the Adult, Larval, and Pupal Stages of a

new Mcsquito from Lord Howe Island, South Pacific,” and Prof.

U. L. Bovtencsr’s paper on “ Filariid Worms from Mammalia

aod Bircs in the Society’s Gardens, 1914-1915,” were taken as

The next Meeting of the Society for Scientific Business will

be held on Tuesday, February 8th, 1921, at 5.30 p.m.

A notice stating the Agenda for that Meeting will be circulated

early in February.

The following Papers have been received :—

C. F. Sonntac, M.D., Ch.B., F.Z.S.

The Comparative Anatomy of the Tongues of the Mam-

malia.—IJ. Fam. Simiide.

(oye)

Aveusta ARNBACK CuristTIp- LINDE.

On the Reproductive Organs of the Ascidian Avikenthalia

borealis Gottschaldt.

S. Moncxron Copeman, M.D., F.R.S.

Note on the Capture of a rare Parasitic Fly, Hammomyia

(Hylephila) unilineata Zett, on Primrose Hill.

E. DuxKINFIELD Jongs. F.Z.S., F.E.S.

Descriptions of New Moths from South-East Brazil.

D. M.S. Watson, F.Z.8.

The Basis of Classification of the Theriodontia.

B. P. Uvarov.

The Geographical Distribution of Orthopterous Insects in

the Caucasus and in Western Asia.

Miss Joan B. Proctor, F.Z.S8.

On the Variation of the Scapula in the Batrachian Groups

Aglossa and Arcifera.

E. G. BouLenceEr, F.Z.8.

Experiments on Colour-changes of the Spotted Salamander

(Salamandra maculosa), conducted in the Society’s Gardens.

J. SrepHenson, D.Sc., F.Z.S.

Morphology, Classification, and Zoogeography of Indian

Oligocheta.—I. The Affinities and Systematic Position of the

Genus Hudichogaster Mchlsn., and some related Questions.

II. On Polyphyly in the Oligocheta. III. Some general

Considerations on the Geographical Distribution of Indian

Oligocheta.

R. Broom, D Sc., F.R.S., C.M.Z.S.

On the Structure of the Reptilian Tarsus.

Cuas. F. Sonntag, M.D., Ch.B., F.Z.S.

A Contribution to the Anatomy of the Three-toed Sloth

(Bradypus tridactylus).

W. T. Cauman, D.Sc., F.Z.S.

Notes on Marine Wood-boring Animals.—II. Crustacea.

The Publication Committee desire to call the attention of

those who propose to offer Papers to the Society, to the great

increase in the cost of paper and printing. This will render it

necessary for the present that papers should be condensed and

be limited so far as possible to the description of new results.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Socrery or Lonpon,

ReEcGeENt’s Park, Lonpon, N.W. 8.

November 23rd, 1920.

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PAPERS.

On some Results of Ligaturing the Anterior Abdominal Vein in the Indian Toad (Bufo

stomaticus Liitken), By W. N. F. Woopianp, D.S8c., F.Z.8., Indian Educational

Service, Senior Professor of Zoology, Muir Central College, ‘Allanatag. U.P., India.

(Text- figure Te) SS hele cid Date a ORE AO EROA OKO Gio eA ICH Leats PB MicwmI MMe cid alh B Sieh i Che

Some Observations on the Structure and Life-History of the Common Nematode of

the Dogfish (Scylliwm canicula). By J. H. Lioyp, M.Se., F.Z.8., Assistant Lecturer

and Deimonstrator in Zoology at University College, Cardiff. (Text-figures 1-3.) ..

On the Sexual Phase in certain Indian Naidide (Oligocheta). By Harv Ram Munra,

M.S8c., Professor of Zoology, Hindu University, Benares. (Text-figures 1-3.) ......

Observations on the Flight of Flying-Fishes, By E. H. Hanxry, M.A., Se.D., Agra,

ithe, (CearsreRh)) lod an ao pobpes dope on Coan GoommooE Bs ial a) stale usa Marae

Observations on the Life History, Biology, and Genetics of the Lady-bird Beetle,

Adalia bipunciata Mulsant. By Onéra A. Murrirr Hawgus, M.Se.(Birm.), B.Se,

(Ubon) ane Sosin s Gb a Nob erad hn Aero encic HO Rod OOO Ur Ot > Ue TOU O REDS iY Ais hi

Filariid Worms from Mammals and Birds in the Society’s Gardens, 1914-1915. By

Pniniest, Maliore sn (bext-tipresy les) )ycciks avatsurte yates ais oe tcsthe Stat epee eet sale le

The Fauna of the African Lakes: a Study in Comparative Limnology with special

reference to eee By Wm. A. Ounnineron, M.A., Ph.D., F.Z.S. (Text-

figures 1-2.).. Re air ree MI Ne cA rend cs SAS a en RUN RN

Descriptions of the Adult, Larval, and Pupal Stages of a New Mosquito from Lord

Howe Island, 8. Pacific. By Henry F. Carrer, Liverpool School of Tropical

Mieditennerra Mext=tia ures il 32). aye ob tale is qe apalena cee sieve aacareroucbeuevsce ePelsneieinie orelisisieremete

The Life-History and Habits of the Yellow Dung-Fly (Scatophaga stercoraria): a

possible Blow-Fly Check. By G. 8. Correretn With a Preface by Professor

MaxwrnnmneROv, HK. Z.S8. (Dext-floures L=04))) ci tsare sieve mistress toners orareo tele

Remarks on the Respiratory Movements of Nectwrus and Cryptobranchus. By

A, Witty, M.A., D.Se., F.R.S., F.Z.S.

UMTS ARSE IS Ge cae Eke Ome ee ATTEN rE Cees By ae ab MT ar te aos aa

List of Council and Officers

Alphabetical List of Contributors ..--....

Index of Illustrations ..

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