bas 2

OF THE

COUNCIL AND OFFICERS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

1923.

Patron.

His Masrstry Tur Kine.

COUNCIL.

His Grace Tar Duxe or Beprorp, K.G., F.R.S., President.

Cot. 8. Moncrron Copeman,

M.D., F.R.S.

CHARLES DruMMoND,

Treasurer.

ALFRED Ezra, Esq., O.B.E.

Huexn §. Guapstonn, Ese.,

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

Tue Rr. Hon. 'tHr Viscount

Grey or Fatiopon, K.G.,

P.C., Vice-President.

Sir Srpney F. Harmer, K.B.E.,

MEAS. Sc.D e Ransse, iiace=

President.

Pror. James P. Hitz, D.Sc.,

E.R.S., Vice-President.

‘“Witt1am Huntsman, Esa.

Lr.-Cou. THe Lorp ALAsratr

Ropert Innes-Kur, D.S.O.

Sir Watrer Roper LAWRENCE,

Br..G.C.1.E., G.C.V.O., C.B.

Ksq.,

| Con. Str Henry McManon,

G.C.M.G., K.C.L.E.

HE. G. B. Mrapz-Watpo, Esgq.,

Vice-President.

P. Cuatmers MrrcHety, Esq.,

C.B.E., M.A., D.Sc., LL.D.,

F.RB.S., Secretary.

THe Hart or Onstow, O.B.E.,

Vice-President.

Masor Atperr Pam, O.B.E.

Lyr.-Cot. Str Dayrp Prain,

C.M.G., C.1.E., F.RB.S8.

Tue Lorp QuEENBOROUGH.

Harotp Joun Hastines

RussEtt, Esq., K.C.

RicwarD STEPHENS TAYLor,

Esq.

A. Smurra Woopwarp, Esq.,

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

dent.

PRINCIPAL OFFICERS.

P. Cuaumers Mircuent, C.B.i., M.A., D.Se., LL.D., E.RS.,

Secretary.

G. M. Vevers, M.R.C.S., L.R.C.P., Superintendent of the

Gardens.

D, Seru-Surru, Kesident Curator of dfammals and Birds.

Epwarp G. BouLencEr, Curator of Reptiles.

Miss L, EK. Cunusman, F.E.S.

, Curator of Insects.

C. F. Sonntac, M.D., Anatomist.

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

Prof. G. H.Woo.tpripez, F.R.C.V.S., Hon. Consulting Veterinary

Surgeon.

Dr. R. W. A. Saumanp, O.B.E., M.D., Hon. Consulting

Radiologist.

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

F. W. Bonn, Accountant.

W. H. Coun, Chie? Clerk.

“ TAST OF CONTENTS.

1923, pp. 1-481.

EXHIBITIONS AND NOTIUKS.

Tue Secrurary. Report on Additions to the Society's

Menagerie during the months of November and

Deere, SPAN Bec cennentaidees sy oscceuednends Oeneenanae

Lord Rorascuttp, F.R.S.,F.Z.8. Exhibition of adult male

Mtonmata ian Gorell to. aM Bee aber aver eg Bn'aisictone spe

Mr. Ouprietp Tuomas, F.B.S., F.Z.S. Exhibition of a new

Rock Kangaroo from Northern Queensland ............

Exhibition of the Skull of a Pigmy Fruit-Bat from

Sian attra eee ea ree cet Ale sicelntie wa gteaa ees

Mr. J. B. Scrtvenorn, M.A. Exhibition of photograph

showing native method of breaking-in captured

Beye raitis) 053 Se Atak male tian Bia ples «sale aisha nol

Mr. E. G. Boutencerr, F.Z.8. Account of his recent visit

ROWAN TESTA! pesca acts 1s Abe cht ehcp ame ee rs SR staas ta ai

The Secretary. Report on Additions to the Society’s

Menagerie during the month of January, 1923 ......

My. D. Sera-Smiru, F.Z.8. Exhibition of photographs and

Skins Ghebindis2@n- Paradise tient chess tagescceinue sacs

Prof. H. M. Lerroy, F.Z.S8. Exhibition of cinematograph

film of the life-history of the House-fly ...............

The Szcretary. Exhibition of a photograph of the Polar

Teisaras) CS? aya) 8 TBarelope,”” canada aolseueooeedoncene

178

Leg

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

remarks upon, drawings of the feet and nose of the

HOlar Bear < ack iak Genie 6 6. SARC: + « Ceeetete See OES.

The Secrerary. Note on a case of apparent melanism in

Tippelskirch’s Giraffe

Mr. R. I. Pococg, F.R.S., F.Z.S8. Exhibition of drawings

of the feet and spurs of the Echidna

Sir G. ABERcRomBy. Exhibition of head of a Kob

Prof. KE. W. MacBrips, F.R.S., F.Z.S. Exhibition of

photomicrographs of sections through nuptial callosi-

ties of ame amd Algyids |... cemeees. J: Pee dee mana. ne 8,

Mv. F. M. Swynnerton, C.M.Z.8. Exhibition of stomach-

contents of a Crocodile. (Text-figure 1.)...............

The Secrerary. Report on Additions to the Society’s

Menagerie during the month of February, 1923 ......

Mr. F. Marrin Duncan, F.R.M.S., F.Z.S. Exhibition of

a Caterpillar infected with the Entomogenous fungus

Comdypeeps robortsta 12. we sie, o5. fsa th-io8 « waa nae arene

Mr. M. A. C. Hinton, F.Z.S. Exhibition of skin of a

WRNOMIES See etre ere uate ae ie ee he voc Cie eee ne

The Sscrerary. Nxhibition of photographs of deep-level

reservoirs of the Society’s new Aquarium

Mr. G. C. Rowson, M.A., F.Z.S. Exhibition of the Snail

Pianorbis du joury Graelis). ...... a. eae eee eee

Dr. G. M. Vevers, F.Z.S. Account of his recent visit to

Zoological Gardens in Holland and Belgium............

The Secrerary. Report on Additions to the Society’s

Menagerie during the month of March, 1923 .........

Capt. R. B. Murray, F.G.8., F.R.G.S. Exhibition of living

specimens of a Giant Centipede from Trinidad ......

Lt.-Col. S. Moncxron Coprman, F.R.S., F.Z.S., and Major

E. E. Austen, D.S8.0., F.Z.S. Exhibition of a rare

British: Brpterom sy her. ceesnscsne aes ce os eae eee

Page

US)

179

180

479

480

10.

12.

PAPERS.

. Coat Colour in Greyhounds. By Avarr Diceuton,

WBC Dy, GaSe. 0 eS ARREAD, Meets ae. aRUIDL DIN.

. Note on the Zoxa of a Land Urab, Cardisoma armatum.

By H. Grauam Cannon, B.A., F.Z.8. (Text-figures

MNO) pees csacesYa cha drtarecostclore ter ER ER OM a RL is Soe

On the Linguatalid Arachnid Raillietiella furcocerca

{ Diesing, 1835] Sambon, 1922. By Grorce S. GicLioxt,

M.D. Pisa

. The Permian Fishes of the Genus Acentrophorus. By

K. Leonarp Gitt, M.Sc. (Text-figures 1-16.).........

The Postorbital Articulation of the Palatoquadrate

with the Neurocranium in the Ceelacanthid Fishes.

By Epwarp Puetes Autis, Jr., F.Z.8.

- Some Microfilariz found in the Blood of Birds dying

in the Zoological Gardens, 1920-1921. By Rrra

MarKBREITER, B.Sc., Research Assistant in the

Helminthological Department of the London School

of Tropical Medicine, (Text-figures 1-6.)

. On the Vagus and Sympathetic Nerves of the Terres-

trial Carnivora. By CuHartes F. Sonntac, M.D.,

¥.Z.S., Anatomist tothe Society. (Text-figures 1-14.)

. Remarks on some Palearctic Bears. By Hinar Lonn-

BERG, F'.M.Z.S., &e. (Plates I. & II.)

. Some Notes on Leander longirostris M. Edwards, and

other British Prawns. By Roserr Gurney, M.A.,

F.LS., F.Z.S. (Text-figures 1-6.)

Report on the Deaths which occurred in the Society’s

Gardens during 1922. By N.S. Lucas, M.B., F.Z.S.,

Pathologist to the Society

The Comparative Anatomy of the Tongues of the

Mammalia.—VIIT. Carnivora. By Crarurs F.

Sonnrac, M.D., ¥.Z.8., Anatomist to the Society.

(ext - figures mM ae Nise dete agit. dec dct aetae alias sean +.

Notes on the Tunicate Rhizomoigula globuiaris Pallas.

By R. Kirxparricx, F.Z.8. (Plate I.)

eee eee eee eee oes

Page

125

129

13.

14,

15.

18.

iS),

Zalbe

On the Feet and Rhinarium of the Polar Bear (Zhal-

arctos maritimus). By R. I. Pocock, F.R.S., F.Z.S.

(escbe tise: 2). eee oa 2 ae ee

The Embryonic Development of the Porbeagie Shark,

Lamna cornubica. By EH. W. Suan, B.A., F.Z.S.

(Text-figures 1 & 2.)

Ce i i i i i i a er ed

Notes on the Pairing of the Land Crab, Cardisoma

armatum. By Miss L. KH. Cuzxsman, F.E.S., F.Z.S..

On the External Characters of Hlaphurus, Hydropoies,

Pudu, and other Cervide. By R. I. Pococs, F.R.S.,

EZ. (i (ilext-heures 21 72) \ peers ee sane eens Ree

The Classification of the Sciuride. By R. I. Pococx,

F.R.S., F.Z.S. (Text-figures 18-29.)

On the Mammals obtained in Darfur by the Lynes-

Lowe Expedition. By Oxuprienp Tuomas, F.R.S.,

F.Z.S., and Martin A. C. Hinton, F.Z.8

Coe te tee ee week

On a Remnant of the Omphalo-mesenteric Arteries in

the Manatee. By K. Kosranecki1, M.D., LL.D.,

F.Ac.Sc. Cracow. (Text-figure 1.)

. Cuckoos’ Eggsand Evolution. By E.C, Stuarr Baker,

F.Z.8. (Plates I-IV.)

A Comparative Study of the Buccal Glands and Teeth

of the Opisthoglypha, and a Discussion on the Evolu-

tion of the Order from Aglypha. By Susum Cu.

Sarkar, F.Z.S. (Text-figures 1-29.)

. On the Anatomy, Physiology, and Pathology of the

Chimpanzee. By Cuarues F. Sonntag, M.D., F.Z.S.,

Anatomist to the Society. (Plates I.-III.; Text-

figumes 20-49.) cyte. A it ee Nie Pe eee tie

. The Elephant-Seals of Kerguelen Land. By T, P. A.

Rane SpE aibes lh i Au) See ee eremnese ears RN as 2)

. The Skeleton of Lepidosieus, with remarks on the origin

and evolution of the lower Neopterygian Fishes. By

C. Tare Rucan, M.A., F.R.S., F.Z.8., Keeper of

Zoology in the British Museum (Natural History),

(esst mene AS ae eer ee reer ee nC cc. ee

. On the Origin of Flight in Birds, By Baron Francis

INNORCSA. 7 (@ext-tommres pla). ee. 2.0.) alo ce eae

Page

159

161

181

209

247

273

277

295

323

431

445

ALPHABETICAL LIST

OF THE

CONTRIBUTORS;

With References to the several Articles contributed by each.

1923, pp. 1-481.

ABERCROMBY, Sir G.

1Dpdanil ortinain, Git ey LaveACl OP 6, IOs no edasococnsdonuoonoor dab aey. 180

Autis, EpwarpD Puetrs, Jr., F.Z.8.

The Postorbital Articulation of the Palatoquadrate

with the Neurocranium in the Coeelacanthid Fishes

Austen, Major E. E., D.S.0., F.Z.8. See Copeman, Lt.-Col.

S. Moncxk ton.

Baker, E. C. Sruart, F.Z.S.

Cuckoos’ Eggs and Evolution. (Plates I.-IV.)......... 277

Boutencer, E. G., F.Z.S.

Account of his recent visit to Vienna

Cannon, H. Granam, B.A., F.Z.S8.

Note on the Zocea of a Land-Crab, Cardisoma armatum.

(Mext= fie umes Oeics ean .cca- remnants seleannnaegcynencde esi, 11

CuerEsmMAN, Miss L. H., F.E.S., F.Z.8.

Notes on the Pairing of the Land-Crab, Cardisoma

OTT SHED oe eo SSCA A Se CARRS SO CEE SAG ne ens ga 173

vill

Copeman, Lt.-Col. 8. Moncxton, M.D., F.R.S., F.Z.S., and

Austen, Major H. H., D.S.O., F.Z.8.

Exhibition of photographs and specimens of a rare

British Dipteron (6.0.2... !.. cae ee eee ee

Dieuton, Aparr, F.R.C.S., F.Z.8.

Goat (Colour in ‘Greyhounds --seeeeeteee eee eerie

Duncan, F. Martin, F.R.M.S., F.Z.8. (Librarian to the

Society).

Exhibition of Caterpillar infected with the Entomo-

genous fungus Cordyceps robertstt ...........0.ececeeeee eee

GIaLIoLI, GrorGcE 8., M.D. Pisa.

On the Linguatulid Arachnid Raillietiella furcocerca

(iesine, US83o) Sambon, 1022 7 oie -weese ee aeereentee ny.

Git, E. Lzonarp, M.Sc.

The Permian Fishes of the Genus Acentrophorus. (Text-

figures 12062). ice esc Bees c sso. saa et. eee ts

Gurney, Rosert, M.A., F.L.S., F.Z.8.

Some Notes on Leander longirostris M. Edwards, and

other British Prawns. (Text-figures 1-6.)..................

Hinton, Martin A. C., F.Z.8.

Exhibition of the skin of a Lioness. See THomas,

OLDFIELD.

Kirxpatrick, R., F.Z.S,

Notes on the Tunicate Rhizomolgula globularis Pallas.

(iBlaite 1s) eae eon 5G. cs ee WodjoSeteee eee ;

Kosranecst, K., M.D., LL.D., F.Ac.Sci. Cracow.

On a Remnant of the Omphalo-mesenteric Arteries in

the Manatee. (Text-figure 1.) G20. ayenee.. +. +... sslerabiwees

Lerroy, Prof. H. M., M.A., F.Z.8.

Exhibition of Cinematograph film of the life-history

Ol Ge WLOUSE-HY., «6 scaei- ( sod ois sa + ables oeeB Etats © eee eee

Page

48]

AT9

JES)

Si)

155 ©

273

Lonneere, Einar, F.M.Z.S., &e.

Remarks on some Palearctic Bears. (Plates I. & II.)

Lucas, N.8., M.B., F.Z.8., Pathologist to the Society.

Report on the Deaths which occurred in the Society’s

Gardens during 1922

Bem em eee cet weet semen ses ees e eee eee t ese ses ee ees

MacBripg, Prof. E. W., F.R.S., F.Z.8.

Exhibition of photomicrographs of sections through

nuptial callosities of Rana and Alytes

eee cee ee wee eee OBL ee ee

MarKpreIver, Riva, B.Sc., Research Assistant in the

Helminthological Department, London School of

Tropical Medicine.

Some Microfilariz found in the Blood of Birds dying in

the Zoological Gardens, 1920-1921. (Text-figures 1-6.)

Mircuett, Dr. P. Coaumers, C.B.E., M.A., D.Se., LL.D.,

F.R.S. (Secretary to the Society).

Report on Additions to the Society’s Menagerie during

the months of November and December, 1922

mu wee een e ne

Report on Additions to the Society’s Menagerie during

the month of January, 1923

Pe mee ew eee ere weer eer ees eseseessceee

Exhibition of a photograph of the Polar Bears “Sam”

and “ Barbaia”

Note on a case of apparent melanism in Tippelskirch’s

Giraffe

‘ale}eiteilals}eiejeiele/s aia iisiae,a\'s\ o/c «| ea \e\esfaila\els) olelalalals/atelsimielelulalwleinic\ sath telaveretarcicreioreve

Report on Additions to the Society’s Menagerie during

the month of February, 1923

DOO IOIO DOO 60 ON OOO0 GUOORO.oo0 COCO

Exhibition of photographs of the deep-level reservoirs

of the Society’s new Aquarium

Bee Reet a ete c esse eeees cenwica veces

Report on the Additions to the Society’s Menagerie

during the month of March, 1923

Murray, Capt. R. B., F.G.S., F.R.G.S.

Exhibition of living and mounted specimens of a Giant

Wencipede) from irimicladyy ioe. cy wade eka, CR oi Ok

Proc. Zoon. Soc.— 1923, b

Page

125

180

178

179

Norcsa, Baron F.

On the Origin of Flight in Birds. (Text-figures 1-7.)

Pocock, R. I., F.R.S., F.Z.8.

Exhibition of, and remarks upon, drawings of the feet

eval Taare Our (Haves Ietolbae IBY ON Woe oe bdo cess osaecoceennecebunen.

Exhibition of drawings of the feet and spurs of the

I DYE UIC FAVE Atel. ieee oe a PRM oar oreo ANID Nn a

On the Feet and Rhinarium of the Polar Bear (Thal-

Orcosumaratineus))...) Nextt ure we) maa sane see eee

On the External Characters of Hlaphurus, Hydropotes,

Pudu, and other Cervide. (Text-figures 2-17.)............

The Classification of the Sciuride. (Text-figures 18-29.)

Reean, C. Tare, M.A., F.R.S.. F.Z.S.

The Skeleton of Lepidosteus, with remarks on the

origin and evolution of the lower Neopterygian Fishes.

(ert fieumes wL ESS ame ThE cet, Mad Se ete eee

Rine, T. H.

The Hlephant-Seals of Kerguelen Land. (Plates I. & II.)

Roruscuitp, Lions. Water, Lord, D.Sc., F.R.S., Ph.D.,

E.Z.S.

Exhibition of adult male Mountain Gorilla...............

SARKAR, On. Susuit, F.Z.8.

A Comparative Study of the Buceal Glands and Teeth

of the Opisthoglypha, and a Discussion on the Evolution

of the Order from Aglypha. (Text-figures 1-29) .:.......

SORIVENOR, J. B., M.A.

Exhibition of photograph showing native method of

breaking in captured Elephants ......................, dia Nee

Sera-Smita, D., F.Z.8.

Exhibition of photographs and skins of Birds-of-

Rarer chi Se Mee Muna aici iiayaNMumeune ban ceiuy A013 sce een eo:

Page

463

179

445

431

295

178

SHann, H. W., B.A., F.Z.8.

The Embryonic Development of the Porbeagle Shark

(Lamna cornubica). (Text-figures 1 & 2.) ...............4-.

Sonnrac, Cuarues F., M.D., F.Z.S., Anatomist to the

Society.

On the Vagus and Sympathetic Nerves of the Terres-

trial Carnivora. (Text-figures 1—14.)............-..-.:s020+

The Comparative Anatomy of the Tongues of the

Mammalia.— VIII. Carnivora. (Text-figures 15-24.)...

On the Anatomy, Physiology, and Pathology of the

Chimpanzee. (Plates I.-III.; Text-figures 25-49.)......

Swynnertron, Ff. M., C.M.Z.8.

Exhibition of stomach-contents of a Crocodile. (Text-

dior dliaeh) Wee Ven Hecate). SAR unas uO, NIM Hse Daten Verte vere ete tte

‘THOMAS, OLDFIELD, F.R.S., F.Z.8.

Exhibition of a new Rock Kangaroo from Northern

COU EVSOIS ICE ahah nBee Sele aA SAP yaa ante et mneeee eee ai sceriaa

Exhibition of the skull of a Pigmy Fruit-Bat from

RS UINIT ATG ete e sta Ie UP a Reo. OE ps Ui a ib cetacean kate eal

Tuomas, OLDFIELD, F.R.S., F.Z.S.,and Hinton, Martin A.C.,

E.Z.S.

On the Mammals obtained in Darfur by the Lynes-

dower Ecpe dition js o.WA Nie Seem tne at ccrlacicisemai lalla aueie

Vevers, Dr. G. M., F.Z.S. (Superintendent of the Gardens).

Account of his recent visit to Zoological Gardens in

Holland) anclgellsinina,\)!.. se risen hdd teatime te cee sccaeeted

Page

161

129

323

180

247

INDEX OF ILLUSTRATIONS.

Acanthopneuste

PY TT, p. 279:

Acentrophorus abbsi, Fig. 14, p. 36.

altus, Figs. 2, 4, 6, 12, 13, pp. 23,

25, 27, 34, 35.

-—— glaphyrus, Figs, 2, 9, 15, pp. 28,

30, 37.

—— varians, Figs. 1-3, 5, 7-11, pp. 21,

23, 24, 26, 28-32.

Ailurus fulgens, Figs. 1, 10, pp. 66, 76.

Amia calva, Figs. 6-8, pp. 452-454,

Ammopus, Fig. 2, p. 466,

Anchisauripus, Fig. 3, p. 467.

Anomepus, Fig. 3, p. 467.

Anthropopithecus troglodytes, Pls. I.—

TIL., figs. 25-49, pp. 323, 324, 326,

327, 329, 331, 332, 3384, 336, 340,

342, 350, 355, 366, 377, 381-383,

392, 393, 400, 404-408, 410.

Antrodesmus, Figs. 4, 5, pp. 468, 469.

Apatichnus, Fig. 3, p. 467.

Archeopteryx, Figs. 5, 6, 7, pp. 469,

470, 472.

Argyra caudata, fig. 4, Pl. 1., p. 277.

subrufa, fig. 6, Pl. 1., p. 277.

Atilax paludinosus, Fig. 1, p. 66.

Atlantoxerus getulus, Fig, 24, p. 231.

Axis avis, Hig. 4, p. 185.

occtpitalis, fig. 2,

Belomys trichotis, Wig. 29, p. 245.

Caccomantis merulinus, figs. 1-4, 10-14,

20-23, 25-27, Pl. IV., p. 277.

Proc. Zoot. Soc.—1923.

Callosciurus castaneoventris, Big. 21,

p. 221.

notatus, Hig. 18, p. 218; Fig. 21,

p: 221.

prevosti, Fig. 18, p. 213; Fig. 21,

p: 221.

Canis mesomelas, Fig. 22, p. 142.

thous, Bigs. 5, 14, pp. 70, 80.

Cardisoma armatum, Figs. 1-6, pp. 12,

13.

Cerberus rhynchops, Big. 5, p. 303.

Cercoleptes caudivoluulus, Fig. 22,

p- 142.

Cervus canadensis, Fig. 2, p. 183.

—— edi, Fig. 3, p. 184.

-— hortulorum, Figs, 3, 17, pp. 184,

205.

—— (Sika) hortulorum, Fig. 17, p. 205.

Cettia cantans, fig. 6, Pl. IIL., p. 277.

Chrysopelea ornata, Figs. 17, 18, 27,

pp. 314, 315, 322.

Cisticola cursitans, fig. 19, Pl. LYV.,

p. 277.

Citellus leursi, Fig, 26, p. 235.

—— mexicanus, Fig. 26, p. 235.

-—— -— mongolicus, Fig. 26, p. 235.

-—— 13-lineatus, Fig. 26, p. 235.

Civettictis civetta, Figs. 2,7, pp. 67, 72.

Clamator coromandus, fig. 8, Pl. I.,

1s 2 Cle

—— jacobinus, figs. 1, 3, Pl. I., p. 277.

Celurus, Fig. 6, p. 470.

Compsognathus, Fig, 4, p. 468.

X1V

Corvipes, Kig. 1, p. 465.

Corvus coronoides culminatus, ficeels

JEL, 1, fos Ae

—— intermedius, fig. 9, Pl. IL.,

p. 277.

—— splendens protigatus, fig. 6, Pl. IL.,

oe ils

1s hee

Crossarchus obscurus, Fig. 22, p. 142.

Cuculus poliocephalus, figs. 1, 3, 5, 7,

TAG OU a Pare

Cynelurus jubatus, Fig. 19, p. 137.

Cynictis penicillata, Fig. 10, p. 76.

Cynomys ludovicianus, Figs, 25, 26,

pp. 232, 235.

Cystophora cristata, Fig. 24, p. 146.

proboscidea, Fig. 24, p. 146.

splendens, fig. ‘7, Pl. IL,

Dapedius orbis, Fig. 16, p. 39.

Dendrophis pictus, Figs. 8, 9,

pp. 307, 308, 320.

Dipsas trigonata, Fig. 6, p. 304.

Dremomys dawsoni, Fig. 22, p. 223.

lokriah, Fig. 22, p. 223.

—— rufigenis, Fig. 22, p. 223.

HSCS, Viney, OD, 0), 3).

Dromeus, Fig. 6, p. 470.

Dryoplis mycterizans, Figs. 19, 20, 28,

pp: 316, 522.

23,

Hlaphurus davidianus, Figs. 5, 6, 15, 17,

pp- 187, 189, 203, 205.

Hoglaucomys fimbriatus, Bigs. 27, 29,

pp. 242, 245.

Hudynamis scolopaceus, figs. 3, 4, 5, 8,

10, Pl. IL., p. 277.

Hupalamopus, Fig. 2, p. 466.

Huxerus erythropus, Figs.

pp. 231, 232.

Felis bengalensis, Figs. 3, 4, 12, pp. 68,

69, 78.

caffra, Fig. 20, p. 138.

earacal, Big. 20, p. 138.

leo, Hig. 18, p. 136.

—— nebulosa, Fig. 19, p. 137.

pardaiis, Fig. 19, p. 137.

24, 25

INDEX OF ILLUSTRATIONS.

| Helis pardus, Fig. 18, p. 136.

| sylvesiris, Fig. 20, p. 138.

—— tigris, Fig. 18, p. 136.

Franklinia gracilis, fig. 7, Pl. TV.,

p. 277.

Funambulus palmarum, Figs. 18, 20,

pp. 213, 218.

Funisciurus congicus, Fig. 19, p. 215.

Fig. 23,

| —— leucostigma niveatus,

p. 228.

Garrulaz moniliger, fiz. 11, Pl. I.

p. 277.

-—— pectoralis, fig. 12, Pl. L., p. 277.

Genetta felina, Figs. 1, 7, pp. 66, 72.

Geosciurus capensis, Figs. 24, 25,

pp. 281, 282.

Glaucomys volans, Fig. 28, p. 244.

Gorgosaurus, Big. 7, p. 472.

Graliator, Fig. 3, p. 467.

Grammatoptila striata, fig. 10, Pl. 1..

Ds AU lc

Halicherus gryphus, Big. 24, p. 146.

Heliosciurus vrufobrachium hardyi,

Fig. 23, p. 228.

| Herpedactylus, Fig. 1, p. 465.

| Hierococeyx« sparveroides, figs. 9-11, 16-

18, Pl LET ps 20%

varius, figs. 7, 9, Pl. 1, p. 277.

Horornis fortipes, fig. 8, Pl. 11L., p. 277.

Hyena striata, Fig. 21, p. 139.

Hydropotes inermis, Figs. 7-11, pp. 192-

196.

Aylopetesalbsniger, Figs, 27, 28, pp. 242,

244.

— phayret, Big. 28, p. 244.

| Lanthocincla cineracea, fig. 19, Pl. 111.,

p. 277.

Ictonyx zorilla, Figs. 1, 9, pp. 66, 75.

| Lamna cornubica, Figs. 1, 2, pp. 165,

167.

Lariscus jalorensis, Fig, 22, p. 223.

INDEX OF ILLUSTRATIONS.

Leander adspersus, Bigs. 1, 2, pp. 100,

107.

—— longirostris, Figs. 1, 2, 3, 5, pp. 100,

107, 109, 120.

serratus, Higs. 1, 4, pp. 100, 117.

—- squilla, Figs. 1, 2, 4, 5, pp. 100,

107, 117, 120:

Lepidosteus osseus, Hig. 1, p. 446.

platystomus, Figs. 2-0, pp. 446-

450.

tristoechus, Hig. 1, p. 446.

Lutra maculicollis, Hig. 8, p. 75.

Lycodon aulicus, Figs. 14, 15, 25,

pp. 311, 312, 321.

Manatus inunguis, Fig. 1, p. 274.

Marmota marmota, Bigs. 25, 26, pp. 232,

230.

Massospondylus, Figs. 4, 5, pp. 468,

469.

Mazama tema, Figs. 13, 14, pp. 200,

202.

Meles meles, Kigs. 2, 14, pp. 67, 80.

Melursus ursinus, Bigs. 1, 11, pp. 66,

ale

Mephitis mephitica, Fig. 11, p. 77.

Mungos ichneumon, Wig. 3, p. 68.

Muntiacus muntjak, Big. 15, p. 203.

Mustela martes, Wig. 2, p. 67.

Nandinia binotata, Figs. 1, 21, pp. 66,

139.

microtis, Fig. 22, p. 223.

whiteheadi, Fig. 22, p. 228.

Nasua narica, Fig. 28, p. 144.

Nothura, Big. 7, p. 472.

Odocotleus virginianus, Fig. 12, p. 197.

peruvianus, Figs. 15, 17,

pp. 203, 205.

spinosus, Kigs. 12, 14, pp. 197,

202.

Ornitholestes, Big. 5, p. 469.

Orthotomus sutorius, figs. 5, 6, 15, 16,

Vee 20 te

| Rhinosciurus

Otaria californiana, Big. 24, p. 146.

—— gillespii, Big. 24, p. 146.

Oxybelis fulgica, Figs. 16, 26, pp. 313,

o21.

Palemonetes varians, Figs. 1, 2, 4, 6,

pp. 100, 107, 117, 122.

Paradoxurus larvatus, Figs. 1,6, 13, 14,

pp. 66, 71, 79, 80.

Paraxerus cepapt, Wigs. 19, 28, pp. 215,

228.

—— palliatus, Fig. 23, p.

Petaurista philippensis, Fi

sp, Fig. 29, p. 245.

Petinomys fuscocapillus, Fig. 28, p. 244.

Phoca vitulina, Fig. 24, p. 146.

Phylloscopus affinis, fig. 4, Pl. IIL.,

p. 277.

Platypterna, Big. 2, p. 466.

Plectopterna, Fig. 1, p. 465.

Polemarchus, Fig. 2, p. 466.

Prinia socialis, figs. 24, 28, Pl. IV.,

1 PAU

Procyon otor, Bigs. 3, 9, pp.

228.

gp. 29, p. 245.

Proteles eristatus, Hig. 21, p. 139.

| Protoxerus stangeri, Figs. 19, 23,

pp. 215, 228.

| Psammophis sibilans, Figs. 21, 29,

pp. 317, 322.

Pudu pudu, Figs. 13, 15, pp. 200, 202,

208.

Putorius vison, Fig. 8, p. 73

| Rangifer tarandus, Pigs. 14, 16, pp. 202,

204.

| Ratufa gigantea, Fig. 20, p. 218.

—— indica, Fig. 20, p. 218.

robinsont, Hig, 22

’

p. 223.

Rhizomolgula globularis, P}. I., p. 155.

Seiurus (Parasciurus) niger, Fi

0 PAU).

vulgaris, Fig. 18, p. 213

Stomach contents of a Crocodile, Fig. 1,

p. 180.

19,

XV1

Struthio, Figs. 6, '7, pp. 470, 472.

Struthiomimus, Fig. 5, p. 469.

Suya crinigera, figs. 8, 9, 17, 18,

EV icnp iets

Tamiasciurus hudsonicus, Fig. 18, p. 213.

Tamiops mactlellandi barbei, Fig. 22,

p- 228.

tristriatus, Figs. 18, 20, pp. 213,

218.

Tarbophis variegatus, Fig. 7, p. 805.

Thalarctos maritimus, Fig. 1, p. 160.

Tomeutes hippurus, Fig. 21, p. 22).

—— lokrioides, Fig. 21, p. 221.

— mimatus, Wig. 21, p. 221.

—— robinsont, Fig. 21, p. 221.

INDEX OF TLLUSTRATIONS.

Tomeutes tahan, Fig. 21, p. 221.

— vittatus, Fig. 18, p. 2138.

Tragulus, Fig. 15, p. 208.

Trichechus rosmarus, Fig. 24, p. 146.

Tropidonotus stolatus, Figs. 14, 10-18,

24, pp. 298-301, 309, 310, 321.

Turdoides griseus striatus, fig. 5, Pl. I.,

p. 277.

—— terricolor, fig. 2, Pl. 1., p. 277.

Ursus arctos, Fig. 24, p. 146.

-—— pruinosus, Pls. I., I1., p. 85.

Xerus rutilus, Fig, 24, p. 231.

Xiphopeza, Fig. 1, p. 465.

INDEX.

1923.—Pages 1-481.

[New names in clarendon type.

Systematic references in italics.

(z.8.L.) indicates additions to the Society’s Menagerie. |

Acanthopneuste magnirostris, 288.

oceipitalis, 283, 292.

Acentrophorus, 19.

abbsi, 19, 33, 34.

altus, 19, 38, 34.

chicopensis, 58.

glaphyrus, 19, 20, 27, 33, 39.

varians, 19, 23.

Acomys witherbyi, 46.

Alurus, 150.

4Ethalops alecto, gen. et sp. n.,

178.

Athopyga seherie, 292.

Aithosciurus poensis, 226.

Agama hispida, 50.

Alactaga, 470.

Alcelaphus lelwel tschadensis, 270.

Amia, 41, 45, 451, 455.

Amiurus, 43.

Ammopus, 464.

Andrena fulva, 481.

Anomepus, 464.

Anthropopithecus troglodytes, 323.

Anthus pratensis, 288.

richardi rufulus, 291.

striolatus, 291.

—— trivialis, 288.

Proc. Zoou. Soc.— 1923.

Apatichnus, 464.

Arachnothera magna, 284, 292.

Archeopteryx, 463, 468, 471, 475.

Archegosaurus, 49,

Argya, 280, 282.

Artiodactyla, 470.

Arvicanthis testicularis, 266.

Asillia tridens, 249.

Atelerix albiventris, 251.

Ateles ater (2. s. u.), 480.

Atilax paludinosus, 65, 66, 75, '7'7, 150.

Atlantoxerus getulus, 232.

Atractus trilineatus (u. s. u.), 179.

Auchisauripus, 464.

Awelia, 41, 51, 53-55.

Axis axis, 184, 480.

porcinus, 184.

Belomys (2?) trichotis, 244.

Bos bubalis, 474.

-—— depressicornis, 474.

gourus, 474,

Budorcas taxicolor (z. 8. u.), 178.

Cacatua ophthalmica (2. 8. u..), 175.

Caccomantis passerinus, 286.

XVili

Callosciurus atrodorsalis, 219.

caniceps, 219.

—— castaneoventris, 219,

erythreys, 219.

—— notatus, 219.

pluto, 219.

prevostt, 217, 219.

-—— sladeni, 219.

Canis anthus soudanicus, 254.

bengalensis, 65.

—— faumiliaris, 65.

thous, 65, 67, 72, 74-76, 79.

Capitosaurus, 49.

Cardisoma armatwm, 11, 173.

guanhumi, 11.

—— hirtipes, 11.

Castor fiber, 275.

Centropus, 278.

Cerberus rhynchops, 302.

Cereoleptes, 150.

Cercopithecus tantalus marr-

ensis, subsp. n., 248.

Cervus canadensis, 182.

——- elaphus, 182.

eldi, 184.

—— hortulorum, 183.

sika, 188.

-— xanthopygus, 182.

Cettia cantans, 283.

Chaleococcyx, 291, 292.

Chrysopelea ornata, 304, 314.

Cisticola, 286, 290, 292.

Citellus leursi, 234.

mexicans, 234.

mongolicus, 233.

——- 18-lineatus, 234.

Civettictis civetta, 65, 66, 76, 78, 80,

150.

Clamator, 2717.

—— glandarius, 285.

jacobinus, 279, 292,

Coccysies, 277.

Celogenys paca (4.8. L.), 175.

Cordyceps robertsit, 479.

Corvipes, 464.

Crocidura aridula, sp. n.. 252.

—_— darfurea, sp. n., 251.

INDEX.

Crocidura hindei marrensis,

subsp. n., 252.

—— marita, sp. n., 253.

Crocuta crocotta, 254.

Crossarchus, 150.

Cuculus, 277.

cancrus bakeri, 277, 288, 292.

canorus, 288.

—— -— telephonus, 288, 290.

poliocephalus, 283.

Cynelurus gubatus (2. 8. u.), 175.

Cynictis penicillata, 65, 74, 81, 150.

Cynomys ludovicianus, 236.

Cyon dukhwensis (4. 8 u.), 480.

Cystophora, 151.

Dama dama, 186.

Danis horribilis, 87.

Dapedius orbis, 39.

Dendrophis pictus, 307.

Desmodilliscus brauert, 264.

Dipas trigonata, 303.

Diphyllodes magnifica hunsteinr, 179.

Diplocercides, 51, 53-55.

Diplotriena, sp., 62.

Dipodillus lowei, sp. n., 261.

— muriculus, sp. n., 263.

—— principulus, sp. n., 262.

Dorypterus hoffmanni, 38.

Dremomys dawsoni, 224.

lokriah, 224.

——- rufigenis, 224.

Juscus, 224.

Dryonastes, 282.

Dryophis mycterizans, 302, 315.

Eidolon helvwm, 249.

Elaphurus davidianus, 186.

Enicurus maculatus maculatus, 290,

Eoglaucomys, 243.

— jimbriatus, 241.

Evethizon, 471.

Erythrocebus pyrrhonotus, 248.

Huarctos, 87.

Hudocimus ruber, (uz. 8. u.), 175.

Hudynamis scolopaceus, 281.

Eugnathus, 461.

Hupalamopus, 464.

INDEX.

Eusthenopteren, 50.

Huxerus, 233.

chadensis, 255.

erythropus, 230.

—— —— limitaneus, subsp. n.

255.

Hxocampe, 464.

Felis bengalensis, 65, 72, 73, 77, 149.

caffra, 149.

--— caracal, 149.

—-—- concolor, 149.

—— domestica, 65, 68, 80, 149.

leo, 149.

— nebulosa, 149.

ocreata, 253.

onca, 65, 149.

—-- pardalis, 149,

pardus, 149.

serval, 253.

sylvestris, 65, 149.’

tigris, 149.

——— viverrina, 149.

yaguarundi, 149.

Funambulus palmarum, 215.

Funisciurus congicus, 227.

—— leucostigma, 226.

niveatus, 226,

—— pyrrhopus, 227.

Galago, 469.

—— sennaariensis, 249,

Garrulax moniliger, 280.

—- pectoralis, 280.

Gazella dorcas, 271.

—— rufifrons, 270.

—— subgutiurosa (2. 8.u.), 1709.

Genetta, 150.

—— felina, 66, 72, 75.

Geosciurus, 232.

capensis, 230,

Gerbillus agag, 260.

—— nancillus, sp. n., 260.

—— pygargus, 260.

Gigandipus, 464.

Giraffa camelopardalis tippelskirchi,

179.

Glaucomys volans, 243.

Proc. Zoou. Soc.—1923.

Gorilla gorilla beringeri, 176.

—— —— castaneiceps, 176.

—— —-— diehli, 176.

gorilla, 176.

—— —— matschiei, 176.

Grallator, 464.

Grammomys aridulus, sp.

268.

Graphiurus orobinus, 257.

Haliattus athicilla (a. s.%.), 175.

Halicherus, 151.

Hatlopus, 464.

Helarctos, 88.

Heliosciurus bongensis

canaster, subsp. n., 256.

punctatus, 229.

—— rufobrachium, 229.

hardyt, 227.

Heptanchus, 41, 42, 44, 47, 51.

Herpedactylus, 464.

Hlerpestes sanguineus, 253.

Heteroxenicus nipalensis, 284.

Hexanchus, 41, 42, 44. 47, 51.

Hierococeyx, 277.

Jugaz, 291.

nisicolor, 291.

—— sparveroides, 283, 284.

varius, 279, 282.

Hipposideros caffer, 249.

Hyena hyena, 254.

Hydropotes ineriis, 192.

Hyleocherus, 474.

Hylephitas obtusa, 481.

Hylopetes alboniger, 242.

phayret, 245.

Hyphepus, 464.

Hyrax capensis, 68.

Hystria, 471.

Ictonyx striatus sudanicus,

subsp. n., 254.

--— zorilla, 65, 66, 73, 74, 81,

Jaculus gordont, 269.

jaculus, 269.

Lacerta, 50.

Lamna cornubica, 161.

Lanius schach erythronotus, 290.

Lariscus jalorensis, 223.

Larvivora brunnen, 290.

Leander adspersus, 98, 106.

longirostris, 97, 98, 108.

serratus, 98, 99.

sguilla, 98; 102.

— varians, 98, 114.

Leggada tenella, 269.

Lemuiscomys dunnt, 267.

—— nubalis, sp. n., 267.

—— lynesi, sp. n., 267.

Lepidosteus, 41, 42, 44, 46, 51, <H5,

451, 45, 474.

-— ossens, 445,

-—-— platystomus, 445.

tristoechus, 445.

Lepidotus, 23, 31.

Lepus hawkeri, 269.

Liothrix lutea, 291.

Lutra maculicollis, 72, 150.

Lycaon pictus, 65.

Lycodon auticus, 310.

Macropus giganteus (z. 9. .), 179.

Manatus inunguis, 273,

Marmota marmota, 236.

Mazama tema, 200.

Meles meles, 63, 66, 150.

Melursus ursinus, 65, 66, 73, 80, Sl.

Menetes berdmoret, 219.

Mephitis mephitica, 65, 66, 71, 75, 80.»

Mesia argentauris, 291.

Mungos, 150.

ichneumon, 63, 67.

mungo, 69.

Mustela martes, 65, 66, 150.

Mylarctos, subgen. n., 91.

Mystax illigert (4.8. .), 479.

Nandinia binotata, 65, 71, 75, 150.

Nannosciurus exilis, 225.

micrates, 225.

whiteheadi, 225.

Nasua narica, 65, 150.

—— viltata (2. s. L.), 479.

Niltava macrorigorie (z. 8. u.), 17%.

Nyctes scandiaca (z. 8. u.), 175.

Nyctinomus tongaensis, 251.

INDEX.

docoileus virginianus, 196,

peruvianus, 198.

spinosus, 198.

Opisthocomus, 475.

Oreicola ferrea, 290.

Ornitholestes, 468.

Orthotomus, 286.

| Osteoglossum, 44, 46, 51.

Otaria, 151.

Otogyps auricularis (a. 8. u.), 479.

Ourebia ourebi splendida, 270.

Oxybelis fulgida, 312.

Palemonetes varians var, mesogenittor,

121.

Paleoniscus, 461.

=== aioe, WY).

altus, 19.

—- yarians, 19.

Palamopus, 464.

Papio anubis, 249.

Paradoxurus larvatus, 65, 66, 72,

77, 150.

Paraxerus cepapt, 227.

Penthoceryx sonnerati, 291.

Petaurista, 240,

——— philippensis, 245,

Petinomys fuscocapillus, 243.

Petrogale godmani, sp. n., 177.

Phacocherus elhiopicus, subsp., 270.

Phoca, 151. ‘

Phocena communis, 83..

Phractolemus, 450.

Phylloscopus, 283.

Pipistrellus marrensis, sp. n., 249.

Planorbis dufouri, 480.

Platypterna, 464.

Plectoterna, 464.

Polemarchus, 464.

Polyodon, 57.

Polypterus, SG.

Prinia tnornata, 286.

socinlis, 286. —

Proavis, 468, 464, 467, 469, 475.

Procavia ruficeps, 271.

—— —— marrensis, subsp. n..

Procompsognathus, 468.

Procyon lotor, 65, 72, 156.

73;

z INDEX.

Protoxerus stangeri, 226.

Psammophis sibilans, 305, 317.

Pudu pudu, 201.

Putorius vison, 65.

Raillieticlla furcocerea, 15.

—- geckonis, 18.

indica, 18.

Rana, 48.

Rangifer tarandus, 204, 479.

Rattus (Mastomys) macrolepis, 265.

ugande, 266.

Ratufa fellii, 217.

gigantea, 217.

indica, 217.

Rheithrosciurus macrotis, 214.

Rhinoceros, 474.

bicornis, 175, 474.

sumatrensis, 474.

Rhizomolquia globularis, 150.

Rhopocichla nigrifrons, 293.

Rhopodytes, 278.

Sauropus, 464.

Saxicola torguata indica, 290.

Schistosoma (Bilharzia) hematobium,

480,

Scturopterus russicus, 243.

Sciurus carolinensis, 212.

-—— ingramt, 212.~

—— langsdorffi, 212.

a= lig, BZ

—— nayoritensis, 212.

—— (Neosciurus) carolinensis, 212.

-— niger, 212.

persicus (¥.8.L.), 179.

variabilis, 212.

vulgaris, 212.

Scomber, 43.

Scoteinus schlieffeni, 250.

Scotophilus nigrita, 250.

Sphenodon, 50.

Steatomys aquilo, sp. n., 264.

—— gazelle, sp. n., 269.

Struthiomimus, 468.

Suricata, 150.

Surniculus, 291.

Sus, 474.

— barbatus, 474.

cristatus, 474.

scrofa (Z. 8. u.), 175.

Sylvicapra grimmia pallidior, 270.

Tamiasciurus hudsonicus, 213.

Tamiodes, gen. n., 215.

—— tristriatus, 215.

Taphozous mauritianwus, 251.

Tarbophis variegatus, 305.

Tarsius, 469.

Tatera benvenuta, 257.

robusta, 257.

Taterillus, 257.

butleri, 260.

—— clivosus, sp. n., 25S.

-—— perluteus, sp. n., 259.

rufus, 259.

Tetrapteryx, 463.

Thalarctos maritimus, 159.

Tomeutes hippurus, 222.

—.- lokrioides, 220.

—— miniatus, 222.

rohinsont, 222.

— tahan, 222.

vittatus, 220.

Tremarctos ornatus (az. s. u.), 179.

—— thibetanus, 88.

Trichechus, 151.

Trochalopterum, 282.

lineatum, 290.

Tropidonctus stolatus, 808.

Turdoides, 280, 282.

Ursus, 150.

lastotus, 91.

pruinosus, 85.

Viverra civetta, 65, 150.

Viverricula malaccensis, 65, 150.

pe , ORS

Vulpes pallida, 254.

vulpes, 6d.

Wimania, 41, 51, 53, 53.

Xerus rutilus, 232.

Xiphopeza, 464.

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XX1

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: LIST OF CONTENTS

1928, Parr I. (pp. 1-180).

EXHIBITIONS AND NOTICES.

Tue Secretary. Report on Additions to the Society’s Menagerie during the months of

November and December, 1922

Pr Ce eC ee Cc CC Cr

Lord Roruscuity, F.R.S., F.Z.S. Hxhibition of adult male Mountain Gorilla ......--

Mr, Ouprieip Tuomas, F.R.S., F.Z.S. Exhibition of a new Rock Kangaroo from Northern

Queensland

eo resem Ree ee te ee ee Foe eee ee ee ee OH OOOH Ee ee essere sess e Bes erseee

Exhibition of the Skull of a Pigmy Fruit-Bat from Sumatra

Mr. J. B. Scrivenor, M.A. Exhibition of ook. showing native method of breaking

in captured Elephants

seth ew ee ei te eh et ee eee ew tO ee Ose wee eeese Foe - sc 'e te oore

Mr, E. G. Boutuyerr, F.Z.S. Account of his recent visit to Vienna

se ee seeveerce e@ oe

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

January 1923

weet ee ee ee FP Pe SO rere een ee OFF seers eesrssereseeseeseee ee eeS8ss veer:

Mr. D. Sern-Smiru, F.Z.8. Exhibition of Photographs and skins of Birds-of-Paradise . -

Prof. H. M. Lerroy, F.Z.S. Exhibition of Saunas film of the life-history of the

MEV O USC atliy mundi oeans een area aret ay veer uate te answer SC oke ey sia sta NaS. Bayete te Sea at oiel LOT ae EO Boab es

Tun Secretary. Exhibition of a photograph of the Polar Bears “ Sam” and ‘“ Barbara ”

Mr, R. I. Pococs, F.R.S., F.Z.S. atubiien of, and remarks upon, drawings of the feet

and nose of the Polar BOP oe ntusg obo un CIOd at on 4 a ofan oeVehisiaire/ siete Tevellnlai|lets) siclel atekersyelel ors)

Tue Secrerary. Note on a case of apparent melanism in Tippelskirch’s Giraffe ...

Mr. BR. I. Pocock, F.RS., F.Z.S. Exhibition of drawings of the feet and spurs of the

Echidna.......... PraUerTaleKeiaistosss) = WekeNhe: ites ots Saale seta ee Pe EST Ae RT landcare

Sir G. ABERCROMBY. Ltamiion on hendtohs Kobi ee ae Bei rare ge oe MRC

Prof. HE. W. MacBripz, F.R.S., F.Z.S. Exhibition of photomicrographs of sections

through nuptial callosities of Rand and Alytés 10.05.06 veces ecb cu css cbse cscs aa

Mr, F. M. Swynnzrron, C.M.Z.8. Exhibition of stomach-contents of a Crocodile.

(Text-figure 1.) . a a

180

Contents continued on page 3 of Wrapper.

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PROCEEDINGS

OF TIE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OV THE

ZOOLOGICAL SOCIETY OF LONDON.

PAPERS.

1. Coat Colour in Greyhounds.

By Aparr Dicuton, F.R.C.S8., F.Z.8.

[Animal Breeding Research Department, University

of Edinburgh. |

[Received October 13, 1922: Read February 6, 1923.]

Some months ago, my friend Mr. J. R. Robertson, knowing

that I was interested in both the problems of Heredity and in

Greyhounds, suggested that I should make extractions of their

various coat colours from the stud books, and tabulate them on a

definite plan with a view to ascertaining whether the results

accorded with Mendelian principles and with previous data

obtained by him from direct experiment. At the time Mr.

Robertson suggested this, I was too busy with other affairs te

undertake it, but at the end of the last coursing season | began

what has been a hard but fascinatingly interesting work.

By way of preface, I should explain that the National Coursing

Club is the ruling body in the coursing world, and under its

direction the Greyhound Stud Book is issued annually. A rule

of the Club stipulates that every litter of greyhounds must be

registered within two months of the date of whelping, with the

names of the sire and dam, and the colour (subject to correction

within six months of the date of whelping), sex, and number of

the puppies. In this way there is in the Greyhound Stud Book of

the present day an authentic record of every greyhound born.

Proc. Zoou. Soc.-—1923, No. I. 1

HY MR. ADAIR DIGHTON ON

From the point of view of accuracy in the present statistics it

is unfortunate that colour registration should have to be made

at so early an age, for there is no doubt that in many cases

colour changes with age, and there may be, in fact are, cases in

which the colour would have been registered differently had the

date of registration been postponed. There is one other point

to which I would draw your attention. Under the heading of

“ Registration Regulations” there is the following instruction :—

“It is not desirable to register a colour which is not seen

by the judge when following the dogs. Thus it often

happens that a black dog has an insignificant patch of white

on his chest, it may be a few white hairs on the tip of his

tail or a white toe, The entry of such a dog as black and

white simply leads to confusion, and should be avoided.”

From the practical standpoint this is an excellent rule, but it is

not conducive to that accuracy in description which is of such

vital importance when dealing with colour from a scientific or

statistical point of view.

Bearing these points in mind, I began with the Grey-

hound Stud Book for the year 1907, and have worked through the

fifteen succeeding volumes up to, and including, that pub-

lished in 1921. In this way the records of 16,260 litters have

been examined and the colours of 25,767 whelps have been

tabulated. The pigmentary factors of greyhounds resemble those

of other animals in that they consist of black, chocolate (known

as red), and yellow. In addition, the greyhound has a composite

colour known as brindle, which is due to a pattern factor

combining with one, or more, of the other colours. The results,

generally, may be taken to represent the actual colours of the

various individuals, but personal observation shows that there

are certain discrepancies. Thus there are blacks, so registered,

that show brindle markings, and the returns do not differentiate,

in the majority of cases, between red and fawn, the usual descrip-

tion being “red or fawn,” which explains why it has been

necessary for me to group these two colours together under one

heading. Pure albinism of the skin and hair is very rare, but

partial albinism is common in the pied patterns. Other colours,

or combinations of colours, such as blue-brindle, blue-fawn, black-

brindle, etc., are often met with, but these are too ambiguous to

be taken into account and are not included in my figures.

Take first the matings of black to black. On the assumption

that black and blue are dominant over all other coat colours,

the blacks can be of two kinds: pure dominant blacks, containing

nothing but black factors, and impure dominant blacks, con-

taining one black factor and one alternative factor, which may

be either brindle, or red, or fawn. Now, though pure dominants

do occur in all colours, they are rare. Greyhounds are bred

for speed and work. Colour is a minor consideration, and of the

theusands of dogs born, only a very small percentage of them

COAT COLOUR IN GREYHOUNDS. 3

ever stand at stud, and, of these, only very few have a sufficient

number of mates upon which an assumption of pure dominance

could be based. For these reasons it may be taken that the

parents concerned in the 500 black to black matings were impure

dominants in gametic composition. The 500 matings yielded

3603 whelps, and, on the basis of the classical Mendelian ratio

of 3:1, the expectation is that there would be 2700 blacks to

900 brindles, red or fawns, and whites. The observed results

gave 2697 blacks and blues to 906 brindles, red or fawns, and

whites.

In the black to biue matings, of which there were 231, there were

1338 whelps, and the expectation is the same as in the black to

black matings: namely, three blacks and blues to one other colour.

This would give 1003 biacks and blues to 335 brindles, red or

fawns, and whites. The actual results were 998 to 340.

These figures clearly establish the premise that black is the

epistatic colour, and is dominant over all others.

Take next the matings of black to brindle. The case here

is one of an impure dominant to a recessive, so that the

resulting offspring should be an equal number of impure domi-

nants and of recessives. That is to say, there should be an equal

number of blacks and blues to brindles, red or fawns, and whites.

The number of whelps in the 500 matings was 3053, so that there

should have been 1526 of each. The actual result was 1533

blacks and blues to 1520 brindles, red or fawns, and whites.

In the black to red or fawn matings the case is again one

of impure dominant to recessive. Again equality would be

expected, so that of 3039 whelps there should be 1519 of each.

The actual result was 1525 : 1514.

These figures still further conclusively prove that black is

epistatie to brindle and to red or fawn.

Take now the blue matings. I have taken up the black

to blues. In the blue to brindle matings the case is one of

impure dominant to recessive, and the expected result from

1293 whelps would, therefore, be 646 blacks and blues to 646

brindle, red or fawns, and whites. The actual result was

649 : 644.

Blue and red or fawn matings fall into the same scheme, and

from 1822 whelps the result should have been 911: 911, whereas

it was actually 915: 907.

The blue to blue matings only numbered 25, and resulted in

132 whelps. Here the scheme is impure dominant to impure

dominant, which should give 3 blues and blacks to 1 brindle,

red or fawn, and white. The expected result was therefore

99 blues and blacks to 33 brindles, red or fawns, and whites.

The observed result was 93 to 39.

In the table it will be seen that in these matings there were

6 blacks and 1 black and white, and this, or these, raise a

difficulty. Blue is undoubtedly a dilute black, and the previous

small incidence of blues clearly shows that if there is a special

4 MR. ADAIR DIGHTON ON

dilution factor, it cannot be epistatic to the saturation factor.

In these circumstances it must be recessive, and being so, there

should be no blacks in the blue to blue matings. Very possibly

the blacks were dark blues verging on black, but, anyhow, the

blue to blue matings are too few to form a reliable guide.

Turning next to the brindle and brindle matings. On the

hypothesis that brindle is recessive to black and blue and domi-

nant over red or fawn and white, there should be no blacks or

blues from brindle to brindle matings. Theoretically, with the

blackness stripped off, the brindles become dominant or, as

dominants are rare, impure dominant to the recessive red or

fawns and whites. Thus the mating is one of impure dominant

to impure dominant, and the expected result from 2829 whelps

would be 2121 brindle to 707 red or fawns and whites. The

actual result was 30 blacks and blues: 2118 brindles and 681

red or fawns and whites.

The 30 blacks were made up of 14 blacks; 9 black and whites ;

6 blues and 1 blue and white. The error of deviation is so

small that I think it may be laid down to inaccuracy of

description.

In the case of brindle to red or fawn, there were 2932 whelps,

and as it was a case of impure dominant to recessive the expec-

tation was that there would be equality or 1466 brindles to

1466 red or fawns and whites. The actual result was 1465 toe

1458, but there were, in addition, 9 blacks, which once again,

though unsatisfactory, must be attributed to descriptive error.

The last of the more simple results are reached in the matings

between red or fawn and red or fawn. On the hypothesis that

red or fawn are hypostatic to everything but white, the expec-

tation is that there would be no colours in the 2925 whelps but

red or fawns and a few whites. Actually, the figures observed

were 10 blacks, blues and brindles: 2910 red or fawns and

5 whites. ‘I'he incidence of the epistatic colours black, blue, and

brindle is so low that one is justified in assuming that their

presence in this mating is due to personal error rather than to

genetics.

Reviewing the above results, I think I have proved that

greyhound coat colours follow the sequence of black, which is

epistatic, blue, brindle, red or fawn, and white, which is hypostatie,

and that, secondly, blue is a dilute black.

From supplementary statistics I find that there are undoubtedly

pure dominants for both black and brindle, and, as I have shown,

ved or fawns breed true, which is invariably the case with a

recessive character. Thus it follows that if such were of any

practical use to the coursing man, he would have no difficulty in

establishing pure breeding strains of either black, brindle, red

or fawn.

IT now turn to the secondary results of the matings, and these

are somewhat more complicated than the primary data already

discussed.

COAT COLOUR IN GREYHOUNDS. 5

In the black to black matings, if the blacks are taken‘as

impure dominants and of two sorts:

(1) blacks carrying a brindle factor as recessive,

(2) blacks carrying a red or fawn factor as recessive,

and the incidence of both blacks are equally numerous, the ratio

arising should be:—black, 12 : brindle, 3 : red or fawn, 1.

Further, if every black dog or bitch carried both brindle and red

or fawn factors, the incidence would be the same, as red or fawn

could only appear in the absence of brindle.

It will be observed that the incidence of black came out cor-

rectly in the black to black and blue to blue matings, but in the

place of the brindles being in the ratio of three to one red or fawns,

the colours are almost equally distributed in the black to black

matings, whereas in the blue to blue matings the red or fawns

ave almost twice as numerous as the brindles. These results

eannot be explained by the law of probability, based on the inter-

action of factors avising out of the hypothesis, already proved,

that brindle is dominant over red or fawn,

In the mating of black to brindle, the Mendelian expectation

is:—black, 4: brindle 3, red or fawn 1; and the 3053 whelps

should therefore give 1526 blacks : 1145 brimdles and 381 red or

fawns and whites. The observed results gave 1533 blacks and

blues: 1018 brindles and 484 red or fawns and whites.

Here the results are sufficiently close—having regard to the

fact that the actual incidence of the recessive factors in the

DR blacks is unknown—to be taken as in accordance with

probability.

On the other band, the Stud Book returns for black to red or

fawn show, as regards the incidence of brindles and red or fawn,

a totally unexpected result. Im the 3039 whelps the expected

result was that there would be 1519 blacks: 789°5 brindles and

789-5 red or fawns; whereas the actual result was 1525 blacks:

492 brindies, and 1007 red or fawn and whites. At the moment

the explanation of these figures is beyond me, but I think it is

worth noting that in the black to black matings the incidence of

brindles and red or fawns is practically equal, and the same holds

true in the black to blue matings. In the black to brindle and

the black to red or fawn matings there is again a similarity, as

in the former the ratio of brindle to red or fawn is as 2 to 1,

whilst in the latter the ratio is practically reversed. This may

be nothing more than a coincidence of figures, but somehow I

think it holds the key to the explanation. My own idea was

that black carries a factor which produces either brindle or red

according as to the factor introduced by the other parent, but

if this were so, in the black to red or fawn matings there

should be no brindles, whereas there were actually 492 in

3039 whelps.

In passing to the mixed colours, it is interesting to note that

6 MR. ADAIR DIGHTON ON

throughout the series the mixed combinations very generally

follow the same incidence of ratio as the whole colours with

which they are associated.

Let me now devote a few words to the mixed matings—that is

to say, the “ pied ” colours, in which white plays a leading part

on both sides. These unions, on the whole, fall into line with

those of melanistic character. One would have expected a higher

incidence of whites in these matings, but it is interesting and

important to note that in the red or fawn and white to red

and fawn and white, the hypostatic colours in the scale, there

are more whites than in any other matings. It seems probable

that the mixed colours tend to segregate into the conditions of

melanism and albinism of the skin and hair, giving 1 whole

colour : 2 mixed colours : 1 white. When we recall the instruc-

tion concerning the registration of colours, it is self-evident

that there is a very appreciable chance that some mixed colours

are nearly white. This being so, there should be about 3 ‘‘mixed”

to 1 whole colour in the pied matings.

Adding the ‘“‘ white” to the “ pied” in the whole series, it

will be found that the total is 2413 “ pied” and white to 689

whole colours, which gives a ratio of 3°5 to 1, which is sufficiently

close for all practical purposes.

In conclusion, I must thank Mr. Robertson for all the

help he has given me in drawing up this paper and Mr. Martin

Dunean, our Librarian, for his kindness in furnishing me with

what references on the subject there are.

WHOLE COLOUR MATINGS.

Brack X BLAcK.

Matings. 500,

Whelps. 2603.

Bk. 1607 Be. 196 Bd. 314 R. or F 271 W. 21

REW. 796 BeW. 98 Baw. 162 R. or FW. 138

2403 294, 4.76 409 2

2697 906

Hupectations 2700 900

Brack X Brove.

Matings. 231.

Whelps. 1338.

Bk. 616 Be. 226 Bd. 112 Ivor sh (62 Wert

BkW. 118 BeW. 38 BdW. 32 Ror FW. 27

COAT COLOUR IN GREYHOUNDS.

Brack X BRINDLE.

Matings. 500.

Whelps. 3053.

Bk. 1115 Be. 123

BkW. 258 BeW. 37

1373 160

1533

Hexpectation. 1526

Brack X Rep or Fawy.

Matings. 500.

Whelps. 3039.

Bk. 1139 Be. 146

BkW. 223 Bew. 17

1362 163

1525

Heapectation. 1519

Buvue X Buve.

Matings. 25.

Whelps. 132.

BiG Be. 76

BkW. 1 BeW. 10

Z 86

Expectation. 99

BLUE X BRINDLE.

Matings. 213.

Whelps. 1293.

Bk. 369 Be. 175

BEW. 73 BeW. 32

442 207

649

Hepectation. 645

Buvue X Rep or Fawn.

Matings. 291.

Whelps. 1822.

Bk. 519 Be. 249

BkW. 116 BeW. 31

635 280

915

Hex pectation. 911

Bd. 869 R. or I’. 395 W. 18

BdW. 149 Li. or FW. 89

1018 484. 18

1520

1526

Bd. 400 R. or F. 877 W. 15

BdW. 92 R. or FW. 130

492 1007 15

1514

1519

Bd. 12 18 @ie 18, BBh Ww. —

BadW 1 R. or FW. 3

13 26

Bd. 349 Ven, Oe IM. Nets) W. 2

Baw. 74 R. or FW. 30

423 219 2

644,

646

Bd. 164 R. or F. 598 W. 6

Baw. 50 R.or FW. 89

214. 687 6

907

911

8 MR, ADAIR DIGHTON ON

BRINDLE X BRINDLE.

Matings. 500.

Whelps. 2829.

Bk, 14 Bes Bd. 1898 IR, Ow 18, | BRUT Ww. 18

il BadW. 220 R. or FW. 139

23 i 2118 663 18

30 2118 681

Expectation. 0 2121 On

BRINDLE X RED or Fawn.

Matings. 500.

Whelps. 2932.

Ite @) Be. — Bd. 1343 R.orF. 1287 W. 4

BkW. — BeW, — BdW. 122 R.or FW. 167

9 — 1465 1454 4,

9 1465 1458

Expectation. 0 1466 1466

Rep or Fawn X Rep or Fawn.

Matings. 500,

Whelps, 2925.

Bk. 2 Ba, @ Bd. 1 R. or F. 2550 W. 5

BEW. 4 BeW. 1 Baw. — R.or FW. 350

6 3 1 2910 5

10 2910 5

Expectation, 0 2920 5

MIXED COLOUR MATINGS.

Brack and Wuirr X Buack and WHITE.

Matings. 100.

Whelps. 532.

Bk. 92 Be. 16 Bd. 24 R. or F, 32 W. 4,

BkW. 280 BeW. 12 BdaW. 56 R. or FW. 16

Hapectation, 399 133

Brack and WHITE X BRINDLE and WHITE,

Matings. 100.

Whelps. 552.

Bk. 48 Be. 16 Bd. 64, R.or PF. . 44 W. 4

BkEW. 176 BeW. 32 BdW. 116 R. or FW. 52

ee eeeSSSSsSsSsSsSsSsSssssssFsssesFseseEeEeee

224, 48 180 96 4,

COAT COLOUR IN GREYHOUNDS,

Brack and Waits X Rep or Fawn and WHITE.

Matings. 100.

Whelps. 564.

Bk. 72 Be. 10 Bad. 299, R. or F. By) W. 22

BkW. 184 BeW. 14 Baw. 78 R. or FW. 110

256 24 100 162 22

280 284

Expectation. 282 282

BLveE and Waite MatTrines too few to be used.

BrinpiLeE and WuITE X BRINDLE and WHITE.

Matings. 100.

Whelps. 430.

i es Be. — Bd. 50 R.ork. 34 W. 5

BkW. — Be Wim BdW. 270 R. or FW. 71

= = 320 eels 5

320 110

Hxpectation. 321 107

BrinpiE and WHITE X Rep or Fawn and WHITE.

Matings. 100.

Whelps. 502.

Bk. Sie Bes Bd. 32 R. or F. 68 W. 20

BkW. 14 BeW. 4 BaW. 220 R. or FW. 128

22 12 252 196 20

252 216

Buxpectation. 0 0) 251 2a1

Rep or Fawn and Wuite X Rep or Fawn and WHITE.

Matings. 100.

Whelps. 631.

BE = a, Bd. — R.or EY 53 W. 62

BkW — BeW. 1 BdW. 2 R. or FW. 513

= 2 566 62

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Hepectation. 0

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ON THE ZOHA OF CARDISOMA ARMATUM. 11

2. A Note on the Zoza of a Land-Crab, Cardisoma armatum.

By H. Granam Cannon, B.A., F.ZS.

[Received October 24, 1922: Read March 6, 1923. ]

(Text-figures 1-6.)

Among the Land-Crabs it is known that the females of the

genera Gecarcoidea and Gecarcinus resort to the sea in order that

the young aquatic larve, on hatching from the eggs, may swim

away into the water. According to Moreira [1912] ripe females

of Cardisoma guanhumi also resort to salt or brackish water for

this purpose, whereas, according to Calman {1909!, neither

Ortmann nor Andrews observed any specimens of the Cardisoma

hirtipes in the neighbourhood of the sea.

During September 1922 a female land-crab of the species

C. armatum Herklots, kept in captivity in the Society’s Gardens,

was noticed to be carrying a large mass of eggs underneath the

abdomen. The eggs proved, on examination, to be fertile, and in

a late stage of development. It was obviously of interest to find

out whether the eggs of this species would hatch in fresh or in

salt water.

Some eggs were removed from the female and placed in pure

artificial sea-water, while others were placed in tap-water. A

third lot were put into brackish water.made of equal parts of sea-

water and tap-water. The next day all the eggs appeared healthy,

and the embryos in most showed a fairly regular heart-beat.

After two days these eggs in the tap-water had become opaque,

more especially in the central part of the egg, while those in

brackish water and sea-water appeared to be developing normally.

After three days many of the sea-water eggs hatched out and the

young appeared quite vigorous. In the brackish water only a

few hatched out, but these also appeared quite healthy. In the

fresh water some of the eggs had burst the shell, but the young

had disintegrated. It appears very probable from this that

C’. armatum also resorts to the sea, or at least to brackish water

for spawning-purposes.

The larvee only survived a few days after hatching and did not

develop further, although they were being offered the diatom

Niteschia as food. It could not be seen whether the larve were

actually taking the diatoms into their stomachs.

The larvee were typical Brachyuran zoe. The frontal spine

slopes downwards and then curves upwards, and ends by re-

curving down again (text-fig.1). There isa pair of lateral spines

projecting outwards and slightly forwards (text-fig. 2). All the

spines of the carapace are simple. The antenne are very simple

12 MR. H. G. CANNON ON THE ZOMA

(text-figs. 3 & 4) and the second antenna does not bear a lateral

spinous process (text-fig. 4). The lst abdominal segment bears

at each side a short forwardly directed spine, the other segments

being unarmed (text-fig. 2), The endopodite of the 1st maxilliped

is five-jointed, the second joint being the longest (text-fig. 5).

The exopodites of the Ist and 2nd maxillipeds end in a group of

four short-pointed setz arranged side by side (text-fig. 5). There

is a large branched pigment-spot on the ventral side of the

abdomen between each pair of segments and a small circular

patch of pigment on the distai end of the protopodite of the

Ist maxilliped (text-figs. 1 & 2).

Comparison with the description given by Moreira [1912] for

Text-figure 1. Text-figure 2.

Lateral aspect of Ist zozea of Cardi- Dorsal aspect of 1st zowa of Cardi-

soma armatum. X 88. soma armatum. X 88.

the zoza of C. guankwmi shows only one marked difference. In

C. quanhuma it is stated that the cephalothorax is provided solely

with a dorsal spine, the anterior and lateral spines being absent.

Between two such nearly related forms as ©. guanhumi and

C. armatwm it would not be expected that such a difference would

occur. However, the figures illustrating Moreira’s paper suggest

that a re-examination of the South-American form might be

desirable.

An account of the Ist zoxa of Gelasimus by Hyman [1920]

indicates a close similarity with the zozxa of C. armatum. In this

form, however, there are no lateral spines on the cephalothorax,

aithough the anterior spine is present.

OF A LAND-CRAB, CARDISOMA ARMATUM. 13

Among twenty or thirty larve preserved for examination it

was noticed that two specimens diifered from the rest in the form

of the Ist and 2nd maxillipeds. In these two specimens the

exopodites, instead of ending in four short spines as in all other

specimens, ended in a brush of four plumed hairs, each about the

jength of the remainder of the endopodite. In both C. guanhumi

Text-figure 3.

oes

Ist antenna.

Text-figure 4.

2nd antenna.

Text-figure 6.

Text-figure 5. \y //

af iN if Hi *

Se = | if \ | .

Ist and 2nd maxillipeds,

seen from inside.

Telson.

and in Gelasimus the exopodites of the maxillipeds normally end

in a similar structure. It cannot be said with any degree of

certainty what this represents. It may be that in these two

specimens the Ist zoxa has already developed characters that

would normally appear on a later instar, but, as no specimen was

14 ON THE ZOMA OF CARDISOMA ARMATUM.

reared to the second larval stage, this cannot be verified. It is

interesting to note in this connection that Miss Webb [1919], in

the larve of Upogebia, found two distinct forms. The difference

in this case concerned the number of exopodites furnished with

swimming sete occurring on the thoracic Jegs, and became first

evident at the commencement of the 2nd larval stage. Possibly

the case recorded here is an analogous case of polymorphism.

LITERATURE.

Catman, W.T. 1909. P.Z.S. 1909, p. 703.

Hyman, O. W. 1920. Journ. Morph. xxxiii. p. 485.

Moreira, C. 1912. Mém. de la Soc. Zool. de France, xxv.

p. 155.

Wess Gab e!919) “Journ MB Ae <p molle

THE LINGUATULID ARACHNID RAILLIETIELLA FURCOCERCA. 15

3. On the Linguatulid Arachnid Raillietiella furcocerca

[Diesing, 1835] Sambon, 1922. By Grorce §8.

GieLroL1, M.D. Pisa,

[Received November 1, 1922: Read February 6, 1923.]

This species was described by Diesing in 1835 under the name

of Pentastoma furcocercum, from specimens collected by Natterer

in South America. His original description, translated from the

German, runs as follows :—

“The worms are from 9 to 10 lines in length; 1 line thick

at the anterior extremity, 4 at the posterior. They are grey in

colour, while their internal organs, which can be seen in trans-

parency, are brown and somewhat marked in white and reddish.

The body is spindle-shaped and presents a linear annulation,

which is most marked at the middle part of the body, while it

gradually disappears towards the two extremities. On each ring

are numerous stigmata disposed along a line at regular intervals.

The cephalic extremity is somewhat flattened and is obtusely

triangular. At its apex it presents the oval oral opening. The

hook-pouches, which are situated symmetrically behind the mouth,

are borne by obtusely conical structures. Hach pouch is pro-

tected by three rounded pads, two lateral and one median, which

is situated over the hook. Hach pouch contains two hooks, pro-

bably united at their base; they are situated one over the other.

The upper one is nearly straight and short; the lower one is

longer and more medially directed. The posterior extremity is

bifid. The male can be distinguished from the female by the

penis, situated on a papilla on the mid-ventra! line on the first

anterior ring. I have found the two caudal points divergent in

a female, but the two sexes are similar in size.”

Later, in his ‘Systema Helminthum,’ Diesing again describes

the same species under the name Pentastoma bifurcatwm :—

“Corpus subfusiforme retrorsum attenuatum, extremitate

caudali bifurcatum, annulato-plicatum, annulis linearibus. Caput

obtuse triangulare depressiusculum. Os ovale margine lato cal-

loso apice truncato, inter bothria obtuse triloba in formam coni

truncati disposita situm. MHamulis inequalibus. Long. fem.

9-13'"; crass. antice ultra 1’”; postice 4°". Femina cauda sua

bifurcata, in aliis emarginata, marem multo minorem et gracili-

orem sub coitu probabiliter excipit.”

In 1898 Sir Arthur Shipley, in his monograph on Lingua-

tulide, reproduced Diesing’s description under the name Poro-

cephalus bifurcatum.

In 1910 Sambon included this species in the genus Raillietiella,

and, following the rule in zoological nomenclature, again adopted

16 DR. G. S. GIGLIOLI ON THE

the original specific denomination given by Diesing in 1835, so

that this tongue-worm is now known under the name of Raillie-

tiella furcocerca.

Through the kindness of Dr. Khalil, of the Department of

Helminthology of the London School of Tropical Medicine, I

have been able to examine two female specimens of this parasite,

collected from the lungs of an Amphishena alba, which died in

the London Zoological Gardens, and I would now submit the

following as a more accurate description of the external mor-

phology of this species :—

Length 26 mm.; breadth 1 mm. (anteriorly), The cephalo-

thorax is the bulkiest part of the body, and from it the abdomen

tapers gradually towards the caudal extremity. The body is

straight and delicate. The cephalothorax is roughly triangular

in shape, with the generic characters of Raillietiella very well

marked. The mouth is terminal, but situated on the ventral

surface. The oral papilla is shield-shaped and large, with a well-

marked chitinous outline. It is surmounted by two small

rounded papille, which are terminal. Laterally,and slightly dor-

sally to these, other two long finger-shaped papille are situated,

which are directed laterally. These structures occupy the apex

of a large fleshy cone which forms the anterior part of the

cephalothorax, and is very slightly flexed ventrally. At its base

the anterior pair of hooks are situated. The whole chitinous

hook-apparatus is remarkably small, especially the root. They

are supported, or rather enclosed, by a voluminous fleshy struc-

ture, which protrudes laterally, so that when the hook is at rest

not even its tip is free. The hooks are, in fact, only visible by

transparency. Their fleshy base presents a somewhat complicated

form, in which three parts can be distinguished: an external

rounded pad or mass situated laterally to the hook and slightly

behind it; a similar but smaller structure symmetrically situated

on the internal aspect of the hook; and a compressed, trans-

parent hood, situated over the two preceding organs, and which

shrouds the hook, both root and blade. The hood is open at its

lower end, so as to allew the extrusion of the hook-blades, but

these are so small that only a very little part of the blade can

ever be freed from its fleshy sheath. The hooks are smooth and

simple, the posterior pair slightly larger, and supported by a

much larger and stronger base. As in other species of the genus

Raillietiella, the large sensitory papille, corresponding to the

anterior or medial pair of hooks, are finger-shaped and of unusual

size, and situated on the latero-dorsal surface of the cephalo-

thorax. Behind these and a little nearer to the median dorsal

line two other smaller papille are visible. These I have not

observed in other species (2. bouwlengeri, R. geckonis).

The abdomen is fairly cylindroid in its anterior third, and then

tapers gradually to the posterior extremity, which is acuminated

and bifid. The annulation, though present, is practically effaced,

so that the number of rings cannot be given. On the first five

LINGUATULID ARACHNID RATLUIETIELLA FURCOCERCA. lied

body-rings very distinct acuminated papillz are symmetrically

situated on the latero-ventral margin. The genital opening is on

the mid-ventral line on the first body-ring. The cuticle covering

the body is studded with small eminences which give it a rough

appearance, particularly evident on the large papille. The body-

wall being very fine and transparent, some anatomical characters

can be made out :—

The intestine is straight and dorsal; it receives the short

arenate oesophagus slightly behind its anterior end, and opens

dorsally by the anal orifice, which is situated between the two

caudal appendages. The spermathece are rounded and symme-

trically situated a little dorsally to the vagina, which opens

through the anterior genital pore. The uterus is an ample sac

which fills the whole body-cavity from the spermathece to the

caudal extremity and partly embraces the intestine. It is full of

eggs. On the internal surface of the body-wall the parietal

gland-cells can be seen arranged in small clusters or nests; they

are situated in a more or less regular annular series, two or more

to each ring. The specimens described by Diesing evidently were

not mature, as appears from the double hooks. Moreover, many

of them were found not in the lungs, but in other organs of their

hosts; they were therefore probably still migrating through the

tissues to the lungs.

The hosts in which Natterer found the specimens described

by Diesing are not many, but they belong to different orders ;

Diesing, in fact, gives three snakes: Ooluber lichtensteini, C. flavt-

ventris, Boa constrictor, and a lizard, Amphisbena flavescens.

As I have already observed in other papers dealing with this

group of parasites, the genus Raillietiella, differing from all the

others of the family Linguatulide, comprehends species of para-

sites coming from hosts of very different families: from Ophidia,

Lacertide, and Amphibia.

According to Diesing 2. furcocerca is to be found both in several

snakes and in a lizard of very peculiar habits, the Amphisbena.

Although until now there is no direct evidence to the contrary,

1 think it improbable that the same species should inhabit such

different hosts. For this reason I have thought it well to give

a more complete description of this parasite, so that comparison

may be made with specimens obtained from snakes.

Although several species of Raillietiella have been described,

both in the adult and semi-adult forms, we have as yet no know-

ledge of the younger developmental forms, nor of the intermediary

hosts in which they live. Hitherto the majority of intermediary

hosts for those members of the family, at least, which are found

in snakes, have been mammals; a large variety of mammalia

have often been found to act as intermediary hosts to the same

species of parasite. Armillifer amillatus, according to Noc, has

been found at its larval stage in 36 species of mammals. Other

species, parasitic in crocodiles, have in all probability fish as their

intermediary host. It is probable that all these immature forms,

Proc. Zoou. Soc.—1923, No. II. 2

18 THE LINGUATULID ARACHNID RAILLIETIELLA FURCOCERCA.

given under a variety of names, such as P. gracile, P. crocodili,

and others, generally found in fish of the families Gymnotide,

Characinide, and Siluride represent larval stages of the Sebekini.,

Moreover, the feeding-habits of their definite hosts render this

hypothesis extremely probable. The same is probably the case

with the genus Reighardia.

Many animals, in which tongue-worms of the genus Raillietiella

have been found, are purely insectivorous in their feeding-habits.

For example, the lizards of the genus Calotes and Gecko (R.

geckonis), Amphisbena alba (hk. furcocerca (Diesing)); and the

Amphibian, Bufo melanostictus (R. indica (Gedoelst, 1921)).

If even in the genus Faillietiella, as in all the others, there

exists a biological cycle which evolves in two different hosts, it

will be interesting to see in what intermediary host the larval

forms of these parasites of insectivorous animals pass their

existence, and how they manage to get into their definite host.

Could the intermediary host of Raillietiella be an invertebrate

animal ?

PERMIAN FISHES OF THE GENUS ACENTROPHORUS. 1g

4. The Permian Fishes of the Genus Acentrophorus*. By

HK. Leonard Ginx, M.S8e., The Royal Scottish Museum,

Hdinburgh.

[Received June 6, 1922: Read February 6, 1923. ]

(Text-figures 1-16.)

Introductory.

As the earliest Known members of the family Semionotide and

the only Paleozoic Actinopterygians of a higher grade than the

Chondrostei, the small fishes named Acentrophorus by Traquair

(1877), found in the Permian of the County of Durham, have fre-

quently been referred to and discussed in paleontological writings.

An examination of the abundant material in the museums at

Newcastle and Sunderland showed that it would be possible to

add considerably to what was hitherto known of the structure of

these fishes, and the main results of such an examination are

here given.

The first species of Acentrophorus to be named and described

was A. glaphyrus, which Agassiz published (1835) as a species of

Paleoniscus. It is found in the Marl Slate, the lowermost bed

of the Permian series in Durham and Northumberland, but as it

is rare and not well preserved it has furnished little information

as to the structure characteristic of the genus. A much more

abundant and better preserved fish is A. varians from the Upper

Magnesian Limestone. It was described as Palceoniscus varians

by J. W. Kirkby (1862 and 1864), who at the same time described

two much scarcer species occurring in the same beds, ‘‘ Pale-

oniscus” altus and ‘“ P.” abbsi. Reasons are here given for

doubting whether the species abbsi ever had a real existence.

In 1873 Liitken pointed out that these fishes were wrongly

assigned to Palwoniscus; and in 1877 Traquair established for

them the genus Acentrophorus.

Kirkby’: s aceourt (1864) includes a careful description of the

general proportions, the squamation, and the fins. Traquair

added many details, especially in regard to the osteology of the

head. Ina few minor points, however, the material at his dis-

posal evidently led him astray, and it does not appear to have

shown him anything of the teeth, the axial skeleton, or the bones

supporting the fins. Few figures of the fishes now included

under Acentrophorus have been published, and none give a very

clear representation of their structure. The specimens, from

both the Magnesian Limestone and the Marl Slate, seem to have

* Communicated by D. M.S. Watson, M.Sc., F.R.S., F.Z.S.

20 MR. E. LEONARD GILL ON THE PERMIAN

been fossilised under quiet conditions; the parts are rarely much

disturbed and practically never scattered. On the whole this isa

disadvantage. The scales and the few strong bones (frontal and

angular for example) are often well preserved, but most of the

bones are delicate, and as they are crushed one upon another it is

usually very difficult to make out their boundaries. In the case

of the palate it is practically impossible. The distal parts of the

fin-rays, too, are hardly ever shown satisfactorily. It is only

through having abundant material for comparison that it has

been possible to make out even so much of the skeleton as is here

described.

Deccription of the Genus.

The descriptions already given by Kirkby, Traquair (1877,

pp. 563-4), and Dr. A. Smith Woodward (1895, p. 51), together

with the restorations and other figures accompanying this paper,

render it unnecessary to go into much detail as to the main

external features characterising Acentrophorus. The general

shape (text-fig. 1), including the excavation of the back along

the base of the dorsal fin, is much as in Lepidotus, except that

the snout is blunter. To judge by the number and appearance

of the specimens that have been compressed dorso-ventrally, the

body was fairly rounded. The scales, compared with those of

most Semionotids, are thin; they overlap deeply, nearly half of

each scale being covered by the one in front of it. Their only

ornament is a series of concentric lines near the hinder margin ;

these are presumably annual growth lines, and as many as five

can sometimes be counted. The absence of a row of dorsal ridge-

scales with projecting points is the character upon which Traquair

separated the genus from Semionotus; in place of points these

scales often even show a concave hind border. But the most

striking characteristic of the squamation is the great though

perfectly graduated variation in the size of the scales on different

regions of the body, and particularly the great depth of the anterior

scales of the lateral line. In shape the lateral line scales are not

unlike those of Dapedius. The lateral line can usually hardly be

followed on the outer surface in A. varians and altus, except by

the notching of the scales. In A. glaphyrus it is more prominent ;

and on the inner face of the scales it is marked by a conspicuous

groove in all the species. Peg-and-socket articulation is well

developed on the inner face of the anterior scales of the upper part

of the flanks (text-fig. 2, A); but it is absent in 4. varians from

the smaller scales of the lower flank, and dies out gradually even

in the upper scales as they are traced backward towards the tail-

shaft (text-fig. 2, B,C, D).. In A. aléus and A. glaphyrus the

scales have on the inside a low rounded ridge, which is continued

above into the peg (text-fig. 2, E), and in these species the peg-

and-socket articulation extends almost or quite to the tail. In

all the species the scales on the jugular region are very small and

rounded (text-fig. 7, sc.). As in many other Semionotids, there

FISHES OF THE GENUS ACENTROPHORUS. 21

are three or four enlarged scales in the mid-ventral line im-

mediately in front of the anal fin. Enlarged ridge-scales mark

the mid-dorsal and mid-ventral line of the tail-shaft. As they

near the caudal fin they become pointed posteriorly, and pass

without a break into the upper and lower series of tail fulcra.

Restoration.

Text-figure 1.

About one-and-a-half times natural size.

Acentrophorus varians.

The massive development of the fin fulcra is a striking charac-

teristic of the genus. In the two later species, 4. varians and

A. altus, each fulerum fits closely to the point over the one next

behind it; in the Marl Slate species, 4. glaphyrus, the points are

considerably more drawn out and tend to project more freely.

22 MR. E. LEONARD: GILL ON 'THE PERMIAN

The smaller fulcra are paired, but the larger ones at the base of

each series are single, the transition taking place through the

deepening of a median groove. ‘The pectoral fin has only three

(possibly sometimes four) fulcra, the distal half of its upper

border being unprotected (text-fig. 9).

Of the fins themselves, two features are particularly note-

worthy: the situation of the pectoral fin and the degree of

neterocercy of the caudal. The pectoral fin is placed rather high

on the flank, and (at least in A. varians and A. alius) its lobe is

so formed that the fin can only be spread over or abreast of the

lower flank scales. It could not in life be spread below the level

of the body as in other fusiform Semionotids, with the pre-axial

border turned forwards and downwards. It is occasionally fos-

silised in this position, but it is then seen to be unnaturally forced

down so as to point forward over the jugular region or the bran-

chiostegal rays.

The dorsal body-lobe of the tail is in a particularly interesting

stage of reduction. It still extends to the extreme upper tip of

the tail, but for about the latter half of the distance it is repre-

sented by only a single line of small scales. ‘Towards the tip

these scales are so small that but for their slightly different

shape they closely resemble the joints of the minute first dermal

ray lying beside them. (The appearance suggests the possibility

that on the withdrawal of the notochord these little scales might

be directly converted into the joints of an additional dermal

ray.)

In other respects the fins greatly resemble those of Lepidotus.

As in that genus, a considerable length of the proximal part of

the rays is unjointed. In the best-preserved examples these

unjointed lengths and some of the joints immediately following

them show a flange-like lateral expansion, which led Kirkby to

state that they were covered by an accessory scale. The flanges

are especially conspicuous in the anal fin, where the rays are few

(only five or six behind the fulcra) and widely spaced.

It is impossible to make out completely the bony structure of

the head, since none of the specimens show the elements scat-

tered, and all the more delicate parts are crushed together into

a fragile, semi-transparent plate no thicker than fine notepaper.

Thus of the chondrocranium it can only be said that in the hinder

part it was considerably ossified, and that in one or two examples

a pair of exoccipitals are seen, and in another a club-shaped bone

in the position of the epiotic. A small bony rod occasionally

seen in front of the upper end of the pre-operculum may very

probably be the alisphenoid (text-fig. 7, asp.).

Other internal bones of the skull are often shown, but never

in such a way that they can be completely described. The front

of the parasphenoid is very commonly seen as a straight red

crossing the orbit (text-figs. 6 and 13); it has a lateral flange

beginning a short distance behind its anterior end, but its hinder

portion remains unknown. ‘The vomers are separate, in front

FISHES OF THE GENUS ACENTROPHORUS. 23

at least, but their connexion with the parasphenoid is not clearly

seen. Below the parasphenoid are the crushed remains of the

palatal bones, of which it can only be said that a thickened lower

border represents the palato-pterygoid bar, and that the shape

of the metapterygoid and quadrate can sometimes be roughly

made out. The metapterygoid has a central, upwardly directed

process (as in Lepidotus, Amia, etc.) for articulation with the otic

region of the brain-case. Traquair stated that “a well-marked

quadrate is distinct enough,” but if he was not mistaken he must

have seen some exceptional specimens. A symplectic with a rod-

like stiffening lies just inside the lower end of the preoperculum,

Text-figure 2.

Outlines of scales of Acentrophorus seen from the inside ; about twice natural size.

All the figures except F include scales of the lateral line. A-D. Acentrophorus

varians : A, anterior scales (//.6. sixth lateral line-scale) ; B, half-way between

occiput and dorsal fin; C, between dorsal and anal fins; D, last scales of

lateral line, with others above and below. EH. A. altus, scales above anal fin

(wa., remains of neural arches and spines). F. A. glaphyrus, scales below

lateral line between pelvic and anal fins,

its anterior end overlying the quadrate. Projecting from under

the upper end of the preoperculum some part of the hyomandi-

bular is often seen; it is well ossified, and in shape closely

resembles the hyomandibular of Lepidotus. All these bones, in

so far as they can be made out, are shown in text-fig. ll. Of

the hyoid apparatus, the anterior part of the ceratohyal and the

small hypohyal are frequently shown. The ceratohyal is deep,

especially at its fore end; the hypohyal is knobbed and rather

strongly ossified. In one small specimen of A. varians from the

Kirkby collection the operculars are somewhat displaced, and in

the gap thus left three delicate branchial arches are seen.

The membrane bones forming the exterior of the skull are

24 MR. HE. LEONARD GILL ON THE PERMIAN

much more satisfactorily preserved. Those of the cranial roof

are shown in outline in text-fig. 3. The pair of parietals together

roughly form a square. One well-preserved example of A. altus

shows two parietal ossifications on each side (text-fig. 4), but as

other specimens have only one, this is presumably no more than an

occasional aberration. The parietals are flanked by ‘“‘ squamosals”’

(pterotics, supratemporo-intertemporals) which have a rounded

expansion of the outer border anteriorly. The frontals are fused

into a single plate, the strongest bone in the whole skeleton with

the exception of the cleithrum; it is nearly always found either

turned outwards clear of the head or folded down beneath it,

rarely crushed together from side to side. It is wide behind and

much narrowed in front, the very large orbits producing deep

excavations of its lateral margins, Three of the circumorbitals

adjoin each of these excavations. The nasals are so delicate that

though their extent is seen, their exact shape cannot be made

Text-figure 3,

Outline restoration of skull-roof of Acentrophorus varians. About one-and-a-half

times natural size. c.or., circumorbitals; f., frontal ; m.tm., median temporal

plate; na., nasals; pa., parietals; p.mex., premaxille; p.tm., post-temporal ;

sq., squamosal; s.tm., supratemporal.

out, chiefly because they are crushed down on the comparatively

strong backward processes of the premaxille. In some specimens

there is a suggestion of a small median ‘“‘ethmoid” and of

separate lateral ossifications (either prefrontals or adnasals).

The edge of what may be a postfrontal sometimes appears under

the anterior border of the squamosal. A pair of very large scale-

like post-temporals adjoin the hinder margin of the parietals and

squamosals, the suture being overlaid by a transverse pair of

supratemporals (extrascapulars). A smaller median plate (text-

figs. 3 and 4, m.dm.) is wedged in between the post-temporals ;

it usually overlaps one of them, and is overlapped by the other.

It perhaps belongs to the supratemporal series.

The large orbit is surrounded by a chain of about ten circum-

orbitals, of which those forming its posterior boundary are much

narrower than the rest. Between the circumorbitals and the

preoperculum there were probably thin cheek-plates, but the ir

FISHES OF THE GENUS ACENTROPHORUS. 25

presence is suggested only by faint radiating grooves crossing

the bones of the palate ina few specimens. One or two examples

show a triangular bone which apparently covered the space in

front of the orbit between the nasal and the maxilla.

The opening of the mouth is very small. It is bounded above

by fairly strong: paired premaxille, each supported by a strong

process running backwards beneath the nasals to the frontal

(text-fig. 5, r.pma.). The maxille are short and delicate; their

slender anterior portion passes inward behind the premaxille ;

posteriorly they are expanded to overlie the lower jaw about the

middle of its length. The lower jaw is remarkable chiefly for

the fact that in this early genus it has already acquired in the

fullest degree the peculiar shape characteristic of the Mesozoic

Semionotids and Macrosemiids. As in these later fishes it is

very deep behind and shallow in front, with a downward curve

in the region of the symphysis. The agreement extends also to

Text-figure 4.

ptm mtm

Acentrophorus altus. Skull-roof of a specimen with four parietal plates. In the

Kirkby Collection, Hancock Museum. The plates behind the frontal are

slightly displaced to the right. About one-and-a-half times natural size.

Lettering as in text-fig. 3, with op., operculum.

the shape of the constituent bones, at any rate to those encasing

the outer side of the jaw—the angular, coronoid and dentary

(text-figs. 1, 5 and 6). An articular ossification was probably

present, but though it is sometimes suggested it is not clearly

shown. Many specimens show something of the inner side of

the jaw, but the splenial was apparently very thin, and its boun-

daries are lost. The most conspicuous feature exhibited on the

inner face of the jaw, as i+ is actually preserved, is a row of pro-

tuberances near the lower margin corresponding to the sensory

pits in the outer face of the dentary (text-figs. 5 and 13).

The only mention of teeth in Acentrophorus occurs in Dr. Smith

Woodward’s Catalogue (1895, pp. 51 and 52). Traquair states

that he found none, and Kirkby’s account makes no mention of

any. It is evident, however, that Kirkby had discovered them

subsequently, for the fine specimen represented in text-fig. 5 bears

26 MR. &. LEONARD GILL ON THE PERMIAN

a label in his writing with the word “teeth.” As a matter of

fact, a considerable proportion of all the specimens exhibit teeth

or some trace of teeth when they are examined with sutfticient

magnification and suitable lighting. Text-figs.5 and 6 taken

together show the dentition fairly completely. Long styliform

teeth are borne by the premaxilla, the narrow shaft of the

maxilla, and the anterior half of the dentary. The foremost

teeth on the dentary are especially long and curved, and it is

these that are most frequently found preserved. A single speci-

men suggests, though not conclusively, that there were minute

teeth on the splenial. The vomer is bordered by short, very

Text-figure 5.

COr

Acentrophorus varians. Head showing dentition; Kirkby Collection, Hancock

Museum. About four times natural size. an., angular; ¢.or., cireumorbital ;

dn., dentary ; hm., hyomandibular (part) ; z.op., interoperculum (space left by

it); Z.po., left preoperculum ; 7.pme., left premaxilla ; l.sy., left symplectic ;

mx., maxilla; op., operculum; 7.j., right ramus of lower jaw; 7,j.s.,

probably right splenial, displaced; r.pma., right premaxilla; +.po., right

preoperculum ; 7.sy., right symplectic; sa., coronoid (supra-angular); s.op.,

suboperculum ; v¢., vomerine teeth; x., possibly the right ceratohyal.

stout teeth ; they usually have the appearance of rounded knobs,

but the best examples show them to have had short conical

points. In one specimen that seems to show the buccal surface

of the vomers, there is a suggestion of inner rows of stud-like

tritoral teeth, but they are too obscure to warrant a positive

statement. A few specimens, notably the one from the Sunder-

land Museum represented in text-fig. 6, show that the palato-

pterygoid also bore small conical teeth.

The opercular apparatus is remarkable in the same way as the

jaws, namely in that it is so typically of. the Semionotid pattern.

Text-fig. 6 and the restorations make its eomposition clear

FISHES OF THE GENUS ACENTROPHORUS. 27

without further description. One or two remarks only are

necessary. The suture between the opercular and subopercular

plates varies considerably in position, but nearly always leaves

the suboperculum as the larger bone, except in A. glaphyrus.

The branchiostegal rays number nine or ten on each side (text-

fig. 7). (Traquair thought there were “few, about seven.”)

Traquair suspected the presence of a gular; if there was a gular,

however, it must have been a very delicate plate, for in the

numerous specimens that ought to show it there is no definite

trace of it. The only example which does show anything that

could well be taken for a gular is Kirkby’s type-specimen of

Text-figure 6.

Lop.

Acentrophorus altus. Head showing dentition; Sunderland Museum, c.hy.,

ceratohyal ; c.ov., cireumorbitals (three bordering frontal, three more between

parasphenoid and lower jaw); dn., dentary ; fr., frontal ; h.hy., hypohyal;

hm., hyomandibular; z.op., interoperculum ; ma., maxilla ; op., operculum ;

p.op., preoperculum ; p.pt., palato-pterygoid with teeth; p.sp., parasphenoid ;

s.op., suboperculum; v.é., vomerine teeth; w., left premaxilla, or possibly

the body of the vomer. About four times natural size.

A. altus, the head-of which is represented in text-fig. 13. The

plate marked in this figure is a thin, detached bone, of much

the size and shape that the gular would have to possess.

The opercular bones and the roofing bones of the head are

ornamented with fine vermicular raised lines. The course of the

rincipal sensory canals of the head can be traced fairly com-

pletely. The nasal bones are too thin and badly preserved to

show pits or grooves, but the frontal often shows a line of four

pits on each side near the orbital margin, and grooves and pits

earry the line backwards along the outer side of the squamosal

to cross the outer end of the supratemporal and the antero-

28 MR. E, LEONARD GILL ON THE PERMIAN

external angle of the post-temporal. From thence it crosses the

supraclavicle obliquely to the first scale of the lateral line series.

The under surface of the frontal is marked by a conspicuous

ridge on either side parallel to the curved orbital margin. It

underlies the line of sensory pits and may be connected with their

canal, or it may only be a boundary wall for the chondrocranium

Text-figure 7.

S.Op. ee S.cl

Lait Weatssivocts (Sn eam ad MS aie

IID 1. CH. tas ua NE BE teen a

pal Fel.

Acentrophorus varians. Head crushed obliquely from the left side and from below,

showing branchiostegal rays and parts of shoulder-girdle ; Kirkby Collection,

Hancock Museum. About four times natural size. asp., alisphenoid ; c.hy.,

right ceratohyal; c.or., circumorbitals; fi., frontal, left side from below;

h.hy., right hypohyal ; 7.op., interoperculum ; 1.07, left branchiostegals; 7.cZ.,

part of left cleithrum; 7,7., left lower jaw; ma., maxilla; op., operculum ;

pal., bones of palate;’ p.ma., premaxilla; p.op., preoperculum; 7., ribs;

7.br., right branchiostegals; 7.c/., point of right cleithrum; 7.j., right

lower jaw; se., small scales of jugular region ; s.c/., supraclavicle, displaced ;

s.op., Suboperculum.

in this region. As in Amia, one of the pits on the squamosal

forms a notch in the margin of the bone at the point where the

main canal was joined by the branch traversing the preoperculum.

The supratemporals are strongly grooved and pitted by the

transverse branch of the canal. The situation of all these pits

FISHES OF THE GENUS ACENTROPHORUS. 29

and grooves is indicated in text-fig. 3. On the side of the head

the sensory canals can be followed even more clearly. Three or

four pits are usually conspicuous in the length of the pre-

operculum, and from the lower end of this bone the line is con-

tinued by a groove across the angular, and by a chain of six or

seven well-marked pits along the dentary near its lower border

(text-figs. 5 and 6). Finally, there are longitudinal grooves on

the inferior circumorbitals, probably marking the course of a

suborbital canal.

The pectoral girdle is rarely seen at all completely, being

largely covered as a rule by the operculars and branchiostegal

rays. Its general shape, however, can often be seen through

these bones (text-figs. 7 and 14), and more or less of the thickened

portion of the cleithrum (‘clavicle ”), ornamented by fine trans-

verse raised lines, appears normally along the lower edge of the

Text-figure 8.

Outline restoration of shoulder-girdle of Acentrophorus varians. A., right side,

external view; B., left side, internal view. Double dotted line shows course

of lateral line. c/., cleithrum; p.tm., post-temporal; s.c.c., position of

scapulo-coracoid cartilage ; s.cl., supraclavicle. About three times natural

size.

suboperculum. Occasional specimens show some portion of the

girdle more fully, especially from the inner side, and make

possible such a restoration as is given in text-fig. 8. The large

seale-like post-temporal has already been referred to. The supra-

clavicle, which normally appears in part behind the operculum, is

of simple, slightly curved form, strengthened along its anterior

edge. The cleithrum is massively thickened externally in a

boomerang form, pointed at either end, and with a band of fine

transverse raised lines marking the portion which sometimes

appears at the surface below the suboperculum. Its inner face

is expanded and flattened, especially about the middle of its

length, but this face also bears a rod-like thickening, bent almost

30 MR. E. LEONARD GILL ON THE PERMIAN

at a right angle just above the area of attachment of the scapulo-

coracoid cartilage. This latter cartilage has left a distinct trace

of its form in two or three of the specimens (text-fig. 9). Many

specimens show the pointed lower ends of the ‘two cleithra

meeting (or slightly displaced from their normal contact) in the

midventral line, and prove the entire absence of an infra-

clavicle.

The skeleton of the pectoral fin of A. varians is shown in text-

fig. 9. The radials are about eight in number. They appear as

a single series, though slight cups shown in their outer ends may

have lodged minute distal cartilages. The first fin fulcrum,

which is large, seems always to be based directly on the scapular

cartilage in series with the radials. There is no sign of the

Text-figure 9.

A. Skeleton of left pectoral fin of Acentrophorus varians. Dotted lines show the

approximate shape of the scapulo-coracoid cartilage and of three pustaxial

radials which have only been seen as impressions. From a specimen in the

Kirkby Collection and one in the Sunderland Museum.

B. Part of preaxial edge of pectoral fin of Acentrophorus glaphyrus, showing the

long, slender fulcra. From a specimen in the Kirkby Collection, Hancock

Museum. Both figures about four times natural size.

presence of a metapterygium. The dermal rays are about twice

as numerous as the radials, and where it happens that two

adjacent rays are supported by one radial, the bases of these rays

are noticeably inclined together. ‘The two first (pre-axial) rays

are unbranched, and the three last (post-axial) are both un-

branched and unjointed. The remaining rays are all forked

twice in the distal third of their length.

The pelvic fins are very short-based, and are remarkable exter-

nally for the great length attained by one of the fulera (text-

fig. 10, B). Specimens in which the inner surface of the scales

is exposed not infrequently show the supporting bone of the

FISHES OF THE GENUS ACENTROPHORUS. 31

pelvic fin. The best example of this kind, showing both supports

together, is represented in text-fig. 10; the bones are seen to be

of the same general pattern as in Lepidotus, but of a stouter

form. Nothing is seen of any radials intervening between the

supports and the dermal rays, though the parts are so small and

difficult to see that no definite statement on the point can safely

be made.

Previous investigators do not appear to have noticed any traces

of the axial skeleton of the trunk, except that Kirkby refers to

the supports of the median fins. The distal ends of these

supports fairly often appear, where scales are missing along

the bases of the fins. But much more than this can be made out

in certain specimens, particularly where, as occasionally happens,

Text-figure 10.

Haney

a, v fi)

ey /)

BS ; iy

CL os

A. Pair of pelvic fin-supports of Acentrophorus varians, lying on the inner face of

disturbed scales.

B. Part of pelvic fin of Acentrophorus varians, showing basal fulcra. Both figures

from specimens in the Kirkby Collection, Hancock Museum ; about four times

natural size.

the splitting of the matrix has cleanly separated a right and a

left half, leaving the inner surface of the scales completely

exposed. (Such specimens sometimes show by a difference of

colour the exact outline of the abdominal cavity, which is pre-

cisely as ina herring or similar fish.) Jf a specimen such as this

is examined with a suitable lens and in a strong, very oblique

light, any break in the regularity of the scale pattern being care-

fully scrutinised, it will almost always be found to show more or

less extensive traces of the axial skeleton. Kirkby’s collection in

the Hancock Museum at Newcastle contains three specimens of

this nature in which the skeleton is particularly well shown, and

the information they give is set out in text-fig. 11, where the

unbroken lines represent the parts which are actually seen. The

Ay MR. E. LEONARD GILL ON THE PERMIAN

bones are represented in the fossils by little flattened, semi-

transparent pieces of a substance resembling sugar-candy. They

were probably not very strongly calcified, and the flattening may

hm., hyomandibular; mpt., metaptery-

Text-figure 11.

About one-and-a-half times natural size.

goid; qu., quadrate; sy., symplectic.

Axial skeleton and other internal bones of Acentrophorus varians, from specimens in the Kirkby Collection,

Hancock Museum.

have expanded their shape to some extent, but a comparison of

different parts does not suggest that much allowance need be

made for alteration of this kind.

FISHES OF THE GENUS ACENTROPHORUS. 33

The number of vertebral segments seems to agree closely with

the number of rows of scales—ahout forty in each case from the

head to the middle of the base of the caudal fin—but the longi-

tudinal spacing does not correspond by any means. In the caudal

region the segments are so widely spaced as to suggest some sort

of “foreshadowing of the alternate pleuro- and hypo- -centra found

in the same region in such fishes, for example, as Amia. But

there is no sign of actual centra; the notochord evidently per-

sisted complete. The interpretation of what is seen of the

vertebral column is not easy. In the segments close behind

the head it rather strongly suggests the vertebral column of the

sturgeon. It appears as though in the anterior segments the

neural spines may have been separate, but further back they

were almost certainly fused with the arches. The spines below

the dorsal fin-supports are markedly widened from front to back,

a peculiar feature, which is confirmed, however, by all the speci-

mens in which these parts are shown. In the anterior caudal

region the spines seem to be wide and short; further back they

are narrower again and are lengthened to support the dorsal

ridge-scales. Their continuation into the tail-lobe is not shown

in any of the specimens. Well ossified ribs appear in many

specimens in the anterior abdominal region (cf. text-fig. 7),

extending more than half-way from the level of the lateral

line to the ventral border; but in the posterior abdominal

region they are very short. They are succeeded by four or five

fairly long hemal spines, inclined strongly backwards across the

upper ends of the anal fin-supports. In the posterior part of

the tail-shaft the hemal spines, like the corresponding neural

spines, are lengthened to support the ventral ridge-scales, and

the succeeding spines serve to support the caudal fin-rays in the

usual way. The supports of the dorsal and anal fins are well

shown. ‘There is one to each fin-ray. At the front of the dorsal

fin there is also a separate support for each of the fulera that stand

in series with the rays, but the corresponding fulcra of the anal

fin, which arz more crowded, have only one support for them all.

The distal ends of all the supports are notched or forked ; in the

case of the dorsal fin the two heads so formed appear to share in

the support of two adjacent rays. The internal ends are more or

less expanded, and seem to come into close contact with the

neural or hzemal spines, except perhaps in the case of those from

the back of the fin. The great proportionate size of the anal fin-

supports is a peculiar feature of the internal skeleton of

Acentrophorus.

The Species of Acentrophorus.

There are known at present three indisputable species of

Acentrophorus: A. varians and A. altus (Kirkby sps.) from the

Upper Magnesian Limestone, and A. glaphyrus (Agassiz) from the

Marl Slate. A fourth species, d. abbsi (Kirkby), papeats to be

Proc. Zoon. Soc.—1923, No. IIT. 3°

34 MR. E, LEONARD GILL ON THE PERMIAN

ill-founded. Text-figs. 1, 12, 13, 14, 15 illustrate the differences

distinguishing these species, and will be sufficiently supplemented

by the following brief notes :—

Acentrophorus varians (Kirkby). (Text-fig. 1.) The type-

Species and very much the most abundant. In spite of its name,

not more variable than many modern fishes, e. g., trout, roach.

Specimens range in length from 5-11 cm.

Acentrophorus altus (Kirkby). (Text-fig. 12.) Very much

searcer than A. varians but occurring in the same beds. Differs

only in its deeper form and, as already stated, in the character of

the inner surface and articulation of the scales (text-fig. 2).

Length 4-8 cm.

Acentrophorus abbsi (Kirkby). Also from the Upper Magnesian

Limestone of Co. Durham. Supposed to be distinguished from

the other species by its extraordinarily lengthened form. Close

examination of the two or three examples in Kirkby’s collection

seems to show that they are no more than distorted specimens of

Text-figure 12.

) CEE

‘ aX XX HS NG GEE

AIS

4 \ AW wy =

LOE Sh SAS

Orme Woy UWA

ea eA HDGaRR CER Bamps

A) pe . \\

z ay 4 rs

Bae : te

SO SSSI

Acentrophorus altus. Restoration.

About natural size.

A. varians. Text-fig. 14 shows the head of the type-specimen.

Tt will be seen that the parts are dragged out toa great degree

longitudinally. The opercular apparatus has held together, but

is lying far behind its prover position in relation to the squamosal

and lower jaw, leaving the hyomandibular region widely exposed.

Similarly the squamosal, supratemporal, and post-temporal are

dragged apart, and the pectoral fin pulled from its connexion

witn the shoulder-girdle. A similar distortion is apparent in the

whole length of the body, the scales hardly or not at all over-

lapping and in places separated altogether, Kirkby’s figure

(1864, pl. 18. fig. 3) shows the posterior half of the body

separately ; in the actual specimen the two halves are parted by

a very shallow horizontal fault or slip-line which has carried them

some distance asunder, with a trail of scales as their only

connexion. The eondition is difficult to account for, but it is

evident that movement took place in the surface layer of lime-

stone before it was quite consolidated but after the fish had

FISHES OF THE GENUS ACENTROPHORUS. 35

become completely macerated. In the Marl Slate occasional

examples of Paleoniscus Jreteslebent may be found in precisely

the same drawn-out condition. Unless other specimens exist in

which the attenuated form can be shown to be that of the living

fish the species 4. abbsi should be dropped.

Acentrophorus glaphyrus (Agassiz). (Text-fig. 15.) Found in

the Marl Slate at the base of the Magnesian Limestone, and

Text-figure 13.

be gape © \

mx. chy. a

Acentrophorus altus. Head of Kirkby’s type-specimen (Quart. Journ. Geol. Soc.

vol. xx. 1864, pl. 18. fig.1). About four times natural size. c.hy., ceratohyal ;

el., cleithrum ; f1., frontal (left side broken away) ; h.Ay., hypohyal; J,7., lett

lower jaw ; J.pm., left premaxilla (part) ; mx., maxilla; pa., parietals; p.ti.,

post-temporal ; 7,7., right lower jaw, inner surface; r.pm., right premaxilla ;

sq., squamosal ; s.ém., supratemporal; x., possibly a gular plate.

therefore the earliest species. More variable in form than

A. varians ; some examples, the type-specimen among them, deep

in the body, others shallower than the form represented in text-

fig. 15. The chief diagnostic character hitherto attributed to the

species is a pronounced fine denticulation of the free hinder

border of the scales. This denticulation is, however, so very

MR. E. LEONARD GILL ON THE PERMIAN

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seum (the Yorkshire

Philosophical Society’s Museum at York); here the finely to

CUS9qQM snioydo..zUaoy ,,

which shows

the Yorkshire Mu

The only example known to me

-specimen in

rarely to be detected that it is of little practical use in identifi-

edge is still preserved on three or four scales which are seen

cation.

the type

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Most specimens of

less completely.

A. glaphyrus show no clear trace of it at all.

Such thin scales

FISHES OF THE GENUS ACENTROPHORUS. 37

as those of this fish are not well preserved in the Marl Slate; on

the exposed side of a specimen they are nearly always variously

splintered at the hinder border, and it is usually impossible to find

a perfect scale. A single example in the Hancock Museum

retains undamaged scales, and in this example, as in some in the

British Museum (Smith Woodward, 1895, p. 54), the scales have

the hind margin entire or at the most slightly frilled. It 1s

therefore evident either that the denticulation is an inconstant

character or that more than one species is included under the

name glaphyrus. Against the latter possibility is the fact that

the two examples showing the clearest denticulation are respec-

tively among the deepest and the shallowest in body-form. In

any case the denticulated scales do not form a reliable or useful

specific distinction. Nevertheless, 4. glaphyrusis distinguishable

from A. varians by several well-marked characters. The caudal

fin in A. glaphyrus is considerably longer and more deeply forked

(much like that of Palceoniscus in fact, whereas in the later

species the tail is shaped as in Lepidotus); the fulera on all the

Text-figure 15.

Acentrophorus glaphyrus. Restoration.

About natural size.

fins are much longer and slenderer, with freely-projecting needle-

like points; the pectoral fin is set a little lower on the body and

its basal lobe is so shaped as to permit of its being turned down-

wards with the preaxial border in front; in the skull the frontal

is wider anteriorly than in A. varians, and the maxilla is shorter

and stouter; the lateral line is prominent externally; the inner

face of the scales is ridged and the articulating pegs are narrow

and finely pointed—the inner face of the scales, in fact, resembling

that of A. aliws, and still more closely that of Palwoniscus.

A. note seems called for regarding the puzzling second figure of

A. glaphyrus in King’s monograph (1850, pl. 22. fig. 4). The

original specimen is in the Hancock Museum. Behind the

pectoral region it is particularly well preserved, but the head

(which appears very large and “‘ Palzoniscid”” in the figure) is in

reality broken up, part of the area being occupied by a good

impression of the skull-roof.

A fish found in the Trias of Chicopee Falls, Mass., was

described by Newberry (1888, p. 69, pl. 19. figs. 3, 4) under the

38 MR, E. LEONARD GILL ON THE PERMIAN

name of Acentrophorus chicopensis. The caudal lobe, which his

figures show to be as short as in Semionotus, is enough, however,

to prove that the fish is not an Acentrophorus. In addition, the

scales are too uniform in size, and the general outline does not

suggest Acentrophorus either in head or body. It should also be

said that Newberry’s discussion of the genus and of its relation-

ship to “ Ischypterus”’ (pp. 67-69) is on the whole misleading,

since it is founded only on the obscure figures accompanying

Kirkby’s paper of 1864.

Another fish assigned to the genus is Acentrophorus dispersus

Fritsch (1894, p. 81, pls. 113, 114) from the Lower Permian

(Schwartenkohle) of Kounova, Bohemia. Its remains are very

imperfect, but Fritsch’s figures show enough of the upper and

lower Jaws, the opercular apparatus and the pectoral girdle to

prove that in all these essential elements of its structure the fish

was fundamentally different from any Semionotid. Indeed, at the

end of his description of it, Fritsch himself appears to have

concluded that it was a Paleoniscid.

SUMMATYs

The propriety of placing Acentrophorus in the family Semiono-

tide was not in doubt, but it is amply confirmed by the additional

information now brought forward. In the exact correspondence

ot the dorsal and anal fin-rays with their endoskeletal supports,

in the reduced maxilla, the narrow preoperculum and the whole

plan of the opercular apparatus, in the absence of an infra-clavicle

as in all the details of the pectoral arch and the pelvic bones,

Acentrophorus is a characteristic representative of the Proto-

spondyli and of the family Semionotide.

At the same time, Acentrophorus does in certain respects retain

primitive characters. It seems, in fact, to provide us with some-

thing that is comparatively rarely found among fossil fishes,

namely a really early representative of a new group which still

shows definite traces of its origin from older types. The most

striking character of this kind in Acentrophorus is undoubtedly

the upper caudal lobe. As compared with the caudal lobe of the

contemporary Paleoniscids it is certainly greatly reduced, but the

reduction is only in depth; the caudal lobe of Acentrophorus is

still of the maximum length. It is interesting to find in another

contemporary fish, the extraordinary form Dorypterus hoffmanni,

a caucal lobe in precisely the same stage of reduction, except

that, in place of the single line of scales found in Acentrophorus,

the lobe is marked externally to the very tip by two lines, the

scales as they are traced backwards becoming linear and almost

microscopic. That a stage such audal lobe

extremely attenuated in its hinder portion but still more or less

of the full length—may have heen passed through pretty regularly

in the production of a ‘‘ hemi-heterocercal” tail is suggested not

only by the parallelism between Acenirophorus and Derypterus, but

FISHES OF THE GENUS ACENTROPHORUS. 39

also by at least one fact of fossil ontogeny. A young example of

Dapedius, 9 em. long and beautifully preserved (text-fig. 16),

which has been kindly lent me by Prof. D. M. 8. Watson, has a

caudal lobe extending just two-thirds the length of the upper

border of the tail, instead of only one-third as in the adult ; and

for some distance towards its apex the lobe is marked by a single

line of very small and narrow scales.

The head of Acentrophorus also, though in side view almost

entirely typical of a Semionotid, shows when seen from above

(text-fig. 3) some decided primitive traits. The shape of the

Text-figure 16.

i ah

Dapedius orbis. Tail of a young example, total length 9 cm., showing extended

caudal lobe. Lower Lias; locality uncertain. D., M.S. Watson Collection.

About four times natural size.

frontal shield and of the supratemporals, and the large leaf-

shaped post-temporals, are all strongly reminiscent of Palconiscus.

So also is the internal pattern and articulation of the scales,

especially in the earliest species, 4. glaphyrus.

On the other hand, there are certain points in the structure of

Acentrophorus which seem to be peculiar to this genus among the

Semionotide. The orbit is very large and encircled by only a

single chain of bones; the. scales, compared with those of other

Semionotids, are thin, deeply overlapping and of unusual range

of difference in size, and do not produce the typical “ tessellated

40 PERMIAN FISHES OF THE GENUS ACENTROPHORUS.

pavement” effect; the fin-fulcra in the later species are of par-

ticularly massive form ; the pectoral fin is placed higher on the

flank than in other fusiform Semionotids, and its attachment

gives it a peculiarly ‘‘ modern” appearance. Fused frontals are

also unusual in the family; and of the characters of the axial

skeleton which are shown clearly enough to be worth discussion,

the expanded neural spines under the dorsal fin and the great

size of the supports of the anal fin-rays seem to be quite peculiar

to Acentrophorus.

In view of all these primitive and individual characters it can

no longer be said that Acentrophorus differs from Semionotus only

in lacking the median row of acuminate scales in advance of the

dorsal fin. It is seen to be very much the most primitive known

member of the family Semionotide, and is therefore difficult to

incorporate in any scheme on which the genera may be classified.

On the whole, however, it would appear to be nearest to the line

of ancestry of Lepidotus. Such a relationship is suggested in

many points, in particular in the shape of most of the fins and in

other details of their structure, in the excavation of the back

along the base of the dorsal fin, and in the tendency towards a

tritoral form exhibited by the teeth on the vomers.

My thanks are due to Prof. D. M. 8. Watson, who first sug-

gested the desirability of re-examining the material of Acentro-

phorus in the Hancock Museum at Neweastle; to Mr. J. A.

Charlton Deas and Mr. T. R. Goddard for giving me facilities

for studying some fine specimens in the Sunderland Museum ;

and to Dr. W. E. Collinge for similar privileges at York.

Literature cited.

Agassiz, L. 1835.—Poissons Fossiles, vol. i.

Fritscu, A. 1894.—Fauna der Gaskohle etc. Bohmens, vol. iii.

Kine, W. 1850.—Permian Fossils (Pal. Soc.).

Kirxsy, J. W. 1862.—Ann. & Mag. Nat. Hist. ser. 3, vol. ix.

p. 267.

Kirksey, J. W. 1864.—Quart. Journ. Geol. Soc. vol. xx. p. 353.

(Reprinted, 1865, in Nat. Hist. Trans. Northld. & Dhm.

vol. 1.)

Newsurry, J.S. 1888.—Fossil Fishes, Trias, New Jersey and

Connecticut (Monographs U.S. Geol. Surv., vol. xiv.).

Traquair, R. H. 1877.—Quart. Journ. Geol. Soc. vol. xxxiii.

. 562.

Woopwarb, A. Smite. 1895.—Cat. Fossil Fishes Brit. Mus.,

part 3.

ARTICULATION OF PALATOQUADRATE IN C@LACANTHIDS. A]

5. The Postorbital Articulation of the Palatoquadrate with

the Neurocranium in the Coelacanthid Fishes. By

Epwarp Puetps ALLIs, Jr., F.Z.S.

[Received October 25, 1922: Read February 6, 1923.)

In arecent work (Allis, 1922), I came to the conclusion that

the postorbital articulation of the palatoquadrate with the neuro-

cranium in Wimania and Aelia, two of the Coelacanthid, was

homologous with that in the recent Hexanchus and Heptanchus,

and not with that in Lepidosteus. This conclusion was based

largely on my assumption that the vena jugularis of these fishes

ran posteriorly between the articular process of the palatoquadrate

and the cranial wall, instead of accompanying the ramus

ophthalmicus lateralis, as suggested by Stensid (1921), and

traversing, with that nerve, a foramen that lay between the

alisphenoid and basisphenoid bones of the latter author's

descriptions. Further consideration of the subject, in connection

with a later work by Stensid (1922), has led me to conclude that

the vein did not have the course ascribed to it by either Stensi0

or myself, but, accompanying the ramus ophthalmicus profundus,

traversed a foramen that lay anterior, instead of posterior, to the

alisphenoid, and so entered a trigemino-facialis chamber in the

cranial wall. ‘This conclusion modifies radically my conception

of the conditions in these fossil fishes, but I, nevertheless, still

think that the articulation of the metapterygoid with the cranium

was of the type found in the recent Hexanchus and Heptanchus,

and not of that in Lepidosteus.

There are, in recent fishes, two processes of the palatoquadrate

that are here concerned, as well as two processes of the neuro-

cranium. One of the processes of the palatoquadrate is the

processus basalis, or pedicle, and the other the processus

metapterygoideus, and both of them are found in typical condition

in the adult Amia. The processus basalis of this fish forms the

dorso-anterior corner of the metapterygoid, and lies in the plane

of the peey of the palatoquadrate. It is directed toward the

lateral edge of the basis cranii, in the region of the orbital

opening of ‘the myodome, and is there connected with it by a band

of connective tissue. The floor of the orbital opening of the

myodome is the homologue of the processus basipterygoideus of

the Sauria (Allis, 1913), and it is lined ventrally, and supported,

by the anterior portion of the ascending process of the

parasphenoid. The processus metapterygoideus projects dorso-

mesially from the external surface of the metapterygoid, is

directed toward the projecting dorso-lateral corner of the

postorbital process of the neurocranium, and connected with that

process by a membrane that covers the external surface of the

42 MR, E, P. ALLIS ON POSTORBITAL ARYICULATION OF

musculus levator arcus palatini (Allis, 1897), It lies between

the musculi levator arcus palatini and adductor mandibule,

giving insertion, on its internal surface, to certain fibres of the

former muscle and origin, on its external surface, to certain

fibres of the adductor. It forms the lateral boundary of the

surface of insertion of the levator on the palatoquadrate, the

mesial boundary of that surface being formed by the processus

basalis together with that portion of the dorso-mesial edge of the

metapterygoid that lies between that process and the processus

metapterygoideus. That branch of the nervus trigeminus that

innervates the levator arcus palatini turns posteriorly around the

anterior edge of that muscle, and runs posteriorly on its external

surface, thus lying, morphologically, between the processus

basalis and the processus metapterygoideus, dorsal to the one and

ventro-mesial to the other. That branch of the external carotid

that becomes the secondary afferent pseudobranchial artery

(Allis, 1912) has similar relations to these two processes, as has

also the vena jugularis. The efferent pseudobranchial artery

runs upward and forward ventro-mesial to the processus meta-

pterygoideus, but its relations to the processus basalis are not

positively evident. It lies external to the ascending process of

the parasphenoid, and hence would seem to lie morphologically

ventral to the processus basipterygoideus.

In embryos of this fish up to 19°5 mm. in length, the two pro-

cesses here under consideration are shown by Pehrson (1922,

figs. 3-12) apparently both lying on the dorso-mesial edge of the

palatoquadrate, the metapterygoid process lying posterior to the

basal process and extending farther dorsally in all stages

excepting the very earliest. In a 56 mm. specimen, the meta-

pterygoid process has apparently begun to assume its adult

position, and the metapterygoid bone has begun to ossify, and it

is to be noted that Pehrson shows (J.c. fig. 14) the unossified

portion of the pterygoquadrate cartilage forming a V-shaped

mass, the basalis and metapterygoideus processes forming the two

arms of the V, and the metapterygoid bone lying between them,

in the hollow of the V, and not extending upward into either

process. The otic process of this fish, and that of the Teleostei

as well, is represented in the lateral wall of the trigemino-facialis

chamber (Allis, 1914).

The characteristic features of these two processes of this fish

thus are: that one of them lies postero-lateral to the other; that

the antero-mesial one is directed toward the lateral edge of the

floor of the cranium, where the palatoquadrate articulates with it

in Lepidosteus; that the postero-lateral one is directed toward

the lateral edge of the roof of the cranium, where the palato-

quadrate articulates with it in Hexanchus and Heptanchus; that

the musculus levator arcus palatini has its insertion on the

palatoquadrate between the two processes; that the musculus

adductor mandibule lies external to the processus metaptery-

goideus and is cut off, by it, from any origin or insertion on the

PALATOQUADRATE WITH NEUROCRANIUM IN CQ@ALACANTHIDS. 43

processus basalis; and that the nerve that innervates the levator,

the secondary afferent pseudobranchial artery, and the vena

jugularis all le actually, or morphologically, between the two

processes.

In Scomber (Allis, 1903) and the mail-cheeked fishes (Allis,

1909) these two processes (or flanges developed in relation to

them) have acquired contact with the hyomandibula and become

firmly attached to it, the processus basalis with its internal

surface and the processus metapterygoideus with its external

surface. But, notwithstanding this marked difference in the

direction of the two processes, and also in their relations to

the cranium and the hyomandibula, the levator areus palatini,

the nerve that innervates it, and an artery that corresponds to

the secondary afferent pseudobranchial artery of Amia, still all

lie actually, or morphologically (Cottus), between them, and the

adductor mandibule has its origin on the external surface of the

processus metapterygoideus and not at all on the processus

basalis. The vena jugularis, also, still hes morphologically

between the two processes, for the dorso-mesial edge of the processus

basalis is connected, by tissue, with the lateral edge of the basis

eranil.

In Amiurus the conditions appear, at first sight, markedly

different from those in the fishes above referred to, but they are,

nevertheless, quite certainly strictly homologous. In a 32 mm.

specimen of this fish, Kindred (1919) shows the pterygoquadrate

and hyomandibular ‘cartil ages completely fused with each other

and forming a V-shaped mass with the point of the V directed

ventrally. The hyomandibula has begun to ossify on the posterior

limb of the V, the metapterygoid on the anterior limb, and the

quadrate on the ventrally directed point. Nothing whatever is

shown in the hollow of the V, but it must certainly be spanned

by membrane, for it is in exactly this place that the metaptery-

goid bone first appears in Amia, as above stated, and in the adult

Amiurus the corresponding place is occupied by an anteriorly

projecting flange of the hyomandibula which sutures anteriorly

with the hind edge of the metapterygoid and ventrally with the

dorsal edge of the quadrate. Along the dorsal edge of this

flange, and of corresponding length, there is a horizontal ridge

on the hyomandibula, and the levator arcus palatini is inserted in

art on the anterior end of the ridge and in port along the full

length of its dorsal surface (McMurrich, 1884). Certain of the

deeper fibres of the musculus adductor mandibule have their

origins on the Jatero-ventral surface of this ridge, the superficial

portion of the muscle extending upward external to the ridge.

The anteriorly projecting flange of the hyomandibula of this fish

thus corresponds both topographically and functicnally to the

posteriorly projecting flange on the hind edge of the metaptery-

goid process of the fishes Just above considered, and is quite

certainly the homologue of that flange; and that it has become

a flange on the anterior edge of the hyomandibula, instead of on

44 MR. HE, P. ALLIS ON POSTORBITAL ARTICULATION OF

the hind edge of the metapterygoid, is no more extraordinary

than the incorporation, in this fish, of the dorsal portion of

the preoperculum in the hind edge of the hyomandibula (Kindred,

1919). The metapterygoid, the projecting dorso-mesial portion

of which forms the processus basalis, is said to be attached to

the lateral surface of the cranium by a sheet of muscular and

ligamentous tissue. It is not said at what point on the lateral

wall of the cranium this sheet of tissue has its attachment, but

two sectional views given through the postorbital region would

seem to show that the attachment is either to the trabecula or

to the lateral edge of the parasphenoid.

In all of the above-mentioned fishes the palatoquadrate has no

direct articular contact with the neurocranium. this articulation

being acquired through the intermediation of the hyomandibula,

to which the palatoquadrate is firmly fixed. In Lepidosteus,

Osteoglossum, Hexanchus, and Heptanchus this articulation with

the cranium through the intermediation of a hyomandibula still

oceurs, but in each of them there is, in addition, direct articular

contact with the postorbital portion of the cranium; and these

are the only recent fishes I know of in which the latter articu-

lation is found, for the palatobasal articulation, found in many

of the Selachii, is orbital in position and not postorbital. Gaupp

says (1905, p. 767) that an articulation of the palatoquadrate

with a processus basipterygoideus is found in many fishes, but, as

he does not mention the particular ones, I have not been able to

control the statement. Luther (1913) describes the articulation

in Lepidosteus, and then says that it is also found in Osteoglossum

but in no other teleost. It apparently occurs in certain fossil

fishes, as in Boreosomus, one of the Paleoniscide described by

Stensid (1921).

In the adult Lepidostews, Parker (1882) describes two processes

related to the metapterygoid, one of which he calls the pedicle

and the other the otic process. The pedicle is the larger of the

two, and is shown forming the dorso-postero-mesial corner of the

metapterygoid, the so-called otic process projecting ventro-

posteriorly from the ventral edge of the base of the pedicle. The

hind end of the pedicle articulates with the anterior edge of a

processus basipterygoideus which projects antero-laterally from

the lateral wall of the cranium near its ventral edge, the so-called

otic process projecting posteriorly ventral to the processus

basipterygoideus. Ina 205mm. embryo of this fish, these two

processes of the metapterygoid form (Veit, 1911, fig. 17, pl. C.)

the dorsal and ventral corners of the actual hind edge of the

palatoquadrate, but as this edge of the palatoquadrate of this fish

corresponds to the metapterygoid portion of the dorso-mesial

edge of the palatoquadrate of Améa, the two processes lie,

morphologically, one posterior to the other on the posterior

portion of the dorso-mesial edge of the palatoquadrate. They

accordingly agree exactly, in this, with the basalis and metaptery-

goideus processes of Ama, and I shall hereafter so refer to them.

PALATOQUADRATE WITH NBUROCRANIUM IN C@LACANTHIDS. 45

The musculus levator arcus palatini of the adult Eepidasicus is

said by Luther (1913) to have its origin on the sphenotic, and to

run from there antero-mesially to “have its insertion on the

metapterygoid bone and adjacent portions of the palatoquadrate

cartilage. Luther says (/.c. p. 13) that: ‘ Dabei umgreift der

Muskel das Metapterygoid vor dem Basipterygoid- Gelenk und

reicht auch an der Medialseite bis zu etwa 2 der Breite des

Knochens hinauf.” This would seem to mean that tbe muscle

had its insertion in considerable part on the internal surface of

the processus basalis, which would be so unusual for a levator

muscle that I have looked it up in transverse sections of an

80 mm. specimen of this fish. In this specimen the muscle, as it

approaches the hind end of the palatoquadrate is wrapped around

(‘“umegreift”) the lateral edge of the processus basipterygoideus,

and when it reaches the hind end of the processus basalis, that

part of the muscle that lies ventral to the processus basiptery-

goideus is inserted on the lateral surface of the processus basalis,

this surface of the latter process being presented ventro-laterally.

The processus metapterygoideus lies against the ventral surface

of this part of the muscle, partly imbedded in it, and apparently

gives insertion to some of its fibres, but the larger part of the

fibres of the muscle continue onward and are inserted on the

external surface of the palatoquadrate anterior to the bases of the

two processes, none of them being inserted on its internal surface.

That part of the adductor muscle that Luther calls the musculus

adductor mandibulz postorbitalis ies ventral, and hence external,

to the processus metapterygoideus, and certain of its fibres have

their origins on that process. The action of the levator muscle

is said by Luther (J.c. p. 61) to be to pull the palatoquadrate

almost directly laterally, the motion at the basipterygoid joint

accordingly being a sliding one, in a latero-mesial direction, and

this is in accord with the position of the long axis of this joint,

which is practically at right angles to that of the long axis of

the hyomandibulo-cranial articular joint. The nerve that inner-

vates the levator muscle penetrates it from its external surface,

and hence lies morphologically between the metapterygoid and

basal processes, as it does in the fishes above considered.

The processus basipterygoideus of Lepidosteus is said by Parker

(1882, pp. 453 & 461) to be developed in part from the trabecula

and in part from the parachordal. Veit first (1907) concluded

that it 1s developed from the hind end of the trabecula and

corresponds to a part of the floor of the orbit of the Selachii, but

he later (1911) says that it arises in connection with the trabecula

and the floor of the trigemino-facialis chamber. I first concluded

(Allis, 1909) that it is of trabecular origin and, later (1913), that

it is the homologue of the floor of the orbital opening of the

myodome of Amia; but it is to be particularly noted that there

is, in Lepidosteus, no parasphenoidal leg of the alisphenoid related

to the process, as there is in Amia, and that there is no membrane

that I can recognize replacing it. Veit says that the process

4.6 MR. E. P. ALLIS ON POSTORBITAL ARTICULATION OF

develops late in Lepidostews, and he considers its articulation with

the metapterygoid to be a secondary acquisition, and Luther

holds the same opinion, The ventral surface of the process, its

anterior edge, and the adjacent portions of its dorsal surface are

covered and supported by a process of the parasphenoid which

would seem to correspond to an anterior portion of the ascending

process of the parasphenoid of Ama, and it is apparently this

process of the bone that alone gives articulation to the meta-

pterygoid in Osteoglossum, as explained below. The process of

Lepidosteus lies ventral to the ramus ophthalmicus profundus and

the vena jugularis, and if it be the homologue of the floor of the

orbital opening of the myodome of Amia, as I concluded, it would

seem as if it must he, morphologically, between the profundus

and trigeminus nerves, for its homologue in Amia supports the

parasphenoidal leg of the alisphenoid, which actually lies between

those two nerves. In its relations to these nerves this process

thus differs radically from the postorbital process, which always

lies, so far as I know, posterior to all branches of the nervus

trigeminus, and usually between the foramen for that nerve and

the secondary, or definitive, foramen for the nervus facialis. The

processus metapterygoideus also lies morphologically between the

trigeminus and facialis nerves, while the processus basalis, which

articulates with the processus basipterygoideus, should have,

morphologically, the same relations to those nerves that the

processus basipterygoideus has.

In Osteoglossum a laterally projecting process of the para-

sphenoid articulates with a semicylindrical groove on the dorsal

edge of an antero-dorsally projecting portion of the metapterygoid

(Bridge, 1895), which, as it has exactly the position of the pro-

cessus basalis of Ama, is evidently that precess, and when the

palatoquadrate swings outward and upward, under the action of

the levator arcus palatini, the groove on the metapterygoid has a

sliding motion on the process. Posterior to this process, there is

a short dorsally projecting process of the metapterygoid which

lies against the external surface of the hyomandibula, and hence

evidently represents the processus metapterygoideus. The

laterally projecting process of the parasphenoid has its origin

from the base of a pretrigeminus portion of the ascending

process of that bone and corresponds to that part of the ascending

process of Lepidosteus that underles and supports the basiptery-

goid process of that fish, and hence is, functionally, a processus

basipterygoideus. It cor responds also to that anterior portion of

the ascending process of the parasphenoid of Amia that underlies

the floor of the orbital opening of the myodome of the latter fish,

and the pretrigeminus portion of the ascending process of the

parasphenoid of Osteoglossum corresponds to one of the two legs

of the alisphenoid bone of Ama, but which one is uncertain, for

the relations of the nerves and the vena jugularis to it are not

given.

The conditions in Heptanchus and Hexanchus I have not

PALATOQUADRATE WITH NEUROCRANIUM IN C@ILACANTHIDS, 47

personally examined, but they must be similar to those in

Chlamydoselachus. In the latter fish the most dorsal portion of

the dorso-mesial edge of the palatoquadrate forms the processus

metapterygoideus, a low ridge on the internal surface of this edge

of the apparatus forming the processus basalis (Allis, 1913). In

the groove between these two ridge-like processes, the musculus

levator maxillee superioris, which is the homologue of the levator

arcus palatini of the Teleostei and Ganoidei, has its insertion, the

outer edge of the processus metapterygoideus lying between the

surface of insertion of the levator muscle and the surface of origin

of the adductor mandibule. That branch of the nervus tri-

geminus that innervates the muscle runs posteriorly along its

external surface, and hence lies morphologically between the two

ridge-like processes. In Heptanchus and Hexanchus the con-

ditions unquestionably here differ only in that the processus

metapterygoideus has acquired articular contact with the laterally

projecting dorsal portion of the postorbital process, and that, in

consequence, the anterior portion of the levator maxillee superioris

runs antero-ventrally beneath the overhanging portion of the

postorbital process, and hence internal to the processus meta-

pterygoideus. The nerve that innervates this muscle also runs

posteriorly internal to the latter process. The levator muscle of

these three fishes, and particularly that of Heptanchus and

Hexanchus, can accordingly have but little effect in swinging the

palatoquadrate outward and upward, and Luther (1909) says that

this motion is impressed upon it by the musculus preorbitalis of

his descriptions, which is Vetter’s (1874) muscle Add 8, This

action of this muscle is said to also throw the anterior end of the

palatoquadrate downward, the palatobasal (orbital) process sliding

downward along its articular contact with the orbital wall. The

levator muscle then pulls the palatoquadrate back into place, its

action thus being largely, if not entirely, to pull the palatoquad-

rate dorso-mesially in the plane of the apparatus. The articular

attachment of the processus metapterygoideus to the postorbital

process, in Heptanchus and Hexanchus, would evidently have to

be somewhat loose to permit of this motion, and Luther (1909)

assumes that there is here a certain dorso-ventral sliding motion,

and, also, that the levator muscle must act in part as an adductor

palatoquadrati. Furthermore, the attachment of the palato-

quadrate to the hyomandibula must be such as to allow a certain

amount of motion between them.

In certain others of the Plagiostomi this metapterygoid ridge

of the Notidanide has become a pronounced process, and both

Gegenbaur (1872) and Luther (1909) have called it a muscle

process. It apparently hes directly upon the dorso-mesial edge

of the palatoquadrate, thus having in the adults of these fishes

the position that it has in larve of Amia. Where present it has

the same relations to the levator and adductor muscles that the

ridge-like process of the Notidanide has, and because of its

articular relations to the cranial wall, in the latter fishes, it has

48 MR. E. P. ALLIS ON POSTORBITAL ARTICULATION OF

been quite generally considered to be an otic process; but if, as L

have endeavoured to show, it is the homologue of the processus

mnetapterygoideus of the Teleostomi, this cannot be. Goodrich

considers it to be an otic process and says (1909, p. 98) that it

‘‘appears to have been established very early, since there is

reason to believe that it existed not only in the Jurassic Cestra-

ciontide (Hybodus), but also in the Cladoselachii, Acanthodu,

and Pleurocanthodii.”

In certain of the higher vertebrates a pterygo-cranial articu-

lation is found between processes which are apparently the strict

homologues either of the processus basalis and basipterygoideus

of Lepidostews, or of investing bones related to those processes,

the motion between the two processes, however, being a dorso-

ventral swinging one instead of a latero-mesial sliding one.

In recent Amphibia a cartilaginous processus basipterygoideus

seems not to be found (Gaupp, 1900, p. 537), but it is found im

certain of the Stegocephali, as explained immediately below. The

palatoquadrate of the Amphibia is said by Gaupp to present

four typical processes: a processus oticus, processus ascendens,

processus pterygoideus, and processus palatobasalis, the latter

process being also called by him (1905, p. 736) simply the pro-

cessus basalis. In the Urodela, the latter process is said to lie, at

first, in contact with the ventral surface of the auditory capsule,

at the point where the capsule is connected with the basal plate

of the skull, and to later there fuse with it, The process is said

to seem to correspond to the palatobasal process of the Selachu,

but this seems improbable, for the processus basalis of the

Amphibia arises from the pterygoquadrate portion of the palato-

quadrate, is directed mesially, and lies against the ventral surface

of the floor of the auditory capsule, while the processus palato-

basalis of the Selachii arises from the palatine portion of the

alatoquadrate, is directed dorsally, and lies against the orbital

wall, which forms part of the lateral surface of the cranium.

In Rana, of the Anura, the processus basalis is said hy Gaupp

(1893, p. 349) to first appear in his fourth stage (young frog

about 2cm. long), there arising from the pars metapterygoidea

of the quadrate, close above the root of the processus pterygoideus.

It is directed toward the ventral surface of the anterior portion

of the auditory capsule, and an articular joint is later there

formed, this being considered by Gaupp (1905, p. 737) to be a

more primitive condition than the fusion of the process with the

auditory capsule in the Urodela. In earlier stages of Rana there

is a processus muscularis rising from the upper edge of the

palatoquadrate, strongly developed in the earliest stage described,

but gradually diminishing in importance until its complete

disappearance during the metamorphosis. This process lies, in

larve (Gaupp, 1893, p. 292), external to the musculi temporalis

and pterygoideus, and gives origin, on its mesial surface, to the

masseter, and, on its lateral surface, to the depressor cartilaginis

hyoides and the depressor mandibule. The relations of the

PALATOQUADRATE WITH NEUROCRANIUM IN CGELACANTHIDS. 49

process to the adductor muscles of the mandible are thus not the

same as in fishes, but this is probably not important, for the

superior and inferior branches of the nervus trigeminus both run

forward dorsal or mesial to the process and then turn outward

anterior to it. The process lies between the pars articularis

quadrati and the processus basalis, in the same relation to those

two processes that the processus metapterygoideus of fishes has,

and hence is quite certainly the homologue of the latter process.

In the Apoda there is said to be no processus basalis (Gaupp,

1905, p. 753).

In the Stegocephah, the palatoquadrate is said to be movable

in certain forms, but fixed in by far the larger number. In

Hryops there is, according to Von Huene (1912), a basipterygoid

process which is apparently a definite process of the basisphenoid

bone, but as there is no suture separating the basisphenoid and

parasphenoid, the latter bone may also enter into the process.

The pterygoid is said to be attached (befestigt sich) to the process

by a broad articular surface, the palatoquadrate thus apparently

being movable. In Archegosawrus, the parasphenoid has, according

to Watson (1919), a narrow, outstanding basipterygoid process

on either side, and the pterygoid articulates with it by a freely

movable joint; and similar basipterygoid processes are apparently

found in several others ef the Rachitomi (/.c. p. 53). In

Laccocephalus the mesial process of the pterygoid suturates with

the lateral edge of the parasphenoid in a region that corresponds

to that from which the ascending process of the bone of fishes

has its origin, and cartilaginous extensions of the basisphenoid

are said (J.c. p. 54) to “‘seem to have passed outward above the

flat parasphenoid expansions to the epipterygoid and pterygoid.”

At the hind end of the suture between the pterygoid and

parasphenoid there is a foramen which leads into a canal in the

parasphenoid and transmits the internal carotid artery. In

Capitosaurus the mesial process of the pterygoid suturates both

with the parasphenoid and the exoccipital, and there is no basi-

pterygoid process. The quadrate ramus of the pterygoid is said

(Watson, 1919, p. 27) to form “a thin plate, rising nearly

vertically, to have a long and close connection with the supra-

temporal and squamosal. Its upper inner corner has a sutural

union with the prootic, with which bone and the supratemporal

it forms a large foramen leading forwards over the prootic and

epipterygoid to the anterior part of the skull. This opening

must transmit the vena capitis dorsalis and a lymphatic duct.

Just above its articulation with the parasphenoid and below the

prootic the inner margin of the quadrate ramus of the pterygoid

is notched for the passage of the vena capitis lateralis and the

seventh nerve.” This part of the pterygoid of this amphibian

thus has to this part of the skull the topographical relations that

the ascending process of the parasphenoid of fishes has, or that

the piscine processus metapterygoideus would have if it were

to persist and acquire contact with the cranium after the

Proc. Zoon. Soc.—1923, No. LV. 4

°50 MR. E. P, ALLIS ON POSTORBITAL ARTICULATION OF

disappearance of a musculus levator arcus palatini. The processus

oticus 1s part of the epipterygoid bone and lies anterior and

internal to this process of the pterygoid.

In the Sauria the processus basalis of fishes must certainly be

represented in the meniscus pterygoideus, but whether the

processus basalis of recent Amphibia is represented in the meniscus

alone, or in that structure together with the processus basiptery-

goideus, would seem to be an open question. Regarding the

development of these structures in the Sauria, Gaupp says (1905,

p. 762) that they are both represented, in young embryos of

Lacerta, by a mass of dense tissue which lies between the foot of

the processus ascendens and the root of the trabecula, and is

more closely connected with the former than with the latter.

When chondrification takes place, this mass separates into two

parts, the mesial one chondrifying as a laterally directed process

of the trabecula (the processus basipterygoideus) and the lateral

one becoming the meniscus. ‘This manner of development of

these two structures in this reptile, and their relations to the

ascending process of the palatoquadrate, led me to suggest, in an

earlier work, that the mass of tissue referred to might represent

the pharyngeal element of the mandibular arch.

In the adult Lacerta, the meniscus lies on the mesial aspect of

the Os pterygoideum (Gaupp, 1905, p. 767), and between it and

the processus basipterygoideus an articular joint is formed, the

palatoquadrate being movable. In Sphenodon, the meniscus and

the processus basipterygoideus are also both found (Howes and

Swinnerton, 1901), and there is an articular joint between them,

as in Lacerta, but whether the palatoquadrate of this reptile is

movable, or not, I do not find definitely stated. A mesial process

of the Os pterygoideum is shown by Fuchs (1901), in an embryo

of this reptile, projecting mesially along the ventral surface of

the processus basipterygoideus, and it would seem as if it must

interfere with any movement, if it does not actually prevent it.

Broom (1922) says that Parker described a meniscus, in 1878, in

a fairly advanced specimen of Zootoca, and he himself describes it

in a larval Agama hispida, where it lies, as in Lacerta, between a

cartilaginous processus basipterygoideus and a dermal pterygoid

bone. The articular joint is between the meniscus and the

processus hasipterygoideus, and Broom considers the meniscus to

be the homologue of the mesopterygoid of fishes, basing this

conclusion on the conditions found in Husthenopteron, a fossil

Rhipidistid described by Bryant (1919), The so-called meso-

pterygoid of the latter fish, as shown in the figure reproduced by

Broom, is, however, called by Bryant a metapterygoid, and that

part of the bone that has the marked upward extension shown in

the latter author’s text-fig. 6 would seem to be a processus

metapterygoideus metapterygoidei.

From the above references to the conditions in certain of the

Amphibia and Reptilia it is seen that, in these animals, the

existing postorbital articulation of the palatoquadrate with the

PALATOQUADRATE WITH NEUROCRANIUM IN C@ALACANTHIDS. 51

cranium was primarily, in all probability, that of a processus

basalis metapterygoidei with the lateral edge of the basis cranti

at or near the hind end of the trabecula. The processus basalis

has, in certain of them, later there fused with the basis cranii,

while still retaining its connection with the palatoquadrate. In

others it has apparently lost its connection with the palatoquadrate

but retained that with the neurocranium and then separated into

two parts, thus giving rise to a meniscus metapterygoideus and

a processus basipterygoideus. The metapterygoid cartilage is

always here lined ventrally, and supported by a dermal pterygoid

bone, the processus basipterygoideus being similarly jined and

supported by a lateral process of the dermal parasphenoid, and in

those animals in which the metapterygoid and _ basipterygoid

cartilages are resorbed, or fail to be developed, the two dermal

bones replace them and acquire sutural connection with each

other.

The conditions in the three Ccelacanthide so fully and so well

described by Stensié (1921, 1922) may now be considered, and

the question is: Is the postorbital articulation of the palato-

quadrate of these fishes with the cranium the homologue of that

in Heptanchus and Hexanchus, or of that in Lepidosteus,

Osteoglossum, and higher vertebrates ?

The metapterygoid of these fishes is a substituent bone, as it is

in most recent fishes, and at either end of its dorsal edge there is

a pointed process. Between these two processes the dorsal edge

of the bone is concave, and articulates with a process of the

neurocranium that Stensid considers to be a processus basi-

pterygoideus and which he refers to as the process e. In both

Wimania and Aelia this latter process is a process of what

Stensio considers to be a median basisphenoid bone. It is

directed dorso-antero-laterally, and in Wimania extends so far

dorsally that its dorso-anterior corner is shown in contact with

the dermal bones of the roof of the skull. In Avelia it is shorter,

but still extends above the middle line of the lateral surface of

the cranium. In Diplocercides it is still shorter, and is a process

of a median sphenoid bone and not of a basisphenoid. The

metapterygoid would seem, from the figures given, to have

articulated either with the lateral surface of “the process é

immediately beneath its laterally projecting dorsal edge, or with

that edge itself, and as the dorsal edge of the metapterygoid is

longer than the process @ is wide, the dorso-anterior end of the

metapterygoid would seem to have projected anteriorly somewhat

beyond the process e.

The metapterygoid has an even outer surface, without process

of any kind, and there is no posteriorly directed process, or other

feature, either on this bone or at any place along the hind edge

of the palatoquadrate that would indicate a rigid attachment

to the hyomandibula. This latter attachment was there-

fore probably by ligament only, and in this, as also in the

general configuration of this part of the palatoquadrate, there is

52 MR. E. P. ALLIS ON POSTORBITAL ARTICULATION OF

marked resemblance to the conditions in the Selachii. Stensio

says (1921, p. 73) that there was a large mandibula in Wimania

(and hence probably in the other two fishes also), and that this

presupposes a powerful adductor muscle. Comparison with

recent fishes would then indicate that the surface of origin of

this muscle on the palatoquadrate must have extended upward

at least to the dorsal edge of the metapterygoid, and Stensié

considers it probable that the muscle extended beyond that edge

and had its origin in part on the lateral surface of the neuro-

cranium, The levator arcus palatini, if present, would then

necessarily have had its insertion, as in Heptanchus and Hexan-

chus, either along the dorsal edge of the metapterygoid or partly

on that edge and partly on its mesial surface.

On the internal surface of the palatoquadrate there is a large

dermal pterygoid bone, and a process of this bone extends to, or

nearly to, the point of the dorso-anterior process of the meta-

pterygoid, thus supporting that process. The dorso-posterior

process of the metapterygoid, on the contrary, extends consider-

ably beyond the dorsal edge of the pterygoid, this leaving a

considerable portion of the internal surface of the metapterygoid

exposed beyond the pterygoid. On the lateral surface of the

pterygoid there is a strongly pronounced and rounded ridge

which extends upward to the point of the dorso-anterior process

of the metapterygoid, and strongly suggests that that point

represents the dorsal end of some element of the mandibular

arch.

The relations of the two processes on the dorsal edge of the

metapterygoid to each other, to the palatoquadrate, to the

pterygoid, and to the muscles of the arch, thus all strongly

suggest that the posterior one is a processus metapterygoideus,

and the anterior one either the entire processus basalis er the

anterior end of a ridge-like process similar to that in the Selachii,

the posterior portion of the ridge lying along the internal surface

of the palatoquadrate. There was quite probably a spiracular

canal in these fishes, as there is in the recent Polypterus, and

not simply a diverticulum of that canal, such as is found in the

recent Holostei and Teleostei; and, where this canal is present,

there is, in recent fishes, no articulation of the processus basalis

with the neurocranium.

There is thus strong presumptive evidence that, in these fishes,

it is a processus metapterygoideus that articulates with the

cranium, and, in recent fishes, this process never articulates with,

or even approaches, a processus basipterygoideus. This is

evidently in favour of the assumption that the process e is a pro-

cessus postorbitalis, and the relations to it of the cranial nerves is

also in favour of this interpretation.

There is, in Wimania and Awelia, a median bone which, as

already stated, Stensiéd considers to be a basisphenoid, and it is

said by him to consist of a body (corpus), and three processes on

either side, one of these processes being dorsal, one anterior, and

PALATOQUADRATE WITH NEUROCRANIUM IN C@LACANTHIDS. 93

one ventral. The body of the bone is said to be triangular in

shape, with the point belew, and from there to extend dorso-

posteriorly to its thickened base, which is said to le strikingly

high, leaving only a narrow space between it and the roof of the

cranium. ‘I'he bone, as shown in the figures given, has a shape

and position which show that it must have occupied, and have

completely filled, the hollow of a large cephalic flexure of the

brain, both its anterior and its posterior surfaces sloping antero-

ventrally. The dorso-posterior edge of the bone is concave, with

a short process on either side, and looks like the dorsum sell of

the skulls of certain vertebrates turned upward and backward.

The dorsal process of either side projects dorso-antero-laterally

from the corpus, and, in Wimania, is, as already stated, so tall

that its dorso-anterior corner may be in contact with the postero-

lateral corner of the dermosphenotic portion of the fronto-

dermosphenotic. The dorsal end of the process thus lies at or

near the level of the roof of the cranial cavity, and it is apparently

with the ventral surface of the outer end of the process that the

metapterygoid articulates. The anterior process of the bone is

lamellar in form with its basal portion extending upward along

the internal surface of the dorsal process, and it gives support on

the dorso-anterior edge of its basal portion to the alisphenoid of

Stensio’s descriptions. Dorsal to these two processes the lateral

wall of the cranium must have been of cartilage, and this

cartilage and the dorsal process must together have formed a

laterally projecting dorsal portion of the posterior wall of the

orbital fossa, and hence have formed a postorbital process in the

sense in which that term is here employed. Stensid says there is

no postorbital process in these fishes, but it is possible that he

employs the term in a somewhat different sense. The basi-

sphenoid is traversed, on either side, by but a single canal, which

traverses the anterior process and will be later considered.

In Diplocercides the anterior portion of the basisphenoid of

Wimania and Awelia is replaced by a sphenoid bone, which

extends forward through the orbit to the ethmoidal region and

is everywhere in contact, dorsally, with the ventral surface of

the dermal bone, or bones, that here form the roof of the

cranium. The process e of Wimania and Aselia is, in this fish,

apparently simply a protuberance on the lateral surface of the

hind edge of the sphenoid, at about the middle of its height.

From it a ridge runs upward along the hind edge of the lateral

surface of the dorsal half of the sphenoid, and, as it is considered

to be the homologue of the alisphenoid bone of Wimania and

Axelia, it is called the alisphenoid wulst. The postclinoid

wall of this fish does not project dorsally above the level of the

floor of the posterior portion of the cranial cavity, as it does in

Wimania and Axelia.

The truncus maxillo-mandibularis trigemini is considered by

Stensid to have issued from the cranium, in each of the three

fishes here under consideration, posterior to the alisphenoid, and

54 MR. H. P, ALLIS ON POSTORBITAL ARTICULATION OF

in Wimania posterior also to the dorsal process of the basi-

sphenoid. ‘Che ramus ophthalmicus profundus of Avelia is said

to run forward internal to the base of the alisphenoid, and to

issue from the cranium anterior to that bone, and in Diplocercides

it has similar relations to the alisphenoid wulst. In Wimania

it is said to probably have issued through that small canal,

above referred to, that traverses the anterior process of the

basisphenoid, but, as will later appear, it is much more probable

that this canal was traversed by the efferent pseudobranchial

artery. The ramus ophthalmicus lateralis trigemini is said to

run forward, in Wimania, over the dorsal process of the basi-

sphenoid, close to the lateral wall of the brain-case, thus

evidently issuing from the cranium between the dorsal process

and the alisphenoid, and then running forward lateral to the

latter bone. In Awelia this nerve lies ina groove that crosses

the external surface of the alisphenoid, and in D¢plocercides

in a canal that traverses the alisphenoid wulst and is continued

onward, a certain distance beyond that wulst, in the body of the

sphenoid bone. The vena jugularis is said to have had, in

Wimania, a course similar to that of the ramus ophthalmicus

lateralis trigemini. In Awelia and Diplocercides its course 1s

not given.

The profundus and trigeminus nerves of Aelia thus have, in

the courses ascribed to them by Stensid, exactly the relations to

the alisphenoid bone of that fish that the corresponding nerves

have, in recent fishes, to the parasphenoidal leg (pedicle) of the

alisphenoid, the trigeminus nerves all issuing posterior to that

leg and the profundus nerve anterior to it; and as the vena

jugularis of recent fishes always accompanies the profundus

nerve in this part of its course, it seems exceedingly probable

that it accompanied that nerve also in Awelia, and hence passed,

as in recent tishes, antero-mesial to the alisphenoid. In Diplo-

cercides the conditions differ from those in Awelia only in that

the ramus ophthalmicus lateralis trigemini traverses a canal in

the alisphenoid part of the sphenoid bone, instead of- passing

lateral to it; but even in recent fishes, as in Ama, this nerve

may traverse a foramen that perforates the parasphenoidal leg of

the alisphenoid.

In Wimania the conditions, as described by Stensi6, differ from

those in Awelia and Diplocercides in that the ramus ophthalmicus

profundus is said to have probably traversed the canal in the

anterior process of the basisphenoid, that the truncus maxillo-

mandibularis trigemini presumably ran outward in a notch at

the base of the hind edge of the dorsal process of that bone, and

that the vena jugularis accompanied the ramus ophthalmicus

lateralis trigemini, and hence passed postero-lateral instead of

antero-mesial, to the alisphenoid bone. ‘This course for the latter

vein would be most exceptional in fishes, and it seems much

more probable that both it and the ramus ophthalmicus profundus

passed internal to the alisphenoid, as they both apparently did in

PALATOQUADRATE WITH NEUROCRANIUM IN CQELACANTHIDS. 55

Axelia and Diplocercides. ‘The vena jugularis would then con-

tinue posteriorly and, having joined the nervus facialis, issue

with it through the facialis foramen at the hind edge of the

basisphenoid. In this part of its course it must have lain in a

canal in the cranial wall, for that it perforated the dura mater and

entered the central cranial cavity seems wholly improbable. ‘This

canal would thus, in this, exactly resemble the trigemino-facialis

chamber of recent fishes, and it seems wholly impossible that the

truncus maxillo-mandibularis trigemini did not run forward in

it, and issue from it between the process ¢ and the alisphenoid.

If the truncus issued posterior to the process e, as suggested by

Stensio, it would be separated from its ophthalmicus lateralis

branch by the entire width of that process, which would be

wholly exceptional, as compared with recent fishes, whether the

process be a postorbital one or a basipterygoideus. The notch

at the base of the hind edge of the process e would then probably

transmit the ramus palatinus facialis, which is not otherwise

accounted for.

If the trigeminus and profundus nerves of Wimania and Axelia

had the courses that [ have above ascribed to them, they must have

first run upward along the posterior surface of the basisphenoid,

then have passed, on either side, lateral to that posteriorly project-

ing dorsal end of the basisphenoid that resembles a dorsum selle,

and then have run forward along the dorsal surtace of the body of

the basisphenoid, there either lying free in the cranial cavity or,

much more probably, being enclosed in the canal traversed by

the vena jugularis. In recent fishes these nerves run forward

across the ventral end of the hollow of the cephalic flexure, instead

of upward and forward over it. ‘That part of the basisphenoid

bone of Wimania and Awelia that oceupies the hollow of this

flexure could not then have arisen from the conditions found in

recent fishes by the simple chondritication or ossification of tissues

that filled the hollow of the flexure, for the nerves would then

have been enclosed, i st#w, in the cartilage or bone so formed

and have traversed canals or foramina init. And it seems equally

improbable that the bone could have grown upward in the hollow

of the flexure, pushing the nerves before it. It must then be

that, at a certain. stage in the development of embryos of

Wimania and Aelia, the anterior end of the notochord was

bent upward and slightly backward, as it is in 33 mm. embryos

of the recent Acwnthias (Goodrich, 1918, fig. 14, pl. 2), and

that, in the former fishes, this curvature was not later reduced.

The cartilago acrochordalis, which, in recent fishes, develops in

tissues related to the anterior end of the notochord, was there-

fore, in Wimania and Aelia, directed dorso-posteriorly, and its

morphologically ventral surface, presented dorso-anteriorly, lay

directly beneath the membrane that, in recent fishes, extends

from the anterior edge of the cartilago acrochordalis to the top

of the preclinoid wall. The hind ends of the trabecule, and the

bases of the alisphenoid cartilages, would accordingly be carried

56 MR. E. P. ALLIS ON POSTORBITAL ARTICULATION OF

upward to the positions the corresponding bones of the adult

actually occupy, and that part of each anterior process of the

median basisphenoid on which the base of the related alisphenoid

rests would correspond to the outer end of the processus basiptery-

goideus of Amia and Lacerta, the remainder of the anterior

process apparently corresponding to that little lamellar process that,

in Amia, forms the lateral wall of the basal portion of the orbital

opening of the myodome and is traversed by the foramen for the

efferent pseudobranchial artery. ‘There would have been, internal

to the alisphenoid, a space traversed by the vena jugularis and

the ramus ophthalmicus profundus, the mesial boundary of this

space being formed by cartilage that represents the basisphenoidal

leg of the “alisphenoid. This leg of the alisphenoid would have

been continuous, posteriorly, with the mesial wall of a trigemino-

facialis chamber, that wall being either of cartilage or of

membrane, and the base of the leg of one side of the head would

have been connected with that of its fellow of the opposite side

by the preclinoid wall. The optic chiasma must have lain upon

the dorsal surface of the latter wall, and the lobi inferiores in a

depression posterior to it, on the dorsal surface of the membrane

(more or less ossified) that covers the dorso-anteriorly presented

ventral surface of the cartilago acrochordalis. A median opening

in this membrane would lead into a large pituitary sac, which

would extend ventrally into the space enclosed between the

ventral processes of the basisphenoid, these latter processes thus

corresponding to those antero-ventral processes of the parachordals

of recent fishes that bound laterally the pituitary fossa. The

basisphenoid of these fishes would thus apparently correspond to

a bone formed by the fused prooties of recent fishes.

There are thus several features in the cranial anatomy of these

fishes that indicate that the so-called dorsal process of the basi-

sphenoid corresponds to some part of the postorbital process of

recent fishes, and not to the basipter ygoid process, the most

important being that the process gives articulation to a processus

metapter yeoideus metapterygoidei and not to a processus basalis,

that 1t has no supporting relations to the alisphenoid, and that

the truncus maxillo-mandibularis trigemini quite certainly ran

outward anterior to it, between it and the alisphenoid. The

basisphenoid of these fishes is such a peculiar bone that all com-

parisons with recent fishes are more or less tentative, and there

is one further feature in their cranial anatomy that seems equally

peculiar. In all recent bony fishes (Crossopterygii, Holostei,

Teleostei) it is, so far as I know, an invariable rule that the dorsal

section of the hyomandibular latero- -sensory canal runs upward

toward, or actually to, a postotic portion of the main infraorbital

latero-sensory canal, while in the Plagiostomi this canal turns

forward and falls into the infraorbital portion of the main infra-

orbital canal at the ventro-posterior corner of the orbit. Now,

whether correlated to the course of this canal or not, it is a fact

that, in all the former fishes the hyomandibula articulates with

PALATOQUADRATE WITH NEUROCRANIUM IN C@LACANTHIDS. 57

the neurocranium dorsal to the vena jugularis, is either traversed

by the truncus hyomandibularis facialis or lies anterior to that

nerve (Polypterus), and is, if I am correct in my conclusions

(Allis, 1918), developed from the branchial rays of the hyal arch,

while, in the Plagiostomi, the hyomandibula articulates with the

neurocranium ventral to the vena jugularis, and is developed

either from the pharyngeal or the epal element of the hyal arch

(Allis, 1915). In Polyodon, where the hyomandibula articulates

with the neurocranium dorsal to the vena jugularis, and is of the

teleostome type, it nevertheless differs from that in the bony

fishes in that it les posterior to the nervus facialis, and in this

fish the hyomandibular canal has a position intermediate between

the two above referred to. There may be no morphological

significance in this, for there is in certain of the Plagiostomi a

line of surface pit-organs that has a position similar to that of the

preopercular canal of the bony fishes, and in certain of the latter

fishes there are surface lines of pit-organs that correspond in

position to that of the hyomandibular canal of the Plagiostomi;

but it is to be noted that in the Celacanthide, where a hyoman-

dibula, although presumably present, has not yet been found, the

hyomandibular canal has the position of that in the Flagiostomi,

while in the Paleoniscide, where the hyomandibula was well

developed and of the teleostome type, the canal has the position

of that in the bony fishes. In the Stegocephali, the canal on the

cheek would seem to be formed by the horizontal portion of

the canal of the Plagiostomi together with the dorsal portion

of the preopercular canal of the bony fishes.

LITERATURE CITED.

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Fe Ff —~ 1912. The Pseudobranchial and Carotid Arteries in Hsoa,

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1914. The Pituitary Fossa and Trigemino-facialis Chamber

in Ceratodus fosteri. Anat. Anz. Bd. xlvi.

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58 ARTICULATION OF PALATOQUADRATE IN CCELACANTHIDS.

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MICROFILARIH FOUND IN THE BLOOD OF BIRDS. 59

6. Some Microfilarie found in the Blood of Birds dying

in the Zoologicai Gardens, 1920-1921. By Riva

MARKBREITER, B.Sc., Research Assistant in the

Helminthological Department of the London School

of Tropical Medicine *.

(Text-figures 1-6.)

[Received November 6, 1922: Read February 6, 1923.]

From October 1920 till June 1921 the blood of 183 birds

which died in the Zoological Gardens of London was examined

and 22 were found to contain Microfilarie, roughly 12 per cent.

Of these 22 birds, one, a King Bird of Paradise (Cicinnurus

regius), contained two species of Microfilaria, and these two

species each occurred once again separately in other King Birds

of Paradise, so that in all 25 Microfilariz of 21 different species

were found.

‘Lhe birds were of 21 species belonging to 15 different families,

and came from the following parts of the world :—Mexico, Brazil,

S. and N. America, various parts of India, Chili, Java, Mada-

gascar, New Guinea, Malacca, and 8. and W. Africa, which

shows how wide the distribution of these parasites is. lt-is not

possible to name the embryo parasites unless the adults have

been simultaneously found; in these cases they were only found

three times, namely :—(1) In the Mexican Hangnest (Cassiculus

melanicterus), the adult being unidentified; (2) in the Occipital

Blue-Pie (Urocissa occipitalis); and (3) in the Wandering ‘lree-

Pie (Dendrocitia vayabunda): in both (2) and (3) the adult was a

Diplotrizna, and it is interesting to note that the corresponding

Microfilariz are of the same type though they differ in size.

Asin the Microfilariz of man, the avian Microfilariz fall into

two distinct types, sheathed and unsheathed, and in both groups

many different forms are found, long and short, thick and thin,

coiled and straight. Three of the Microtilariz listed were also

found by Plimmer, namely, the short form, occurring in Cicin-

nurus regius, and those in Urocissa occipitalis and Cittocincla

macrura; the others are reported for the first time. In the

detailed lists which are appended the adjectives short, medium

length, long and thin, medium thickness, and thick are repre-

sented by the following measurements:—Short = 0-100,;

thin = under 3 4; medium length = 100-200; medium thick-

ness = 3-4; long = over 200; thick = over 4p.

Reference.—Plimmer, ‘‘On the Blood Parasites found in Animals

in the Zoological Gardens,” Proc. Zool. Soc., 1912.

* Communicated by Prof. R. T. Lererr, M.D., D.Sc., F.Z.S.

60 MISS R. MARKBREITER ON SOME MICROFILARIA

Yext-figure 1.

Oe Lyeorece 2B, Lens &.

eS eS ES TIS aT nd}

(6) 05 Jimm

No, 773

NO. /022

ra ee EN

Text-figure 2.

No, 54

ee) \ Wo. 50

Lens Zeiss D. Eyepiece 2

No. 157 SS er ee ef eee (ees | eee eee | oe]

ea 0 05 mim.

ot No. 370 Wee

No. 249

No. 139

eee

No. OF

Outlines of Microfilaria to show comparative sizes and forms. Camera lucida

used. Leitz microscope: eyepiece 2B; lens + for drawings 773 (1920) to 1022

(1920) and 50 (1921) to 73 (1921).

Zeiss microscope: eyepiece 2; lens ZD used for drawings 92 (1921) to 370 (1921).

The numbers referring to Microfilarie in text-figs. 1 and 2 are those of the

Death Book of the Zoological Society for the years 1920 and 1921.

Trxt-Fie. 1. TEXT-FIG. 2.

773. Host. Cassiculus melanicterus. ~ 54. Host. Serinus scotops.

821. ,, Lrene turrosa. 50. ,, Quiscalus quiscalus.

848. ,, Caccabis chukar. 73. ,, Bubuleus coromandus.

889. ,, Cicinnurus regius. 92. 4, Otogyps calvus.

887. ,, Psittinus incertus. 139. ,, Hlanus ceruleus.

870. ,, Cicinnurus regius. 157. ,, Paradisea minor.

904. ,, Urocissa occipitalis. 182. ,, Turacus corythaix,

918. ,, Dendrocitta vagabunda. 249. ,, Steganura paradisea.

1022. ,, Agapornis cana. 303. ,, Cittocincla macrura.

51. ,, Hypochera nitens. 370. ,, Ramphocelus brasilius.

FOUND IN THE BLOOD OF BIRDS. 61

Text-figure 3.

Freehand drawings to show details.

Text-fig. 3. Microfilaria out of Irene turrosa (Fairy Blue-Bird).

Ba A, Microfilavia out of Caccabis chukar (Chukar Partridge).

st 5. Microfilaria out of Cicinnurus regius (King Bird of Paradise).

5 6. Microfilaria out of Cicinnurus regius (King Bird of Paradise).

Ex.P. = excretory pore. N.Ex.P. = nucleolus of excretory pore. Ex.M.C.

external muscle cells. Int.C. = internal cells, G,R. = genital region. N.R,

nerve ring. S, = sheath, ,

62 MISS R. MARKBREITER ON SOME MICROFILARIA

Admission

Fost. Habitat. to Helminth.

Gardens.

Iorprip 2. Mexico 9.6.20 | Microfilaria. Sheathed, short, medium

Cassiculus thickness, blunt tail, no cells at one

melanicterus | end of the body. Adult found by

(Mexican Hang- Dr. Vevers.

nest). Length with

SINGIN conepoue 86 p 84 86

Length —with-

out sheath 84 82 82

Breadth 4, 4 4

TURDID®. Java 13.10.20 | Microfilaria. Sheathed, long with

Trene turrosa sheath, body of raedium length;

(Fairy Blue-Bird). thick, graceful. Blunt tail; cells in

end of tail.

Length with

sheath 22.2... 2672 214 198

Length with-

out sheath 162 133 =119

TESHERKELN sce cconne 8 8 7

PHASIANIDA. N.W. India| 16.7.23 | Microfilaria. Sheathed, medium

Caccabis chukar | length, thick. Pointed tail; cells to

(Chukar Part- end of tail.

ridge). Length with

sheath 22) cen. 168 8 =176 93

Length = with-

out sheath 164 174 92

Breadth with

shheatilieeneeee U 8 6

Breadth with-

out sheath ... 5 6 4,

PARADISEID®. New Guinea | 13.10.20 | Microfilaria. Unsheathed, — short,

Cicinnurus medium thickness. Pointed tail,

regius nearly straight.

(King Bird of Length ......... 6lu 61 73

Paradise). breadtinwerereres: 4, 4, 4,

PsITTACIDS. Malacca 13, 10.20 | Microfilaria. Unsheathed, medium

Psitiinus length and thickness.

incertus Meng higeeenseses: 1304 170 148

(Blue-rumped Breadth ......... 4, 4, 4,

Parrakeet).

PARADISEID®. New Guinea | 13.10.20 | Microfilaria as in previous King Bird

Cicinnurus of Paradise.

regius enc hieeeseeeee 604 . 66 62

(King Bird of Breadiihiyeeeessee 4 4, 5

Paradise). Also Microfilaria, long and slender,

with pointed end. Unsheathed.

WEEN soe oso25e 218 220

| Bread this eee 15 2

CoRvID#. Himalayas | 16.7.20 | Microfilaria (a Diplotriena sp., adult

Urocissa found by Dr. Vevers). Unsheathed,

oceipitalis medium length, and thick. Pointed,

(Occipital Blue- tilted tail.

Pie). enc thwaener 143 158 150

Breadth eee 4-5 4-5 4-5

Convip2. India 4.6.20 | Microfilaria (a Diplotriena sp., adults

Dendrocitta found by Dr. Vevers). Unsheathed,

vagabunda short, thick. Pointed tail. Similar

(Wandering Tree- in type to preceding microfilaria.

Pie). Lens this eee 904 100 90

Breadth .3.....- 5 4, 4

FOUND IN THE BLOOD OF BIRDS, 63

Admission

Host. Habitat. to FHelminth.

Gardens.

PsITTACIDH. Madagascar | 24.9,17 | Microfilaria. Unsheathed, medium

Agapornis cana length, thin. Pointed tail.

(Love-Bird). Weng thier 128i tod 130

Bread bene yess 2 2-3 3

PARADISEIDE. New Guinea} 13,8.20 | Microfilaria. Unsheathed, medium

Cicinnurus length to long, thin, pointed at one

regius end.

(King Bird of Iie |) con cabooe 198u 184 205

Paradise). Breadtheeeeee 2 2 2

PLOCEID®. W. Africa 8.9.19 | Mierofilaria. Unsheathed, long,

Hypochera thick, pointed at one end, blunt

nitens end coiled.

(Shining Weaver- ENERO Soodcoone 3044 250

Bird). ; Breadth ee eesee. 5-6 5

FRINGILLID. S. Africa 21.1.2i | Microfilaria. Unsheathed, medium

Serinus scotops length and thickness, one end

(Seed-eater). pointed.

ene ieee 150 128 104

Breadth......... 4 4. 4

IcTERID®. N. America | 9.6.20 | Microfilaria. Unsheathed, short,

Quiscalus medium thickness, one end pointed.

quiscalus JOBINEANN son nhe nes 78 78 78 76 86

(Purple Grackle). Breadth......... 3-4 3-4 3-4 3-4 3-4,

ARDEIDZ. India 28.4.20 | Microfilaria. Unsheathed, long and

Bubulcus thick, end tilted and very pointed.

coromandus ene thee. 287 w —- 308 337

(Cattle Egret). Breadth......... 6-7 6-7 6-7

VULTURID®. 2 16.7.20 | Microfilaria. Unsheathed, thick,

Otogyps calvus | medium length, slightly pointed at

(Pondicherry | one end.

Vulture). Weng he eees ss: 140 pu

Breadthy ses... 8

Fanconip2. W. Africa 3.5.20 | Microfilaria. Unsheathed, medium

Hlanus length and breadth.

ceruleus Length ......... 200 u 162 180 152

(Black-shouldered Breadth......... 3) sion 4 275

Kite).

PARADISEIDZ. New Guinea} 27.10.19 | Microfilaria. | Unsheathed, short,

Paradisea medium thickness, nearly straight,

minor ends blunt.

(Minor Bird of Wemert hese ee 100

Paradise). Breadtheensnss 4

MusopHaGip™. S. America | 29.9.20 | Microfilaria. Unsheathed, short,

Turacus thin.

corythaix ene ime ee 72 w

(Louracou). Breadth......... 2

CoLUMBIDE. Chili 24.8.17 | One microfilaria, partially hidden by

Zenaida corpuscles and badly seen, medium

auriculata length and thickness.

(Auriculated ener ee ene 180 pu

Dove). rend ieee 4,

PARADISEID®. W. Africa | 7.7.19 | Microfilaria. Unsheathed, long and

Steganura thick, blunt one end, pointed the

aradised other.

(Paradise Whydah- hen sthyereerer: 316m 260 245

Bird). Breadth......... Beene oy 6

64 MICROFILARIM FOUND IN THE BLOOD OF BIRDS.

Host.

TURDIDA.

Cittocincla

MACVULA

(Shama).

TANAGRIDE.

Ramphoceelus

brasilius

(Scarlet Tanager).

Habitat.

India

Brazil

Admission |

to | Helminth.

Gardens. |

16. 7.20 | Microfilaria. Unsheathed, long,

graceful, thick and blunt at the

anterior end, tapermg gradually to

a slender pointed posterior end.

iplenc:ihiee seers 230 wp 244 244.

| Thickest

breadthivwnerse 5 5 5

18.12.19 | Microfilaria. Unsheathed, graceful,

blunt at one end which contains a

clear refractory body; other end

pointed and frequently curled,

medium length and thickness.

Length ......... 1204 100 #90

Breadth......... 6 6 5:5

ON THE VAGUS NERVES OF THE TERRESTRIAL CARNIVORA. 65

7. On the Vagus and Sympathetic Nerves of the Terres-

trial Carnivora. By Carus F. Sonntac, M.D.,

F.Z.8., Anatomist to the Society.

[Received October 2, 1922: Read February 6, 1923.]

(Text-figures 1-14.)

The course, relations, and branches of the vagus and sym-

pathetic nerves in the dog and cat are described in text-books,

and Swan (1) has recorded some of the conditions present in the

fox, Jaguar, and porpoise. But no account of these nerves in

the entire group of terrestrial Carnivora has been published.

In the present paper, which is based on the examination of

animals which died in the Society’s Gardens, the conditions

present in representatives of all families except the Hyznide,

Protelide and Cryptoproctide, are described. The course and

relations in all forms resemble those in the dog and cat.

The cervical parts of the vagus and sympathetic nerves may be

fused or separate, or both forms may be present in the neck of

the same animal.

The cervical parts are fused in :—

Family Felide :—Velis domestica, F. onca, F. sylvestris, F. ben-

galensis ; Family Viverride :—Nandinia binotata, Viverra civetia,

Civettictis civetta, Viverricula malaccensis, Paradoxurus larvatus,

Mungos mungo, M. ichneumon, Atilax paludinosus, Cynictis peni-

eillata; Family Canidee:—Canis familiaris, C. thous, CO. ben-

galensis, Vulpes vulpes, Lycaon pictus; Family Mustelidee :—

Mustela martes, Meles meles, Putorius vison, Mephitis mephitica,

Ictonyx zorilla; Family Procyonide:—Procyon iotor, Nasua

narica; Family Urside:—Melursus wrsinus. The degree of

fasion varies. It may consist of an intimate mingling of the

fibres, or the nerves may be easily separated when the fused

sheaths are divided. The fusion may occur low down or high up.

It is most extensive in Vandinia binotata, in which the ganglion

nodosum of the vagus, and the superior cervical ganglion of the

sympathetic are fused to form a lobulated mass (text-fig. 1A).

The vagus and inferior cervical ganglion are fused in Canis thous

(text-fig. 5), Paradoxurus larvatus (text-fig. 6), and Civettictis

civetta (text-fig. 7B). The cervical parts are separate in Genetta

felina, Paradoxurus hermaphroditus, and Ailurus fulgens. Both

forms are present in the same animal in a specimen of Lutra

maculicollis, the nerves being fused on the left side and separate

on the right*.

* A similar condition has already been recorded in Tamandua tetradactyla (4)

and in Anthropopithecus troglodytes.

Proc. Zoou. Soc,— 1923, No. V. 5

66 DR. C. F. SONNTAG ON THE VAGUS AND

The Vagus Nerves.

The ganglion nodosum is absent in Adlurus fulgens, Ictonyx

zorilla, Paradoxurus larvatus, Melursus ursinus, Atilax palu-

dinosus and Genetta felina (text-fig. 1 B-G). It is represented by

a thin, flat, expanded part of the nerve in Civettictis civetia,

Mustela martes, Meles meles, and Mephitis mephitica (text-fig. 2) ;

and in other genera it is a well-marked, round, oval, fusiform or

pyriform swelling (text-fig. 3). Communications run as usual

between it and the other nerves in its vicinity.

The vago-sympathetic cord (text-figs. 1, V-S.-11, V-S.) usually

Text-figure 1.

The upper cervical parts of the vagus and sympathetic nerves in: A. Nandinia

binotata; B. Ailurus fulgens; C. Paradoxurus larvatus ; D. Atilax palu-

dinosus; E. Melursus ursinus; F. Genetta felina; G. Ictonyx zorilla;

C.S : cervical sympathetic; X: vagus nerve; IX: glossopharyngeal nerve ;

XI: spinal’ accessory nerve; XII: hypoglossal nerve; G.N: ganglion

nodosum; S8.C.G: superior cervical ganglion; V-S: vago-sympathetic;

C.C.P; communications to cervical plexus; a: pharyngeal nerve; 6:

internal laryngeal nerve; é.c.v: internal carotid nerve.

separates again into vagus (V.) and sympathetic (S.) in the

posterior third of the neck, but they separate in the anterior

part of the thorax in Canis thous (text-fig. 5). I believe the

large swelling seen on the nerve in that species is a fusion of the

vagus and inferior cervical sympathetic ganglion. It represents

a greater degree of fusion than that seen in text-figs. 6 and 7B.

Pharyngeal Nerve (text-figs. la, 2a, 3a):—This nerve arises

from the ganglion nodosum or from the nerve above it, the latter

SYMPATHETIC NERVES OF THE TERRESTRIAL CARNIVORA. 67

being more frequent; and it runs to the pharyngeal plexus where

it meets branches of the spinal accessory and sympathetic nerves.

The accessory filaments usually pass through the vagus, but in

Canis familiaris (1) they run separately. In Mungos ichneumon

(text-fig. 3B) the pharyngeal nerve is given off from a laryngeal

nerve. In Procyon lotor (text-fig. 3D) the pharyngeal nerve

communicates with the superior laryngeal nerve; and in Mustela

martes (text-fig. 2B) and Civettictis civetta (text-fig. 2 A) the sym-

pathetic communicates with it before it reaches the pharyngeal

plexus.

Text-figure 2.

The upper cervical parts of the vagus and sympathetic nerves in: A. Civettictis

civetta; B. Mustela martes; C. Meles meles; D. Anastomoses between

the recurrent nerves in Meles meles; L1.V. and R.V : left and right vagi;

d. and e: right and left recurrent nerves; ¢: communicating nerves.

Other letters as in text-fig. 1

The superior laryngeal nerve (text-figs. 1b, 26, 36) is the

largest branch given off from the ganglion nodosum, or from the

upper part of the vagus when no ganglion is present. In Mungos

ichneumon (text-fig. 3B) the superior cervical ganglion of the

sympathetic gives off a large nerve (L-P-N.) which receives three

filaments from the ganglion nodosum, and supplies the pharynx

and larynx. The superior cervical ganglion, or sympathetic cord,

communicates in some cases with the superior laryngeal nerve

by small branches, or by a thick cord, as in Canis thous

68 DR. C. F. SONNTAG ON THE VAGUS AND

(text-fig. 30). The nerve gives off the external laryngeal nerve

and enters the larynx through the thyro-hyoid membrane, or

through an opening in the thyroid cartilage. It is sometimes

seen to communicate with the recurrent nerve, but never in such

a complete manner as that described in Hyraw capensis (5). No

trace of a depressor nerve was seen in any of the animals described

in the present paper, although it is described by several authors

as existing in Felis domestica.

As the middle cervical sympathetic ganglion is absent, no

Text-figure 3.

The upper cervical parts of the vagus and sympathetic nerves in: A. Felis

bengalensis; B. Mungos ichneumon: C. Canis thous; D. Procyon lotor ;

G.R: root ganglion of the vagus; a.n: Arnold’s nerve. Other letters as in

text-fig. 1.

communicating twigs are given off from the vagus till the root

of the neck is reached . There the sympathetic is given off and

communicating branches run again between vagus and sym-

pathetic. They run to the sympathetic cord itself, or to the

inferior cervical ganglion of the sympathetic (text-figs. 4-11).

In Procyon lotor they run from the vagus to the annulus of

‘Vieussens.

The cervical part of the cesophagus is supplied by the pharyn-

geal and, in some cases, the recurrent laryngeal nerves,

SYMPATHETIC NERVES OF THE TERRESTRIAL CARNIVORA. 69

Vhe right recurrent laryngeal nerve (text-figs. 4,d-11,d) has the

usual origin, course, and relations. It may communicate with

the sympathetic and cardiac plexuses. In Meles meles (text-

fig. 2D) a thick nerve unites it to the left recurrent nerve, and

in Mephitis mephitica (text-fig. 11A) it forms a loop with the

right vagus, whence cardiac branches arise.

The left vagus usually runs straight across the aortic arch, but

in Canis thous (text-fig. 5) it is very tortuous, and its branches

are also undulating, so the aortic arch is covered by sinuous

nerves.

Text-figure 4.

L.V.

The lower cervical and thoracic parts of the vagus nerves in Felis bengalensis.

L.V. and R.V: left and right vago-sympathetic nerves; V: vagi; S :

sympathetics; A.P.P. and A’.P.P’: anterior pulmonary plexuses; C.B.S :

cardiac branches of the sympathetic ; I.C.G: inferior cervical ganglion of

the sympathetic; P.G: plexus gule; T.C.S: thoracic sympathetic cords;

d and e: right and left recurrent laryngeal nerves ; £: cardiac branches of

the vagus; g and g’: pulmonary branches of the vagus.

The left recurrent laryngeal nerve (text-figs. 4,e-11,e) arises

from any point in the vagus above, in front of, or below the

aortic arch. It has the usual course, relations, and termina-

tions. It gives twigs to the cardiac plexus, and it may com-

municate with the left anterior pulmonary plexus.

Tracheal and csophageal branches arise in the thorax, and the

former make a more or less complex plexus.

Cardiac Nerves (text-figs. 4, f.f—11, f.f'):—In none of the

terrestrial Carnivora did I observe any cervical cardiac branches

70 DR. C. F. SONNTAG ON THE VAGUS AND

of either vagus; and all the branches arose within the thorax

from :—

a. The vagus nerves (constant).

6. 'The left recurrent nerve (constant).

c. The right recurrent nerve (not universal).

In all forms the vagus cardiac branches contain some sympa-

thetic filaments, but in a few species they contain the entire

Text-figure 95.

The thoracic parts of the vagus nerves in Canis thous. D.C.P. and S.C.P: deep

and superficial cardiac plexuses; R.I: first rib; s.v.b: sympathetic nerves

on subclavian arteries. Other letters as in text-fig. 4.

sympathetic supply to the heart. The cardiac plexuses anastomose

with the anterior pulmonary plexuses, and with the nerves round

the branches of the aortic arch. In some cases (text-figs. 4, 5, 6)

the vagus gives off well-marked nerves, whose bifureating

terminal twigs run to both cardiac and anterior pulmonary nerves.

The actual components of the plexuses are described on p. 76.

SYMPATHETIC NERVES OF THE TERRESTRIAL CARNIVORA. 71

Anterior Pulmonary Nerves (text-figs. 4, A.P.P.-1]1, A'.P’.P’.)

arise from the vagi anterior to the pulmonary roots, or as bifur-

cations from erie: -pulmonary branches.

Posterior Pulmonary Nerves (text- -figs. 4, 9. Vise —11, g.9'.), which

are two to four in number, arise from the vagi as they run along

the dorsal surfaces of the roots of the lungs. They may be very

short and minute, or long and thick.

Plexus Gule (text-figs. 4-11):— Between the roots of the lungs

and the diaphragm there is a more or less intricate anastomosis

between the vagi across the csophagus. There may be only

Text-figure 6.

v. AVP.

The lower cervical and thoracic parts of the vagus nerves in Paradoxurus

larvatus. s.v.b: sympathetic nerves on the subclavian arteries. Other

letters as in text-fig. 4:

branches of communication, but in most forms there 1s a mingling

of divisions of the vagi. From the plexus two cords emerge ; one

runs through the ventral aspect of the cesophageal orifice in the

diaphragm and the other through its dorsal aspect. The arrange-

ments observed by me were:

In Wandinia binotata, Mephitis mephitica (text-fig. 11 A),

72 DR. C. F. SONNTAG ON THE VAGUS AND

Oynictis penicillata (text-fig. 10 B), and Melursus ursinus (text-

fig. 11 B) the ventral cord is formed by the left vagus, the dorsal

cord by the right one, and communications run between the

nerves.

In Felis bengalensis (text-fig. 4) and Paradoxurus larvatus

(text-fig. 6) the ventral cord is a branch of the right vagus and

the dorsal cord is a combined trunk of the vagi. In the former

communications run between the vagi. In Genetta felina

(text-fig. 7) the ventral cord and left vagus are united xby

communications.

Text-figure 7.

The thoracic parts of the vagus nerves in: A. Genetta felina; B. Civettictis

civetta. Letters as in text-fig. 4.

In Civettictis civetta (text-fig. 7B), Ailurus fulgens (text-

tic. 10 A), Lutra maculicollis (text-fig. 8 A), and Putorius vison

(text-fig. 8 B) the dorsal cord is a combined trunk of the vagi,

and the ventral cord is a branch of the left vagus.

In Procyon lotor (text-fig. 9B) the ventral and dorsal cords

are produced by both vagi after forming a complex plexus gule.

In Canis thous (text-fig. 5) both vagi divide. The mesial

halves unite to form the ventral cord, and the lateral halves fuse

to form a very thick dorsal cord.

The ventral and dorsal cords end in different ways in the

SYMPATHETIC NERVES OF THE TERRESTRIAL CARNIVORA, TiS)

abdomen. In Felis bengalensis (text-fig.. 12) the ventral cord

sends branches over the ventral gastric wall and lesser curvature.

They anastomose with branches of the right vagus and off-shoots

of the solar plexus accompanying the gastric and duodenal

arteries. The right vagus supplies dorsal gastric nerves and

divides into two bundles of fibres which end in the cceliac ganglia

(C.G.), gastric (G.P.), splenic (S.P.), superior mesenteric (S.M.P.),

and left renal (L.R.P.) plexuses.

In Civettictis civetta, Atilax paludinosus, Genetta felina, Ictonyx

zorilla, Mephitis mephitica, Melursus ursinus, and Ailurus fulgens

the ventral cord divides into branches which supply the ventral

Text-figure 8.

1.C.G.

7.C.S.

The thoracic parts of the vagus nerves in: A. Lutra maculicollis; B. Putorius

vison; G.W: ganglion of Wrisberg; (2S: cesophagus. Other letters as

in text-fig. 4.

surface of the stomach and course along the lesser curvature to

the pylorus. The dorsal cord sends a few branches to the lesser

curvature and dorsal wall of the stomach and ends in the solar

plexus; it always remains large and thick.

In Paradoxurus larvatus (text-fig. 13) the conditions are very

complex. The ventral cord (V.C.) supplies the stomach, as in

the preceding forms, by thin gastric branches (g.}.), but it also

gives off the hepatic plexus (H.P.). The dorsal cord (D.C.) gives

off a large gastric nerve (G.P.), several twigs to the hepatic

plexus (H.P.), the splenic plexus (S.P.), and many communicating

74 DR. C. F. SONNTAG ON THE VAGUS AND

branches to the solar plexus (c.8.P.). It is continued into the

superior mesenteric plexus (S.M.P.), one branch in particular

being thick and reaching the cecum (Ca). These sympathetic

plexuses even if traversing the vagus cord have come from the

solar plexus.

In Canis thous the ventral cord runs along the lesser curvature

of the stomach to the pylorus, but it does not give off the hepatic

plexus. The dorsal cord supplies the dorsum of the stomach,

communicates with the solar plexus by a large cord, and sends

many branches into the solar offshoots. Many branches can be

traced through the superior mesenteric plexus to the cecum.

In Cynictis penicillata the right vagus replaces the ventral cord

in the species described above, but has a similar course. The

left one runs through the dorsal aspect of the oesophageal opening

in the diaphragm, supplies the stomach, and breaks up into fila-

ments which pass to the offshoots of the solar plexus.

It is now believed that the Arctoid Carnivora have genetic

relations to the Cetacea, and Swan (1) described the conditions

in Phocena communis as follows:—‘*In the porpoise the par

vagum communicates with the sympathetic, but is otherwise

separate from this, as in the baboon, rabbit, and others; it gives

off a small recurrent which winds round the subclavian artery

on the right side, and the arch of the aorta on the left; it sends

filaments to the cesophagus; it gives several branches to the

heart, and copiously supplies the lungs; it then passes to the

cesophagus, where its branches are more deeply imbedded in

the muscular fibres than in other animals; the greater portion

corresponding with the posterior trunk supplies the first, or

cuticular, and the second, or villous, stomach; after forming a

corona or ring on the lower part of the cesophagus, it sends fila-

ments to the diaphragm and to the left semilunar ganglion; it

also sends branches towards the other three stomachs or duodenal

pouches, a branch to the liver, and others to communicate with

branches from the cceliac plexus on the branches of the coronary

artery passing to the stomachs, and with some of the branches of

the hepatic plexus as this passes to the liver; the smaller portion

corresponding with the anterior trunk passes down and sends

some filaments to the lower portion of the csophagus and the

first stomach, but its principal part divides to join both semilunar

ganglia.”

It is, therefore, evident that there is a greater degree of com-

munication between the vagi and solar plexus in the porpoise

than in the Arctoid Carnivora, both trunks being connected to

the semilunar ganglia. Complications are introduced owing to

the differences in the characters of the stomach.

The Sympathetic Nervous System.

The superior cervical ganglion (text-figs. 1, S.C.G.-3, 8.C.G.) is

absent in Atilax paludinosus, Ictonyx zorilla, and Canis thous,

SYMPATHETIC NERVES OF THE TERRESTRIAL CARNIVORA. 75

and I could neither observe it in the neck nor in the carotid

canal. In other species it is round, oval, or pyriform, but its

posterior pole is never bifurcated. It communicates with the

ninth, tenth, and twelfth cranial nerves, and it sends branches

to the pharyngeal plexus. The internal carotid nerve (7.c.n.) is

single or multiple. Rami communicantes also run to the upper

cervical spinal nerves.

The superior cervical ganglion and ganglion nodosum are fused

in Nandinia binotata (text-fig. 1 A), but in other animals the

former rests on the vagus nerve, or is connected to it by a

sympathetic cord (S.) of variable length. In only a few species

(p. 65) is the sympathetic always separate from the vagus.

Text-figure 9.

A. Thoracic parts of the vagus nerves in Ictonyx zorilla; B. Plexus gule

in Procyon lotor. Letters as in text-fig. 4.

In no ease is there a middle cervical ganglion.

When a vago-sympathetic cord is present it is usually resolved

again into its component elements in the posterior third of the

neck. But in Canis thous (text-fig. 5) the vago-sympathetic

passes into the thorax, and expands into an oval mass whence

the sympathetic runs cranio-laterad. The sympathetic enters

the inferior cervical ganglion or the first dorsal ganglion when the

former is absent.

The inferior cervical ganglion (text-figs. 4, I.C.G.—11, I.C.G.) is

absent in Putorius vison, Atilax paludinosus, Genetta felina, and

76 DR. C. F. SONNTAG ON THE VAGUS AND

it is, im my opinion, represented by the swelling on the vagus in

Canis thous (text-fig. 5). In Civettictis civetta (text-fig. 7 B)

it is fused with the medial border of the left vagus, but it is

absent on the right side. It may communicate with the right

recurrent nerve. Branches run to the brachial plexus, and vaso-

motor filaments accompany the vessels to the fore-limbs.

In Canis thous (text-fig. 5, s.v.b.) the latter arise independently

from the vagus enlargement. Branches accompany the vertebral

artery through the cervical vertebre.

Cardiac branches reach the cardiac plexuses in three ways. In

most species they are contained within branches of the vagus.

Text-figure 10.

Thoracic parts of the vagus nervesin: A. Ailurus fulgens; B. Cynictis peni-

cillata; V.P: nerve plexus round branches of the aortic arch. Other

letters as in text-fic. 4.

In some forms they reach the cardiac plexuses through the plexus

round the aortic arch and its branches. In others they run

directly to the cardiac plexuses.

The inferior cervical ganglion is united to the first thoracic

ganglion by one or more nerves, or by an annulus of Vieussens.

Pulmonary branches may be included within the vagus, or fine

filaments may run to the lungs from the upper thoracic ganglia.

It may, however, be difficult to trace the latter.

Cardiac Pleaus:—The cardiac plexus lies between the aortic

arch and trachea, and in some species there is an aggregation

of nerves with or without ganglia, lying in the position of the

SYMPATHETIC NERVES OF THE TERRESTRIAL CARNIVORA. WE.

superficial cardiac plexus. It is connected to the anterior pul-

monary plexus on one or both sides, and to a plexus around the

main branches of the aortic arch. In some species branches of

the vagus and sympathetic reach the cardiac plexus through the

latter. The composition of the plexus in the species described in

this paper are :—

felis bengalensis (text-fig. 4):—-Two branches from the left

vagus (f), a twig from the left anterior pulmonary plexus

(A.P.P.), and a thick branch from the inferior cervical ganglion

of the left sympathetic (C.B.S.). These form a superficial plexus

without ganglia. The deep plexus receives a branch from the

right vagus (f’), a branch from the right anterior pulmonary

Text-figure 11.

Thoracic parts of the vagus nerves in: A. Mephitis mephitica ;\ B. Melursus

ursinus; V.P: nerve plexus round branches of the aortic arch. Other

letters as in text-fig. 4.

plexus (A!.P’.P’.), and a branch from the left recurrent laryngeal

nerve; but it gets no separate sympathetic filaments.

Paradoxurus larvatus (text-fig. 6):—Two offshoots of the left

vagus unite to form a cardiac nerve (f) which runs to the plexus.

No branches of the left sympathetic run directly to the plexus,

but they enter the plexus round the branches of the aortic arch

(A.V.P.). The right vagus gives two branches (/") and the left

recurrent laryngeal nerve contributes; but no branches come

from the right sympathetic. The plexus communicates with the

pulmonary plexus.

Atilax paludinosus (text-fig. 8C):—Each vagus contributes

78 DR. C. F. SONNTAG ON THE VAGUS AND

two branches (f,f’) and the left recurrent laryngeal nerve (e)

sends a twig. No sympathetic filaments run directly to the

plexus on either side.

Genetta felina (text-fig. 7 A) :—The left vagus and its recurrent

laryngeal branch send each a branch to the plexus, and the

former bifurcates (f). The right vagus sends two large cords (f'),

Text-figure 12.

Abdominal parts of the vagus and sympathetic nerves in Felis bengalensis

C.G: celiac ganglia; D.C: dorsalcord of the vagi; D.P: duodenal plexus;

G.P: gastric plexus; H.P: hepatic plexus; L.R.P. and R.R.P: left and

right renal plexuses; P.N: left phrenic nerve; P.P: phrenic plexus;

S.M.P: superior mesenteric plexus; S.N: splanchnic nerve; S.P: splenic

plexus; V.C: ventral cord of the vagi. ,

which both divide before reaching the plexus. No separate

sympathetic filaments run to it on either side.

In Civettictis civetta (text-fig. 7B) the plexus is formed by a

branch from each vagus (f.f') and a brushwork of fibres from the

left nerve. Filaments from the left inferior cervical sympathetic

SYMPATHETIC NERVES OF THE TERRESTRIAL CARNIVORA. oe)

ganglion reach it via the plexus round branches of the aortic

arch.

In Canis thous (text-fig. 5) the superficial and deep plexuses

are very well marked. The former (S8.C.P.) receives a thick cord

from the left vagus (f), several twigs from the right vagus (/"),

communications from the left anterior pulmonary plexus, and

a thick sympathetic filament (C.B.S.). The latter (D.C.P.)

is formed by branches of the right vagus (f’), left recurrent

Text-figure 13.

V.C. D.C.

A.P.

Abdominal parts of the vagus nerves in Paradoxurus larvatus. A.P: aortic

plexus; C.A: cecum; ¢: communicating nerves. Other letters as in

text-fig. 12.

laryngeal nerve (¢), and communications with the right anterior

pulmonary plexus ; but there are no separate sympathetic

i nts.

su, Cynictis penicillata (text-fig. 10B) the cardiac plexus

receives a branch from each vagus (f./'), one from the left reeur-

rent nerve (e), and communications from the anterior pulmonary

plexuses (AGBER) i yNlo branches come separately from the right

sympathetic, but the left inferior cervical ganglion (1.C.G.) sends

80 DR. C. F. SONNTAG ON THE VAGUS AND

numerous fine filaments to the aortic plexus, the latter being

connected to the cardiac plexus.

In Mephitis mephitica (text-fig. 11 A) a marked plexus grouped

round the great vessels (A.V.P.) receives branches from the left

vagus and left inferior cervical ganglion. The cardiac plexus

receives a twig from each vagus (f.f’), one from the leit recurrent

laryngeal nerve (e), and a brushwork of filaments from a right

vago-sympathetic loop.

In Melursus ursinus (text-fig. 11 B) superficial and deep cardiac

plexuses are present and communicate with pulmonary plexuses.

The superficial plexus gets a branch from the left vagus (f), but

both vagi and the left recurrent laryngeal nerve supply the deep

one. Neither plexus gets separate sympathetic filaments.

Text-figure 14.

A." Paradowurus larvatus, showing the thoracic sympathetic cord (T.C.S.)

becoming the splanchnic nerve (S.N.); B. Canis thous, showing the thoracic

cord (T.C.S.) becoming the great splanchnic nerve, the small splanchnic

filaments ending in the left suprarenal plexus (L.S.R.P.) and a sympathetic

ganglion giving rami communicantes to spinal nerves (D. 14 and L. 1);

C. Solar plexus in Meles meles: 1.M.P: inferior mesenteric plexus. Other

letters as in text-fig. 12.

The Thoracic Gangliated Cord does not always possess the same

number of ganglia as intercostal nerves. When ganglia are

absent the rami communicantes meet the sympathetic cords in

T- or V-shaped junctions. Ganglia may be minute or well-

marked but are larger in marine than in terrestrial Carnivora.

The splanchnic branches vary as follows :—

a. One thick cord arises on each side in the thorax and breaks

up into several filaments before entering the solar plexus :—

Felis bengalensis (text-fig. 12), Civettictis civetta.

6. One large thoracic cord and some fine abdominal nerves

constitute the splanchnic nerves:—Velis domestica, Cynictis

SYMPATHETIC NERVES OF THE TERRESTRIAL CARNIVORA. 81

penicillata, Ictonyx zorilla, Mephitis mephitica, Ailurus fulgens, and

Melursus ursinus.

ce. The thoracic cord becomes the main splanchnic nerve, and

the others come from the abdominal sympathetic. The remainder

of the sympathetic cord appears as a branch of the main splanch-

nic in Paradoxurus larvatus (text-fig. 14A). In Canis thous

(text-fig. 14.B) the main splanchnic cord gives a branch to a

ganglion whence the abdominal sympathetic passes caudad, and

two long rami communicantes run to the last dorsal and first

lumbar nerves.

Branches of the thoracic cord to the aortic plexus vary in

prominence.

The Solar Plexus varies considerably in details, but it is built

on the same general plan in all. It lies on one side of, or is

wrapped sound the ceeliac axis. It has always at least one large

ganglion, but smaller ones may be present in addition. It receives

the dorsal vagus cord and splanchnic nerves, and its offshoots

accompany the various abdominal arteries, interlacing at their

terminations. Sometimes the hepatic and splenic plexuses are

apparently given off from the right vagus. Superior and inferior

mesenteric ganglia may be present, and these are most marked

in Meles meles (text-fig. 14C). As filaments of the right

vagus get into many, if not all, of the offshoots the abdominal

organs get a rich supply of both vagus and sympathetic fila-

ments. The following list contains an enumeration of the

offshoots :-—

1, Phrenic plexus. 7. Renal plexuses.

2. Gastric i, 8. Suprarenal plexuses.

3. Hepatic ,, 9. Spermatic Ms

4, Splenic _,, 10. Ovarian As

5. Duodenal ,, 11. Inferior mesenteric plexus.

6. Superior mesenteric plexus. 12. Aortic plexus.

All the offshoots are not equally obvious in all the Carnivora.

The most variable is the duodenal plexus, which depends on the

presence or absence of a large duodenal branch of the superior

mesenteric artery (text-fig. 12).

The Sympathetic Nervous System in Phocena communis.

Swan (1), in his deseription gives the following data :—

1. The pyriform superior cervical ganglion of the sympathetic

nerve sends one portion upwards and one downwards.

2. The sympathetic communicates with the superior laryngeal

nerve.

3. The inferior cervical ganglion on the right side communi-

cates with the vagus, but the left one does not.

4, The thoracic ganglia are well marked, and on the whole

larger than in other mammalia; they give off aortic and

Proc. Zoon. Soc.—-1923, No. VI. 6

82 DR. C. F, SONNTAG ON THE VAGUS AND

splanchnic nerves, rami communicantes and late ral branches to

the rete mirabile on the thoracic parietes*.

5. The aortic plexus has a large ganglion.

6. The rectum and bladder receive small nerves.

7. There is no perceptible difference in the nerves to the upper

and lower part of the intestines, just as these parts of the gut

merge gradually into one another, and there is no capacious,

cecum.

It is, therefore, evident that the sympathetic nerves differ in

many respects from those in the Fissipede Carnivora.

Swan has also made several generalizations on the sympathetic

nerves in the Mammalia, and the conditions described in this

paper furnish illustrations of his remarks which, as far as the

Carnivora are concerned, were made on a very limited series of

animals.

Summary and Conclusions.

1. In most Carnivora Fissipedia the cervical parts of the vagus

and sympathetic nerves are fused. In most cases fusion only’

affects the cords, but in a few species the ganglia are also united.

Fusion consists either of an intimate mingling of fibres or

adhesion of sheaths only.

2. The ganglion nodosum is frequently absent in both the

neck and the foramen lacerum posticum; and the superior

cervical ganglion of the sympathetic is likewise absent in a few

species.

3. No middle cervical ganglion is present, and inferior cervical

ganglia are frequently absent.

4. Sympathetic cardiac nerves are frequently contained

entirely in branches of the vagi.

5. The cardiac plexus is sometimes divisible into superficial

and deep parts, but ganglia are uucommon.

6. Pulmonary nerves may arise from cardiac branches of the

vag,

7. The plexus gule varies in complexity. It may be absent.

In most species ventral and dorsal cords produced by a varying

degree of intermingling of the vagi pass from thorax to

abdomen.

8. The ventral vagus cord supplies the ventral gastric wall

and branches run along the lesser curvature to the pylorus.

9. The dorsal vagus cord supplies the dorsum of the stomach.

It ends directly in the celiac ganglia or in one of the offshoots of

the solar plexus, or it is diffused throughout the plexus.

10. The number of ganglia on the thoracic sympathetic cord

does not always correspond to that of the intercostal nerves.

* The rete disseminated throughout the body enables the animal to accommodate

itself to variations in the water pressure when it dives or rises to the surface; and

the sympathetic nerves running to the thoracic rete form part of the accommodating

neuro-vascular mechanism.

SYMPATHETIC NERVES OF THE TERRESTRIAL CARNIVORA. 83

11. The superior and inferior cervical ganglia ave variable, but

the middle cervical sympathetic ganglia are absent.

12. Thoracic sympathetic ganglia are absent, smal] or large.

13. The splanchnic nerves may be branches or continuations of

the thoracic cords, and a branch of the great splanchnic becomes

the abdominal sympathetic in the latter case.

14, Well-marked nerves are traceable from the vagus and solar

plexus to the cecum.

15. The vagus nerve may give off plexuses which arise from

the solar plexus in most mammals.

16. The vagus and sympathetic nerves of the terrestrial Car-

nivora differ in many details from those of Phoceena communis,

the latter being more complex in several respects.

Bibliography.

1. Swan, J.—Illustrations of the Comparative Anatomy of the

Nervous System. London, 1864.

List of former papers on the vagus and sympathetic nerves :—

2. Sonntag, C. F.—P. Z. 8S. 1921, pp. 572-575 ( Phascolurctos).

: .Z.8. 1921, pp. 873-876 (Marsupialia).

.Z.8. 1922, pp. 99-108 (Hdentata).

.Z. 8. 1922, pp. 149-156 (Hyras).

.Z.8. 1922, pp. 444-449 (Wandrillus).

> Gr & OO

pas lachae

PZ. S993) “CONNBERG ebolr

PALEARCTIC. BEARS.

‘SavV4ad = OILOYVATVd

py! YOSsfe!ueg Wy suog'ajeg uyor ms) ‘'S

I'ld “SYSENNOT ‘E2EI “SZ ‘d

REMARKS ON SOME PALEARCTIC BEARS, 85

8. Remarks on some Palearctic Bears.

By Einar Lonnzere, F.M.Z.S., ke.

(Plates I., I1.*)

[Received November 15, 1922: Read February 20, 1923. |

Among the collections which the Royal Nat. Hist. Museum in

Stockholm recently has received from China, chiefly through the

courtesy of Professor J. G. Andersson, there is also material of

two different kinds of Bears which are likely to arouse great

interest. Therefore I take the pleasure of laying before the

Society the following notes, in which I endeavour to prove that

the pruwinosus Bears must be regarded as so different from

other Bears that they should form a separate group of sub-

‘generic value, and also what is to be understood by Ursus lasiotus

Gray.

A Bear of the pruinosus Group.

On the 14th of August, 1921, Mr. D. Sjolander obtained a young

Bear, evidently of this group, in the Min-Shan Mountains, South-

Western Kansu. With regard to its colour, it does not closely

correspond with Lydekker’s plate of U. pruinosus (Proc. Zool. Soc.

1897), but there is an agreement in pattern which may be of more

importance. ‘The present specimen has the snout pale yellowish

grey, with a dark brown area around and especially below the eye.

Forehead and sides of head rather rich buff, but with the con-

cealed parts of the hair blackish brown; on the occiput the

colour is rather more cinnamon-rufous. The ears are richly

clothed with long, shaggy fur, blackish brown in colour, and the

same colour extends also over an area below them. A broad

white band extends across the chest and up in front of the

shoulders so as to meet dorsally and form a collar around the

neck; but a branch also extends backwards across the upper part

of the shoulders, so that by this and the collar, a large oval patch

of blackish colour (but partly with yellowish tips) on top of

the withers (interscapular region) is surrounded, except on the

posterior side. From the posterior end of the posterior white

branch a rather narrow yellowish-grey stripe continues down-

wards, and thus helps to define the black fore limb from the

body. The back and flanks are black, with more or less numerous

yellowish tips to the hairs. The hind limbs are black like the

fore limbs. Although the colour differs in the different individuals

* For explanation of the Plates, see p. 95.

86 PROF. Ee. LONNBERG : REMARKS ON

(which is a common thing among Bears), the pattern thus described

may be recognized on Lydekker’s plate, and on Sven Hedin’s

photos, published by Leche in his report on the zoological speci-

mens collected by that explorer (‘Scientific Results of a Journey

in Central Asia, 1899-1902,’ vol. vi. part 1, Stockholm, 1904).

It will appear from this, as well as from the descriptions by

various authors, that, in spite of the differences in colour

which have been observed on Bears named “ pruinosus” or

“lagomyiarius,” there is a certain pattern common to all. Our

knowledge about these Bears is very unsatisfactory, and it is for

the present impossible to say whether they constitute more than

one species or subspecies; but, nevertheless, they appear to form

together a systematic unit, which differs from the common Ursus

arctos or the genus Ursus s. str., to which they usually have been

referred, This may be proved by the structure of the feet and

the teeth, as will be shown below.

Through the investigations of Mr. R.1. Pocock, it has been proved

that the Bears form several natural groups (by him considered as

genera), which may be distinguished by means of the different

structure of their feet. The disposition of hairiness and naked-

ness, and the degree in which the digital pads are free from or

connected with each other, give the distinguishing characteristics.

An examination of the feet of this pruinosus Bear gives the

following results :—

The digital pads of the second, third, and fourth fingers are

basally closely connected, so that there are only shallow depres-

sions between them. The corresponding depression between the

fourth and fifth fingers is somewhat broader and more pronounced,

while the cleft between the second and first is still deeper. ‘The

interspaces between the digital pads are, however, in all cases

completely naked. The digital pads of the fifth and, somewhat

more narrowly, the first fingers, are connected by a naked area

with the plantar (palmar) pad. Between the latter and the

digital pads of the second, third, and fourth fingers there is a

matting of rather thick and long hairs which are directed for-

ward so as to partly cover the basal parts of the digital pads.

This matting appears continuous, but a closer examination proves

that it really consists of four patches, because, if the hairs are

divided, there is found a hairless tract connecting each digital

pad with the plantar pad, although this is concealed by the over-

lying hairs (¢f. Pl. I. figs. 1 & 2).

The plantar pad is transverse and somewhat broader on the

lateral side. In the present specimen there is only a slight crease

visible, opposite the interdigital space between the second and

third fingers. It is very widely separated from the carpal pad

by a broad and thickly hairy area. The latter pad is rather

small, somewhat elongate in shape, but not transversely

expanded.

With regard to the relation between the plantar pad and the

earpal pad, our pruinosus Bear thus resembles Huarctos, Ursus,

SOME PALEARCTIC BEARS. 87

and ‘“ Danis,” and differs from “ Arcticonus” (=Selenarctos *),

Tremarctos, Helarctos, &e. he structure of the anterior portion

of the fore feet of pruinosus is, however, very different from

that of Ursus, because the interspaces between the digital pads

are hairy in the latter; in fact, the digital pads of Ursus are

entirely surrounded by hair, because the naked strips between

the digital pads and the plantar pad are so exceedingly narrow

that they are even difficult to find when one is looking for

them; still less are they plainly visible like those between the

digital pads of tie first and fifth fingers on one hand and the

plantar pad on the other, as is described above, in the case of

pruinosus. With regard to this latter detail, the anterior portion

of the fore feet somewhat resembles the corresponding parts of

thibetanus (as figured by Pocock, 1914), but the digital pads of the

latter are much more distinct and free from each other than in

the prwinosus.

The structure of the fore feet of Hwarctos is essentially similar

to that of Ursus (at least in a specimen of the Alaska race

which I have had for comparison), and it differs thus in the same

way from pruinosus. The fore feet of the Grizzly Bears appear,

to judge from Pocock’s description, to differ from those of Ursus

in haying the digital pads more connected with each other and

without hair between them, in which respect they resemble those

of pruinosus. In Pocock’s figure (1918) of the “right fore foot

of Danis horribilis” there are not to be seen any naked tracts

connecting the first and fifth digital pads with the plantar pad,

which is so characteristic for the fore feet of the pruinosus. The

latter appears thus to differ from ‘“ Danis” as well as from the

other Bears with regard to the structure of its fore feet.

The digital pads of the hind feet of our pruinosus are quite

fused together basally, although the notch between the first and

second toe is deeper than the others. They are not at all sepa-

rated from each other by hairy tracts. The first and fifth digital

pads are broadly connected with the plantar pad by means of a

naked area (in the same manner as on the fore feet). Between the

three middle toes and the plantar pad is a transverse area thickly

covered with hair. If these hairs are divided with the aid of a

pair of pincers, there is, however, to be seen a naked strip of skin

connecting also these digital pads with the plantar one, although

this is not visible without such a proceeding (cf. Pl. I. figs. 3

& 4). The plantar surface is naked to the heel, and there is no

notch or depression covered with hair on the hallucal side, only a

slight superficial crease indicating the limit between plantar and

heel pads. In the absence of this hairy depression on the hallucal

side, the hind foot of the pruinosus difters from Ursusand EHuarctos.

Both these genera are also different from prwinosus in having the

* Sowerby has drawn attention to the fact that Heude already (1901) gave the

name Selenarctos to an assemblage of Black Bears, among which also thibetanus is

found; and Sowerby, as “first reviser”’ of the group, selects this one as the type for

Heude’s genus (cf. Journ. Mamm, 1920, pp. 216-17).

88 PROF. E. LONNBERG : REMARKS ON

digital pads surrounded by hair and quite free, not fused basally.

The hind foot of ‘ Danis” appears more similar to that of prui-

nosus by having at least two of the digital pads fused, and the

others more closely connected than in the genera just mentioned,

and without hair in the interdigital spaces, as well as with regard

to the weakness of the hallucal depression and the absence of hairs

in the same, The likeness between ‘“ Danis” and the pruinosus

group is, however, not complete, because judging from Pocock’s

figure (1918) it has no naked connection between the first and

fifth digital pads on one side and the plantar pad on the other.

Such a connection is, however, visible in the figure of the hind foot

of “ Tremarctos thibetanus,” figured by the same author in 1914;

and in fact this figure exhibits several features similar to those of

the pruinosus, but the hind foot of the Thibetan Bear has quite

free digital pads and hair in the interdigital interspaces. The

fore feet of the latter are also very different in structure when

compared with those of the pruinosus group, as they have a very

large carpal pad expanded across the whole plantar surface and

only separated by naked and soft skin from the plantar pad.

A comparison between the feet of Tremarctos and those of the

pruinosus group is scarcely needed. The fore feet of the former

have entirely free digital pads entirely surrounded by hairs, so

that a careful examination is needed to reveal the narrow and

incomplete connections with the plantar pad. The plantar pad is

broadly connected with the rather large carpal pad by means of

naked skin on the ulnar side; it extends also backwards on the

radial side, and is there connected with a small pad. On the hind

feet of Tremarctos as well the digital pads are free and rather

thickly surrounded by hairs, so that the connections between the

digital pads and the plantar pad are entirely concealed until the

hairs are artificially divided.. With the feet of Helarctos those of

the prainosus group have no resemblance, as the former are much

less hairy, and this is, of course, still more the case with those of

Melursus.

It is thus evident that the structure of the feet of the pru-

nosus Bears differs from that of all other Bears, and most certainly

from that of Ursus s. str. The question is then, whether this

difference is also connected with some other morphological

differences.

Pocock has demonstrated that the noses of different Bears are

different in structure. It is very difficult to judge only from a

dry skin, but it appears as if the naked tract between the rhina-

rium and the upper lip was broader in the present pruinosus

specimen than in, for instance, U. arctos, Huarctos, Selenarctos, and

Tremarctos, but of course not so broad as in Helarctos. In the

present specimen, dry as it is, it measures about 1 em., and is

equal in breadth to the narial septum.

As the pruinosus Bear from Kansu is rather young, the

measurements of its skull have only relative value; but, thanks to

the courtesy of Professor N. Holmgren, I have been able to

SOME PALEARCTIC BEARS. 89

measure also the skull of a very old male which was brought

home from Thibet by Dr. Sven Hedin. The latter skull has the

teeth extremely worn, so that their dimensions are of little value,

but the other measurements are useful for comparison :—

Old specimen Young specimen

from Thibet. from Kansu.

mm. mm.

Greatest length of skull ........0....c0.c0cccces eee eee 351 292

Condylobasalileng tht wuss. 5.5... heieee een eeeeeeE 331 285

Basicramialéleng thw wacecer ccc cs cece econ eee ene 311 =

/IEXDITENAKS VDL Coo ease asaEBEHe SOR don dhe caaksoo: tines cos 217 153

Mengthyonmasals yi) .. sa). Gen cceeeeee ee eae — 90°5

Length of palate from gnathion .................. 169 153

Width of palate inside middle of m? ............ 515 4A,

Distance from foramen lacrymale to gnathion. 134°5 118°5

BreadEhot brain=case)es.csesc1 seeneeni renee eee 101 101

Imteronbitallibreadthi. ss .se--)..4 eva eneeeeeeeetne 69 63

Mastoidh breadth .fcciiis.. cosas eavem tue anon 157°5 =

Hind margin of m? to front of # ............... 6. 150 147

Combined length ot pt, m1, and m? ...........5... 79 83'2(84)

RE eT ALS 8. cg ad odie ve suis aeRO 18°3X14 18°7<15°5

TO ob gab ocd ER SERED RERERACERCT REE SBS Ui Gio oon codebase: 2419 26°5 x 19°2

Urs Soho EO LOG aA EOC LOR RCE REE SER ER ee Sen oeBoe mobos 39X21 Al &22°2

Combined length of p4—2g......... 66... cece eee eee 88 95

Gin a Ae ec = 13°8X8°7

Bear skulls are very variable in shape, and it is thus rather

difficult to say whether a certain characteristic exhibited by an

individual is of any taxonomic value or not. In the pruinosus

specimen from Kansu the nasals are rather long, and their pos-

terior ends extend much beyond the frontal processes of the

maxillary. The mesial length of the nasals is also longer than

the mesial frontal suture. In Swedish Bears the opposite as a rule

prevails, and I believed, therefore, that this might constitute a

characteristic of some importance. This is, however, not the case,

because I found later that, in a young Swedish Bear from the

province of Jamtland, the nasals were just as in the pruinosus

specimen, being considerably longer than the mesial frontal

suture,

For the present it appears very difficult to point out any

definite cranial characteristic, which in every case holds good as

distinctive between the pruinosus and arctos groups, except those

that are derived from the teeth, or stand in connection with their

development.

The teeth of the prwinosus group are, especially the molars,

very much larger than the corresponding ones of the arctos

group (cf. the table above). Thesame is also the case with p* and

p, On m* of the former the cingulum is quite well marked

90 PROF. E. LONNBERG : REMARKS ON

off, even on the inner side of the tooth and not only on the

outer. On m? it is very strongly developed on the inner side,

where it forms a distinct shelf (Pl. II. fig.5). On the outer side it

is weaker, but quite traceable. p, has a well-developed antero-

interior cingulum cusp.

The ereab size of the molars is relative as well as absolute (of.

Pl. IT. figs.5 & 6). The greatest length of m° that I have ever

seen when examining a considerable number of Swedish Bears is

35 mm., while the same tooth in the present specimen of the

pruinosus group measures 41 mm. The difference in breadth is

still more striking, because m° of Swedish Bears is seldom more

than about 17 mm. broad, while in the present prunnosus s(eear

men it even exceeds 22 mm. The combined length of p*, mi, and

m* is in Swedish male Bears, as a rule, not more than about

70 mm. and often less, and among the skulls examined by me it

was only once 73 mm.; in the young pruimosus, however, it is as

much as 83 (84) mm. The dimension in question is, in adult

males of the former kind, less than the distance hetween m? and

the processus postglenoideus, and also less than the interorbital

breadth, but in the prutnosus the former dimension is larger than

ie wORO thers: | (i ALWN El Ayodteaitnlae sl esiey nay es

In pruinosus the combined length of these three teeth is more

than half the mastoid breadth, but. in arctos considerably less.

The superior size of the teeth of prainosus may be proved by still

more comparative measurements, but the samples mentioned may

be enough. It is, however, of interest'to observe that the teeth

of the prwuimosus specimen are not only absolutely and compara-

tively larger than those of the typical arctos, but also than those

of the big Black Bear from Mongolia, which I consider identical

with U. lasiotus Gray (conf. below), and which belongs to the

arctos group. This holds good, although this Mongolian Bear is

very much larger than pruinosus; and if the comparison between

the combined length of the three teeth mentioned and the other

dimensions quoted above is repeated with regard to the Mon-

golian Bear, the same result is obtained as with arctos. By this

it appears to be proved that the difference between the pruinosus

and the arctos groups is distinct enough in this respect.

The enormous size of the last premolar, and the molars of the

lower jaw can also be seen from the table of measurements above,

so that further comments on this may not be needed. The big

Mongolian Bear is also as regards the teeth of the lower jaw,

very much inferior to the prauinosus.

As the ie recorded measurements prove, the combined length

of p,, m,, m,, and m, is considerably greater than half the length

of the ae in the adult pruinosus *; but in the Bears of the

arctos group—the big Mongolian one (cf. below) included—the

former measurement is even less than half the length of the palate

in the adult males. In the latter the combined length of the

* The much worn teeth of the very old specimen do not give very satisfactory

measurements, but the corresponding dimension of the young animal must be

compared with the palatal length of the older one.

SOME PALEARCTIC BEARS. | 9]

teeth mentioned is about equal to half the mastoid breadth, or

perhaps hardly that, but in the pruinosws group the former

measurement is about from 56 to 60 per cent. of the latter.

In consequence of the great size of the mandibular teeth

and the resulting great length of the tooth series in pruznosus,

m, has been pushed backward so to speak, so that the posterior

portion of the same is concealed by the processus coronoideus when

viewed from the side. This characteristic, which also has been

observed and mentioned by Leche when he described the mammals

brought home from Thibet by Dr. Sven Hedin (1. c.), serves easily to

recognize a mandible of the prucnosus group, because in the now

living Bears of the arctos group, m, is in its whole extent visible

in front of the processus coronoideus.

Of the Grizzly Bears, I have unfortunately no material for

comparison, but with regard to the skull of a fossil Cave Bear

I have had the opportunity of stating that it exhibits the same

relative dimensions as the recent arctos in the cases mentioned

above. Thus the combined length of p*, m', and m? is shorter

than the distance between m” and the processus postglenoideus,

and likewise the former dimension is shorter than the prevrbital

width and less than haif the mastoid breadth. The combined

length of Py My My and m, of the same skull is even contained

45 dames in the length of the palate, and is less than half the

mastoid breadth.

In consequence of these facts, I am inclined to consider that the

Bears of the pruinosws group (whether it consists of only one or

more species or subspecies) are so different from other Bears that

they are entitled to subgeneric rank. This new subgenus I

propose to call Mylarctos; it is characterized by its very large

molariform teeth and foot structure, as described above.

Ursus nasiotus Gray.

A fine, big male Bear, procured in Northern Mongolia by the

Swedish missionary, Mr. Larsson, and through the courtesy of

Professor J. G. Andersson presented to the Stockholm Museum,

must, according to my opinion, be named as above. It agrees with

Gray’s short description, bemg black with brown nose and some-

what brownish on the head in front of the ears, in consequence of

such tips to the hairs. The ears agree with the specific name,

being richly covered with long black Wages as well inside as outside.

On the sides of the neck there is in some shades of light a faint

chestnut-reddish lustre. The under-fur is well developed and

dark brown. The claws blackish horn-coloured. In a mounted

state the specimen stands a little more than 1 m. at the fore

quarters.

Gray’s name (/asiotus) of 1867 has by later authors been more or

less discarded. Even in 1869 Sclater identified Gray’s Bear

with U. piscator Pucheran, which latter name referred to a

Bear from Kamtschatka. It is true ‘that not much is known

concerning Gray’s lasiotus. It was “imported from Northern

92 PROF. E. LONNBERG : REMARKS ON

China, and was stated to come from the interior of that country”

(Sclater). There appears, however, to be little reason to believe

that an animal with such a history came from Kamtschatka.

On the contrary, it must be considered far more probable that it

really has come from the northern or interior parts of the Chinese

empire, e.g. from Mongolia.’ As it is now proved that in fact a

Bear with an exterior appearance agreeing with Gray’s descrip-

tion of U. lasiotus lives there, am inclined to identify it with

Gray’s Bear.

It is therefore a matter of secondary importance to make out

whether this big Black Bear of Mongolia and the interior of

China is identical or not with the Fishing Bear of Kamtschatka.

As long, however, as nothing is known about this, all kinds of

guessings are unnecessary, and Gray’s name U. lasiotus may stand

for the big and Black Mongolian Bear *.

Dimensions of the skull of the Mongolian Black Bear,

Ursus lasiotus Gray.

mm.

Greatest lengthy: pease iiis eee Sec ateeer en cts ene cases 387

5 width . Foe SE ACA CET te UICC Sane Seo eae jucen are oe 218

Interorbital auth etna tian Rae Mer M nics mateo. 80

Tenth at mumcale chon endl oP apace TAR Reh Tee anne: 65

5 :. FREI LOT THONG ChE CPFOM Bc co anaces oon Dad ove eed an5 83

Width of muzzle across alveoles of canines ................. 78

ss palate inside m! ............ eh he Ree eR 48°3

a posterior part of e Bega seman Beste 4.7

Tesuk Grdth ot ualave Bend mn oles bits MASA EE a SU 44,

Width of palate at the premolar diastema ................-.... 60 —

5 skull outside middle of m1! ...........e ccc cee cee eee ee 82°5

3 VAIN CASOM chee eee shire date Rane Come ice eee aie 106

IPEIAEHEN OP THOSENS TEERANK oon obeo0 con soa en 2noawticns quo daaaneoseorA 117

Greatest combined breadth of nasals .............0000000e see ee 37

Distance from hind margin of palate to erosion Hadicersetees 188

MS Fey) CHONG 0) ATE NINO) coonebion soudeodoppaedasogue aut 14:7

Krontioncaninemoubackaoten alee aren nereee eee enenee ee ee eet 132

Combined length of 4, m1, and m2) (20... ... 0. see eseee eee 75

1Dfssa¥ss ol Sona oy RaEHERER Re AA cA Sees MANA Am KBR MANIsER see trae asec sand. aa coc 165

I BIRSANG UH] Teorey ain een wale Maoh hecamtedcste secbieaciedinns aac san ght 13°3

Pismetiot el = eit ek ve bh oe oer bel seis Shee sania Mee eaa a ys aes TE 24

Breadth: ofa ly shies Vel aaenlins Diyala. ce bes Pack nen n ana anette ies

Meng bla oss are be Mek) uhh ieee: chy Wal ae dha enl apt eek eS Se Reta a ae 35'3

TESe=e 6 6 NEO) 7a ee ee Bee iis eth Base Ar dete be Ae 185

Length of lower jaw ....... GAP emansuntaseake vattsions 253

Depth of lower jaw at aRalitls ee TS RR LU Rist ce aay 59

Combined length of y4, 1, mo, and mz... .. cece eee eee ee 845

* Since writing the above I have had the opportunity of seemg some Bears from

the Kamtschatka Peninsula which I suppose must be regarded as Ursus piscator

Pucheran. There has been no time for a thorough examination of the skulls, but

the skins certainly look very different from the specimen which I consider to be

SOME PALEARCTIC BEARS. 93

Mr. A. Sowerby has recently published a review of “ Heude's

Bears in the Sikawei Museum and on Bears of Palearctic Kastern

Asia.” In this paper he accepts Heude’s specific name cavifrons,

and attaches the same to a Bear of N.W. Manchuria. Sowerby

himself has shot a Bear of this kind in N. Kirin, Manchuria.

According to the description, the exterior of this Bear must be

very similar to that of the present specimen, as it is said to be

“generally black, merging into brown on the muzzle; brownish

ony themheadE yy was... So far there is nothing which prohibits

the specific identity of this Bear with Gray’s U. lasiotws and the

present specimen,

Mr. Sowerby has also published some measurements of his Bear

from Kirin, which may be compared with the corresponding ones

of the present specimen as recorded in the accompanying table.

The greatest length of the skull of Sowerby’s Bear is recorded as

16 in., or about 405 mm., thus only 18 mm. more than the pre-

sent specimen. The greatest width of the former is 9°25 in. or

about 234 mm.; the interorbital width is about 88 mm. If

“‘ovreatest width of cranium” is to be understood as width of

brain-case, this dimension, about 108 mm., is rather similar to

that of the present specimen. Some of the other measurements

recorded by Sowerby are less easily understood, and some are

certainly larger than those of the present specimen. ‘This is

especially the case with the length of the lower jaw. As the

Bears generally are very variable, it is difficult to decide whether

these two are to be regarded as belonging to the same species or

not, for the negative conclusion emphasizes the fact that Sowerby

refers Heude’s cavifrons to ‘‘Speleus.” As characteristic of the

latter he mentions ‘‘ very high foreheads so that the cranial out-

line at this point is strongly concave.” Heude’s figure of the

type shows also such a condition. In opposition to this our

Mongolian Bear shows a cranial outline which at the forehead is

nearly straight. The question then presents itself: How much

value can be attributed to such a characteristic as a more or less

concave or straight facial profile line? With my knowledge

about our Brown Bears in Sweden, I am not inclined to overrate

this characteristic, because I have found that it is very variable in

them. We have, for instance, from the same tract of Southern

Lapland, Bear skulls with straight profile and others with the

profile just as concave as Heude’s figure of cavifrons. This fact’

does not, of course, prove that the variability of the Bears of

Mongolia and Manchuria in this respect is as great as in Europe,

but there is always the possibility or even probability that this is

the case.

The important cranial characteristic which Gray mentions as

U. lasiotus Gray. They are all much paler than the latter—brown, brownish grey

or lighter,—but even if they bad been black, and I am told there are also very dark

or black Bears in Kamtschatka, I think that such specimens must be easily recog-

nized by their softer and much more shagey fur than the Mongolian Bear. Bears

which I have seen on several occasions in Zoological Gardens under the name of

U. piscator were also similar to the present Kamtschatka skins.

94 PROF. E. LONNBERG: REMARKS ON

distinguishing the Grizzly Bears from the true Ursus of the arctos

group, viz. “the palate narrow and contracted behind,” is not

mentioned for ‘‘cavifrons,” and it is not known if its palate has

this characteristic shape or not. If such should happen to be the

ease, our Mongolian Bear has nothing to do with it, because the

latter has a broad palate, which is not more contracted behind

the molars than is the case with true arctos; and it therefore

certainly belongs to the same group as the latter.

Mr. Sowerby also mentions another of Heude’s Bears, called by

the latter ‘“ Ursus mandchuricus.” It is also a large and dark

Bear, although not so black as “cavifrons,” and is said to. have

a “fairly straight cranial outline.” It is possible that this is

identical with Gray’s lasiotus, and, if such is the case, the latter

name of course has priority. The question about the identity can

hardly be decided for the present, but so much ought to be

certain, that nothing prevents Gray’s name from being Jaid on

the present big and Black Bear from Mongolia, which evidently

is a member of the arctos group.

With regard to the general size, the Mongolian Bear appears to

be larger than the European, and the greatest length of the skull

of the former (387 mm.) is larger than that of any Swedish Bear

skull I have seen. The three largest specimens of the latter

kind that I have had the opportunity of measuring, have had

maximum lengths of 362, 367, and 372 mm. With regard to

the zygomatic width, the Mongolian Bear is not superior to some

of the largest Swedish Bears. On one occasion I found the same

breadth, viz. 218 mm., in one of the latter, but several times still

greater dimensions, é. g. 223, 225, 229, and even 231 mm. The

latter is thus similar to Sowerby’s specimen in breadth. With

regard to the interorbital width (80 mm. in the Mongolian Bear),

IT have seen several Swedish Bear skulls as large or still broader,

e. g. 80, 83 (twiee), 84. and 89 mm.—the last thus fully equal to

Sowerby’s specimen. Otherwise the Mongolian Bear is greater

in most dimensions or near the maximum. ‘The palate is, how-

ever, often broader in male Swedish Bears.

The comparative size of the teeth is to be seen from the

following :—

p* in the Mongolian specimen 16°5 mm.; in Swedish male Bears 15-165 (once 17%

and once 18) mm.

Te xp ” 39 24 mm. 3 3 » 21-23 (once 20)

mm.

We og » » 35°3 mm. ES 35 5, 82-85 (once resp,

31, 30, and 29

mm.).

The difference in this respect is therefore not so very great

Unfortunately, Sowerby has not given any measurements of the

teeth of “cavifrons,” but Heude’s figures of the teeth, which are

said to represent the actual size, do not indicate that the teeth of

his Bear are larger than those of an average Swedish Bear, and

m* appears to be rather narrow.

Fig. 1.

» 2.

33 oe

Bs eke

Fig, 5.

» 6.

SOME PALEARCTIC BEARS. : 95

EXPLANATION OF THE PLATES.

Puate I.

Photograph of the lower side of the right fore foot of a pruinosus Bear

(skin) from Kansu.

Diagram of the lower side of the right fore foot of a pruinosus Bear to

show the distribution of the hairy areas and the connections between

the digital pads and the plantar pad.

Photograph of the lower side of the right hind foot of a pruinosus Bear

(skin) from Kansu.

Diagram of the lower side of the right hind foot of a pruinosus Bear to

show the connection between the digital pads and the plantar pad.

Puate II.

Upper jaw with dentition of a young prauinosus Bear from Kansu.

Lower jaw with dentition of a young pruinosus Bear from Kansu.

Fini ie: Genes

Titties

eatiae al i

TaN

LEANDER LONGIROSTRIS AND OTHER BRITISH PRAWNS. 97

9. Some Notes on Leander longirostris M. Hdwards, and —

other British Prawns. By Roperr Gurney, M.A.,

F.LS., F.Z.S.

[Received November 18, 1922: Read February 20, 1923.}

(Text-figures 1-6.)

ConTENTS.

Page

Tmo duc bionic: manic nea ee Ree eee TI

Characters of British Prawns ............s00000000000 = 98

TE COMM EI ISET TALUS eh ya ek Ee oe

Ns EDIE fe Bier Bele aeeiar oie cian Gate a boo aie Spoe sata eum Bord

ay QUSPETSUS icc) deck ae eee ee LOO

OUT LR OS He Sogn soscccoon one coocabcosedeneccaesjons |. Was

5 Bs (Qnionl hing) eee eee 3

IPAVEMONELES) UAT ICIS eee eRe eee eeEEE Eero oer 114

Modification of Pleopods for Egg-bearing............ 120

Palemonetes varians var. mesogenitor ............... 121

The European Prawns of the genus Leander have been very

thoroughly revised by De Man*, who has shown that Z. longi-

rostris occurs commonly on the Dutch coasts, where it has hitherto

been recorded under the name of ZL. squilla. The two species

closely resemble one another in respect of the form of the rostrum,

but differ greatly in other respects, and De Man’s very careful

and detailed description has made discrimination of the species

an easy matter.

In Norfolk a prawn has long been known to occur in the

lower reaches of the rivers flowing into Breydon Water, and has

been recorded by Mr. A. H. Patterson ¢ and myself { under the

name of L. sqguilla. Waving obtained a number of specimens of

this prawn from Breydon Water in June 1921, a careful examina-

tion of these specimens was made, with the result that it became

perfectly clear that they could not be referred to any species

hitherto recorded as British. Subsequent reference to De Man’s

paper at once proved them to belong to Leander longirostris

M. Edw. An examination of old material and of specimens

since collected in the Norfolk rivers and at various points on the

coast has shown that LZ. squilla does not normally occur at all in

any part of the rivers, but that it is quite common between tide-

marks on the coast.

T have also visited the estuaries of the Kast Coast from Nortolk

to the Thames, and have not been able to obtain any evidence of

the occurrence of ZL. longirostris anywhere except in Breydon

Water and the rivers entering it.

* De Man, Tijdschr. Nederl. Dierk. Vereen. (2) xiv. p. 115 (1915-16).

+ Zoologist, (4) ii. p. 178 (1898).

{ Trans. Nort. & Nor. Nat. Soc. vil. p. 637 (1904).

Proc. Zoou. Soc.—1923, No. VII,

98 MR. R. GURNEY ON LEANDER LONGIROSTRIS

It may be of interest not only to give some account of the dis-

tribution and habits of Z. longirostris in Norfolk, but also to add

some notes on the remaining four species of British Prawns.

The structural specific differences have been dealt with so

thoroughly by De Man that it is not necessary to repeat them in

great detail, but I have added some account of the colour of

living specimens, since the species may readily be distinguished

when alive by colour alone, and this is a character to which

sufficient importance has not been attached. Some account is

also given of the range of variation of certain characters which

are regarded as of specific importance.

In the following table, measurements are given of typical

examples of the five British species. For purposes of comparison

the absolute measurements of the antennule and second leg have

been converted into percentages of the peduncle and dactylus

respectively. There is much individual variation, and the pro-

portions of these parts are very different in immature specimens,

so that such selected examples can only be taken as a general

guide; but they illustrate very well the characters of the differ-

ences generally to be observed.

TABLE I,

Measurements of typical individuals of the British species of

Prawn. The measurements for 2nd leg and antennule are

converted to percentages of the dactylus and peduncle

respectively.

2xnp Lze. Ist ANTENNA.

By a i

2 | a)

By iilie ¢| 58/8] >

: a) | te 5 5 E as) ee i}

Locality. Demy eG Bl | ¢ a g = se is ze

=) Wis | cen rete || ees | ee Mrs eee lives ies

a SlAlolre|ola | a | ew A |<

LL serratus § ...| Burnham. | 86 mm. i 100; 191} 91) 100) 133 100 | 83 | 16] 66

L. squilla 2 ...|Thornham. Bimes : 100 | 308 | 203 | 290| 258 || 100) 106 | 47 | 57

| '

L, adspersus .... Poole. | 68, : |100! 229/129] 195|183//100| 122] 39 83

LZ. longirostris.... Breydon.| 69 ,, i 100 | 266 | 166 | 246 | 233 || 100 65 | 17 48

\ |

P.varians ...... Cley. 43 ,, - 100 | 300 | 200 | 410 236 | 100 80 | 58 | 21

SIP atc yD 38, is | L00 253 | 153 | 286 244) 100} 107 | 77 30

AND OTHER BRITISH PRAWNS. 99

Key for determination of the Species.

1. Carpus of 2nd leg much shorter than merus-..................... DL. serratus.

Carpusionger than: mreruis us. cee ee eee ree en ee ee 2.

2. Carpus of 2nd leg longer than chela_ ..................... Palemonetes varians.

Canpusyshoxrtersuhantcliel ar: see: ote een eeraa nen eee

3. Dactylus of 2nd leg one-third the length of the chela ......... L, squilla.

Dactylus nearly or quite one-half length of chela ............... 4.

4. Short flagellum of antennule as long as or longer than _

peduncle SR er eMC ee nerves on od abereece On LL. adspersus.

This flagellum much shorter than peduncle ...................... Li. longirostris .

1. LEANDER SERRATUS (Pennant).

Colour.—Thorax and abdomen strikingly banded with brownish

red, the lines on the thorax running almost horizontally, or

obliquely forwards and upwards. Rostrum covered with small

red chromatophores. Legs banded with purple and yellow.

Rostrum slender and greatly exceeding the length of the

antennal scales. It is conspicuously upturned and devoid of

spines in its distal third. The apex is bifid, and the usual number

of spines is 7 dorsally and 5 on the ventral margin. The first

spine is situated well behind the eye, and the second either above

or slightly behind it.

De Man (ibid. p. 169) has described three specimens in which

the rostrum was of abnormal form, and I have had the opportunity

of examining two specimens in the museum of the Marine

Laboratory at Plymouth which are of some interest. In both

these cases (measuring 57 and 54 mm.) the rostrum is straight,

without the upturned toothless portion characteristic of Z. serratus,

the apex undivided. The spine-formula is s and q.

In both cases the general resemblance of the rostrum to that

of L. squilla was rather striking, and they also agreed more with

L. squilla in having the short flagellum of the antennule approx-

imately equal in length to the peduncle. On the other hand, the

form of the second leg, and in one case the palp of the mandible

(that of the other was not seen), left no doubt that these were two

abnormal examples of Z. serratus.

In quite young specimens up to about 25 mm. the rostrum

alone is by no means a safe guide to identity. In the autumn on

the South Coast young JZ. squilla and L, serratus are found

commonly in rock-pools mingled together, and their separation is

not altogether easy. In such specimens the length of the dactylus

of the second leg is also an unreliable character, since this joint

in specimens of ZL. sguilla up to 15 mm. may nearly equal the

length of the palm, and the adult form is only gradually assumed.

On the other hand, the antennule provides, as I believe, a safe

means of separating the two species. In such small specimens of

L. squilla the free part of the shorter ramus is much shorter than

100 MR. R. GURNEY ON LEANDER LONGIROSTRIS

the fused* part, whereas in ZL. serratus it is considerably

longer.

Antennule.—The short flagellum is about one-seventh shorter

than the peduncle, and the fused part one-quarter or one-fifth of

its total length.

Text-figure 1.

Rostrum.

A. Leander longirostris. B, C. ZL. longirostris with unusual number of

teeth. D. Palemonetes varians. E. LZ. squilla. F. L. adspersus.

G. L. serratus (adult). H. ZL. serratus (young, 15 mm. long).

Second Leg.—Dactylus nearly half the length of the chela;

carpus shorter than the chela or the merus.

Distribution.—This, the so-called ‘“‘Common” Prawn, is a

littoral species, preferring, but not confined to, weedy and rocky

ground. It is abundant on the south coast of England and on

* The inner, shorter, branch of tho outer flagellum is usually described as being

fused to the outer branch for part of its length, and the terms “fused ” and “ free ”

part are convenient and intelligible. But the “free” part is clearly shown in

development to be a secondary or accessory outgrowth of the basal, sensory, part

of the flagellum. The basal part, plus the accessory flagellum, is here spoken of as

the “shorter flagellum” simply for convenience of description,

AND OTHER BRITISH PRAWNS. 101

some parts of the Irish coast, but it is by no means common on

the Hast Coast north of the Thames. Murie* has given an

excellent account of the distribution of this prawn, to which

T can add little. The shrimpers from Southend and from

Burnham-on-Crouch bring in fair numbers of them, but at West

Mersea they are much more rare. I have been to sea with one

of these shrimpers and saw only one prawn in a catch of 12

gallons of ‘‘ Pink Shrimps” (Pandalus montagui). At Harwich

also the prawns brought in are so few as to be hardly worth the

trouble of separating them from the shrimps. They are said to

be taken at times in some numbers in the Orwell and. Deben, and

I have myself taken them as far up the Deben as Woodbridge.

At Aldeburgh the species is so rare that a fisherman who took

one among his shrimps in 1921 had never seen one before! Off

Yarmouth the capture of prawns is exceptional, though a few are

sometimes taken on the sandy ground close inshore, and J have

myself seen specimens taken on Breydon Water. At Lynn it

appears to be almost unknown.

L. serratus has been recorded from Oresund (Denmark), and is

found on the coasts of Holland, Belgium, France, and the Channel

Islands. In the Mediterranean it occurs in “prodigious

quantities” on the coast of Algeria (Lucas), and inhabits the

shores of Italy, Greece, and the Bosphorus.

It is therefore a southern species, which is only a straggler in

the North Sea.

Breeding-period.

Whereas the other species of British prawns breed during a

well-defined period in summer, the breeding-period of L. serratus

seems to extend through winter and to continue till midsummer.

In the List of the Plymouth Marine Invertebrate Fauna? it is

recorded as breeding from November to June, but egg-bearing

females may still be foundin July. I have little personal acquaint-

ance with this species, since it is so rare off the Norfolk coast,

but it seems to me that the few published records indicate that ‘

LL. serratus may prove to have a breeding-bahit somewhat similar

to that of Crangon vulgaris. In this species Hhrenbaum ¢ found

two main periods of egg-laying—namely, A pril—J une and October—

November. The autumn-laid eggs took 4 to 5 months to develop,

and hatched from February to April, while those Jaid in summer

hatched in about 4 weeks—.e., from May to August.

Larve of Z. serratus are found in very small numbers in the

plankton from December onwards.

Owing to the difference in the breeding period, any Leander

larve found off British coasts from December to nearly the end

of June may confidently be assigned to ZL. serratus.

* Report on the Sea Fisheries and Fishing Industries of the Thames Kstuary,

p. 247. London, 1903.

+ Journ. M. B. A. vii. (1904).

£ Mitth. der Sekt. fiir Kusten und Hochsectischerci, Jg. 1890.

102 MR. R. GURNEY ON LEANDER LONGIROSTRIS

2. LEANDER sQuiLua (Linn.).

Colour.—Both thorax and abdominal’ segments bear dark

yellow-brown bands, which are usually very conspicuous, and are

retained for a long time even in specimens preserved in formol,

though rapidly disappearing in spirit. The rostrum is sometimes

quite colourless, but) generally small red chromatophores are

scattered over it or arranged in a median row. Kemp states

that the rostrum is without chromatophores, but this is excep-

tional in my experience of Norfolk specimens. The eye-stalks

and peduncles of the antennules are deeply pigmented with

purple-brown, and the same is the case with the basipodite and

ischium of the third, and sometimes of other, legs. The joints

of the legs are marked by bands of yellow pigment, and the palm

of the chela of the second legs is bright blue. The intensity of

the colour seems to vary to some extent with locality and season.

In the summer of 1921 all the prawns, of all ages, taken in

Wells Harbour, were brilliantly coloured as described above, the

blue of the chele being particularly conspicuous. But others

taken from rock-pools at Whitsand Bay in the spring of 1922,

though showing the same distribution of colour, were by no

means conspicuously banded. They could, however, be im-

mediately distinguished from JZ. serratus of the same size by

their darker colour. Again, the colouring of the prawns in

Wells Harbour during the summer of 1922 was far from being

so pronounced as in the previous year,and many, particularly the

males, were found to be almost colourless. The blue colour which

was so striking a feature of the chele in 1921 was seldom

brilliant, and often absent, in 1922.

Length. Mate 28-50mm. Female 30-63 mm.

The great range in size is due to the fact that maturity 1s

reached in the first year at an average size of about 40 mm. for

females and 30 mm. for males. Females over 50mm. may be

assumed to be two years old, and it is probable that those of

60 mm. and more are in their third year.

Rostrum broad, very slightly upeurved, armed dorsally with

7-9 teeth, two of which are placed behind the eye, and the third

above or slightly behind it. A minute apical tooth is almost

invariably present in addition. Ventral teeth usually three.

The number of these teeth varies within very narrow limits.

For 114 females from Wells, in Norfolk, the number of teeth

was as follews :—

Dorsal teeth : 9 8 7 6

No. of individuals: {2 65 36 1

10:5 %/, 57%, 31:5 %/) 86 9/)

Ventral teeth : 4 3 2

No. of individuals : il 112 1

"86 9/) 98°2 %/, "86 9/)

AND OTHER BRITISH PRAWNS. 103

Adding 62 specimens from various localities on the East Coast

south of Norfolk the frequency is somewhat changed :—

Dorsal teeth : 9 8 7 6

Individuals : 22, 101 52 1

12°59, 57°49/) 29°5 9/5 GUT

There seems to be a somewhat higher frequency for 9 dorsal

teeth than there is in Norfolk, but the number examined is not

sufficient for a definite conclusion. De Man’s figures for 106

specimens from various localities are as follows :—

Dorsal teeth : 9 8 7

Individuals : 20%) 66 °/) 13 %/,

In the typical form of ZL. squilla from Scandinavia, De Man

found seven dorsal teeth in 31 per cent. of specimens and nine

teeth in 18 per cent.

Such figures as these seem to indicate a definite local variation

in respect of this character, but in my opinion the material

examined is only sufficient to indicate a probability that such

variation occurs.

Mandible palp—L. squilla differs from all other European

species of Leander in having the mandible palp two-jointed.

This difference was first pointed out by Dr. W. T. Calman%*,

but the palp was correctly figured by Ortmann in 19017,

though not alluded to in his definition of the genus Leander.

This character cannot be used in the determination of young

specimens 20 mm. or less, since, in L. longirostris at all

events, the mandible palp is still often two-jointed at that

size.

Antennule.—The short flagellum is approximately equal in

length to the peduncle, but may be either shorter or longer

than it. This flagellum exceeds the length of the peduncle

more frequently in the male than in the female. ‘The free

part generally exceeds the fused part in the proportion of

5 to 4.

Second Leg.—The second leg reaches, when extended, beyond

the antennal scale by the whole chela or even by part of the

carpus as well. The dactylus is conspicuously shorter than in

the other species, being usually about one-third the length of the

whole chela. The carpus nearly always slightly exceeds the

length of the merus.

De Man, as the result of the examination of large numbers of

* See Kemp, “ The Decapoda Natantia of the coasts of Ireland.” Fisheries, Ireland,

Sci. Invest. i. 1908, p. 127 (1910).

‘+ “Die Klassen und Ordnungen der Arthropoden.” Abth. Crustacea Malacostraca,

Taf, Ixxiv. fig. 2 e (1901).

104 MR. R. GURNEY ON LEANDER LONGIROSTRIS

L. squilla from many localities, has come to the conclusion that

three varieties or geographical races should be separated :—

(1) L. squilla, typical form.

Scandinavia and the Baltic Sea.

(2) L. squilla var. intermedia De Man.

Holland, British, and probably French coasts.

(3) L. squilla var. elegans Rathke.

Mediterranean, Black Sea, Azores, Madeira, Canaries,

Cape Verde Islands.

These varieties are separated by very slight differences, the

most important and constant of which relate to the antennule—

thus :—

(1) Fused part of short flagellum a little shorter or a little

longer than the free part, rarely equal to it.

Typical form.

(2) Fused part distinctly shorter than free part.

var. intermedia.

(3) Fused part distinctly longer than free part.

var. elegans.

Norfolk specimens agree with the description of the variety

intermedia, but it is possible, as De Man suggests, that L. squilla

from Scotland may prove to belong to the typical northern race.

I have not had the opportunity of examining specimens from the

east coast of Scotland, but a male received from Millport on the

west undoubtedly belonged to the var. intermedia, since the free

part of the flagellum exceeded the fused part in the proportion

of 5:4.

Distribution in Britain.—L. squilla appears to be distributed

all round the coasts of England, Ireland, and Scotland, even as

far as the Shetlands. It is a littoral species living between tide-

marks, and I have found it to be abundant all along the Norfolk

coast from Hunstanton to Cley. At Wells it can be caught in

quantities by working a hand-net along the wooden quay-heading

at high tide. The pools on the salt marshes at Wells are tenanted

for the most part by Palemonetes varians, but some were found

in August to contain LZ. squilla in addition, At Thornham

numbers were taken on the woodwork of a sluice, and I have

found that such sluices, where a pool of water remains even at

low tide, are favourite resorts for this species. In such situations

it is quite commonly associated with P. varians, and it runs far

up the East Coast estuaries, but it appears to be none the less

intolerant of fresh water, and to abandon a sluice when a large

quantity of fresh water is being discharged. It does not nor-

mally occur on Breydon or in any part of the rivers Yare, Bure,

or Waveney. A single small specimen was, however, taken in

AND OTHER BRITISH PRAWNS. 105

1921 by Mr. O. Hunt at Acle at a time of exceptionally high

tides. I have taken it myself at the following places :—

R. Crouch at Burnham and Battlebridge.

Blackwater at Maldon.

Mersea Island.

R. Stour at Harwich and Wrabness.

R. Deben at Woodbridge.

I was unable to find it in the Alde between Aldeburgh and

Tken, or in the Orwell at Pinmill.

Breeding-period.

In Norfolk the first eggs are laid about the end of May or

beginning of June. One female, taken on June 1 witn eggs

apparently very recently laid, hatched her young on July 5. the

eggs having therefore been borne 35-40 days. This result is in

general agreement with Mr. Hlmhirst’s figures*. Mr. Elmhirst

kept observation on L. squilla in rock- pools at Millport, and

found the period of development to depend on temperature as

follows :—

Period of development. Average temperature.

LOOT ee 30 days 23° C.

BOM LW Te ASR ai cetthers 40 days 14:°5° C.

OBZ) os: cece. 56 days ENC.

Development may therefore be taken as requiring about 6 weeks

under usual conditions.

Breeding continues actively through June and July. In 1921

it ceased about the middle of August, but in 1922 it was con-

tinued into the first week of September—a period of about

100 days. Hach breeding female seems to produce two broods

in the season. In July females with eggs in an advanced stage of

development always have the ovary distended with eggs of the

second brood, while in August the ovary is usually empty. The

production of two broods in the year was established by

Mortensen? for Z. adspersus and by Khrenbaum for Crangon vul-

garis, but Mortensen found that it only applied to the larger

prawns. In Norfolk, on the other hand, not only does practically

every female breed down to a size of about 30 mm., but the

majority,sat all events, produce second broods. In July the

population of females may be separated into two groups—namely,

a small number of large prawns from 63 mm. to about 48 mm.,

and the remainder of smaller prawns among which sizes of

36-39 mm. are the most frequent. These gr groups no doubt

comprise prawns of two years’ and one year’s growth. The

former on July 25 for the most part bore eggs in early stages

having hatched their first brood, while a large proportion of the

* Scottish Mar. Biol. Assoc. Ann. Rep. 1921, p. 7.

+ Vid. Undersog. paa Fiskeriernes omraade udgivne af Dansk Fiskeriforening,

i. (1897).

106 MR. R. GURNEY ON LEANDER LONGIROSTRIS

one-year group carried eggs approaching hatching or showed

signs of having recently hatched young. It is probable, therefore,

that the older prawns spawn before the younger ones.

The period of larval development has been ascertained by

Mortensen to be about 4 weeks in Z. adspersus, and is probably

much the same for Z. sguilla. I have not been able to keep the

larve through more than one moult, so have no direct evidence

to offer. No post-larval prawns are to be found at Wells in July,

but about the middle of August they begin to appear, and become

abundant both in the marsh-pools and in the fucus growing on the

woodwork of the quay. In 1922, young did not become abundant

till the middle of September, but some then measured 22 mm.

and must have been in the harbour for some time. For the

most part the smallest young taken measure about 12mm.

Since the young in the first and second post-larval stages do not

exceed 9 mm.,it seems that metamorphosis occurs out at sea, and

that the young do not usually reach the shore till after three or

four moults. A very small proportion of the young prawns

found in September 1922 were either in the first or second

post-larval stages,and these must certainly have been brought in

by the flood-tide and have metamorphosed on the spot. On the

other hand, in spite of much search, I have only taken one larva

in Wells Harbour, and there can be no doubt that the whole

larval life is normally spent out at sea. ‘This is in agreement

with Mortensen’s conclusions with regard to L. adspersus.

The proportion of adult’ males to females was only noted

accurately on two occasions—in Wells Harbour on July 25,

1922, and in Blakeney Harbour on August 10. In the former

case 37 males were found among 114 females, while in the latter

the males exceeded the females, the numbers being 69 males and

56 females.

3. LEANDER ADSPERSUS Rathke.

Palemon rectirostris Zaddach, 1844.

Palemon leachit Bell, 1853, p. 307.

Leander adspersus var. fabricit De Man, 1916.

Colour.— Unlike the other British species of Leander,

L. adspersus has no bands of colour on either thorax or abdomen,

but the body appears of a uniform yellowish grey, due to small

black or reddish-black chromatophores scattered irregularly. The

rostrum is covered with chromatophores, which are concentrated

on the lower half in the form of red or sometimes purplish-red

blotches, which, as Kemp has pointed out, provides a conspicuous

feature by which the species may be recognized ata glance. The

long flagella and the peduncle of the antennule are also very red

and the legs banded with yellow, but without the blue on the

chela, which is so striking in Z. squilla.

Rostrum.—The rostrum usually extends well beyond the

antennal scales, is nearly straight, not very deep, and provided as

a rule with 6 spines above and 3 below in addition to the small

terminal spine. Only one of the dorsal spines is situated behind

AND OTHER BRITISH PRAWNS. 107

the eye, the second usually slightly in front of the orbital notch.

The dorsal teeth differ slightly from those of L. longirostris and

L. squilla in being more depressed.

Mandible palp.—Three-jointed, the second joint about one-third

the length of the last joint.

Antennule—The shorter ramus exceeds the length of the

peduncle, and is ‘‘fused” to the longer flagellum by only about

one-third of its length. The length of the free part is a very

noticeable character of the species. There are certain minor

Text-figure 2.

Bia:

ict

Ti ee

Speco facie

= ae

MME

Antennule of female.

A. ZL. longirostris, drawn from moulted skin.

B. L. longirostris. C. L. adspersus. D. L. squilla. E. P. varians.

differences in the structure of the peduncle between the species of

Leander and Palemonetes. These relate to the proportional

length of the joints and the form of the stylocerite and terminal

plate, but they are less easily described than illustrated. In text-

fig. 2 the antennules of the different species are drawn side by

side to such scales that the peduncle is represented as of the same

length in each case. The proportions of the joints and of the

flagella are in this way made clear.

Second leg. — Hxtends beyond the antennal scales by the

108 MR. R. GURNEY ON LEANDER LONGIROSTRIS

dactylus and part of the palm. The dactylus is long and slender,

more than three-quarters of the length of the palm. The carpus

usually considerably exceeds the length of the merus, but may be

of the same length.

De Man has separated this species into two forms :—

L. adspersus (Rathke). Black Sea.

L. adspersus var. fabricii (Rathke). Scandinavia, Baltic,

Denmark, France, Adriatic, Mediterranean, British

Isles.

The chief differences are as follows :—

Rostrum usually with 4 ventral teeth; shorter ramus of anten-

nule usually projecting by 4 to 4 its length beyond rostrum.

L. adspersus.

Rostrum usually with 3 ventral teeth; shorter ramus projecting

usually by more than 4 its length beyond rostrum.

L. adspersus var. fabricii.

Distribution in British Isles.—Bell described the species under

the name of P. leachii from specimens taken in Poole Harbour.

Mr. J. Omer Cooper has kindly sent me a collection of prawns

from this estuary, which proved to be made up as follows :—

DG NSCRBATUS. See It. eRe aot ee 528

DL SAASPETSUS Worcs cote nade saee pis. 9

LA SQULUGY BSR eed cectik tants o es 5

It is evident that Z. adspersus is by no means an abundant

species. Mr. Kemp has recorded it from two localities in Co.

Galway, in one of which it occurs in company with LZ. squilla and

L. serratus as it does.at Poole, and he notes that it has been taken

also at Weymouth and in the Thames estuary. I have myself

taken two small specimens from between tide-marks at Burnham-

on-Crouch, and have had others sent to me from West Mersea,

where it is known as the ‘‘ Mud Prawn,” and is taken in some

numbers by eel-catchers. It is an estuarine species, preferring

a muddy bottom, but Mersea seems to be about its northern limit,

as I have not found it in the Stour or the Orwell, and it certainly

does not occur in Breydon Water in Norfolk, where conditions

would seem to be favourable.

4. LEANDER LonGiRostRis Milne Edwards. (Text-fig. 3.)

P. longirostris M. Edwards, Hist. Nat. des Crustacés, ii. 1837,

p. 392.

P. edwardsi Heller, 1863.

LL. tongirostris De Man, 1916, p. 149.

There has been some confusion in the use of the name

L. longirostris, since not only did Milne Edwards describe two

distinct species under the same name, but, by a misplacement of

a footnote reference, even the authority of the name was wrongly

referred to Say. Milne Edwards himself corrected his second

AND OTHER BRITISH PRAWNS. 109

P. longirostris to P. styliferus, which applies to an Indian prawn,

but Miss Rathbun is undoubtedly right* in maintaining that

the name L. edwardsi Heller must give place to L. longirostris

of M. Edwards.

Colour.—The colour is rather variable, but the majority are,

Breydon, 21. 7. 21.

Text-figure 3.

Leander longirostris M. Edw., 2.

in life, almost colourless, and of an opaque white immediately

after death. A close examination shows that the whole body,

including the rostrum, is speckled with small red chromatophores.

In certain areas these chromatophores are surrounded by a halo

of blue pigment, which may sometimes be greatly developed and

* Proc. U.S. Nat, Mus. xxvi. p. 50 (1903).

110 MR. R. GURNEY ON LEANDER LONGIROSTRIS

entirely obscure the red. In such eases, which are rare, the

animal appears of a very dark purple-black colour. Occasionally

the red pigment alone is present, the blue being suppressed, such

specimens appearing of a beautiful rosy colour, the margins of

the abdominal segments being more deeply coloured. An

unusually large female (75 mm.) of this type was taken in the

River Bure in July 1922, and has lived in a fresh-water

aquarium for over three months without losing its red colouring.

There is a tendency for the chromatophores to become arranged in

more or less conspicuous lines and patches (text-fig. 3), but these

do not give the appearance of distinct bands as in ZL. squilla and

L. serratus. 'The limbs are usually colourless, except for a few

scattered red chromatophores, but there is sometimes a faint blue

colour on the chele.

Length.—Female (bearing eggs) 50 to 77 mm. Male 35 to

77 mm. © Usually much smaller than the female.

Rostrum.—The rostrum projects considerably beyond the

antennal scales, and is deep and nearly straight, but commonly

slightly upcurved. It is usually armed with 8 dorsal and 4

ventral teeth, the dorsal teeth being rather prominent, as in

L. squilla. The first two teeth are situated behind the eye, the

third just in front of the orbital notch. As has been pointed

out by De Man, it is particularly characteristic of this species

that the space between the first two teeth is one-and-a-half times

as great as that between the second and third. The following

figures show the variation in the number of rostral teeth in 191

females from Norfolk :—

Dorsal teeth : 10 9 8 7

Individuals: 2 18 108 61

1:04 9/, 9°49/, 513 %/, 3179/97,

Ventral teeth: 6 5 4 3

Individuals: 2 6 167 16

1:049/, 3149, 87:49, 84/,

In Norfolk, therefore, the usual formula is = but De Man

found? the usual formula for Dutch specimens to be mt and

that the dorsal teeth varied from 12 to 6. I have excluded

males from my table, but have no reason to believe that they

differ in this respect from females. The number of teeth does

not increase with age or size, and the smaller size of the male is

not of itself likely to reduce the average number of rostral teeth.

I have not in all cases noted the number of small apical teeth,

but in 46 specimens only 3 had two of these teeth, whereas

De Man found this number in 42 per cent. The difference may

perhaps be explained on the assumption that the proximal

apical tooth, when present in Norfolk specimens, is larger and

less separated from the others, and has therefore been counted in

AND OTHER BRITISH PRAWNS. LY

with them. On the other hand, the general frequency of occur-

rence of the various number of teeth differs so greatly that

T am of opinion that a real local difference is shown. De Man’s

figures are as follows :—

Dorsal teeth: 10 9 8 f( 6

Individuals: 8 °/) 6-6 %/p 37% D2) Of * ded! 8/6

Ventral teeth : 6 5 4 3 fi

Individuals: 1°06: 9/6 i eae ige ian Git ig on 29/91,

Mandible palp.—Three-jointed, the second joint about half as

long as the third in the adult, but only one-third of it in young

specimens.

Antennule.—The short flagellum is two-thirds, rarely a little

more, of the length of the peduncle, and is fused to the longer

flagellum by about one-third of its length. Taking the average

of seven measurements, and regarding the total length as 100,

the result is as follows:—Fused part 31; free part 69; the free

part being therefore about twice the length of the fused part.

Second leg.—The second leg reaches beyond the antennal scale

by the whole of the chela and usually about one-third of the

carpus. The dactylus is usually nearly one-third of the length

of the whole chela, but the proportion is very variable, and it

may sometimes be nearly equal to the palm. The chela of this

leg does not therefore provide means for distinguishing this

species from L. adspersus. The chela exceeds the length of the

carpus by about one-tenth, and the carpus is usually slightly

longer than the merus. Carpus and merus are, however, so

nearly equal that very careful measurement is necessary to

determine which is the longer. An average of 20 measurements

gave the figures :—Carpus 101; merus 100!

Distribution.—L. longirosiris has been recorded from Liberia

(Rathbun), Corsica, and the French coast at Noirmoutier. It

has also been found in the River Gironde as far up as Bordeaux,

and in the Loire up to Vertou (30 miles). In the British Museum

there are specimens from near Seville, about 50 miles up the

River Guadalquivir. De Man found it to be common at certain

points on the Dutch coast. In the estuary of the Meuse it is

abundant in the Hollandsch Diep, and has been taken as far up

as Werkendam, which is just above the Biesbosch. It occurs

also in the Zuider Zee, the Ij, im the Rhine near Katwijk, and

in the Scheldt up to Antwerp. It therefore inhabits estuarine

regions, but within the range of sea-water. De Man only records

L squilla from the outer Scheldt on the Zeeland coasts, so that it

seems to be replaced in Holland by L. longirostris.

In Norfolk this prawn is abundant in Oulton Broad, and in

Breydon Water at certain times, and it is known to local fisher-

men as the “Jack Shrimp” or “‘ White Prawn.” In Oulton it is

‘used as bait for perch-fishing, but not for food. In Breydon it is

said to be more numerous when there is much fresh water passing

112 MR. R. GURNEY ON LEANDER LONGIROSTRIS

down, but it is found in abundance, at all events in the breeding-

season, even when the water is entirely salt. On the other hand,

the numbers are generally found to increase towards the upper

end of the estuary. It is never taken at sea by the shrimpers,

though it is said to be caught occasionally just at the mouth of

Yarmouth Harbour. Mr. Patterson has sent me specimens taken

at St. Olaves on the Waveney, and probably its distribution is

continuous from Yarmouth to Oulton Broad via the Waveney.

No doubt it also oecurs in the Yare, at least as far as Reedham,

but I have not been able to search for it there. In the Bure it is

common, probably at all states of the tide, as far up as Acle,

14 miles from the sea, and I have even taken it in Heigham

Sounds, 22 miles from the sea. It is well known to the eel-

catchers, and numbers are taken in the eel-nets at certain times.

At South Walsham eel-set they are usually found in the net after

there have been high salt tides in the river and the salt water is

beginning to run down again. At such times they are also

commonly taken in an eel-set near Hickling Broad.

To judge from its distribution in Norfolk, Z. longirosiris is

essentially a river prawn capable of existing either in salt or in

fresh water, but preferring that part of the river in which the

water is generally brackish. Their indifference to salinity is such

that they may be repeatedly transferred directly from fresh water

to salt and vice versa without any apparent ill-effects, and I have

at this moment a number which, taken originally in salt water,

have been living for months in pure fresh water. One of these,

after hatching her young in salt water in June 1921, was trans-

ferred to a fresh-water aquarium in which she moulted 10 days

later, and is still (November 1922) flourishing. This indifference

to salinity can be equalled by very few animals and surpassed by

none.

Breeding.

The fact that, in order to catch ZL. longirostris with certainty

and in any numbers, it is necessary to use a trawl either in the

lower reaches of the rivers or in Breydon Water, has rather

restricted my opportunities for observation, and I have not been

able to get representative collections throughout the year. It

is, however, certain that breeding begins at the end of May or

beginning of June, as it does in L. squilla, and that two broods

are hatched in the season. A number of specimens taken far up

the River Bure on May 16, had ripening ovaries, but no eggs

had been laid, but on June 15, 1922, out of 98 females taken

in Breydon Water, 48 bore eggs in various stages, 42 had hatched

put had not moulted, while 8 had hatched and moulted, and were

ready to lay another lot of eggs. All, except one with freshly-

laid eggs, had the ovary full. As development probably takes at:

least a mouth, and the moult following hatching occurs usually

4 or 5 days later, the first eggs must have been laid this year

about May 10. No doubt the larger two-year-old prawns

spawn about the middle of May, and are followed towards the

AND OTHER BRITISH PRAWNS. 13

end of the month or in June by the younger females. During

June and the first half of July every female taken either bears

eggs or can be shown to have hatched young; but about the

middle of July, though a large proportion bear eggs, many have

evidently ceased to breed. For example, out of 103 specimens

taken on July 14 there were 33 with eggs, 46 without eggs, and

24 males. In all the females the ovary was empty.

Egg-bearing females migrate down into salt water as the time

of hatching approaches, and, so far as my observations go, the

larvee are very rarely hatched in the river itself. While I have

had the larve hatch in aquaria in salt water on several occasions,

they have never done so when the parent has been kept in fresh

or slightly brackish water. In such cases the eggs are eventually

stripped off. In more than one case a female kept in fresh water

past the time at which the young were expected to hatch has

been put directly into salt water, with the result that the young

have hatched during the following night. Probably hatching

takes place always at night under normal conditions, and the

young are carried out to sea by the ebb-tide. I have only once

caught a single larva in an early stage of development, in spite of

much search in Breydon Water at the height of the breeding-

season. A few larvee in the last stage are occasionally found in

Breydon, and doubtless metamorphose there, but the great

majority must complete their development at sea and migrate up

the rivers in the post-larval condition. Young prawns have been

taken 12 miles up the river towards the end of August between

19 and 25 mm. long, but it is by no means easy to obtain these

young stages, and it is probable that they are to be found in

Breydon much earlier, On the other hand, a very careful search

for them on August 28, 1922, in the shallows and among

Zostera, was entirely unsuccessful, so that it is not improbable

the immigration is delayed to a later stage than is the case in

L. squilla. Mortensen found that the young of Z. adspersus,

though appearing in the shallows about the middle of July, do

not reach the innermost parts of certain fjords during the first

summer.

Moulting.

The process of moulting takes place usually at night, but I

have been fortunate in having been able to witness it on two

occasions during the daytime. On each occasion the prawn was

found in a peculiar position, the body greatly flexed, the head

bent sharply downwards. The cuticle breaks between the thorax

and abdomen, leaving the anterior sclerite of the latter attached

to the first segment. The thorax then bulges out through the

opening, and the animal draws the whole of the thorax and

appendages out evenly, without pause. Immediately after freeing

the eyes and antenne, the animal gives a sudden leap forwards,

freeing the abdomen instantaneously. The whole process took a

Proc, Zoou. Soc,— 1923, No, VIII, 8

114 MR. R. GURNEY ON LEANDER LONGIROSTRIS

surprisingly short space of time, probably not more than half a

minute, but no doubt there were preliminary movements which

were not observed. After the moult the movements are most

erratic. The prawn may leap about with violent movements of

the abdomen or lie on one side in apparent discomfort, moving

its appendages continually. Some hours elapse before normal

progression is resumed. The very erratic movements immediately

after the moult are very likely due in part te the fact that the

otocyst is empty, and I was not able to see the process of inserting

new grains of sand. The cast skin is often eaten, at least in part,

while the newly-moulted prawn frequently falls a victima to its

companions.

Breeding females do not, as is commonly the case in Caridea,

moult immediately after hatching of the eggs. Both this species

and also Z. squilla and P. varians moult 4 or 5 days after hatching,

but the moult may-be delayed even longer. For example a female

about 70 mm. long was taken on July 17, the condition of her

pleopods showing that she had recently hatched young; but she

did not moult till August 19, or 33 days later.

Tam unable to say at what intervals adult prawns normally

moult, since this can only be ascertained by keeping single

individuals for long periods under as nearly as possible natural

conditions. Warrington’s observations on this point are, in my

opinion, quite unreliable, since several prawns were kept in the

same aquarium. The female mentioned above moulted a second

time on October 5, an interval of 47 days, no growth having

taken place. On the other hand, another adult female kept for

over a year in fresh water has only moulted once during the

whole period.

5. PALHMONETES VARIANS (Leach).

Colour.—To the naked eye P. varians is generally almost

colourless and translucent, except for traces of yellow-orange

colour at the end of the abdomen and on the joints of the legs.

Under a lens the whole body is seen to be speckled with small

blackish chromatophores. On the thorax these are generally

arranged in lines, and have a faint yellow halo, while a few pure

yellow chromatophores are scattered among them. The rostrum

is colourless, except for a row of orange-yellow chromatophores

below and of black ones along the middle line. The eye-stalks

and antennules are richly pigmented. The abdomen is speckled

with black and yellow like the thorax, but there is also an orange

spot at the junction of segments 4 and 5, and 5 and 6. The

uropods and telson have orange and black spots, but the pleopods

are colourless. The chele of the second legs have an orange patch

at the base and at the end of the dactylus, while a similar

patch is seen on the merus of the remaining legs. Barrois* has

* Bull. Soc. Zool. France, xi. (1886).

AND OTHER BRITISH PRAWNS. 115

described the colour of this species, and has drawn attention to

the orange colour of the chelz as a striking character.

Length. Female 29-43 mm. Male 18-25 mm.

Mandible—Palp absent.

Rostrum.—The rostrum is narrow and straight, or even may

be somewhat depressed, and it is armed with a variable number

of teeth. I have examined a large number of specimens from

Norfolk and from other localities, and find that the formula 2 is

by far the most usual. A small apical tooth is present in more

than half the individuals. Four or six dorsal teeth are not

uncommon, but it is exceptional to find more or less than two

ventral teeth. Only one tooth is situated on the carapace behind

the eye.

The number of rostral teeth has long been known to be variable,

and this variation has been studied by Weldon *, Brozek + and

others. The latter has made a statistical study of the numbers

of teeth in specimens from various localities in fresh and brackish

water, and concludes that the fresh-water form from south

Europe has on an average a larger number of dorsal teeth than

the brackish northern form. His figures are as follows :—

Elymouth (Weldon)... .seeee one 4°3

Lago di Castello (Italy) ...... be icaisd: 4-9

mee Ubardl (2. 35207.2als A ae een 6:4

Montenegro.) .:. Ge ae 6:3

Monfalcone’ (Istria) ie. iy. sae ee a7

As in a later paper he has given the frequency for 134 specimens

from Copenhagen to have been 6:22, there seems to be no ground

for supposing that the number of teeth has anything to do with

geographical situation or salinity.

It is, however, not improbable that a statistical study of the

rostral teeth based on a large material would show constant

local variations, since many populations of this species must be

isolated for long periods and subject to intense selection. The

result is hardly likely to justify the great labour necessary, but

it may be of interest to summarize such facts as I have collected

bearing on this point (Table II.). The figures given in this

table express the frequency of ocenrrence of various numbers of

dorsal teeth as percentages of the individuals examined. It

should, however, be mentioned that Weldon’s figures alone are

drawn from a really adequate material (915 specimens). So far

as the figures go, they show that six dorsal teeth is far more

frequent on the East Coast and in Scotland than it is at

Plymouth, though the examples from the Stour are an apparent

exception. The range of variation is also less.

Antennule.—The shorter flagellum is about four-fifths, or

* Journ. M. B. A. n.s. i. p. 459 (1890).

+ SB. K. Bohm. Ges. Wiss. Jg. 1907, 1909, and 1912.

116 MR. R. GURNEY ON LEANDER LONGIROSTRIS

between 70 and 80 per cent. of the length of the peduncle, and

the fused basal part is almost three-quarters of its total length

(between 70 and 75 per cent.).

TABLE II,

Dorsal rostral teeth of P. varians.

Number of teeth.

Locality. Arathor) | ete) 2e5 eS ues NGS Ivan! 18:

Monfaleone, Istria sogricpaue oa)) JESHOVACIEE pie abe ie: “a 31 ‘67 Giga Paty

Ved pines Waroeecescnracsenadenses AMENGLNE|| AA Tess ek |) sth yl |

Forth Estuary, Scotland 2 Evans.| 5/ 9/| 2] 9 | 42/28] 2 | 2

River Bure, Norfolk ......... TR Geol soo |) poof aon | PHO) SS) Peto! af

Wells, Norfolk.................. o weft |e 15 | 51 | 34 |

Maldon, Essex .................. a A teste (OAS a ae |

Stour Estuary, Suffolk ...... 3 sop vob Il) coe. {Ss 1 IS G

}

Second leg —Reaches, when extended, beyond the antennal

scales by the dactylus or part of the dactylus only. The dactylus

is about half the length of the palm. The carpus is very long,

and greatly exceeds the length of both the chela and the merus.

The Telson.—The form and armature of the telson is very

constant throughout the Palemonide, and that of P. varians is

normally armed in precisely the same way as is that of the

British Leanders—namely with two pairs of small spines on the

dorsal surface, two pairs of large terminal spines, and a pair of

feathered sete springing from beneath the median triangular

prolongation of the telson. There are, in addition, one or two

(normally one) small hairs on either side of the projection

dorsally.

This arrangement of spines and sets is very variable in

P. varians from Norfolk. Out of 30 specimens taken in the

River Bure only 17 were entirely normal, three more differed only

in having an additional pair of dorsal sete, and one in having an

additional minute ventral seta. The remaining nine were

strikingly abnormal in respect either of the number of terminal

spines or of ventral feathered setz, as follows :—

One had only a single feathered seta in the middle line.

Four had three of these sete.

One had four, one had five, and one had six feathered setz.

One had three pairs of terminal spines.

The specimen with six sete (text-fig. 4B) is evidently a case of

the retention of the whole of the original seven pairs of larval

setze,

AND OTHER BRITISH PRAWNS. a7

These deviations from the normal are of some interest, since

systematic importance has been attached to the numbers of these

feathered sete, and the presence of more than two of them has

been given as one of the principal characters of the genus <Allo-

caris Sollaud. It is evident that it possesses neither generic nor

specific importance. Pesta* has already shown that Sollaud’s

genus is untenable on this and other grounds. and states that he

has found four setz in the telson of P. varians from Albufera.

Text-figure 4.

Telson.

Palemonetes varians with supernumerary pair of dorsal setze.

B. 55 Bs » four supernumerary ventral sete.

C. P. varians from Algeria,

D. LZ. squilla with three ventral sete.

EH. LZ. serratus with additional spine and four ventral seta.

I possess specimens from Algeria which likewise all have four

sete,

The frequent presence of supernumerary sete in P. varians is no

doubt a primitive character, or perhaps more precisely a retention

of a larval character comparable to the non-development of

a mandible palp.

In the British species of Leander the telson varies very little,

but supernumerary feathered sete are occasionally met with.

* Ann. K.K, Naturh. Hofmus. Wien, xxvii. (1913).

118 MR. R. GURNEY ON LEANDER LONGIROSTRIS

For instance, among 30 specimens of LZ, squilla from Wells one

possessed three of these sete. Among ten ZL. serratus from

Poole one was found not only with four sete, but with an

additional terminal spine (text-fig. 4 H).

Distribution.—As is well known, P. varians inhabits fresh water

in the Mediterranean region and brackish or salt water on the

North Sea and Atlantic coasts, but it is not exclusively a fresh-

water species even in the Mediterranean region, since it is

recorded by Gourret* in salt water in the Etang de Thau and

Etang de Berre on the south coast of France, in company with

strictly marine species such as Leander wiphias.

The distribution has been fully described by Barrois T, but may

be briefly repeated here with some additions.

In fresh water it is recorded from several of the Italian lakes,

Venice, Dalmatia, Montenegro, Scutari, Corfu, Turkey, Egypt,

aud Mesopotamia. According to Pesta, Allocaris sinensis 1s only

a variety of this species, its distribution being thereby extended

to China (Pekin). It has been found in a hot spring at San

Giuliano, near Pisa, at a temperature of 28°75° C. In Tunisia

it is said to be very common in fresh waters in the south

(Gabes, Tozeur, &c.), and has even been found in water at a

temperature of 25° to 27°C. I have taken it myself in a stream

near Biskra in the Algerian Sahara.

In salt or brackish “water it has been found in the Etangs de

Berre and Thau (S. France), the Black Sea, Seine Estuary,

Boulogne and Pas-de-Calais, coasts of Holland and Friesland,

Portugal, Denmark, North Germany and Sweden.

In Britain it is probably widely distributed. It is recorded

from several localities on the coast of Devon, Dorset, and Corn-

wall, and seems to occur all round Ireland. It has also been

taken in the Isle of Wight, on the Northumberland and Durham

coasts, and by the estuaries of the Forth and Clyde in Scotland.

In Norfolk the great stretch of salt marsh between Thornham

and Salthouse is eminently suited to the requirements of

P.varians. ‘These marshes are seamed with muddy creeks which

are empty at low tide, but the marsh between lies for the most

part above the level of spring tides. Dotted over this marsh are

innumerable small pools full of brackish water, and in almost

every one of them these prawns are to be found. Late in autumn

the young of the year swarm in these pools, having reached a size

ef about 20 mm., but in June, when maturity is reached, the

numbers have decreased most strikingly, showing a great mor-

tality during winter and spring. When breeding begins, the

water in some pools may become almost thick with larve, very

few of which probably survive even to the post-larval condition,

since the available food must be rapidly used up. In two pools

at Blakeney examined in June large numbers of larvee were found

together with quantities of the Copepod, Hurytemora lacinulata.

* Ann. Mus. Hist. Nat. Marseille, v. (1897).

+ Bull. Soc. Zool. France, xi. (1886).

AND OTHER BRITISH PRAWNS. 119

When the same pools were visited again in August all the

Eurytemora had disappeared, and the numbers of young Palc-

monetes were far less than of the larve previously present.

The marshes are occasionally flooded by exceptionally high

tides, which may sometimes occur in early summer. In such

cases the young must be washed out to sea and spread about

along the coast. If it were not for such floods, every pool would

become an isolated unit in which a struggle for existence of

extraordinary intensity would operate, and the circumstances

would be most favourable for the establishment of local races.

How far the population of the various pools are in fact isolated

it is difficult to ascertain, but there are pools which are probably

not flooded by any but very exceptional tides such as occur at

intervals of several years. Probably, however, isolation in this

sense rarely, if ever, persists for very long, and the sea carries

the larve up and down the coast, intermingling the various

populations.

Breeding-perrod, ke.

The breeding-period begins, in Norfolk, about the middle or

end of May and ceases about the end of July. Larve were first

met with this year near Yarmouth on June 16, all being in the

first stage and hatched since about four days, as most of them

moulted within the next two or three days. On June 17 zozas,

nearly all in early stages, were found in great abundance at

Wells, so that about the middle of June may be regarded as the

main hatching-season. The records of the Marine Laboratory at

Plymouth show breeding to begin there in April and to continue

till July. The eggs are carried 5 or 6 weeks before hatching, if

I may judge from a single female kept in an aquarium, but pro-

bably both the rate of development of eggs and larve would be

rather more rapid in natural conditions. After July I have not

found egg-bearing females, and larve in early stages are not

met with.

I have not found any direct evidence of females producing a

second brood, as is usually the case with ZL. sqwilla and L. longi-

rostris. Examination of large numbers of young taken in

September bears out the conclusion that second broods are the

exception. That second broods are, however, produced occasionally

there can be no doubt, for in some cases the young are found to

fall distinctly into two age-groups. For instance, in a pool at

Cley on September 6 it was possible to draw a clear distinction

between young of 17-22 mm. representing the first brood, and

others of 11 mm. or less, some of which were still in the second

post-larval stage and must have been hatched about the end

of July.

Probably the production of a second brood depends a good deal

on local conditions, and is likely not to take place in such small

pools, which must be extremely overcrowded, whereas it may be

the rule in other and more favourable conditions.

120 MR. R. GURNEY ON LEANDER LONGIROSTRIS

Modification of the Pleopods for egg-bearing in Leander and

Paleemonetes.

The pleopoda of the British prawns (I have not been able to

examine JL. adspersus in this respect) are subject to special

modification for the purpose of egg-bearing which, so far as I can

find, has not been previously referred to.

Text-figure 5.

Second pleopod of female.

A. L. squilla after moult following hatching of eggs.

B. The same, proximal inner corner more enlarged.

C LZ. squilla. Moulted skin after hatching of eggs.

D. ZL. longirostris. Before laying of eggs.

AND OTHER BRITISH PRAWNS: 121

The condition of the pleopods in the female before and after the

breeding-period is shown in text-fig. 5. The shaft or basipodite

is broad and flattened on its outer edge into a thin flange, which

is either bare or carries a few irregularly-placed set. Along the

inner face of this joint is a series of circular markings which

appear to be spots where the cuticle 1s thin or pierced by a pore.

In the centre of some of them is a short conical projection or

spine, and the arrangement of these ‘spinous discs” is definite

and apparently almost constant.

At the moult immediately preceding the laying of eggs a great

change takes place. A dense row of feathered sete appears along

the outer side of the basipodite and on its posterior surface, being

slightly overhung by the thin outer flange. At the same time,

on the inner edge of the joint the place of the “ spinous discs ” is

taken by a series of long, smooth sete, while a variable number of

shorter sete spring apparently from the remaining circular spots.

These long sete are not the same in number in each pair of legs,

but have a remarkably regular arrangement. They may be

described as forming three groups as numbered in the figure,

while an additional seta may be present on the anterior face in a

position corresponding to the letter ‘‘ A” in the figure, and may

be regarded as accessory to Group 3. The following arrangement

of these sete was found in a specimen of L. squilla, and may be

taken as typical for all the species :—

Pleopod.

Ie ES ES EVE V.

Number of setze: Group 1...

99 eoe aed

s Sie 3

BIO 5 i

me ep e OO

— eo — 69

3 a

It is to these sete: on the inner face that the eggs are attached,

and the outer feathered sete are no doubt developed partly as a

protection to the eggs, and partly to assist in promoting a current

of water through them.

The presence of these ovigerous sete may be taken as a certain

indication that eggs are about to be laid, or that young have been

hatched ; while their loss marks the end of the breeding-season.

Palemonetes varians var. mesogenitor Sollaud.

The peculiarities in the reproductive habit of the brackish and

fresh-water forms of P. varians are well known, and ‘ physio-

logical” varieties have been distinguished, namely var. microgenitor

Boas, var. mesogenitor Sollaud, and var. macrogenitor Boas,

according to the size of the eggs laid and the stage of development

at which the young are hatched. These varieties are, however,

said to be structurally indistinguishable. ;

122 MR. R. GURNEY ON LEANDER LONGIROSTRIS

The variety mesogenitor Sollaud* oceurs in Tunisia, and I have

a few specimens, taken in the stream flowing from the Sources

D’Oumach, near Biskra, in Algeria, which would probably belong

to this variety. They are all small, not exceeding 24 mm., but

include four adult males, while the females appear also to be

fully grown. While the femaies differ to some extent from

Norfolk specimens in the relative length of the joints of the

second leg and the proportions of the short flagellum of the

Text-figure 6.

—S SS Se eee

Palemonetes varians.

A and B. Posterior and anterior views of appendix masculina of two

specimens from Algeria.

C. Appendix masculina of specimen from Essex. 34 mim.

antennule, the material is not sufficient to justify any definite

conclusion as regards varietal or specific differences. It is other-

wise with the males. In all these four males there are two

characters which very distinctly separate them from the brackish-

water representatives of the species. These are :—

(1) The appendix masculina of the second pleopod is slender,

and extends by nearly half its length beyond the inner branch of

the pleopod. There is a group of strong spines at its apex, but

the sides and anterior face are bare or armed with only one or two

* Sollaud, C. R. Acad. Sci. Paris, t. 155, p. 1268 (1912).

AND OTHER BRITISH PRAWNS. 123

spines (text-fig. 6 A, B) In Norfolk specimens (text-fig. 6C) the

appendix is always very much shorter than the inner branch of

the pleopod, and is provided with a row of spines along the inner

and anterior side. This difference is very striking, and easily

seen without dissection.

(2) The short flagellum of the antennule is almost or quite as

long as the peduncle, and the free part is nearly half as long as

the fused part. .

Tasxe LIT,

Comparison of measurements of Norfolk and Algerian

specimens of P. varians.

Srconp LzEe. ANTENNULE.

Ee = eee

Sle Sle eielseleg| 2

RO; h|O/a] A |B See es

Norfolk, 8 eescccsscs., 100/ 250/150 | 925/964//100) 62) 45 | 17

ltiae) oa |, [269 169 | 331| 254|| _,, 10 | 49 | 21

Meera ude yy hilt boen 167 329 236|| ,, | 100 | 64 | 36

We i Samo a Ps | eee | ey pce) | 67 | 29

e © aspse lates es 04 s (209 jee ee > | Ge) 6a 9

In all the specimens, male or female, there are either three or

four feathered sete on the telson.

I have not been able to examine any representatives of the

South European fresh-water form, and, so far as I know, no

attention has previously been paid to these two characters of the

male, so that it is possible they may be found to be distinctive

not of var. mesogenitor alone, but also of var. macrogenitor. I

consider them of such importance as to justify the separation of

the Algerian form as a distinet species, which, if it should prove

to be identical with the Tunisian form and not to share its

characters with var. macrogenitor, should take the name P. meso-

genttor Sollaud *.

* Since the above was written M..H. Chevreux has been guod enough to send me

specimens of P. varians from Lake Fetzara in Algeria, but unfortunately they

proved all to be females. I have also, through the kindness of Dr. W. T. Calman,

been able to examine specimens of the var. macrogenitor from several localities in

south Europe and find that the appendix masculina does not differ from that of the

brackish-water variety.

i Fibs “iota ) as iongarsabes % soni

} Panda ‘h cine neclinanblst iY

Meas ale ced to t Feat

ida ay en ee sid re

yey

A) aT eke che it py st. 4

Silanes Pe i) ovina ot Wide: ged diet mea” tans

AP SHE OS. bith eed nae oe meek Bhi.

i aeed sy an tb weal es Fagg tpl Kaa ay i

Bibs ad eave) te yay Aha: lebie

r Fe tt

ne ee sas: ?

Ph eel | TO Fis ape es

Peay, Heike. Rig,

. x tae mldicid ght

be be ii eA 4

i) |

ON DEATHS. IN THE GARDENS IN 1922. ]

10. Report on the Deaths which occurred in the Society’s

Gardens during 1922. By N.S. Lucas, M.B., F.Z.8.,

Pathologist to the Society.

[Received January 26, 1923: Read February 20, 1923.]

On January Ist, 1922, there were 696 mammals in the

Gardens, 1878 birds, and 759 reptiles.

During the year 459 mammals, 1026 birds, and 334 reptiles

have been added, making a total for the year of 1155 mammals,

2904 birds, and 1093 reptiles.

Of the 1155 mammals 320 have died, giving a percentage of 27-7.

2904 birds 583 a i 5 20:0.

» lO93 reptiles 237 33 ‘3 99 21°6.

The percentages for 1921 were 21°4, 20-4 and 24:5.

¥ a 9 L920 9) Bogie oes 201:

If only those animals are considered which have lived in the

Gardens for more than six months 167 mammals have died and

95 reptiles.

This gives a percentage for mammals of 14:4. —

” ” ” ” reptiles ” 8°6.

The percentage for 1921 was 10-0 for mammals.

be)

9) 39 ” 1920 ? 16°9 19 .

” ” ,, 1921 ,, 8:1 for reptiles.

jy O20) ae es x

7 9

For the reasons given last year, it is impossible to give this

percentage accurately for birds. -

From these figures it would appear that the warm, dry summer

of 1921 was distinctly beneficial to the mammals.

It has been felt for some time that the death rate in the

unacclimatized mammals has been unduly high. It is hoped

that the reservation of a special room in the Sanatorium, where

small mammals and birds can be kept under conditions of extra

care and attention, will do something to lower the death rate

among new arrivals.

It is satisfactory to ncte that the death rate among the

mammals from tuberculosis is still falling. The number of

deaths among birds from this disease has risen. This is probably

due to the large numbers of birds now in the Gardens.

Especially is there crowding in the Western aviary, for of the

31 deaths from avian tuberculosis, 17 have occurred in that

aviary. he figures for nephritis still continue Jow. The two

diseases which have accounted for the highest numbers of mammals

and birds are congestion of the lungs and enteritis.

Of these, the former represents a state where the lungs contain

a great quantity of blood and are darker than normal. They

may even reach the stage of being a dull red purple, but they are

126 DR. N. §. LUCAS ON

still aerated, in contradistinction to pneumonia, where they are

consolidated.

The latter represents a state of affairs where the alimentary

tract shows various stages of inflammation, varying from bright

red to purple, with sloughing of the mucous membrane.

The rarity of pleurisy among mammals referred to in my

report for 1920 is still maintained. There have been no cases

of this disease in either 1921 or 1922.

Table of Causes of Death.

Reptiles Wamibers

Causes of Death. Mammals.| Birds. ra >| yveferring to

i Notes.

Pulmonary disease.

IB AGUOTOTEY, W-cosbauscoboos osu osecesea: 35 30

Congestion of lungs ............... 51 73

@idemayotluneseece ee ee te 1

PN ORCS Orb IDNs <ooobnsssdesoceseos Bit 1

Acariasis of lungs .................. iL a ies

IBN Weenie Sain aoe niere eon iece aie 2 Bi ate 1

J BROS) CUUNIS). “yodne syenecuccecetosoasane 22

Cardiovascular.

Pericarditis ies ctosaiceene tes ene i 7

PASAT A ii heeds Mescsce eee APE 2

Ruptured mesenteric artery ...... 1

fae

Renal disease.

INGphTibis' ees ccteete nessa testes cei. 9 11 2

Granuloma of adrenal ............ 1 fi Be 2,

Alimentary Tract.

Abscess of mouth ..............00..

SHHOMMRNTNS so con gousemnod epena wanes oe det

Acute dilatation of proventriculus a5 2 ne 3

RG astritisuedyenenass ccc com ane rel

|. Gastro-enteritis 2.20.0. coceee... ee |

JOSAUT UIA) esas pon coaceMeee cree a |

| Constipation acme seeaptets

Intussusception .....................

CESHHO WIG coccecusaccehe sen anabec |

| Strangulated hernia

| Prolapse of bowel

pat

PRE EEO oo

185 24,

ors

Hepatic.

| Fatty degeneration of liver ......

Cinhosistotmliversese- eee eee

Him Necrosisrotllivermees teen een. ae ae

| Granuloma of liver.................. is age il

Portal pyzemia Ree marca Reis f.

VATINGICOMMCATISe REE En ene 1

eres

ars

Pancreatic.

JERNDOIREE AINE) sco Sonat cacsebaddedsson ene A

Genital organs.

IRE OACHUAE. «coo one noo onaoee pebeeeeee

Iniflamed oviduct... ............-.-...

Prolapse of oviduct ............... An

Accidents of labour ............... 3

ro Or

DEATHS IN THE GARDENS IN 1922.

Table of Causes of Death (cont.).

127

Causes of Death.

Nervous System.

Convulsions ? cause

Acute Infections.

Tuberculosis

| ‘Tick imfection .......

Malignant disease.

Carcinoma of thyroid

Disease of Thyroid.

Adenomatous cyst ...............0..

CWollordicorbreW ent dassseeseneee

Miscellaneous.

Inanition P cause

Scurvy ?

Kallledieaereeees.

Paralysis Peausetys acl sieiae

Melavedish oc kaerrmeen. joss en:

IN GOSS: sandandaceanadepeess

SI DIOFETTIID, 55 boc ongcoscoosan sooteones

General serosibis'. <2... ess.essescec.s.

Je lViaaVOTIE VEINS 45) 5 245 segeoneeceoseee

Osteomyelitis of jaw ......

Reribonibisiee secs e ne

|) Protozoal infection .........:........

Sarcoma of liver ...............

Carcinoma of liver .........

THOS ee ee!

Mente eet ie

Wrawmedy asc scones.secsesets

OSPOSUTC ete terc se stavsneceee asec ee

Under anesthetic .........

Destroyed by rats ...............05.

Notidiasnosedienstee.7-s-see ety ee eee

Not examined ...............

Wecomposedenaemesisc eee er|

Mammals.

See

/ powe

Birds.

tt he

Reptiles,

etc.

Numbers

referring to

Notes.

Explanatory Notes on the Causes of Death.

1. One case was that of a hyena.

A sharp piece of bone had

perforated the stomach, the diaphragm, and pleura so

that the gastric contents had escaped into the pleural

cavity.

The other case was that of a Napu mouse-deer.

. In a rufous-tailed fire-back pheasant.

gorged them selves.

In a green cercopitheque.

These were small craterous ulcers in the rumen of a gazelle,

There was also one ulcer in the reticulum,

. In a bear,

6. This was in a Caucasian ibex,

3. Both of these cases were birds of prey which had over-

128

ON DEATHS IN THE GARDENS IN 1922.

A fishing-cat on post-mortem showed inflammation with

exudation of clear fluid in all its serous cavities. It was

apparently chronic, as the peritoneum over the liver

showed ciliated epithelium.

. This was a chronic condition in a jungle-cat. The stomach

showed extensive secondary ulceration.

. A sarcocystus infection in a Chinese quail.

. Ina hamadryad.

Two of these cases were in wolves and one ina jackal. All

showed secondary deposits in the lung.

In a hybrid pheasant and Wyandotte hen.

. In a wolf, apparently primary.

. Ina Cayenne rail. It was spherical and 2 em. in diameter.

It caused death by compression of the thoracic viscera.

. Ina Mandarin duck.

. In a blackbuck. This had been bred in the Gardens and

had produced a young one, which had died a short time

previously and shown similar but much less extensive

signs of the disease. For a long period there had been

no change in the feeding, which seems to have been satis-

factory from the vitamine point of view.

THE ANATOMY OF THE TONGUES OF THE MAMMALIA. 129

11. The Comparative Anatomy of the Tongues of the

Mammalia. — VIII. Carnivora. By Cuaries F.,

Sonntac, M.D., F.Z.S., Anatomist to the Society.

[Received November 20, 1922: Read March 6, 1923. ]

(Text-figures 15-24.)

ConTENTS.

Page

Introduction Aisa se ees eee tee ee recente Lao

Subordensbssipedtaymceneeseree seete este een eeerese eee meee)

SectioneAulunoideay a. ere een eee eect LZ

wee Cymoideat, sacacdete Mitac eo eee acc) LA

ee.) Hr ChOld eal: 2:sas/araconseceeee omer ee La

Suborder Bimmipediay nereeeeere. eects aceasta

Systematiey).c5. des sccsseaeee oes 148

Comparisons with the Cetacea ...............2..:.-....:. 151

Bibliosraphyeyescees sees eee eee eee coer Oy

INTRODUCTION.

The materials on which the present paper is based consisted of

preserved and fresh specimens in the Society’s Prosectorium and

the Museum of the Royal College of Surgeons. Nearly five

hundred tongues, representative of most genera were examined.

Characters were observed which are of considerable value for

purposes of classification, and a systematic arrangement of the

tongues of the Felidae, based on these features, agrees closely

with Pocock’s system (24), which was founded on external and

cranial characters. It will be seen that the tongues differ from

those described in my papers on the Primates (29), Ungulata,

Sirenia, and Cetacea (30). Speaking generally the mechanical

power is frequently greater and the gustatory organs are fewer

than in many animals already described. The conical papille

have strong, horny sheaths.

The tongues of the Fissipedia differ in many respects from those

of tne Pinnipedia.

Suborder FISSIPEDIA.

Section AILUROIDEA.

The tongue is long and comparatively narrow in all species, but

it is wider in Felis leo, F’. tigris, and F. onca than in other cats.

It thickens progressively from before backwards, but the free part

is never very thick. he sides are parallel or tapering, but the

tongue is spatular in a few cases. The apex is truncated or

rounded, but it is usually devoid of a notch. It is beset with

Proc. Zoou. Soc.— 1923, No. IX. 9

130 DR. C. F. SONNTAG ON THE ANATOMY

small conical and fungiform papille, but these do not stand up

prominently. And there’is in no case a large cluster of promi-

nent fungiform papillae containing large taste-buds, such as are

present in the Primates. The lateral borders are rounded, but

lateral organsare variable. ‘They have many backwardly-directed

conical papille; these usually lie flat in the Felide, but they

stand up prominently in some of the Viverride. No lobules are

present such as occur in the Cetacea.

The oral and pharyngeal parts of the tongue can be easily dis-

tinguished from one another when the vallate papille form a V

or a triangle, and when there is a marked contrast between their

conical papilla. It is, however, difficult to distinguish them when

the vallate papille form rows which do not meet, and when the

conical papillze maintain the same characters from the apex back

to the epiglottis. The papillose base of the tongue is close to the

epiglottis in all Adluroidea except Lelis leo and Ff. onea, in which

an extensive, smooth tract of mucous membrane intervenes

between the base of the tongue and the epiglottis.

The conical papille in the Felide, Hyznide, and Proteles,

have thick, sharp or rounded, horny sheaths, which can exert

a powerful mechanical action on the food. This character is

less marked in many of the Viverride. In some of the papille

in Proteles cristatus they form blunt caps for rounded papille

(text-fig. 21).

Median dorsal sulci and transverse dorsal ridges and sulci are

absent. And there is no trace of frenal lamelle, sublingua, plicze

fimbriatee, or foramen cecum.

Circumvallate Papille (text-fig. 15), The following patterns

are present :—

1. A pair of papille.

2. Two or more pairs of papille arranged in converging rows,

which do not meet in an apical papilla.

3. Three papille in a triangle with the apex behind.

4, Several papille in v-formation.

5. Papille form a double V.

The converging rows and the v can be regarded as incomplete

or complete chevrons. There is no trace of T or Y types, or of

fields of papille. And in no case were papille absent. It will

be seen that pairs of papille are present in the Felide, Hyznide,

and Proteles, but all types exist in the Viverride. As some

tongues have no lateral organs, and few fungiform papille with

taste-buds, the vallate papille are frequently the main gustatory

organs.

It may be difficult to count the number of papille, for they

are sometimes very small and inconspicuous, or concealed by

overhanging conical paypille. They may be all equal or unequal

in size. They are circular, oval or dumbbell-shaped when

viewed from above, and they are cylindrical or conical on elevation.

One or more may be included in one fossa. The vallum and

OF THE TONGUES OF THE MAMMALIA. 131

fossa vary in prominence, and the papillary surface is smooth,

finely granular or papillated. In the Felide} Protelide, and

Hyeenide there are one or more pairs of papille in rows which

converge, but do not meet. In the Viverride there are in

Text-figure 15,

@ 3) ie

® g

ve eo eo

@7° ror f

® @ @

@°o o

@ oS.

A. iN 2

f= a cy rm Vx Vi

The circumvallate papille ; a-k: papille on’plan; 7: papille on elevation.

addition to this pattern three papilla in a triangle, or several

papille in a single or double Vv.

The arrangements seen by myself and recorded by others are:—

Family FEevipz.

Felis leo :—One to five pairs of papille ; seven papille in two

rows.

L”, tigris :—One or two pairs ; seven papille in two rows.

F. pardus :—-One or three pairs.

F. domestica, F'. sylvestris, F. caffra, F. viverrina, F. nebulosa,

LF, bengalensis, F. concolor :—T wo pairs of papille.

F. lynv. F. caracal, &. rufa, F. pardalis :—Three pairs of

papill.

Cynelurus jubatus :—Two or three pairs of papilla.

Family VIVERRID&.

Viverra civetta :——Three papille in a triangle.

» fusca :—Six papille ina y.

Oivettictis civetta :—Three papille in a triangle.

5 5; Five papille in a V.

132 DR. OC. F. SONNTAG ON THE ANATOMY

Viverricula malaccensis :—Three papille in a triangle.

Genetta felina :—Four papille in a v.

» pardina :i— ,,

» tegrina:—LIwo papilles on the right side and one on

the left.

Hemigale hardwicku :—Three papille in a triangle.

:—Four .. CaN

Par nanan: Us lee watus:—Three ,, , triangle.

- hermaphroditus :—Three papille in a triangle.

4 typus :—Vive papille in a V.

Mungos mungo, M. ichneumon (several examples of each

species) :—Three papille in a triangle.

Atilax paludinosus :—A. pair of papille.

Nandinia binotata, Suricata tetradactyla, Galidia elegans :—

Three papille in a triangle in several examples of each

species.

Arctictis binturong :—Highteen papille in a double y (text-

fig. 15h).

Cynictis penicillata :—Three papille in a tr iangle.

Crossarchus obscurus:— _,, 8 a My

Family Hy@nipa@.

Aycena striata :—A pair of papille (text-fig. 21).

:—Two pairs of papille.

H. crocuta UN pair of papillee.

Family PRoreLipa.

Proteles cristatus:—A pair of papillee (text-fig. 21).

Fungiform Pupille :—-These are numerous or scanty in the

Afluroidea; and in some tongues they are well represented, but

indistinct. They may stretch right across the dorsum, or they

may be absent from the centre, ther eby forming a dorsal bound-

ing zone. They are numerous, scanty or absent on the ventral

papillary zone. They have the usual arrangement in clusters

and rows of varying degrees of obliquity. They are hemispherical

or pedunculated, and the surfaces are smooth or granular.

Their distribution on the oral part of the dorsum is determined

by the presence or absence, and characters, of the patch of papille

spinose. As a rule they are absent from the latter, but some

are present on it in F. caracal (text-fig. 20).

In many Carnivora lateral organs are absent, and it is probable

that fungiform papille lying on the postero-lateral parts of the

dorsum replace them. This has been definitely proved to be the

case in Felis domestica. And it is shown below that club-shaped

fungiform papille with well-marked taste-buds lie in the position

of Jateral organs in some Felide.

In the Felide there may be many fungiform papille just behind

the apex, but they are never as large and prominent as those in

OF THE TONGUES OF THE MAMMALIA. 133

the Primates. The cluster contains many papille in Felis pardus,

fF. nebulosa, F. bengalensis, F'. concolor, and Cynelurus jubatus.

It contains few papille in /. leo, &. tugris, F. viverrina, and

F. sylvestris. No apical fungiforms are present in J. pardalas.

The fungiforms situated along the sides of the oral part of the

dorsum also vary. They are absent in some examples of /. leo,

moderately developed in /. sylvestris, F. pardalis, and F. concolor,

and numerous in /. pardus, F. nebulosa, F. viverrina, F’. benga-

lensis, and Cynelurus jubutus. No papille are present in the

spinous patch, except in 7’. caracal (text-fig. 20). On the dorsum

behind the spinous patch there are insignificant or prominent

fungiform papille ; they are poorly developed in JF. leo, F. tigris,

F. viverrina, Ff’. pardalis, and Cynelurusjubatus ; they are present

Text-figure 16.

“OF Say

« < AW et

s 5 . A i! | wh AG

V spinous 2 hak id AA GAT Ses

x PATCH = FE domest. FE. caracal.

Seis

NSS

ea) Se | aren

N 8 = zg 2 sO Kk cattra.

Se UN ISU Se

a/g 8 8 § & B.

af ue Diba ae

Site of cluster

of fungitorm

papillae in some

specles x Ba

papillae

@ ate @ clavatae

@ @ and lateral

e © organs

CONICAL PAPILLAE

ON B

The divisions of the Aluroid tongue (A.) and the characters of the papille

clavate (B.). F. domest.: domestic cat; F. sylv.: wild cat; a papille

with a taste-bud greatly magnified is seen in the lower right-hand corner.

in moderate numbers in /. pardus, numerous in /. nebulosa and

F. caffra, and very prominent in £. sylvestris.

On the posterior parts of the lateral parts of the dorsum there

ig sometimes a row of finger-like or clubbed papille for which I

propose the name of papille clavate (text-fig. 16 A and B). They

lie in the position of the lateral organs, and they appear to be

modified fungiform papille. Histological examination reveals the

presence of taste-buds in their epithelium. They are useful for

purposes of classification, being present only in Felis domestica,

F. sylvestris, F. caffra, and F. caracal. Until material comes to

hand Iam unable to state whether they occur in /. chaus and

F. lynx. They are relatively largest in /. caracal.

Watson and Young (33) stated that the fungiform papille are

134 DR. CG. F. SONNTAG ON THE ANATOMY

numerous in Hyena crocuta. In H. striata they are not very

numerous.

in Proteles cristatus (text-fig. 21) the papille form a marked

cluster in front of the vallate area, and others are all over the

dorsum.

In the Viverridz the conditions of the fungiform papille are

more varied than in the Felide, for the patch of papille spinose

is absent, weak or strong; and the distribution of fungiform

papille depends on its characters. In Nandinia binotata (text- .

fig. 21), Veverra civetta, and Civetiictis civetta there is no spinous

patch, and fungiform papille cover the dorsum from the apex of

the tongue to the vallate area, and from one side to the other.

A few papille are present on the ventral papillary zone. In

Paradoxurus larvatus the patch is small, and its component

conical papille are slender; fungiform papille encircle it, and a

Text-figure 17.

j y 24 Pla) \)\ saya

The lingual papille seen under a hand-lens. 1-26: conical papille ;

27-30: fungiform papilla.

few are found on it. In other species the arrangement of the

fungiform papille and the patch are as in the Felide.

The characters of the fungiform papille are shown in text-

fig. 17.

No papille clavate are present in the Hyenide, Protelide,

and Viverride.

The value of the clavate and other fungiform papille for

purposes of classification is shown on page 149.

Conical Papille:—As the mechanical function predominates

over all others in the Ailuroidea, the conical papille are strongly

developed. And they are provided with sharp-pointed horny

sheaths. In the fore part of the oral division of the dorsum

there is, in most cases, a pateh of large conical papille which

have been termed papille spinose; and the area on which they

stand is called the spinous patch in this paper (text-fig. 16 A).

OF THE TONGUES OF THE MAMMALIA. iB ys)

The patch may be completely surrounded by a zone of small

papille, or it may extend laterally till it covers the whole width

of the tongue. The histology and chemistry of the horny

epithelial sheaths of the papille have already been described by

Severin (28), Klein and Verson (17), Csokor (7), Podwisotsky (25),

and Ranvier (28). ‘The ventral papillary zone may be composed

of small conical papille alone, or of both conical and fungiform

types.

Family Felide:—In all forms the spinous patch is surrounded

by a zone of small papille, but it varies in size, and in the

prominence of its papilla. The points of the papille are directed

backwards, and are straight or recurved to a variable degree ;

but those of the papille on the encircling zone face backwards

and inwards. The lateral papilla may increase or decrease from

before backwards, but the papille spinose increase in size from

without inwards.

In all Felide except /. leo and #. onca the conical papille

extend back to the epiglottis, but in these species the most

posterior papille are separated from the epiglottis by a smooth

mucous surface. The conical papille on the pharyngeal part may

be smaller or larger than those on the oral part of the dorsum.

The characters to which special attention must be paid are :—

1. The extent of the smail band of conical papille encircling

the patch of papilla spinose, and the relative quantities of fungt-

form papille situated on it.

2. The posterior limit of the spinous patch, and the distance

between the anterior extremity of the patch and the apex of the

tongue.

3. The characters of the papillee spinose, particularly the shape

of their spines and their closeness or separateness.

4, Characters of the conical papille behind the spinous patch.

5. Presence or absence of a row of papille clavate in the region

of the lateral organ on each side.

6. The relative sizes of the conical papille on the oral and

pharyngeal parts of the dorsum.

7. Character of the ventral papillary zone.

In Felis leo (text-fig. 18) the small papillary zone girdling the

spinous patch extends back along the anterior third of the tongue.

The spinous patch begins close to the apex of the tongue, but

does not reach the middle of the oral part of the dorsum. The

papille spinose have sharp, recurved, closely-set points so their

bases are not much exposed. Behind the patch the conical

papille form a central band ef small ones and two lateral tracts

of large ones. No clavate papille are present, and the papille on

the pharyngeal part of the dorsum are small. In /. tigris (text-

fig. 18) the conditions are in general similar to those in the lon,

but their development is not so great. The papille spinose have

recurved, but not very prominent points, but the conical papille

en the pharyngeal part of the tongue are prominent. Fungiform

136 DR. CG. F. SONNTAG ON THE ANATOMY

papille are more marked behind the spinous patch. The tongue

is not separated from the epiglottis by a smooth mucous tract. In

F. sylvestris, F. domestica, and F. caffra (text-fig. 20) the zone of

small conical papille surrounding the spinous patch extends far

back along the lateral aspects of the dorsum; it hag many fungi-

form papille at the sides of the tongue, but few behind the apex.

The spinous patch begins a considerable distance behind the apex

of the tongue, and extends to the middle of the oral part of the

dorsum. The papille spinose are discrete, their points are

straight, and their granular bases are visible. Papille clavate

are well-marked. The conical papille behind the spinous patch

are divisible, aceording to size, into a central and two lateral

Text-figure 18.

“glee

ran

mde

pa teat

hal aarr sta ef

Arh a py )

Sepilbns

Saggy te

, 4 Feabe

wives ee

iy wiry

VV NN VY?

Nouv {

lava

SUG ye

Felrs pardus.

Felis tigris.

leo.

Felis

The tongues of the Pantherine.

tracts; and the papille on the pharyngeal part of the tongue are

large. In F. pardus (text-fig. 18), of which several varieties were

examined, the dorsal bounding zone of small conical papillae only

extends along the apex and anterior third of each lateral aspect

of the dorsum; and it is well-studded with fungiform papille; it

is also very narrow when compared with the zone on other

tongues. The spinous patch begins close to the apex of the

tongue, but does not reach the middle of the oral part of the

dorsum. The papille spinose are discrete, and have sharp re-

curved points. No papille clavate are present. The conical

papille behind the spinous patch are divisible into a central and

two lateral tracts; and the papilla on the base of the tongue are

OF THE TONGUES OF THE MAMMALIA. 137

large, soft and not very closely aggregated. The tongue of

F’, nebulosa (text-fig. 19) differs somewhat from that of the last

species. The wide dorsal bounding zone of small papille contains

more fungiforms. The spinous patch begins far from the apex of

the tongue and reaches nearer the middle of the oral part of the

dorsum. In other respects the tongues are similar. In /, viver-

rina the dorsal bounding zone extends along the anterior third of

the oral part of the dorsum; it is narrow and has few fungiforms.

Clavate papillae are absent. The spinous patch begins a short

distance behind the apex of the tongue, but lies entirely in the

anterior third of the tongue. Its constituent papille are slender

and inconspicuous. The papillae behind the spinous patch are

arranged in a central and two lateral tracts, and the conical

Text-figure 19,

Hic

Cynae furus Ju Aztus. Felis nebu/osa, Felis pardal/s.

The tongues of the Acinonychine and Feline (No. 1).

papille on the base of the tongue are narrow, pointed, and discrete.

Inthe tongue of /’. bengalensis the dorsal bounding zone is relatively

wide and has well-developed papille fungiformes. No papille

clavate are present. The spinous patch begins some distance

behind the apex of the tongue and extends to the middle of the

oral part of the dorsum; its papille have strong points. The

central and lateral tracts of papille behind the spinous patch are

just distinguishable; and the papille on the pharyngeal part are

long, slender, and discrete. In F. pardalis (text-fig. 19) the

dorsal bounding zone is narrow and has few fungiform papille.

The spinous patch does not extend as far back as the middle of

the oral part of the dorsum. In other respects it is similar to

138 DR. C. F. SONNTAG ON THE ANATOMY

that of 2. bengalensis. In F. caracal (text-fig. 20) the bounding

zone is wide. The spinous patch is small, and begins far from

the apex of the tongue. The conical papille behind the patch are

divisible, according to size, into a central and two lateral tracts.

Papille clavate are very well marked, and form a row of discrete,

prominent structures. The conical papille on the base of the

tongue are pointed and discrete, being separated by wide areas of

the pharyngeal part of the dorsum.

In Cynelurus jubatus (text-fig. 19) the bounding zone extends

half-way back along the dorsum, and it lodges a few small

fungiform papille. The spinous patch begins far from the apex,

Text-figure 20.

ar} ae?

(iti Hose oe

Why Z

UNA ARRAN Meee 2 Bead

NG ae

: ; x

Felis carfra.

Felvs carscal.

The tongues of the Felinz (No. 2).

and does not reach the middle of the dorsum; and the points of

the papille spinose are slender and sharp. No papill clavate

are present. The conical papille behind the spinous patch are

divisible into a central and two lateral tracts, and the conical

papille on the base are small.

It is, therefore, evident that the tongues of the Felide are

divisible into two groups, according to the presence or absence of

papille clavate. In the former are Felis domestica, F. sylvestris,

FP. caffra, and FP. caracal. As no examples of F. chaus and F. lyna

have come to hand the above list cannot be regarded as complete.

OF THE TONGUES OF THE MAMMALIA, 139

In the latter are /. leo, F. tigris, F. pardus, F. pardalis, F. onca,

F. serval, Ff. viverrina, F. bengalensis, and Cynelurus jubatus.

The subdivision of the Felide by other characters is given on

page 149.

Family Hyende:—In H. striata the dorsal bounding zone of

small papilla is wide. The spinous patch begins far ‘trom the

apex, and is less extensive in proportion to the length of the

tongue than that of any of the Felide; the points of the papille

spinose are small. The division of the conical papillze behind

the patch into central and lateral tracts is not perceptible. The

Text-figure 21.

Nandinia

Sinotata.

‘gv ni ‘0d!

Hyaena Striata.

Pra pales cristatus.

The tongues of the Hyznide, Protelide, and Viverridz (No. 1).

conical papille on the base of the tongue are divided into a

central band of large elements, and two lateral tracts of small

ones. No papille clavate are present (text-fig. 21),

Family Protelide : —The tongue in Proteles cristatus is spatular,

and the spinous patch covers the whole anterior expanded part

(text-fig. 21). The papille on the patch are circular or blunt-

pointed cones. The conical papilla behind the patch are very

small and are not divisible into central and lateral tracts. Basal

conical papille are scanty, and no papille clavate are present.

Family Viverride :—The spinous patch is absent, small or well-

140 DR. C. F, SONNTAG ON THE ANATOMY

developed, and the development of the points varies as shown in

the following table :—

Greciee Development Development

i of patch. of points.

Ciwettictis civetta ............ — —

(IO CTIRIBN CO SCULAC POL REAPER SA SBOE DA. 4 +- +

Viverricula malaccensis ...... ++ +

Genelia folima ......eccceccb.0s +-+ sr

penloe CEGIONUG: A ce He Oras aR ar f=

Paradoxurus larvatus ...... + Pate

et hermaphroditus = + oF

Arctictis binturong.. ......... += +

Nandinia binotata ............ _ — (text-fig. 21).

Mungos ichnewmon...... . ... is argh

os UOOULGLD Sass kostpancr 5 45 “hat

Atilax paludinosus............ ++ ap aR ==

Galidia elegans ............05. ++ +

Cynictis penicillata ......... +44 ++

Suricata tetradaciyla......... + ot

Crossarchus obscurus ......... ++ +++ (text-fig. 22).

The conical papille have the usual direction, and increase in size

in the usual manner. ‘They usually lie flat on the lateral borders,

but they project from them in Nandinia binotata and Mungos

ichneumon. Behind the spinous patch they are not divisible

into central and lateral tracts as in the Felide. The conical

papille on the base of the tongue are usually small, and they are

close together or discrete. They are small and discrete in Van-

dinia binotata and Viverricula malaccensis; but they are small

and close together in Mungos mungo, M. ichnewmon, Civettictis

civeita, and Viverra civetta. They are large in Paradoxurus

larvatus, Genetia felina, Suricata tetradactyla, and Arctictis bin-

turong.

No papille clavate are present.

Lateral Organs ave poorly developed or absent. Various

authors have described them in Felis domestica (19,17, 7, 12, 31),

F. tagris (19), F. pardus (12), Viverra civetia (19), V. fusca (20),

Genetta tigrina (22), and Hyena striata (19). In my specimens

they were arranged as follows :—

Family Felide:—Organs absent in V/elis leo, F. domestica,

Ff, sylvestris, I. caffra, Ff. caracal, F. bengalensis, F'. nebwlosa, and

Ff. pardalis. They ave represented by slight fissures in F. par-

dus, f. concolor, F. viverrina, and Cynelurus jubatus.

Family Viverride:—Organs absent in all except Wandinia

binotata, Viverra civetta, Cynictis penicillata, and Suricata tetra

dactyla.

One or two foliate clefts are present in the Hyenide anc

Protelide.

Glands :—Lymphoid nodules and the orifices of ducts and pits

are scanty or absent on the base of the tongue. No apical gland

OF THE TONGUES OF THE MAMMALIA, 141

of Nuhn is present. In the cat mucous glands predominate over

Ebner’s glands.

Lytia:—The lytta lies in the free part of the tongue close to

the apex. It is short, and thick or slender; and it is not of

much mechanical advantage. Oppel describes its histology, and

recapitulates the views of Codronchi, Casserius, and Cuvier as to

its nature. It has been described in the cat by Flower (10) and

by Ludwig, Prince of Bavaria (18). In Felis leo it is three-

quarters of an inch long and half an inch thick.

The ventral surface of the tongue is plain. It has-a well-marked

frenum, and there is frequently no trace of a ventral mesial

sulcus and ventral papillary zone.

Section CYNOIDEA.

The form of the tongue and the characters of the apex and

lateral borders are essentially similar to those in the Atluroidea.

But mesial dorsal sulci are sometimes well marked.

The cireumvallate papille are arvanged in converging lines which

do not meet in all specimens examined by myself and recorded by

others. They are round or oval on plan, and cylindrical or conical

on elevation ; the fossa is usually well marked, and the vallum is

smooth, granular or lobulated. All papille may be equal in size,

or the members of a row may vary. In some cases the papille

are concealed by overhanging conical papille. The number of

papille in my specimens, and those in the Museum of the Royal

College of Surgeons are :—

Canis familiaris :—One to six pairs.

Cams lupus :—'Two or three pairs.

Canis cinereo-argentatus :— Five papille in two rows (81).

Canis aureus :—One pair.

Canis mesomelas and C. azare :—Two pairs of papille.

Camis thous, C’. occidentalis, C. juhatus:—Three pairs of papille.

Vulpes vulpes, V. bengalensis, V. leucopus, V. fulvus, Alopex

lagopus, Cyon dukhensis, Lycaon capensis, and Nyctereutes pro-

cyonides :—Three pairs of papille.

As the number of papillze is variable in those species of which I

was enabled to examine several examples they are useless for

purposes of classification.

The fungiform papille may be very obvious, but they are

usually minute, and it may be necessary to employ histological

examination to detect their presence. Ina specimen of Vulpes

leucopus I was unable to detect them at all. They have the usual

arrangement, but the apical cluster is not prominent. Some are

present on the ventral papillary zone. Well-developed taste-buds

are present in Canis familiaris and Vulpes vulpes. Papiile

clavatee are absent.

Conical Papille :—No spinous patch is present. The papille

have the usual arrangement in rows and clusters, but those on

the oral part are much smaller than those on the pharyngeal

142 DR. C. F. SONNTAG ON THE ANATOMY

division of the dorsum. A clear line of demareation, which is

concave forwards, separates the two groups. This arrangement

was also described in my paper on the tongues of the Lemu-

roidea (29). The forms are shown in text-fig. 17, 1-6 and 12-15.

Lateral Organs :—Vavrious authors have described them in

Canis familiaris (23), C. lupus (5), C. cinereo-argentatus (81),

C. mesomelas (31), C. jubatus, Vulpes vulpes, V. leucopus, and

V. fulewus. In Canis familiaris they are variable, being present

in some specimens, but absent in others. Authors have described

the histology of these structures in that species. In the speci-

mens at my disposal J found organs in some specimens of Canis

Samiliaris, but not in others.

In the specimens at my disposal I found the organs always

very slightly developed when present. No traces were present

Text-figure 22,

Cans Mmesaneas. Cercoleptes

Crossarchus O0bSCUFUS. ceudivolyuslus.

The tongues of the Viverridze (No. 2), Canidge, and Procyonidx (No. 1).

in (. lupus, C. mesomelas, O. jubatus, Vulpes vulpes, V. lewcopus,

V. fulvus. But they were present in all other Canide enume-

rated in this paper.

Glands :—In the dog and common fox serous glands surround

the vallate papille and lateral organs, and mucous glands cover

the base of the tongue. But there is no apical gland of Nubn.

Lymphoid nodules and orifices of ducts and pits are scanty on

the base of the tongue.

The frenum is well marked, but there are no traces of frenal

lamellee, plice fimbriate, and foramen cecum. The median

glosso-epigiottic fold may run forwards for a considerable distance

on to the pharyngeal part of the tongue and separate the conical

papille into two groups.

OF THE TONGUES OF THE MAMMALIA. 143

The lytta is well developed, and its histological characters

have been described by many authors, whose observations have

been collected by Oppel (23). In Canis aureus it is continued to

the hyoid bone by a long, slender fibrous thread.

The characters of the tongues of the Cynoidea are so similar in

many species that they cannot be used for purposes of classifi-

cation, as they can in the case of the Aluroidea.

Section ARGTOIDEA.

The tongue is long, narrow and thin. The apex is rounded

and may or may not have a notch. Notches are absent in the

Mustelide and MAdluride, but are well marked in some of

the Procyonide and Urside. Minute conical papille clothe

the apex, and the fungiform papille in that region are small

in the Mustelide and Atluride. The apical conical papille

are long and prominent, but the fungiform papille are small

in the Urside. The apical fungiform papille are large, but the

conical papillz are small in the Procyonide. ‘The lateral borders

are acutely or widely rounded, but lateral organs are variable.

The conical papille thereon lie fat in the Mustelide, Alluride,

and Procyonide, but they stand up prominently in the Urside.

It was shown in a previous paper (80) that lateral projections are

present in Sus and the Cetacea. Median dorsal sulci are present

in many species, but they are very prominent in Mustela martes,

M. erminea, and some species of Ursus. Transverse ridges and

sulei are very well marked in J/ustela erminea. The entire dorsum

is clothed with papille which may be inconspicuous or very

prominent, and the conical papille on the base may be much

larger, or smaller than those on the oral part of the dorsum.

Circumvallate Papille :-The following arrangements exist :—

1. One or more pairs of papille in two converging rows which

do not meet.

2. Several papilla in y-formation.

3. Several papillee in a semicircle.

The following list shows the arrangements observed by myself

and recorded by others:

Family Musrenipa.

Mustela martes :—Two pairs of papille.

», erminew:—Three pairs of papille in Miinch’s 27

specimens.

Ictonyx zorilla :-—A pair of papille.

Putorius vison :—Two pairs of papille in three specimens.

Meles meles :—Seven papille in a semicircle (text-tig. 23) or five

to nine papille in a V.

Arctonyx collaris :—Hight papille in a vy.

Lutra vulguris :—Five or eight papille in a vy.

Mephitis mephitica :—A pair of papille.

Conepatus proteus :—Five papille in a Vv.

Galictis vittata ;— Hight papille in a v,

144 DR. C. F. SONNTAG ON THE ANATOMY

Family Procyonipa.

Procyon, lotor :—Five pairs of papille, or twelve papille in a Vv

(text-fig. 23).

Nasua narica :—F¥our to eight pairs of papille in ay (text-fig. 23).

Cercoleptes caudivolvulus :—Ten papile in y-formation (text-

fig. 22).

Hlurus fulgens :—Ten or twelve papill in a vy (text-fig. 23).

Family Urstpa.

Ursus maritimus :—Two pairs of papille, or ten papille in a Vv.

» arctos:—16 papille in a vy.

» americanus :—20 papille in a semicircle.

», juscus:—Ten pairs of papille.

» malayanus:—Seven to thirteen papille.

», labtatus :—16 or 17 papille ina y.

Melursus ursinus :—Ten papille in a vy.

The papillz are round or oval on plan and cylindrical or conical

on elevation in all except Meles meles in which globular forms are

present. The vallum is smooth, granular or lobulated, and the

Text-figure 23.

Nasua rarica.

The tongue of Naswa and the arrangements of the vallate papille in the

Mustelide and Ursidee.

fossa is closed or patulous. Ina specimen of Meles meles (text-

fig. 23) the mucosa over the papille is brown and tesselated; but

this appearance was absent in five specimens. The vallums may

be in contact (text-fig. 23) or separated by intervals. Numerous

taste-buds are present.

The fungiform papille may be just visible to the naked eye, or

OF THE TONGUES OF THE MAMMALIA. 145

they may stand up prominently. They are inconspicuous in

Mustela erminea, M. martes, and some species of Ursus. They are

small, but obvious, in Meles meles, Lutra vulgaris, Arctonyx

collaris, Galictis vittata, Procyon lotor, and dilurus fulgens. They

are larger and more prominent in Vasua narica than in any other

Carnivore examined by myself. They have the usual arrange-

ment in clusters and rows of varying degrees of obliquity; but

they usually stretch from the mid line to the lateral borders of

the tongue. Those at the posterior parts of the lateral borders

are very prominent in Cercolepies caudivolvulus, and may replace

the lateral organs. ‘Taste-buds are present in the fungiform

papilla in many species. Papille clavate are absent.

Conical papille cover the entire dorsum from the apex back to

the epiglottis, and they frequently form a ventral bounding zone.

But there is no trace of papille spinose. The conical papille

have the usual arrangement into apical clusters and rows of

varying degrees of obliquity. But their distribution according

to size differs considerably as follows :—

1. Papille on the oral part of the dorsum large, but those on

the pharyngeal part are small :—Mustela, Arctonyx, Galictis.

2. Papille on oral part small, but those on the pharyngeal

part large: —Wasua, Procyon, Hlurus, Ursus, Melursus, Ictonyx.

3. Papillz small all over :—Meles.

4, Papillee large all over :—Lutra.

The papillae make the edges of the tongue rough in Mustela,

Lutra, and Nasua, but the edges are smooth in Meles. In Ursus

they stand up prominently on. the apex of the tongue and the

anterior ends of the lateral borders.

Lateral Organs :—There is considerable difference of opinion

as to their occurrence-in the Arctoidea. Various authors have

described them in Mustela martes, Putorius vison, Procyon lotor,

Cercoleptes caudivolvulus, and several species of Ursus. I observed

no trace of these structures in Mustela erminea, M. martes,

M. foina, Ictonyx zorilla, Mephitis mephitica, Conepatus proteus,

Meles meles, Lutra vulgaris, Procyon cancrivorus, Naswa narica,

Ailurus fulgens, Cercoleptes caudivolvulus, Arctonyx collaris,

Galictis vittata. Some small foliate clefts were seen in Procyon

lotor, Ursus maritimus, and Melursus ursinus.

There is no trace of frenal lamella, sublingua, plice fimbriate,

or foramen cecum. Lymphoid nodules and the orifices of ducts

and pits are scanty on the base of the tongue

The lytta is present, and it reaches its highest degree of

development in Cercoleptes caudivolvulus, in which it is long,

ligamentous and ensheathed.

Suborder PINNIPEDIA.

The tongues are shorter and wider than those of the Fissipedia,

and they taper more rapidly.

Proc. Zoot, Soc.— 1923, No. X, 10

146 DR. C. F. SONNTAG ON THE ANATOMY

The apex is entire or cleft, and may be smooth or covered with

lobules and papille. It is round, massive, smooth, and entire in

Trichechus rosmarus. In Cystophora proboscidea it is rounded,

with a deep median cleft, and is covered with coarse lobules. In

other forms it has a wide apical sulcus separating its apex into

two lateral parts, which vary in appearance. Both halves are

covered with papille which may be restricted to the apices or

extend along the borders. As the specimens examined were not

at my disposal for histological purposes I am unable to state

whether the papille and lobules in the Pinnipedia are homologous

with those in the Cetacea and Galeopithecus. The halves are

rounded in Otaria californiana, and they are rounded and

laterally projecting in O. gillespii. They are sharper in Phoca

vitulina, Halicherus gryphus, and Cystophora cristata.

Text-figure 24.

oN anti, aye

\ ey

ry %, {

Trichechus Cystophora \ | Otaria | f Otaria

TOSINEFUS - proboscidea. it calforniana.\ giilesps.

' Halichoerus “

gry phus.

Phocea

Cystophora ‘

vitulina. A eda!

cristata.

iy APS

t RSShy q

: SSH

Ursus arctos.

The tongues of the Pinnipedia and Ursus.

The lateral borders ave rounded in all forms, but are elevated

in some examples of Phoca vitulina. They are massive in

Trichechus rosmarus and Cystophora proboscidea ; they are smooth

in the former, but covered with papille and lobules in the latter.

In other forms they have many small papille. In some forms,

such as Otaria californiana, there are rows of elevations at their

posterior extremities, but these must not be mistaken for lateral

organs.

Sulci and Ridges:—No ridges are present on either surface.

Mesial dorsal and ventral sulci are faint or absent, as in the

Cetacea, but transverse sulci were only observed in Cystophora

proboscidea.

In all forms except Trichechus rosmarus the tongue has no inter-

molar elevation, and the oral part can usually be distinguished

OF THE TONGUES OF THE MAMMALIA. 147

from the pharyngeal division. The former is covered with

papillae in all forms except Cystophora proboscidea, and the

latter is folded and covered with orifices of glands and pits, as

in the Cetacea.

The circwmvallate papille are the chief gustatory organs in

Trichechus rosmarus and Cystophora proboscidea, which have no

lateral organs. They are sometimes absent as well, so the gusta-

tory function is poor or absent in some Pinnipedia. The

following table shows the arrangement of the papille and lateral

organs :—

Species. Circumvallate papille. Lateral organs.

Otaria californiana.................. Absent. Absent.

SIO CULES Priam escent s S.o ete rece 5 or

Trichechus rosmarus ... 001... .11 0s. TinaV. 3

Halicherus gryphus .......00.....- Absent. Fe

LEIPOEE COAMICDE ono coe qeapnn nen cpacosane 5ina V. Well marked.

e 1 LE a ee 10-12 ina V. 2 s

i 35 EM eset eshiaat Seatataes 3 in a triangle. A re

Fs 4 USE OR ee UE a Fe 8-9 ina V. 5 x

Cystophora cristata ... .......6.10 Absent. - ss

Hs proboscided ..........0 8inaV. Absent.

It is, therefore, evident that gustatory organs attain their

greatest development in Phoca vitulina, and they are poorest in

Otaria californana, O. gillespii and Halicherus gryphus. The

conditions are intermediate in T’richechus and Cystophora.

Fungiform Papille :—These are numerous, scanty or absent;

and when many are present they are thickly clustered on the

centre or sides of the dorsum, or spread all over it. They are

hemispherical or pedunculated, and the surfaces are smooth or

granular.

Papille are absent in Halicherus gryphus. They are numerous,

especially on the centre of the oral part of the dorsum, in Cysto-

phora cristata, but they are aggregated around the edges of the

dorsum in Zrichechus rosmarus. In Otaria californiana they are

scanty, they stretch right across the dorsum, and they have the

usual arrangement. In Phoca vitulina the usual mammalian

arrangement is present. In the Cetacea fungiform papille are

absent.

Conical papille are absent in T'richechus rosmarus and Cysto-

phora proboscidea, but they are present in all other forms. And

they have their usual arrangement in clusters and rows on the

oral part of the dorsum. They also increase in size in the usual

manner from before backwards and without inwards. The

pharyngeal part is folded and devoid of papille standing up from

the surface in Cystophora, Phoca, and Halicherus, but large

ND} 2

148 DR. C. F. SONNTAG ON THE ANATOMY

pedunculated or tapering papille are present in Trichechus,

Otaria californiana and O. gillespit.

No papille clavate are present in the place of lateral

organs,

Glands:—The orifices may be numerous, scanty or absent.

In Trichechus rosmarus they are thickly clustered on the oral part

of the dorsum, but in other forms they are mainly present on the

pharyngeal part.

The frenum is slight or absent, and there are no traces of

frenal lamelle, sublingua, plice fimbriate, or foramen cecum.

The ventral papillary zone is narrow in Oéaria and

Cystophora.

Tuckerman (31) described how many fat cells are present in

the mucous membrane, submucous tissue, and subpapillary zone

in Phoca vitulina.

No lytta is present in the specimens of Phoca vitulina and

Otaria californiana examined by myself. And the median

glosso-epiglotitic fold is well marked in the former, but absent in

the latter.

The comparisons between the tongues of the Fissipedia and

Pinnipedia are shown in the following scheme :—

Suborder Fisstpep1A:—Tongues long, narrow, thin. Apex

entire. Few have laterally-prejecting papillae on margins.

Pharyngeal part not folded. Vallate papille always present. No

gland orifices seen on oral part. Frenum and lytta present.

Spinous patch and papille clavate sometimes present. Lateral

organs variable.

Suborder Prnnrpep1A:—Tongue shorter, wider, thicker. Apex

cleft in all except Z’richechus. Edges lobulated or have laterally-

projecting papillae. Mucosa of pharyngeal part folded. Many

glandular orifices present. Vallate papille frequently absent.

Lytta absent. Frenum slight. No trace of a spinous patch or

papille clavate. Lateral organs variable.

SYSTEMATIC.

When the characters described above are arranged in systematic

order the following results are obtained :—

Suborder FISSIPEDIA.

Section ATLUROIDBEA.

Spinous patch present in all Hyzenide, Felide, Protelide, and

most Viverridee.

Family Felide :—Conical papille behind spinous patch divided

into central and lateral tracts. Patch never reaches edges of

tongue. Basal conical papillae nevi r form a median strip.

Family Viverride :—Conical papille behind the patch not

divided into central and lateral tracts. Spinous patch never

OF THE TONGUES OF THH MAMMALIA, 149

reaches edges of tongue. Basal conical papille never form a

median strip:

Family Hyenide:—Conical papille behind spinous patch not

divided into central and lateral tracts. Patch never reaches edges

of the tongue. Basal conical papille form a prominent central

strip.

Family Protelide :—Conical papille behind spinous patch not

divisible into central and lateral tracts. Patch reaches margins

of tongue. Basal conical papillae never form a median strip.

Family Fevipa.

A. Spinous patch begins close to apex of tongue, and is restricted

to the anterior part of the dorsum. Papille spinosz have

recurved points. No papille clavate.

i. Tongue separated from epiglottis by a long, smooth tract

GOL TONOAO WS) WANS) FVD’ conecodan soocdeacn aoseae ka nA AOS Felis leo, F. onca.

ii. Tongue not separated from epiglottis by a long. smooth

mucous tract.

Gap eapilica: Spinosee) Weaks ccc -ea-eceer een eoeeareenesiees ete OL CAG 2S.

Hapbapilleespinosce strony ces. eeeeeepa chee etaerciehersieessoa ti F’, pardus.

B. Spinous patch begins close to apex of tongue and reaches

middle of oral part of dorsum. Points of papillz spinosz

recurved and strong. Basal conical papilles moderately

large or small. No papille clavate.

i. Basal conical papillz moderately long and close ............... F. yaguarundi.

ii, Basal papille small and discrete........................ F. concolor.

C. Spinous patch begins far from apex of tongue. Papille

spinose have strong, straight points. Papille clavate

present.

a. Fungiform papille on spinous patch ............60. cee F.. caracal.

b. No fungiform papille on patch.

i. Large fungiforms behind patch .................0-00..- 2000-00

ii. Small fungiforms behind patch ..................... F. caffra, F. domestica.

D. Spinous patch begins far from the apex of tongue. Papille

spinosze have strong, straight points. No papilla clavate.

FE. sylvestris.

a. Basal conical papillee scanty or absent .................. ++ FE. pardalis.

b. Basal csnicals long, sharp, discrete ......... F. viverrina, F. bengalensis.

c. Basal conicals large and close ............s00cseeeeeeesee ss. HY, nebulosa,

E. Spinous patch begins far from apex of tongue. Papille

spinose have slender points. No papille clavate. Basal

conicals numerous, minute, close...... ...... Cynelurus.

Pocock’s classification (24) agrees with the above as follows :—

Subfamily Pantherine =Group A.

Feline =Groups B, C, D.

Acinonychine = Group EH.

As regards the Hynide and Protelide the characters given

at the beginning of this section are all that can be obtained from

the tongue for purposes of classification. And the lingual

150 DR. C. F. SONNTAG ON ‘THE ANATOMY

characters do not show any aflinity between Proteles and the

Hyznas. Moreover, they do not show that Proteles resembles any

of the Herpestine Viverride as stated by Weber.

Family VIvERRID#.

A. Spinous patch absent or moderately developed.

a. Patch absent.

i, Conical papilla on oral part of tongue strong, but those

on base moderately developed ..,............ Nandinia.

ii, Conicals on oral al small, those on panel strongly

developed ............... Civettictis.

ili. Conicals on oral part strong, ee on age vealr or absent Viverra.

6. Patch slight .. Aas .... Paradoxrurus

c. Patch Pnodemntely iesetoped: Hane: “is papilla are meauler aA Viverricula.

d. Patch moderate, papille pojnted.

i. Vallate papille in a single V..........0...0..000022 cece eeeeeeees — Genetta.

iy Weller sin ey GO MDIG Wogsusscobccecteoee sun sdtupoaescen | sosbbduseens | AURBFIAHIG,

B. Spinous patch strongly developed.

a. Points of papille sharp ...............0..000..... Oynictis, Wungos, Atilax.

6. Points broad and blunt .................ccceece cee eeeceeeeseseesessseee Orossarchus.

é)7Papillee globular ei ata oo cecrie eM ee he ae Meee en ISULPLCO EOS

Weber divided the Viverride into Viverrine and Herpestine

subfamilies. The former corresponds to Group A, and the latter

to Group B in the above scheme.

Section CYNOIDEA.

No trace of a spinous patch in any species. Circumvallate

papille always in converging rows which do not meet. Fungiform

papille insignificant. Conical papille on oral part of tongue

small, but those on pharyngeal part large. Characters so similar

in all species that they cannot be used for purposes of classifi-

cation.

Section ARCTOIDEA.

No spinous patch. No papille clavate. Fungiform papille

more obvious than in Cynoidea. Conical papille variable.

A. Conical papille on oral part of dorsum large, but those on

pharyngeal part small... ...................... Mustela, Arctonyx, Galictis.

B. Con cal papille all small fr om apex of doneue back to epiglottis. Meles.

C. 3 5) 29 large 9 2” ” oy) oy) Lutra.

D. a 3 on oral part of dorsum small, but those on

pharyngeal part large.

i. Conical Baile on anterior ends of lateral borders stand out

prominently................ . Ursus.

ii. Conical papille on Part anee ail of idee porate a not Sent

out prominently.

a’. Fungiform papille on posterior parts of lateral borders

immense. Lytta very large... ..........c.0cecceessecseseee Cercoleptes.

6’. Fungiform papille large all over ...........0.....cccc0) wees. Nassua.

c! 3 i small ,, 0 5) veeceestssneseseceeseeee Procyon, Aulurus.

OF THE TONGUES OF THE MAMMALIA. 151

Suborder PINNIPEDIA.

A. Apex entire. Posterior part of tongue elevated. Mine fom

papille aggregated round edges of dorsum ........... Trichechus.

B. Apex cleft. Posterior part of dorsum not Besse. Tuner

form papillz not aggregated round edges of dorsum.

a. Halves of apex rounded and blunt. No vallate papille nor

lateral organs. Rare eee conical papille on base

of tongue ........ BHA nemca Non nceciaolass cud oj oor aM OREM aR OLLIE e

6. Halves of apex shai per. valare papillz or lateral organs or

neither present. No large pedunculated papillie on base.

i. No fungiform a No vallate papille. No lateral

HERING. soacdoce: Halicherus.

i. Fungiform papill, Fallatey papill, tral Tatoralic penn swell

marked ...... Phoca.

ii. Fungiform papille “present Vallatey - papill or ine

organs present.................. Cystophora.

It is, therefore, evident that Phoca has a fuller complement of

gustatory organs than all other Pinnipedia described above.

CoMPARISONS WITH THE CETACEA.

It 1s now believed that the Cetacea have affinities with the

Arctoidea and Pinnipedia, so it is necessary to compare the

tongues. A full description ef the Cetacean tongue is contained

in the previous paper of this series (29).

The tongue is long, narrow and thin in the Arctoidea, but

wider and thicker in the Pinnipedia and Cetacea. The apew is

entire in most Arctoidea and all Cetacea, but is deeply cleft in all

Pinnipedia except Zrichechus. The lateral borders have papille

in the Arctoidea; they are lobulated in the Pinnipedia and

Odontoceti, but entire in the Mystacoceti. Vallate papille are

present in all Arctoidea, many Pinnipedia, but no Cetacea.

Conical papille are present in all Arctoidea and most Pinnipedia,

but they are scanty or absent in the Cetacea. Lateral organs

are variable in the Arctoidea and Pinnipedia, but they are

absent in all Cetacea. The onfices of glands are innumerable

in the Cetacea, fewer in the Pinnipedia, and scanty in the

Arctoidea. The frenuwm is well marked and the lytia is present

in the Arctoidea, but the former is slight and the latter is

absent in the Pinnipedia and Cetacea. The lingual fat is well

developed in the Mystacoceti, in smaller quantity in the Odonto-

ceti and Pinnipedia, and very scanty in the Arctoidea. The

lingual muscles are well developed in the Arctoidea and Pinni-

pedia, but they are poor in the Cetacea.

It is, therefore, evident that the tongues of the Arctoidea

differ markedly from those of the Cetacea, but the conditions

are intermediate in complexity in the Pinnipedia. The condi-

tions in the Cetacea are a mixture of degeneration and hyper-

trophy. The mechanical and gustatory organs have almost dis-

appeared, but the glands have increased. And the pharyngeal

glands are very numerous as well. The secretion of these glands

152 DR. 6. F, SONNTAG ON THE ANATOMY

may contain a viscid or glairy substance which protects the

mucous membrane from being injured by prolonged immersion

in the water. I also observed numerous pits on the roof of the

mouth and on the tongue in fresh-water and marine fishes.

But they are not so numerous in the Pinnipedia, as might be

inferred from a comparative study of the habits of these groups

of animals. Poulton (26) believes that the large glands in the

bill of Ornithorhynchus secrete a substance which protects the

delicate integument, The glands may also be numerous in

the Cetacea because salivary glands are absent.

BIBLIOGRAPHY.

1. Asrar, A. K. v.—Avch. f. mikr. Anat. vol. viii. 1872, pp. 455-

460.

2. Bepparp, F. H.—Proc. Zool. Soc. 1895, pp. 430-437.

3. Bovunartr and Pruuinr.—cC. R. de la Soc. de Biol. vol. 36, 1884,

pp: 626-627.

4. Bounart and Pinuier.—Journ. de l’Anat. et de’ la Physiol.

1885, pp. 337-345.

5. Briicuur, C.—Deut. Zeit. f. Tiermed. u. vergl. Path. 1884,

pp. 938-111.

6. Cuatin, J.—Legons d’anat. et de la physiol. comparées.

Paris, 1880.

7. Csoxor.—Oest. Vierteljahrsschr. f. wiss. Veteriniirkunde,

1884, pp. 117-163.

8. Cuvier, G.—Legons d’anatomie comparée.

9. ELLENBERGER and Kunze.—Bericht iiber das Veteriniswesen

in K. Sachsen, 1884, pp. 148-164.

10. Fuower, W. H.—Medical Times and Gazette, 1872.

11. Garrop, A. H.—Proe. Zool. Soc. 1878, pp. 373-377.

12. Guetin.—Arch. f. mikr. Anat. 1892, pp. 1-28.

13. Honiescumiep, J.—Med. Centralb. 1872, pp. 401-403.

14. a * Zeit. f. wiss. Zool. 1877, pp. 255-262 ;

ibid. 1873, pp. 414-434 ; ib. 1880, pp. 452-459; ib. 1888,

pp. 190-200.

15. Kipp, P.—Quart. Jour. Micros. Sci. 1876, pp. 386-388.

16. Kien and Smiru.—Atlas of Histology. London, 1880.

17. Kuerin and Verson.—Art “‘ Darmkanal” in Stricker’s Hand-

buch der Lehre von den Geweben des Menschen und der

Tiere, 1869.

18. Lupwie, Ferpinanp.—Zur Anatomie der Zunge. Miinchen,

1884.

19. Mayer, F. J. C.—Nov. Act. Acad. Leop. Carol. Nat. Cur.

1844, pp. 721-748.

20. Mecken and Reicuman.—-Quoted by Miinch (see no. 22).

21. Merxet, F.—Arch. f. mikr. Anat. 1875, pp. 636-652.

22. Mion, F.—Morphol. Arb. 1896, pp. 606-690.

23. OppeL, A.—Lehrb. der vergl. mikr. Anat. vol. iii. pp. 365-

380.

OF THE TONGUES OF THE MAMMALIA. 153

. Pocock, R. 1—Ann. & Mag. Nat. Hist. 1917, vol. xx.

pp. 3829-350.

. Popwisotsky, V.—‘‘ Anat. Untersuch. iiber die Zungen-

driisen.” Diss. Dorpat, 1878.

. Pouxtron, E. P.—Quart. Journ. Micros. 8ci. vol. 36, p. 143.

. ScHwatBe, G.—Arch. f. mikr. Anat. 1868, pp. 154-187.

. Severrn.—Arch. f. mikr. Anat. 1885, pp. 81-88.

. Sonnac, C. F.— Proc. Zool. Soc. 1921, pp. 1-29, 277-322,

497-524, 741-767.

. Sonntag, C. F.—Proc. Zool. Soc. 1922, pp. 639-657.

. TuckeRMAN, F'.—Several papers quoted by Oppel (see no. 23).

. EBNER, von.—Several papers quoted by Oppel (see no. 23).

. Watson and Youne.—Proe. Zool. Soc. 1879, pp. 79-107.

. Scuacut, KH. C.—‘‘ Zur Kenntnis der secernierenden Zellen,

ete.” Diss. Kiel, 1896.

. Youne and Ropinson.—Journal of Anatomy and Physiology,

1889, pp. 187-200.

Had

res Bi

ren

ed vis HRV we i,

ete

ie)

/ Wi

Nays Cae iH

Bon es

Whee

PZ oO 2o. MIRKPAT RIG Karine

p i ) “EN UN

one

Highley del.

RHIZOMOLGULA GLOBULARIS PALLAS.

ON THE TUNICATE RHIZOMOLGULA GLOBULARIS. 155

12. Notes on the Tunicate Riizomolgula globularis Pallas *.

By R. Kirxpartricg, I.Z.8.

(Results of the Oxford University Expedition to Spitsbergen,

1921, No. 24.)

[Received December 1, 1922: Read March 6, 1923.]

(Plate I.)

Among the Tunicata collected by the Oxford University Expe-

dition to Spitsbergen, and sent to the Natural History Museum

for identification, were 26 specimens of the small sandy Simple

Ascidian Rhizomolgula globularis Pallas. As the material is

fairly well preserved, the writer believes a few anatomical notes

may be of interest.

Description.—The specimens (Pl. I. fig. 1) are somewhat tri-

angular in shape, and covered with fine sand; the largest are

about 10 mm. high, not including the stem-like basal process,

10 mm. broad and 3 mm. thick. The two orifices, one at each

end of a flat oval area, are usually barely discernible.

Oral tentacles (Pl. I. fig. 2): there are 8 large primaries,

8 secondary, 16 tertiary, and 32 quaternary, 7.¢. 64 in all.

Primary tentacles sickle-shaped, compressed laterally.

Dorsal tubercle horseshoe-shaped with opening forwards and

to the left.

Branchial folds six on each side, each with five inner longi-

tudinal vessels.

Infundibula deep, with the long double spiral gill-clefts wound

in the same direction ; accessory gill-slits rather simple, commonly

with a single curve nearly flat.

The hermaphrodite sexual gland in the loop of the intestine,

the minute oblong kidney, and the paired peduncular secretory

glands near the base of the endostyle, all on the left side excepting

the right secretory gland.

Locality. Klaas Billen Bay, just above low-tide mark, 7. e. ex-

posed to the air at low tide; near by a line of large boulders

close to the Nordenskiold Glacier (C. S. Elton).

The history of the genus Rhizomolgula is interesting. It was

founded by Ritter in 1891 for a new species of Molgulid from

Prince William Sound, Alaska, 20 fms., 7. e. outside the Arctic

Circle. The generic characters were the existence of the

peduncle, the presence of six folds on each side of the branchial

sac, the deep infundibula with spiral gill-cleft, and the left-sided

position of the intestines and hermaphrodite sex gland, and the

* (This title replaces the title published in the Abstract No. 236.—Eb. |

156 MR. R. KIRKPATRICK ON THE

presence of the peculiar glands supposed to be renal, one on each

side of the posterior end of the endostyle. In 1903 Hartmeyer

described a second new species, &. ritteri, from Baffin Bay,

20 fms.; the new species differed in having fewer inner longi-

tudinal vessels, and in having accessory gill-clefts supposed to be

absent in Ritter’s species. Hartmeyer showed that the paired

glands were not renal, but organs for supplying a secretion for

fixing the animal to the bottom; the true renal organ was found

by him to be a minute oblong body placed below the stomach.

In 1907-8 Redikorzev described two more new species, #. gi-

gantea from Kotelnyi Id. and New Siberian Ids., 3-9 metres ;

and R. warpachovskii from Nova Zembla, 40 metres.

In 1908 Michaelsen added a further new species 2. intermedia,

based on some specimens found in the Hamburg Museum, and

from an uncertain locality.

In 1911 Huntsman * made the interesting discovery that in

1776 Pallast had described an undoubted Rhizomolgula, viz.

Ascidia ylobularis from Kara Sea, and he identifies some speci-

mens from Herschel Id., Canadian Arctic Ocean, with this

species.

Lastly in 1916, Redikorzev{ arrived at the conclusion that

all the forms hitherto described belonged to one species, Ascidia

globularis Pallas, the supposed specific differences being merely

variations such as might be found in a large number of specimens

from any particular locality ; or, again, that certain variations in

external appearance (especially as regards the peduncle) might be

due to methods of collection and preservation.

The writer adopts Redikorzev’s view with the reservation that

a more detailed study of the oral tentacles might show genuine

specific differences. ‘The number of primary tentacles might be

regarded as a specific character, but not the number of multiples

of the primary number.

The specimens from Spitsbergen have eight primaries; but

there appears to be some uncertainty concerning the number

of primaries in the other supposed species [R. arenaria, 12-14

tentacles ; R. ritteri, 12 large ones, also with smaller (about 18 in

all); R. warpachovskii, 12; R. gigantea, 18 in three sizes; A. anter-

media, 40 in three orders].

Distribution. Pallas’s original Ascidia globularis was gathered

in 1770 from Kara Bay, Kara Sea. Over 130 years later the

same species (named R. warpachovski Rdkrazv.) was got from

Matochkin Strait, Nova Zembla, practically the same locality ;

for Nova Zembla curves round Kara Sea like a sickle separated

from the mainland only by another island. The species is

recorded also from New Siberia and from two American Arctic

* Trans. Canadian Inst. 1911-21, 1x. p. 127; also Contrib. Canadian Biology,

Ottawa, 1912, p. 136.

+ Reise, 1776, iii. p. 709, Appendix ; and Nova Acta Acad. Petropol. 1787, 11. p. 247,

Tab. vii. figs. 39, 40.

t Faune de la Russie et des pays limitrophes, 1916. Tunicata, livr. 1. pp. 126-137.

TUNICATE RHIZOMOLGULA GLOBULARIS. 157

localities (Baffin Bay and Herschel Id.), all within the Arctic

Circle; and from a place in Alaska outside that circle. The

material from Spitsbergen fills in a gap in the circumpolar

distribution.

Depth. Shallow water to 40 metres.

There are present in all the examples examined numerous

specimens of a species of parasitic protozoa of the group Suctoria

with a globular sessile body 0:5 to 0-1 mm. in diameter and with

one or more tufts of granular slender, vase-shaped tentacles

(Pl. I. figs. 2-4, 9, 10).

When the Ascidian is cut in half transversely and the front

half turned back, the Swctoria appear—under a lens—like snow-

balls scattered over the tentacles (fig. 2). They are distributed

over the anterior aspect of the tentacles and front wall of the

peripharyngeal groove, and never on the pesterior wall or the

avea behind it; they are especially abundant near the upper end

of the endostyle and dorsal lamina. This curious distribution

led the writer into mistaking the Swctoria for sensory organs of

the Ascidian, and he has to thank Dr. W. Tl’. Calman for suggesting

that the “ knobs’ were probably parasitic Protozoa.

In 1894 Dr. R. Gottschaldt (Jenaische Zeitsch. Bd. xxviii.

p. 343) described and figured Suctoria found behind the tentacles

of the Ascidians Polyclinopsis haeckeli Gottsch. and Synoicwm

turgens Phipps from Spitsbergen. The Suctorian figured by

Gottschaldt differs from the present one in having knobbed

tentacles. Dr. Gottschaldt regards his species as symbiotic

rather than parasitic, because the remains at least of the sucked-

out Protozoa brought to the Ascidian fall to the lot of the latter ;

whereas some of the ciliated Infusoria might, in the absence of

the Suctoria, have escaped altogether.

EXPLANATION OF PLATE I.

Figures of Rhizomolgula globularis (Pallas).

Fig. 1. A rather large specimen. Nat. size.

Fig. 2. Anterior part reversed, of a specimen cut across transversely, showing the

posterior aspect of the tentacles. 1, primary tentacles; 7'U, secondary ;

M, mouth ; At, atrial opening; P, peripharyngeal bands and groove :

dt, dorsal tubercle; d7, dorsallamina; Kn, endostyle; f, folds of branchial

sac; bs, cut edge of branchial sac. > 10.

Fig. 3. Groups of Suctoria on or near the anterior peripharyngeal band, near the

end of the endostyle. Hn, endostyle; P’, posterior pharyngeal band;

P"', anterior ditto; Tyl, the knob-lke Suctoria. X 18.

. Group of Suetoria in epibranchial area near dorsal tubercle, dt. 18.

. Part of tentacular ring and tentacles of four orders (mainly diagrammatic).

x 30.

Primary tentacle, antero-lateral aspect. > 60.

Primary tentacle, posterior aspect. Partly diagrammatic. x 40.

Primary tentacle, anterior aspect. Partly diagrammatic. x 40.

A Suctorian, with a nearly central nucleus-like body, and one tuft of

sensory hairs. X 280.

Fig. 10. Part of a Suctorian. X 1400.

OOD NE

FEEL AND RHINARIUM OF THE POLAR BEAR. 159

13. On the Feet and Rhinarium of the Polar Bear

(Thalarctos maritimus). By R. I. Pocock, F.R.S.,

F.Z.8.

[Received February 20, 1923: Read February 20, 1923. ]

(Text-figure 1.)

The fore feet resemble in a general way those of all the

northern species referred to Ursus and Huarctos in having a

single small isolated carpal pad entirely surrounded by hair and

separated from the plantar pad. The pads of the hind foot are

very similar to those of the fore foot, but the digitals are a little

smaller and the plantar a little larger, although of the same

width. A point in which the hind foot of Zhalarctos differs

from that of all other genera of Urside is the extent to which

the metatarsal area is overgrown with hair, reducing the meta-

tarsal pad to a small elliptical pad occupying nearly the same

position as, and only a little larger than, the carpal pad of the

fore foot. In the other genera the metatarsal area is either

wholly naked or is merely invaded on the inner side by a

narrow tract of hair along the groove marking the division

between the plantar and metatarsal pad. As subsidiary differ-

ences it may be noticed that the carpal pad is altogether smaller,

and the plantar pads of both fore and hind foot shorter, than in

the rest of the genera.

The digits of both fore and hind foot are separable by tolerably

equal spaces as in Ursus arctos, and, as in that species, the inter-

digital integument extends approximately half-way along the

digital pads.

The rhinarium resembles closely that of Ursus arctos, horribilis,

Euarctos americanus, and Selenarctos tibetanus. It is everywhere

sharply circumscribed by hair, though less so on the upper lip

than above. In profile view the internarial septum is not

concealed by the lateral border of the nostril; and there is a

deep, smooth infranarial area on each side, marked by a shallow

groove which diverges outwards and upwards from the middle

line to the nostril. Their point of union in the middle line is

crossed by another shallow groove, which descends vertically

from about the middle of the internarial septum to the ill-

defined philtrum, dividing the hairs of the upper lip.

The ears are in no respect degenerate, and resemble those of

160 FEET AND RHINARIUM OF THE POLAR BEAR.

Ursus and Huarctos in having well-developed supratragus and

basal ridges, with a narrow notch between the angular tragus

and the much less prominent antitragus *.

Text-figure 1.

A. Lower side of right fore foot of Thalarctos maritimus.

B. The same of the hind foot.

C. Hind foot of the same with hairs omitted to show the separation and

interdigital integument of the digits.

D. Rhinarium of the same from the front.

* Wor descriptions of the ears, rhinarium, and feet of other Bears, see Proc. Zool.

Soc. 1914, pp. 929-941, and Ann. Mag. Nat. Hist. (9) i. pp. 875-384 (1918),

EMBRYONIC DEVELOPMENT OF THE PORBEAGLE SHARK. 161

14. The Embryonic Development of the Porbeagle Shark,

Lamna cornubica. By HE. W. Suaxy, BSe., F.Z.8.,

Biology Master at Oundle School.

[Received November 13, 1922: Read February 20, 1923. ]

(Text-figures 1, 2.

ContTENTS.

Page

I. Recapitulation of Previous Work ..................... 161

LisRecordiof Material 2021p eee eee eee eae BS

EMR xtenmalMeatures: \. . 20.0: eee nee se OL

NVeelinbeynalobrucbrie\:.co.cse essen eereneeenaerrees ae tecaen ees

Wer ehysiologyot Nutritions |p. eee eereee eee) LOS

I. Recapitulation.

In 1910 I investigated the anatomy of the advanced embryonic

stage of Lamna cornubica. My vesults were published in the

Twenty-eighth Annual Report of the Fishery Board for Scotland.

Simultaneously with this publication there appeared a much

fuller account of the same subject, profusely illustrated, by

Lohberger (4). As this author placed an entirely new interpre-

tation upon the anatomy of the alimentary canal, as well as upon

the mode of embryonic nutrition, I was desirous of repeating

my investigations; I accordingly asked Dr. Williamson of the

Scottish Fishery Board, who had supplied my original material,

if ever he obtained more embryos of Zamna to give me the

opportunity of dissecting them. Cases of the capture of Lamna

embryos are few and far between, and it was not until 1922 that

Dr. Williamson was able to comply with my request. In

February of this year a female landed at Aberdeen was found to

contain four embryos, each measuring about 35 cm. in total

length. These embryos, together with the entire oviducts and

ovaries of the mother, were placed at my disposal; for which I

take this opportunity of thanking Dr. Williamson. One of these

embryos was a male, which is of interest as being the first

recorded case of the capture of a male embryo of Lamna.

As the result of my investigation of the latest material, J am

able to confirm Lohberger’s observations almost in their entirety ;

and this despite a strong preconceived scepticism regarding the

possibility of his view of the nutritive process, a scepticism which

was shared by all the zoologists with whom I had discussed the

subject. Briefly, the facts regarding the development of the

embryo are as follows :—

The original yolk-sac is absorbed at a very early period, when

the embryo measures about 6 cm. Thenceforward the cardiac

portion of the stomach becomes filled with semi-solid matter,

Proc. Zoou. Soc.—1923, No. XI. 11

162 MR. EB. W. SHANN ON THE EMBRYONIC

which process distends the abdominal wall of the body to an

incredible extent. The yolk-stomach so formed was erroneously

described by previous writers (including myself) as a yolk-sac.

The semi-solid matter is derived from the ovary of the mother

and is actually swallowed by the embryo in the uterus. Feeding

continues in this manner for a long time, certainly fur more than

a year, during which time the yolk-stomach continues to increase

in size (unlike a yolk-sac) until it assumes gigantic proportions.

It will be shown that there is a regular increase in the bulk of

the yolk-stomach from the embryo of 35cm. to that of 55cm.

Embryos have been found in the uterus measuring as much

as 70 cm. (Pennant, 5); unfortunately, no record is available

of the measurements of the paunch, though it is said to be

very large.

The smallest free-living specimens of Zamna measure 82 cm.

(Day) and 87cm. (Williamson); these, however, have lost all

trace of the yolk-stomach, and in external appearance possess 1n

every detail the character of the adult. Birth presumably takes

place when the young shark measures about 80cm. But to what

purpose is the great accumulation of food in the stomach put ?

Certainly it is not used in body-building, for the free-living

young are only a few cm. longer than the paunched embryos. It

seems possible that the accumulation of nutrient matter in

the embryo is used for the expensive purpose of building the

reproductive organs, and that the young at birth are already

mature*.

Such a condition would be unique among chordate animals ;

but, indeed, the very mode of nutrition of the embryo is unique.

Viviparity among Elasmobranch fishes is by no means uncommon,

and three general methods of nutrition are in vogue: either the

yolk-sac forms a pseudo-placental connection with the uterine

wall; or the latter secretes a nutrient fluid which is absorbed by

means of external gill-filaments; or, again, the uterine wall itself

produces long secretile ville which enter the alimentary canal of

the embryo by way of the spiracles. From all these recognized

methods of embryonic nutrition the condition in Zamna forms a

fundamental departure ; moreover, whereas in other forms the

maternal nutriment is used up at once in body-building, here

the vast majority of it accumulates in the stomach as a reserve

store.

After consulting the ‘‘ Zoological Record,” as well as from con-

versations with several eminent ichthyologists, I believe that I am

now in possession of all the outstanding facts at present known

regarding the embryonic development of Zamna cornubica. I

have attempted in the following pages to arrange these facts in

** At the same time Dr. Williamson, in a letter to me, says: “A female 3 ft. 6in.

long, in October, was immature. Further, a Porbeagle 6ft. long, in December,

appeared to be a male, but it had only a ‘slight indication of the external male

characteristics.”

DEVELOPMENT Of THE PORBEAGLE SHARK. 163

logical sequence, and to interpret them in the light of my recent

observations.

II. Record of Material.

Table I. contains a record of the captures of all Lanna embryos

known to me, and in addition of the smallest free-living specl-

mens. It is apparent at the outset that from the time when the

embryo has attained a length of 25cm. down to the time when it

is approaching readiness for birth (75 em.), we have a fairly

complete series of records. None of the embryos in this series,

however, differs in any important developmental character from

TABLE I,

q | Embryos.

s Authority. ae Length | Number roan

cS Capture. (cm.). | ae (Gay

A Swenander. Jan. — 2? 55 46:0

B_ | Calderwood. | -- Sai 2 25°

Cc Collet. Dec. — p

D Collet. Jan. ae 2? 29°5

E Swenander. — — | 4? 30°0

F Shann. Feb. 150 lg, B82 350

_G Collet. Feb. 256 4? 42°5

H Lohberger. — = | 22 | 4:98

5°33

on Shann. Mar. — 1?¢ 54-4

K Shann. — 150) | 49 45°4,

60°5

L Shann. June? -— 2? AT5

M Pennant. — — P 70°0

75°0

N Day. : — 82 (Smallest recorded free-

O Williamson. Nov. 87 living young.)

the others; the period which they represent, in other words, is

mainly characterized by growth in bulk. The only record of an

earlier phase is that of Swenander (7), who states that the

stalk of the original yolk-sac was present as a mere shred in

embryos 5°5-6:0 cm. long.

From the fact that the smallest known embryos (A) were

obtained in January, and that they measured not more than

6em., it may be inferred that fertilization takes place towards

the end of the year. In December and January again we

find embryos measuring 29°0 and 29°5 cm. (C, D); these have

164 MR. E. W. SHANN ON THE EMBRYONIC

presumably been in the uterus for just over a year, since it is

incredible that they are of the same age as the minute specimens

noted above. Allowing for the incompleteness of the record,

subsequent measurements bear out this supposition remarkably

well. In February the embryos may measure from 35 to 42cm.,

while at the end of March we find one measuring 54cm.

The last named is, according to our supposition, well on in

the second year of intra-uterine development. The figures

obtained for the months of December to March are sufficiently

consecutive to rule out the supposition that they represent the

lapse of a second year; in which case we may suppose that the

rate of growth during this period is more rapid than during

the first year, namely, about 8cm. per month. This estimate of

the rate of growth of the embryos during their second year, how-

ever, must be treated with caution; for an embryo (lL) said to

have been captured in June only measured 47°5 cm., which would

reduce our hypothetical rate of growth to about 4 cm. per month.

The rate of growth during this period is probakly liable to

variation; indeed, we find differences in length of 12 to 15cm.

between embryos in the same uterus (H, K). On the whole it

seems reasonable to infer that the average rate of growth during

the second year of intra-uterine development is 5 to 6cm. per

month.

Dates of the capture of the larger embryos (M) are unfor-

tunately lacking ; but if growth continues at the rate indicated,

we may suppose that they were captured about July and were then

about 21 months old. This supposition, when taken with the

observed fact that a free-living young specimen (O) was cap-

tured in November, seems to indicate that birth may take place

in September or October, i.¢., approximately two years from

conception. In what state the young are born is a matter

of conjecture. The Table shows conclusively that the young

at birth are not much longer than the largest intra-uterine

specimens; moreover, they are approaching maturity, if not

already mature. If the young are born with the yolk-stomach

still distended it seems unlikely that in their unwieldy con-

dition they could have evaded capture hitherto. Hence the

suggestion offered above, namely, that the yolk store is used in

building up the reproductive system. It is extraordinarily diffi-

cult to believe that the huge paunch is lost in the course of a few

months without any apparent effect upon the growth of the

young fish.

Ill. Haternal Features.

The external features of the advanced embryo are amply

illustrated in two earlier papers (Lohberger, 4, and Shamnn, 6).

In order to save trouble in reference to other works, and at the

same time to render the descriptions in this paper more readily

DEVELOPMENT OF THE PORBEAGLE SHARK. 165

intelligible, my drawing of an embryo measuring 454mm. is

reproduced in text-fig. 1. As there is no noteworthy difference

in the external features between embryos of 350 to 605mm.,

except in point of size, it is possible to make a general summary

of the distinctive characters.

In its natural position the cylindrical body is curled around

the massive paunch. The snout is extremely blunt and the

whole head dorso-ventrally flattened, both of which characters

are in strong contrast with the adult condition. There is an

internasal groove. The pit beneath the chin shown in text-fig. 1

I now believe to be due to the shrinkage of tissues and not a

normal character, for [ have not observed it in any other speci-

men, nor does it appear in Lohberger’s excellent photographs.

The eyes are well developed. Teeth are prominent in both jaws,

even in the smallest specimens examined by me (350 mm.),“but

Text-figure 1.

Embryo of Zamna. X 2.

1.G., internasal groove; Y.S., yolk-stomach (paunch).

they are devoid of the lateral cusps which are found on the teeth

of the adult. Vestiges of spiracles are usually, but not invariably,

present as minute pores situated midway between the eye and

the first gill-slit; in no case yet examined do they communicate

with the pharynx. Spiracles in the adult are either absent, or,

if present, minute and functionless. The five pairs of large eill-

slits are fully open, and the gills well developed (there are no

traces of external gills). The lateral line is well marked. The

lateral keels, so characteristic of the adult, in the caudal region

are prominent, as is the notch in the back at the base of the

caudal fin. The cloaca is open. The fins are fully developed

and resemble closely those of the adult, excepting the caudal,

whose dorsal and ventral lobes have not yet expanded so that

they present the chelate appearance seen in the figure (at the

166 MR. E. W. SHANN ON THE EMBRYONIC

same time the notch on the inner border of the upper lobe is

distinctly visible).

The ground-colour is slate-grey fading to cream underneath

and upon the paunch. The skin in the younger examples

(350 mm.) is perfectly smooth, but in slightly larger ones (430 to

450mm.) there is a slight roughness due to ‘the developing

scales. The skin passes without interruption from trunk to

paunch (7.e., without the deep crease which the illustration seems

to indicate).

Table II. shows the detailed measurements of a series of

embryos. The outstanding feature of this Table is the clear

Tasre LT.

(All measurements are in millimetres.)

| Trndexa(See yal less) came ees seen eG, H K H K

Meng hh ee ccenscecceeseet teaser nee seo eeren|| PSDO) a) eet 28 ek ota ob asaloUS

Tle arene lon ivasie: ceeteee neces 105 | 186 | 149 | 900 | 911 | —

‘oso Ushort axis ...00.0..1..] 1524) == 94) SOS") MISS mieaesia| =

| |

Diameter of Eye ee SPAS as [Sere lian gre hl eae 12 14

Tip of Tail to Y oneeneen see senses oe5h) 0 r se [SRI fy Se 267 —

Anterior of Cloaca to Root of Tail ...) 95 | — — | 100 | — | +125

| Snout to ist Dorsal Pim ....c..5.....-..|) 125 Kostas | 1381 | 166 178 | 220

| | | | }

| Ist Dorsal to 2nd Dorsal Fin ............) 107 — 123 | 159 | 178 | 2038

| Pectoral to Pelvic Fin ............... 230) WON | ae OM a aA Os als _—

lPelvicito fA nal aRiin utter oeeteiaee eee) SO ne ae 65 | —

: PAG BWC DRRGL oeetaeee haces 25 — 25 44, AS | 57

dst, Borsa BS srerchheee ewe Sealy gh ey cot mca tice2 al ueag

| { |

- os. POPS Genaneraacmeconssvascaoall | 240) Ilys 21 26 30 40

| Begin! Fin | oveth Mtge amt ost tare) SOIA| AAV Al tO in Gy Tuned Re

«a1: ( base (transverse) -........ 10 — 10 11 16 21

eine Fin { Tensth F daedao lca #20 | — | 22 | 299} 99 | 365

Caudal Fin ( upper lobe . Peres et 1 121 | 115 | 145 | 147 | 168

(outer rim) ¢ lower lobe ..........0./..605 45 54 | 54 75 82 85

demonstration that the paunch (due to the presence of the yolk-

stomach) increases in bulk as the fish grows in size; we are

thus forced to conclude that growth at this stage is not due

to the use of reserve material, but to some external cause. The

other measurements, apart from a few discrepancies (accountable

either to normal variation or to individual methods on the part

of different observers), give a very fair representation of normal

growth.

In the male specimen of 350mm. the claspers of the pelvic

fins are manifest even to a casual glance. The fins themselves

showed the same measurements as those of the female twin (F).

DEVELOPMENT OF THE PORBEAGLE SHARK. 167

IV. Internal Structure.

A full account of the internal structure of Zamna embryos

ean be obtained by reference to Lohberger (4) and Shann (6);

thus it will be necessary here to give only a summary of the

outstanding features of the alimentary system, together with

certain corrigenda of previous statements. Text-fig. 2 is

reproduced from Lohberger to illustrate my description of the.

alimentary system.

On dissection the outer skin in the abdominal region is found

Text-figure 2,

_ --6e.

LY O: Vv.

Sp. PY.

Alimentary canal of Zamna embryo (after Lohberger).

an., anus; b.d., bile-duct; c.s., cardiac portion of stomach (“ yolk-stomach ”) ;

oe., esophagus ; py., pylorus ; py.o., origin of pyloric portion; py.s., pyloric

portion of stomach; 7.d., rectal diverticulum; s.z., spiral intestine; sp.,

spleen; v., blood-vessels.

to be underlain by a thin layer of muscle, which is continuous

with the lateral muscle of the trunk. The cavity within is a

normal celom, The greater part of the ccelom is occupied by the

swollen cardiac portion of the stomach, which in an embryo of

350 mm. had a volume of 150 ¢.c.; it is this organ, in fact, which

causes the great ventral distension of the body-wall alluded to

above as the “paunch.” The wall of the cardiac stomach is

richly supplied with blood-vessels. (The statement in my previous

paper that the yolk “‘ found its way into every interstice of the

body-cavity, investing completely the abdominal organs” is, of

168 - MR. E. W. SHANN ON THE EMBRYONIC

course, erroneous; it arose through my having been content to

dissect a specimen whose stomach had already been punctured.)

Anter iorly the cardiac stomach tapers somew hat abruptly to the

wide opening of the esophagus. ‘The latter organ is thick-walled

and its lumen is longitudinally ridged ; it communicates freely

with the pharynx. Leaving the cardiac stomach in the region of

the posterior third and sligh tly on the right side a narrow, but

very muscular, tube runs forwards to enter the anterior end of

the spiral intestine close to the point where the csophagus

debouches ito the stomach. This tube is undoubtedly the

pyloric portion of the stomach ; it opens by a minute tunnel-like

aperture into the cardiac portion, in whose wall it is embedded for

some distance. The lumen, though minute. is nevertheless con-

tinuous from the cardiac portion to the spiral intestine. The

latter organ is fully developed and passes posteriorly into the

short rectum, which bears the characteristic dorsal diverticulum.

The spleen is well developed (this is the “lobed tissue” whose

significance I was unable to determine in my former paper).

The pancreas is smaller and does not appear in the illustration,

since it is situated on the left side. The liver is of large size and

typical shape; it communicates by an apparently functional bile-

duct with the apical intestine, entering it from the ventral aspect

close to the pyloric opening.

In the youngest specimens examined by me (F) the muscles of

the head presented a curious condition. The normal muscles (e. g.,

constrictor, coraco-mandibularis, coraco-hyoideus, and coraco-

branchiales) were recognizable, but instead of being composed

of well-defined masses of compact muscle-fibres, the fibres were

few in number and embedded in a mass of gelatinous non-cellular

material. In older specimens this gelatinous matter gives place

to true fibre ; the process is correlated with considerable skrinkage

in the relative girth of the head of the embryo.

There are five pairs of different branchial arteries, as figured

and described by Lohberger. My previous deseription of six pairs

T now consider to have been due to an error in dissection, for I

have found only five pairs in the specimens examined since my

first dissection was made.

V. Physiology of Nutrition.

The cardiac portion of the stomach (yolk-stomach) contains a

mass of pale yellow-coloured pulp. The latter is finely granular as

seen under the microscope; I failed, however, to find the dumb-

bell-shaped granules described by Lohberger. The pulp is not

uniform in consistency, for intermingled with it are irregular

aggregations of skin-like matter, which Lohberger regards as

portions of egg-membrane; indeed, he states that Swenander

found two entire egg-capsules in the yolk-stomach of a 30cm.

specimen. :

DEVELOPMENY’ OF THE PORBEAGLE.- SHARK. 169

The uterine portion of the parental oviduct is thick-walled and

highly vascular; its inner walls, moreover, are thrown into deep

folds which have a glandular appearance. There is no evidence,

however, that the uterme wall secretes a nutrient material.

The uteri of specimens examined by Dr. Williamson contained,

as he assures me, no substance apart from the embryos them-

selves. Swenander (7) found in the uterus of his specimen forty

pieces of material, which proved to be groups of eggs surrounded

by a common membrane. The portion of both oviducts imme-

diately above the uterus in specimen F contained matter

resembling the contents of the yelk-stomach, the lumen of the

shell-gland was replete with it, and it was also present in the

uppermost portions of the oviducts as far as their source; the

skinny content, as might be expected, was not found above

the level of the shell- alana The oyaries themselves were large

but contained no ripe ova; their contents much resembled in

consistency the matrix in the yolk-stomach, but was darker

in colour owing apparently to the accumulation of blood.

Not only did the matter in the yolk-stomach resemble that in

all parts of the oviduct as regards physical properties, but a

general chemical analysis conducted in the school laboratory

revealed no outstanding difference in this respect.

Similar material was found in the mouth and csophagus of

each of the embryos of the I and H groups.

The only conclusion acceptable on this evidence is that nutrient

material is derived from the ovary in the form of immature eggs

or partially degenerate ovarian tissue—that this is taken up by

the oviduct, 1s partially or completely covered with a thin mem-

brane in the shell-gland, and is passed thence into the uterus,

where it is swallowed by the embryos.

The contents of the spiral intestine are of a uniform semi-fluid

consistency and of a greenish colour; they have, in fact, every

appearance of having undergone the process of digestion. The

rectum also contains greenish matter of a somewhat darker

shade, and very slight compression of the trunk of the embryo

(before dissection) causes this feecal matter to exude from the

cloaca.

These observations seem to show that food supplied by the

ovary is taken into the alimentary system of the embryo through

the mouth and digested in the manner characteristic of free-

living animals. Owing to the sedentary nature of the embryo

there is little wastage of tissue to repair; and, since the supply

exceeds the demand, a vast sur plus of potential food acevmulates

in the cardiac portion of the stomach. Whether all the waste

products of digestion normally accumulate in the intestine of the

embryo until birth, or whether a portion of them is voided into

the maternal uterus requires investigation. If fecal matter finds

its way into the uterus it must be got rid of in some way. The

passage of the uterus to the cloaca is short and wide; it is possible

170 MR. E. W. SHANN ON THE EMBRYONIC

that water may enter and be expelled through this channel.

Such a supposition offers an explanation of why no matter

(whether nutritive or fecal) is found as a rule in the uterus on

capture. It also offers a solution to the problem of respiration ;

for, as has been shown, the gills are fully developed and apparently

functional, The uterine wall is highly muscular, so that a

potential mechanism for pumping water in and out of the uterus

is present. Dr. Williamson, in a letter to me, says: ‘‘ Mr. Ennson

observed that the pregnant female dogfishes (piked dogs) when

brought up in the trawl] had the abdomen distended and sea-water

poured out of the cloaca on to the deck.”

Although the mode of embryonic nutrition described above, so

far as I can ascertain, is unique among Hlasmobranch fishes,

certain observations by Gudger (3) on the Batoids of Beaufort,

N.C., are worth consideration in this connection. Speaking of

Dasyatis say Gudger remarks :—

“The young are found bathed in a substance of the color and

consistence of rich yellow Jersey cream.’

“The older embryos had the large intestine filled with a

chlorine-yellow substance, evidently the milk-like food secreted

by the villi and taken in probably through the spiracles.” In

the younger stages it is taken in by the long external gill-

filaments.

“‘ Notwithstanding the fact that the umbilical cord entered the

alimentary tract at the junction of the small with the large

intestine, and that the material in the anterior part of the large

intestine was lighter in colour than in the middle and hinder

regions, it is reasonably sure that it was not yolk.” The material

is described as ‘finely divided flocculent grading to large plate-

like masses.”

In this Ray and in Pteroplatea maclura when the uteri are

gravid the ovaries are insignificant. ‘The lumina of the ovaries

were filled with “an abundant yolky material which probably

came from the breaking down of some of the ova.”

Gudger does not describe the contents of the upper parts of

the oviducts: the question arises, did they contain any of the

‘“‘yolky material” found m the ovaries? If not, how are we to

account for the “large plate-like masses’ which were found’in

the intestine: surely “they do not come from a purely milky

nutrition? Whether the nutrient matter is taken in through

the spiracles (which in a Ray have sufficiently large openings) or

through the mouth is a minor consideration ; the important

point is that food is actually swallowed and digested, according to

Gudger, instead of being merely absorbed through gill-filaments.

In the Rays, however, the supply does not exceed the demand,

and, consequently, no yolk stomach is produced. The fact that

in these Rays two batches of young are produced each season

precludes the possibility of a protracted intra-uterine develop-

ment such as occurs in Lamna cornubica.

DEVELOPMENT OF THE PORBEAGLE SHARK, 171

Titerature.

. CALDERWooD, W. L.—‘‘ Notes on an Intra-uterine specimen

of the Porbeagle (Zamna cornubica).” Sixth Annual

Report of the Fishery Board for Scotland, p. 263.

. Cotter, R.—Meddelelser om Norges Fiske, 1 Aarene, 1884-

EOI ps Cte

. GupeEr, E. W.—‘‘ Nat. Hist. Notes on some Beaufort, N.C.,

Fishes, No. 1. Elasmobranchii with Special Reference to

Utero-gestation.” Proc. Biol. Soc. Washington, vol. xxv.

pp. 141, 156.

. Lonperenr, J.—‘‘ Ueber zwei riesige Embryonen von Lamna.”

Beitr. zur Nat. Ostasiens, Miinchen, 1910.

. Pennant, T.—Brit. Zool. vol. iii. p. 118.

. SHann, E. W.—“ A Description of the Advanced Embryonic

Stage of Zamna cornubica.” Twenty-eighth Annual

Report of the Fishery Board for Scotland, p. 73.

. SWENANDER, Gusr.—‘“‘ Ueber die Ernihrung des Embryos der

Lamna cornubica.”’ Zool. Stud. Festschr. fur Tullberg,

Upsala, 1907.

muh

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A a A)

NOVES ON THE PAIRING OF THE LAND-CRAB. es}

15. Notes on the Pairing of the Land-Crab, Cardesoma

armatum. By Miss L. EH. Cunzsmay, I.E.S., F.Z.8.,

Curator of Insects to the Society.

[Received March 5, 1923: Read March 6, 1923.]

As nothing is recorded of the life-history of this species, it

was desirable to encourage the pairing of those specimens under

observation in the Insect House. Few females, however, have

been collected. Of the three consignments of these crabs kindly

sent to us by Captain Arpzitage from the neighbourhood of the

mouth of the River Gambia, the first, received in September

1921, consisted of nine males; the second, received in March

1922, consisted of fourteen males, and two small females which

died in less than a fortnight. The third consignment, received

in July 1922, consisted of thirteen males and two females, one

small and one of medium size.

The larger female was placed with a male in a separate bay

containing sand and coarse gravel a foot and a half in depth, and

a tank of fresh water. The male dug a vertical burrow in one

corner, and the female dug a horizontal tunnel in the centre of

the bay. The male fed in the daytime, but the female was

rarely seen until an hour after closing-time. Both crabs took

boiled rice, boiled and raw potatoes, and dry leaves; the male

(but not the female) fed also on pieces of raw meat and fish.

In the first week of August the female’s tunnel had a second

opening on the side nearest the male’s burrow, and for the next

fortnight they shared the tunnel: neither of the crabs were seen

in the daylight during that time, unless disturbed. At the end

of a fortnight the male dug a new burrow, and was not after-

wards found in the female’s burrow.

On August 26th the male was removed, and two tanks were

placed in the bay, one of fresh water, the other of sea-water

renewed weekly. The female only emerged at night after that

date. All the females we have had under observation have been

reluctant to show themselves above ground before dusk; this

probably accounts for their being less easy to procure than the

males. This female came out every three or four nights to feed

and bathe, but any movement, or the switching on of the electric

lights, caused her to retire to her burrow; if she was out on the

other nights she did not feed, for the food was untouched. She

was only twice seen in the sea-water tank. If she did not

appear for three days the burrow was opened, otherwise she

was not disturbed.

On September 18th, 1922, the crab was discovered to be in

spawn.

On September 22nd a portion of the eggs was severed from

her and sent to Professor McBride at the Imperial College of

Science. The crab cast the rest of the eggs that night.

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THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 17

EXHIBITIONS AND NOTICES.

February 6th, 1923.

Sir S. F. Harmer, K.B.E., F.R.S., Vice-President,

in the Chair.

The Srcrerary read the following Report on the Additions to

the Society’s Menagerie during the months of November and

December, 1922 :—

NOVEMBER.

The registered additions to the Society's Menagerie during the

month of November were 53 in number. Of these 31 were

acquired by presentation, 6 were deposited, 8 were purchased,

1 was received in exchange, and 7 were born in the Menagerie.

The following may be specially mentioned :—

1 Cheetah (Cynelurus jubatus), from Northern Nigeria,

presented by John Holt & Co. on November 21st.

3 Spotted Cavies (Celogenys paca), from 8. America, purchased

on November 17th.

1 Great Kangaroo (Macropus giganieus), from Australia,

presented by Capt. Daniel and Mr. Urquhart on November 27th.

1 Snowy Owl (Vyctea scandiaca), captured at sea, 300 miles

from Cape Race, deposited on November 23rd.

A Scarlet Ibises (Hudocimus ruber), from South America,

purchased on November 20th.

DECEMBER.

The registered additions to the Society’s Menagerie during the

month of December were 55 in number. Of these 28 were

acquired by presentation, 13 were deposited, 11 were purchased,

1 was received in exchange, and 2 were born in the Menagerie.

The following may be specially mentioned :—

1 African Rhinoceros (Rhinoceros bicornis), 2, from Arusha,

Tanganyika Territory, purchased on December 15th.

1 Wild Boar (Sus scrofa), 9, and 1 Gmelin’s Sheep (Ovis

orientalis), 3, from Mesopotamia, presented by Major Dickson,

C.1.E., on December 6th.

2 Persian Gazelles (Gazella subgutturosa), 2 9, and 1 White-

tailed Sea-EHagle (Haliaétus albicilla), from Mesopotamia, pre-

sented by Sir Percy Cox, K.C.M.G., K.O.S.1., and Capt.

Cheesman, C.M.Z.S., on December 6th.

1 Blue-eyed Cockatoo (Cacatua ophthalmica), from New Britain,

received in exchange on December 19th.

176 LORD ROTHSCHILD ON A MOUNTAIN GORILLA.

Lord Roruscuizp, F.R.S., F.Z.8., exhibited a fine adult male

Mountain Gorilla (Gorilla gorilla beringert Matschie), on behalf

of Messrs. Rowland Ward, Ltd. He said :—

‘Since my notes on Anthropoid Apes in the P.Z.8. 1904,

413, our knowledge of Gorillas has increased consider-

ably. At that time the only material of the Mountain Gorilla

was the type-skull, and of the remaining races much more

material has come to hand, and it is therefore possible definitely

to acknowledge three distinct subspecies, or local races, showing

decided structural—especially cranial—differences. Three other

so-called species have been separated, but insufficient material

prevents any final decision on at least two of these.

“ The specimen exhibited, together with a much older solitary

male now in the Tring Museum and an adult female, were

obtained by Mr. T. A. Barnes at between 6000 and 10,000 feet

on the volcanoes near Lake Kivu while collecting butterflies for

Mr. J. J. Joicey. In addition to these, there are in England at

this moment an adult male obtained by a Belgian officer and an

adult male, female, and a baby obtained by the brothers Foster.

“The three following subspecies. or local races, are well defined

and distinct :—Gaboon Gorilla (Gorilla gorilla gorilla Savage &

Wyman); Cameroon Gorilla (Gorilla gorilla dichli Matschie) ;

Mountain Gorilla (Gorilla gorilla beringeri Matschie). The

Gaboon and Cameroon Gorillas, as distinguished from the

Mountain Gorilla, are dimorphic, 7. e. they have a black and a red

phase; in the Gaboon race the red phase does not differ in colour

on the body, but has the whole crown chestnut rufous, whereas

the Cameroon race has a rufous phase in which the red crown is

less sharply defined, and the body-colour brown or more mixed

with rufous hairs. ‘These rufous phases have been described

respectively as distinct species or races as follows: Gaboon race,

Gorilla castaneiceps Hack., and the Cameroon race Gorilla gorilla

matschiei Rothsch.; they must, however, stand as Gorilla gorilla

gorilla form. dimorph. castaneiceps, and Gorilla gorilla diehli

form. dimorph. matschiet.

“The Mountain Gorilla is at once distinguishable externally

from the two other races by the much stouter and more stocky

build, by the much thicker pelage, by the intense shining black

of the hair, and by the large fleshy callosity on crest on the top

of the head. ‘This crest was first noticed by My. Barnes, and his

photographs of the animal in the flesh were the first intimation

to systematists of this peculiarity. This callosity is similar in

its nature to the cheek callosities of the Orang Outan, but unlike

these, appears to be common to all adult males, and not a sign of

senile impotence as in the Orang.

“The most essential differences of the three races are, however,

in the skulls. G. g. gorilla has the occipital region narrow and

appearing almost triangular, owing to the lambdoidal crest

running up to a sharp point in the centre. G. g. diehli has the

occipital region very broad, and the lambdoidal crest in the

MR, OLDFIELD THOMAS ON A ROCK KANGAROO. 177

centre only rises to a low blunt point. Lastly, G. g. beringeri

has the occipital region very broad, and the lambdoidal crest is

quite flat and horizontal to support the fleshy callosity.

“ The remaining two races Gorilla jacobsi Matschie and Gorilla

manyemda (Alix and Bouvier) are more than doubtful, but the

material is too scanty to decide definitely. Matschie and Oscar

Neumann have definitely applied the name manyema to the low

country Gorilla of the Congo and, as far as I can see, there are

no osteological characters to separate this race from the typical

Gaboon race; the two or three adult males examined, however,

appear to have the pelage brighter in colour and more sharply

contrasted, Of jacobs: only the type male is known, and the

extremely sharp facial angle of the skull may be an anomaly.

“Adult males of the three well-defined races vary in height

from 5 ft. to 6 ft., and there is no specimen preserved over 6 ft.

in height, but in the ‘Illustration’ for February 14th, 1920,

p. 129, is a photograph of a gorilla 9 ft. 4 in. in height, according

to M. Villars-Darasse, and the photograph certainly shows a

gigantic animal. This individual is said to have been killed in

the Forest of Bambio, Haute-Lobaze.”

Mr, Ouprietp THomas exhibited a new Rock-Kangaroo

(Petrogule) which had been obtained in Northern Queensland by

My. T. Sherrin, the collector employed on behalf of the Godman

Exploration Trustees, by whom it had been presented to the

National Museum. It was described as follows:

PETROGALE GODMANI, Thos.*

General characters about as in P. assimilis Ramsay, of which

a fine series from Inkerman, N. Queensland, was available for

comparison. Black axillary patch less extensive. Forearms

more strongly buffy. Tail, instead of being black for its terminal

third or half, drabby whitish, its basal fourth only grizzied with

black, the rest dull whitish to the end. In a young specimen

there was an indication of the upper side being darker, though

not so dark as in assimilis, but in the adult even this was absent.

Skull of the same length as that of assimilis, but it was more

convex in the frontal region, and the nasals were decidedly

broader and heavier, especially anteriorly, where they had not

the marked narrowing found in assimilis. Teeth as in assimilis

except that the secater was larger, 7-8 mm. in length as com-

pared to a nearly uniform length of about 6-7 mm. in six

specimens of assimilis.

Dimensions of the type, measured in flesh :-—

Head and body 465 mm.; tail 502; hind foot 140; ear 59,

Skull, greatest length 101; condylo-basal length 92; zy g0-

matic breadth 52; nasals, length 41, anterior breadth 8°5,

posterior breadth 14:5; teeth, length of 7 4, secator 7:8,

Gils AOS

* Abstr. P.Z.S, 1923, No. 235, p. 13.

Proc. Zoor, Soc.— 19238, No, XII. 12

178 THE SECRETARY ON ADDITIONS TO THE MENAGERIE.

Hab. North Queensland. Type from the Black Mountain,

16 miles 8.W. of Cooktown.

Type. Adult male. B.M. No. 23.1.5,19. Original number 196.

Collected July 20th, 1922, by T. V. Sherrin. Two specimens,

adult and young.

This Rock Kangaroo was readily distinguishable from P. asst-

milis by its whitish tail, broader nasals, and larger secator.

Mr. Thomas expressed his pleasure at being able to bring

forward so fine a discovery as one of the first-fruits of the

Godman Exploration Fund, which had been recently founded by

Dame Alice Godman in memory of her husband, Dr. F. Ducane

Godman, F.R.S., the well-known naturalist.

The Trustees of the Fund were now to be a permanent body,

with their headquarters at the British Museum (Natural History),

and would be glad to take charge of any further funds which

persons wishing to benefit the National Museum might give or

bequeath to them. Such funds would always be administered

under the names of, and in accordance with the wishes of, the

donors.

Mr. Tuomas also exhibited the skull of a Pygmy Fruit-Bat

from Sumatra, upon whicn he had recently founded a new

genus (Ann. Mag. N. H. (9) xi. p. 251, Feb. 1923). The generic

name had since proved to be preoccupied, and he therefore

now proposed to substitute for it that of Mthalops. The type-

species would thus bear the name of Zthalops alecto.

Mr. J. B. Scrtvenor, M.A., exhibited, and made remarks

upon, a photograph showing the method adopted by Malay

natives in breaking in recently-captured Hlephants.

Mr. BK. G. Boutencer, F.Z.S., gave an account of his recent

visit to Vienna, and of the experiments carried on there by

Dr. Kammerer and others upon Amphibians and Insects.

February 20th, 1923.

Dr. A. SmirH Woopwarp, F.R.S., Vice-President,

in the Chair.

The Secretary read the following Report on the Additions te

the Society’s Menagerie during the month of January, 1923 :—

The registered additions to the Society’s Menagerie during the

month of January were 156 in number. Of these 55 were

acquired by presentation, 15 were deposited, 85 were purchased,

and 1 was born in the Menagerie.

The following may be specially mentioned :—

1 Takin (Budercas taxicolor), 9, from Bhutan, presented by

Major F. M. Bailey, C.I.E., F.Z.8., on January 25th.

_ MELANISM IN TIPPELSKIRCH'S GIRAFFE. 179

1 Spectacled Bear (Zremarctos ornatus), from the Andes of

Peru, presented by H. Fox Strangeways, Hsq., on January 16th.

1 Persian Squirrel (Sciurus persieus), new to the Collection,

from Mosul, presented by Major Dickson, C.1.E. .

1 Lesser Niltava (Viliava macgrigorie), from the Himalayas,

new to the Collection, presented by E. W. Harper, F.Z.S., on

January 24th.

A large collection of N. American reptiles, including 2. Helo-

derms, 2 Confluent Rattlesnakes, 2 Moccasins, a Copperhead,

and a Three-lined Snake (Atractus trilineatus), from Trinidad,

new to the Collection, purchased on January 25th.

Mr. D. Seru-Smurru, F.Z.8., exhibited a number of skins of

Birds-of-Paradise and gave an account of the various forms of

sexual display, drawing special attention to the display of the

Magnificent Bird-of-Paradise (Diphyllodes magnifica hunsteini),

a living specimen being now in the Society’s collection of tropical

birds. i f

A Cinematograph record of the life-history of the House-fly,

taken under the direction of Prof. H. M. Lurroy, F.Z.S., was

exhibited. ee ag

The Srcrerary exhibited a photograph of the Polar Bears

“Sam” and “ Barbara.” he

Mr. R. I. Pococr, F.R.S., F.Z.S., exhibited, and made remarks

upon, a series of drawings of the feet and nose of the Polar Bear.

March 6th, 1923.

Sir S. F. Harmer, K.B.E., F.R.S., Vice-President,

in the Chair.

The Secretary read the following note from Mr. Caldwell, of

the Game Warden’s Office. Nairobi :—

‘“‘' You may be interested to hear of a case of apparent melanism

in Tippelskirch’s Giraffe, Girafia camelopardalis tippelskirchi.

The animal, a bull, was recently seen in the South Game Reserve;

its markings are said, by a competent Huropean observer, to

have shown through the black in the faint manner that the

rosettes show in a black leopard. Wakamba and Masai natives

who saw it were much excited, and said they had never seen

such a thing before.”

Mr. R. I. Pococs, F.R.S8., F.Z.8., exhibited, and made remarks

upon, a series of drawings of the feet and spurs of the Echidna.

180 ON THE STOMACH CONTENTS OF A CROCODILE.

In the absence of Sir G. Abercromby, Mr. R. I. Pocock

exhibited the mounted head of a Kob from East Africa.

Prof, E. W. MacBrips, F.R.S., F.Z.S., exhibited, and made

remarks upon, a series of photomicographs of sections through

the nuptial callosities of Frogs.of the genera ana and Alytes.

Mr. GC. F. M. Swynyzrton, C.M.Z.S., exhibited, and made the

following remarks upon, the stomach-contents of a Crocodile :—

«The crocodile, the contents of the stomach of which are

shown in the photograph (text-fig. 1), was shot far up the Duma

River, flowing into the Speke Gulf of the Victoria Nyanza from

the south, on June 6th, 1922.

“Natives came to me to complain of a notorious crocodile, that

had taken many people, and, as it was several miles out of my

way, I sent a scout to deal with it. According to his own

statement, the crocodile made an attempt on himself as he sat

beside the water, but, at any rate, he shot it, and ib was carried

Text-figure 1.

Stomach-contents of a Crocodile.

whole into my next camp by a large number of natives. It

measured only about twelve feet in length, but was of astounding

girth. The objects shown in the photograph, all of which were

taken by me, or in my presence, from the stomach of this one

crocodile, were some heavy rings, some lighter bracelets, a blue

bead necklace, a piece of dark cord, the tortoiseshell from the

carapace of a tortoise (the bones having apparently been digested

first), and the quills of a porcupine (eaten last) and bones.

Some of the quills still lay in the crocodile’s throat, though

most appeared to have been swallowed with the porcupine.”

No. 235.

ABSTRACT OF THE PROCEEDINGS

OF THE-

ZOOLOGICAL SOCIETY OF LONDON.*

February 6th, 1923.

Sir 8. F. Harmer, K.B.E., F.R.S., Vice-President,

in the Chair.

The Srecrerary read a Report on the Additions to the Society’s

Menagerie during the months of November and December 1922.

Lord Roruscuitp, F.R.S., F.Z.8., exhibited an adult male

Gorilla gorilla beringert from near Lake Kivu, and further

illustrated his remarks by a series of lantern-slides.

Mr. OupFrieLD THomas, F.R.S., F.Z.8., exhibited a new Rock-

Kangaroo, which he diagnosed as follows :—

Petrogale godmant, sp. a.

Like P. assimilis, but with a whitish tail, broader nasals, and

larger secator. j

Hab. Black Mountain, near Cooktown, N. Queensland.

Type. B.M. No. 23.1.5.19. Presented by the Trustees of

the Godman Exploration Fund.

Mr. Thomas explained to the Meeting about the Godman

Exploration Trust, which had been founded in memory of her

husband by Dame Alice Godman, and from which it was hoped

much benefit would be gained by the Museum.

* 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. THomas also exhibited the skull of a Pygmy Fruit-Bat

from Sumatra, upon which he had founded a new genus. He

proposed to substitute Wthalops for the preoccupied name

thalodes, which he had inadvertently given to the new genus

of Fruit- Bats described in the ‘Annals aud Magazine of Natural

History’ for February.

Mr. J. B. Scrivenor, M.A., exhibited, and made remarks upon,

a photograph showing the method adopted by Malay natives in

breaking in recently captured Elephants.

Mr. U. A. Aparr Dicuton, M.B., F.R.C.8., F.Z.8., gave a

résumé of his paper on ‘‘ Coat-colour in Greyhounds.”

Mr. EK. G. Bounencer, F.Z.8., gave an account of his recent

visit to Vienna and of the experiments carried on there by

Dr. Kammerer and others upon Amphibians and Insects.

In the absence of the Author, Professor Watson, F.R.S., F.Z.S.,

gave an account of a paper by Mr. E. Leonarp Ginn, M.Sc.,

entitled ‘* The Permian Fishes of the Genus Acenérophorus.”

The following communications were taken as read :— CHARLES

F. Sonnrac, M.D., F.Z.S., ‘On the Vagus and Sympathetic

Nerves of the Terrestrial Carnivora”; Epwarp Pairs ALLIs,

Jv., F.R.M.LS., F.Z.8., “The Postorbital Articulation of the

Palato-quadrate with the Neurocranium in the Celacanthide ” ;

GeorGE 8. Giexion1, M.D., “On the Linguatulid Arachnid,

Raillietiella furcocerca (Diesing, 1835), Sambon, 1922”; The late

Mrs. Rrra MarxksREITER, B.Sc., “Some Microfilaria found in the

Biood of Birds dying in the Zoological Gardens, 1920-1922.”

The next Meeting of the Society for Scientific Business will be

held on Tuesday, February 20th, 1923, 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 1923.

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

Exhibition of a Cinematogiaph film of the House-fly.

Dr. N. 8. Lucas, F.Z.S.

Reports on the Deaths which have occurred in the Society’s

Gardens during 1922.

Prof. Erxar LoONNBERG, F.M.Z.S,

Remarks on some Palearctic Bears.

KE. W. Swann, B.S8c., F.Z.S.

The Embryonic Development of the Porbeagle-Shark,

Lanna cornubiea.

Roserr Gurney, M.A., F.Z.S.

Some Notes on Leander longirostris, M.-Edwards, and other

British Prawns.

The following Papers have been received :—

Cuas. F. Sonntac, M.D., F.Z.8.

The Comparative Anatomy of Tongues of the Mammalia.—

VIII. Carnivora.

R. Kirrxratricr, F.Z8.

On a new Species of the Tunicate Rhizomolgula with

remarkable Sensory Organs. No. 24. Results of the Oxford

University Expedition to Spitzbergen, 1921.

T. H. Rive.

The Elephant-Seals of Kerguelen Land.

SusHit Cu. Sarkar, F.Z.8.

= ee

A Comparative Study of the Buccal Glands and Teeth of the

Opisthoglypha, and a Discussion on the Evolution of the Order

from Aglypha.

H. G. Cannon, B.A., F.Z.S.

A Note on the Zowa of a Land-Crab, Cardisoma armatum.

JosEPH ConrAD CHAMBERLIN.

A Systematic Monograph of the Tachardiine or Lac Insects

(Hemiptera——Coccide).

Ouprivtp Tomas, F.R.S., F.Z.8., and M. A. C. Hivton.

On the Mammals obtained in Darfur by the Lynes-Lowe

Expedition.

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

(1) On the External Characters of Hlaphurus, Hydropotes,

Pudu, and other Cervide.

(2) The Classification of the Sciuride.

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 SocigEry oF Lonpon,

Recent’s Park, Lonpon, N.W. 8.

February 13th, 1923.

No. 286.

ABSTRACT OF THE PROCEEDINGS

ZOOLOGICAL SOCIETY OF LONDON.*

February 20th, 1923.

Dr. A. SmrrH Woopwarp, 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 January 1923.

Mr. D. Sern-Surrn, F.Z.S., exhibited a number of skins of

Birds-of-Paradise and gave an account of the various forms

of sexual display, drawing special attention to the display of the

Magnificent Bird-of-Paradise (Diphyllodes magnifica hunsteint),

a living specimen being now in the Society’s collection of tropical

birds.

A Cinematograph record of the life-history of the House-fly,

taken under the direction of Prof. H. M. Lrrroy, F.Z.S., was

exhibited.

The Secrerary exhibited a photograph of the Polar Bears

‘Sam and Barbara.” ;

Me. R. I. Pocock, F.R.S., F.Z.8S., exhibited, and made remarks

upon, a series of drawings of the feet and nose of the Polar Bear.

Dr. N. S. Lucas, F.Z.S., communicated his Report on the

Deaths which occurred in the Society’s Gardens during 1922.

* 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 2f Sta Shillings per annum, payable in advance.

The following papers were taken as read :—Prof. Einar Lonn-

BERG, F.M.Z.S., ‘‘ Remarks on some Palearctic Bears”; E. W.

Suann, B.Sc., F.Z.8., “The Embryonic Development of the

Porbeagle-Shark, Lamna cornubica”; Ropery Gurney, M.A.,

F.Z.S., ‘“‘Some Notes on Leander longirostris, M.-Kdwards, and

other British Prawns.”

The next Meeting of the Society for Scientific Business will be

held on Tuesday, March 6th, 1923, at 5.30 p.m., when the

following communications will be made :—

H. G. Cannon, B.A., F.Z.S.

A Note on the Zoxa of the Land-Crab, Cardisoma armatum.

Miss L. EH. Corrsman, F.E.S., F.Z.S.

_. Notes on the Pairing of the Land-Crab, Cardisoma armatum.

Cuas. F. Sonnraa, M.D., F.Z.S.

The Comparative Anatomy of Tongues of the Mammalia.

VIII. Carnivora.

R. Kirkpatrick, F.Z.S.

On a new Species of the Tunicate Rhizomolgula with

remarkable Sensory Organs. No. 24. Results of the Oxtord

University Expedition to Spitzbergen, 1921.

T. H. Rivne.

The Elephant-Seals of Kerguelen Island.

The following Papers have been received :—

SusHit Cu. Sarkar, F.Z.8.

A Comparative Study of the Buccal Glands and Teeth of

Opisthoglyph Snakes, and a Discussion ‘on the Evolution

of the Order from Aglypha.

OLpFIELD Tuomas, F.R.S., F.Z.S., and M. A. C. Hinton, F.Z8.

On the Mammals obtained in Darfur by the Lynes-Lowe

Expedition.

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

(1) On the External Characters of Hlaphurus, Hydropotes,

Pudu, and other Cervidee.

(2) The Classification of the Sciuride.

JosEPH CoNRAD CHAMBERLIN.

A Systematic Monograph of the Tachardiine or Lac Insects

(Hemiptera— Coccide).

Prof. K. Kostanscxi, M.D., LL.D.

Ona Remnant of the Omphalo-mesenteric Arteries in the

Manatee.

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 LonpDon,

Recent’s Park, Lonpon, N.W. 8.

February 27th, 1923.

aie ie

ae lice

patsy

No. 237.

ABSTRACT OF THE PROCEEDINGS

ZOOLOGICAL SOCIETY OF LONDON.*

March 6th, 1923.

Sir 8. F. Harmer, K.B.E., F.R.S., Vice-President,

in the Chair.

The Secrerary read a note from Mr. Caupwetu of the Game

Warden’s Office, Nairobi, on a case of apparent melanism in

Tippelskirch’s Giraffe (Giraffa camelopurdalis tippelskircht).

Mr. R. I. Pocock, F.R.S., F.Z.S., exhibted, and made remarks

upon, a series of drawings of the feet and spurs of the Echidna.

In the absence of Sir G. ABpeRrcromsy, Mr. R. I. Pocock

exhibited the mounted head of a Kob from East Africa.

Prof. EH. W. MacBrips, F.R.S., F.Z.S., exhibited, and made

remarks upon, a series of photomicrographs of sections through

the nuptial callosities of Frogs of the genera Rana and Alytes.

Mr. F. M.Swynnerton, C.M.Z.S., exhibited, and made remarks

upon, the stomach-contents of a Crocodile.

Mr. H. G. Cannon, B.A., F.Z.S., communicated his paper ‘‘ A

Note on the Zowxa of the Land-Crab, Cardisoma armatum.”

* 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 Sir Shillings per annum, payable in advance,

Miss L. E. Cusssman, F.E.S., F.Z.S., communicated her

paper “Notes on the Pairing of the Land-Crab, Cardisoma

armatum.”

Dr. Cuas. F. Sonnac, F.Z.S., gave a résumé of his paper on

“ The Comparative Anatomy of the Tongues of the Mammalia.—

VIII. Carnivora.”

Mr. W. P. Pycrart, F.Z.S., gave a résumé of Mr. T. H. Rine’s

paper on “ The Elephant-Seals of Kerguelen Land.”

Mr. R. Kiregparricx, F.Z.S., communicated his paper “‘ On

the Tunicate Rhizomolgula globularis Pallas. No. 24. Results

of the Oxford University Expedition to Spitzbergen, 1921”*.

The next Meeting of the Society for Scientific Business will be

held on Tuesday, March 20th, 1923, at 5.80 p.m., when the

following communications will be made:—

Susan Ca. Sarkar, F.Z.S.

A Comparative Study of the Buccal Glands and Teeth of

Opisthoglyph Snakes, and a Discussion on the Evolution

of the Order from Aglypha.

OLDFIELD Tuomas, F.R.S., F.Z.S., and M. A. C. Hinton, F.Z.S.

On the Mammals obtained in Darfur by the Lynes-Lowe

Expedition.

R. I. Pococs, F.R.S., F.Z.8.

(1) On the External Characters of Hlaphurus, Hydropotes,

Pudu, and other Cervide.

(2) The Classification of the Sciuride.

* [Title changed from that announced in Abstract No. 236.—Ep1rTok. ]

The following Papers have been received :—

JosePH CoNRAD CHAMBERLIN.

A Systematic Monograph of the Tachardiine or Lac Insects

(Hemiptera— Coccide).

Prof. K. Kosranrcxk1, M.D., LL.D.

On a Remnant of the Omphalo-mesenteric Arteries in the

Manatee.

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.

March 13th, 1923.

10.

Wale

12.

13.

14.

15.

PAPERS.

. Coat Colour in Greyhounds. By Aparr Dranton, F.R.O.S., F.Z.S........022-0.00,

. A Note on the Zosea of a Land Crab, Cardisoma armatum. By H. Grauam Cannon,

B.A., F.Z.8. (Text-figures 1-6.)

CC eC Ce ee i ay

. On the Linguatulid Arachnid Raillietiella furcocerca [Diesing, 1885] Sambon, 1922.

By Grores 8S. Gieniou, M.D. Pisa

Ce er ee ee cy

. The Permian Fishes of the Genus Acentrophorus. By BE. Lronarp Ginn, M.Sc.

(TLext-figures 1-16.)

. The Postorbital Articulation of the Palatoquadrate with the Neurocranium in the

Celacanthid Fishes. By Epwarp Pusurs Au.is, Jr., F.Z.8..... 2.220. ce ce eens

. Some Microfilariz found in the Blood of Birds dying in the Zoological Gardens,

1920-1921. By Riva Marxsrmiter, B.Sc., Research Assistant in the Helmintho-

logical Department of the London School of Tropical Medicine. (Text-figures 1-6.).

. On the Vagus and Sympathetic Nerves of the Terrestrial Carnivora. By Cuarues F.

Sonnraa, M.D., F.Z.8., Anatomist to the Society. (Text-figures 1-14.)

. Remarks on some Palearctic Bears. By Ernar Lonnper@, F.M.Z.S.,&c. (Plates L., II.)

. Some Notes on Leander longirostris M. Edwards, and other British Prawns. B

Rosert Gurney, M.A., F.L.S., F.Z.S. (Text-figures 1-6.) ........--2. 22-0 Be

Report on the Deaths which occurred in the Society’s Gardens during 1922. By N.S.

hucas, MiB; B-Z:S85 Pathologist to the Society... )...2..2 cece secre» cosecece

The Comparative Anatomy of the Tongues of the Mammalia.—VIII. Oarnivora. By

Cuaruss F, Sonntac, M.D., F.Z.S., Anatomist to the Society. (Text-figures 15-24.) .

Notes on the Tunicate Rhizomolgula globularis Pallas. By R. Kirxrarricx, F.ZS8.

(Plated: )egsse my creee hot cote aetiers dat enamote ny gsc ron rater Rok aain Mono,

On the Feet and Rhinarium of the Polar Bear (Thalarctos maritimus). By R. I.

Pocock, B.R:S:, F428. (Textefigure l.).c6s5. 6 ise sc i ctsaccee. cbckevenaten

The Embryonic Development of the Porbeagle Shark, Lamna cornubica. By BE. W.

Suid, Cone Ih ret) (Clerdiscetnyas Iles) Goes Sonecconeaung docuec on ddodueucer

Notes on the Pairing of the Land Crab, Cardisoma armatum. By Miss L. E.

@RWESMAI, TH Si ZiS Stan clcterees etter operon sysascv'iaeareve ct cre gmnyehesiale a, sone, naeeetnr a atlanta eat ote

Page

129

155

173

ONG ES TOW PALA S.

19238, Part I. (pp. 1-180).

‘ Page

Einar Loynsura: Pls. 1-II. Palearctic Bears -........ ........ 85

R. Kirxratrick: PI. 1. Rhizomolgula globularis Pallas ..... 155

NOTICE.

The ‘ Proceedings’ for the year are issued in four parts, paged consecutively,

so that the complete reference is now P. Z. 8. 1922, p.... The Distribution

is usually as follows :—

Part I. issued in March.

TT. Sone thane’

Seo wl Ait be 3 September.

OEE ee December.

‘Proceedings,’ 1922, Part LV. (pp. 887-1276), was published on February

13th, 1923.

The Abstracts of the ‘ Proceedings,’ Nos. 235-237, are

; contained in this Part. :

The dates of Publication of ‘Proceedings’ 1830-1858 will be found in the

* Proceedings’ for 1893, page 436.

Tho dates of Publication of ‘ Transactions’ 18338-1869 will be found in the

‘ Proceedings’ for 1913, page 814.

Wey

PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCIETY

OF LONDON,

1923. |

PART II. . |

conTaINiInG Paces 181 ro 481, wita 9 PLaTEs aND

98 ‘l'Ex1-FIGURES.

ee PC 7 1093

PRINTED FOR JHE SOCIETY, ,

oF Bs.

SOLD AT ITS HOUSE GkNirs

LONDON:

MESSRS. LONGMANS, GREEN, AND CO.,

PATERNOSTER ROW. :

[Price Twelve Shillings. |

“LIST OF CONTENTS.

1923, Part II. (pp. 181-481).

EXHIBITIONS AND NOTICES.

Page

THE Szcrutary. Report on Additions to the Society’s Menagerie during the month of

Ee BQ QB” Mad 16 sh ety ph ataaave etistere oases ts be ce era e tal che eee ene eRe Gen ane 479

Mr. F. Martin Duncan, F.R.M.S., F.Z.8. Exhibition of a Caterpillar unrecten with the

Entomogenous fungus Condeyceps movertsiii ys ose its eae tee peer yee 479

Mr, M. A.C. Hinton, F.Z.S. Exhibition of skin of a Lioness .......-.-- + .ccaeecees 479

Tz Secrerary, Exhibition of photographs of deep-level reservoirs of the Society’s new

aA] AR LUNIA’ presse "ait a a Cheilensy pe etle Slate ate eee Sip amt aiden Spies Acie Baan (eats te ane eR eR

Mr. G, C, Rosson, M.A., F.Z.S. Exhibition of the Snail Planorbis dufouri Graells .... 480

Dr. G. M. Vevurs, F.2Z.S. Account of his recent visit to Zoological Gardens in Holland

anciebsel ritnnien sahara eee wets ee eveter boise shah iaie ere RIG A NSO Seeo as dd Sts 480

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

AVERT Chis M26 iwi citays air clo utswis oe) emotes i shee iratretat ot crete fain a alinfes @ Sar tat nies a cr 480

Capt. R. B. Murray, F.G.S., F.R.G.8. Exhibition of living specimens of a Giant

Centipede from Trinidad ............... Mialelstexcpde sins hateile el-eke initia reacts Cele yorciome etree 480

Lt.-Col, 8. Moncxron Copeman, F.R.S., F.Z.S., and Maj. E. E. Ausren, D.S.O., F.Z.S,

Exhibition of a rare British Ti Gee OL revere sah cararena un laieeer adele ar eqanter cnc tolertte eoee 481

PAPERS.

Page

16. On the External Characters of Elaphus, Hydropotes, Pudw, and other Cervide. By

Real Pocotsst RS, EZ.S, (Lextsoures: 2-17,)) eae en eee eI OIL

17. The Classification of the Sciuride. By R. I. Pocock, F.RS., F.ZS. (Text

SHES VC ZO )\iin crepe sve secre sier serene etal gcceenet whee siete) sreMteme te Sere torsraies Siocs |aim'atatata 209

Contents conitsnued on page 3 of Wrapper,

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EXTERNAL CHARACTERS OF ELAPHURUS, EC. 181

16. On the External Characters of Elaphurus, Hydropotes,

Pudu, and other Cervide. By R. I. Pocock, F.R.S.

[Received February 5, 1923: Read March 20, 1923. ]

(Text-figures 2-17.)

CONTENTS.

Page

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Classification of the Cervidee... 2.0.0.0. -...s sees 206

Introduction.

In a paper “On the Specialised Cutaneous Glands of Rumi-

nants” (P. Z.S. 1910, pp. 840-986) I described in some detail these

glands in a considerable number of genera and species of

Cervide, pp. 939 to 967 of the memoir being devoted to this

family. Several well-marked forms, however, were not at that

time available for examination, Hlaphurus, Hydropotes, and Pudu

being three of the most important. Since that time examples of

Elaphurus and Pudu, exhibited in the Zoological Gardens, came

after death into my hands in the Society’s Prosectorium ; and the

Duke of Bedford, knowing my interest in the subject, very kindly

sent tome from Woburn the body of a freshly-killed male example

of Hydropotes. Iam also greatly indebted to His Grace for the

chance to examine a second specimen of Hlaphurus, the body of

which he was good enough to send to me. I must also take this

opportunity of thanking Major A. Pam, O.B.E., for securing

for the Society at my special request the example of the Pudu

on one of his trips to Chili. To the subjoimed account of the

external characters of these three rare and isolated genera of

Cervide, [ have added descriptions of some less interesting

species which I had not actually examined previously, although

the characters of related species belonging to the same genera had

been recorded by me. The following paper therefore is supple-

mentary to the one I publishedin 1910. To facilitate comparison

between the two papers, I have in the following pages quoted

after each specific heading the page in my previous memoir where

the genus or species was referred to.

I have also included in this paper some notes on the rhinarium,

ears, facial vibrissee, and penis, which I did not describe in 1910,

Proc. Zoou. Soc.— 1923, No. XIII. 13

182 MR, R. I, POCOCK ON THE EXTERNAL

Finally, I must express my regret that, when writing on

the glands in 1910, I inadvertently omitted to consult and

quote Caton’s valuable work on the deer of North America.

In thig he fully described the glands and other external

characters of the species of that country, including those now

known as Rangifer tarandus, Alces alces, Odocoileus virginianus,

O. columbianus, O. hemionus, and O. acapulcensis, the last-

mentioned being regarded at the present time as a subspecies of

O. virginianus.

To Caton also we owe, I believe, the discovery of the very

remarkable fact that in Odocoileus the sheath of the penis, instead

of being confluent with the skin of the abdomen, so that the

short prepuce is situated about halfway between the groin and

the umbilicus, is long, pendulous, quite free from the skin of

the groin, except at its base, which is close to that of the

scrotum.

Genus Cervus Linn.

Cervus ELAPHUS Linn. (p. 941).

The feet and glands of a second specimen are identical with

those described in 1910.

The rhinarium has the infranarial portion mesially sulcate and

slightly narrower than the area between the nostrils; it is not,

however, visible in profile view, owing to the forward extension

of the hairs of the upper lip. | On the upperside the hairs of the

muzzle encroach in the middle line some distance beyond the

posterior angle of the nostrils. The buccal and ocular vibrissze

are well developed ;. the interramal tuft is represented by a single

vibrissa and the genal tuft by two placed close together and set

halfway between the line of the mouth and of the eye, beneath

the posterior angle of the latter.

CERVUS CANADENSIS Hrxl.

The rhinarium resembles tolerably closely that of C. elaphus,

except that the upper surface is more overgrown with hair,

which extends nearly as far as the anterior border of the nostrils,

and the infranarial portion isalso narrower, being much narrower

than the area between the nostrils, owing to the forward and

foe extension of the hairs of the upper lip. (Text-fig. 2,

Its 13%

The Asiatic Wapiti (Cervus xanthopygus) has a very similar

rhinarium.

In these two species of Cervus the rhinarium is more reduced

by hair-growth from the muzzle than in any of the typical deer,

except Dama, although these genera are surpassed in that respect

by Alcesand Rangifer.

CHARACTERS OF ELAPILURUS AND OTHER CERVID. 183

CERYUS HORTULORUM Swinh.

As was to be expected, the feet and glands of this species ditfer

in no respect from those of the Japanese species (Cervus sika)

I described in 1910.

In one example the labial and ocular vibrisse are normally

developed ; but the genal and interramal tufts are absent. The

rhinarium has a considerable portion of its upper surface, nearly

half, overgrown with hairs from the muzzle, which form a convex

curve between the nostrils; the infranarial portion is tolerably

wide, a little wider than the space between the nostrils.

(Text-fig. 3, A.)

Text-figure 2.

SOT! f

I Wh Hy

A. Rhinarium of Wapiti (Cervus canadensis), from the front.

B. The same from above.

Two examples showed an interesting variation in the structure

of the penis. In both the urethra was inferior and marked

distally by a distinct ridge ending in a slightly upturned point

as in other typical Cervine deer; and in one specimen the

truncated apex above the urethra was, as usual, lobate, the tips

of the lobes folding inwards so as to make a groove where they

meet above the point of the urethra. But in the other specimen

this terminal portion of the glans was simple and unlobed—a

very unusual occurrence, (Text-fig. 17, C, D.)

In an example of C’. sika the glans penis resembled that of the

first-described specimen of C. hortulorwm.

US}

184 MR. R. I. POCOCK ON THE EXTERNAL

CErvus ELDI Guthr. (p. 944).

In two additional specimens the feet and glands resembled

those of the example described in 1910.

In both specimens the buccal and ocular vibrisse were well

developed. In one of them there was a single long genal and a

single long interramal vibrissa. In the other there were two

rather widely separated genals, but no interramal *.

Text-figure 3.

ASATW ION;

WU My

NARA

) fi)

A i}

A. Rhinarium of Cervus hortulorum, from the front.

B. The same of Dama dama.

C. The same from above.

D. The same of Cervus eldi.

The rhinarium is large. On its upper side the bair of the

muzzle advances in the middle as an angular growth, which reaches

to a point approximately in a line with the middle of the nostrils.

The infranarial portion is wide, considerably wider than the area

between the nostrils. (Text-fig. 3, D.)

Genus Axis H. Smith.

Axis axis Erxl. and A. porcinus Zimm. (pp. 948-950).°

Two examples of the Chital or Axis Deer (A. avis) examined

since 1910 show some interesting variations in the hairiness of

the glandular depression on the front of the pastern of the hind

foot.

In the specimen originally described the walls of the depression

were covered with short, rather sparsely scattered hairs and the

* I have noticed in other species of Cervide that the genal and interramal

yibrissse ave of incoustant occurrence within specific limits.

CHARACTERS OF ELAPHURUS AND OTHER CERVIDA, 185

interungual tie was naked. In both the other examples the walls

of the depression were clothed with long hairs running down-

wards and forwards, following the curve of the floor of the pouch,

but capable of being raised so as to project from its orifice.

In one of these two specimens the interungual tie was naked,

but in the second it was covered with short hairs. (Text-

fig. 4, B.)

One of the interests of these variations is that they break

down the differences I recorded in 1910 between the feet of the

Chital and of the Hog Deer (A. porcinus), in which the inter-

ungual tie is hairy and the glandular pouch provided with long

projecting hairs. On the strength of these differences I separated

porcinus from avis, retaining for it the subgeneric name Hy-

elaphus ; and this opinion was adopted by Lyddeker. Owing to

the identity in the structure of the hind foot in the two species,

Text-figure 4.

A. Longitudinal section of the fore foot of Axis axis.

B. The same of the hind foot.

Hyelaphus now falls as a synonym of Awisand the latter will con-

tain two species A. avis and A. porcinus, the genus Amis being

distinguished from the supposedly allied form Musa, which has

similar simple antlers, by the presence of the glandular pouch on

the hind foot.

In the two examples of A. auis above referred to, the creamy

or waxy secretion sticking together the hairs of the glandular

pouch smelt like tailow candles, with an admixture of cheese.

The fore feet differed in no important respect from those of the

specimen described in 1910. (Text-fig. 4, A.)

Ina fetal Axis Deer, measuring sixteen inches (about 400 mm.)

from the snout to the root of the tail and weighing 3 lb., the

skin although naked showed faintly all the spots characteristic of

the adult, The tail was relatively much longer than in the adult.

186 - MR. R. I, POCOCK ON THE EXTERNAL

The eyes were closed. ‘Phe only hairs developed were the sensory

vibrissee on the head, the mystacials, submentals, superciliaries,

and suboculars being very evident. On the cheek there were two

genals and the interramals arose from a warty excrescence. ‘The

early development of all the tufts of vibrisse characteristic of the

Mammalia is interesting. The position of the antlers was marked

by a depression of the integument on each side of the crown of

the head. The preorbital ‘gland was represented by a shallow

depression, opening by a short linear orifice a little distance below

and in front of the eye. The metatarsal gland was marked by a

small pale oval patch on the skin of the leg below the hock.

The pedal giands were as in the adult, a deep pit on the hind foot

and a shallow pit on the fore foot.

Genus Dama Frisch.

Dama DAMA Linn. (p. 950).

The rhinarium in the Fallow-Deer is much reduced by the

encroachment of hair from the muzzle and lips. On its upper-

side the hair extends nearly as far as the anterior end of the

nostrils, leaving only a narrow strip bordering the nostrils

laterally above. The infranavial portion is very narrow inferiorly,

being at its narrowest point much less than the width of the

space between the nostrils. (VText-fig. 3, B, C.)

Genus ELapnurus M.-Edw.

ELAPHURUS DAVIDIANUS M.-Hdw. (p. 945).

My notes upon this species in 1910 were based upon the

observations of others and upon the examination of a dried skin

in the British Museum. Since then I have seen two examples in

the flesh, both from the Duke of Bedford’s herd at Woburn.

The first of these, a male, died in the Zoological Gardens from

impaction of the psalterlum in August 1917; the second, a

female, died. at Woburn and was kindly sent to me for exami-

nation by the Duke in January 1923.

The facial vibrisse are represented by well-developed buccal

and orbital bristles; by a small genal tuft on each side below the

corner of the eye and above the line of the mouth, and by an

interramal tuft of a few short bristles arising from a small

elevation. In the male there is an interramal dew-lap which is

absent in the female. Perhaps this is seasonally developed.

The rhinariwm is large and naked with the nostrils narrow and

widely separated, but with the space between them a little

narrower than the infranarial portion, which has a short inferior

median sulcus. On the upperside the hair of the muzzle

encroaches to a great extent between the nostrils, reaching beyond

CHARACTERS OF ELAPHURUS AND OTHER CERVID. 187

the middle of their length, but leaving a wide naked rim above

them laterally. Although the rhinarium differs considerably in

its nakedness from that of Cervus canadensis or Dama dama,

it is not very unlike that of Cervus eldi. (Text-fig. 5, A, B.)

The preorbital gland is a deep and long pit, with a naked floor,

its orifice when closed being about as long as the eye.

The ear is moderately large and thickly clothed with long

hairs inside. It has three vertical ridges, the anterior and

posterior being stronger than the median. The basal ridge is

bilobed, the posterior lobe is much longer than the anterior, but

not very much higher, and has a convex edge. Beneath it there

Text-figure 5.

A. Rhinarium and edge of lower lip of Hlaphwrus davidianus,

from the front.

B. Rhinarium of the same from above.

is a curved rounded ridge leading down to the ovifice. (Text-

fig. 15, A.)

The feet are long and broad with the digits widely separabie,

owing to the shallowness of the interdigital web arising from the

depth of the depression between the heels of the hoofs. The

upper interdigital depression is approximately as deep as in

Cervus elaphus or maral, and is scantily covered with hairs in its

proximal half, naked in the distal half. The hoofs themselves

are moderately wide and pointed, but are remarkable for the

exceeding length of the heels, which extend backwards almost to

188 MR. R. I. POCOCK ON THE EXTERNAL

the false hoofs—the latter being exceptionally long, particularly

on the fore feet. The area between the false hoofs is naked,

except for a narrow strip of hair running on each side from the

back of the leg to the heel. The deep depression between the

heels is entirely naked, The hind feet differ from the fore feet

in being smaller and in having relatively shorter false hoofs.

The feet of the female in the winter were more hairy below than

those of the male in summer described above. ‘The wide

separation of the digits and the length of the heels and of

the false hoofs suggest adaptation to swamp-life. (Text-fig. 6,

A, B.

Fae metatarsal gland may, it appears, be present or absent.

It is present on the skin in the British Museum; but I could

find no trace of it in the female, and have no note on the subject

in the male.

The penis is truncated apically and provided with five pairs of

lobes which normally fold over the urogenital orifice, but are

capable of spreading out like the petals of a flower. Except,

however, that the lobes are somewhat better defined, the penis

does not differ essentially from that of Cervus xanthopygus

and of other typical deer of the Old World. (Text-fig. 17,

KE, F, G.)

It may be recalled that, according to Garrod (P. Z.58. Sie

p- 9), the penis of the American deer now referred to the genera

Odocoileus, Mazama, Pudu, and others differs from that of such

Old World deer as Cervus, Aais, Dama, etc., in being attenuated

at the apex, the urethral canal being produced into a median

slender process, unlike the upcurled termination of the urethra,

which does not project beyond the truncated tip of the glans in

the Old World forms. Hence the discovery of the structure of

the penis in Hlaphurus is the final piece of evidence required to

establish the affinity of that genus with the typical Old World

deer and to sever it from the American genera with which

Lyddeker affiliated it.

Notes on the Seasonal Colour-change and Antler-growth of

Elaphurus.

Even as recently as 1915, the colour of Hlaphurus was described

as “reddish tawny with a tinge of grey” by lLyddeker, who

judged apparently from the mounted specimen in the British

Museum, which was presented to that institution by the Duke of

Bedford in July 1899. Asa matter of fact, the reddish tint is a

transient phase lasting some three or four months of the summer

season, roughly from May or June to August or September.

For the remaining eight or nine months of the year the animal

is the colour of a grey donkey; and this grey coat is thick and

luxuriant in the winter, whereas the reddish coat is comparatively

short and sparse.

CHARACTERS OF ELAPHURUS AND OTHER CERVID. 189

Text-figure 6.

A. Lower side of fore foot of Hlaphurus davidianus.

B, Section of fore foot of the same.

190 MR. R. 1. POCOCK ON THH EXTERNAL

The stag presented to the Society on January 26th, 1916, by

the Duke of Bedford was in the grey pelage. ‘This was replaced

by the red pelage in June, and the grey phase was assumed in

September and carried through the winter until May 1917, wnen

the red summer coat was substituted. The first clear signs of

the shedding of this red coat were noticed on July 23rd. It

came away with great rapidity and was all gone in about a week’s

time, except on the crown of the head, where it persisted until

the stag’s death in August.

The red phase was therefore about a month earlier in its

appearance and disappearance in 1917 than in the previous

year.

With regard to the antlers, the stag was carrying a burnished

pair on its “arrival on J anuary 26th, 1916. These were shed on

February Ist. The new antlers grew rapidly and were burnished

by the time the grey pelage was replaced by the red pelage in

midsummer, when rutting set in. These antlers were shed on

October 22nd, and a new pair started at once; but instead

of reaching their full size im the winter and being shed in

February as in the previous year, they continued to grow through

the winter and spring, began to peel in May, coincidently with

the shedding of the grey winter pelage and the appearance of the

red summer coat, end were unshed at the time of the stag’s

death in August. Being the result of about seven months’

growth, these antlers were much larger than the preceding pair,

which were developed in about four and a half months.

But, as recorded by Lord Tavistock in a letter to me, the antler-

shedding in Hlaphurus is a much more complicated and variable

phenomenon than those who have seen single stags are aware.

The following is the substance of his observations made upon the

herd at Woburn :—

No immature stag grows more than one pair of antlers in a

year. The antlers ‘of a yearling are not clean till the end of

June and are shed in midwinter or even later. The antlers of a

four-year old stag are shed early in November, and may not be

clean for three weeks after those of the old stags have lost their

velvet. All old stags clean and shed their summer antlers with

as little variation as to date as red deer. But the date of the

cleaning and shedding of their winter antlers is exceedingly

variable. In the case of late shedding of the winter antlers,

stags will sometimes be found in May with but very few inches

of new velvet-covered antlers; but, no matter how small the

growth, these antlers will harden and clean in time for the

rutting season. These stumpy-horned stags, however, have very

poor chance against their better-armed companions, and it is

only when the latter ave exhausted at the end of the season that

the former have any chance of collecting a harem at all.

Very large antlers of Hlaphurus are always the result of

a winter erowth by an adult animal; but the production of a

CHARACTERS OF ELAPHURUS AND OTHER CERVIDA. 191

single pair of antlers of large size instead of two pairs of small or

medium size becomes increasingly rare the longer the deer remain

in this country. The result is now that the only really handsome

heads are those carried by stags five and six yearsold. As arule,

there is no attempt to resume the rut when the winter antlers

harden, the stag remaining as lazy and peaceful as if still in

velvet ; but one or two stags were known to Lord Tavistock which

for several seasons recommenced calling in midwinter, November

was the latest month in which he heard stags, still carrying their

summer horns, calling; and he has seen a six-year old stag, in.

full summer coat and carrying a large pair of horns, herding his

hinds with great energy and calling in the first week in February.

The antlers in this case were the continuous growth of the whole

autumn and winter.

The majority of the calves are born in April and May. Very

late ones are not common.

In 1922 Ludwig Zukowsky, Hagenbeck’s assistant, published a

paper * upon Hlaphurus davidianus, based upon a specimen that

was exhibited for nearly two years at Stellingen. Unfortunately,

the author was not acquainted with the published literature on

this subject. He described in detail the mode of growth of the

antlers, and quite correctly spoke of the anterior branch as the

“brow-tine.” In this he confirmed in every particular my account

published in 1912 (P. Z. 8. pp. 777-780) from sketches kindly

supplied by Lord Tavistock, the accuracy of which I was subse-

quently able to verify on the specimen exhibited in the Zoological

Gardens. Zukowsky also recorded the succession of the antlers

in this aberrant species. The specimen at Stellingen cast its

antlers on the following dates :—March 8, 1913, Sept. 18, 1913,

March 17, 1914, Oct. 5, 1914, and died on Dec. 30, 1914, with

fully developed antlers in velvet.

When Hagenbeck’s specimen died it was sent to be mounted to

the Natural History Museum, Hamburg, and Herr Gast, Super-

intendent of that institution, subsequently wrote to Zukowsky

to inform him that he had found just beneath the edge of the

under lip ‘‘a deep sheath or vagina let into the skin, which seemed

to point to a glandular duct, one inch in length.” I failed to find

any trace of this structure in the female specimen of Elaphurus

which the Duke of Bedford sent to me for the special purpose of

investigating this gland. Possibly there may be such a gland

restricted to the male; but, so far as is at present known, the

cutaneous glands in the Cervide are alike in the two sexes aeG

* J am indebted to the Duke of Bedford for kindly sending me a copy of this

paper, which was published in Arch. Naturg. vol. Ixxxviti., May 1922.

+ When I first suggested in 1910 that the anterior and posterior branches of the

antlers of Elaphurus are strictly homologous with the * brow-tine ” and “ beam”

respectively of the antlers of ordinary Cervidee, I was not aware that I had been

anticipated in this view by Dr. Theodore Gill, who came to the same conclusion.

This was published I believe in ‘ Forest and Stream,’ but I do not recollect the

yeference. 4

+ Following Garrod, I exclude Moschus from this fainily.

192 MR. R. I. POCOCK ON THE EXTERNAL

Genus Hypropores Swinh.

Hypropores INERMIS Swinh. (p. 956).

Some of the external characters and the visceral anatomy of

this aberrant genus were described by Garrod (P. Z. 8. 1877,

pp. 789-792). Additional information was later supplied by

Forbes (P. Z. 8. 1882, p. 637).

The head is remarkable for the complete absence of all trace of

antlers—a feature in which this genus is unique in the Cervide,—

the long, slightly curved, pointed and movable upper canines, and

the narrow muzzle. (Text-fig. 7.)

The lightly areolated rhinarium has a deep wide infranarial

portion, which is wider than the supranarial portion and therefore

much wider than the internarial area. The latter is of normal

Text-iigure 7.

a —

Head of Hydropotes inermis.

width. ‘The supranarial portion is unusually high, so that the

height of the entire rhinarium, as described by Garrod, is about

equal to its width. (‘Text-fig. 8, C.)

The mystacial vibrisse ave tolerably numerous and of average

length. The superciliaries and infraorbitals are also normally

developed ; but in the thick hairs of the cheek and throat I can

find no trace of genal or interramal vibrisse, which are some-

times present but never abundant in other Cervide.

The preorbital gland is represented by a small shallow pouch

lodged in a naked area of skin just in front of the eye. (Text-

figs. 7; 8, D.)

The ear is short and broad, and densely clothed within with

hairs, which meet in the middle line of the hollow of the pinna.

CHARACTERS OF ELAPHURUS AND OTHER CERVIDA. 193

The overfolded edges of the base of the ear meet at an acute

angle. There are two vertical cartilaginous ridges, an anterior

and a posterior, supporting the pinna; and the prominences

on the basal ridge are very unequal in size, the anterior being

quite small and slender, whereas the posterior rises as a high

triangular peak. These prominences are quite unlike those of

any other species of Cervide that I have examined. (Text-

fig.) 8; A, Bs)

Feet.—The hoofs of the fore foot are long and pointed, and

tolerably widely separable; the soft inferior cushion, constituting

the greater part of the sole, is continued backwards some distance

behind the smooth heel-tie which joins the heels together. On

the front of the pastern there is a tolerably deep and smooth

Text-figure 8.

A. Base of ear of Hydropotes inermis.

B. The same cut open.

C. Rhinarium of the same from the front.

D. Section of preorbital gland of the same.

glandular depression, and the skin all along the back of the

pastern from the heel-tie to the area between and beyond the

false hoofs is also glandular. ‘This area is more scantily covered

with hair than the area of the leg above the false hoofs, and the

false hoofs themselves, which are short, are basally encircled by

an area of naked skin divided inferiorly by a narrow scantily

hairy strip of skin. (Text-figs) 9, A; 10, A.)

The hind foot is very similar to the front foot, but the hoofs

are rather more widely separable, and the heels are narrower and

a little longer. As in the front foot the back of the pastern is

hairy down to the heels, and the heel-tie is naked. The glan-

dular depression is considerably deeper and longer, and has a

more abruptly upturned anterior rim. The widely separable

194 ; MR. R. I. POCOCK ON THE EXTERNAL

hoofs with their long heels are adapted for progression upon soft

marshy ground. (‘T'ext-figs. 9, B; 10, B.) :

There is no trace of metatarsal or tarsal gland.

The anus opens in the upper half of an oval naked area of

skin, broader above than below and extending from the root of

the tail nearly to the scrotum. The orifice is surrounded by a

thickened glandular rim. The edge of the smooth oval area is

sharply defined by the thick growth of hair, which is as luxuriant

here as elsewhere on the body. The tail is short, only long

enough to cover, when depressed, the smooth anal area, and is

thickly covered with hair below as well as laterally and above,

(Text-fig. 11, B.)

The inguinal region has a single pair of teats—not two pairs

us recorded by Garrod in the newly-born fawn. The area be-

tween them is scantily clothed with long hairs; but on the outer

Text-figure 9.

A. Lower side of fore foot of Hydropotes inermis.

B. The same of the hind foot.

side of each there is a large nearly naked area of skin extending

a considerable distance outwards and backwards; and towards

the outer edge of this there is a shallow, curved, glandular depres-

sion, recalling the inguinal glands of many of the Bovide. This

is the first record of the presence of inguinal glands in the

Cervide. (Text-fig. 11, A.)

The prepuce is a naked button of skin surrounded by long

hairs and occupying the normal abdominal position in front of

the inguinal region and remote from the scrotum. The penis is

perfectly simple, straight, and slightly attenuated, with the orifice

terminal. It runs between a couple of ridges in the dorsal wall

of its sheath. (Text-fig. 11, A.)

CHARACTERS OF ELAPHURUS AND OTHER CERVIDA. 195

The absence of antlers and the presence of large upper canine

teeth in Hydropotes naturally suggested the possibility of kinship

between it and Moschus. Since Garrod definitely disposed of this

view, it need not be further discussed. He closed his account

of the anatomy of Hydropotes as follows:—‘ To what group of

the Cervidee Hydropotes is most allied there is still considerable

uncertainty. That it is not allied to the New-World type is

evident from its vomer not extending downwards to join the

osseous palate posteriorly. hat it is not Cervuline [related to

the Muntjacs, which also have large canines] is equally certain on

Text-figure 10.

A. Section of fore foot of Hydropotes inermis.

B. The same of the hind foot.

account of its cuneiform bones being free from the naviculo-

cuboids. Its large Spigelian lobe favours the view suggested by

Sir Victor Brooke, that it is most closely allied to tie Rusine

Deer” (p. 792). ‘This view, | think, may be dismissed without

hesitation. On the other hand, Forbes’s opinion (P. Z. 8. 1882,

p- 637) that Hydropotes is related to Capreolus is worth more

consideration. The two, at all events, agree in being “ telemeta-

carpalian ” and in the structure of the vomer and apparently of

the glans penis. But the differences between them are too many

and too deep-seated to admit of close affiliation. In the first

place, Hydropotes is the most primitive of all existing Cervide: in

196 MR. R. I. POCOCK ON THE EXTERNAL

the complete absence of the antlers and the presence of long

tusk-like canines in the male. Capreolus, on the other hand, has

well-developed branched antlers, and has normally lost all trace of

the canine, being in the latter particular more specialised than

Cervus. Again, Capreolus has lost the preorbital and retained the

metatarsal gland. Hydropotes has retained the preorbital and

lost the metatarsal gland; and has acquired inguinal glands, a

new feature in the Cervide. The differences between the two in

the length and separability of the hoofs are also marked; but,

considering the differences of habitat, they are not obviously an

indication of remoteness in kinship. The deep and long inter-

digital glandular depressions in Hydropotes, on the contrary, are

Text-figure 11.

A. Inguinal and genital area of Hydropotes inermis g, showing the nearly

naked area of skin on each side, with a single pair of teats and the shallow

pouches of the inguinal glands. The sheath of the glans penis cut open

along the middle line, showing the glans turned towards the right and the

two ridges forming the groove along which it runs. .

B. Anal area of the same with the tail raised, showing the anus insunk in th

surrounding glandular area.

much more primitive than the pouch-like gland of the hind foot

of Cervulus.

Tn view of the peculiarities of Hydropotes, I propose to separate

the genus from the rest of the Cervide as the type of a special

subfamily, Hydropotine.

Genus OpocortEus (= Dorcelaphus).

ODOCOILEUS VIRGINIANUS Bodd. (p. 962).

In 1910 I had seen no fresh example of this species, quoted as

Dorcelaphus americanus, and was only able to describe the glands

in the feet, as shown by specimens in the Museum of the College

CHARACTERS OF ELAPHURUS AND OTHER CERVIDA. 197

of Surgeons. Since then I have examined an adult female that

died in the Gardens on August 28th, 1911.

The rhinarium is large and naked, the hairs of the muzzle

encroaching but little on its upper surface ; the infranarial portion

is wider than the space between the nostrils.

The preorbital gland is represented by a very small, shallow

pit.

Text-figure 12.

A, Longitudinal section of the metatarsal gland of Odocoileus virginianus

(typical form).

B. Transverse section of the same.

C. Section of fore foot of O. virginanus spinosus.

D. Section of hind foot of the same.

H. Section of fore foot of Moschus moschiferus.

The tarsal gland consists of a patch of thickened vascular skin

covered by a thick pad of long hair stuck together with secre-

tion, but without any under-fur ; the hairs are umber-brown with

white bases.

The metatarsal gland shows externally as an elongated patch

of naked, horny, granular skin overlapped by the long, mostly

Proc. Zoou. Soc.—1923, No. XIV. 14

198 MR. R. I. POCOCK ON THE EXTERNAL

white hairs growing round it. The secreting glands lie beneath

these long hairs, the skin being thicker and more vascular there

than elsewhere. The naked area between the long hairs seems

to serve as a receptacle for the secretion, since it collects and is

encrusted there. (Text-fig. 12, A, B.)

The feet and pedal glands are as described in 1910, except that

the pouch on the hind foot is rather more capacious, has a wider

orifice and naked walls. The pouch on the fore foot is similar,

but only about half the size.

ODOCOILEUS VIRGINIANUS SPINOSUS Gay & Gervais (p. 962).

In 1910 I referred to a living example, identified as Dorc-

elaphus americanus savanunarum, which came from Venezuela and

was presented to the Society by Major Albert Pam. The animal

died in 1914, and I was able to examine it in a fresh state.

The rhinarium resembles very closely that of the typical

North-American form, but from my sketches it seems that the

infranarial portion is somewhat narrower inferiorly and the

upper surface a little more overgrown with hair in the middle.

(Text-fig. 14, A.)

The vibrissz on the upper and lower lips and above and below

the eye are well developed; but there is a single long genal

bristle arising beneath the posterior angle of the eye and far back

in a line with the mouth. There is no interramal tuft.

The preorbital gland is small as in the typical form.

The tarsal gland is marked by a thickened bunch of hair,

brown and white in colour, and covered at the base with

secretion.

The metatarsal gland is a very small patch of thickened skin

overgrown with white hair.

The glands are well-developed pouches, both on the front and

the hind foot. The hind foot hardly differs at all from that of

the typical O. americanus, the walls of the pouch being naked ;

but in the fore foot the walls of the pouch are thickly covered

with hair and the orifice is much longer than in the typical

O. virginianus, with the heel-tie shallower. There is also a small

naked patch at the posterior inferior angle of the heel-tie.

(Text-fig. 12, C, D.)

In this animal, as in the above-described example of the typical

O. virginianus, the false hoofs of the fore foot are much longer

than those of the hind foot.

ODOCOILEUS VIRGINIANUS PERUVIANUS Gray.

A male example from Iquique, presented by Miss Peggy Lomax

on April 24 in 1911, died in Jan. 1923 from pneumonia, the

worn condition of the teeth showing it to be an old animal. The

donor informed me that it was born in 1909. It was therefore

CHARACTERS OF ELAPHURUS AND OTHER CERYVIDA. 199

in its fourteenth year. This was the unidentified specimen of

Odocoileus, of which I described the antler-growth in 1912 (Proc.

Zool. Soc. pp. 781-783) to indicate the homology between the

so-called “ subbasal snag” of the genus Odocoilews and the “ brow-

tine” of Cervus.

The description given above of the external characters of

O. v. spinosus applies very closely to this form. The only

variations noted were the presence in O. v. peruvianus of a pair of

long interramal vibrisse, of longish hairs on the sides of the

glandular pouch of the hind foot, and the reduction in the size of

the metatarsal gland. This gland was merely represented super-

ficially by an inconspicuous patch of hairs slightly longer and

slightly different in tint from those of the surrounding area of

the lower half of the metatarsal area. The skin beneath this

felt slightly thicker to the touch, but the only indication of

secretion was a small scab on the gland of the right side. This

gland was not visible to me in the living animal, and I thought it

was absent. It might very easily be overlooked in prepared

skins; and this would account for O. v. peruvianus having been

described as without metatarsal glands. Not improbably they .

may be sometimes altogether aborted in this subspecies.

The tarsal gland, on the contrary, is represented by a large

thick mat of longish hairs, which, upon being separated, showed

as a mixture of grey and black with yellow secretion at the

base.

The car is tolerably large and nearly naked, as in the related

race O. v. gymnotis. It has two vertical cartilaginous ridges, of

which the posterior is much the stronger. Inferiorly it forms

the posterior border of a deepish pit, the anterior border of which

is a rounded ridge descending from the rounded posterior lobe of

the basal ridge. The anterior lobe of this is a little smaller than

the posterior; there is a low short longitudinal crest on its outer

side, and the anterior vertical ridge terminates on its inner side.

A thickened rounded crest, defined in front and behind bya

hollow, descends to the bottom of the cavity of the ear and the

orifice opens in the anterior hollow. (Text-fig. 19, B.)

The penis, as pointed out by Caton for the North-American

forms of Odocoileus, was pendulous from a point just in front of

the scrotum. The tip of the prepuce was almost naked, but just

within its ovifice were some hairs arising from definite papille.

The glans, as in other American deer examined, agrees exactly

with Garrod’s description, ending in a narrowed point with ter-

minal orifice. (Text-fig. 17, A, B.)

Within the limits of the genus Odocoilews the preorbital

gland and the pedal glands appear to be always present, the

latter occurring on both hind and front feet. According to

Caton, they are, however, relatively smaller in O. hemionus than

in O. virginianus, O. columbianus coming between the two in this

particular.

14*

200 MR. R. I, POCOCK ON THE EXTERNAL

The metatarsal and tarsal glands are variable. The former

attains its maximum in development in O. hemionus, where,

according to Caton, the bare patch of skin overlapped by long

hairs may be five or six inches long. In North American ex-

amples of O. virginianus it varies from less than one inch to an

inch and a half in length, whereas in some of the southern forms

(such as O. v. toltecus, acapulcensis, etc.) it is absent. The tarsal

gland varies in size and colour according to the species ; but I do

not know whether it is absent or not. According to Caton, its

hairs expand under excitement, like the hairs of the tail and

rump, in the North American species.

Genus Mazama Raf. (p. 962).

The two species of this genus previously described were

M. nemorivagus (p. 962) and MM, bricentt (p. 964).

MAZAMA TEMA Raf.

A single male example from Guatemala, which died on April

26th, 1914.

The rhinarium is naked above almost as far back as the

Text-figure 13.

A. Longitudinal section of the fore foot of Mazama tema.

B. The same of the hind foot.

C. The same of the hind foot of Pudw pudw.

posterior angles of the nostrils and the infranarial portion is very

wide, showing laterally as far back as the middle of the nostrils,

as in M. nemorivagus. (Text-fig. 14, C.)

CHARACTERS OF ELAPHUBUS AND OTHER CERVIDA. 201

The normal labial and ocular vibrisse are present; the genal

tuft is represented by two bristles beneath the posterior corner of

the eye and as low as the line of the mouth; the interramai tuft

is composed of three bristles a little behind a line joining the

corners of the mouth.

The preorbital gland is a small shallow depression, as in

M. nemorivagus and MM. briceniti.

The tarsal gland is represented by a small tuft of hair, as in

MM. nemorivagus.

The metatarsal gland is absent as in MW. nemorivagus and

M. bricenit.

The pedal glands on the fore foot are much larger and deeper

than in M. nemorwagus and M. bricenii, resembling those of

O. virginianus spinosus in depth, but having a shorter orifice.

The walls are thickly hairy. The heel-tie also is lower than in

the other species of Mazama and is altogether naked, and the

heels of the feet are long and the hoof moderately so. (Text-

fie) NS, Av.)

The pouch on the hind foot is larger than on the fore foot and

has thickly hairy walls as in WV. bricent?. In this foot also the

heel-tie is naked, the heel is long, but the hoof is short. (Text-

fig. 13, B.)

This species differs from the other two species of Mazama in

the large size of the gland of the fore foot. The nakedness of the

heel-ties on both fore and hind feet is a character unrecorded

elsewhere in the Telemetacarpalian deer. :

Genus Pupu Gray.

Pupu pupu Mol. (p. 967).

The head of this species is remarkable for the forward growth

of the hair from between the ears to the summit of the crown,

where it meets and forms a crest with the backwardly growing

hair of the muzzle and forehead.

The rhinariwm is tolerably large and lightly areolated; the

supranarial portion is mostly naked above, the hair only en-

croaching upon it toa slight extent posteriorly; the infranarial

portion is mesially grooved ; it is narrower than the supranarial

portion, but a little wider than the space between the tolerably

widely separated nostrils. (Text-fig. 14, B.)

The facial vibrisse are represented by some shorter mystacial

and submental and longer superciliary and subocular bristles,

there being no trace of genal or interramal tufts.

The preorbiial gland is a comparatively small and shallow

pouch, opening on a naked area a little in front of the eye.

The ears are rounded, the expanded portion being supported by

202 MR. BR. I. POCOCK ON THE EXTERNAL

four vertical ridges ; the basal internal ridge has an angular pro-

minence, which is as high as its posterior rounded termination.

(Text-fig. 16, D.)

The feet have pointed hoofs with a tolerably short heel and

short false hoofs. On the hind foot there is a deep and long

interdigital glandular depression, the integument of its floor

being almost in contact with that of the hack of the pastern.

The distal or anterior edge of this depression is not elevated, the

heel-tie being shallow and smooth. ‘The walls of the depression

Text-figure 14.

A. Bhinarium of Odocoileus virginianus spinosus, from the front.

&. The same of Pudw pudu.

OC. The same of Mazama tema.

D. The same of Rangifer tarandus.

are clothed with hair, which is rusty yellow towards the mouth

and white near the bottom. The depression on the fore foot is

very similar, but a little shallower. Both tarsal and metatarsal

glands are absent. (Text-fig. 13, C.)

In the brief description given by Flower (P. Z. 8. 1875, p. 160)

of the feet of this species it is said that the pedal glands are not

represented by distinct pouches, ‘‘ but the skin in the depression

between the toes... is bare and evidently has a free sebaceous

CHARACTERS OF ELAPHURUS AND OTHER CERVIDA. 203

secretion, representing, in the author’s opinion, the most rudi-

mentary or earliest stage of an interdigital gland.” How little

this account accords with the actual facts may be seen by com-

paring it with the description given above. I was completely

misled by Flower into imagining that the interdigital areas in the

Pudu resemble those of Cervus, surprising as such a conclusion

was. ‘There is in reality no such resemblance. The depressions

are like those of the hind foot of the Fallow-Deer (Dama dama)

and of the Muntjacs (Muntiacus = Cervulus) and Hlaphodus,

which I described and figured in 1910: that is to say, they

Text-figure 15,

A. Base of ear of Hlaphurus davidianus, cut open.

B. The same of Odocoileus virginianus peruvianus. ‘

C, The same of Tragulus.

D. The same of Pudu pudu.

EK, The same of Muntiacus muntjak.

belong to what I believe to be the most primitive type of pedal

gland in the Cervidee—a long deep depression the floor of which

is in contact, or nearly so, with the integument of the back of the

pastern. No other genus of Cervide inhabiting America has

feet of this type, so far asis known. Cervus canadensis may be

set aside as an alien from Asia; but in Rangifer, Odocoileus, and

Mazama the glandular depression of the hind foot is a deep pouch

with constricted orifice, whereas in the front foot it is a small

shallow pit, the heel-tie in both cases being deep.

204 MR. R. I. POCOCK ON THE EXTERNAL

Genus Raneirer H. Smith.

RANGIFER TARANDUS Linn. (p. 960).

In 1910 I was only able to examine the dried limbs of the

North American race of this species, 2. tarandus caribou. That

account I can now supplement by observations upon a female

specimen that died in the Gardens on July 27th.

The muzzle of the Reindeer has been described as covered with

hair, as if there was no trace of rhinarium. It is true that the

Text-figure 16.

A. Section of fore foot of Rangifer tarandus.

B. The same of hind foot.

area above, between, and below the fleshy valvular nostrils is

entirely overgrown with short soft hair; but along the edge of

the upper lip there is a naked areolated tract, wider than the

space between the nostrils, with convex upper edge and similar

in sculpture and width to the corresponding tract on the edge of

the lower lip, but about twiceas deep. (Text-fig. 14, D.)

This is the inferior portion of the rhinarium retained. In

this particular the Reindeer is unique amongst the Cervidee,

CHARACTERS OF ELAPHURUS AND OTHER: CERVID. 205

The external appearance of the well-developed tarsal gland was

fully described by Caton, and I have nothing to add to his

description or to my account of it published in 1910. The

metatarsal gland, as Caton and others have stated, is absent.

My first account of the feet, taken from dried specimens, requires

some modification. On the fore foot of the fresh specimen there

is a deeper depression on the front of the pastern; and on the

Text-figure 17.

A. Penis and scrotum of Odocoileus virginianus peruvianus, showing the

long pendulous prepuce close to the scrotum.

. Glans penis of the same.

. Glans penis of the same from the side.

. Apex of glans penis of Hlaphurus davidianus, with lobes folded, from

the front.

E. The same with lobes expanded.

C. Apex of glans penis of Cervus (Sika) hortulorum, from the front.

G. Glans penis of the same from the side.

back of the pastern the depression is deeper still, the integument

forming the floor of the posterior depression being almost in

contact with that of the anterior depression, so that the two

digits are joined together by a very shallow fold or loop of

integument. This loop expands towards the hoofs, and the

expanded portion is filled with a mass of soft, fatty, somewhat

gelatinous material. But there is no trace of a glandular pouch

206 MR. R. J. POCOCK ON THE EXTERNAL

on the fore foot *. It is to the remarkable shallowness of the

integumental fold between the digits and the depth of the

depression on the back cf the pastern that the wide separability

of the hoofs in Rangifer is due. The skin of the interdigital

depressions is everywhere covered with long thick hair. In the

hind foot, the posterior depression of the pastern is not so deep

as on the fore foot, but the integumental fold is very similar,

although even shallower proximally. On the front of the

pastern the depression is deeper and forms a deep glandular

pocket very like that of Odocoileus. (Text-fig. 16, A, B.)

A species which even surpasses Rangifer in the extreme

shallowness of the heel-tie and the depth of the depressions

between the digits is the Musk-Deer (Joschus), which has the

hoofs similarly separable for progression on soft snow and for the

sure-footed descent of steep slopes. (Text-fig. 12. E.)

Classification of the Cervide.

In 1910 I divided the Cervide into two subfamilies, Capreo-

line and Cervine, corresponding respectively to Sir Victor

Brooke's divisions Telemetacarpalia and Plesiometacarpalia. It

appears to me that we must adopt Brooke's primary grouping of

the family ; but that the genera of both groups, more particularly

of the Telemetacarpalia, are too diversified to be assigned to the

two subfamilies proposed. It may be claimed at all events,

I think, that the subfamilies tabulated below are as well defined

as the subfamilies of the Bovide :—

a. Distal ends of Jateral metacarpals retained, proximal end aborted

(Telemetacarpalia).

6. Vomer dividing the posterior nares.

e. Naviculo-cuboid and external cuneiform bones of tarsus

united; pedal glands a deep long cleft, with shallow heel-

tie. Pudine.

ce’. Naviculo-cuboid and external cuneiform bones “separated ;

feet with pouch-like pedal glands and deep heel-tie.

d. Antlers present in males only; prepuce long, scrotal in

position ; feet compact, tightly tied at the heels; rhi-

narium large and normal........ Odocoileine.

d’. Antlers in both sexes; prepuce “short, “abdominal; ‘feet

widely separated at the heels; only the labial portion of

the rhinarium retained: .-........csesc eeceecetensteceecerceseens eangifentna.

6’. Vomer not dividing the posterior nares.

e. Antlers absent, male with large upper canine tusks; inguinal

glands present ; tarsal and metatarsal a absent; pedal

elands deep long clefts.. beeen .. Hydropotine.

. Autlers present in males ; ‘upper _ canine teeth absent or

ninute; no inguinal glands; tarsal o1 metatarsal glands

goa pedal glands a deeper shallow pouch on hind

oot.

* Caton put on record the very interesting fact that in reindeer calves there are

traces of a glandular pouch on the fore foot. This suggests that the pedal elands

in Rangifer were originally present on both fore and hind feet, as they are in

Odocoileus.

CHARACTERS OF ELAPHURUS AND OTHER CERVIDA. 207

Ff: Muzzle short, not swollen; rhinarium large and normal;

preorbital and tarsal glands absent, etc. .... Capreoline.

f’. Muzzle very long and much sw ollen; rhinarium reduced

to a small triangular patch between the anterior ends of

the nostrils; preorbital and tarsal glands present, etc.... Alecine.

a’. Distal ends of laterai metacarpals lost, their proximal ends

usually retained, rarely vestigial or absent (Plesiometacarpalia).

g. Naviculo-cuboid and external and median cuneiform bones of

the tarsus fused into a single bone. Large upper canine tusks

present in males; antlers supported on a long hairy pedicel... Muntiacine.

g'. Naviculo-cuboid and cuneiform bones separated. Upper

canines small or ahsent; antlers supported onashort pedicel. Cervine.

Many of the subfamilies mentioned above—namely, Rangi-

ferine, Hydropotine, Capreoline, an:1 Alcine—are monotypical.

The Pudinge contains the two genera Pudu and Pudella, the

latter differing from the former in the loss of the preorbital

gland and the lacrymal pits, and in having the first lower incisor

much larger than the second.

To the Odocoileine I refer, in addition to the typical genus,

Mazama, Hippocamelus, and Llastocerus, although the inguinal

position of the prepuce and the structure of the feet are un-

known in these three. If one or more of them prove to have

the prepuce abdominal, I should separate such forms as a distinct

subfamily on that character alone.

The Muntiacine* contain three genera—I/untiacus (= Cer-

vulus), Procops T, nov., and Elaphodus. The type of the new genus

. Procops is Cervulus fee, Thomas & Doria, which is generi “ically

separated from Muntiacus by the absence ‘of the frontal glands,

a character in which it resembles Hlaphodus.

The Cervine, containing most of the Old World deer, are a

highly diversified group, composed of the following well-defined

genera :—Dama, Axis (+ Hyelaphus), Cervus, and Elaphurus.

But Cervus itself is subdivisible into several minor groups—Lusa,

Sika, Rucervus, and Cervus itself—which in the future will

probably take full generic rank.

* Established by Garrod under the name Cervuline (P. Z. 8.1876, p. 757), based on

Cervulus, which is antedated by Muntiacus now in general use.

+ From Prox, a generic name applied by Ogilby to the Muntjacs.

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ON THE CLASSIFICATION OF THE SCIURIDA. 209

(7. The Classification of the Sciuride,

By R. I. Pocock, HRs.

[Received February 5, 1923: Read March 20, 1923. ]

(Text-figures 18-29.)

ConTENTS.

Page

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Description of the Penis and Baculum ..................... 212

1. The Palsearctic and American Species ............ 212

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Sh INOS AD ANCHE SHAE oocouhvocuerpapy ouaoscesaccone EAS

4, The African Ground-Squirrels........................ 230

5. The Sousliks and Marmots ........................... 238

6. The Subfamilies of Sciuride ........................ 286

Phe Hays) ID Kyavayse Kio fHyb ORNS oe coh ohanateacusedecoacoseconsseone LO

INTRODUCTION.

In the following brief review * of the more important attempts

to classify the Sciuride, it is unnecessary to do more than quote

Gray’s papers, published in 1867 (Ann. Mag. Nat. Hist. (3) xx.

pp. 270-286, 323-334, 415-436). The results he achieved were

of no great moment from my present point of view, but he

introduced a number of new sectional names which have to be

borne in mind. In 1880, Trouessart (Le Naturaliste, i. pp. 290—

293, 315) made a similar but more successful and useful effort,

proposing several new names but ignoring those of Gray as

connoting groups composed of unrelated elements. He divided

Sciurus into seventeen subgenera, amongst which appear such

well-defined forms as Lheithrosciurus and Xerus, which he regarded

as equivalent to Neosciurus, Parasciurus, Hchinosciurus, and

Tamiasciurus, dismembered from Sciurus on certain comparatively

trivial characters presented by some of the American species.

The most important contribution to the subject was made in

1893 by Forsyth Major (Proc. Zool. Soc. 1893, pp. 186-190) from

a study of the teeth and skulls. He recognised the three sub-

families Sciurine, Pteromyine, and Nannosciurine. The Sciurine

* T have not here taken into account the numerous papers on American Sciuridie

which belong to four well-marked and universally admitted types—namely Sciwrus,

Tamias, Citellus, Cynomys, and Marmota. As compared with the true arboreal

Squirrels of the Old World, the American species of Sciurus are singularly uniform

in essential characters, although a large number of subgenera are admitted. Citellus

has been similarly broken up into subgenera, and Hutamias has been dismembered

from Lamias.

210 MR. R. I. POCOCK ON THE

he divided into six genera :—(1) Rhetthrosciurus; (2) Xerws; (3)

Sciurus; (4) Citellus (Spermophilus)*; (5) Marmota (Arctomys) ;

(6) Cynomys. Of these, Rheithrosciurus, Citellus, Marmota, and

Cynomys were not subdivided; but both Yerus and Seirus

contained several subgenera.

Xerus was divided into five:—(1) Protoxerus for stangeri,

aubinnii and other African species ; (2) Xerus for rutilus, capensis,

and erythropus; (3) Atlantoxerus for getulus; (4) Paraxerus for

cepapi, congicus, isabella, lemniscatus, aud other African Squirrels ;

(5) Funambulus (Howerus) for palmarum, tristriatus, and other

related Oriental Squirrels. To systematists the interesting point

to notice in connection with Forsyth Major’s conception of the

genus Xerus is the inclusion in it of certain soft-furred arboreal

species of African Squirrels (Profoxerus and Funisciurus) and of

the Oriental Palm Squirrels (Yunambeulus), which were previously

regarded as more nearly related to Sevurus.

Sciwrus, according to Forsyth Major, comprised three sub-

genera :—(1) Ratufa (Hosciurus) for the large Oriental Squirrels

indica, bicolor, etc. ; (2) Sciurus, sensu stricto; (3) Zanias for the

Chipmunks or Chipping Squirrels. atufa and ZYamias were

undivided; but the species of Sctwrus were classified in four

unnamed groups: (a) comprising certain African species, annu-

latus, punctatus, rufobrachiatus, and others; (3) Oriental species,

prevostt, caniceps, notatus, ferrugieus, etc. ; (y) vulgaris, syriacus ;

(6) carolinensis, estuans, abertt, and other American species.

In 1897 (Proc. Zool. Soc. 1897, p. 933) Thomas revised

Majov’s classification in nomenclature and other points—so far as

the genera Rheithrosciurus, Xerus, and Sciwrus were concerned.

Granting generic value to all Major’s subgenera, with the ex-

ception of Atlantoweruws which remained a subgenus of Xerws, he

assigned the Squirrels to the following eight genera :—Lheithro-

sciurus, Protowerus, NXerus, Funisciurus, Funambulus, Ratufa,

Sciurus, and Zamias. Under each of these generic names, apart

from Rheithrosciurus and Protoxerus, one or more subordinate

names appear; but it is not in all instances clear whether these

were cited as connoting subgenera or merely as synonyms. In

the case of Yerws it seems certain that Geosciurus and Atlantoxerus

stood for subgenera; and probably that value was assigned to

some at all events of the named divisions of Seizes, such as

Callosciurus and Tamiasciurus. But it is not likely that Rukaria

and Hosciurus were looked upon as subgenera of Ratufu; and it

is certain that Palmista was quoted merely as a synonym of

Funambulus. Nevertheless great service was done by nailing

each name so quoted to a type-species.

In 1898, De Winton (Ann. Mag. Nat. Hist. (7) ii. pp. 12-13)

attempted to show that Forsyth Major’s genus Protowerws was

heterogeneous, some of the species being Xerus-like and belonging

** In the case of nos. 4 and 5, I have adopted the names now in use for the genera,

those employed by Forsyth Major being put in brackets.

CLASSIFICATION OF THE SCIURIDE. 211

to Funisciurus (= Paraxerus Major) and others to Major’s

section a of the genus Sciwrus, for which Trouessart’s name Helio-

sciurus was available. The interest of this paper lies in the

circumstance that two authors working on the same material and

using the same characters, namely skulls and teeth, came to very

different conclusions. De Winton also commented on the

curiosity and inconvenience of the fact that in the African

Squirrels that approach Yerws in harshness of fur the skull and

teeth are Sctwrus-like, whereas those with softer fur approach

NXerus in cranial characters.

In a paper upon the African Squirrels (Ann. Mag. Nat. Hist.

(8) iu. pp. 467-475, 1909) Thomas revised de Winton’s and

Major’s conclusions with respect to the Ethiopian species. As

the result of a more exhaustive examination, he came to the

conclusion, mainly on the evidence of skulls and teeth, that no

fewer than twelve genera should be admitted :—WSciurus for

poensis and two others; Heliosciurus for gambianus, punctatus,

rufobrachiatus, ete.; Myrsilus for aubinniw and one other ; Muni-

sciurus for isabella, leucostigma, lemniscatus, etc.; Paraxerus for

cepapi, palliatus, pauli, etc.; Protowerus for stangeri alone;

Epixerus for ebit and wilsoni; Atlantowerus for getulus; Xerus

for brachyotus and rutilus; Geosciurus for capensis; Huxerus for

erythropus and microdon; Myosciurus for minutus.

With the exception of Myosciwrus, retained in the subfamily

Nannosciurine, all the rest were referred to the Sciurine. But

the genera of Sciurinze were divided into two sections, A and B,

B containing the four genera <dAélantoxerus, Xerus, Geosciurus,

and Huwerus, distinguishable from the rest of the Sciurinee by

their coarse fur and cranial characters.

In connection with these attempts to arrive at the truth with

regard to the affinities of the species of Squirrels, it may be noted

that both Forsyth Major and Thomas retained a certain number

of typical African and Asiatic forms in the genus Sciwrus, of

which S. vulgaris in the type. This affiliation resulted from the

evidence supplied by skulls and teeth; but the conclusion very

forcibly suggested by the literature of the subject is the un-

trustworthiness of such characters.

This conclusion as applied to typical Squirrels was confirmed

by Thomas in 1915 (Ann. Mag. Nat. Hist. (8) xv. pp. 383-387)

from a study of the penis bone or baculum. To the details of

this paper I shall refer in the following pages; but attention

may here be drawn to: one or two important points that were

clearly brought to light :—(1) None of the tropical African * or

Asiatic species, except Rheithrosciurus, shows any near affinity to

Sciurus. (2) All the American Squirrels ranging from Canada

* To the African species assigned to Sezurus in 1909, Thomas subsequently gave

the name AMthosciurus, with poensis as the type (Ann. Mag. Nat. Hist. (8) xvii.

p. 271, 1916), and in 1918 (Ann. Mag. Nat. Hist. (9) i. p. 33) he dismembered the

black-striped species of Parawxerus from that genus, calling them Tamiscus.

212 i MR. R. I. POCOCK ON THE

to South Brazil are closely related to Seiwrus. (3) The subfamily

Nannosciurinze must be abolished as composed of heterogeneous

elements, alike only in convergent characters *.

In 1923 (Proe, Zool. Soc. 1922, pp. 1171-1212) I published a

paper on the external characters of all the species of Sciuride

available to me as fresh or spirit-preserved specimens. But for

want of adequate material I was compelled to omit consideration of

the penis and baculum. ‘lo Mr. Thomas's kindness in lending me

the collection of bacula prepared under his orders at the Natural

History Museum, I now owe the opportunity to figure and

describe, not only the specimens of this bone deseribed in general

terms in his paper, but also those of a few Petauristide, of

Citellus, Xerws, and some others which he intended, granted time,

to work out himself. To the account of these bacula I have added

descriptions of the penis and bacula of such specimens of Sciuridee

as I had preserved for the Zoological Society's collection.

DESCRIPTION OF THE PENIS AND BacuLum.

1. The Palearctic and American Species.

In Seiwrus vulgaris the apex of the penis is compressed and

upturned where the terminal blade of the baculum runs into it.

The orifice opens on the right side behind and below the tip, and

is surmounted by a small fleshy lappet; and the small process of

the blade of the baculum forms a marginal projection just below

and in front of the orifice. The soft swollen tissue enveloping the

apex of the penis is continued backwards as a curved crest from

above the orifice to terminate near the middle of the dorsal

surface of the glans; and laterally and inferiorly it is defined by

a groove from the wrinkled, more slender, and rod-like proximal

portion of the glans. My observations on this species agree with

those of Tullberg, who figured the penis and the extracted

baculum from two points of view. (Text-figs. 18, A; 19, A, B.)

I find, moreover, that the glans penis of Sctwrus (Neosciurus)

carolinensis is of precisely similar structure. Thomas also

described the baculum of S. vulgaris, and pointed out that the

type of baculum found in that species occurs also in a number of

other forms from Asia and North and South America. In the

collection of the British Museum the bacula of the following

species demonstrate this truth:—W. vulgaris from Europe, persicus

from Asia Minor, lis from: Japan, niger from North America,

carolinensis from Tennessee and England (imported), nayoritensis

from Jalisco in Mexico, langsdorfi from the Sierra de Chapada,

ingrami from Rio Janeiro, and variabilis from South America.

And there is no reason to doubt that the glans penis in all these

species conforms to the shape of the baculum. In view of these

* True at all events of the African Myosciwrus and the Oriental Nannosciurus,

which respectively fall into line with other species of the regions they inhabit. But

the baculum of the South American genus has yet to be examined.

CLASSIFICATION OF THE SCIURIDA. 213

facts, Tullberg’s statement that the North American species

LTamiasciurus hudsonicus, always regarded as closely akin to the

other Squirrels of that country and as ranking merely as a sub-

genus of Sciwrus, has an elongated apically attenuated glans penis

Text-figure 18,

Ay i a Hi

A. Glans penis of Seiwrus vulgaris.

B. The same of Tamiasciurus hudsonicus.

C. The same of Funambulus palmarum.

D. The same of Tamiodes tristriatus.

KH. The same of Callosciurus prevosti. ?

F, Penis of the same, with baculum attached, showing position of blade.

G. Glans penis of Callosciurus notatus.

H. Apex of same from right side, showing orifice.

I, K, L .The same of Tomeutes vittatus trom right side, above, and left side,

o. Orifice.

without a baculum, appeared to me to be so remarkable and

contrary to all expectations that I wrote to Mr. Gerrit S. Miller,

of the Smithsonian Institution, Washington, to ask if he would

be good enough to verify the statement for me, In reply, he very

Proc. Zoo. Soc.—1923, No. XV. 15

214 MR. R. Ie POCOCK ON THE

kindly arranged for two male examples of the species, preserved

in alcohol, to be sent to me*. From these two specimens I find

that Tullberg was perfectly correct, and had not, as I supposed,

made an error of identification. The glans penis in the Alaska

specimen is slender and tolerably evenly tapering, with the

orifice terminal and just beneath the apex. The upper surface

has a median groove, and a pair of crests of minute serrula-

tions running along it, and the sides are finely striolate. The

distal portion below the bend is somewhat flattened, and provided

on each side with a narrow laminate expansion. In the New

York specimen, assigned to the subspecies loguas, the general

form and sculpturing of the penis are much the same, except that

the striole are coarser, the intervening spaces forming fine ridges,

a difference possibly due to greater contraction of the epithelium,

and the distal portion beyond the bend is not so evenly tapering,

but shows an excrescence on each side. Nevertheless the two

are of essentially the same type, and in both the distal portion

is soft and flexible without a vestige of baculum that I can detect.

The structure of the penis sharply differentiates hudsonicus from

the Squirrels with the vulgaris type of penis and baculum, and

compels the admission of Tamuasciurws as a well-defined genus

without any near allies in the family. (Text-fig. 18, B.)

On the other hand, although Thomas stated that the baculum

of the Bornean species Rheithrosciurus macrotis is like that of

Sciurus vulgaris, he tells me that he is so impressed by the

improbability, on geographical and other grounds, of the two

species being alike in that respect T, that he considers confirmation

of the fact necessary before definite acceptance be accorded it.

There is always, of course, the possibility of confusion between

the bones by taxidermists during the cleaning process.

The bacula of the S. vulgaris group are relatively well

developed, but vary in size with the species, from about 14 mm.

in S. niger to 8 mm. in S. ingrame.

The foregoing account shows that the Palearctic and American

Squirrels fall into two categories by the structure of the penis

and baculum :— ,

1. Sciwrus, embracing the Palearctic species, all the Nearctic

species except the one quoted below, and the Neotropical

species.

2. Tamiasciurus, represented by hudsonicus and possibly related

species.

* My best thanks are due to the authorities of the Smithsonian Institution for the

generous loan of these two Squirrels. One, in winter pelage (January), was collected

at Nulato in Alaska by W. H. Dall, the other, m summer pelage (September), at

Tamarak Swamp, New York, by E. A. Mearns.

+ Although Rheithrosciurus, with its grooved incisors, has always been regarded

as an isolated genus, the discovery that it is really related to Sciwrus (s.s.), on the

evidence of the baculum, would surprise me much less than did the discovery of the

wide divergence between Tamiasciurus and Sciurus in the structure of the glans

penis and the occurrence of the baculum.

CLASSIFICATION OF THE SCIURIDA, 215

2. The Oriental Species.

In Funambulus palmarum the glans is long, subeylindrical

proximally, swollen and spongy in texture distally, and terminates

in a longish, narrow, attenuated process, slightly or considerably

inclined upwards and rising from the dorsal extremity of the

swollen portion. The orifice lies at the base of this process at

the right side. The baculum isa simple curved rod, concave on its

upper side, convex below and tolerably gradually attenuated from

the thick base to the narrow apex, which reaches to the extremity

of the above-described terminal process of the glans. The

Text-figure 19.

A. Baculum of Sciurus (Parasciurus) niger, from the left side.

B. Apex of the same from the right side.

C. Baculum of Protoxerus stangeri, from the left side.

D. Baculum of Funisciurus congicus, from below, with the soft, dried,

shrivelled tip of glans adherent.

K. The same of Paraxerus cepapi, with the tip of glans removed.

I’. The same from the left side.

curvature varies apparently individually. The bone measures

10°5 mm.—that is to say, it 1s actually as long in this small

Squirrel as in the giant Ratufa. (Text-figs. 18, C ; 20, D.)

In Tamiodes (gen. nov.) tristriatus, regarded by Blanford as

possibly nothing but a local race of palmarum, the glans is very

different. It is relatively shorter and thicker, and ends in

two well-marked subequal labia, one above and one below the

terminal orifice. The upper labium is unmodified ; but the lower,

- which has its inferior edge inclined upwards, is subdivided into a

number of subordinate labia by deep lateral clefts, which are

15*

216 MR. R. I. POCOCK ON THE

represented on the lower side of the tip by grooves running

obliquely backwards and inwards towards the middle line. It

was clearly, I think, to a Squirrel of this genus, possibly indeed to

tristriaius, that the glans penis figured and described by Tullberg

as that of palmarum belonged. His figure, however, does not

show such a definite lower lip as was present in the example from

Malabar above described, the area beneath the upper lip being

supplied with two rows of lobules representing the subdivisions

of the lower lip in my specimen. (Text-fig. 18, D.)

The difference between /. palmarum and T. tristriatus in the

structure of the penis is reflected in the shape of the baculum,

which in 7’. éristriatus, instead of being gradually curved upwards

from base to apex, has the proximal three-fourths curved towards

the right with the right side concave, the left convex, and the

distal fourth bent up at a right angle to form a vertical branch

with an expanded tip, a convex crested posterior edge and a

concave anterior edge. Erom the inferior angle of this arises a

much smaller branch directed downwards. ‘his description is

taken from a baculum, measuring 11°5 mm., taken from a specimen

of tristriatus from Helwak, Satara, and preserved in the British

Museum. Another baculum in that collection, taken from an

unidentified specimen without locality, differs in having a less

expanded apex to the upper branch, no crest on the latter ard no

inferior branch, the shaft being straight. It measures 11mm., and

may represent a different race or species. (Text-fig. 20, A, B, C.)

In the example from Malabar of which the glans penis was

described, the apex of the upper branch projected a little way

into the upper lip of the glans, and the lower branch formed the

bulge at the inferior part of the base of the lower lip. The

baculum, measuring 12 mm., is similar in curvature and other

particulars to that of the example from Satara, except that the

inferior branch is more quadrate and not hook-like.

‘The two genera of Oriental Squirrels described above are a most

instructive instance of close superficial resemblance associated

with great differences in the glans penis and baculum. They are

so much alike that Blanford was doubtful if trzstriatws was more

than what he would have called a ‘‘ variety” of palmarum. Yet

it is impossible to believe in close kinship between them; and the

differences explain the occurrence of the two in the same districts

in India without ever intergrading—that, at all events, I believe

to be the case.

The structural characters of the two may be recapitulated as

follows :—

a. Glans penis ending distally ina narrow elongated point supported

by the distal portion of the baculum; the orifice on the right

side near the base of the slender portion and remote from the tip

of the glans. Baculum a simple upcurved attenuated rod ...... Funambulus.

a’. Glans penis thick, blunted, and bilabiate distally, the orifice

terminal between the upper and lower lips, the lower lip

secondarily subdivided. Baculum with its terminal portion

bent vertically upwards and curved forwards, with a _pro-

THOUS WIHT OND A NVeR She he ace ne doAcderer.cbieshoaeGacdoncooban conoicctenccnny, ol Meme kes.

CLASSIFICATION OF THE SCIURIDA. nla

The baculum of an example of Ratufa gigantea from Gokteik,

N. Shan States, in the British Museum, is simple and shaped in a

general way like that of Funambulus palmarum, but is relatively

much shorter and stouter. It is expanded at the base, and in

lateral view its upper and lower edges are respectively concave

and convex, but with a sinuous outline. The distal end of the

lower surface is irregularly bevelled up to the apex, which is

narrowed but not pointed, and there are four subsymmetrically

arranged rows of small irregularly spaced bony spicules on this

distal portion, which from the dorsal or ventral aspect is seen to be

slightly asymmetrical. It measures10mm. (Text-fig. 20, E, F.)

The baculum of a specimen, referred to the same species, from

Darjiling is very similar, but more strongly bevelled and upcurved

distally, the apex being a little expanded; and one of R. fellii

from Yin, L. Chindwin, is also very like it. But a baculum of

a captive specimen * of &. indica, without locality, also in the

British Museum, is very different. It is a much larger and

longer bone, about twice the length of the others if straightened,

and has its distal half bent strongly upwards at an obtuse angle,

a shoulder-like excrescence on each side marking the bend. This

distal upturned portion is quite symmetrical, expanded basally

and apically, concave laterally, hollowed inferiorly, and tridentate

at the tip, the median tooth being a little longer and set further

back than the laterals. This baculum, measuring in a straight

line from base to apex 14 mm., may be perfectly normal. Its

strongly upturned distal portion evidently corresponds to the

slightly upturned denticulated, bevelled distal pertion of the

other bacula of the genus. (Text-fig. 20 G, H.)

Setting aside Lhetthrosciurus macrotis, above referred to as

probably having a baculum shaped like that of Sciurus vulgaris,

the rest of the Oriental genera of Sciuride, as Thomas showed,

have bacula specialised by the development, on the upper side, of a

blade which is hinged to the shaft in such a way as to be slightly

movable from right to left. The blade projects backwards, and a

longer or shorter portion of its posterior end is free from the

shaft. Bacula of this kind belong to two main categories—one,

characteristic of Callosciuraus, the other of Tomeutes. But distinct

as the bacula of these two kinds are in their extreme forms,

there are indications of intergradation between them +.

In Callosciurus prevosti, the type of Callosciurus, the glans is

* Bacula of mammals kept in confinement are sometimes abnormal.

+ In his preliminary description of these bacula Thomas wrote :—“ In position in

the penis the blade points to the right, its edge cutting outwards.” Again, the baculum

of Callosciurus is said to have “a narrow blade set on the side of it, in the con-

cavity of its general curvature... . the greatest breadth [of the blade] is only about

one-fourth to one-sixth of its length.” This, however, does not agree with my

observations, for I find in all the fresh or spirit-preserved material examined that the

concavity of the shaft of the baculum and the blade are on the dorsal or upper side of

the bone. If the blade were on the right side, the asymmetry of the baculum would

be extreme and the edge of the blade would cut the right side of the vulva. I have

no doubt that the function of the blade is to make a vertical cut of the integument

(hymen) covering the orifice of the vagina.

218 MR. R. I. POCOCK ON THE

long, slender, nearly straight and compressed, narrowed and

pointed at the end, with the orifice just near the tip on the right

side. There is comparatively little fleshy tissue enveloping the

baculum, and the upper edge of the blade of the latter forms a

Text-figure 20.

ial

A. Baculum of Tamiodes tristriatus, from Satara, from the left side.

B. The same from above.

C. The same of another specimen, without locality.

D. Baculum of Funambulus palmarum, from the left side.

Hi. The same of Ratufu gigantea, from the left side.

E. The same from below.

G. Baculum of captive specimen of R. indica, from the left side.

H. Distal end of the same, from the front.

hard, sharp ridge, covered merely by a coating of fine membrane,

at the distal end of the dorsal side of the glans. The baculum

itself is long and slender, gradually attenuated from base to

point and with a sinuous curvature, being slightly concave in the

CLASSIFICATION OF THE SCIURID. 219

middle below and above in its distal third. The blade, about one-

third of the length of the entire bone, is lodged in the distal con-

cavity. It has a straight upper edge, not rising above the median

dorsal convexity of the shaft. Its posterior third is pointed and

free from the shaft. Its anterior end is attached immediately

behind the slightly expanded tip of the shaft. Hence the blade

is at the distal end of the shaft. The baculum above described

measures 26 mm., the longest in the Sciuride. (Text-figs. 18,

Ss) 2B)

Callosciurus notatus, from Java, has the glans shorter than in

C. prevosti and with a pronounced dorsal curvature in its distal

half, so that its upper surface is concave, its lower surface convex.

The sharp distal edge formed by the blade is more pronounced,

and the orifice is situated as in C. prevosti. The baculum differs

from that of C. prevosti in being relatively stouter, in having

a larger thickening at the proximal end, a strongly upcurled

distal half with a concave upper and a convex lower edge; but

the blade, although similarly placed distally, is not so pointed

and its upper edge is above the line of the edge of the shaft

preceding it. (Text-figs. 18,G, H; 21, C.)

In specimens of C, castaneoventris from Ningpo and of C. airo-

dorsalis from Siam the shaft of the baculum resembles that of

C. notatus, although it is less strongly curved upwards; but the

blade, which is considerably longer, being nearly half the length

of the shaft, is not attached to its distal end, which projects freely

beyond the anterior end of the blade, the blade itself lying nearly

in the middle of the dorsal concavity of the shaft. In C. castaneo-

ventris the baculum measures 19 mm. (Text-fig. 21, A.)

In Thomas’s preliminary account of the bacula of the Oriental

Squirrels, (. prevosti was one of the species in the list of un-

examined forms, whereas C. notatus, castaneoventris, and atro-

dorsalis appear in the list of species of which the baculum was

examined, The remaining species of this list were caniceps,

erythreus, pluto, and sladent.

The differences between the bacula of C’. prevosti and C. notatus

on the one hand and of C. castaneoventris and C. atrodersalis on the

other suggest the possibility of usefully splitting Calliosciwrus

into two genera. The characters appear to me to be at least as

valuable as the presence or absence of a degenerate premolar.

But it would perhaps be premature to take that course, at all

events until it has been shown that the bacula of other species do

not show intermediate stages in the position of the blade.

The baculum of a specimen of Menetes berdmorei from Kokareet

(Tenasserim) in the British Museum very closely resembles that

of C. prevosti described above, except that the shaft is straight

up to the concavity containing the blade. It is convex below

only at its distal end and correspondingly concave above, the

blade lying in the distal concavity as in C’. prevosti and C. notatus.

The baculum of this species, measuring 11 mm., was held by

Thomas to have a blade-somewhat connecting the types of bacula

220 MR. R, I. POCOCK ON THE

characteristic of Callosciurws and Tomeutes. It does not seem

to me that its baculum is more Zomeutes-like than that of

C. notatus. The partial intergradation between the bacula of

O. prevosti, the type of Callosciwrus, and of 7’. lokroides, the type

of Tomeutes, is effected, as explained below, by C. notatus and

T.. vittatus. (Text-fig. 21, D.)

The bacula of what may be called the Tomeutes type differ from

those of the Callosciurus type above described in having the shaft

of the bone shorter and thicker, and the blade deeper and longer,

with a shorter hinged area and a longer portion free from the

shaft. ‘The shaft is variable in length, being sometimes twice the

length of the blade, the point of which falls considerably short of

its proximal end or base, or sometimes a little shorter than the

blade, the point of which then overlaps the base of the shaft. In

all cases probably the glans penis is shorter and thicker than

in Callosciurus.

In Zomeutes vittatus the glans penis is very like that of Callo-

sciurus notatus, but.is relatively shorter and thicker. Its upper

and lower edges are parallel ; along its upper runs a sharp carina

formed by the edge of the blade of the baculum and ending

distally in a short point. On the right side about half-way

between this point and the inferior rounded angle of the distal

end opens the orifice, which has a little fleshy lobe on its outer

side; and from this a groove runs backwards along the right side

of the glans nearly to its proximal end. (Text-fig. 18, I, K, L.)

The baculum, measuring 14 mm., although conforming in a

general way to the typical Zomeutes type of baculum, is more like

that of Callosciurus notatus than any I have seen. The shaft is

about twice the length of the blade, very thick at the base, slender

beneath the blade, with lightly convex but sinuous inferior

border, and upturned at the apex, which is furnished with a

small button-shaped thickening projecting an appreciable distance

beyond the distal end of the blade. The latter, lying in the con-

cavity of the upper side of the shaft, has a nearly straight, smooth

upper edge, the lower edge, also nearly straight, running obliquely

upwards and backwards to meet it in a point. The hinged

portion ends inferiorly in an expansion on each side which, as it

were, clamps the blade to the shaft, and corresponds to the

lateral expansion of the blade seen in Callosciurus notatus and

castaneoventris. This species, Tomeutes vittatus, appears in

Thomas’s list of unexamined species of Callosciwrus. From the

superficial resemblance between vittatus and notatus, he probably

inferred that their bacula would be alike. That, however, is not

the case. (Text-fig. 21, H, F.)

The baculum of a specimen of the type of the genus Tomeutes,

namely 7’. lokroides, from Kursiong in Sikkim, preserved in the

British Museum, has a much shorter shaft and bigger blade than

that of 7’. vitiatus, and is altogether more specialised. The shaft

has a thickened proximal end with the distal third sharply

upturned, making a deep, short concavity. The blade, measuring

CLASSIFICATION OF THE SCIURIDZ. DIAN

8 mm., is longer than the shaft, its blunted point overlapping

the thickened proximal end of the shaft, and the upper edge of

the blade rises high above the concavity of the bone. In this

respect it differs from the baculum of 7. vitiatus, where the

upper edge of the blade does not project above the concavity

lying between the apex and base of the shaft. (Text-fig. 21, G.)

Text-figure 21.

. Baculum of Callosciurus castaneoventris, from the left side.

. The same of C, prevosti.

. The same of C. notatus.

. The same of Menetes berdmorei. —

. The same of Tomeutes vittatus, from the left side.

. The same from above.

. The same of 7. Zokroides, from the left side.

. The same of 7. hippurus.

. The same of T. robinsoni.

. The same of 7. miniatus.

. The same of 7. tahan.

EAH Deets Sem

The baculum of a specimen of 7. hippurusfrom Bukit Tangga,

Negri Sembilan, also has the blade overlapping the proximal end

of the shaft, but not to the same extent as in 7’. lokroides. The

shaft, moreover, is relatively longer, with a longer concavity ; the

blade is not so deep and does not project so high above the con-

cavity of the baculum as in 7. lokroides. In a measure this

22% MR. R. I. POCOCK ON THE

baculum serves to connect those of 7’. vittatus and J. lokroides.

(Text-fig. 21, H.)

The baculum of a specimen of Yomeutes miniatus from Bukit

Kutu, Selangore, has the shaft tolerably similar to that of

7’. lokroides, but the blade is slightly shorter than the shaft and

much thinner, being concavely emarginate inferiorly so that

there is a larger space between its lower border and the subjacent

upper edge of the shaft. There are also a few bony spikes on

the inferior edge of the shaft at the base of the upturned distal

portion. The bone measures7-5 mm. This species, Mr. Thomas

teils me, comes near 7’. vittatus in general characters; but the

great differences in their bacula preclude the view of close kinship

between the two species. (Text-fig. 21, K.)

In a specimen of Yomeutes robinsoni—a species akin,

Mr. Thomas tells me, to 7’. lowii—from Bukit Tangega, Negri

Sembilan, the baculum has the shaft short, stout, and strongly

curved, being very like that of 7’. miniatus but somewhat thicker

proximally, shorter, and without trace of the bony spicules. The

blade, too, is tolerably similar but shorter, with a lightly convex

upper edge, with a notch and tooth. The bone measures 6 mm.

(Text-fig. 21, I.)

Inaspecimen of 7’. tahan (allied to 7. tenwis), from Bukit Kutu,

Selangore, the shaft is relatively longer and thinner than in the

previously described species, except 7’. vittatus, and its inferior

angle, very obtuse, is near the middle of the lower surface

approximately beneath the point of the blade. The blade itself

is short and deep, with a long hinge, a lightly convex upper edge,

a strongly concave lower edge, and a narrow apex. The bone

measures mm. Judging from the baculum alone this species

would be referred to Dremomys. (Text-fig. 21, L.)

The main distinctive features of the bacula of the species of

Tomeutes above described may be summarised as follows :—

a. Shaft comparatively long, about twice as long as blade, its lower edge

lightly convex, its apex projecting well beyond blade ; upper edge of

blade not higher than line tangential to upper proximal and distal

ends of shat. ai. ua 2. i deat aeey coer ann Meany Sk ain eM Leet or 768

a’. Shaft comparatively short, much less than twice as long as blade,

its lower edge strongly curved or geniculate, its apex only surpassing

blade slightly ; upper edge of blade higher than line tangential to

upper proximal and distal ends of shaft.

6. Angle of lower edge of slender shaft set far back almost beneath

point of blade, distal portion beyond bend inclined gradually

upwards; hinge nearly as long as blade ............cccceececcceaeeeeeeee” EUAN.

b'. Lower edge of stout shaft with pronounced angle in front of lower

end of hinge, distal portion beyond angle tipped steeply upwards ;

hinge much shorter than blade.

ce. Blade with apex not overlapping proximal end of shaft, its lower

edge deeply and widely emarginate towards hinge ... miniatus, robinsoni.

e’. Blade with apex overlapping proximal end of shaft, its lower

edge nearly straight up to small concave notch by hinge.

lokroides, hippurus.

According to Thomas the following species belong to

Tomeutes :—phayrei, blanfordi, pygerythrus, janetta, pryeri,

CLASSIFICATION OF THE SCIURID”. PHYS

philippinensis, melanogaster, tenuis, brookei, lowii, murinus. I

have not seen their bacula; but those described above are

sufficient to show how variable this bone is within the limits of

the genus as admitted by Thomas. The variation, indeed, is so

great that it is impossible to affirm any character by which the

baculum of Tomeutes can be distinguished from the bacula of

Dremomys, Rhinosciurus, T amiops, and Lariscus. It is not

improbable that a careful comparison between the bacula of all

the species assigned at present to Zomeutes will give useful indi-

cation of interspecific relationship.

Text-figure 22.

A. Baculum of Dremomys lokriah, from the left side.

B. The same of D. rufigenis.

C. The same of D. rufigenis fuscus.

D. The same of D. dawsoni.

KH. The same of Lariscus jalorensis.

K. The same of Rhinosciurus robinsoni.

G. The same of Tamiops maclellandi barbei.

H. The same of Nannosciurus microtis.

I. The same of IV. whiteheadi.

K. The same of XW. evilis.

The baculum of a specimen of Lariscus jalorensis from Bukit

Tangga, Negri Sembilan, is represented by the blade and the

piece of the shaft to which it is attached. This portion of the

blade is compressed and shows no angulation of its inferior

border, which suggests that the upcurvature was gradual as in

Tomeutes Mitatus or Dremomy ys dawsoni, Its apex is produced

as an acute angle beyond and in the same line as the upper edge

of the blade, which is nearly straight, with a slightly depressed

apex, and forms an acute angle with the line of the hinge very

22.4 MR. R. I. POCOCK ON THE

much as in Tomeutes vittatus. In the latter, however, the apex

of the shaft is expanded and button-shaped, not compressed as in

Lariscus jalorensis. (Text-fig. 22, H.)

The baculum of a specimen of Rhinosciurus robinsoni from

Tiernan Island, in the British Museum, is very like that of

Tomeutes lokroides. The shaft is short, proximally thick, distally

upturned, with a short concavity, with its large blade pro-

jecting high above it and overlapping the proximal end.

The upper edge of the blade is nearly straight, but is

irregularly notched in its posterior half. The bone measures

10mm. (Text-fig. 22, F.)

Ina specimen of Zamiops maclellandi barbei from Kekareet,

in the British Museum, the shaft of the baculum gradually nar-

rows from base to apex, its distal half being upturned at an obtuse

angle. The blade has a long hinge but is short, the point being

very narrow, with a notch and tooth near its apex. Its upper

edge is only slightly higher than a line touching the upper edge

of the proximal and distal ends. The bone measures 5 mm.

(Text-fig. 22, G.)

In fom species of Dremomys, namely, D. lokriah from Sikkim,

D. rutigenis from H’Kauti, Chindwin, D. rufigenis fuscus from

Annam, and JD. dawsoni from Tatken, Chindwin, the shafts of

the bacula are longish and longer than the blades. In that of

D. rufigenis the proximal end of the shaft is very thick and

raised, the median part is straight, and the upcurved distal end

has an evenly convex edge. The upper edge of the blade is

markedly convex to the apex, which falls far short of the proximal

end of the shaft; its lower edge is straight with a pronounced

concavity near the hinge. The baculum of D. rufigenis fuscus is

sufficiently different to suggest specific distinction between the

two forms. The shorter shaft is comparatively narrow proxi-

mally, its inferior edge is evenly convex, the upcurvature beginning

beneath the inferior end of the hinge; the blade is larger as

compared with the shaft, has a longer hinge, an evenly concave

lower border, and a less convex upper border. In D. dawsoni

the shaft overlaps the blade proximally to an even greater extent

than in D. rufigenis, but it is not so stout, and its ‘lower edge is

inchned upwards from a point below the inferior end of the

hinge as in D. rufigenis fuscus, but more abruptly, and the con-

cavity of the lower edge does not extend up to the bottom of the

hinge, the edge at this point being slightly convex. In D. lok-

riah the shaft is very like that of D. rufigenis fuscus, but is a

little stouter distally, and the lower edge of the blade is straight

up to the point where it runs into te thickening forming the

lower end of the hinge. (Text-fig. 22, A—D.)

The bacula of the species of Dremomys above described may

be contrasted as follows :—

a. Lower edge of shaft tolerably straight for the greater part of its

length, distal end of shaft expanded with strongly rounded edge;

blade thinner, with wide emargination close to short hinge ......... rufigenis.

CLASSIFICATION OF THE SCIURIDA, 225

a’. Lower edge of shaft tolerably evenly convex throughout or convex

beneath lower end of long hinge, distal end not expanded, without

strongly convex edge.

b. Apex of blade some distance from geet end of shaft, lower

edge of blade with sigmoid curvature.. gad . dawsoni.

b’. Apex of blade a little behind proximal éndl of shaft, aes alee of

blade without sigmoid curvature.

c. Lower edge of blade evenly emarginate, distal end of shaft not

thickened . pee ee eee mIUILOCIUES US CUSs

. Lower edge us tide es up ig i eeliae notch by hinge;

distal end of shaft to which blade is “hinged somewhat ex-

panded .. gga IRAN SOURS ea ented omenea tine bene tata LOMPOC L

In three species of Vannosciurus—lN. whiteheadi from Kina

Balu, WV. exilis from Borneo, and WV. microtis from Sarawak—the

proximal end of the shaft of the baculum is not thickened and

elevated, and its upper edge, as far as the hinge of the blade, is

almost straight, and is parallel in a general way to the lower edge

of the blade; the tip of the blade “just reaches the base of the

shaft. In NV. whiteheadi and NV. exilis the lower side of the shaft

has a rounded swelling near the middle, and the distal end is only

a little raised ; the blade is arcuate with serrated upper edge and

a narrowed point. In WV. whiteheadi the apex of the shaft, to

which the blade is hinged, is thicker, and the blade itself is more

arcuate and more str rongly serrated than in W. ewilis, In

NV, microtis the shaft has no definite inferior thickening, its

distal half is bent up at an obtuse angle, the blade is not arcuate

nor serrate, and the tip is bluntly rounded and scarcely narrowed

in profile view. (Text-fig. 22, H-K.)

The measurements of these bacula. are as follows:—W. white-

headi 5 wm., NV. exilis 4 mm., V. microtis 3 mm.

In the following table the characters of the bacula of the three

species of Vannosciurus, here described, are briefly summarised :—

a. Shaft with its distal half hent up at an obtuse angle without a

rounded thickening near the middle of its lower surface; blade

with tolerably straight, smooth wpe Ge and ey rounded

LO oageascor ce: ... microtis.

a’. Shaft toler only, alts one axially fomn cal ho ie he cgtremiens apex

a little elevated, a conspicuous thickening near ‘the middle of the

lower surface ; the blade long, apically attenuated , with upper edge

convex and serrulate.

b. Blade high and arched, with wide hace between its lower edge

and the shaft.. ocean 5 , ..... whiteheadi.

. Blade lower, ibsee creed a en narrower space Rhcoreen His

edge and Sic Ate ey ee Ue amma nC ne cE. are exilis.

The Oriental Squirrels thus fall into three main groups, judged

by their bacula :—

1. Rheithrosciurus, belonging to the same group as Sciwrus of

the preceding section.

2. Ratufa, Funambulus, and Tamiodes, with simple bacula.

3. Callosciurus, Tomeutes, Tamiops, Lariscus, Nannosciurus,

etc., which have compound bacula provided with a blade.

MR. R. I. POCOCK ON THE

3. The Hthiopian Species.

In Protoxerus stangert, as identified by Tullberg, the glans

penis is elongated and nearly straight, but exhibits at the distal

end of its proximal half a large swelling, showing a pair of low

crests above and a pair of distally directed pointed processes

below. Beyond the swelling the glans is gradually narrowed

towards the apex, but the apex itself is shapéd like an arrow-

head, with the point slightly Hpbunned and a pair of small

swellings just before it. (Text-fig. 23, M.) _ ...

The baculum of a specimen of this species from the Como

River, in the British Museum, is a symmetrical bone, slightly

concave above, convex towards the middie below, a little upturned

at the apex, which ends in a blunt condyle-like button. On each.

side of the shaft in its distal third towards the apex there is a

faintly defined longitudinal crest. The proximal end, or base, is

much thickened, and carries a coronet consisting of five sym-

metrically arranged bony lobes, one being in the middle line

above and two on each side, the two lower lobes being separated

by a notch in the middle line below. There is no doubt, I think,

that the thickened proximal end of this bone is imbedded in the

tissue forming the submedian thickening of the penis described

and figured by Tullberg, and, from analogy, I suspect the button-

like tip of the baculum is lodged in the thickened portion of

the arrow-headed apex. (Text- “fig. LS Gp)

The actual length of the bone, despite the large size of the

Squirrel, is 7-5 mm.

In Athosciurus poensis, as identified by Tullberg, the glans

penis is quite short, and consists of a thickened proximal portion,

composed of two transverse folds of soft tissue of which the

posterior has a finely serrated posterior border on the upper side,

and of a slender distal portion which gradually narrows from the

base to the simple apex. According to Tullberg, the glans penis

of this species has no baculum, but from analogy I suspect a

small baculum, imbedded in the tissues, was overlooked.

In an example of Munisciurus leucostigma from Bibianaha the

glans penis is not at all unlike that of @. poensis. Its proximal

portion is quite short and thick, consisting of soft grooved tissue,

but the distal portion, abruptly differentiated from the proximal,

is a comparatively slender somewhat flexible rod, tapering apically

but furnished on each side with aserrulated crest starting distally

behind the tip and ending proximally on the upper side of the

thickened base. These two crests define fe upper from the

lateral surfaces of the process. (Text-fig. 23, E-G.

In funisciurus leucostigma niveatus the glans penis differs

somewhat from that of 7. lewcostigma above deseribed, although

consisting of a short undifferentiated proximal pertion, a swollen

laminate and grooved median portion, and a thinner longer distal

serrulated flexible portion. The median portion, however, is not

so thick as in typical /. lewcostigma, and the distal portion is

is)

CLASSIFICATION OF THE SCIURIDA. 27

thicker and more irregular in shape, being markedly asymmetrical

and not so sharply marked off from the swollen median portion.

The orifice appears to lie in the swollen median portion beneath

the proximal end of the distal portion. When at rest within

the abdomen the terminal half of the distal portion lies in the

prepuce and is bent like a hook, and the penis does not extend

beyond the prepuce as in Heliosciurus rufobrachium described

below. (Text-fig. 23, C, D.)

The dried distal portion of the glans of a specimen of

Funmsciurus pyrrhopus from Ashanti, in the British Museum,

agrees in all essentials with that of /. leucostigma, except that

the apex is better defined and more like an arrow-head. The

baculum, a simple rod measuring about 1°5 mm., is imbedded in

the serrulated area, its distal end not reaching the tip of the

glans. :

In a similarly preserved part of the glans of F. congicus, with

the tissue of the serrulated area cut away leaving the baculum

exposed, the baculum is a slightly curved rod about 2 mm. in

length, and the dried tissue of the apex of the glans, still attached

to the distal end of the baculum, is lanceolate. (Text-fig. 19, D.)

In Paraverus cepapi the glans is very similar to that of Puni-

sciurus leucostigma. The proximal portion is very thick and

short, and bent on itself, its convex surface having a median line

of serrulations. Distally this thickened portion shows grooved

flaps of soft tissue. The terminal process is very like that of

Funisciurus leucostigma, being broad at the base, pointed at the

apex, and provided dorso-laterally on each side with a serrulated.

crest. The sides of this process are, however, concave, so that

the dorsal area bordered by the crests is much narrower in the

middle than behind and infront. (Text-figs. 23, H, 1; 19, E, F.)

The diied process of the glans of a specimen of Paraxerus

palliatus from Gululand, in the British Museum, is tolerably

similar, but the serrulated area is somewhat sharply constricted

in front, so that the tip of the process is better defined and more

arrow-headed in shape and the middle of the serrulated area is

broad, broader indeed than its proximal end, where it rises into

a crest. The baculum, measuring less than 2 mm., is imbedded

in the process of the glans behind the apex and between the con-

strictions. (Text-fig. 23, K, L.)

The dried condition of the distal portion of the glans penis of

Funisciurus pyrrhopus and of Parazerus palliatus makes their

comparison with that of the spirit-preserved material of F’. lewco-

stigma and Paraxerus cepapi a little unsatisfactory. But the

general similarity between the penes of these two so-called genera

admits of no doubt. I suspect Tullberg overlooked both the

baculum.and the serrulated crests in the specimen he referred to

Sciurus poensis.

In an adult male of Meliosciurus rufobrachium hardyi the

penis is of exceptional length. When extended it reaches as far

as the posterior end of the sternum, and measured from its

228 MR. RB. I. POCOCK ON THE

origin just in front of the anus, it is more than half the length

of the head and body. Within the abdomen its proximal portion

up to the origin of the glans projects some distance beyond the

prepuce. The glans is very sharply divided into two parts, a

Text-figure 23.

Penis of Heliosciurus rufobrachium hardyi, retracted.

Glans of the same extended, from the right side.

-Glans penis of Funisciurus leucostigma niveatus, from above.

The same from the right side.

Glans of Ff. leweostigma, from the left side.

The same from the right side.

Narrowed distal portion of the same, from above.

Glans of Paraxerus cepapi, from above.

The same from the right side.

Narrowed distal portion of glans of Paraxerus palliatus, from above.

. The same from the right side.

. Glans of Protowerus stangeri, from right side (after Tullberg).

BHAHTR Oe ea oad

proximal and a distal, nearly equal in length. The proximal

portion, when at rest, is folded back beneath the distal end of the

preglandular part of the penis, is dorse-ventrally expanded, and

CLASSIFICATION OF THE SCIURIDE. 229

slightly compressed and striolate laterally. Its upper portion is

lightly convex, smooth, and cartilaginous, but not ossified. Its

lower portion consists of softer tissue, and carries the urethral

orifice on a small fleshy process at the distal extremity of its right

side. The distal extremity of the glans is long, slender, flexible,

and filiform, like a rat’s tail. J can find no trace of baculum in

it, unless the remnant of this bone is represented by a slight

opacity at the distal end of the thread. (Text-fig. 25, A, B.)

In an immature male of Heliosciwrus punctatus the penis

resembles in a general way that of HA. rufobrachiwm, being

exceedingly long and distally slender and filiform without trace

of baculum. ‘The specimen is not well preserved, but the filiform

portion appears to terminate in a small arrow-headed tip and to

pass into the main body of the penis without the intervention of

the swollen glandular portion carrying the orifice such as is seen

in H. rufobrachiwm. The position of the orifice I could not

determine.

The differences above described are very considerable, and if

they are found to obtain in adult examples of H. punctatus, they

would, I suppose, indicate a generic difference between that

species and #H. rufobrachium. But since the example in question

is immature and poorly preserved, the examination of fresh

material must be awaited ere a decision on the point is reached.

Temporarily setting aside the penis of the example of Helio-

sciurus punctatus above described, the characters of this organ as

shown in the other genera may be tabulated as follows :—

a. Penis exceedingly long, the distal portion of the glaus forming

a long, flexible thread-like termination without baculum ......... Heliosciurus.

a Penis quite short ; the distal portion of the glans shorter, stouter,

rod-like, not filiform, and provided with a baculum.

6. The narrow rod-like distal portion flexible, with a finely serru-

lated crest on each side, and carrying a minute baculum near

its distalend ......................... Athosciurus, Funisciurus, Parawxerus.

6’ The narrowed distal portion not fiexible, without serrulated

crests, but supported throughout by a relatively large baculum. Pv otoxerus.

It must be remembered, however, that the characters assigned

to the penis of Protoxerus and dthosciurus are taken from

Tullberg’s descriptions and not from my own observations.

Despite the differences between the penes of these genera,

there are certain significant features they have in common. The

glans. is divisible into two parts, a swollen proximal portion

carrying the urethral orifice and a more slender terminal portion.

The most primitive type appears to be that of Protoxerus,

where the terminal portion is relatively stout, moderately long,

less sharply distinguished from the swollen ‘proximal portion,

without serrulated crests, and supported by a well-developed

baculum, the proximal thickened end of which is probably lodged

in the thickening of the glans, while the apex extends to the

arrow-headed tip of the thinner distal portion.

In the next stage, exemplified by Wthosciurus, Paraxerus, and

Proc. Zoou. Soc.—1923, No. XVI. 16

230 _ MR. R. I, POCOOK ON THE

Funisciurus, the baculum is reduced to a small rod imbedded

near the apex of the distal portion, which is consequently

flexible. This portion, however, is specialised by the development

of a finely serrulated crest on each side, and is more sharply

marked off from the swollen portion of the glans, upon which it

is capable of being moved up and down.

Finally in Heliosciurus, the highest type, the distal portion is

developed into a flexible filiform process without baculum, the

thickened portion of the glans is longer and simpler, and the

preglandular portion of the penis is greatly lengthened.

Returning to Protoverus, it may be recalled that ‘Thomas drew

attention to “the likeness between its baculum and the baculum

of Ratufa. Unfortunately the penis of Ratufa is unknown;

but a similarity may be traced between the glans of Protoxerus

and of Funambulus if, as I suppose, the orifice opens in

Protoxerus on the swollen part of the glans at the base of the

terminal slender portion. The chief differences in that case

between them will be reduction in the size of the baculum in

Protoxerus and specialisation of the glandular thickening from

which the slender terminal portion arises. In any case the like-

ness between the bacula and penes of Protoxerus and Funambulus

is greater than the likeness between those same organs in

Funambulus and Tamiodes.

4, The African Ground-Squirrels.

In the African Bristly Ground-Squirrels the glans penis is

relatively large, turns downwards at the apex, and has a well-

developed termina] baculum.

In Luxerus erythropus the glans is long, and consists of two

parts, a subeylindrical proximal piece composed of soft, striate or

otherwise sculptured tissue, and a terminal smooth, compressed

piece with an elevated, convex upper edge. which curves down-

wards distally to end in a rounded apex, behind which the

inferior border is concave. The upper edge ends posteriorly in a

twist near the middle of the upper side of the glans. At the

distal end of the spongy portion beneath and a little to the right

of the middle line just behind the base of the compressed portion,

the orifice opens. (Text-fig. 25, E-H.)

The baculum, measuring 8-9 mm., conforms to the shape of

the distal portion of the glans, and consists of a stout, short

cylindrical proximal piece and a compressed blade, sometimes

rounded, sometimes hooked backwards at its inferior apex. The

upper side of the bone is slightly concave; but carries a carti-

laginous crest, sometimes partially ossified, which runs backwards

some distance behind the proximal end of the bone and re pre-

sents the distal dorsal crest of the glans. (Text-fig. 24, A—C.)

The glans of Geosciwrus capensis is tolerably similar, with a

similar dorsal crest ending in a twist proximally, but the distal

portion is not compressed, but gradually narrows to end in a

CLASSIFICATION OF THE SCIURIDA. 231

slightly expanded truncated apex. ‘he orifice is in the median

ventral line, nearer the tip of the glans than in Hwaxerus

erythropus. (Text-fig. 25, A—D.)

The baculum, measuring 8 mm., has a much longer proximal

subcylindical portion than that of MHuwerus, and the upper

Text-figure 24.

A. Baculum of Huaerus erythropus, from the left side.

B. The same from above, without the cartilaginous crest.

©. Baculum of another specimen, ticketed EH. erythropus.

D. Baculum of Geosciurus capensis, from the left side.

i. Apex of the same from above.

¥. Apex of the same from the front.

G. Baculum of Atlantoxerus getulus, from the left side.

H. The same from above.

I. Tip of the same from the front.

K. Baculum of Xeruws rutilus, from the left side.

LL. The same from above.

M. Tip of the same from the front.

surface of the terminal portion is an elongated area rather more

than twice as long as broad, with a low median erest, a rounded

spatulate hollowed apex, and distinctly constricted sides. The

lower side of the terminal portion is developed into a median

vertical crest. (Text-fig. 24, D-F.) 4

232, MR. R. 1. POCOCK ON THE

The baculum in Ailantoxerus getulus, measuring 7 mm., has a

still longer proximal portion, or ‘‘ handle,” than in Geosciurus, and

the blade is simpler. Its dorsal area is apically asymmetrical

and not markedly spatulate, and the sides are scarcely constricted,

Text-figure 25.

A. Glans penis of Geosciurus capensis, from the right side.

3. The same from the left side.

3. The same from below.

D. The same from above.

EK. The same of Huverus erythropus, from the right side.

KK. The same from the left side.

G. The same from below.

H. The same from above.

I. The same of Cynomys ludovicianus, from the right side.

Kk. The same of Marmota marmota.

but the median crest seen in Greosciwrws is present. The inferior.

crest of the blade is also asymmetrical, curving towards the left

inferiorly. (Text-fig. 24, G—I.)

In Xerus rutilus the “ handle ” of the baculum is shorter than

CLASSIFICATION OF THE SCIURIDA. 233

in Geosciurus and no longer than in Huxerus; but the upper sur-

face of the blade is very wide, much wider than the “ handle” and

shaped like a spearhead, with convexly rounded sides and a

sharp point, the width being rather more than half the length ;

its dorsal median crest is much less pronounced than in Greosciurus

and its apex is not spatulate and hollowed. The inferior crest is

similar to that of Geosciwrus. The bone itself measures 6 mm.

(Text-fig. 24, K-M.)

Of the above-described bacula that of Aélantowerus comes

nearest to the baculum of typical Sciuride in the length of the

proximal cylindrical portion or handle; Geosciurus comes next,

and Xerus and Huaerus last with the proximal portion quite

short. In the gradual differentiation or specialisation of the

blade the series runs Huwerus, Atlantoxerus, Geosciurus, Xerus ;

but which of the four is the most primitive type I am quite

unable to say. It is interesting, however, to note that the most

widely separated types are found in Xerus and Huxerus, which

until recently were referred to a single genus, although Geosciwrus

and Atlantoxerus had been severed from YXerus

a. Distal portion of baculum a compressed blade not euugided

above, its upper edge narrow.. ae ... Hucrerus.

. Blade of distal portion of tecnieen laterally Nee aed more or

less above, its upper surface spatulate or subspatulate with a

longer or shorter median crest behind.

6. The upper surface forming an elongated oval plate with uni-

tormly convex sides, pointed at the apex and with a short

erest behind ; proximal Dee of baculum only about half

the length of the distal .. bees . Kerus.

. The upper surface much narrower, coin. sinuous patel an a

longer crest ; proximal portion almost as long as the distal

or longer.

c. Sides of the upper surface strongly constricted, the ter-

minal area apically rounded and hollowed .e..c.csccesceseeeeee Gleosciurus.

. Sides of the upper surface neatly constricted, the tecnriaal

area apically narrow and flat.. ee ee a tee eAELCHICOMETILSs

A point of systematic interest connected with the glans penis

and baculum of the Xerine is the complete absence of resem-

blance between them and the corresponding organs in the African

Squirrels, Protowerus, Punisciurus, and Paraxerus. which were

associated with Xerws by Forsyth Major on the evidence derived

from skulls and teeth.

5. The Soushiks and Marmots.

The glans penis of Citellus I have had no opportunity of

examining, but the bacula of several species are preserved in the

collection of the British Museum. Jn all cases the bone consists

of an elongated shaft, broad at its proximal end and narrowing

distally to the apex, which is upturned and expanded into a wide,

sometimes double dise with denticulated margin.

The simplest type is the baculum of a specimen of C. mongo-

licus from Chifu. The shaft is abruptly expanded at the base,

234 MR. R. I. POCOCK ON THE

nearly straight when viewed from above or below, but with the

upper margin concave, the lower convex. The slightly raised

distal expansion, hollowed above like a shovel, is irregularly semi-

circular in shape, and has a continuous unevenly denticulated

edge, without median notch; and the apex of the shaft of the

baculum projects forwards as a process from its lower surface.

The bone measures about 3 mm., and the width of the distal ex-

pansion is about half the length of the shaft. (Text-fig. 26, A, B.)

The baculum of a specimen of C. leursi from Jalisco, in

Mexico, has the shaft very broad at the base, whence it gradually

narrows to the expansion. Its upper and lower edges are nearly

straight from the lateral view, but from the dorsal view the shaft

is seen to bend distally slightly towards the right. The disc is

relatively and actually much wider than in C. mongolicus, and

its right side is produced more than the left; but its edge is

continuous, without median notch and more regularly denti-

culated than in C. mongolicus. There is, moreover, no process

representing the tip of the shaft on the under side of the disc.

The bone measures 5 mm., and the width of the dise is almost

equal to the length of the shaft. (Text-fig. 26, C, D.)

In two examples of C. mexicanas, one from the City of Mexico,

the shaft differs from that of C. mongolicus and lewrsi in having

its upper edge abruptly inclined downwards and distally towards

the disc; and the dise itself differs in having a deep median

notch in its margin and in being much more strongly bent

upwards, so that its hollow is much deeper and more triangular

in shape. As in C. mongolicus, the apex of the shaft is con-

tinued as a short process projecting from the lower surface of

the disc in front a little beneath the notch. The bone measures

about 5°5 mm. and the greatest width of the dise across its

posterior angles is less than half the length of the shaft.

(Text-fig. 26, E, F.)

In a specimen of C. 13-lineatus from Minnesota the shaft is

intermediate in shape between that of C. mexicanus and of

C. leursi; but the dise differs from that of the other species in

having its edge broad and furnished with two rows of teeth. It

is, moreover, very definitely divided into a right and left lamina

by a wide and moderately deep median notch in front, and it is

somewhat abruptly upturned. The tip of the shaft shows as a

low, indistinctly defined rounded boss on the under side of the

dise just below the notch. The bone measures 5 mm. and the

width of the dise is about half the length of the shaft. (Text-

fig. 26, G—I.)

a. Edge of the apically widely notched dise broad and armed with two

rows of teeth <0... 0... 05. viterescseese Ld-lineatus.

a’, Edge of the dise narrow and armed witha single row of teeth.

b. Dise nearly vertically upturned, with a narrow median notch in

front: upper surface of shaft distally geniculate..................... MeLICANUS.

b’ Disc a little upturned, without median notch; shaft not distally

geniculate above.

CLASSIFICATION OF THE SCIURIDA,

_ Text-figure 26.

A. Baculum of Citellus mongolicus, from below.

B. The same from the left side.

C. Baculum of C. leursi, from below.

J). Apex of the same from the left side.

E. Baculum of C, mexicanus, from above.

F. The same from the left side.

G. Baculum of C. 13-lineatus, from below.

H. The same from the left side.

I. Apex of the same from above.

K. Baculum of Cynomys ludovicianus, from the left side.

L. The same from above.

M. The same of another specimen, from below.

N. Baculum of Warmota marmota, from below.

O. The same of another specimen.

236 _ MR. R. I. POCOCK ON THE

e. Disc comparatively small, its width about half the length of

the shaft, the tip of the shaft projecting as a process be-

MOA GAY. M208. hsggce Noses celseececesu enn ceeea eames eameoncae nee . mongolicus.

ce’. Dise very wide, its width almost equal to the length of the

shaft, the apex of which forms no projection beneath it in .

PRONE a hiveetic vbviecsienanctunamoasis diame beeen alaeemuetean eae eae ctr eset: MRL CUT SOS

In Cynomys ludovicianus the glans penis is short, its proximal

two-thirds being stout and subcylindrical and its distal third

forming a quite narrow terminal process sharply defined from

the thicker portion. The orifice lies on the right side close to

the base of the narrowed end. The baculum, measuring 4 mm.

in length, lies in the narrowest portion and reaches its apex.

It is broader at the base and tip than in the middle and is

slightly asymmetrical. Its upper edge is very slightly concave,

its lower edge mesially geniculate, the apex being irregularly

denticulated. (Text-figs. 25,1; 26, K-M.)

In Marmota marmota the glans penis is shaped very much

like that of Cynomys, but is relatively a little shorter. The

orifice opens similarly on the right side at the base of the

narrowed terminal process and the baculum, measuring only

7 mm., and lying in the narrowed piece is a slightly asym-

metrical bone with expanded base and expanded irregularly

denticulate apex. It is not quite alike in two specimens

examined, and I am unable to affirm any character apart from

size by which it may be distinguished from the baculum of

Cynomys. (Text-figs. 25, K ; 26, N, O.)

It may be recalled that Tullberg many years ago described

the glans penis of Marmota [| Arctomys] as very small and asym-

metrical, with the opening on the right side behind the tip, which

carries a small baculum. The male genitalia of Cynomys he

described as like those of Marmota; but, what is more interesting

still, he declared the male genitalia of Zamias to be also very

similar to those of M/armota.

6. The Subfamvilies of Sciuride.

For the following grouping of the genera of non-volant

Squirrels, constituting the Sciuride, I take the glans penis and

the baculum as supplying characters of primary value. It may

be objected that these characters apply only to one sex; but I

do not think it can be doubted that corresponding differences

occur in the females and merely await discovery. The baculum

has been examined either by Mr. Thomas or myself in all the

principal genera, and the penis by Tullberg and myself in a

considerable number of them. As subsidiary characters I have

used the ears and feet, leaving alone cranial and dental

characters, the study of which by Thomas, Forsyth Major, and

many American authors has led to no very satisfactory result

apart from the steady but unavoidable and convenient multi-

plication of genera and subgenera.

CLASSIFICATION OF THE SCIURIDE. 237

Subfamily Scrurinz,

Glans penis apically expanded, compressed and upturned,

enveloped behind the upturned tip in soft swollen tissue.

Baculum large, with compressed terminal blade hollowed on the

right side, which carries inferiorly a short downwardly directed

process.

Ears relatively long, more tubular at the base, with large

angular antitagral flap.

Feet with fourth digit the longest; the plantar, carpal, and

metatarsal pads normal and relatively simple.

Genera: Sciwrus, with subgenus Yenes* for persicus; all the

American subgenera or genera, Weosciwrus, Parasciurus,

Echinosciurus, etc., except TZamiasciurus; % Rheithro-

SCtUrUs.

Distribution. Holarctic and Neotropical Regions and ? Borneo

(Rhetthrosciurus),

I feel compelled to include Rhetthrosciurus provisionally in this

group, pending confirmation or refutation of Thomas’s statement

that the baculum is like that of Sciwrws. In other external

characters supplied by the ears and feet it is certainly unlike

other Oriental species and comes nearer to Sciwrus. On the

evidence at present available I should judge it to be a highly

specialised member of the Sciurinz as above defined.

Subfamily TAMIASCIURINA,

Distinguishable from the Sciurine, which it resembles in

cranial and other external characters, by the structure of the

penis, which is relatively long and slender and tolerably evenly

attenuated from hase to apex, and flexible throughout owing to

the suppression of the baculum.

Genus Vamiasciurus.

Distribution. Nearctic Subregion.

The general resemblance between Zamiasciurus and the typical

Sciurine suggests that it is a specialised offshoot of that group.

It is a very remarkable fact that the extreme differentiation of

the penis does not appear to be correlated with other variations

from the Sciurine type.

In the structure of the penis Z'amiasciurus shows to a certain

extent a convergent® resemblance to the African genus Helvo-

sciurus (see below, p. 238).

Subfamily FuNAMBULINA.

A highly diversified group of genera, with the glans penis

exceedingly variable in size and structure and the baculum either

* Thomas, Ann. & Mag. N. Hist. (8) iii. p. 468 (1909).

238 MR. R. I. POCOCK ON THE

relatively very large (Funambulus, Tamiodes), relatively small

(Ratujfa, Protoxerus), minute (Punisciurus, Paraxerus, ditho-

sciurus), or absent (Heliosciurus). It is, however, when present,

always a simple bone without the spatulate expansion at the

apex seen in the Sciurine, without the accessory blade of the

Tomeutine, and without the compressed downturned lamina

of the Xerine. The ears are always relatively shorter and wider

and less tubular at the base than in the Sciurine, but are without

the peculiarities of the ears of the Xerine and altogether better

developed.

Genera: Vunambulus, Tamiodes, Ratufa, Protoxerus, Htho-

seiurus, Funisciurus, Paraxerus, Heliosciurus, and

probably the other African genera admitted by Thomas,

including possibly even Wyosciurus.

Distribution. Orientai and Ethiopian Regions.

When the genera are better known, this group will perhaps

be split up into several subfamilies. So far as my observations

go, the genera fall into the following sections by the characters

established in this and my previous paper :—

a. Tamiodes, with a thick glans penis with terminal labiate

orifice and a large baculum with its distal extremity bent

upwards at a right angle to the rest of the shaft.

B. Funambulus, with the glans terminally narrowed, the simple

orifice on its right side, some distance behind the tip of

the glans, and, the baculum large and gently curved upwards

distally.

y: Ratufa, with the baculum relatively quite small, curved as in

Funambulus, but stouter and bevelled below distally ; (glans

unknown); the feet in their pad development quite different

from those of the other genera.

6. Protoxerus, with the glans possessing a median swollen area

carrying the orifice, and an elongated narrower distal portion

supported by a relatively small baculum not differing greatly

from that of Ratufa, but with a button-like apex and a

scalloped proximal end.

e. Lunisciurus, Paraxerus, Hthosciurus. Glans with a marked

median grooved or laminate swelling and a slender distal

portion as in Protoxerus; but the distal portion flexible,

laterally serrulate, and carrying a minute undifferentiated

baculum close to the apex.

€. Heliosciurus. Penis and glans exceedingly long, the glans

with narrower, longer swelling, and exceedingly thin, long

filiform distal portion without trace of baculum.

Of other genera that have been established, Zamiscus, recently

dismembered from Paraverus, probably falls into section e. But

whether Hpiverws comes into 6 or e I am unable to surmise.

Probably an examination of the glans penis and baculum will

CLASSIFICATION OF THE SCIURIDA. 239

settle the doubtful question as to the affinities of the genus with

Protoxerus or Funisciurus. Of the genitalia of Myosciwrus, 1

know nothing beyond Thomas’s statement that the baculum is

simple as in other African genera. The position of Myrsilus is

also doubtful.

Subfamily CaLLosciuRIn&.

Glans penis variable in shape and length according to the

baculum ; the orifice close to the tip on the right side. Baculum

long and slender or short and stout, slightly or strongly upturned

distally, and always provided with an accessory blade, movably

jointed to the concavity of the upper surface.

Ears asin Funambuline.

Feet, where known, intermediate in structure between those

of the more primitive types of the Funambulinz (/unambulus)

and the specialised genus Ratufa.

Genera: Callosciurus, Menetes, Tomeutes, Rhinosciurus, Laris-

cus, Dremomys, Tamiops, Nannosciurus, and probably

others.

Distribution. Oriental Region from India’ to China and

Borneo.

This group may perhaps be regarded as a specialised offshoot

from the Funambuline. The simplest type of baculum, ¢. 9.

that of Cailosciurus, may be derived from that of Funambulus by

the addition of the blade. But the subterminal position of the

orifice is as in the Sciurine.

Subfamily XeRinz.

Glans penis, where known, with a strongly convex crested

upper distal edge and a downwardly directed, rounded or truncated

apex. Baculum with a stout, longer or shorter subcylindrical

proximal portion, and a compressed downturned terminal blade

with a median crest above and frequently a laterally expanded

upper surface.

Ears more or less reduced, but with large fleshy antitragal

thickening.

Feet with strong fossorial claws and the third digit the longest.

Fur scanty and harsh. An additional tuft of superciliary

vibrisse over the posterior corner of the eye.

Genera: Atlantowerus, Xerus, Humerus, Geosciuwrus.

Distribution. Africa from Morocco to Cape Colony.

The structure of the penis and baculum alone serves to

negative the idea that this group of African Ground-Squirrels is

closely akin to certain African arboreal Squirrels, an idea which:

prompted the proposal by Major of such names as Protoverus and

Paraxurus for two genera of the latter category. The Xerine,

indeed, are in addition distinguishable from the rest of the

240 MR. R. I. POCOCK ON THE

African genera of the family by their ears, feet, fur, and the

additional tuft of superciliary vibrisse, and by the cranial

characters pointed out by Thomas in his paper on African

Squirrels.

Subfamily Marmorin a».

Glans penis, at least in Marmota and Cynomys, very small, a

diminutive copy indeed of that of Punambulus but with the point

not upturned by the curvature of the baculum; also somewhat

resembling that of Yuniscturus and Paraxerus, except that the

proximal thickening and the terminal narrowed piece are

structurally simple and less sharply differentiated. Baculum

also relatively very small, highly specialised with a symmetrical

terminal, denticulated lamina in Citellus, but ending simply with

a few irregularly placed denticles in Marmota and Cynomys.

Hars simplified, without tragal or antitragal thickenings, but

with a hairy expansion of the anterior rim extending into the

cavity beneath the supratragus.

Feet with fossorial claws and the third digit not shorter than

the fourth.

Genera: Marmota, Marmotops, Cynomys, Citellus with many

subgenera, ? Zamias, and Hutamias.

Mstribution. Holarctic Region.

Although admitted as a subfamily in many text-books, this

group was not granted that rank by some competent judges,

e.g. Forsyth Major and Miller, on account of the closeness of

its kinship with the true Squirrels. It appears, however, to be

sufficiently well characterised to be admitted.

I have provisionally included Zamias and Hutamias mainly on

the strength of Tullberg’s statement that the genitalia are similar

to those of Marmota, but also because Tamias and Citellus are

alike in the presence of large cheek-pouches, a character in

which they differ from all the typical Squirrels.

Nevertheless the feet of the only example of this group I

have seen in a fresh state, namely a female of Hutamias quadri-

vittatus, are not like those of C%éellus, and hardly differ from

those of the Squirrels. The ears, too, do not show the special

features characteristic of those of Citellus, Warmota, and Cynomys,

but are more like those of Sciwrws. More extended observations

on the two genera are required to determine their exact

systematic status. Possibly they will prove to be worth ranking

as a special subfamily—Tamiine.

7. The Flying Squirrels.

The Petauristidee were classified by Thomas in 1908*. Setting

aside Petwurista, he took the genus Sciwropterus, already shown

by Heude and Forsyth Major to be composed of heterogeneous

* Ann. Mag. Nat. Hist. (8) i. pp. 1-6.

CLASSIFICATION OF THE SCIURID. 24)

elements, and pointed out that it is divisible by the structure of

the teeth into six genera :—Sciwropterus, type rwssicus, for the

Palearctic species; Trogopterus for wanthipes from China;

Belomys, type pearsoni, Pteromyscus, type pulverulentus,

Petaurillus, type hosei, and Jomys, type horsfieldi, from various

parts of the Oriental Region, each containing one or more

species besides the type. Furthermore, he divided Scvwropterus

into four subgenera: Sciuropterus itself; Glaucomys for the

North American volans and the Himalayan jimbriatus* ; Zylopetes,

containing many species ranging from Nepal to the Malay

Islands, with everettii as the type; and Petinomys, ranging

from Ceylon into South India and the Malay Islands, with lugens

as type.

‘Owing to lack of material, I can contribute very little to what

has been already established as to the affinities of the genera of

this faraily. The outstanding result of my examination of the

bacula is the complete severance of Hoglaucomys from Glaucomys.

This suggests that further surprises may be in store for him who

has the time and opportunity to carry on the work.

The penis and baculum are as variable in the Petauristide as

in the Sciuride. The penis itself is only known to me in two

genera, Hoglawcomys and Hylopetes, of which one specimen of

each, preserved in alcohol, has been available.

In Hoglaucomys fimbriatus the glans is stout and tolerably

long, a little stouter just beyond the middle than at the base,

and slightly upeurled but not appreciably narrowed at the end.

On the left side above, just before the tip, there are three

processes in a line, the first and third short and semiconical,

the second or middle one much higher and roughly three-sided.

Beyond these the apex of the glans forms a, lamina hollowed on

the left side, and beneath this there is a rounded area with a

fleshy flap defined by a groove which passes backwards on to the

right side, where the area in question is narrowed and runs

backwards, being defined above and below by a groove. ‘The

orifice appears to be situated towards the posterior end of the

upper groove, and from the orifice two more grooves run back-

wards tc the proximal end of the glans. (Text-tig. 27, A-C.)

The baculum, measuring 7°5 mm., is exceedingly short and

occupies only the distal half or less of the glans. It is furnished

with several processes, and agrees tolerably closely in shape to

the shape of the end of the glans, Itis indeed the bony processes

of the baculum, enveloped in soft tissue, which give shape to

the end of. the glans; but the rounded area of the glans below

the distal hollow is wholly composed of soft tissue, no portion of

the baculum entering it. The proximal end of the baculum

forms a short, stout rod, shorter than the terminal toothed por-

tion and emarginate below. Considering the former association,

* This species was subsequently separated from Glaweomys as the type of the

subgenus Hoglaucomys.

242 MR. R. I. POCOCK ON THE

suggested by their names, of Hoglawcomys with the North

American Glaucomys, the differences between their bacula are

very striking and suggest that the resemblances between the

genera, which induced Thomas in his revision of the Petauristide

to refer fimbriatus to Glaucomys, are purely convergent. The

differences between their bacula are apparently indeed greater

than the differences between the bacula of any other two

species. (Text-fig. 29, H-K.)

In Hylopetes alboniger the glans penis is very different from

that of Hoglawcomys. The distal half of the glans is narrowed

and curved considerably towards the left and upwards at an

Text-figure 27.

A, B. Glans penis of Hoglaucomys fimbriatus, from right and left sides.

C. Apex of the same from the front.

D, E, F. Glans penis of Hylopetes alboniger trom right side, left side, and

above.

angle of about 135°. On the left side of the upturned portion

there is a crest running downwards from the apex and ending

near the middle of the lower convex surface in a well-marked

compressed point. Near the base of the glans on the upper

surface there are two crests, one in front of the other and

defining a kind of saddle-shaped area; these crests pass down-

wards on to the right side of the glans. J could not find the

orifice of the organ of the specimen examined. (Text-fig. 27,

D-F.)

The baculum, measuring 13mm., is shaped distally like the distal

portion of the glans, having the same upward and sinistral curva-

CLASSIFICATION OF THE SCIURID. 243

ture. There is a well-developed crest on its left side, passing

from the apex down to the inferior tooth which is the termination

of the crest. But although the baculum is long and extends

throughout the length of the glans, it shows no trace of crests or

processes behind the termination of the crest inferiorly. This

baculum differs from that of Hoglawcomys in its length, its simple

apex, and the lateral inferior crest. (Text-fig. 28, C, D.)

Of this baculum I have seen two specimens, one taken from

the spirit-preserved example from the Himalayas in the Zoological

Society’s collection, the other from an example from Mishmi in

the British Museum.

The baculum of a specimen of Hylopetes phayret from Burma,

in the British Museum, measures 11 mm., and agrees very closely

with that of H. alboniger, but is less markedly upeurled distally

and is provided with a larger crest. (Text-fig. 28, E.)

Hylopetes, it may be recalled, was regarded by Thomas as a

subgenus of Sciuropterus. Since the baculum of Se. russicus does

not appear to have been described, it is impossible to give an

opinion as to the precise status of Hylopetes. But if, as appears

to me probable, the baculum of Sciwropterus russicus resembles

that of Glaucomys volans, full generic value should, I think, be

given to Hylopetes.

The baculum of an example of Petinomys fuscocapillus from

Cevlon, in the British Museum, is a tolerably stout bone with a

sinistral curvature like that of Hylopetes, the right side being

convex, the left side concave, but 1t has no very marked upward

curvature, the upper edge being slightly sinuous and the apex

a little raised, almost as in Hylopetes phayreit. The apex, how-

ever, is broader than in Hylopetes and somewhat spatulate, the

left margin of the shallow apical hollow being raised into an

angular tooth behind. On the lower side of the bone there is a

strong crest divided by a notch, about in the middle of the bone,

into a short proximal and a long distal portion. The latter,

which is angled behind, forms a sinuous curve towards the left,

but does not ascend on to the left-hand side of the bone distally,

as in Hylopetes, but terminates on the lower side of the expanded

spatulate apex. The baculum measures 11 mm. (Text-fig. 28,

F-H.)

In view of the affiliation by Thomas of Petinomys with Hylo-

petes as subgenera of Sciwropterus, it is interesting to note the

broad general resemblance, associated with marked differences,

between their bacula.

The baculum of Glaucomys volans, judging from two specimens

in the British Museum, one of which is ticketed Virginia, is

exceedingly long and slender, slightly sinuous in its proximal

third, and inclined slightly upwards distally. The extreme apex

is bifid, the lower process being rounded, the upper more pointed.

On the left side there is a long crest running from the summit of

the upper terminal process and ending abruptly behind the left

side about one-third of the distance from the proximal end of

244. MR. R. I. POCOCK ON THE

the bone. It lies over a well-marked groove, and there is a

second shallower groove on the right side of the bone. The bone

measures 12°5 mm. (Text-fig. 28, A, B.)

Despite its straightness and apical notch, this baculum is

similar in type to that of Hylopetes, the erests on the left-hand

side of the two corresponding closely. The crest in Glaucomys

terminates on the left side of the bone instead of passing on to

its lower surface as in Hylopetes. The surprising differences

Text-figure 28.

ri G

. Baculum of Glaucomys volans, from the left side. ~

. The same from below.

. Baculum of Hylopetes alboniger, from. the left side.

. The same from below.

. Baculum of A. phayret, from the left side.

. Baculum of Petinomys fuscocapillus, from the left side.

. The same from below.

. The same from above.

Heats oewP>

between the bacula of Glawcomys and Hoglaucomys have already

been mentioned under the description of the latter genus.

The baculum of a specimen of Belomys (2) trichotis from Yin,

Chindwin, in the British Museum, is hardly longer than that of

Foglaucomys, but is otherwise very different from it. The

proximal portion consists of a stout subeylindrical “handle,”

sharply geniculated where it passes into the abruptly upturned

distal portion, which ends in a wide lamina, shaped like a

CLASSIFICATION OF THE SCIURIDA. 245

hammer-head and lying obliquely and transversely, the shorter

right-hand branch projecting farther forwards than the longer

left-hand branch, This baculum is quite unlike that of any

Sie species of the Petauristide examined. (Text-fig. 29,

, G.)

The baculum of an example of Petawrista philippensis from

Kanara, in the British Museum, is a long, stout bone, gradually

Text-figure 29.

. Baculum of Petaurista philippensis, from above.

The same from the left side.

. Apex of the same from the front.

. Baculum of Petaurista sp.? from above.

. The same from the left side.

Baculum of Belomys trichotis, obliquely from behind.

. The same from above.

. Baculum of Hoglaucomys fimbriatus, from the left side.

. The same from the right side.

. The same from above.

AHH Qe et boWwPe

narrowing from the base to the slightly expanded and upturned

distal end, which, when viewed from the front, is seen to be

shaped rather like the widened spout of a jug, the lower rim of

the spout being evenly rounded. From the side this terminal

lamina has a rounded upper border, a rather deeply emarginate

and thickened distal border, and a nearly straight, obliquely

ascending, thickened posterior border ending inferiorly in a small

Proc. Zoou. Soc.—1923, No. XVII. 17

246 ON THE CLASSIFICATION OF THE SCIURIDA.

tooth. Just below the hollow on the left side there is a small

crest ending above in another small tooth. The bone measures

20 mm. in length. (Text-fig. 29, A-C.)

The baculum of another but unidentified and unlocalised species

of Petaurista in the British Museum differs in some well-marked

characters from that of P. philippensis. It measures only

8 mm. owing to the shortness of the proximal portion, which

does not exceed the distal portion in length. The latter has its

hollowed surface looking more to the left, the upper or right-

hand rim being more, and the lower or left-hand rim less

elevated than in P, philippensis, and its distal rim is not

curved over and spout-like. If the ossification of the baculum

proceeds backwards from the distal to the proximal end, it is

possible that the difference in the length of this bone in

P. philippensis and the unidentified species is a matter of age.

(Text-fig. 29, D, H).

The following table shows how the genera examined may be

distinguished by their bacula :—

a. Baculum with crest running from the comparatively small and

simple apex backwards along the left or under side of the

bone.

6, Baculum nearly straight, puna aoe long and slender;

apex bilobed, crest confined to left side.. Fee Glaucomys.

6’, Baculum shorter, stouter, more curved ; apex Pade ‘Giesi

wholly or partially ventral.

e. A single crest curving distally up on to the left side of the

bone to the apex, which is simple, not spatulate.............. Zylopetes.

c', A supplementary crest behind the main crest, which is en-

tirely ventral; apex spatulate ............ . Petinomys.

a’. Baculum without lateral or ventral crest, eh eapandleane com-

plex apex.

d. Distal end of baculum abruptly upturned and ending in a

two-headed or hammer-shaped lamina . be Belomys.

d', Distal end of baculum not abruptly ceenneas My apex ex-

panded, more or less hollowed, and complicated with accessory

processes.

e. Apex of baculum hollowed, any, sea with small

accessory processes ......... Govan hadouaAuacaandee | eS AROOUTAN TE

e', Apex of baculum armed aah uprising peor processes

onthe Wlentisvde Eig sntascataecscecce ce eee eace ere eee LOO LGCCOMIy Ss

ON MAMMALS OBTAINED IN DARFUR. 247

18. On the Mammals obtained in Darfur by the Lynes-

Lowe Expedition. By Oupriznp THomas, F.R.S.,

F.Z.8., and Martin A. C. Hiyron, F.Z.8.

[Received January 23, 1923 : Read March 20, 1923.]

Thanks to the generosity and public spirit of the two explorers,

the British Museum has received as a donation the whole of the

fine collection of Mammals made by Rear-Admiral Hubert Lynes

and Mr. Willoughby P. Lowe during their recent expedition to

Darfur.

The pedis took place during the whole of 1921 and the

early part of last year, and a complete survey of the country

was made, equally of the comparatively flat desert region round

El Fasher, the capital of Darfur, the still more desert area north-

ward to the bare and unpr oductive Jebel Maidob, the zoologically

unknown dominating mountain Jebel Marra, running up to a

height of 10,000’, and, finally, of the lower region of Wadi Aribo,

in the south-western part of Darfur, where the drainage is towards

Lake Chad.

No mammal collection had ever been made in this area, so

that the present fine series (which numbers upwards of 800

specimens) adds very greatly to the material available for the

study of African Mammalia, and we have reason to be most

evateful to the donors for the geuerosity and patriotism which

have resulted in this notable accession to the Museum—the

largest single collection that the latter has ever received.

On the whole, the species contained in the collection are most

nearly related, as is natural, to those of Kordofan and other

parts of the Egyptian Sudan, and are generally different from

those of the more humid Bahr-el-Ghazal.

So far as the mammals are concerned, Darfur would seem to

be just on the southern boundary of the northern desert fauna,

the collection containing quite a number of forms which are

either the most southern records of northern species (Jaculus

jaculus, Dipodillus campestris group, etc.) or the most northern

records of southern ones (Steatomys, &c.).

The great mountain Jebel Marra, isolated as it is from other

high ground, has naturally a number of interesting forms peculiar

to it and different from those of the plains. Thus there is a

mountain species of Striped Mouse (Lemniscomys) found on it,

which we have named in honour of Admiral Lynes, and a

Gerbil (Dipodillus lowei), whose nearest ally is found in Algiers.

In all, the collection proves to consist of 62 species, of which

we have had occasion to describe 19 as new, either as species or

subspecies.

Aliya

248 MESSRS. OLDFIELD THOMAS AND M. A. C. HINTON ON

A few mammals were obtained by Admiral Lynes on a previous

visit to the country, and these have been here incorporated.

Their numbersrun from 1 to 24, while those of the main collection

start at 400.

1. Eryrurocenus pyrReonotus Hempr. & hr.

3. 648. 9. 672. Foot-hills, 8. Jebel Marra. 4000’.

Practically the first satisfactory examples of this species to

reach the Museum.

2. CERCOPITHECUS TANTALUS MARRENSIS, subsp. n.

¢. 631. Foot-hills, 8. Jebel Marra. 4000’.

@. 626, juv. S.W. Jebel Marra. 5000.

A very brightly coloured subspecies.

Face and chin black. White brow-line well defined, with an

anterior edging of black hairs. Whiskers yellowish white, directed

backwards and upwards, completely concealing ears. A sharply

defined black streak from outer canthus of eye to neighbourhood

of ear, separated from the crown-patch by a narrow tract of white

whisker-hairs. Predominant hue of upper parts (crown of head,

back to rump, and flanks) bright buff or golden, darkened on

crown of head and rump by black bair-tips and to some extent

by the hair-bases of slate-grey, which are darkest on the crown

of the head. The golden tint is especially bright and clear upon

the withers and flanks. Under surface of body, with inner

surfaces of limbs, well-haired, pure white ana rather sharply

contrasted with flanks. Outer surfaces of arms and hands

from shoulders, and of the legs and feet, from the thighs, of a

general light grey colour of’ cold tone, produced by a mixture

of pale slate-grey and dirty white, without any trace of buff.

A tuft of white hair on each side of the root of the tail above.

Upper surface of tail, in its proximal two-thirds, like the rump ;

its lower surface in the same region white, becoming yellower

distally. Distal third of tail, above and below, dull yellow.

Dimensions of the type (measured in the flesh) :—

Head and body 830 mm.; tail 1140; hind foot 145; ear 32.

Skull: extreme length 118°6; condylo-basal length 88; zygo-

matic breadth 68:2; external orbital width 62:6; postorbital

constriction 44; width of brain-case 56°5; canine to m° 34:2;

pom 2d°4,

Hab. Foot-hills of Jebel Marra.

Type. Adult male. B.M. No, 23.1.1.1. Original No. 631.

Collected April 1, 1921.

This isa very well-marked subspecies of C. tantalus, a species

known to range from Nigeria and the West Coast eastwards to

the shores of Lake Albert. (. toldti Wettstein, described from

Kadugli, 8S. Kordofan, on the basis of rather unsatisfactory

material, appears to be a member of the “ callitrichus” or

sabeus group.

MAMMALS OBTAINED IN DARFUR. DAY

3. Papio anusis F. Cuv.

3.8. Jebel Marra. 7000’.

Shot and presented to Admiral Lynes by Mr. Cecil McConnel.

4, GALAGO SENNAARIENS?S Less.

6. 781, 804. 2, 783. Kulme, Wadi Aribo, 3300’.

5. EipOLoNn HELVUM Kerr.

No specimens of this bat were obtained. But a stem of a

small tree, which Mr. A. 8. Brown has identified as-being either

an Odina ova Sclerocarya, completely gnawed through, was found

on Jebel Marra and brought home by the collectors as evidence

of the existence of some large rodent upon the mountain. The

tooth-marks, however, are very different from those of any

rodent.

In 1920 Mr. R. H. Bunting, of the Agricultural Department

of the Gold Coast, sent us some specimens of Arawcaria, from a

grove at Aburi, which had been seriously damaged by large bats

in a time of drought. With the timber came some of the bats,

caught in the act, and they proved to be Hidolon heluwm. The

bitten wood from Jebel Marra is exactly like that from Aburi,

and may be regarded, perhaps, as good evidence of the presence

of this bat in certain seasons upon the mountain.

6. HrprosrDEROS CAFFER Sund.

3. 887. 2. 885, 886. Kulme, Wadi Aribo, 3300’.

6. 939. 2. 954." Zalinger. 33007

1174. 170 miles HE. of Hl Fasher.

Forearm 48—49 mm.

(, ASELLIA TRIDENS Geoff,

$. 6, 17, 18. 2. 5, 19, 20, 24. Um Esheishat Well, 104

miles H. of El Fasher. 2200’.

These specimens were collected by Admiral Lynes during his

preliminary visit to Darfur. Two of them were obtained on

February 2, 1920, the others on May 13 following; all are in

the brillant red phase.

8. PIPISTRELLUS MARRENSIS, Sp. 1.

3. 633, 653, 656. Foot-hills of S. Jebel Marra. 4000’.

Kssentially as in P. deserti Thos., but of smaller size and darker

colour.

Colour comparatively dark and rich, much like tnat of Heyptian

specimens of P. kuAli, the general hue of the upper parts being

near “ Dresden brown” of Ridgway. Hars noticeably darker than

back. Wings dark brown, with usual whitish edgings ; .inter-

femoral paler. ‘Tragus shorter and broader than in desertz, with

250 MESSRS. OLDFIELD THOMAS AND M, A. C. HINTON ON

broadly rounded tip and parallel borders; the inner border the

longer. Forearm not exceeding 28 mm. (29°8 in desert).

Apart from its slightly smaller size, the skull agrees with that

of P. deserti. ‘The outer upper incisor and the small upper

premolar p* appear to be a little more reduced than in deseriz,

but the available material shows a rather wide range of variation

in these respects. In the type-skull 2° is both absolutely and

relatively much smaller than in the only known skull of deserti ;

and the point of the tooth does not rise above the cingulum of 7.

The small premolar is also greatly reduced, and so crowded

between the canine and p* that it is not easy to detect. But in

the fragments, all that is left of the two other skulls from Jebel

Marra, the teeth in question, though rather smaller than in

deserti, are considerably larger than in the type.

Dimensions of the type :—

Head and body 37mm.; tail 21; hind foot 5:5; ear 12.

Forearm 26:5; third finger 46 (m.c. 245; phalanges 8°7—

7 5—5: 5); lower leg and hind foot (¢.u.) 14°5.

Skull: greatest length 11-2; median length above 9°6; median

length below 8:1; interorbital breadth 4:3; intertemporal

breadth 3:4; breadth of brain-case 6:7; canine to m? 3°8.

Hab. Foot-hills, 8. Jebel Marra; altitude 4000’.

Type. Adult male. B.M. No. 23.1.1.15. Original No. 633.

Collected April 3,192]. ~~

This interesting little bat is, no doubt, closely related to

P. deserti, described from Tripoli. Its smaller size, darker colour,

shorter tragus, and possibly more reduced dentition seem to

warrant its receiving distinct specific rather than subspecific

rank. The individual variation noticed in the dentition is of

some interest in a group in which, normally, even minute dental

characters are surprisingly constant.

Both marrensis and deserti are apparently closely allied to

P. kuhli; and we can see no good reason for placing them in the

genus Scotozous (cf. Miller, Fam. & Gen. Bats, p. 206, 1907).

9, ScoroPHiLUS NI@RITA Schreb.

6. 401. 60 miles W. of El Obeid.

3d. 639. 9. 660,662. Foot-hills, 8. Jebel Marra. 4000’.

Q. 734, 735, 738, 743, 744, 745, 746, 748, 749. Zalingei.

2800’.

At Zalingei: “ Very common around swampy ground” in

May 1921. In a note dated Oct. 27, Mr. Lowe says “I believe

all these bats are migratory, as I have not seen any since the last

obtained ” (Zalingei, May 31).

10. ScoreInus sCHLIEFFENI Pet.

dS. 411. 70 miles W. of Nahud, Kordofan.

3. 1187,1188. 9.1186. 35 miles E. of Nahud.

&. 1197. 50 miles W. of El Obeid.

MAMMALS OBTAINED IN DARFUR. 2

11. TarHozous MAURITIANUS Geoff.

3d. 794. Kulme, Wadi Aribo. 3300’.

12. NycTINOMUS TONGAENSIS Wettst.

@. 1171. 125 miles K. of Hl Fasher.

Forearm 49 mm.

We are agreed that, on the whole, it would be advisable to

separate generically the species of this group with 4 lower in-

cisors— Vyctinomus (genotype, egyptiacus)—from those with 6—

Tadarida (type, teniotis). No species seem doubtful as to their

allocation in one or other of the genera, and the separation will

clearly be a convenience. And at the same time it will elude

the disputed ‘question as to whether Vadarida or Nyctinomus is

the earlier name for the combined genus.

13. ATELERIX ALBIVENTRIS Wagn.

Gin loos da Gol Um Kedadar

Skull only: 1153. 32 miles HK. of Hl Fasher.

14. CRocIDURA DARFUREA, sp. n.

GAO (30s (OliO4. OD, Gil iG 608, 9092929" Fal, Iai Oae.

945, 946, 950, 955. 2. 733, 739, 758, 876, 893, 934, 941, 944.

Zalingei, Darfur. 2800-3300’.

3. 827, juv. 2. 833, 894, 919, Unsexed. 892, 907, 926.

Kulme, Wadi Aribo. 3300’.

A large species resembling C. surwre in colour, but with

much shorter tail.

Size large (hind foot about 18mm. ; condylo-incisive length of

skull in-adult male about 29). Tail short, about half the length

of the head and body. General colour of dorsal surface pale, in

lighter specimens near ‘“ citrine-drab” of Ridgway, in darker

ones approaching ‘ hair-brown.” Underparts lighter and

greyer, with a decided wash of silverin fresh full fur. No sharp

fHank-line. Flank-gland on each side marked by a small patch of

white hairs. Upper surfaces of hands, feet, and tail essentially

concolorous with back ; but tail often a little darker and some-

times dusky, its lower and upper surfaces similar.

Skull essentially as in xyanse and surure.

Dimensions of the type (measured in the flesh) :—

Head and body 122 mm. ; tail 65; hind foot 18; ear 13.

Skull-measurements of type (with those of an adult female in

parentheses): condylo-incisive length 29-4 (27:3); greatest

breadth 12:2 (11-4); least interorbital breadth 4:9 (5); upper

tooth-row 13 (12°4).

Hab. Darfur.

Type. Adult male. B.M. No. 23.1.1.40. Original No.

755. Collected June 4, 1921, at Zalingei, Darfur. Altitude

2800’.

252 MESSRS. OLDFIELD THOMAS AND M, A. C. HINTON ON

This Shrew is the local representative of O. nyanse and

C. surure. Resembling the latter in colour, it is distinguished

from both by its relatively shorter tail. In the type of surwre,

the tail measures only 64mm. with a head and body measurement

of 111; but the material in the Museum indicates that swrure

normally has a tail of 75 or 80 mm., which is therefore consider-

ably more than half the length of the head and body.

In this species there seems to be a well-marked sexual differ-

ence of size—female skulls being usually less, and males more

than 28 mm. in condylo-incisive length,

15, CRocIDURA HINDEI MARRENSIS, subsp. n.

3. 045. Wadi Kongei, Hast Central Jebel Marra. 6200’.

@. 544. Wadi Barei, N.W. Jebel Marra. 6000".

Closely resembling C. hk. diana Dollman, but darker in colour

and with a longer tail.

Size slightly smaller than in diana, but tail considerably longer

(58 mm. instead of 48). Colour of upper parts darker, near the

‘“ snuff-brown ” of Ridgway. Under parts greyish white. Flank-

lines of demarcation reg rular, though not sharply defined. Hands

and feet yellowish white above. “Tail dark brown above, paler

below.

Skull and teeth nearly as in diana; tooth-row a little shorter.

The unicuspid teeth above without the peculiar narrowing and

elongation of their crowns seen in diana.

Dimensions of the type (measured in the flesh) :—

Head and body 74mm. ; tail 57; hind foot 12:5; ear 13.

Skull (posterior part broken) : condylo-incisive length probably

about 23; breadth of brain-case 9°5; least interorbital width

4:1; tooth-row 9°3,

Hab. Jebel Marra, at about 6000’.

Type. Adult male. B.M. No. 23.1.1.57. Original No. 545.

Collected Feb. 27, 1921, at Wadi Kongei, East Central Jebel

Marra.

C. h. diana, described from Lake Chad, is apparently the

nearest ally of this Shrew. The darker colour, longer tail, and

more normal dentition of marrensis force us, however, to treat it

as a distinct subspecies.

16. CRocIDURA ARIDULA, sp. 1.

6. 846, 861, 928. 9. 872. Kulme, Wadi Aribo. 3300’.

A grey Shrew resembling C. duéleri Thos., but with a longer

and unswollen tail.

Size medium. ‘Tail normal, more than half the length of the

-head and body.

Colour of upper parts cold-grey, near the “deep greyish olive ”

of Ridgway. Under parts white, in sharp contrast with. upper

MAMMALS OBYAINED IN DARFUR. 253

parts along flanks. Hands and feet white above, ‘Tail above

concolorous with back, white below.

Skull essentially as in butleri.

Dimensions of type (measured in the flesh) :—

Head and body 88 mm.; tail 43; hind foot 14; ear 12.

Skull: condylo-incisive length 23:3; breadth of brain-case 9°8 ;

least interorbital width 4:2; tooth-row 9°8.

Hab. as above.

Type. Adultmale. B.M. No, 23.1.1.61. Original No. 928.

Collected Sept. 21, 1921, at Kulme, Wadi Aribo. _ 3300’.

This species is sufficiently distinguished from C, 6ulleri by its

normal tail and somewhat colder colour.

17. CrociDuRA MARITA, sp. n.

©. 673. §.E. Downs, Jebel Marra. 8650’.

A very small Shrew, related to nana and religiosa. Fur

moderately long, hair of back measuring 3-4 mm. in length.

Colour of dorsal surface dark olivaceous grey; underparts

greyish white ; without any hard flank-line of demarcation.

Hands and feet whitish. ‘all indistinctly bicolor, dusky above,

whitish below.

Skull flattened. Second and third unicuspids about equal in

size.

Dimensions (measured in the flesh) :—

Head and body 56°5 mm.; tail 38; hind foot 10°5; ear 7°5.

Skull: condylo-incisive length 16°6; breadth of brain-case 7:5 ;

least interorbital breadth 3°4; tooth-row 7:3.

Hab, as above.

Type. Adult female. B.M. No. 23.1.1.63. Original No, 673.

Collected April 21, 1921. §.E. Downs, Jebel Marra, altitude

8650".

18. Fetis ocrvara Gmel.

3. 450. El Fasher.

3. 1062. 35 miles N. of El Fasher.

@. 655. Foot-hills, 8. Jebel Marra, 4000’.

“Lives in company with Fennee Fox. Said to be rare and

local, living in colonies like rabbits.”

19. Frvis servaw Schr.

©. 855. Kulme, Wadi Aribo. 3300’.

“ Native name Git.”

20. HERPESTES SANGUINEUS Riipp.

6.1141. El Fasher.

3.1061. 35 miles N. of Hi Fasher.

3. 607,611. 92. 610,974,1019. Jebel Marra. 7100’-8000’.

2. 758, 759, 762. Kulme, Wadi Aribo. 3300’

254. MESSRS, OLDFIELD THOMAS AND M. A. C. HINTON ON

Of this strongly-marked species the Museum previously pos-

sessed only a single specimen, which was also obtained by Admiral

Lynes in this same region. It was originally described from

Kordofan.

21, Hyana Hy@na Linn.

go. 412. 100 miles W. of Nahud.

6. 641. Foot-hilis, S. Jebel Marra. 4500’.

@. 874 (skull only), Kulme, Wadi Aribo. 3300’.

“« Does not appear to be found over 5000'.”

22. Crocuta crocorra Hrxl.

3. 911. Kulme, Wadi Aribo. 3300’.

“This is the common species found here. Very destructive

to sheep, goats, and donkeys. This individual stole three sheep

out of our servants’ huts.”

23, CANIS ANTHUS SOUDANICUS Thos,

6. 426. 16 miles EH. of El Fasher, Darfur. Jan. 29, 1921.

3. 596,597. Central Jebel Marra. 10,000’. March 10, 1921.

“Very common, and noisy at night. Usually seen in pairs.”

3. 683. §.E. Downs, Jebel Marra. 8000’. April 25, 1921.

“Very abundant on Downs.”

3. 985 (skull only). Jebel Marra. 9500’. ‘Feeds on

insects which it obtains by scratching in the grass-roots.

Stomach contained a large amount of vegetable matter, no doubt

swallowed in the process.”

2.1001. Jebel Marra. Dec. 6, 1921.

@. 1015. Niurmya, Jebel Marra. 7000’. Dec. 29, 1921.

24, VULPES PALLIDA Rupp.

6. 829, 857. @. 858, and male skulls 914, 923. Kulme,

Wadi Aribo. 3300’.

6. 971. S.E. Jebel Marra. 8000’.

“‘ Native name [at Wadi Aribo] Doctorri.”

‘“¢ Found in large colonies [at Jebel Marra.”

25. LoroNyx SPRIATUS SUDANICUS, subsp. n.

3. 666. Foot-hills of Jebel Marra, 4000’. April 16. B.M.

INOS aig WAS i 5 Shia.

An even lighter-coloured form than intermedius, with which

it shares the narrowing of the black hairs of the nape and fore-

back, and the breaking of the supraorbital black bar. Size

decidedly larger than in intermedius, but only one specimen of

each ayailable for comparison. Fur long, loose, aud shaggy.

MAMMALS OBTAINED IN DARFUR. 255:

Frontal white spots of medium size, the bar separating it from

the temporal one on each side considerably broken by white hairs.

Black lines of nape very narrow, much overlaid by the white

hairs, Posterior black lines also at a minimum, the median one

on the loins scarcely more than an inch broad, and its imtensity

much reduced by the long overhanging white hairs, the lines

throughout less developed and sharply defined than in any other

form. Chin and interramia largely intermixed with white hairs,

a few only being present in intermedius and none in other forms.

Black of under surface considerably narrowed in the ventral

region, the white of the flank-bands encroaching on the belly on

each side, while all across the latter there is an admixture of

white with the black hairs. Tail with the basal portions of the

hairs black as usual.

Skull fairly large, decidedly larger than in the type of inéer-

medius, but smaller than in shoe.

Dimensions of the type (measured in the flesh) :—

Head and body 355 mm.; tail 255 (not quite perfect); hind

foot 58; ear 24.

Skull: greatest length (median) 68-5; condylo-basal length

66°5; zygomatic breadth 38-7; interorbital breadth 18; inter-

temporal breadth 16°3; mastoid breadth 33:7; palatal length

32:4; maxillary tooth-series 20; p' on outer edge 6:5.

Hab. and Type as above.

A form with the white markings at a maximum, in correlation

with the desert characteristics of its habitat. J. erythrec 1s

smaller, show larger and very dark-coloured, and «wtermedius

somewhat similar in colour, but with much smaller skull and the

usual wholly black underside.

26. HUXERUS CHADENSIS Thos.

. 436. Hl Basher.

. 460. 35 miles S.W. of Hl Fasher.

. 1114. Jebel Maidob.

This desert Ground-Squirrel ranges along a strip at about

14° N. lat. for a considerable distance, as Capt. Buchanan

obtained it at Zinder, French Nigeria; the typical series came

from Lake Chad, and the present region is again much to the

east of that lake. It is of very pale colour, and comparatively

small (skull 58-61 mm.), though still much larger than the little

Ki. agadius of the Air region (51:5 mm.).

+O +O O,

27. HUXERUS ERYTHROPUS LIMITANEUS, subsp. n.

. 722. Zalingei, mouth of Wadi Aribo. 2800".

. 803. 92. 837, 854. Kulme, Wadi Aribo. 3300’.

. 657. Foot-hills of Jebel Marra. 4000’.

. 609. Central Jebel Marra. 7200’.

. 1. Dilling, Nuba Land. 2300’.

O,4 +O Os OQ, +O

256 MESSRS. OLDFIELD THOMAS AND M. A. GC. HINTON ON

Larger than chadensis and leucoumbrinus. Colour about as in

the latter.

Upper surface of body very much as in leucowmbrinus, that is

to say near ‘‘sayal-brown,” decidedly darker than in chadensis.

Light lateral line sharply defined white. Dull lateral line darker

than the back.

Skull decidedly larger than in chadensis and leucowmbrinus, the

total length of the skull about 61-65 mm.

Dimensions of the type (measured in the flesh) :—

Head and body 270 mm.; tail 246; hind foot 65; ear 17.

Skull: greatest length 62; condylo-incisive length 58:5 ;

zygomatic breadth 33°5; nasals 19; palatilarlength 31:7; cheek-

teeth exclusive of p’ 12; lower cheek-teeth 13.

Hab. Region of Wadi Aribo, extending eastward to Jebel

Marra. Type from Zalingei.

Tope. Adult female. © IB Mas No: 23.1 oOo ine Orietnall

number 722. Collected 20 May, 1921.

Distinguished both from lewcowmbrinus * and chadensis by its

larger size, and from chadensis by its darker colour, though the

Jebel Marra specimens are a little paler than those from the

Wadi Aribo.

28. HELIOSCIURUS BONGENSIS CANASTER, subsp. n.

6. 671, 705. @. 663, 706. Foot-hills of Jebel Marra.

4000’.

A paler and greyer form of the H. bongensis of the Bahr-el-

Ghazal.

Size as in bongensis, Dorsal colour as in that animal or imcon-

spicuously paler. But sides, forearms, hips, and backs of legs

far paler, whitish or greyish white. Under surface quite white

to the bases of the hairs, a patch of buffy on each side of tarsus.

Top of muzzle greyish white, much paler than the crown. White

supra- and infra-orbital lines well marked; ears prominently

whitish. Upper surface of hands and feet greyish white. Tail

similarly ringed to that of bongensis, but paler throughout.

Skull about as in bongensis, equally smaller than that of mzdliz-

color. Nasals rather longer.

Dimensions of the type (measured in the flesh):—

Head and body 178 mm.; tail 217; hind foot 44; ear 15.

“Skull: greatest length 44; condylo-incisive length 38°7;

zygomatic breadth 26; nasals 14x 6:8; interorbital breadth 13 ;

maxillary tooth-row 8°8.

* With regard to lewcoumbrinus, that species was founded by Rtippell in so vague

and general a manner that it needs pinhing down to some particular form, as 1t may

have included quite a number of distinct races. ‘There is in the British Museum one

of his original specimens, purchased of the Senckenburg Museum (B.M. No. 79 6), and

this we propose to select as a lectotype. It is of a pale cimnamon-brown colour, and

corresponds in size with specimens having a skull-length of about 58 mm., its lower

cheek-teeth 12 mm. It was received as from “ Abyssinia,” but how far this is

trustworthy we are not in a position to state.

MAMMALS OBTAINED IN DARFUR. SY Ti

Hab. as above.

Type. Adult male. B.M. No. 23.1.1.107. Original number

705. Collected 8 May, 1921.

This Squirrel is a whitened desert representative of the Bahr-

el-Ghazal H. bongensis Heuglin, a species distinguished from the

better-known and widely distributed A. multicolor by its

decidedly smaller size.

29. GRAPHIURUS OROBINUS Wagn.

3. 963, 965. W. of Jebel Marra. 4000!

g. 771. ©. 809, 918, 922. Kulme, Wadi Aribo. 3300’.

©. 882,930. Zalingei. 2800’.

Type-locality, Sennaar.

30. TaterA RoBusTA Cr.

3. 462, 463, 464, 466, 467, 499, 1068, 1086, 1128. 9. 424,

443, 447, 456, 457, 458, 459, 460, 461, 465, 470, 480, 481, 482,

483, 484, 485, 490, 494, 495, 1144, 1152, 1154, 1155, 1173, 1175.

Hl Fasher and neighbourhood.

3d. 1185. WNahud, Kordofan.

$6. 1198. @. 402, 1194, 1208. El Obeid.

g. 1112. @. 1087, 1088, 1089. Jebel Maidob. 2750’.

9. 1088. ina, Wells.

31. TarerRA BENVENUTA Hint. & Kersh.

3. 784, 807, 832, 836. 2. 729, 801, 873. Kulme, Wadi

Aribo. 3300’.

3. 549. 2. 548,550. Wadi Kongei. 6200’.

dg. 529. 9. 508,523, 531, 585, 586, 649, 658, 700. Jebel

Marra. 4000'—7900’.

TTATERILLUS.

The genus Z'aéerdllus is represented by nearly 70 specimens,

covering the whole area explored, except the upper part of Jebel

Marra, where its place is taken by the special local form of

Dipodiilus described below.

In the district round HE] Fasher, two forms are found—the

large bright-coloured 1. rufus, with its naked whitish soles, and

also a smaller species.

The other members of V'aterillus in the collection all have the

hair-band on the soles, and are related to 7. buileri of Bahr-el-

Ghazal, which seems to extend northwards into Kordofan, and

we confess we do not see any reason to distinguish from it either

T. kadugliensis or Taterina lorenizi of Wettstein. The distinction

of Laterina trom Taterillus as a genus is based on a character—

58 MESSRS. OLDFIELD THOMAS AND M. A. CG. HINTON ON

the presence of an extra cusp in m,*—-which is far from constant,

and many specimens of Yaéerillus have a rudimentary cusp in

the same position. Major Graham has sent examples of Taterillus

from §S. Kordofan, which we may take as representing both

kadugliensis and lorentzi, and these quite agree with the type of

butlert from Bahr-el-Ghazal.

32. 'TATERILLUS CLIVOSUS, Sp. 0.

3. 509, 516, 522, 525, 539, 645, 707. 9. 519, 520, 521, 524,

647, 698, 699, 708, 1032. Jebel Marra. 4000'—5500'.

$. 770, 830, 847, 860. 2. 820, 848, 850, 862, 864, 865,

869. Kulme, Wadi Aribo. 3300’.

Q. 726. Zalingei. 2800".

6.473. 35 miles W.S.W. of El Fasher.

©, 1102,1106. Jebel Maidob.

Like 7. butlert, but with longer and rather more bushy tail.

Size about as in bwilert, or a little larger. General colour

buffy brown, a little lighter than in butleri, but decidedly darker

than in the plains form to be deseribed below. Face without

special white markings. Hars of medium size, buffy brown,

scarcely different from the back. Hands and feet white, soles

blackish, generally with well-marked and often with very ‘broad

hair-bands. Tail decidedly larger than in buéleri, and more

heavily pencilled, its terminal two-fifths with blackish hairs

upwards of 13 mm. in length, its upper surface brown, its lower

buffy whitish. The type has a white tail-tip, but this is

evidently abnormal.

Skull without special peculiarities, about 35-37 mm. in length.

Dimensions of type (measured in the flesh) :-—

Head and body 126mm.; tail 182; hind foot 33; ear 20. ~

Skull: greatest length 37:5; condylo-incisive length 34; pos-

terior palatine foramina 4:2; bulla 10; upper molar series 5:4.

Hab. Slopes of Jebel Marra below about 6000’, and region of

Wadi Aribo. Type from Jebel Marra, south, 5300’.

Type. Adult female. B.M. No. 23.1.1. 127. Cugizal number

‘698. Collected May 5, 1921.

* We are quite agreed that in systematic work it is, at least for the present and

for a long time to come, better to use the conventional notation of m}, m*, m® for the

three cheek-tecth of Muride, rather than any notation which attempts to indicate

their real homologies as compared with the cheek-teeth of other rodents. Mr.

‘Kellogg has recently, im a paper dealing with certain Californian Voles (Univ. Calif.

Publ. Zool. xxi. p. 245, 1922), described and figured the anterior tooth as the last

premolar (p*), the second tooth therefore as m}, and the third m?, a method which is

liable, in systematic work, to give rise to a good deal of confusion. AJl the more

that his revised nomenclature is by no means acceptable to other writers, as will be

seen froma paper by Hinton (Ann. & Mag. N. H. (9) xi. p. 162, 1923), who reviews

the whole question, and has reason to confirm the view that the anterior tooth is a

milk-premolar (mp*) and not a permanent one, a conclusion which Thomas also

thinks may be the true one.

But in either case the conventional notation m1, m2, m? would seem to be the best

for current systematic work.

MAMMALS OBTAINED IN DARFUR. 259

This Gerbil, which is found in a more elevated region than

T’. butleri, resembles that species in its general colour, but differs

by its longer and more bushy tail.

The determination of the last three specimens in the list is a

little doubtful, but they may be provisionally assigned to the

present form.

33. TATERILLUS PERLUTEUS, Sp. n.

6-417. Q. 416, 418, 419, 1160, 1161, 1178. Um Kedada,

100 miles H. of El Fasher. 2400’.

Q. 444, 446, 491, 492, 501, 1069, 1151, 1180. Neighbour-

hood of Kl Fasher.

Oy 1084, 1122) 1123, Tagbo Bilis) 795) amilesw iN. Bae of

Kl Fasher.

A smaller species of vivid buffy colour.

Size decidedly less than in bwtleri and clivosus. General colour

bright clear buffy, about asin rufus and in gracilis angelus, much

paler and clearer than in butleri and clivosus. Upper surface

near ‘‘ warm buff,” darkened a little along the median basal area,

clearer on the sides. A patch behind eye and another behind ear

white. Hars buffy. Hands and feet white; soles brown with

well-marked hair-band. ‘ail pale buffy above, white below, its

terminal third with a brown pencil.

Skull distinctly smaller than that of 7. elivosus.

Dimensions of the type (measured in the flesh) :—

Head and body 106mm.; tail 149; hind foot 30; ear 18.

Skull: greatest length 34; condylo-incisive length 30; nasals

13°3; palatine foramina, anterior 5°4, posterior 3°7; bulla 10;

upper molar series 5.

Hab. Plains of Darfur, round and to the east of El Fasher.

Type from Um Kedada, about 100 miles H. of the capital.

Type. Adult female. B.M. No. 23.1.1.143. Original number

1160. Collected March 15, 1922.

The bright buffy colour of this Gerbil distinguishes it from

butleri and clivosus, while from the equally bright rufus it differs

by its smaller size and the presence of hair-bands on its soles.

34. TATERILLUS RUFUS Wettst.

Tatera rufa Wettstein, ¢.c. p. 111.

dg. 479, 493, 506, 1080, 1169. 9. 507. W. of El Fasher.

©. 1166. Um Kedada, 100 miles E. of El Fasher.

@. 1090. Jebel Maidob. 2750’.

Dr. Weitstein has in our opinion laid too much stress on the

presence of the hair-band on the sole as diagnostic of the genus

Taterillus, while quite ignoring the length of the posterior palatal

foramina. It is clear from his excellent photographs that his

Tatera rufa has the long foramina characteristic of Taterillus,

2.60 MESSRS. OLDFIELD THOMAS AND M. A. C. HINTON ON

while, as Thomas has shown, Z'aterillus is often without the

hair-band. '

In this species the sole is whitish and generally completely

naked, the ears are unusually large, and the general colour is.

particularly bright and vivid.

7'. gyas Thos., from the Dinder River, is related to T. rufus,

but is larger.

“ Soles of feet white.”—W. P. L.

35. TATERILLUS BUTLERI Wrought.

3S. 1200, 1202, 1206. &. 404, 404%, 1199, 1201. El Obeid

and neighbourhood.

©. 409, 1189. Near Nahud, Kordofan.

A rather dark form with comparatively short tail.

386. GERBILLUS PYGARGUS FEF. Cuv.

g. 1091. - 9. 1092, 1104. Jebel Maidob. 2750'.

©. 1083. Hl Fasher.

37. GERBILLUS AGAG Thos.

2. 406, 408. Nahud, Kordofan.

@, 413, 414, 415. 100 miles W. of Nahud.

@. 423. 65 miles H. of Nahud.

6. 445, 476, 1060, 1129, 1148. ©. 451, 474, 475, 476, 486.,,

Ne NUR. MOTs Jk, Wes WG, MIRO, Sa, WSs NaS. JUS,

1146, 1156, 1157.. El Fasher and neighbourhood.

©. 1168. Um Kedada, Darfur.

Hitherto only known from the type, a male obtained in

November 1902, at Agageh Wells, by Capt. Dunn.

38. GERBILLUS NANCILLUS, Sp. n.

@. 425. 16 miles EH. of El Fasher.

©. 1078. 45 miles N. of Hl Fasher.

A minute species smaller than any hitherto known.

Size even less than in Dipodillus henleyi. General colour

above very pale gerbil-colour; only a narrow dorsal area with the

usual dark bases to the hairs, those of the flanks with white

bases and bufiy tips. Under surface snowy-white, the white area

taking in the whole of the fore limbs. Middle of face buffy ;

cheeks, a ring round the eyes, patch at base of ears, and another

behind them white. Hars small, their proectote contrasted

brown, with the tip whitish, their metectote buffy. Soles of

hind feet partly naked behind, but with the hairy terminal part

of metatarsus characteristic of Gerbillus. ‘Tail practically white,

the upper surface very slightly more buffy, the end lightly

pencilled with brown.

MAMMALS OBTAINED IN DARFUR, 261

Skull with proportionally large brain-case and short muzzle.

Bulla fairly large, much smaller than in Dipodillus henleyi.

Dimensions of the type (measured in the flesh) :—

Head and body 54mm.; tail 79; hind foot 17-5; ear 11.

Skull: greatest length 20°5; condylo-incisive length 17;

nasals 7:1 ; breadth of brain-case 10-2; anterior palatine fora-

mina 3:2; posterior palatine foramina 2:2; bulla 7; upper molar

series 3°2.

Hab. Plains of Darfur. Type from. 45 miles N. of El Fasher.

Pype. Adult female. B.M. No. 23.1.1.169. Original number

1078. Collected February 6, 1922.

This pretty little Gerbil reminds one of the tiny Dipodillus

henley of Lower Heypt, but the soles, though partly naked

behind, are distinctly those of Gerbillus, in which genus this is

much the smallest species known.

39, DIPODILLUS LOWET, sp. n.

3. 566, 577, 583, 587, 595, 600, 601, 603, 605, 625, 676, 677,

678, 679, 684, 685, 975, 979, 981, 986, 990, 992, 995, 1005, 1016,

1105.

@. 564, 565, 578, 581, 582, 584, 588, 591, 592, 593, 594, 602,

604, 618, 620, 624, 674, 675, 680, 681, 688, 689, 693, 694, 906,

973, 976, 978, 980, 982, 983, 984, 987, 988, 989, 991, 993, 995,

997, 998, 999, 1000, 1002, 1004, 1007, 1008. Jebel Marra.

6650'-9750'. )

A dark-coloured mountain form of the D. campestris group.

Size about as in VD. campestris. General colour dark, much

darker than the clear gerbil-colour of most members of this genus,

though occasional or younger specimens of campestris are similar.

Back dark buffy grizzled with the brown tips of the hairs, the

general tone near ‘“ Brussels-brown,” while some specimens even

approach sepia. Sides a little clearer buffy. Under surface as

usual wholly white. Kars brown, with the conspicuous whitish

markings behind eyes and at base of ears. Legs buffy brownish ;

hands and feet white. Tail with a fairly well-developed blackish

tuft on its terminal third, its hairs ordinarily attaining 8-9 mm.

in length ; under surface of tail dull buffy brownish.

Skull, on the whole, like that of D. campestris, but rather more

robust, and the zygomatic plate projecting slightly further for-

ward, so that its anterior point stands about 3:5 instead of

3°0mm. in front of the base of the notch, as viewed from above.

The bulle are of about the same length, but, apparently owing to

a less convex inner side, the slit between them and the front part

of the basioccipital is nearly invariably broader. Molars slightly

heavier.

Dimensions of the type (measured in the flesh) :—

_ Head and body 110mm.; tail 143; hind foot 25:5; ear 17.

Proc. Zoo. Soc.— 1923, No. XVIII. 18

262 MESSRS. OLDFIELD THOMAS AND M. A. C. HINTON ON

Skull: greatest length 31:5 mm.; condylo-incisive length 28°5 ;

zygomatic breadth 15:7; nasals 12°5; palatal foramina 6:3;

bulla 9; upper molar series 4°4.

Hab. Jebel Marra, above 6000’. Type from 9500’.

Type. Adultfemale. B.M. No. 23.1.1.212. Original number

982. Collected December 3, 1921.

‘‘ Lives under rocks; feeding on grassand flower-seeds. These

animals appear to be strictly nocturnal.”—W. P. L.

This Gerbil is of considerable interest, for while it is nearly

allied to the Algerian and Saharan D. campestris, no velative of

it is recorded from anywhere near Darfur; nor did the Lynes-

Lowe expedition obtain a single specimen of the group in all the

low country round Jebel Marra, so that we may be confident that

it is really isolated high up on the mountain. In the plains its

place would seem to be taken by Yaéerillus buileri, which, like

other Gerbils, is pale sandy-coloured, while the present animal

has become darkened to suit its rocky environment, just as has

happened with Lemniscorys lynest and other members of the

mountain fauna.

We have much pleasure in naming it after Mr. Willoughby

Lowe, to whose energy the great extent of the Darfur mammal

collection is mainly due.

40. D1poDILLUS PRINCIPULUS, sp. n.

@. 1119. El Malha, Jebel Maidob, N. Darfur. 2700’.

3.1127. 90 miles N.E. of El Fasher.

A bright buffy species with long tail and swollen brain-case.

Size medium, about as in D, stigmonya. General colour bright

sandy buffy, without the slight dorsal darkening generally found

in stigmonyx. Sides and posterior part of rump above tail without

slaty bases to the hairs, these being white with buffy tips. A

white patch behind eye and another behind ear. Ears buffy.

Hands and feet white, the soles naked. Tail longer and rather

more bushy than in stagmonyx, the upper surface lined buffy

brown, the lower side white; terminal two-fifths with a well-

marked brownish pencil.

Skull about as long as in stigmonyx, but much more swollen

posteriorly, the brain-case more convex above and considerably

broader ; bulle much larger. Palatal foramina shorter. Molars

about as in that species.

Dimensions of the type (measured in the flesh) :—

Head and body 73mm.; tail 115; hind foot 21; ear 11.

Skull: greatest length 26°35; condylo-incisive length 23;

zygomatic breadth 14:5; breadth of brain-case 13; height from

crown to lowest point of bulla 10°8 ; palatal foramina, anterior 4:1,

posterior 2°4; bulla 10x55; bi-meatal breadth 14; upper

molar series 3°5.

Hab. N. Darfur. Type from El Malha, Jebel Maidob.

2700’. Nk

MAMMALS OBTAINED IN DARFUR. 263

Lupe Adult) female. 4) bev Nowe 2on ol Als, /<Orieimnal

number 1119. Collected 10 February, 1922.

The members of the genus Dipodillus may be arranged accord-

ing to the extent to which the brain-case and bulle are swollen up

in proportion to the face, or are of more or less normal Murine

development. This species and the much smaller D. henley: are

at the swollen-headed end of the series; then among Kgyptian

species follows amenus; then stigmonyx with its allies or

synonyms duteolus and botéw, and the smaller-toothed watersi.

Then comes campestris and its ally dowei; and finally, and most

extremely murine, the small species next to be described.

This pretty and distinct species is one of the few novelties

resulting from the visit of the Expedition to the black rocky

mountain Jebel Maidob, in N. Darfur, whose mammal fauna

has proved far less striking than that of Jebel Marra.

41. DIPoDILLUS MURICULUS, sp. n.

6. 1082. Madu, 80 miles N.E. of El Fasher. 2900’.

A small species with a very murine-shaped skull.

Size very small, considerably less than in princepulus, little

larger than in henleyz. General external characters as in prinet-

pulus, except for the markedly shorter tail. Colours as described

in principulus, with similar white bases to the lateral and anal

hairs. The general tone is, however, more strongly buffy, but

how far this is affected by the only specimen being tinted with

the reddish sand of its locality, we are unable to say. Hye

and ear markings present. Hinder soles quite naked. ‘Tail

much shorter than in principulus, scarcely pencilled, the proximal

portion thinly haired and showing the scales.

Skull, for a Gerbilline, of a remarkably murine shape, the

cranial region neither broadened nor inflated and the bullz small,

so that at first sight, at least from above, the skull might be

taken for that of a J/us. Supraorbital edges well marked.

Palatal foramina narrow. Bulle comparatively small, oval, the

mastoid portion almost without inflation.

Teeth apparently as usual, but much worn in the only specimen.

Dimensions of the type (measured in the flesh) :—

Head and body 65 mm.; tail 82; hind foot 18; ear 12.

Skull: greatest length 23 mm.; eondylo-incisive length 20 ;

aygomatic breadth 12; nasals 8:5 ; interorbital breadth 4; greatest

breadth of brain-case 10°3 ; height from crown to lowest point of

bulla 8:8; palatal foramina, anterior 3°7, posterior 2:1; bulla

75x45; bi-meatal breadth 10°6; upper molar series 3:1.

fab. as above.

Type. Old male. B.M. No. 23.1.1. 220. Original number

1082. Collected 3 February, 1922.

A very striking little Gerbil, distinguishable by its short tail

and very murine skull.

18*

264 MESSRS. OLDFIELD THOMAS AND M. A. C. HINTON ON

42. DESMODILLISCUS BRAUER Wettst.

3.1192. 75 miles W. of El Obeid.

@. 1172. 140 miles E. of El Obeid.

The type of this remarkable little Gerbil was described from El

Obeid. These almost topotypical specimens are most welcome

additions to the collection.

They show that the Nigerian species D. buchanani is more

nearly allied to braweri than we had supposed, as the difference.

that we had observed in the first lower molar seems to be only

due to Dr. Wettstein’s photograph having been taken at an angle

which gave a misleading outline to the tooth. And as to the

difference in the bulle, those of buchanani had had the meatal

inflation broken off, an accident which was only perceived on

the examination of the present specimens. But the two species

are, however, still distinguishable by the greater size of the

molars of buchanani—3°2 mm. in length above as compared with

2-9 *

ad e& .

43. STEATOMYS AQUILO, sp. n.

©. 1013. Niurmeya, Jebel Marra. 7000’. Collected 23

December, 1921. B.M. No, 23.1.1.438. Type.

A small species related to the Nigerian cuppedius.

Size about as in cuppedius. General colour above pale wood-

brown, washed laterally with buffy. Under surface white, less.

sharply defined than usual. Face rather more buffy than back.

Ears brown with a small whitish patch behind them. Forearms.

and hands, and hind feet white. Tail brown above, whitish

below.

Skull very much as in cuppedius, but the palatal foramina

longer, extending nearly to the level of the middle lamina of m'

and narrower.

Incisors unusually strongly bevelled, so much so that in side-

view the part of the tooth behind the groove appears little

broader than that in front. In the type the upper incisors are

white, but we suppose this to be due to some accidental chemical:

deterioration ; the lower ones are yellow as usual.

Dimensions of the type (measured in the flesh) :—

Head and body 69 mm. ; tail 39; hind foot 14; ear 13.

Skull: greatest length (c.) 19:5; nasals 7:5 ; interorbital breadth

4; front of incisors to back of m° 9-7; palatal foramina 4:5 =

upper molar series 3:7,

fab. and Type as above.

“ Found dead on native trail with throat cut.”—W. P. L.

This represents the most northern occurrence of the genus.

Steatomys, although on the west side of Africa it ranges nearly

as far—to Kano, Nigeria. The species would seem to be nearly

allied to S. cuppedius, unless the peculiar white coloration of the-

* Alveolar lengths—buchanani 3°7, braueri 3:°2-3°4 (Wettstein 3°5).

MAMMALS OBTAINED IN DARFUR. 265

upper incisors is a normal character, a point which can only be

settled on the arrival of further specimens.

[The description of the following Steatomys, distinguished and

labelled some years ago, seems accidentally never to have been

published :-—

Steatomys gazelle, sp. n.

A large species, with large bulle and very large molars.

General colour above dark wood-brown, darker than in most

species. Under surface white, but only along a narrower area

than usual. Hors dark brown, without postaural whitish patch.

Hands and feet dull whitish.

Skull of about the same large size as in pratensis and caurinus,

but the bulle larger than in either, and the molars very much

larger. Palatal foramina long, widely open.

Dimensions of the type (measured on the skin) :—

Head and body (c.) 100 mm.; tail 52; hind foot 18:5.

Skull: greatest length 26mm.; condylo-incisive length 24:5 ;

palatiiar length 12:2; palatal foramina 5-4x 2-4; bulla 6°3;

upper molar series 4:7.

flab. Bahr-el-Ghazal; type from Tamburas.

Type. Adult male. B.M. No. 5.10.7.4. Collected and

presented by Maj. H, Bray.

Readily distinguishable by its large size and very large

molars. |

44, Rarrus (Mastomys) MAcroLEPIs Sund.

3. 453, 471, 505. 9. 452, 472, 496, 497, 500. Hl Fasher.

3d. 510, 518, 519... 2. 511, 512, 514, 515, 532, 533, 534, 535,

936, 538, 540, 1021, 1022, 1026, 1033, 1034, 1035. N.H. Jebel

Marra. 5500’.

3. 042. 9. 543, 1023, 1027. N.W. Jebel Marra. 6000’.

Sg. 637, 668, 703, 704. 2. 634, 635, 636, 640, 646, 650, 651,

652, 654, 659, 662, 667, 669, 701, 702. Foot-hills, 8. Jebel

Marra. 4000’.

3. 970. &. 962,972. S.H. Sebel Marra. 8000’.

6. 621, 623. 2. 622. S.W. Jebel Marra. 6650’.

g. 504, 561. &. 551, 555, 556, 558. Niurmya, Central

Jebel Marra. 7100’.

GeO 10, (lee 71s @ 5 BAG. G7, 370, DSO, Ola lee fale,

716, 717, 964. Central Jebel Marra. 5000'-7900'.

ie (OE Baa Oh | OTs). alo, Dats Ee eka:

Zalingei. 3300’.

Gi O34, 163,169) (OM MSniO2e COD OON Monel oon loo, 090.

798, 805, 806, 810, 812, 813, 816, 822, 825, 831, 835, 838, 839,

840, 842, 851, 863, 867, 868, 871, 912.

OME TOS Cole Val holon (Ost hay ihm aOOmaIUn Ia, Olio, :O2i,

828, 841, 845, 858, 870, 901,921. Kulme, Wadi Aribo. 3300’.

©, 1182. Nahud, Kordofan.

3. 1203. @. 1204. Abu Heraz, Kordofan.

266 MESSRS. OLDFIELD THOMAS AND M. A. C. HINTON ON

A very fine series of these difficult field-mice, which will be

of great use when a closer study can be made of the group.

There is considerable variation in size among them, and perhaps.

more than one form is included.

The two members of the subgenus Masfomys in this region are

respectively this wild-living form, more or less rufous, with a

white or whitish sharply detined underside, and a house-haunting

race, dark smoky grey, with little-defined dull greyish-bufty

belly—indeed, corresponding closely in colour to bouse-haunting

Mus musculus. A co-type of Sundevall’s us macrdlepis in the

British Museum (No. 46. 6. 2.58) certainly agrees more with the

former than the latter, and we have therefore used that name for

the species. We also consider that Wagner's Mus limbatus

should be placed as a synonym of it.

45. Rarrus (Mastomys)—ugande group.

6. 779, 866, 898, 899, 900, 904, 908, 913, 916, 917, 924.

@. 780, 843, 853, 895, 897, 902, 905, 910, 925, 927. Kulme,

Wadi Aribo. 3300’.

Gols. 649. 1ON7, 1030; L03I TONG, LOS a2 eS

Jebel Marra at various altitudes.

3. 879. ©. 938,953. Zalingei. 2800’.

The house-haunting members of Mastomys are very much alike

all over Northern Africa, and may even be all of one species.

For the present, however, we do not venture to give an exact

determination of these specimens.

46. ACOMYS WITHERBYI de Wini.

6. 420, 428, 448, 477, 1059, 1138, 1142, 1177. @. 421, 429,

430, 431, 432, 433, 487, 458, 449, 455, 1070, 1076, 1093, 1125,

1139, 1140, 11438,1145. Hl Fasher and neighbourhood.

. 1036. Kurra, Jebel Marra. 5200’.

. 606. Niurmeya, Central Jebel Marra. 7100’.

. 1121. Near Togato Hills.

. 407, 1184. ©. 1119, 1183. Neighbourhood of Nahud.

. 403, 405, 1190, 1195, 1196. @. 1191. Hl Obeid and

neighbourhood.

3. 1020. N.E. Jebel Marra. 6500’.

@ = 1096, 4.099; - I100e W107 4 1108," LUIS 1a Gl Mie:

@. 1087, 1101, 1103, 1109, 1111,1113. Jebel Maidob. 2700’.

Os Os +O Os +O

47. ARVICANTHIS TESTICULARIS Sund.

6. 1021, 1042, 1044, 1045, 1046, 1050, 1051, 1052, 1053, 1054,

1055, 1057, 1066. @. 434, 439,440, 442, 454, 1039, 1040, 1041,

1048, 1047, 1048, 1049, 1056, 1074. El Fasher.

6. 1067, 1072, 1075. ©. 1073. 35 miles N. of El Fasher.

3. 630, 664. 9. 629, 697, 709. South Jebel Marra.

©. 562. Niurmya, Central Jebel Marra.

©. 517, 1025. N. Jebel Marra.

MAMMALS OBTAINED IN DARFUR. . 267

S20) AI (42, ORs SAD GG Oe 20 Nn (21, 980,0947),

Zalingei.

3. 768, 833, 896. 9. 767, 884. Kulme, Wadi Aribo.

3d. 1110, 1120. Jebel Maidob.

©. 1205. El Obeid, Kordofan.

48. LEMNISCOoMYS DUNNI Thos.

6. 1158,1170. Y&. 435,441. El Fasher.

3. 1009, 1029. ©. 541. Jebel Marra. 5200'-7700'.

©. 1162, 1163, 1164,1167. Um Kedada.

Fully agreeing with the type, which was obtained by Col.

Dunn in the Kaga Hills in 1902. The species is apparently

common on the plains of Darfur including the lower slopes of

Jebel Marra, while it is replaced higher up on that mountain by

LL. lynesi.

Nearly related to it is the following form from the Nuba region

of Southern Kordofan :—

LEMNISCOMYS DUNNI NUBALIS, Sp. ND.

Similar in essential characters, in size and spininess of form, to

true dunnit, but the colour throughout much intensified, the stripes

and other parts, which in dwnzi are pale buffy, very strong and

rich buffy, almost ochraceous. Hars very deep buffy. Under

surface buffy laterally instead of white.

Skull as in dwnne.

Dimensions of type :—

Head and body 103 mm.; tail 180; hind foot 24; ear 14.

Skull: greatest length 29:5; condylo-incisive length 2677;

bulla 6; upper molar series 5.

Hab. Nuba Country, Southern Kordofan. Type from Talodi.

1300’.

Type. Adult female. B.M. No.18.7.2.14. Original number

5. Collected 21 February, 1917, and presented by Maj. C.

Graham. Three specimens.

Lemniscomys zebva Heugl., of which Thomas has examined the

type, has a wholly white underside, and is clearly a member

of the other group of this genus, with much darker-coloured

fur and smaller bullez. To this latter group the next species

belongs.

49, LEMNISCOMYS LYNESI, sp. 0.

6. 617, 690, 967. ©. 563, 590, 598, 691, 968, 1003, 1006,

1011. Jebel Marra, at altitudes from 7700’ to 9750’, and to the

crater.

A dark species with comparatively long soft fur.

Size fairly large. Fur comparatively long, and much softer

and more spineless than in any allied species. General colour

268 MESSRS. OLDFIELD THOMAS AND M. A. C. HINTON ON

dark, the light lines dull whitish, with a distinct tendency

to break into spots. Under surface buffy whitish, the bases of

the hairs broadly slaty, except on the chin. Ears dull ochraceous,

a richer ochraceous tuft at their anterior base. Hands and feet

dull buffy whitish. Tail black above, dull buffy whitish below.

Skuil with distinctly smaller bulle than in Z. dunn.

Dimensions of the type (measured in the flesh) :—

Head and body 112mm. ; tail 125; hind foot 25; ear 17.

Skull: greatest length 29; condylo-incisive length 26-6; palatal

foramina 5:1; bulla 5; upper molar series 5:3.

Hab. Jebel Marra, at various altitudes, from 7000’ upwards to

the crater. Type from Central Jebel Marra on western slope,

8000’.

Type. Adult male. B.M. No. 23.1.1.313. Original number

617. Collected March 22, 1921.

This distinct mountain species is readily distinguished by its

comparatively long soft fur, which is practically without spines,

in marked contrast with both LZ. dunni and zebra. Also by the

slaty bases to its belly-hairs and by its small bulle.

On freshly-killed specimens the attention of both collectors was

specially attracted to this animal by the greenish iridescent

sheen on its fur, though, as with so many mammals, this sheen

fades away in the dried skin.

We have much pleasure in naming this well-marked species

in honour of Admiral Lynes, to whose enthusiasm and scientific

spirit the National Museum owes the present magnificent acces-

sion to its collections.

50. GRAMMOMYS ARIDULUS, Sp. n.

do. 826. Kulme, Wadi Aribo. 3300'.

An unusually pallid, desert-coloured species.

General colour of dorsal surface dul! ochraceous lined with

dark brown, brightest and clearest upon the flanks and rump.

At the rump the tint is near Ridgway’s ‘“‘ochraceous-tawny.” Top

of head and nape duller and greyish. Region between eye and

ear on each side grey. A small white patch behind each ear.

Ears bright ochraceous tawny within and without, save fora mar-

ginal band of dark reddish brown extending from the base of the

proectote to the tip of the ear. Underparts cream, in sharp

contrast with flanks. Outer surface of forearms and upper

surfaces of hands and feet cream-buff. Tail bicolored, dusky

above, whitish below.

Skull most nearly resembling that of G. buwntingi, described

from Liberia, in form, but distinguished chiefly by its larger

bullee and heavier molars.

Dimensions (measured in the flesh):—

Head and body 115 mm.; tail 175; hind foot 24; ear 18.

Skull: condylo-incisive length 26°6; extreme length 29°4;

MAMMALS OBTAINED IN DARFUR. 269

zygomatic breadth 14:4; interorbital construction 4:3; cranial

width 12:8; dental length 13:5; cheek-teeth (crowns) 4:2.

Type. Old male. B.M. No, 23.1.1.324, Original number 826.

Collected July 22, 1922, at Kulme, Wadi Aribo, Darfur.

Altitude 3300’.

Hab. as above.

This is a very well-marked species distinguished by its bright

coloration from all forms previously described. In outward

appearance it is perhaps more like G. s. elgonis than any other,

while in skull it resembles the outwardly very different

G. bunting?.

. L&GGADA TENELLA Thos.

- 400. 30 miles W. of El Obeid.

. 627.. 2. 628. Foot-hills, S.W. Jebel Marra. 4500’.

. 712, 718. South Central Jebel Marra. 4000’.

. 599. Central Jebel Marra. 9750’.

. 587. Niurmya, Central Jebel Marra. 7150’.

. 695, 818, 824. 9. 903. Kulme, Wadi Aribo. 3300’.

Wtoaeagaa F

. JACULUS GoRDONI Thos.

@. 14. Near El Fasher. 2600’. May 1920 (20.7.30.17).

Incisor-buila length 35:0 mm.

53. JAcuLus gAcuLUS Linn.

go. 1135. 55 miles N.H. of El Fasher.

While it is not unnatural that the Kordofan J. gordont should

be found in Darfur, the capture of this specimen, clearly

referable to J. jaculus, so close to where the same collector found

the first-named species in 1920, is worthy of note. Moreover,

within the species J. jaculus, it has not the warm colour of the

Khartoum J. 7. buéleri, but is very similar to the more buffy

J. j. jaculus, which therefore would seem to range along the

deserts to the west of the Nile as far south as Darfur. The

nearest previous record for this form is Dongola.

54. Lupus HAwKERI Thos.

@. 1198. 50 miles W. of El Obeid.

¢. 410. 70 miles W. of Nahud.

©. 422. 75 miles KE. of El Fasher.

Gs (21, 136, 747,0(54, 90la, 2 iiatattee eaéalinge:, Darfur:

2800'—3000'.

3. 692. 2. 632. Foot-hills, 8. of Jebel Marra. 4000’.

©. 1014 juv. Niarmya, Jebel Marra. 7000’.

3. 696. Deriba Lakes, Jebel Marra. 7400’.

©. 977. Jebel Marra. 9500’.

2. 765 juv., 808. ?sex. 796 juv. Kulme, Wadi Aribo. 3300!.

270 MESSRS. OLDFIELD THOMAS AND M. A. C. HINTON ON

These agree very well with the type of L. hawkeri, described

from Kaka, Sudan. JZ. hawkeri would seem to be the repre-

sentative of isabellinus (“ ethiopicus”) in the open country west

of the Nile.

55, PHACOCHGRUS HTHIOPICUS subsp.

6. 800. Kulme, Wadi Aribo. 3300’.

@. 728 juv., skull only. Kulme, Wadi Aribo, 3300’.

Not improbably referable to P. e@thiopicus fossor Schwarz.,.

described from the Bagirmi District.

56. ALCELAPHUS LELWEL TSCHADENSIS Schwarz.

é. 760. Kulme, Wadi Aribo. 3300’.

$. 777, skull and head-skin only. Kulme, Wadi Aribo.

3300’.

3. 856. Kulme, Wadi Aribo. ‘Skull, tail,and fore and hind

legs only-—the latter to show the black markings, which are

unusually pronounced for these parts. Chin always black.”

57. SYLVICAPRA GRIMMIA PALLIDIOR Schwayrz.

¢. 644, 638. Foot-hills,S. Jebel Marra. 4000’. “ Tolerably

common on dry stony hill-sides.”

©. 823, 849. Kulme, Wadi Aribo. 3300’.

@. 752. Zalingei, Darfur, Sudan. 2800’. ‘Native name

‘ Digi-dig.’”

A very distinct form, of which we have had no satisfactory

material previously. Characterized by its pale colour and large

size.

58. OuREBIA OUREBI SPLENDIDA Schwarz.

6. 852. 9.791. Kulme, Wadi Aribo. 3300’.

“Found very sparingly in wocded country. Native name,

‘Gazella amma.’ ”

The female captured on July 2 contained a well-developed

foetus.

As the collectors noted, the female differs from the male in

having a dark patch on the head.

59. GAZELLA RUFIFRONS Gray.

©. 589. Central Jebel Marra. 10,000’. ‘Shot from a lot

of 5 on plateau.”

S$. 811. Kulme, Wadi Aribo. 3300’.

6.1181. Head only. 205 miles H. of El Fasher. “‘ Rarely

seen here and further H. along the road.”

We are not at present prepared to determine this material

subspecifically.

MAMMALS OBTAINED IN DARFUR. OAL

60. GAzELLA DoroAS Linn.

3. 1058. 10 miles N. of El Fasher. Horns 113”.

¢. 1094, 1095. 150 miles N.E. of El Fasher.

3S. 1147, skull only. 19 miles N. of El Fasher.

3g. 469. @. 470. Skulls only. 35 miles EH. of El Fasher.

Q. 427. 10 miles HE. of El Fasher.

Q. 1149, 1150. Near El Fasher.

@. 915, no skin. Wadi Aribo.

61. Procavia RuFIcErs H. & HE.

¢. 1065. 2. 1063, 1064. 35 miles N. of El Fasher.

@. 1131. 60 miles N.H. of El Fasher.

@. 773, 819. Kulme, Wadi Aribo.

3. 957, 958. @. 880, 959, 960. Zalingei.

¢. 1098. Jebel Maidob.

Native name, ‘‘ Bogos.”

62. PROCAVIA RUFICEPS MARRENSIS, subsp. n.

¢6. 665. 92.670. Foot-hills of Jebel Marra. 4000’.

6. 552, 559, S16, 1012. ©. 553, 566, 608. Jebel Marra.

7100’.

S$. 527. 2. 526, 528. Jebel Owi, N.E. Jebel Marra.

6500’.

Rather larger on the average than true ruficeps; the fur

longer, the colour darker, and the dorsal patch less conspicuous,

more overlaid with the dark ends to the dorsal hairs. General

colour approximating to ‘‘Saccardo’s umber,” but often darker,

and always darker than the average in true rujiceps. Nape dark

brown, like the crown. MHairs of back broadly tipped with -

blackish brown in a specimen in good pelage. Hind back and

rump and the thighs dark brown. Under surface dull buffy,

darker than in rujficeps.

Skull averaging larger than in rujiceps.

Dimensions of type (measured in the flesh) :—-

Head and body 504mm.; hind foot 71; ear 30.

Skull: condylo-basal length 91:5 ; zygomatic breadth 55 ; front

of p' to back of m° (teeth much worn) 33.

Hab. Jebel Marra. Type from the central part of the mountain

at 7100’.

Type. Oldmale. B.M. No. 23.1.1.486. Original number 616.

Collected March 19, 1921.

This is a dark mountain race of the widely spread P. rwficeps

of the Soudan. It is unusually variable in colour, but certainly

averages considerably darker than the Dassie of the plains.

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OMPHALO-MESENTERIC ARTERIES IN THE MANATER. Dis

19. On a Remnant of the Omphalo-mesenteric Arteries in

the Manatee. By K. Kostanrcxt, M.D., LL.D.,

F.Ac.Se. Cracow *.

[Received February 19, 1923: Read April 10, 1923.]

(Text-figure 1.)

In a paper published in 1897 in the Proceedings of the Zoological

Society (Notes upon the Anatomy of a Manatee, Manatus

imunguis), Beddard describes and represents in a drawing the

eecum of this animal. The general shape of the cecum is

precisely like that of the other species of Manatee, especially of

Manaius latirostris, one specimen of which Beddard has also.

examined, and which previously had been described by Murie 7.

Beddard adds: “I should not have had the drawing prepared

were it not for a peculiar fold of mesentery which it is the main

purpose of that sketch to illustrate. This fold, which is not

referred to by Dr. Murie, lies on either side of the mesentery

supporting the ileum and runs nearly to the cecum. It does not

bear a blood-vessel, and the fold of either side is continuous

with its fellow by a complete bridge over the front side of the

ileum as indicated in the sketch. Both species are precisely alike

in the presence and in the relations of these two mesenteries.”

(The drawing in Beddard’s paper is also reproduced in Weber's

‘ Siiugetiere.’) d

Neither Beddard nor Weber gives an explanation of this.

fold.

I have had the opportunity of examining a Manatee feetus

(Manatus imunguis) 35 cm. in length in the collections of the

Royal Museum of Natural History at Brussels, thanks to the-

kindness of Prof. Dollo.

The cecum presented the aspect known in the adult animals

from the papers of Cuvier, Owen, Home, Murie, Flower, Rapp,

Huntington, Beddard, Waldeyer, and others.

I noticed immediately a few cms. away from the cecum the

bilateral folds described by Beddard, which ran from the dorsal

mesentery to the wall of the ileum, uniting on its ventral,

antimesenteric wall. However, they did not terminate there

in a narrow bridge, as in Beddard’s drawing, but the central

part was markedly elevated above the surface of the ileum,

and was elongated into a narrow strip stretching towards the

umbilicus and terminating there (as can be seen in the text-

figure, p. 274).

* Communicated by THE SECRETARY.

+ Murie, “On the Form and Structure of the Manatee,” Trans. Zool. Soc. viii.

p. 127.

“274 DR. K. KOSTANECKI ON A REMNANT OF THE

The connection of these folds with the umbilicus furnishes an

immediate explanation of the condition in the adult animal

described by Beddard. Without any doubt, we have before us a

remnant of the involution of the vitelline duct and its blood-

vessels, especially its arteries. I have not been able to dissect the

precious specimen and make microscopical sections, but examining

the strip and its relation to the ileum, I noticed that the wall

of the ileum between these folds was smooth; the vitelline duct

had completely disappeared, and blood-vessels were not apparent

under the peritoneum forming these folds; it may be that

microscopical sections would have disclosed degenerated remains

of the walls of these vessels. In spite of that, there can be no

doubt, considering the course of this strip, that it owes its

existence to structures connected with the yolk-sac apparatus.

Text-figure 1.

Cxceum and umbilicus of Manatee.

b.c.=bifid cecum.

Especially regarding the symmetrical origin of the two folds

from the dorsal mesentery of the terminal portion of the ileum,

there can be no doubt that these are produced by the omphalo-

mesenteric arteries. The omphalo-mesenteric or vitelline vein

or its remains cannot play any part here; in feetal life it runs

differently, namely, from the umbilicus to the mesoduodenum.

In the foetus which I have examined it had vanished completely.

The changes of the omphalo-mesenterice arteries are described

by several authors, especially in the very exact paper by

Broman*, who, together with the resuits of previous researches,

gives many new data.

It is known that the omphalo-mesenteric arteries are originally

paired and run on either side of the ileum to the umbilicus, then

unite, both in the part which runs through the mesentery,

* Broman, “ Uber das Schicksal der Vasa vitellina bei den Siugetieren,” Ergebnisse

.der Anatomie und Entwickelungsgeschichte, Bd. xxi. (1918).

OMPHALO-MESENTERIC ARTERIES IN THE MANATEE. 275

forming one arteria omphalo-mesenterica, the future superior

mesenteric artery, and in the part where, ventrally to the ileum,

they run towards the umbilicus; Broman calls this last part—

which runs outside of the gut together with the vitelline duct,

towards the umbilicus and in the umbilical cord—the vitelline

artery.

In some animals the arteriz omphalo-mesenterice, though not

paired in their proximal and distal parts, remain for some time

paired in the part which runs on both sides of the ileum, thus

forming a ring around the ileum, which, starting dorsally from

the single arteria omphalo-mesenterica, passes on the ventral

side of the ileum into the single arteria vitellina, which runs

toward the umbilicus in the mesodermie vitelline stalk.

In most animals this part does not remain paired, the artery

on one side disappearing (the left in some animals, in others

the right), the persisting artery of the other side running on the

lateral surface of the ileum, but deviating from its wall, so as to

produce a peritoneal fold; usually this fold later on loses its

connection with the wall of the ileum and thus, as the beginning

of the arteria vitellina, passes on the mesoileum.

When after the disappearance of the vitelline duct, the vitelline

artery also disappears, the mesodermal yolk-sac-stalk grows

thinner and thinner, ruptures, and finally vanishes.

Broman in the paper I have cited proves that during this

process we meet with variable conditions in different animals.

Thus, in some animals until birth the vasa omphalo-mesenterica

can remain visible always, in others abnormally; but usually

they disappear in the feetus, and leave behind them folds,

running towards the umbilicus, which persist in some animals

only for a short period of thei: foetal life, in others until birth

and sometimes even in extra-uterine life in the adult animal.

These folds, running towards the umbilicus, rupture, and either

vanish completely or (either only in foetus or new-born or adult

animals) form bands adherent to the dorsal mesentery, brt

hanging freely down into the abdominal cavity.

The remains of the vitelline vein which run towards the

mesoduodenum, Broman calls “appendix mesoduodeni,” the

remains of the vitelline artery ‘‘ appendix mesilei”; any part of

the ruptured yolk-duct-stalk remaining near the umbilicus,

‘appendix umbilicalis,”

Broman noticed in an adult Castor fiber in the place where in

other new-born animals, or their feetuses, could be seen the

appendix mesilei, a remarkable fold, running from one side of

the mesileum to the wall of the ileum; he explains it as the

remains of the fold, formed by the artery joining the omphalo-

mesentevic artery in the mesileum and the arteria vitellina in the

yolk-sac-stalk by separating from the wall of the ileum. «

I see in this case of Castor fiber a complete analogy with the

condition in the Manatee, except that in the Manatee, in the place

of the appendix mesilei of other animals, such a fold appears on

276 OMPHALO-MESENTERIC ARTERIES IN THE MANATEE,

both sides. This shows that in the Manatee the paired parts of

the vitelline arteries, which enclose on both sides the ileum, must

have persisted for a longer time than in other animals, and,

deviating symmetrically from the wall of the ileum, must have

formed symmetrical peritoneal folds. As in Castor jiber the

one-sided fold, so here the paired folds persisted in the adult.

animal.

The fact, that those paired folds meet on the ventral wall of

the ileum, forming a sort of bridge, appears quite natural,

considering that the omphalo-mesenteric arteries joined on the

ventral wall of the ileum and from there ran toward the

umbilicus in the yolk-sac-stalk, which in the foetus which I

examined was still preserved.

P.Z.S. 1923; STUART BAKER, Pi. |

H.Gronvold, pinxit. John Bale,Sons & Danielsson, L“4

CUCKOOS’ EGGS AND EVOLUTION.

PZ /-S.Si23, sh UART BAKER ae eile

jie!

H.Grénvold, pinxit. i John Bale.Sons & Danielsson.

CUICKOOS) TECCS, JAN DEVO UmlON:

BP Zs lZSs SnUAR Ty EAE Se eats

[lass

H.Grénvold, pinxit. John Bale.Sons & Danielsson.

CUCKOOS’ EGGS AND EVOLUTION.

P.Z.S. 1923, STUART BAKER, PI. IV.

H.Gronvold,pinxit. ; John Bale,Sons & Danielsson, L*?

CUCKOOS’ EGGS AND EVOLUTION.

CUCKOOS’ EGGS AND EVOLUTION. Die

20. Cuckoos’ Eggs and Evolution.

By E. C. Sruartr Baxesr, F.Z.8.

[Received March 19, 1923: Read April 24, 1923.]

(Plates T= hve)

Much has recently been written on Cuckoos and Cuckoos’

eggs, in some cases the writers having laid down laws and

accepted facts as proved on very insufficient data. Certain of

these writers, who have watched the actions of individual

Cuckoos and who have spent much time and money on their

valuable observations, have theorized on the presumption that

what one Cuckoo does all Cuckoos do. Other writers, also from

watching one bird, essay to prove that the former are wrong

because their observations do not have the same result as their

own. Asa matter of fact both are equally correct, the result of

their observations are of great value, and their work worthy of

all praise so long as they do not pose as authorities, able to prove

or disprove theories, until their experiences are far wider.

For over forty years now I have studied Cuckoos, and the

older I grow and the more I know about these birds, the more

certain Iam that I have not yet sufficient data to enable me to

do more than roughly outline certain theories and point out what

there is in support of them.

Curiously enough, the most interesting of all Cuckoo subjects

is the one which seems to have been most avoided. I refer to

the evolution of Cuckoos’ eggs resembling the eggs of their foster

parents sufficiently well to enable them to escape destruction.

Perhaps this is because our British Cuckoo, Cuculus canorus

canorus, is only a very recently evolved type of Cuckoo and

within its few thousand years of existence has not yet had time to

eliminate those members of its species which lay eggs dissimilar

to those of the birds it selects as fosterers. To find really satis-

factory instances of perfected or far advanced evolution, or of

evolution in an obvious stage of transition, we have to go abroad

and examine the eggs of such genera as Clamator, Coccystes,

Hierococcyx, etc. Amongst these we may hope to find sufficient

evidence to enable us to show what has already taken place and,

by analogy, to learn what is probably taking place at the present

day amongst other genera and species.

The material from which I have drawn my deductions consists

of some 2500 Cuckoos’ eggs collected from Europe, Asia, and

Australia (not Africa), but principally from India where Cuckoos,

of many species, are exceedingly common. As I have already

* For explanation of the Plates see pages 293, 294.

Proc. Zoou. Soc.—1923, No. XIX 19

278 MR. E. C. STUART BAKER ON

said, this material is insufficient for me to pretend that I have

learnt all that there is to know about Uuckoos and their eggs, but

it seems sufficient to prove that :—

(1) In some genera and species evolution is one in its earliest

stages 5

(2) Tn others it has progressed much further; and

(3) In certain genera and species it has arrived at perfection

or finality.

Before proceeding to examine these three conditions in detail,

it may be as well to propound the theory of evolution as accepted

by me in this connection.

It will be probably generally admitted that the original pro-

genitors of Cuckoos laid white eggs and that they continued to

do so for a very long period after all trace of their reptilian

ancestry had disappeared. In fact, this persists to the present

day amongst those Cuckoos of the genera Centropus, Rhopo-

dytes, etc., ete., which are not parasitic. The belief of course

entails the acceptance of the theory that parasitic Cuckoos have

acquired their curious habit and not that non-parasitic Cuckoos

have grown out of it. If we accept this theory of an original

white or nearly white egg, it will be necessary also to accept the

fact that there has been some degree of evolution in almost all

parasitic Cuckoos’ eggs, for white eggs are very rare amongst

Cuckoos and even those which are white are quite different in

character to the glossless dead white or yellow-white of most

reptiles’ eggs. If, however, we accept the theory that all

Cuckoos’ eggs were originally white, we have again to accept the

probability that all other birds’ eggs were white and, this being

so, we should expect that the Cuckoo's egg would have developed

pari passu with the eggs of the bird in whose nest it was

deposited. Other birds’ eggs have acquired certain coloration,

shapes, sizes, etc., in each case for some definite purpose which

has aided it to escape destruction. In some instances the

purpose for which the character has been acquired is obvious.

Many birds which lay their eggs on the ground lay such as are

beyond doubt protectively coloured; some lay eggs conspicuously

coloured to enable the owners to distinguish their own eggs

among colonies; others lay eggs so shaped that they cannot roll

off narrow ledges of cliffs; yet others lay eggs so large that the

chicks emerge fledged and able to take care of themselves from

the start, whilst in many other instances we are still too ignorant

of the demands of nature to know why the characters exhibited

have been acquired.

Why then have not the eggs of all parasitic Cuckoos arrived

at the same degree of evolution as those of other birds? Simply

because Cuckoos are a modern ereation and even amongst

Cuckoos the parasitic habit is more modern than the bird and

amongst those again which have acquired this habit, some have

taken to it later than others.

All sorts of fantastic theories have been advanced as to how

CUCKOOS’ EGGS AND EVOLUTION. 279

Cuckoos’ eggs have attained similarity to those of their fosterers

but really this evolution has been brought about in the simplest

manner, We may presume that at one time all the eggs of any

one particular kind of Cuckoo were white and that they were

laid promiscuously in any birds’ nests which were handy. If

among these there had been a suitable fosterer with white eggs,

there would have been little or no evolution or alteration in the

Cuckoo’s egg. What would have occurred would merely have

been that those white eggs placed in fosterers’ nests with white

eggs would have been hatched in far greater numbers than those

which were placed with contrasting eggs and gradually the latter

would have been eliminated whilst the former would have sur-

vived. This is, indeed, what seems to have happened in a few

cases as I will show later on. But in the majority of instances

it is evident that there were either no suitable fosterers laying

white eggs or that they were not in sufficient numbers to be able

to act both as foster-parents to the Cuckoo and also perpetuate

their own race. Amongst the Cuckoos’ eggs, however, there

were probably some which were abnormally coloured with faint

blue, red, or some other colour. Now, if these were placed

alongside the eggs of foster birds with somewhat similarly

coloured eggs, they would escape detection in greater proportion

than the quite pure white ones, whilst among the former again

those eggs with the deepest tinge would escape more often than

the others and so, just by the gradual elimination of the most

unfit, we finally obtain a coloured egg so close to that of the

fosterer that no further elimination is required.

Evolution in the Cuckoo's egg has, in fact, proceeded as in all

other phases of evolution. Destruction and creation are synony-

mous terms in evolution ; the unfit have perished and the more

fit survived, the latter in their turn being wiped out as yet

better adapted individuals took their place.

In discussing the three degrees of evolution, referred to above,

it will probably be more convenient if we examine the third

degree, 7. ¢., that of perfect evolution first, and bring forward

whatever proofs are possible in support of my theories.

Perfect evolution in a Cuckoo’s egg must mean that a degree

of resemblance in colour, combined with size and shape, has been

arrived at which is sufficient to ensure the continuation of the

Species in its proper numerical proportion with, or without, other

controlling factors. There will, therefore, be no variation in

such egg and comparatively little variation in the genera and

species of birds selected as fosterers. Finally, such a state of

evolution also infers that there is an ample supply of suitable

fosterers laying eggs of the type to resemble which the Cuckoo’s

ege has been evolved.

No finer examples can be given of perfected evolution than the

eggs of Clamator jacobinus, the Pied Crested Cuckoo, and Hiero-

coccyx varius, the Common Hawk Cuckoo, the two most common

19*

280 MR. E. GC. STUART BAKER ON

Cuckoos in India. These Cuckoos lay deep blue eggs which they

deposit in the nests of birds of the genera Argya and Turdoides,

two of the most common and universally distributed genera of

birds in the Empire of India, which also lay similarly coloured

eggs of about the same size and shape. Here we have two

Cuckoos of different genera laying eggs so closely alike that it is

practically impossible to separate them except by their size and

the colour of the yolk. Even in size, however, they constantly

overlap whilst, though the Hawk Cuckoo generally lays earlier

in the year than does the Crested Cuckoo, in many places both

birds are laying simultaneously. The genera Argya and T'ur-

doides both lay unicoloured blue eggs, very glossy, hard-shelled

and broad, blunt ovals in shape. The Cuckoos both lay eggs

very similar in colour but with a softer, more satiny surface,

less glossy and with much thicker shells. In shape they are more

spherical or elliptical and in size generally a trifle larger than

the eggs of Turdoides and much larger than those of Argya.

In some instances both these eggs, as I have already said, are

practically indistinguishable from those of the fosterer when

lying side by side, though when being blown they are very easily

distinguishable by the great thickness of the shells of the

Cuckoos’ eggs and by the colour of the yolks which are a much

paler yellow, especially in the eggs of Clamator.

In these instances we have all the factors which seem to be

necessary to hasten evolution. In the first place we have as

suitable foster-parents two nearly ailied genera containing

numerous species of birds which are very numerous from the

level of the plains up to about 4000 feet over practically the

whole of India, Burma, and Ceylon. In addition to this, their

breeding-season is very protracted and they are excellent

mothers and fathers. Finally, they are very clever birds, and

possibly elimination of dissimilar eggs would proceed with these

fosterers faster than with most.

Another and almost exactly similar instance of perfect evolu-

tion is that of Clamator coromandus, the Red-winged Crested

Cuckoo. This bird has a different breeding range to that of the

last two, and is found from the foothills of the Eastern Hima-

layas, throughout the hills of Burma up to about 5000 feet, but it

is most common as a breeding species at 1000 to 2000 feet. The

most common birds within this area are the Laughing Thrushes

of the family Zimaltide. Amongst these the genera Dryonastes

and Garrulax contain the most numerous and most widely spread

species and amongst these again the two species Garrulax mont-

liger and G@. pectoralis are the two which are found throughout

this great extent of country in the greatest numbers. These two

birds, which are outwardly extremely alike, are the ones we

should expect to find acting as fosterer to the Red-winged

Crested Cuckoo, and so it is. These two Laughing Thrushes both

lay pale blue eggs, and the Cuckoo lays an egg exactly similar

in colour and about the same in size but differing not only in

CUCKOOS’ EGGS AND EVOLUTION. 281

texture but also in shape for, whereas the Laughing Thrushes

normally lay rather long, oval-shaped eggs, this Cuckoo lays the

most spherical of all eggs in the family.

This is rather interesting, for we find that not only is a

spherical egg a very common form in reptiles, but that the

more primitive non-parasitic Cuckoos also often lay spherical

eggs. It would rather seem, therefore, as if shape was not a

characteristic of much importance in birds’ eyes, a theory which

is confirmed by the difference in shape obtaining between other

Cuckoos’ eggs and those of their fosterers. Probably we may

take this as a case of “complete evolution ” if not of ‘“ perfect

evolution,” as all that is necessary has been achieved by the

production of an egg similar in size and colour to that of the

fosterer.

Yet another instance of complete, though not perhaps perfect,

evolution may be found in the eggs of the Koel, Hudynamis

scolopaceus, which lays its eggs in the nest of various species of

the Corvide. Here we have birds, looked upon as the most

astute of all the Passerine birds, regularly duped by a smaller

bird, although everyone who knows India and who has taken

the most superficial interest in birds, is fully aware that the

Crow, whatever his species or race, has the bitterest instinctive

hatred of the Koel.

The Koel is a large bird, and really belongs to a section of the

Cuckoo family, the other members of which are non-parasitic.

She naturally requires fosterers who are in a position to obtain

a very large quantity of animal food for the voracious youngsters

and, probably, in the area she occupies, the Crow is the only bird

found from Ceylon to the Malay Archipelago whose habitat is

as widespread as her own and who could furnish such food.

Accordingly we find that an egg has been evolved by the

process of elimination sufficiently perfect for the purpose of

hoodwinking the Crows and ensuring perpetuation of the

Cuckoo. Crows’ eggs are too well known to need much descrip-

tion but, roughly, they are pale bluish in ground-colour, with

profuse blotching of dull red all over their surface, the

general effect of the colour-scheme being a dark greenish egg.

The Cuckoo’s egg has been improved and perfected, until we get

a type very similar in general colour to that of a Crow’s egg

but it is not alike in shape, for it is generally a shorter, blunter

oval and in size it is much smaller.

The above three examples may all be said to be instances of

complete evolution but I purposely use the term perfect evolu-

tion only in reference to the first two birds’ eggs (C. jacobinus

and H. varius). Here we have eggs so close to those of their

foster-parent in colour, shape and size that often human eye-

sight is not acute enough to distinguish between the two. In

the second case, the eggs of the Red-winged Crested Cuckoo and

those of the Laughing Thrushes are very alike in colour and also

in bulk, though their shape betrays them at once to the human

282 » MR. E. C. STUART BAKER ON

eye. In the third case, although we have an excellent and

apparently sufficient simulation of the colour of the Crow’s egg

by the Koel’s egg, yet any child could tell the difference between

the two.

After reading the above, it will doubtless be asked whether

the Cuckoos referred to do not ever deposit their eggs in birds’

nests other than their normal fosterers.

As regards the first two birds, H. varius and C. jacobinus,

certainly one hundred eggs will be found in the nests of Argya

and 7'urdoides to every one in any other bird’s nest.

Clamator coromandus lays at least ten times in the nests of

G. pectoralis or G'. moniliger to every once in other birds’ nests

and even then she generally deposits her egg in the nest of some

other Garrulax, Dryonastes, or Trochalopterum, which are very

closely allied birds and which lay eggs either very much like, or

rather like, those of the proper fosterers.

The Koel may be said never to lay in any nest but in that of

the true Crows. The only exceptions to the many thousands of

eggs recorded are a few deposited in the nests of Magpies in

Northern Burma and West China and one instance of its being

laid in a Myna’s (Acridotheres tristis) nest.

We may say, therefore, in the three examples given, the first

including two species of Cuckoo, we have eggs which have

arrived at a point in their evolution which is perfect or complete.

Even if not perfect in human eyes, the adaptation in resemblance

between the eggs of the Cuckoo and foster-parent is so great

that a single type of the latter suffices over the whole range of

the former to ensure a sufficient number of young Cuckoos being

annually reared to. replace normal loss. Further, the resemblance

is sufficiently perfect to ensure deception of the fosterers without

there being any necessity for variation in the Cuckoo’s egg

The above may, I think, be taken also as examples of wholly

stabilized evolution, but it is perhaps advisable to explain that

even this stability varies somewhat in degree. ‘Thus in the eggs

of both the Crested Cuckoos and in Hierococcyx varius the colour

is always the same and varies only very slightly in depth of tint.

In the Koel, however, the variation is greater and, in fact, in

this respect it may even be an advance in evolution over those of

the others, for the variation in tint of colour, of extent of

marking, etc., may all be evolved in order to give the possessor a

better chance of escaping rejection. ;

All Crows’ eggs vary greatly in colour ; some are pale, some

dark; some greener, some browner; in some the reddish

markings predominate, in others they are almost absent. To

suit such variation in colour there are two possible types of

evolution which would occur to us as most useful: first, the pro-

duction of an average coloured egg which would agree with an

average number of ‘Crows’ eggs; secondly, a similar variation in

degree of coloration in both Crows’ and Cuckoos’ eggs, so that an

average number of eggs would match. The latter scheme is the

CUCKOOS’ EGGS AND EVOLUTION. 283

one which has prevailed and its value is curiously increased

when we find, as we do, that in certain areas both Crows’ and

Koels’ eggs show a certain tendency to a certain type, or degree

of colour intensity. Thus the eggs of both Crows and Cuckoos

in Assam eastwards average dark; those from the dry areas of

Northern India average paler, whilst those from the dry desert

areas of Sind, etc., are palest of all. Then when we come to

Ceylon and the wet coasts of Malabar and Travancore, we again

come back to dark eggs, whilst the drier regions of Ceylon seem

to have a special reddish type confined to that island.

There are, of course, many more similar instances of perfect

or complete evolution. For instance, there is Clamator glandarius

in Spain and other parts of EKurope which deposits its egg in the

nests of Magpies but the examples given are sufficient, I hope,

to prove that in some cases certain species of Cuckoos always lay

the same type of egg and, normally, always cuckold the same

species of foster-parents.

There is, however, another form of complete evolution which

is even more interesting than the last and this is dimorphic

evolution, or even, if 1 may use such a word, polymorphic

evolution.

There are some species of Cuekoo which cover a far wider

range of country than is covered by any one possible fosterer,

and in some instances of this nature we find in their eggs the

most wonderful contrasting examples of evolution.

The two most striking instances of dimorphic evolution known

to me are to be found in the eggs of the two Cuckoos, Cuculus

poliocephalus and Hierococcyx sparveroides.

Cuculus poliocephalus is a small Cuckoo which ranges from the

Western Himalayas through the mountains of China to Japan.

Over much of her western area she deposits her eggs in the nests

of several of the small warblers of the genera Phylloscopus and

Acanthopneuste. All these little birds lay either pure white eggs

or eggs which are white, sparsely spotted. Moreover, she more

often selects as fosterers those species which lay pure white eggs

rather than those which lay spotted ones and her favourites

seem to be Acanthopneuste occipitalis and A. magnirostris. With

these fosterers a white egg is laid.

In Japan C. poliocephalus is as common as she is in the Hima-

layas, but here she selects as fosterer the equally common little

Cettia cantans, a species which lays a remarkably beautiful deep

terra-cotta or chocolate-pink egg. She seems to deposit her eggs

almost invariably in this bird’s nest and, accordingly, the egg she

lays is similarly coloured, matching extremely well the fosterer’s

egg in everything but size. I believe the Rev. F.C. R. Jourdain

has received specimens of this Cuckoo’s eggs with other fosterers

but he informs me that these eggs were taken by a Japanese col-

lector about whom we know little and the late Alan Owston,

who sent me over 30 of this Cuckoo’s eggs, said he had never

known it lay in anything but nests of Cetéia. Throughout the

284. MR, E. GC. STUART BAKER ON

Chinese Hills I have little evidence of a sufficiently reliable

nature for me to say what kind of egg the bird lays, but what

little there is would lead one to suppose it to be the same as the

Japanese form. In Eastern India our evidence is ample and

here we find that the two types overlap so that from Assam to

Garhwal we get both the pure white and the chocolate type but

the first is the common one to the West, the second the more

common in the Hast. In India, however, we have no true

Cettia and accordingly we find that the normal fosterers in this

country for the red eggs are the various species of the genus

Horornis, all of which lay dark chocolate eggs which go well with

the dark form of Cuckoo’s egg. Other nests in which the Hima-

layan Cuckoo deposits her terra-cotta eggs are those of Testa and

Oligura, both genera (if they are divisible from one another) of

birds which lay pink eggs not contrasting greatly with those of

the Cuckoo.

It is true that with this last Cuckoo we have a great many

instances of both red and white eggs being placed in nests

containing eggs which are quite unlike their nermal fosterers

but this is what oceurs more or less with every Cuckoo. It

occurs least of all with those Cuckoos in which evolution has

proceeded farthest, most with those Cuckoos in which the

destruction of the unfit has only just commenced to operate. In

the present case, however, neither of these factors can be con-

sidered alone. Here we have a bird, the eggs of which in each

extreme of its habitat have by the process of elimination arrived

at one definite type, which is in every way complete in so far as

its local needs are concerned, whilst in the neutral country

between the two extremes we have both types of eggs laid and

consequently we obtain frequent instances of mistaken depositing.

Eggs so deposited are probably not hatched in many instances

and we shall thus, by the gradual elimination of the unfit, finally

get two definite types of egg laid by the same species of bird

within the same area, but always normally placed in its proper

type of foster-parents’ nest.

The next example of Cuckoo shows a further advance in this

direction. Hierococcyx sparveroides lays two types of egg which

contrast quite as strongly in colour as do those of Cuculus polio-

cephalus and we here also have a coincident variation in the size

of the Cuckoos’ eggs.

This bird, the Large Hawk Cuckoo, lays two types of eggs.

The first is a dark olive-brown egg, varying somewhat in depth

of colour and in the degree of olive tint. The surface is hard

and glossy and the shape a very elegant, pointed oval. This egg

is laid in the nest of the Great Spider-Hunter, Arachnothera

magna and at least 20 eggs of the Cuckoo will be found in this

bird’s nest to one elsewhere and even of these latter the great

majority will be taken in the nests of the Little Shortwing,

Heteroxenicus nipalensis. The eggs of both the Spider-Hunter

CUCKOOS’ EGGS AND EVOLUTION. 285

and the Shortwing are very like those of the Cuckoo, the former

more especially.

The second type of egg is a bright pale blue, similar in other

respects to the brown egg, but much bigger. This egg is deposited

in the nests of the Laughing Thrushes which lay eggs of a

similar, sometimes almost indistinguishable, type to those of the

Hawk Cuckoo. Another favourite fosterer is the Whistling

Thrush, Myiophoneus h. temminckti, which lays eggs very like

pale, washed-out specimens of those of blackbirds.

Now the blue eggs are laid by this Hawk Cuckoo all over its

range in the Himalayas from the North-West, where they were

taken by Col. Rattray, A. E. Osmaston, and others, right away

to the Burmese Hills, where a great number have been taken

by Messrs. K. Macdonald, J. M. D. Mackenzie, C. Hopwood,

P. Wickham, and others. The brown type, on the other hand,

appears to occur only in Assam, and even there practically only

in the hills south and east of the Brahmaputra. The reason,

however, does not require much searching for, as it is only in

this tract of country that the Spider-Hunter occurs in sufficiently

large numbers to ensure a constant supply of fosterers. Even

in Assam, however, the blue-egg laying Cuckoo is almost as

common as the other, so that here we get about an equal number

of both eggs. There is, in spite of this, no intermediate form of

egg in this area, nor do we ever find a blue egg in a Spider-

Hunter’s nest or a brown egg in that of a Laughing Thrush.

The last two examples are, as I have already pointed out, not

quite parallel. C. poliocephalus Jays two contrasting types of

egg. In each extreme of its habitat only one form is laid,

agreeing closely with the one type of fosterer selected but in the

centre of this Cuckoo’s habitat the two types of egg occur in

equal numbers and the process of elimination has not yet arrived

at a point where it has succeeded in wiping out all individuals

which lay their eggs in wrong nests. H. sparveroides has

advanced a stage further; it also lays two strongly contrasting

types of egg, but it lays one type of egg which is blue from

extreme east to extreme west of its habitat, whilst in one com-

paratively small district, only some 700 miles long, it lays a

second brown type of egg which has been evolved to suit

exclusively one particular fosterer, which is sufficiently numerous

and, presumably, exceptionally suitable in other respects.

In both the above examples evolution seems to have advanced

to a stage of completion or stability in regard to coloration and

size of egg but in the one, C. poliocephalus, there still remains a

complete degree of stability to be acquired in the selection of

fosterers over a portion of the area inhabited. I should, how-

ever, say that, even in this instance, I personally place little

importance on the fact that numerous eggs are laid in the nests

of more or less abnormal fosterers, as this can be accounted for

in many rational ways. This point I deal with more fully later

on.

286 MR. E. C. STUART BAKER ON

Turning now to instances of evolution which are incomplete

but which show considerable advance, there can be no better

example than that of the little Cuckoo, Caccomantis passerinus.

This Cuckoo is found in different geographical races from Ceylon,

through practically the whole of the Indian Empire. In many

portions of its area we do not yet know what are its normal

fosterers or we have not yet sufficient evidence to show to what

degree evolution has progressed. We may say, however, that

over most of its range it lays principally in the nests of Cisticola,

Orihotomus and Franklinia ; in the Nilgiris and adjoining hills

it breeds freely, making use of the nest of Prinia inornata ; in

Assam and the North-East, where it is possibly more common

than anywhere else, the fosterers selected are of the genera Suya

and Cisticola; whilst in Hyderabad (Deccan) Prinia socialis seems

to form the sole fosterer.

Has anything been evolved towards perfecting the resemblance

of the eggs of Caccomantis to those of its many fosterers? The

answer is “ Yes, much”; and if we take them seriatim as given

above, we shall find that this is so.

Roughly speaking, the eggs of Cisticola, Orthotomus and

Franklinia are either white or pale blue in ground-colour and

are speckled with dark brown in the first-named; blotched,

spotted, or speckled with various shades of red or brown in the

second and third. Orthotomus is the most common of the three

genera and also the most universally distributed and with the

longest breeding-season. Caccomantis over all this area lays an

egg very like that of Orthotomus and not strikingly unlike that

of the two other genera. The ground-colour varies from white

to blue just as it does in the fosterer’s eggs but the blotehes are

generally larger, paler and more smudgy in character than in

any of the other eggs except those of Orthotomus and they are

also decidedly larger.

In the Nilgiris the Cuckoos’ eggs seem always to be pale blue

with blotches of reddish, whilst those of its local fosterer, Prinia

inornaéa, are blue with dark, bold blotches of reddish black or

blood-red.

In Assam the fosterers selected, Swya and Cisticola, lay,

the former, eggs which are white to rather deep blue-green,

marked in many ways with specks, spots, and small blotches of

reddish to blackish brown, the latter, white or very pale blue

eggs speckled with brown. Of the two fosterers, birds of the

genus Swya are the more popular, both khasiana and crimigera

being impartially cuckolded.

Many of the Cuckoos’ eggs agree fairly well with the eggs of

(Msticola but many cannot, unless one tests the shell and weighs

the egg, be distinguished from those of the other foster-parents

of the genus Swya. I have seen as many as fifty eggs of these

birds in one season in the Khasia Hills, but I am quite sure

that many of these would have escaped my notice had I not been

always on the look-out for Cuckoos’ eggs and naturally suspicious

CUCKOOS’ EGGS AND EVOLUTION. 287

of any egg in the least differing from the others. Once suspicion

was aroused and one knew the differences to expect, it was, of

course, very easy to say whether or not the egg was that of a

Cuckoo. The differences are those to which I have already

referred, viz., a very stout, heavy shell of rather coarse texture,

a feebly coloured yolk and a weight far greater than that of

similarly sized eggs of the foster-parent.

Mr. C. F. M. Swynnerton has already referred to the question

of polymorphic colouring of the eggs of Cuckoos (Ibis, 1918,

p- 127) and also of the foster-parents, and I also have com-

mented on it (Ibis, 1913, p. 384).

When we come to Hyderabad we find the most extraordinary

case of nearly completed evolution which I know of in the whole

history of Cuckoos’ eggs. The bird selected as fosterer within a

very small area only a few miles across, round about this city, is

little Prinia socialis, a bird which lays a bright red egg, utterly

unlike the eggs of any of the other fosterers. Yet we find with-

in this tiny area, where the Cuckeo is very common, a red egg

has been evolved in order to ensure the preservation of the

species. Why Prinia socialis should have been selected as a

foster-parent is not very obvious, for though it is extremely

common in Hyderabad, breeding in every garden and hedge, yet

Orthotomus also is not rare and one would have thought would

have sufficed. Evidently, however, Prinia socialis is the better

foster-parent for local needs as it has completely ousted the

Tailor-Bird within Hyderabad, though only a few miles away

from the city the former bird is deserted and the latter again

imposed upon.

It would seem that this evolution of a red egg is comparatively

modern, as it has not reached the stability of any of the eggs

which I have cited as examples of complete evolution. The

original discoverer of this red egg was Professor K. Burnett of

the Hyderabad College, to whom I owe a beautiful series, others

being taken by Col. R. Sparrow, Sergt. F. Kemp, etc. Professor

Burnett having found two red eggs took one and sent it to me.

The second was hatched and the bird, when fully fledged, also

forwarded to me and its identity so made certain. In collecting

a series, however, Professor Burnett found it much easier to get

fully red eggs than such as were intermediate between them and

the usual types of this Cuckoo's eggs. After many years con-

stant work by Professor Burnett aad a final most generous gift

to me of his whole series, I have been able to link up the red

eggs with the others, thus giving a most perfect demonstration

of an egg in the course of evolution. Unfortunately these little

eggs fade very rapidly, possibly because of their recent evolution,

and neither plates, magic-lantern slides, nor the eggs themselves,

give a proper idea of their beautiful colouring when fresh.

With this example before us it hardly seems necessary to

labour further the question of whether Cuckoos’ eggs are still

undergoing some phase of evolution, but in discussing the

288 MR. E. GC. STUART BAKER ON

evolution of the eggs of our Common Cuckoo, Cuculus canorus,

the point will repeatedly come up. Cuculus canorus, which may

be taken as the type of Cuckoo in which evolution is most back-

ward, is the Cuckoo with the greatest range of all Cuckoos,

ranging practically throughout Europe and Asia as well as

through much of Africa. The geographical races with which I

am best acquainted are 0. c. canorus, breeding in Europe, C. c.

telephonus, breeding in Western and Northern Asia, and

C. c. bakert, breeding in the Eastern Himalayas, Burma and,

probably, Western tropical China where the mountains are high

enough, 2.e. 5000 feet and over. The degree of evolution

obtained varies greatly in the different geographical races and,

moreover, it even varies greatly in individual strains in each

race.

Taking first our English Cuckoo, Ouculus canorus canorus,

we do not find that evolution has advanced very far. The com-

mon fosterers cuckolded by this bird are the Wagtail, Motacilla

alba, the Robin, the Reed-Warbler, the Pipits, Anthus pratensis

and Anthus trivialis and the Hedge-Sparrow. There is certainly

no example of perfected evolution in England. If we take the

Hedge-Accentor first, we find that there is no such thing as a

blue Cuckeo’s egg known in England, although one or two

reputed eggs are in collections. The nearest approach to blue

eggs is probably shown by the series collected by Mr. Owen

round about Felsted, which are a rather pale blue-grey, sparsely

spotted and blotched with darker. If we examine large series

of eggs taken by genuine collectors with the three fosterers,

Pipits, Wagtails, and Reed-Warblers, we find that the Cuckoos’

eggs go very fairly well with all of them and also that the con-

trast 1s not very startling as between them and the Robin’s eggs.

This looks, therefore, as if evolution had arrived at a point at

which an inconspicuous egg had been evolved which agrees com-

paratively well with the five normal fosterers, whilst in respect to

the Hedge-Sparrow it would appear that this bird is so extremely

foolish that little or no evolution has taken place, or indeed, is

necessary.

A closer examination, however, of large series of eggs, although

it shows no case of evolution such as is found in the eggs of the

Cuckoos we have previously dealt with, does show signs of some-

thing taking place in the way of elimination of the unfit. In one

hundred sets of each of the five fosterers named, I find that of those

eggs placed in the nests of Wagtails, the majority are rather pale,

whilst the general trend of those placed in the nests of Pipits

and Reed-Warblers are rather dark. Amongst the eggs of

Cuckoos found in Robins’ nests there are certainly a greater

percentage of eggs with a reddish tint, although pale ones do not,

as we should expect, appear to be any more numerous than dark

ones ; finally, with the Hedge-Sparrow we find almost any type

of egg, though even amongst these we get Mr. Owen’s beautiful

series of rather bluish eggs.

CUCKOOS’ EGGS AND EVOLUTION. 289

In England, unfortunately, a large proportion of the Cuckoos’

eggs in collections have been supplied by dealers who, even if

they would not cheat intentionally, are themselves deceived by

the men who supply them, who are generally paid prices according

to the rarity of the foster-parent with which they are supposed to

have been taken. For this reason we have to dismiss from our

calculations about two-thirds of the eggs in private collections.

Outside the five best known fosterers, there are a few others well

authenticated which we may accept as normal fosterers, such as

the Sedge- Warbler, Spotted Flycatcher and one or two others.

As regards the rest, I cannot consider them as anything but

abnormal or casual. Dr. R. Williams, nearly forty years ago,

showed how certain Cuckoos which normally cuckolded Wagtails,

Robins, and Hedge-Sparrows, occasionally deposited their eggs in

the nest of practically any other small bird building within the

area they covered when they could not find a nest of the species

they really wanted.

Perhaps no one in England has personally taken more Cuckoos’

eggs than Dr. Williams, whom I found to be possessed of a wealth

of knowledge on the subject. The greater number of his eggs

were given by him to Dr. Rey but I was so fortunate as to

obtain nearly 150 from him after he had ceased sending them to

that gentleman. From these most interesting sets, I find that

practically all the casual eggs laid in such nests as those of the

Bullfinch, Linnet, Greenfinch, Wren, Chiff-Chaff, etc., can be

matched with those of Cuckoos which year after year deposited

their eggs in the nests of some one of the three birds which in

his vicinity provided the normal fosterers.

Continental Cuckoos seem to have advanced considerably

further in evolution than have the individuals whieh yearly seek

our shores for breeding purposes. True, we cannot say that in any

instance complete or perfect evolution has been arrived at but

many instances show that elimination has already been very busy

and thus certain fosterers have been accepted through sufficient

generations to allow of all those Cuckoos who laid eggs in striking

contrast to their own to be completely exterminated. Thus we

find that the Cuckoos’ eggs now placed in the nests of many birds

such as the Great Reed-Warbler, the Red-backed Shrike and a

few others, approach very closely both in colour and character to

the eggs of their foster-parents. The degree of resemblance varies

greatly, some eggs are an excellent match and some are only fair,

whilst strongly contrasting eggs are in a very small minority and

even these, of course, may have been deposited by Cuckoos

normally cuckolding other fosterers,

Perhaps the greatest advance shown by Continental Cuckoos is

the evolution of a blue egg to resemble those of the Redstart with

which they are placed. This blue egg seems normally to be laid

only by certain individuals which breed in the Baltic States and

East Prussia, more especially in Finland. Even here, however,

blue Cuckoos’ eggs are not always placed with the blue Redstarts’

290 MR. E. C. STUART BAKER ON

eges and it would seem that whilst a few individual Cuckoos

have reached a very high degree of evolution in this respect, the

great majority are still only in its earliest stages.

Leaving the European Cuckoo and moving farther east into

Asia, we find a very closely allied form, Ouculus canorus telephonus,

apparently depositing eggs which are much further advanced in

evolution. Unfortunately, we have not much material to work

on but even among the few eggs I have seen of this bird—under

two hundred—there appear to be many cases of advanced

evolution. The most noticeable, perhaps, are to be found in the

very close resemblance between the eggs of the Red-backed

Shrike, Lanius schach erythronotus, the Spotted Forktail,

Enicurus maculatus maculatus and the eggs of the Cuckoos

which respectively victimize these two birds.

Again, we find that Calliope seems to form a favourite fosterer

for this Cuckoo, and all the eggs placed in the nests of these birds

have a very distinct blue ground also, one or two being but very —

faintly marked.

These eggs were all taken in the extreme North-West of India,

but the farther east we work, the more advanced the evolution.

In the Central Himalayas the two favourite fosterers seem to be

Trochalopterum lineatum and Larvivora brunnea, or, in certain

areas, Saxicola torquata indica and Oreicola ferrea. The first two

birds lay unicoloured bright blue eggs, the last two, bluish eggs

very faintly marked with pale reddish. To go with these eggs

we have a beautiful blue egg evolved and it is seldom one finds a

Cuckoo’s egg of any colour but blue placed in the nests of these

birds. Working yet farther east we come to another race of

Cuckoo, Cuculus canorus bakeri, a very sedentary bird, breeding

at comparatively low levels and migrating for very short distances

into the plains in winter.

As with the other races, so with this species, the number of

fosterers cuckolded is very large, and we have no signs that

evolution has been in existence long enough to eliminate all but

those Cuckoo individuals which deposit their eggs in only the

nests of one, two, or a few of the birds which are the most

suitable for the purpose of bringing up their young. We do,

however, find that normally, the number of species called upon

to rear the young Cuckoo is much more restricted than it is

farther west. For example, in Assam, nineteen out of twenty

eges—perhaps a much greater pereentage even than this—will

be found in the nests of the two genera Cisticola and Suya, both

consisting of small Warblers which swarm all over the hills

above 4000 feet, or in the nests of Pipits. The Cisticolas are of

two species, C. c. cursitans, extremely common, and C. tytleri,

rather rare. The former lays a tiny white or bluish-white egg

speckled or spotted with reddish brown; the latter lays a bright

blue egg boldly spotted with very dark brown. The eggs of the

Suyas vary much more, indeed they may be said to be poly-

morphic, for the ground colour may be white, pink, bluish, blue-

CUCKOOS’ EGGS AND EVOLUTION. 291

green or greyish pink and the character of the markings is almost

as varied as the ground-colour. The most common .type of egg

undoubtedly has the ground white to pink or pinky-grey,

the surface profusely covered with small spots and specks of

reddish brown. Now evolution has produced for this Cuckoo in

Assam, eggs which vary from pure white with numerous small

specks of reddish, to deep pink heavily marked throughout with

various shades of reddish brown. The first type is generally

placed in the nests of Cisticola and agrees with the eggs of that

bird in all respects but size ; the second type of egg is most often

deposited in the nest of Suya and agrees well with the most

common red type of that bird’s egg.

As regards the Pipits, two species are called upon to do duty,

the first, Anthus richardi striolatus, a very rare bird which breeds

on one or two ridges only, and Anthus richardi rufulus which

breeds in considerable numbers up to about 5000 feet or a little

higher. Here the only signs of evolution traceable are in the

fact that the majority of Cuckoos’ eggs deposited in these nests

are dark and therefore do not contrast strongly with those of

the Pipit.

With C7. c. bakeri, blue eggs are much less common than they

are with C. c. telephonus, doubtless because suitable fosterers

laying blue eggs are not available. We, however, do find such

eges occasionally, and these most often in the nests of either

Liothria lutea or Mesia argentauris, birds which lay bright blue

eggs marked boldly with reddish brown or purple-black. In these

cases, too, we often find that the blue Cuckoos’ eggs are more or

less spotted. It would be possible to continue instances of

incipient and partial evolution in the eggs of Cuculus canorus

and its geographical races almost ad infinitum, but perhaps more

than enough has been written to prove beyond all doubt that

the three degrees of evolution taken as a basis for this article

do exist,

From Australia I have not been able to obtain any material to

help me much in my work. This has been because, as in

England, collectors vie with one another not in getting eggs to

prove some theory or to form a basis for another but in

getting eggs with a great variety of fosterer. Future work by

Messrs. Orton, Berthling, Archer, and others should prove of

great assistance and value and material should be available before

long sufficient to enable us to decide what is happening in

Australia.

I have not commented on many other Cuckoos, the eggs of

which are fairly well known, such as various species of Chalco-

coccyx, Hierococcyx nisicolor and fugax, Penthoceryx sonnerati,

Surniculus, ete., etc., as though none of these afford any evidence

contradicting what I have written, yet none of them afford such

good instances of the various stages of evolution as those I have

selected for the purpose.

Perhaps, before leaving the subject of Cuckoos and their eggs,

292 MR. E. C. STUART BAKER ON

I may be permitted briefly to refer to the method adopted by

Cuckoos for placing their eggs in the nests of other birds.

Mr. Chance has proved that sometimes at least, OC. canorus lays

her eggs direct into the nests of Pipits, whilst Mr. Scobey has

almost equally well proved that she does not always do so in the

nests of Wagtails but deposits them therein with her bill.

Now in India, Hierococcyx sparveroides could certainly never get

inside a nest of Arachnothera magna, not only because it is too

small, but the shape of the nest prevents it. On the only occasion

on which I have personally taken an egg of this Cuckoo from this

bird’s nest and also seen the Cuckoo at it, all the latter did was

to cling desperately for a second or so to the outside of the nest,

or to the stalk of the leaf beside it, whilst the parent Spider-

Hunters ineffectively hustled her. Certainly, I cannot say I saw

her place the egg inside the nest with her bill but, equally

certainly, she did not get inside it, yet the egg was there when

Tarrived. Again, I have'seen, and others have seen, Clamator

jacobinus deposit her egg in the nest of Argya with her bill,

though it is probable that both C. jacobinus and C. coromandus

do, not infrequently, lay direct into the nests of these genera.

Cuculus c. bakeri lays the great majority of her eggs in the

nests of Cisticola and Suya, both genera which build tiny egg-

shaped nests of grass far too small to contain the Cuckoo chick

when half grown, much less large enough to allow a Cuckoo to

sit in them. I have seen very many hundreds of these little

birds’ nests with Cuckoos’ eggs in them and it was obvious from

their perfect condition that no attempt had been made by the

Cuckoo to get inside them. When the young Cuckoo hatches

and commences to grow, the grass-work of the nests expands so

as to form a trellis-work round it until finally it bursts and the

young Cuckoo either falls to the ground or remains perched

on a pad of broken-down nest. But Cuckoos’ eggs have been

taken from nests even more impossible of access than these grass

nests. Thus Mr. J. Livesey and Capt. Bates found a Cuckoo’s

egg in the nest of a Phylloscopus built in a hole in a tree, the

entrance to which was only big enough for the tiny warblers to

go in and out. The nest was some way in and the Cuckoo’s egg

‘n rolling in had smashed one of the eggs of the owner in falling.

Twice I have taken eggs of Cuculus c. bakeri from hollow bam-

boos in which an Abrornis had built its nest and in neither of

these cases could the old Cuckoo have got more than its head

into the hole, whilst the young Cuckoo, if hatehed, could certainly

never have got out. Col. R. H. Rattray found an egg of C.c.

telephonus in Danga Gali, near Murree, placed in the nest of

Acanthopneuste occipitalis. This nest was in amongst the roots of

a tree and the Cuckoo had had a desperate struggle to get even

its head to the nest, losing many neck feathers in its final struggle

to overcome the last inch or two. Messrs. Inglis & Primrose

took several eggs of Chalcococcyx from the nests of Wthopyga

seherte and in no case were any of these tiny fragile nests

CUCKOOS’ EGGS AND EVOLUTION, 293

injured in any way, as they must have been had the Cuckoo tried

to force an entry into them. Yet again, Mr. W. E. Wait found

a young Surniculus in the nest of a Rhopocichla nigrifrons in

Ceylon, and this young bird, though not half-grown, completely

filled the nest, into which no full-grown bird could possibly

have crept.

I understand that Mr. Chance suggests that even when

Cuckoos do not enter nests for the purpose of laying their eggs,

they place themselves in such a position that they can shoot the

eggs by natural means into the nest direct from the oviduct but

this suggestion may be negatived from the fact that Cuckoos very

seldom smash the eggs of the tiny foster-parents with their own

heavy eggs, which would assuredly be the result if they adopted

such a method. Moreover, many nests are so placed that no

Cuckoo could obtain a foothold which would enable her to assume

a position suitable for such an action; whilst many other nests

are so fragile that they would never stand the weight of an adult

Cuckoo hanging on to them during her acrobatic performances.

I have dealt in this paper with only one aspect of Evolution

in regard to Cuckoos and their eggs but this subject opens out a

very wide field for observation. Thus certain recent theories of

‘‘age and area” are certainly not confirmed by a study of the

distribution of the Cuculide. Again, many of the tenets of

Mendelism and the heredity of acquired characteristics seem to

be contradicted by the evolution of Cuckoos’ eggs. But these

are big subjects and I merely refer to them here to show how

great an asset a wisely acquired collection of eggs may be in the

study of evolution and connected sciences.

EXPLANATION OF THE PLATES.

Prate I.

Fig. 1. Clamator jacobinus. An oviduct egg.

Ex nest of Turdoides terricolor.

3. 23 33

2. Turdoides terricolor.

eS Tite ews striatus, ¢ Common fosterers to Cjacobinus and H. varius.

6. Argya subrufa. J

7. Hierococcyx varius. An oviduct egg.

8. Clamator coromandus. Ex nest of Garrulax moniliger.

. Hierococcyx varius. Exnest of Argya caudata.

. Grammatoptila striata. Common fosterer in extreme N.W. Range.

ll. Garrulax monilig Cha The two most common fosterers to C. coromandus.

12 oe pectoralis.

Puate II,

. Corvus coronoides culminatus. (Siam.)

. Hudynamis scolopaceus. From nest of 1.

. Corvus splendens insolens. (Burma.)

EHudynamis scolopaceus. From nest of 2.

es ms From nest of 7.

. Corvus splendens splendens. (Behar, N. India.)

. Corvus splendens protegatus. (Ceylon.)

. Hudynamis scolopaceus. From nest ot 6.

. Corvus coronoides intermedius. (Assam.)

10. Hudynamis scolopaceus. Krom nest of 9.

Proc. Zoou. Soc.—1923, No, XX. 20

£00 > STOVE bo 0

294 CUCKOOS’ EGGS AND EVOLU'TION.

Pratve III.

Fig. 1. Cucaulus polioeephalus. White type from nest of 2,

2. Acanthopneuste occipitalis.

3. Cuculus poliocephalus. White type from nest of 4.

4, Phylloscopus affinis.

5. Cuculus poliocephalus. Red type from nest of Cettia cantans.

6. Cettia cantans. (Japan.)

7. Cuculus poliocephalus. From nest of Horornis fortipes.

8. Horornis fortipes. (Assam.)

9 to 11. Hierococcyx sparveroides. Brown type from nests of Arachnothera

magna.

12 to 15. Arachnothera magna. (Assam.)

16 to 18. Hierococeyx sparveroides. Blue type from nests of Garrulax

moniliger and Lanthocincla cineracea.

19. Ianthocincla cineracea. (Burma.)

Puate LV.

Figs. 1 to 4. Caccomantis merulinus. Blue types from nests of 5 to 9.

5, 6. Orthotomus sutorius.

7. Franklinia gracilis.

8, 9. Suya crinigera.

10 to 14. Caccomantis merulinus. White types from nests of 15 to 19.

15, 16. Orthotomus sutorius.

17, 18. Suya crinigera.

19. Cisticola cursitans.

20 to 23. } Caccomantis merulinus. Pink types ranging from the least to the

25 to 27. best developed type, all from nests of Prinia socialis.

24 & 28. Prinia socialis.

BUCCAL GLANDS OF THE OPISTHOGLYPHA, 295

21. A Comparative Study of the Buccal Glands and Teeth of

the Opisthoglypha, and a Discussion on the Evolution

of the Order from Aglypha*. By SusHin Cu. Sarkar,

E.Z.8.

[Received December 22, 1922: Read March 20, 1923.]

(Text-figures 1-29.)

Introduction.

The comparative anatomy of the buccal glands, teeth, and the

problem of the evolution of the opisthoglyphous Snakes forms the

subject of this paper, and was suggested to me by Professor G. E.

Nicholls, under whose supervision the investigation was begun

in the Biological Department of King’s College for Women,

Household and Social Science Department. After the research

had been in progress for three months Dr. Nicholls had to leave

for Australia, and since then the work has been carried on under

Miss Philippa C. Hsdaile, D.Sc., F.Z.S. It is with pleasure that I

acknowledge my indebtedness to Dr. Esdaile for her invaluable

advice and criticism during the progress of the work.

I wish also to express my obligation to Professor Dendy and

the Government Grant Committee of the Royal Society of

London fora grant-in-aid. Lastly, I have to thank Miss Alfreda

Newton for her kind assistance in cutting series of sections of

a large number of heads of snakes, and also for taking photo-

graphs of specimens and sections.

Historical Facts.

The snakes chosen as types in this research belong to the

Aglypha and Opisthoglypha, 7.e. two of the three divisions of

the Family Colubride. These differ from one another by the

fact that in the Aglypha all the teeth in the maxilla are solid,

whilst in the Opisthoglypha one or more grooved teeth are

present in the posterior region. The members of the latter

group are more or less poisonous.

The group Opisthoglypha is of special interest in that it is

considered to be the connecting-link between the Aglypha on the

one hand and the Viperide (with grooved teeth at the anterior

end of the jaw) on the other; the group Proteroglypha is

believed to have developed independently and directly from the

former. Although the possible evolution of the Opisthoglypha

was pointed out by Boulenger, no research to verify his

statements has been attempted in connection with the important

* Thesis submitted for Ph.D. degree, University of London, June 1922.

20*

296 MR, S. C. SARKAR ON THE BUCCAL GLANDS

organs, such as the poison gland, its duct and poison fangs, taken

concurrently, So far as Iam aware, this is the first research on

these lines.

The presence of grooved teeth in the posterior region of the

maxilla was first observed and discussed by Thomas Smith (11)

and later by J. G. Fischer. In the year 1892, Niemann (6)

published an account of the structure and relationships of the

glands of the upper lip in general, but he described only three

types of the Opisthoglypha. His work on this group is very

scanty, while the diagrams are insuflicient and wanting in detail,

but the connection of the duct and the groove of the fang is

well emphasized. In 1895, West (15) published a detailed

description of the buccal glands and teeth of opisthoglyphous

snakes. His paper made a_ considerable advance in the

knowledge of this subject; his observations are still looked upon

as.a good groundwork, and his work is quoted by various authors.

His descriptions, however, relate only to morphology. I have

described four more genera of this group—Chrysopelea, Larbophis,

Psammophis, and Cerderus ; the first two were not mentioned at

all by West, and of the other two he gave only ashort description

of the teeth.

Tt was in 1896 that -Boulenger put forward an account of the

probable evolution of the Opisthoglypha from the Aglypha. He

pointed out that in the higher genera of the Aglypha the series

of teeth in the jaw shows an increase in size from in front

backwards, so that we are gradually introdueed to the opistho-

glyphous condition, with a large fang in the posterior region of

the jaw. In the higher types of both the Opisthoglypha and

Proteroglypha this fang is always distinctly bigger than the

tooth in front of it. Therefore in my discussions of the

probable evolution of the fang of the Opisthoglypha from the

somewhat enlarged posterior tooth of the Aglypha, I have

selected those snakes in the Aglypha in which the posterior

tooth is clearly larger than the preceding one—e. g., Z'repidonotus

and Lycodon.

Among recent papers on the Ophidia, those of Mary Phisalix

are the most important. In her work (9) she discusses the

accounts given by various authors, and she has attempted to

arrange the members of this group in an ascending series on

evolutionary lines, according to the disposition of the teeth. In

this paper I have attempted a like task, but my series differs

from hers in that I have tried to select for consideration only

those Aglyphodont forms in which the last tooth has become

markedly different from the rest. Also I have discussed what

IT consider to be the highest grade of evolution in the Aglypha—

i.e., the condition presented by those types in which the last

tooth has acquired two cutting-edges instead of only one. Mary

Phisalix does not continue her series beyond those with one

cutting-edge. It should be noticed that my diagram of Zropido-

notus stolatus (text-fig. 10) differs from hers (see p. 164, ref. 9),

AND TEETH OF THE OPISTHOGLYPHA, 297

The hinder tooth in my diagram of this species is represented as

larger than the one in front of it. My observation is gathered

from the dissection of several specimens.

Methods and Material.

Most of the snakes used for my observation were brought from

India by myself. Some of them were preserved specimens, well

fixed and suitable for histological purposes.

Unfortunately others which I had hoped to keep alive for

physiological experiments had to be killed unexpectedly, and the

tissues were only imperfectly fixed,

This investigation has necessitated the preparation and exami-

nation of a number of serial sections of the heads of snakes, and

owing to the presence of the scales and hard bones forming the

skull, this proved to be a difficult task, After many experiments

the following method was successful.

The head was separated from the body close to the angle of

the jaw, and placed in the fixing and decalcifying reagent aceto-

bichromate (Bolles Lee, pp. 49 & 50). In order to ensure

thorough penetration of the fluid, an incision in the scales was

made just behind the eye, the lens of which was also removed.

After a fortnight the head was taken out, and was carefully

cut by means of a sharp razor into two halves by a median

longitudinal vertical cut. Hach half was again placed in fresh

aceto-bichromate solution with the addition of one or two drops

per 100 cc. of acetic acid. I found it necessary to leave it in this

for three to four wéeks to get complete decalcification. Each of

the halves was then used for longitudinal or transverse sections,

It was found necessary to leave the head in the paraftin bath at a

low temperature for more than twelve hours. The sections were

cut at a thickness of 10-15. Several stains were tried, and

I found the following most useful: Ehrlich hematoxylin

counter stained with eosin or hemalum and eosin. Hach of

these methods gave a good differentiation between the serous

and the mucous secreting cells in the poison gland, the former

being stained pinkish, the latter bluish. Both these being water

stains, I found that the teeth and bone were often washed off

the slide while being taken through the alcohol. To prevent

this the slides were dipped in a thin solution of colloidin in 90 °/.

alcohol. In order to verify my observations with regard to the

duct and its connections with the fang, I made several

cardboard models of portions of the jaw.

General Discussion on the Teeth in the Mandible and Maxilla

and the Mechanism of the Hinged Teeth.

The maxilla, mandible, and pterygoid bear teeth and these occur

in uninterrupted series, increasing or decreasing in size from

in front backwards. Sometimes a diastema is left, and this is

298 MR. S. C. SARKAR ON THE BUCCAL GLANDS

generally found in front of the tooth, which becomes bigger than

the rest (Dryophis). The maxillary and mandibular teeth are

mostly inclined posteriorly at a considerable angle—i.e., the

proximal part of each tooth for a little distance is almost

at right angles to the bone, while the distal part bends back-

wards. The teeth appear to be of various types when seen

in transverse section—round, triangular, flattened, etc. But more

interesting still is the fact that the alternate teeth are hinged

and fixed. In the dried specimens the hinged teeth frequently

drop out; the dropping-out of the teeth after death has been

frequently noticed by several workers, including Boulenger, and

was accounted for by the accidents of preparation.

Thus, quoting Boulenger :—

“‘Tt very often happens that every alternate tooth having

dropped out, the jaw appears, ona superficial examination, to

possess half the real number of teeth.”

At first, while making dissections of a large number of snakes,

I often found that some of their teeth differed from the others

Text-figure 1.

eee Uibtes

mand.

Tropidonotus stolatus. Portion of mandible seen in lateral view, drawn with

camera lucida froma cleaned specimen. ft. fixed tooth; h.¢. hinged tooth ;

L. ligament; mand. mandible. X 46,

in that they could be bent backwards. I took it to be that

these teeth were being either gradually fixed or were about to

drop out of the jaw. On a more careful examination, I found

that these loose teeth, as a general rule, alternate with the fixed

ones. This regularity led me to suspect that the condition was

not accidental. In order to verify this, I examined a number of

cleaned skulls, and found that these loose teeth either drop

out, leaving spaces or remain somehow fixed to the jaw, so that it

is difficult to distinguish them from the fixed ones. I concluded

from this that there must be some ligament or structure which

AND TEETH OF THE OPISTHOGLYPHA, 299

keeps the tooth attached to the bone, but this hardens when

dried, giving the tooth an appearance of being fixed. In order

to make a further examination, I mounted the decalcified jaw

complete, carefully dissected out from the head of the snake.

In text-fig. 1 it will be seen that there are three fixed teeth,

while the other two are attached to the socket by means of

connective tissue (J), and there is a space, one tooth having

dropped out during the process of dissection. I also cut sections

of the heads of different genera, taking care to cut the jaw as

nearly longitudinally as possible. Externally these hinged

teeth differ from the others by being bent more sharply, and also,

when moved with a needle, they can be made to bend towards

the median line. As regards the forward movement, it is

limited to a certain extent. At the base also the two sets of

teeth differ from one another by the fact that the hinged teeth

have a whitish appearance, due to the presence of the connective

tissue which holds it to the bone. In order to study the

Text-figure 2.

Tropidonotus stolatus. Longitudinal section of mandible with teeth. X 92.

mechanism of its movement, I carefully examined the series of

sections of the tooth and the bone. ‘Text-fig. 3 shows a diagram

drawn from a longitudinal section taken at about the middle of

the tooth. It will be noticed that the facets in the maxilla at

the base of the two ends of the tooth differ in their structure,

and, at the same time, the base of the tooth differs markedly at

the anterior and posterior ends. In the front the base ends in

a peculiar, slightly knob-like structure which also projects

inwards. This fact was also noticed by Tomes (13), who writes

as follows :—

“As the tooth approaches completion, there is a

peculiarity in the form which its base assumes and which I

have not noticed in other animals, namely that the dentine

at the widely open base of the tooth is often abruptly bent

inwards as though the base of the tooth were about to be

closed by a sort of operculum of dentine.”

300 MR. Se Ce SARKAR ON THE BUCCAL GLANDS

In his plate Tomes has given figures of sections of four or

five teeth, and he has shown the bending in one only, and this

also at the anterior side. ‘This evidently proves that this was

one of those hinged teeth.

This knob-like structure at the anterior edge of the tooth fits

into the slightly concave, vertically inclined, smooth facet of the

bone, so that when the tooth is pressed forward it strikes against

the slightly inclined plane and is prevented from further

bending. At the posterior side the distance between the bone

and the end of the tooth is greater. The connective tissue

between these two structures is more prominent, and its

histological structure can be examined. Under the high power it

has the appearance as shown in text-figs. 3 and 4, and it resembles

white fibrous tissue. It has not at all the appearance of develop-

ing bone or enamel. Had it been enamel it would have been

dissolved by the continued action of the acid in the decalcifying

Text-figure 3.

Tyropidonotus stolatus. Longitudinal section of maxilla, showing a loose tooth

with its connective-tissue attachment. J. ligament; max. maxilla; p.c.

pulp cavity. X 260.

solution, to which it was subjected for more than a month.

Text-fig. 4 shows also the connection of the same tooth at the

posterior edge. It will be noticed that the connective tissue (¢)

keeps it attached to the bone. Thus we see that the tooth when

pressed from in front moves so that the hinder end of the

base, which has a greater field in which to move, is pressed down

on to the bone; but it is prevented from slipping further by

the connective tissue; while, on the other hand, if the tooth

is pressed forwards from behind, its power of movement is

restricted by the inclined edge of the socket of the bone and

also by the band of connective tissue at the hinder end, which,

so far as can be ascertained at present, appears to be non-

elastic. This hinged and fixed arrangement I have found to be

extended to the grooved teeth. If there are two grooved teeth,

one of them is hinged and the other fixed. It depends upon the

arrangement of the other teeth whether the anterior or the

AND TEETH OF THE OPISTHOGLYPHA. 301]

posterior one is hinged. It often happens that either the two

or three consecutive teeth drop out from the dried bones, This

is due to the fact that one of the fixed teeth is being replaced.

The function of such an arrangement is difficult to determine,

but it seems to provide a firmer grip for the prey. While the

prey is being taken into the mouth and forced towards the

gullet, these hinged teeth, which point inward and backward,

bend down; so that when the prey is struggling to escape from

the mouth, the teeth, being hinged, allow it to pass downwards

into the gullet but not outwards. Therefore from these

observations I am led to conclude that the hinged teeth are not

part of a series in the course of development which will become

ankylosed to the jaw, but that they will always remain hinged,

and during life will be held in place by the band of connective

tissue, while after death they may drop out. This opinion is

further emphasized by the facts that: firstly, in the maxille and

Text-figure 4.

Tropidonotus stolatus. Longitudinal section of maxilla, showing a loose tooth with

its connective-tissue attachment. 1. ligament; m. maxilla; ¢. tooth. > 260.

mandibles of the dried skull of the adult we can clearly see the

plane facets at the anterior end of the sockets, which show that

it has been subjected to constant friction ; secondly, if the tooth

were to be later ankylosed to the jaw-bone, there would be no

necessity for the inbending of the anterior end of its base;

thirdly, it is inconceivable that in the numerous cases I have

examined, every second tooth should become worn out or drop

out accidentally.

The grooved fangs are always placed at the posterior end of

the maxilla. Sometimes they are situated far back in the mouth,

so that their points are quite hidden within the angle of the jaw.

In this case the teeth are considerably bent (Dipsas), but where

they are a little more forward in the mouth they are generally

straighter. In shape they may be either flattened (Oxybelis) or

circular (Psammophis), or may have a cutting-edge developed at

the posterior region. The grooves are either placed in the

302 MR. S. C. SARKAR ON THE BUCCAL GLANDS

front face of the tooth or on the external surface. In some cases

the groove slightly curves round in its course; it may extend

throughout the length of the tooth or may end a little above the

tip. The groove may be shallow, or may be widely open or

almost closed.

Morphological Description of the Dissection of the Heads

of a few Opisthoglyphous Snakes.

Genus Dryopuis.

DRYOPHIS MYCTERIZANS. (Text-fig. 19.)

The poison gland is distinctly definable from the superior

labial gland, and is large and extends a little further forward

than the middle of the eye and posteriorly a little behind the

angle of the jaw. The superior labial gland reaches to the

anterior extremity of the maxilla, and it extends a little behind

the poison gland. The two portions of the gland are continuous

with one another. Near the region of the fang-like tooth in

front the gland becomes broader. The inferior labial gland

consists of a very thin strip of glandular tissue, and does not

extend as far as the angle of the jaw. There are fourteen teeth

altogether in the maxilla, and they are arranged as follows :—

four small teeth in front, then two large fang-like teeth, followed

by a diastema, then six very small teeth, and finally two very

large, stout, and straight grooved teeth. The groove is on the

outer side and is deep and extends throughout the length of

the tooth. The duct of the poison gland opens near the base of

the tooth. There are eighteen teeth in the mandible. The

first is rather small, the next three very large, the middle of the

three being the largest, then a diastema, which is followed by

fourteen teeth which gradually diminish in size. The Harderian

gland is much reduced, and consists of a thin strip of tissue lying

behind the eyes in the eye-socket.

Genus CERBERUS.

CERBERUS RHYNCHOPS.

The poison gland is clearly distinguished from the upper

labial gland, and is large and oval (text-fig. 5); itis so distinct an

organ that it can be easily picked up from the surface of the

superior Jabial gland without injury to the latter. The superior

labial gland extends from the anterior end of the maxilla and

posteriorly up to the poison gland. Practically the whole of the

posterior portion of the glandular area is occupied by the poison

gland, so that the superior labial gland does not extend far

behind the eye. The lower labial gland also does not extend up

to the angle of the jaw; this is probably due to the sharp

upward turning which the lower jaw takes, thus giving a greater

AND TEETH OF THE OPISTHOGLYPHA, 303

extension to the mouth and placing the fangs in a more exposed

position. There are eighteen maxillary teetl:; the last two are

grooved and large. The grooved teeth are placed slightly

further forward away from the angle of the jaw. The hinder

grooved tooth is hinged. The groove is situated on the anterior

side of the tooth at its proximal end, while on the outer side at

the distal end. The groove does not extend up to the very end

Text-figure 5.

Cerberus rhynchops. Dissection of head from the right side. f. fang; 7.1.9.

inferior labial gland; p.g. parotid gland; s.J.g. superior labial gland. 3.

of the tooth, it is deep and almost takes the shape of a canal.

The mandible contains twenty-two teeth of unequal size.

The Harderian gland is a little peculiar, for the greater part

of the outer portion lies under the poison gland. There are

three lobes; the external part is cylindrical, tapering at the

posterior end, while the other two within the orbit divide into

dorsal and ventral portions surrounding the optic nerve.

Genus DrIpsas.

DIPSAS TRIGONATA.

The poison gland is clearly distinguishable from the superior

labial gland, and anteriorly extends up to about the middle of

the eye. The lobules of this gland are smaller than those of the

superior labial gland and more distinct. The latter extends to

the very end of the mouth anteriorly, and ends posteriorly about

the angle of the jaw. It is continuous all along. The inferior

labial gland is more highly developed than the superior labial

gland, extending far back so that it almost meets the superior

labial gland behind the angle of the jaw. The maxilla con-

tains twelve teeth; the last two are grooved. The teeth are

larger in front except the first one. The poison fangs are bent

considerably backwards and slightly inwards so that the pointed

ends are directed towards the pharynx. At the proximal end

the grooves of the teeth lie on the outer side, while at the distal

end they are in front of it, The grooves extend three-fourths

304 MR. S. C. SARKAR ON THE BUCCAL GLANDS

the length of the tooth. The alternate teeth are hinged and

fixed. The mandible contains about thirteen teeth; they are

also larger in front, gradually diminishing behind. The

Text-figure 6.

Dipsas trigonata. Dissection of head from the right side. f. fang; @./.g. inferior

labial gland; .g. parotid gland; s.J.g. superior labial gland. X 3.

Harderian gland consists of three lobes, two within the orbit

and the larger part outside.

Genus CHRYSOPELEA.

CHRYSOPELEA ORNATA. (Text-fig. 17.)

The poison gland is very small, smaller than in any opistho-

elyphous snake I have examined, and it occupies a small area

posterior to the eye. The smallness of the gland may be due to

the huge growth of the eye. The superior labial gland is well

developed, and extends posteriorly further than the angle of the

jaw and anteriorly to the very end of the maxilla. The lobules

of this gland are larger than those of the poison gland. The

inferior labial gland is also fully developed, and is broader at the

posterior side. The maxilla contains twenty teeth; the first

three are very small, and they increase in size posteriorly; the

last two are grooved. The groove is situated on the outer side

and is widely open. It arises almost from the base of the tooth,

and ends about one-eighth of the length of the tooth from the

apex. In section the teeth are nearly oval in shape. The

mandible contains twenty-two teeth: the first is very small;

there is an increase in size up to the seventh, which is the

largest, and thena gradual decrease. Both maxilla and mandible

show the alternately fixed and hinged arrangement of teeth.

AND TEETH OF THE OPISTHOGLYPHA. 305

The Harderian gland consists of three lobes; two portions lie

within the orbit and one outside.

Genus TARBOPHIS.

TARBOPHIS VARIEGATUS,

The specimen of this snake was in a damaged condition.

The poison gland is large, and quite distinct from the superior

labial gland; it is spindle-shaped, the anterior end lying below

the middle of the eye. The superior labial gland is narrow, and

reaches the anterior end of the maxilla, where it enlarges and

curves round to join the gland of the other side. It is also

continuous below the poison gland. The inferior labial gland

extends from the very end of the mandible in front, and meets

Text-figure 7.

> oan sor

ie CLG.

Tarbophis variegatus. Dissection of head from the right side. f- fang ; i.l.g. 4

inferior labial gland; p.g. parotid gland; s.l,g. superior labial gland, X 3.

the superior labial gland posteriorly behind the angle of the jaw.

Of the nine teeth in the maxilla the last is large and grooved,

and the rest are smaller and subequal in size. The grooved

tooth is situated far behind within the buccal cavity. The

groove is placed on the external surface of the tooth, and does

not extend up to the end of the tip. There are about fourteen

teeth in the mandible, the series decreasing in size posteriorly.

They are alternately hinged and fixed. The Harderian gland

consists of two lobes, one outside and the other within the orbit,

and it is perforated by the optic nerve.

Genus PsAMMOPHIS.

PsAMMOPHIS SIBILANS. (Text-fig. 21.)

The poison gland of this specimen shows the most highly-

developed condition among the opisthoglyphous snakes that I

have examined; it is a distinct organ, and has assumed a definite

shape, being oval. It is comparatively very large. The lobules

of the gland are much smaller than those of the superior labial

306 MR. S. GC, SAKKAR ON THE BUCCAL GLANDS

gland. The latter is continuous below the poison gland, and

extends anteriorly to the very end of the snout, meeting its

fellow of the other side. Behind it extends further back than

the poison gland. The-inferior labial gland is also well developed,

and has the appearance of a compact body reaching as far as the

end of the mandible, and posteriorly it meets the superior labial

gland behind the angle of the jaw. The maxilla possesses four

medium-sized teeth in front, then two large, solid fang-like teeth,

followed by a series of four small equal teeth and finally two large

straight fangs. The groove of the posterior maxillary teeth is

placed on the anterior face of the tooth, and extends throughout

the length of it, ending at the tip. The teeth in the maxilla and

the mandible are straight and slightly directed backwards, and

they also show an alternating arrangement of hinged and fixed

teeth. The Harderian gland consists of two lobes, one outside

and the other within the orbit. The latter shows a superficial

division into two.

Discussion on the Evolution of the Opisthoglypha.

It is generally admitted that the opisthoglyphous snakes are

evolved from the Glauconiide, and that the immediate ancestors

of the Opisthoglypha are the Colubride Aglypha. Boulenger

pointed out in the year 1896 (1) “that from Aglyphodont

forms, in which the teeth increase in size posteriorly, we are

gradually led to the Opisthoglypha, which are only to be distin-

guished by the presence of more or less deep grooves on the

posterior fang-like teeth.”

As far as my own observations go, I find that, apart from the

gradual increase in the size of the teeth, there is a change

concurrently in the structure of the particular tooth which is

situated in the posterior portion of the maxilla near the opening

of the duct of the parotid gland into the mouth. At the same

time, the parotid gland itself undergoes a gradual transformation

from a simple structure, distinguishable only from the superior

labial gland by its colour, to a separate distinct organ lying

detached from it. There. also occurs a change in the connection

of the duct of the parotid giand with the buccal cavity and the

teeth.

My observations are based on a study of the following types,

arranged in order :—

Dendrophis pictus.

Tropidonotus stolatus. _Aglypha.

Lycodon aulicus.

Oxybelis fulgida. >

Chrysopelea ornata. ee i

Dryophis mycterizans. Op isthoglypha.

Psammophis sibilans. _)

AND TEETH OF THE OPISTHOGLYPHA. 307

I had hoped to be able to obtain several specimens of both

Xenodon and Dispholidus, in order that a full study might be

‘made of the characters of the “poison apparatus” in these:

‘peculiar types. But in this I have been disappointed. I have

‘made some dissections, but it was impossible to obtain a good set

‘of serial sections. So I am unable to determine what further

questions might be raised by such a study, or how a complete

knowledge of the conditions exhibited by Xenodon and Dispho-

‘lidus might affect the general lines of my arguments. However,

I hope at some future date to work on these two genera,

comparing and contrasting them with the genera I have had the

opportunity of studying.

Aglypha.

DENDROPHIS PICTUS.

The anatomy of the general dissection of the glands and teeth

of Dendrophis is shown in text-fig. 8. Of the two glands in

the maxillary region the superior labial gland occupies the greater

area, while the so-called parotid gland is confined toa small space.

As will be seen, they are so intimately related that it would be

difficult to distinguish one from the other but for the colour.

This fact struck earlier workers too. Leydig (1873), in his work

: Text-figure 8.

STEER, Sg

x 50 LS aoe Le

RE re Nm an

Peas

Parsee SR SEA SEAN

ey Zee Pe) Pewee Syne

pe ee DY

P9. 6lg.

Dendrophis pictus. Dissection of head from the right side. Letters as before. X:3.

on Tropidonotus, described that the ‘superior maxillary gland

divides itself into two portions which are distinct in the form and

the colour of their follicles; the inferior part is grey and the

principal part is of a yellowish colour, and the follicles are larger

than on the grey part.”

Evidently this yellowish portion becomes specialised to secrete

a fluid different from that secreted by the rest of the gland ; and

we shall see later that this fluid becomes more and more im-

portant in the higher snakes as a weapon of defence, and that

the teeth become modified to allow of its more careful use.

308 MR. S. C. SARKAR ON THE BUCCAL GLANDS

Tn the transverse section of Dendrophis (text-fig. 9) the superior

labial and parotid glands and the ducts arising from them are

figured. On comparing these ducts, it will be seen that the duct

of the superior labial gland opens into the mouth at some distance

from the round. peg-like tooth and has no connection with it,

while the duct of the parotid gland, formerly forming one of the

series of the salivary ducts, has shifted its position in order that

it may open nearer to the hinder tooth. In the maxilla there are

about twenty teeth, small and almost equal in size; but the

Text-figure 9.

Dendrophis pictus. Transverse section of a portion of the head in the region of

the superior labial and parotid glands, showing their ducts. Letters as

before. XX 63.

hinder tooth, referred to above, is bent backwards at a greater

angle than the rest, also a slight cutting-edge has developed on

the posterior face.

Asin all other harmless Batrachians and Reptiles, the teeth are

used to prevent the prey from slipping out of the mouth, and at

the same time to make small punctures in the body while it is

being killed by suffocation. Meanwhile the buccal cavity is

flooded with the secretion from all the buccal glands, and the

mixed saliva covers the animal and gradually finds its way into

the small wounds made by the primitive round teeth,

TROPIDONOTUS STOLATUS.

Text-fig. 10 shows the external features of the gland as seen

after the removal of the skin, and also the maxillary teeth from

the left side. It reveals a condition strangely advanced in some

features, but on the whole distinctly similar to that of Dendrophis

AND TEETH OF THE OPISTHOGLYPHA,. 309

(text-fig. 8). Of the more highly-developed parts the parotid

gland and the posterior tooth are most striking. The gland (p.g.)

is quite well developed, and it extends far behind, as far as the

angle of the jaw, and in front up to the middle of the eye, while

in shape it is peculiar in being rectangular; in fact, as will be

Text-figure 10.

Filigc Ws

Tropidonotus stolatus. Dissection of head from the left side.

Letters as before. X 6.

seen from the diagram (text-fig. 10), more than half the glandular

area in the region of the maxilla has taken up the function of the

secretion of the serous fluid. Apart from its great development

in size, the dissection shows that the parotid gland is really a part

of the superior labial gland, and is only to be distinguished from

it by its yellow colour.

Text-figure 11.

Tropidonotus stolatus. Longitudinal section of the head in the region of the

poison fang and duct of parotid gland. f. fang; duct p.g. duct of poison

or parotid gland; op.t.s. opening of duct into tooth sac; 7.t. reserve tooth ;

ves. vestibule. 63.

The maxilla bears about eighteen teeth, these being subequal

in size, except the last. This (/) is much increased in size, and

is more than twice the length of the tooth in front of it. It is

Proc. Zoou. Soc.—1923, No. X XI. 21

310 MR..§. C. SARKAR ON THE BUCCAL GLANDS

considerably bent backwards. But apart from its being distin-

guished from the others by its size, it is important to notice that

it is separated from them by being enclosed in a special sac

(text-fig. 12, ts.) formed from the folding of the mucous

membrane; in fact, it has all the external appearance of a fang

contained in a tooth sac, and the similarity is so great that it can

readily be mistaken for the fang of the poisonous snakes. Further,

we can see from the portion of the longitudinal section of the

head of Tropidonotus that the tooth is specialised by the growth

of a cutting-edge on the posterior side. Besides, as in other

poisonous snakes, there are reserve teeth (7.t.) lying on the inner

side of this fang-like tooth. Text-fig. 11 shows the connection

of the duct of the parotid gland with tooth and mouth-cavity. It

Text-figures 12 & 13.

Fig. 12.—Tyropidonotus stolatus. Tooth sac and fang. ¢.s. tooth sac; f. fang.

Fig. 13.—Tropidonotus stolatus. Longitudinal section of head in the region of

the duct of the parotid gland, showing its opening into the buccal cavity.

op.t.s. opening of duct into tooth sac; op.d.p.g. opening of duct of parotid

gland. X 63.

is remarkable that the duct of this gland enlarges at first into a

special sac, the vestibule (ves.), which is in communication with

the tooth sac by an opening (op.t.s.). The vestibule opens into

the buccal cavity (text-fig. 13, op.d.p.g.). Hence we see that

the duct, before opening into the buccal cavity, opens into the

vestibule, which becomes confluent with the lower part of the

tooth sac. By such an arrangement it is made possible for the

snake to inject a part of the secretion from the parotid gland into

the severe wound at the time when it is being made by means of

the sharp cutting-edge of the tooth.

LYCODON AULICUS.

' Text-fig. 14 illustrates the general anatomy of the head dissected

from the right side. The general survey will show that it is

more advanced, and of a higher order than Tropidonotus in regard

AND TEETH OF THE OPISTHOGLYPHA. 311

to its teeth and glands. The parotid gland (p.g.) in this snake

is mueh enlarged, and its extension forward goes much further

than in Tropidonotus, almost to the anterior end of the maxilla.

Behind, it extends further than the angle of the jaw, and ends

almost on a level with the superior labial gland; in fact, it has

taken up so much of the glandular area that the superior labial

gland is left reduced to a mere strip of glandular mass, which in

dissection has an appearance more or less like that of the poisonous

snakes. The lobules of the gland are larger than those of the

superior labial gland. ‘This is the only case I have yet met with

in aglyphous or opisthoglyphous snakes where the poison gland

is so highly developed and extends further forward than the

middle of the eye. The most interesting feature to notice is the

point of termination of the gland in front; it gradually narrows,

and ends with a little bend above the enlarged fang-like tooth,

apparently giving an impression in dissection of the duct of

Text-figure 14.

Lycodon aulicus. Dissection of head from the right side. Lettersas before. X 6.

proteroglyphous snakes opening at the base of the fang. I

hope at some future time, when more material is available, to

make a more complete examination of the anterior end of the

jaw of Lycodon, in order to put to the test Boulenger’s sugges-

tion of the probable evolution of the Proteroglypha from the

Aglypha. :

There are eighteen maxillary teeth, the anterior three are large

and are followed by a diastema, the next fourteen are very small,

and the last two are large and fang-like, and are enclosed within

a muscular tooth sac similar to that already noted in Z’ropido-

notus. In the transverse section (text-fig. 15) of the head of

Lycodon will be seen the section of the posterior tooth sac

(f. & t.s.). It will be noticed that the tooth is further spe-

cialised than that of Tropidonotus by the growth of cutting-edges

on both sides. These cutting-edges give the appearance of a

lancet.

21*

312 MR. S. C. SARKAR ON THE BUCCAL GLANDS

The duct of the poison gland, like that of Tropidonotus,

opens into the vestibule which communicates with the tooth sac

at one end, and at the lower end. opens into the buccal cavity

(op.d.p.g.). A comparison of the course of this duct in

Tropidonotus and Lycodon reveals the fact that in Lycodon the

communication of the vestibule with the tooth sac is higher than

in Tropidonotus, so that the secretion of the gland first fills the

Text-figure 15.

Lycodon aulicus. Transverse section of head in the region of the fang, duct of the

poison gland and its opening into the tooth sac and the buccal cavity. ves.

vestibule; other letters as betore. X 63.

cavity of the tooth sac while on its course downwards into the

buceal cavity. By such an arrangement of the duct the tooth-

cavity is kept filled with the poison, and the tooth bathed in it.

In this way a greater amount of poisonous fluid can be injected

into the bigger wound, which is made by the two cutting-edges of

the lancet-shaped tooth.

Opisthogylpha.

OXYBELIS FULGIDA.

This is a very interesting snake because of the fact that

although it has acquired a better and surer means of injecting

poison into the wound by developing a groove in the fang, it still

retains the ancestral method of filling the tooth sac with the poison.

West has described the anatomy of the head in detail, but for

the present purpose of comparison his figures are insufficient.

T have been unable to procure any specimens of Oxybelis to verify

West’s observation, and therefore cannot provide the necessary

AND TEETH OF THE OPISTHOGLYPHA. 313

diagrams. So I have to base my arguments on West’s account

in the Proceedings of the Zoological Society, 1895. According to

West, ‘the poison gland is very clearly defined and the superior

labial gland is exceedingly long and narrow; it reaches to the

anterior extremity of the maxilla, and consists of much smaller

lobules than the poison gland. .. The maxilla possesses more

teeth than that of any other snakes examined, there being twenty

in one uninterrupted series. The first seventeen are equal in size

and much curved; the posterior three are a little larger, com-

pressed laterally, and the external face of each possesses a very

shallow groove. The muscular folds surrounding these three

posterior teeth are not united in front, and in consequence of a

thin muscular fold across the base of the anterior grooved tooth

the poison duct in this snake is placed in communication with the

interior of the mouth before it is with the groove of the tooth.”

(See text-fig. 16.)

Text-figure 16.

Oxybelis fulgida. Transverse section of head in the region of the fang and the

duct of the poison gland opening into the mouth (after West). d.p.g.

duct of poison or parotid gland; f. fang; g. groove; op.d.p.g. opening of

the duct of the parotid gland; #.s. tooth sac.

West has not given any figure of the dissection of the head, nor

any section of the tooth (fang), but from the above description we

may surmise from his use of the words “ very clearly ” that the

parotid gland is even more highly differentiated in Oxybelis than

in Lycodon. With regard to the fang, we may safely conclude

from the above account of its lateral compression that it still

retains some indication of cutting-edges on both sides like those

of Lycodon, and at the same time has developed in “ the external

face . . . a very shallow groove.” Referring now to the tooth-

sac and the duct, I think that the thin muscular fold which West

describes is the wall of what I have called the vestibule. Text-

fig. 16 shows its connection with the buccal cavity, and probably

if another series of sections were examined we should find the

‘openings of the vestibule into the tooth sac. We can infer from

314 MR. S. GC. SARKAR ON THE BUCCAL GLANDS

this that the arrangement of the duct, vestibule, and tooth sac is

more or less like that of Zycodon, and this is emphasized in his

figure, which I reproduce.

Oxybelis, therefore, seems to be one of those types which

connect the Aglypha with the Opisthoglypha. On the one hand

it retains the aglyphous condition of the duct opening into the

mouth, and on the other it has the characteristically opisthogly-

phous grooved teeth.

CHRYSOPELEA ORNATA,

The general description of the glands and teeth has been given

before (p. 304), but for comparison and discussion we will

mention some of the facts again. On comparing the external

appearance of the poison gland of Chrysopelea (text-fig. 17)

with that of Ozxybelis, described by West, and also with that

of Lycodon, one is struck with the relative reduction in size; in

fact, the gland is restricted to a very small area, and in shape and

size resembles that of Dendrophis, which stands lowest in my

series in the Aglypha (p. 307). In other words, it seems to be

a case of retrogressive rather than of progressive evolution. But,

Text-figure 17,

Chrysopelea ornata. Dissection of head from the left side. Hg. Harderian

gland; 7./.g. inferior labial gland; p.g. parotid gland; s.l.g. superior labial

gland. X23.

on the other hand, it must be remembered that the snake has

acquired more perfection by developing a deeper groove, by

means of which it is confident of injecting a sufficient quantity of

the poison into the wound, and that therefore it is obviously

unnecessary to have a large gland and huge amount of secretion.

It also will be seen that the secretion coming from the gland

passes directly into the tooth sac, where greater use can be made

of it than if it were squandered in the buccal cavity.

Now, if we examine the transverse section of Chrysopelea (text-

fig. 18) and compare it with that of Oxybelis (text-fig. 16), we

find the former specimen is more advanced in many ways. The

fang has lost the remains of the cutting-edges on both sides, but

has developed a deeper groove than that in the fang of Oxybelis.

AND TEETH OF THE OPISTHOGLYPHA. 315

Regarding the duct, it no longer opens into the vestibule, but

directly into the tooth sac. The opening of the vestibule into the

mouth cavity is closed, and the tooth sac becomes comparatively

bigger. If in text-fig. 16 the opening of the vestibule into the

mouth is closed, and at the same time the thin membrane, to

which West refers and which is also shown in text-fig. 15,

atrophies so that the cavities of the vestibule and tooth sac are

confluent with one another, the condition, as shown in text-

fig. 18 of Chrysopelea, is arrived at. I have shown that the two

Text-figure 18.

Pg duct. pg. aa

Chrysopelea ornata. Transverse section of the head in the region of the poison

gland, its ducts, the tooth sac, and the fang. 6.v. blood-vessel; duct.p.g.

duct of the poison gland; f. fang; p.g. poison or parotid gland; 7.¢. reserve

tooth; ¢.s. tooth sac. X 63.

cavities really become confluent in 7V’ropidonotus and Lycodon

near to the opening into the mouth. The evidence in favour of

the theory that such changes have been brought about is shown

by the fact that the duct in Chrysopelea opens, not into the

mouth, but into the middle of the tooth sae.

In conclusion, then, we see that Chrysopelea is higher in grade

than Oxybelis, for the reason that the duct of the poison gland

opens directly into the tooth sac and that there isa deeper groove

in the fang.

DRYOPHIS MYCTERIZANS.

This snake is in many respects much more advanced than

Chrysopelea. The parotid gland is more highly developed, and

has become a distinct organ, though it still remains embedded in

the superior labial gland. At the same time, the secretion of the

gland becomes more poisonous; it has been proved that the bite

of this snake is fatal to small animals (West). The fangs (text-

fig. 19) have shifted a little forward in the jaw, and are therefore

316 MR. S. C. SARKAR ON THE BUCCAL GLANDS

in a less concealed position, being rendered capable of inflicting a

more severe wound. There are only fourteen teeth in the

maxilla: this reduction may possibly be explained by the fact

that the teeth, which are present, are large, strong, and more

exposed, and better able to kill the prey or to paralyse it quickly

with the poison that is now injected directly into the puncture

made by the fang.

Text-figure 19.

aa = oi

Dryophis mycterizans. Dissection of the head from the left side. x 3.

Letters as before.

In the transverse section (text-fig. 20) we notice that the duct

of the poison gland has shifted still further up than in Chryso-

pelea (text-fig. 18), so that its opening into the tooth sac is nearer

Text-figure 20.

Dryophis mycterizans. Transverse section of the head in the region of the parotid

and superior labial gland and their ducts, the tooth sac and the fang. m.m.

mucous membrane; other letters as before. X 63.

to the base of the tooth and close to the beginning of the groove.

The poison coming from the gland passes directly into the groove.

In Dryophis the groove is now more or less in the shape of a

canal, as the edges are almost in contact.

AND TEETH OF THE OPISTHOGLYPHA. ld

PSAMMMOPHIS SIBILANS.

This is one of the most highly-developed of the opisthoglyphous

snakes. The very appearance of text-fig. 21 will reveal the

fact that it is more ferocious and is capable of doing more harm

than Dryophis. The structure of the teeth and gland positively

prove that it is so, and this is emphasized in the description in

the first part of this paper.

Looking at the poison gland, one finds that it has assumed a

distinct and definite shape, quite separate from the superior labial

gland. The poison fang has moved still further forward in the

buccal cavity than in Dryophis. It is straighter, and it lies

directly below the eye. The shape of the poison fang is of

importance, 7.e. being straight and not bent backward. The

fang that is bent backward at an angle is of more use in inflicting

Text-figure 21.

Psammophis sibilans. Dissection of head from the right side. f. fang; Hig.

Harderian gland; #.l.g. inferior labial gland; p.g. parotid gland ; 7.¢. reserve

tooth. X3.

a wound on prey that is already in the mouth, while a straight

tooth is of greater use as a weapon of offence and defence, as it

can inflict a wound on prey that is outside the mouth. Another

improvement has taken place; the groove has become deeper,

and extends through the whole length of the tooth, ending at

the very tip. It has also shifted from the side of the tooth

to the front, in order to be still nearer the opening of the duct,

which opens in front of the tooth sac as in the highly-developed

poisonous snakes.

It is unfortunate that I have not been able to examine sections

of this genus to see the connection of duct with the teeth; but I

have seen only one specimen, and it was kindly lent to me by

Miss Procter, of the British Museum, for purposes of observation

but not for sectioning.

318 MR. §. CG. SARKAR ON THE BUCCAL GLANDS

Conclusions.

Taking a general view of this series, it is interesting to endea-

vour to explain the factors which have brought about the gradual

change and complicated modifications leading us to the most

highly-developed snakes.

Our hypothetical ancestor of the Aglypha would in all proba-

bility be one in which the teeth were equal and peg-like, while

the superior labial gland showed no differentiation. The secretion

was used more or less for digestive purposes, like the saliva of

other animals. The next step in evolution was brought about by

the necessity of having a stronger secretion which would have the

action of disintegrating the prey more quickly within the mouth.

A portion of the gland in the region of the maxilla was differ-

entiated to perform this funetion. The tooth nearest to the

opening of the duct of this gland became a little different from

the others by acquiring a cutting-edge, and the duct of this

portion moved nearer to it. By this arrangement it became

possible for the snake to make a deeper and larger scratch and to

pour poison on the wound, as in the case of Dendrophis. How-

ever, by this method the poison from the now differentiated

poison gland is inevitably mixed, and therefore diluted with the

general salivain the mouth. In order to avoid this dilution, the

tooth nearest to the opening of the poison gland became bigger

and separated from the rest by being enclosed in a special § sac.

The sac is kept filled with poison, so that the tooth is always

immersed in it (Lycodon and Tropidonotus). This arrange-

ment acted as a sure means of making a deeper wound, which

may be compared to poisoning the prey by probing the body

with a lancet besmeared with poison. This is undoubtedly

a slow process, as a sufficient quantity of poison could not be

injected.

The next step is a great improvement in the tooth—a groove

appeared which became almost a canal, as we have seen in

Chrysopelea and Dryophis. At the same time, the gland became

a distinct organ (text-fig. 21) from the general glandular mass of

the superior labial gland, and the duct moved upward to open

finally at the base of the tooth. Evidently this is a great advance,

for by this means pure unmixed poison can be injected into the

deeper tissue. We may compare this with the modern method

of injection by a hypodermic or intermuscular syringe, which

acted almost instantaneously. This seems to be the origin of the

opisthoglyphous snakes.

But even this method has its defects in some ways. The

grooved tooth being situated far behind, the prey has to be taken

well into the mouth before the operation can be performed.

Besides, this tooth cannot be utilised as an organ of offence and

defence. Dryophis, in a primitive manner, has overcome this

difficulty by developing an extra solid tooth in the anterior end

AND TEETH OF THE OPISTHOGLYPHA. 319

of the jaw (text-fig. 20) which disables the prey before it is

carried further into the mouth to the region of the poison fang.

In order to do away with unnecessary apparatus and provide one

fang which would serve all purposes, the poison fang has shifted

forward right to the front of the mouth, dragging with it the

duct of the poison gland. This gave rise to vipers, where we know

that the poison apparatus is perfect in all ways and that the

other teeth have disappeared. Probably evolution has taken

place in this way, as already pointed out by Boulenger :

‘the series (in Opisthoglypha) culminating in such forms as

have the maxillary bone much abbreviated, the solid teeth

reduced to two or three, and the fang extremely large and deeply

grooved. If we turn to the skull of the least specialised among

the Viperide, we see that the poison fangs are situated on the

posterior extremity of the maxillary bone, close to its articulation

with the ectopterygoid, a condition which is identical with that

of Opisthoglyphous Colubrids. It is therefore clear to me that

the Viperids have been derived from the Opisthoglyphs.”

SUMMARY.

Text-figure 22.

s. lo tooth

3 et Transverse

Lateral view of head. UNISTS wae an ae

foe tooth.

1. Hypothetical ancestor.

Gland in the region of maxilla undifferentiated : teeth all equal and peg-like.

All the ducts open in the buccal cavity far away from teeth.

Text-figure 23.

duct. s.Lg. ia

2. Dendrophis pictus.

A small portion, near the posterior end, of the gland (p.g.) differentiated. The

duct of this portion moved inward to open near the hinder tooth (f).

The latter has become different from others by being more bent inwards,

and also has developed a slight cutting-edge (c.e.¢.) on the inner side.

320

MR. S. C. SARKAR ON ''HE BUCCAL GLANDS

Text-figure 24.

3. Tropidonotus stolatus.

A greater portion of the gland in the region of the maxilla is differentiated to form

the parotid or poison gland (p.g.). The hinder tooth (f) is much bigger

than those in front. The cutting-edge (c.e.) is more marked. It is also

enclosed within a special sac (¢.s.). The duct of parotid glands (duct p.g.)

enlarges at the end to form a cavity (ves.) which becomes confluent with

the tooth sac before opening into the mouth. Sac is kept filled with the

secretion and the tooth dipped into it.

Text-figure 25.

4. Lycodon aulicus.

The parotid gland is much enlarged, and almost extends up to ? of maxilla. The

duct (duct p.g.) is similar to Tropidonotus, except that it opens a little

higher. The tooth is like a lancet, with a cutting-edge on each side.

Text-figure 26.

5. Oxybelis fulgida.

The poison gland (p.g.) well developed. The arrangement of the duct is similar to

that of Lycodon and Tropidonotus, i.e. enlarges to form vestibule (ves.)

before opening into the mouth (Aglyphous condition). The fang (f/f) is

laterally compressed. Has almost lost cutting-edges, but has acquired a very

shallow groove like Opisthoglypha.

AND TEETH OF THE OPISTHOGLYPHA. al

Text-figure ie

max. SEER

6. Chrysopelea ornata.

The poison gland is reduced in size, due to the fact that the fang (f) has acquired

a deeper groove (g), through which a greater amount of secretion can be

injected into the wound, and there is very little waste. The wall between

the vestibule (ves.) and the tooth sac has disappeared; at the same time

the opening of the duct into the mouth cavity is closed, and hence the

duct ( p.g.) opens directly into the tooth sac.

Text-figure 28.

duct. p.9..

7. Dryophis mycterizans.

The poison gland (p.g.) is much more differentiated. The groove (g.) of the fang is

deeper and almost like a canal. The duct (duct p.g.) has shifted upward to

open into the tooth sac almost near to the base of the fang, which has also

moved forward in the mouth cavity. :

Text-figure 29.

8. Psammophis sibilans.

The poison gland (p.g.) is distinct—almost a separate organ, The fang (/) 1s

almost straight, and has shifted still further in the mouth cavity. The

groove lies in front of the tooth instead of on the side, nearer to the beginning

of the groove.

322°

O93 2

11.

12.

\ 18.

14.

15.

16.

BUCCAL GLANDS OF THE OPISTHOGLYPHA.

References to the more important Literature.

. Bountenaer, G. A.—‘‘ Remarks on the Dentition of Snakes

and on the Evolution of the Poison Fangs.” Proc. Zool.

Soc. London, 1896, p. 614.

. BoutencErR, G. A.—‘ Fauna of British India,’ 1890.

. Fayrer, J.—‘ The Thanatophidia of India,’ 1874.

. Jourpan, J.—“A Yappareil dentaire particulier qu'il a

observé dans le Coluber.” VInstitut, t. ii. p. 222

(1834).

. Martin, H.—“ Recherches sur le développement de l’appareil

venimeux de la Vipera aspis.” OC. R. Assoc. des Anato-

mistes, 4e Session, Paris, pp. 56-66.

. Niemann, F.—“ Beitrage zur Morphologie und Physiologie

der Oberlippendriisen einiger Ophidier.” Arch. Naturg.,

58, 1892 (Bd. 1.), pp. 262-286.

. Parker, W. K.—“ On the Structure and Development of the

Skull in the Common Snake.” Phil. Trans. Roy.

Soc. London, vol. clxix. 1878.

Parker, W. K., & Brrrany, G. T.—‘ Morphology of the

Skull,’ 1877.

Puisatix, Mary.—‘“‘ Modifications que la formation veni-

meuse imprime a la téte osseuse et aux dents chez des

serpents.” Ann. des Sci. Nat. Zool. 9th ser. t. xvi.

pp. 161-205 (1912).

. Putsatrx, Mary.—‘‘ Anatomie comparée de la téte et de

Vappareil venimeux chez les serpents.” Ann. des Sci.

Nat. Zool. 1914.

Smiru, T.—‘“‘On the Structure of the Poisonous Fangs of

Serpents.” Phil. Trans. Roy. Soe. London, vol. eviii.

p. 471 (1818).

Sunxara NaArayAna Pinitary, R.—‘‘ Note on the Structure

of the Teeth of some Poisonous Snakes found in

Travancore.” Ann. & Mag. Nat. Hist. 7th ser. xviil.

p. 238, London, 1904.

Tomes, Cu. S.—* On the Structure and Development of the

Teeth of Ophidia.” Phil. Trans. Roy. Soc. London,

vol. elxv. 1875.

Toms, Cu. S.— ‘On the Development and Succession of

the Poison Fang of Snakes.” Phil. Trans. Roy. Soc.

London, part ii. 1875.

West, G. S.— “On the Buccal Glands and Teeth of Certain

Poisonous Snakes.” Proc. Zool. Soc. London, 1895,

. 812.

een G. S.—* On the Histology of the Salivary, Buccal,

and Harderian Glands of the Colubride, with a note

on their tooth succession and the relationships of

the poison duct.” Journ. Linn. Soc., Zool. vol. xxvi.

pp. 517-526.

PZ. S: 1923, SONNTAG. Pi:

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ANATOMY OF THE CHIMPANZEE.

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ANATOMY OF THE CHIMPANZEE.

ON THE ANATOMY OF THE CHIMPANZER. 323

22, On the Anatomy, Physiology, and Pathology of the

Chimpanzee. By Cuartes F. Sonnrac, M.D., F.Z.S.

Anatomist to the Society, and Demonstrator of Anatomy

University College.

(Plates I.-ITI.* ; Text-figures 25-49.)

[Received March 17, 1923: Read April 10, 1923.]

CoNnTENTS.

Page Page

‘Introduction ......................... 823 | Respiratory Organs ............... 397

Muscular System .................. 324 | Urogenital Organs ............... 399

SUOITIUS Sons non Sod ncobaodoacodsdouduobene ake) I] ONIGIAO LG Shs, | Vesa sd ae asscteeng | CKO:

Organs of Digestion ............... 37 Sense Organsisecees cece eee Lo

Organs of Circulation ...........; 376 | Skin and Hairs .................... 417

Wuetlessi Glands eee eee eee CO Ome batho lo ya es eee ALLS

Blood Gisten cae reneeeesereeccter cere mio Ons) COMMpaLsonsawithe Mant soso 419

Lymphatic System ............... 896 | Bibliography ........................ 426

INTRODUCTION.

Zoological literature contains descriptions of parts of nearly

three hundred Chimpanzees, but the anatomy of one animal only

has been described at any length by Gratiolet (22). His account

omits many special points, which have also been neglected by

other observers, and the same can be said of the works of

Sperino (47) and Vrolik (51), which give accounts of the com-

parative anatomy of all the Anthropoid Apes. It is, therefore,

evident that a full account of one animal is required to serve as a

standard for future workers. The present account is based on

the examination of a young female, Anthropopithecus troglodytes,

which died in the Society’s Gardens after a residence of two and

a half years. And if that species is different from Troglodytes

aubryi it should be a useful companion to Gratiolet’s account of

the latter. The animal had the following measurements :—

Length from supra-orbital crests over head and

baicketovamus!) 32.02 SO a a Ae OT 23°5 inches

Length from supra-orbital crests to inion ...... G74 4

13 1, 2. IMTOn FOAMS LOOT Veh ey 1a

Tip of acromion to centre of antecubital fossa. 9:3 ,,

Centre of antecubital fossa to lower end of

Fads) 3.6.71 9M. BOE ek Oe ean Oe eh

Hand (palm 4 ins.: middle digit 3:2ins.)......... ipa taa

* For explanation of the Plates.see p. 429.

324 DR. OC. F. SONNTAG ON THE ANATOMY,

Total length of pectoral extremity ............... 25°7 inches

Great trochanter to centre of patella ............ SE EN

Centre of patella to lower end of tibia ............ (oman

Foot (sole 4°6 ins.: middle digit 2°8 ins.)......... (RA

Total length of pelvic extremity .................. 2471 ous

Excess of pectoral over pelvic extremity ......... LEO oe

THE MuscuLaR System.

Muscles of the Head, Neck, and Back.

The platysma myoides (text-fig. 25) is much thicker than in

Man. It arises from the fascia over the pectoralis major and

deltoid, and the two muscles are inseparable in the middle line

of the neck. Its lateral parts are more muscular and thicker

than the medial portions. About an inch below the symphysis

menti the fibres of the mesial parts decussate (Ruge 42, Champ-

neys 11), and I observed the fibres of the left muscle lying super-

ficial to those of the right one; but Quain describes the reverse

Text-figure 25.

=== TEMPORAL -

Z.Mi. FASCIA

Hi) y, Z.Nia. \\ \\

Yy thy, > W\\W i)

LEV. LAB.SUP_<¢ Uf 4) AUR.ANT.

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FASCIA.

ORB.

ORIS*-

SS a ~aqg wmQA q_

Superficial muscles of the face. ORB.OC: orbicularis oculi; ORB.ORIS:

orbicularis oris. Other letters in text.

condition in Man. The fibres are attached to the lower border

of the mandible, the skin of the lips, and the muscles of the lips

and angles of the mouth. But no fibres are attached to the

zygoma as described by Champneys. In the face it separates

into upper and lower bundles of fibres ; the former, corresponding

to the risorius in Man, runs to the muscles at the angle of the

mouth ; and the latter, which is much larger, blends with the

skin and muscles of the lower lip. Small fibres, running from

the platysma to the angle of the mouth, correspond to the

triangularis muscle (é77.).

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZER. 325

A fan-shaped muscle separates from the platysma in the neck,

runs upwards behind the auricle and spreads out into bundles

which are attached to the back of the auricle, the occipital crest,

occipitalis muscle, and the deep fascia over the back of the neck.

Ruge (42) has given a very elaborate account of the manner in

which the platysma enters into the other facial muscles.

Oceipito-frontalis (text-fig. 25):—There are many differences of

opinion about this muscle. Tyson (50) and Traill (49) could not

detect it, Owen (39) found a trace of it, and Wilder (53) found

the muscle bellies small, but the aponeurosis was large. Ruge (42)

figured a very extensive muscle and a small aponeurosis.

In my specimen the occipitalis arises from the middle two-

fourths of the occipital crest, but it is not divisible into two bellies

asin Man. The fibres pass forwards for nearly two inches and

end in a well-marked aponeurosis. The frontalis arises from the

supra-orbital ridges and space between, but is not so well-

marked as the occipitalis. It is very easily removed with the

skin. It blends with the orbicularis oculi.

The Orbicularis oculi (text-fig. 27. A) is divisible into orbital

and palpebral parts as in Man. ‘The former arises from the inner

end of the frontal bone and the nasal process of the maxilla; and

both muscles are united across the mid line. As it lies on the

bones bounding the orbit its upper part is strong and compact and

gives offa strong bundle of fibres from its lateral part to enter the

zygomatic mass (Z.M). The fibres on the lower boundary of

the orbit are arranged in loose bundles. The palpebral fibres run

from the internal tarsal ligament to the lateral tarsal raphé, and

are thickened close to the roots of the eyelashes, the thickened

parts being of greater dimensions than the ciliary bundles (C.B)

in Man. At the lateral tarsal raphé the orbital and palpebral

parts are continuous. The nerve-supply from the facial nerve is

shown in text-fig. 26.

The lips and cheeks receive many muscles (text-fig. 25), most

of which, though thin, are of considerable superficial extent.

They are disposed in two layers as in Man, but the characters

are very different in a number of points. The superficial layer

is composed of the risorius, levator labii superioris, zygomatic

mass, orbicularis oris, triangularis and quadratus labii inferioris.

‘The deep layer consists of buccinator, depressor anguli oris,

incisivi, canini, mentales, and premolares. The risorius is com-

posed entirely of the upper part of the platysma, for no fibres

are derived from the fascia over the masseter muscle. It blends

with other muscles at the angle of the mouth. The levator labii

superiorts (Lev. Las. Sup) arises, under cover of the orbital

part of the orbicularis oculi, from the entire infra-orbital border

of the maxilla. It radiates in a fan-like manner and is inserted

into the entire length of the upper lip and upper border of the

ale nasi. The fibres forming the latter insertion correspond to

the levator labii superioris aleeque nasi of Man. Many of the

fibres of the muscle are very thin. Champneys (11) states that it

Proc. Zoou. Soc.—1923, No. XXII. 92

326 DR. C. F. SONNTAG ON THE ANATOMY,

is not well differentiated from the levator anguli oris, but that is

not the case in my specimen; it is only at their insertions that

these muscles are fused. Gratiolet (22) describes a mingling of

the fibres with those of the orbicularis and they cover the malar-

maxillary articulation.

The zygomatic mass (text-fig. 25) in my specimen differs from

the muscles described by Ruge (42), Gratiolet (22), and Champ-

neys (11). In all the descriptions and published figures it is less

powerful, or the parts are more separate. In my specimen it is

the most powerful muscle in the face, and has three powerful

heads of origin. A strong bundle separates from the orbital part

of the orbicularis palpebrarum, the zygomaticus minor (Z.Mi)

Text-figure 26.

GLAND.

COKY

Ba!

ee ae ioe

eae

OTID

PAR

Deep muscles of the face. A.F.V: anterior facial vessels; F.N: facial nerve;

I.C.A: inferior coronary artery; L.S.G: labial salivary glands; MAS:

masseter; ORB.OR: orbicularis oris; §.D: Steusen’s duct; S.T.V and

T.F.V : superficial temporal and transverse facial vessels; T.M: temporal

muscle; Z.F.V : zygomatico-facial vessels. Other letters in text.

springs from the malar bone and temporal fascia, and the zygo-

maticus major (Z.Ma) arises from the anterior end of the zygoma.

The three heads unite to form a strong muscle an inch wide

blending with the muscles of the lips and angle of the mouth,

It probably does much more work than the proper levator anguli

oris. The guadratus labii inferioris (text-fig. 26) has a lower

origin than in Man. It springs from the posterior half of the

lower border of the outer surface of the body of the mandible.

It is in contact with the masseter behind and receives fibres from

the platysma below. The fibres course upwards and forwards

and blend with those of the orbicularis oris. The anterior fibres

are fine and close together, and interlace in the front of the lower

lip with fibres of the opposite muscle. Running through the

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 327

muscle are branches of the inferior labial artery (1.L.A), and

the mental branch of the inferior dental nerve emerges from

the mandible underneath it. Champneys (11) states that this

muscle is not differentiated.

Text-figure 27.

Weg Nahe YY Ly

. t| 4 Vl

L.A.N. LEV LAB. SUP: d ify 4 s

2M. YW

A Y

pe,

CONJ.

a ; BAG Rane

M.L. GDS.

Muscles of the eyelids (A), eyes (C-F) and nose (G): conjunctiva (B). C: cilia;

CONJ: conjunctiva; E: eyelids; E.N.N: external nasal nerve; F.M:

frontalis; I.0: inferior oblique in the Chimpanzee (C) and Man (D); I.T\L:

internal tarsal ligament; L.T.R: lateral tarsal raphé; M.L.GDS: Meibomian

glands; O.P: orbitalis; P.S: plica semilunaris; P.P: palpebralis; L,P.S:

levator palpebrz superioris of Man (E) and the Chimpanzee (F); S.O:

superior oblique muscle.

The triangularis (text-fig. 25, irt.), although figured as a promi-

nent muscle by Ruge (42),. is represented by a few broad fibres

passing from the platysma to the angle of the mouth.

The orbicularis oris:is composed of fibres from all the facial

muscles except the levator labii superioris aleque nasi; and it

22*

328 DR. C. F, SONNTAG ON THE ANATOMY,

has the usual sphincteric arrangement. From its deep surface it

gives off small muscular slips (text-fig. 27) which are attached

to the bones at the bases of the sockets of the incisor, canine and

premolar teeth. These are best developed in the upper jaw.

Between them and the mucous membrane there are numerous

labial salivary glands and branches of the infra-orbital nerve

plexus (text-fig. 26, I.0.P).

Between the levator labii superioris and the levator anguli oris

numerous branches of the facial nerve and infra-orbital branches

of the trigeminal nerve ramify and anastomose, and numerous

labial salivary glands are present. The facial nerve supplies the

muscles, and the infra-orbital nerves can be traced to both skin

and mucous membrane.

The levator anguli oris (text-fig. 26, L.A.O) is a small triangular

muscle. It arises from the maxilla below the infra-orbital fora-

men, and is inserted into the orbicularis oris at the angle of the

mouth. A small slip passes to the skin of the upper lip. The

latter is not mentioned by other authors.

The buccinator (text-fig. 26, Buc) arises from the maxilla and

mandible close to the roots of the last molar teeth and from the

pterygo-maxillary ligament. It emerges from under cover of

the ascending ramus of the mandible and blends with the orbicu-

laris oris in both lips; but the fibres do not decussate as in Man.

Lying on its surface are a pad of fat, several buccal salivary

glands, the buccinator branches of the internal maxillary artery,

and the long buccal branch of the trigeminal nerve. It is crossed

by the anterior facial vein and external maxillary artery.

Nasal Muscles (text-fig. 27 A) :—No nasal cartilages exist *, so

the nasal muscles are inserted into the skin. The upper border

receives a continuous strip of muscles from the combined orbicu-

laris oculi and frontalis and the levator labii superioris. The

former corresponds to the pyramidalis (Pyr) and the latter to

the levator labii superioris aleque nasi (L.A.N). Three small

muscular slips arise from each half of the nasal orifice of the skull

and are inserted into the deep surface of the skin. No depressor

septi nasi is present. The sensory external nasal nerve is seen

emerging from the nasal fossa on each side.

Extrinsic Muscles of the Auricle (text-fig. 25):—Ruge (42)

described and figured a superior auricular muscle descending

from the vertex of the skull to the root of the pinna, and auriculo-

occipital and posterior auricular muscles acting on its posterior

part. A small tragicus runs to the lower and front part of the

pinna. Wilder (53) described an attolens and a combined

attrahens and retrahens. In my specimen the attolens (Aur.

Ant) arises from the epicranial aponeurosis, and the combined

attrahens and retrahens (Aur. Posr) arises from the aponeurosis

and occipital crest, touching the occipitalis above. The fibres

* This statement is based on both macroscopic and microscopic examination.

But future material may show that the conditions here are purely individual in

character.

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 329

of both muscles are continuous on the root of the pinna. Slips

from the platysma (Pua) go to the back of the auricle, and a

small tragicus (TRAG) is present.

The masseter (text-fig. 28 A) consists of the usual superficial

and deep parts. The former (8.P.M) arises from the lower

borders of the malar bone and anterior two-thirds of the zygoma ;

Text-figure 28.

aT AURAL IRN

SS AARNE

TANADMNNY UA RE)

! \ wD

Wey Ved a

MUA We SSy

\ WAS S

Muscles of mastication; A: Masseter; B: Attachments of muscles to the

mandible; C: Muscles, nerves and vessels below the labial mucous

membrane. B.A. and BUC: buccal artery and buccinator muscle; J.O.N.,

L.B.N., and M.B.T: infra-orbital, long buccal, and mental nerves; L.L. and

U.L: lips; M.I. and M.M: incisive and mentalis muscles; L.P. and 8.P:

labial and salivary papille; S.P.F: suctorial pad of fat. Other letters as in

text-fig. 26,

the fibres pass downwards and backwards to be inserted into the

margin of the lower border, angle, and lower half of the posterior

border of the ramus of the mandible. Between the two parts is

a strong fascial sheet into which fibres of both parts are inserted.

The deep part (D.P.M) arises from the entire length of the deep

330 DR, GC. . SONNTAG ON THE ANATOMY,

surface of the zygoma, and the fibres converge to be attached to

the anterior two-thirds of the outer surface of the mandibular

ramus and coronoid process. Numerous large vessels (T.F.V)

ramify between the masseter and parotid gland and supply both.

The actual insertions are shown in text-fig. 28 B.

The temporal muscle is large and powerful. It arises from the

entire temporal fossa from the external angular process of the

frontal bone in front, to about four centimetres behind the concha

and upwards to a point level with the supra-orbital crest. It

also arises from the temporal fascia which covers it. The fibres

are strong, coarse and mixed with tendinous bands; they

converge to be inserted into the anterior border, point and

posterior border of the coronoid process (text-fig. 28B). The

anterior part of the muscle is attached by muscle fibres to the

anterior border of the process, which is of considerable length.

But the posterior part is attached by aponeurosis to the back-

wardly-directed point and short posterior border. On the surface

of the muscle the zygomatico-facial artery ramifies. A piece of

the aponeurotic insertion sweeps over the outer surface of the

coronoid. The deep temporal vessels anastomose within it. The

action of the temporal muscle is described at length by Gratiolet.

The attachments of the masseter and temporal muscles to the

mandibular ramus are shown in text-fig. 28 B. The temporal

fascia is attached to the temporal crest, external angular process

of the frontal bone, malar bone and upper border of the zygoma.

It is overlain by a considerable deposit of fat. It gives an

attachment to the fibres of the zygomaticus minor and extrinsic

muscles of the auricle. A few fibres of the temporal muscle arise

from it.

The pterygoid muscles (text-fig. 29 A) are very similar to those

in Man, and all authors who have described them come to similar

conclusions. The relations of the various nerves in the pterygoid

region are the same as in Man, and the internal maxillary artery

(I.M.A) crosses the outer surface of the external pterygoid

(E.P.M) as in some human bodies. The veins, however, do not

form a large diffuse plexus, but consist of tributaries accom-

panying the large arteries and opening into an internal maxillary

vein. It divides into two veins which unite with the superficial

temporal vein. It communicates with the anterior facial vein

and with deep veins in the neck. No lymphatic glands are

present in the pterygoid region, but much fat is present. It is,

therefore, evident that, with the exception of the characters

of the veins, the pterygoid region is essentially similar to that

in Man.

The sterno-mastoid (text-fig. 30, S-M.M) arises by a long,

gently tapering, strong tendon from the inferior border of the

manubrium sterni, and it does not develop museular fibres till it

reaches the neck. A few small tendinous bundles run from the

tendon of origin to the upper and mesial aspects of the sterno-

elavicular articulation, and strong fascia unites the tendon to the

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 331

Text-figure 29.

wit),

VI/

STN)

1 Wf, 4] 4 LM.A.

LDN. ADA.) FF: A

IOOy

S.C.G.

Pterygoid (A) and Palatal (B) regions. E.P.M. and I.P.M: external and internal

pterygoid muscles: A.P.A: aseending pharyngeal artery; I.M.A: internal

maxillary artery (letters of branches on p. 384); I.C.A: internal carotid

artery; I.D.N: inferior dental nerve; I.J.V: internal jugular vein; C.T.N :

chorda tympani joining the mandibular nerve (M.D.T.); H.P: hard palate

dropped for diagrammatic purposes below its true level (lev. H.P.); P.G:

palatal glands; P.H: pterygoid hamulus; S.C.G: superior cervical sympa-

thetic ganglion; IX. XI: cranial nerves.

332 DR. C. F. SONNTAG ON THE ANATOMY,

inner end of the clavicle. It gives attachment to a few fibres of

the pestoralis major, but it is not enveloped by the latter as

described by Gratiolet and Alix (22). In the neck it forms a

wide, comparatively thin muscle, whose fibres are coarse. It is

inserted into the outer half of the superior curved line of the

occipital bone, overlapping the trapezius. No fibres are attached

to the mastoid process. Between the sterno-mastoid and

subjacent cleido-mastoid there are muscular branches of the

Text-figure 30.

\\'

ANY

é s-M.M\\

Muscles of the middle of the neck. L.G@: lymphatic glands; 0.A.S: opening into

the air-sac from the ventricles of the larynx; O-H.M: omo-hyoid muscle;

S.M.G: submaxillary gland; S-T.M: sterno-thyroid muscle; T.C: thyroid

cartilage; Sy-H.M: stylo-lyoid muscle; Tra: trachea. Other letters in text.

occipital artery. The external jugular vein does not cross the

surface of the muscle, and the transverse cervical nerve, after

emerging from beneath the cleido-mastoid, runs forwards over the

surface of the sterno-mastoid. ‘The sterno- and cleido-mastoids

are separate throughout, though closely apposed and surrounded

by fascia.

The clerdo-mastoid (text-fig. 30, C-M.M) arises from the inner

third of the upper border of the clavicle. As it passes upwards

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 333

in the neck it becomes narrower and is inserted into the outer

surface of the mastoid process. The muscle raises a prominent

ridge on the anterior wall of the air-sac. Many vessels and

nerves pierce the deep fascia at the lateral border of the cleido-

mastoid; and the spinal accessory nerve passes into its deep

surface in the upper third.

When an incision is made through the platysma and deep

fascia the wall of the air-sac makes its appearance. The wall

varies in thickness in different parts, and its lining is smooth and

moist. It consists of a central part with two lateral diverticula.

The central part extends upwards to the hyoid bone, and down-

wards to the lower border of the manubrium sterni between the

tendons of origin of the sterno-mastoid muscles. Its anterior

wall is covered by the platysma, and the larynx, trachea and pre-

tracheal muscles shine through the thin posterior wall. The

lateral parts are very capacious, and have large circular orifices

under cover of the cleido-mastoids. When these are explored

the finger can pass along the greater part of the deep surface of

the pectoral muscles and the inner border of the deltoid; it

palpates the entire length of the clavicle, the head of the humerus,

the glenoid cavity, and borders of the scapula. Many muscles,

nerves, and the carotid sheath form ridges in the walls of

the sac.

The omo-hyoid (text-fig. 32 A) is more complex than in Man,

and it is more complex in my specimen than in others described.

It consists of three bellies. The postero-mesial belly is tapering.

It arises from the back of the first costal cartilage along with the

sterno-thyroid muscle, with which it is considerably fused. The

anterior belly is tapering, and inserted into the lower border of

the hyoid bone at the side of sterno-hyoid. The postero-lateral

belly, which is the strongest, arises from the upper border of the

scapula close to the root of the coracoid process. All three bellies

meet in a Y-shaped junction, and a tendinous thread runs into

sterno-hyoid.

The sterno-hyoid (text-fig. 30, S-H.M) arises from the back of

the upper part of the manubrium sterni, and is inserted into the

lower border of the hyoid bone. The opposite muscles first

diverge and then converge, and fibres pass between them on the

hyoid bone. The sterno-thyroid arises from the back of the

manubrium sterni and first costal cartilage and is inserted into

the upper part of the thyroid ala. Some fibres pass into the

thyro-hyoid muscle.

The digastric muscle (text-fig. 30, D.M) is transitional between

Parson’s first and third types. The anterior bellies are only

separate in front. They are fused behind where they arise from

the front of the body of the hyoid bone. Hach belly is inserted

into the anterior two inches of each half of the mandible. The

posterior belly is bulky, but the tendon (text-fig. 30, P.B.D), which

enters the postero-lateral part of the anterior belly immediately

in front of the hyoid bone is long and slender. It arises from

334 DR. C. F., SONNTAG ON THE ANATOMY,

the depression on the temporal bone corresponding to the

digastric fossa in Man. Chaine (10) has recently described the

digastric muscle. The tendon of the posterior belly tunnels the

stylo-hyoid muscle; it has no direct attachment to the hyoid

bone as described by Gratiolet (22).

Some fibres of omo-hyoid and sterno-hyoid pass into the

anterior bellies of the digastrics.

The stylo-hyoid arises by one large and several small tendons

from the styloid process and bone around. It is long, thin, fleshy,

and wrapped round the digastric tendon. It is inserted into

the upper border of the hyoid bone opposite sterno-hyoid and

omo-hyoid.

The mylo-hyoid (text-fig.31, M-H) arises from the upper

border of the hyoid bone under cover of, but never fused with,

Text-figure 31.

Anatomy of the submental region (No.1). A.B.D: anterior belly of the digastric

turned down; A.F.V: anterior facial vein; H.J.V: external jugular vein;

L.B.T: lingual nerve; P.B.D: posterior belly of the digastric; P.F.V:

posterior facial vein; S.M.G: submaxillary gland; Sy.H: stylo-hyoid

muscle; W.D: Wharton’s duct. Other letters in text.

the anterior belly of the digastric. The level of origin corre-

sponds to the extent of the insertions of the sterno-hyoid and

omo-hyoid on the posterior border. The fibres radiate to be

inserted into the inner surface of the mandible. The posterior

fibres lie just in front of the submaxillary gland. Lying on the

surface of the muscle and supplying it are branches of the sub-

maxillary twigs of the external maxillary artery. I did not

observe any decussation of fibres in the middle line as described

by Gratiolet (22).

No submental lymphatic glands were found behind the

symphysis menti.

The genio-hyoid muscles (text-fig. 31,G-H) are separated

anteriorly close to their origins from the lower part of the genial

fossa on. the back of the symphysis menti. In the greater part of

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 339

the interramal space the two muscles are in contact in the mid

line. They are inserted into the upper border of the body

and part of the great cornu of the hyoid bone. At their sides

lie deposits of fat surrounding the sublingual glands (S8.L.G),

the numerous arteries to these glands, and the large lingual

branch of the trigeminal nerve (L.B.T). The latter is seen

vanishing under cover of the genio-hyoid muscle. The hypo-

glossal nerve (XII) also is seen dividing into branches whick pass

under the muscles and sublingual glands. When the genio-

hyoidei are reflected it is seen that a well-marked bursa, capable

of lodging the tip of the index finger, lies between them and the

hyoid bone anteriorly and the genio-glossi posteriorly ; and there

is a thick fatty septum between the two genio-glossi. The hyo-

glossi crossed antero-posteriorly by the strong, thick, stylo-glossi ;

the sublingual glands and the hypoglossal and lingual nerves are

further displayed. The former is seen giving two branches to its

corresponding genio-hyoid.

The genio-glossi (text-fig. 32, G.G) are two long, narrow, thick

muscles arising from the bottom of the genial fossa. They are

separated in the mid line by a comparatively thick deposit of fat,

and a considerable interval separates each from the mandible.

In that space the entire sublingual gland, the hyoglossus and

styloglossus muscles, the lingual and hypoglossal nerves and the

lingual artery are seen. The artery emerges from under the

hyo-glossus. Some fibres of the genio-glossi reach the hyoid bone

under the hyo-glossus.

The hyo-glossus (text-fig. 32, HY) arises from the lateral part

of the body and the whole of the great cornu of the hyoid bone,

but the origin from the body does not spread over a half as stated

by Gratiolet (22). The fibres pass upwards to be inserted into

the side of the posterior half of the tongue under the stylo-

glossus. It is not fused with the opposite muscle, but fibres of

the thyro-hyoid can be traced into it. A great part is concealed

by the stylo-glossus. The relations are very similar to those

in Man.

The stylo-glossus (text-fig. 32,8.G) is relatively more powerful

than that in Man. It arises by a short, rounded strong tendon from

the outer surface of the base of the styloid process. It describes a

curve as in Man, and its volume increases greatly as it is traced

forwards. It gains an attachment to the side of the tongue from

the level of the outer border of the hyo-glossus behind to nearly the

apex of the tongue in front. It covers the upper half of the

hyo-glossus, and it extends from the side of the tongue above to

the outer border of the genio-glossus below. The part anterior to

hyo-glossus is concealed by the large sublingual gland, with the

lingual branch of the trigeminal nerve curving round its posterior

pole. The connecting loop between the lingual and hypo-glossal

nerves crosses it anteriorly, and Wharton’s duct (W.D) crosses it

obliquely from above downwards and behind forwards.

The superior constrictor of the pharynx is continuous above and

336 DR. C. F. SONNTAG ON THE ANATOMY,

in front with the buccinator, both being attached to the pterygo-

mandibular ligament. It is attached to the mylo-hyoid line on

the mandible, the internal pterygoid plate, the base of the tongue,

the mucous membrane of the floor of the mouth and the bucco-

pharyngeal aponeurosis. The lower border is overlapped by the

middle constrictor, and the stylo-pharyngeus passes between

them as a few separate, but thick, bundles of fibres. Some of

these fibres pass into, and blend with, the outer surface of the

superior constrictor. The upper part of the muscle is separated

by a large sinus of Morgagni from the base of the skull, but the

levator palati and tensor palati, which are situated therein, lie

horizontally, whereas they are more vertical in Man, and of

smaller size. The constrictor is attached above and behind into

Text-figure 32.

The omo-hyoid muscle (A) and the anatomy of the submental region (B). A.B.O.>

P.L.0., and P.M.O: anterior, postero-lateral and postero-mesial bellies of

the omo-hyoid muscle; F.S: fatty septum; L.A: lingual artery; N.G.H:

nerves to the genio-hyoid muscle; H.B: hyoid bone; 8.H. and 8.1: sterno-

hyoid and sterno-thyroid muscles; 1X and XII: cranial nerves. Other

letters as in text-fig. 31.

the basis cranii. It is difficult, and in some places impossible, to

separate the superior constrictor from the stylo-glossus muscle

which courses downwards and forwards on its outer surface.

A well-marked bundle of fibres passes towards the angle of the

mouth. The middle constrictor of the pharynx arises from the

deep surface of the hyoid bone in the angle between the greater

and lesser cornua, and it is inserted into the mid-dorsal line of

the pharynx, its fibres mingling with those of the opposite

muscle, It is overlapped by the inferior constrictor. Some fibres

of the stylo-pharyngeus pass into its outer surface. Between the

superior and middle constrictors there is a non-muscular area

anteriorly. The inferior constrictor of the pharynx arises as in

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 337

Man from the oblique line on the thyroid cartilage and from the

side of the cricoid. Its fibres sweep more or less upwards,

overlap the lower border of the middle constrictor and blend

with the opposite muscle in the mid-dorsal lme. It has no origin

from the first tracheal ring as in Gratiolet’s specimen (22).

The stylo-pharyngeus (text-fig. 32,8.P) arises from the tendon

of the stylo-glossus, but Gratiolet (22) states that it rises from

the base of the styloid apophysis. It splits up into bundles some

of which are inserted into the superior and middle constrictors,

others passing between these muscles and radiating in the wall

of the pharynx. The glosso-pharyngeal nerve hooks round it

and sends it a well-marked branch.

The levator palati and tensor palati (text-fig. 29 B) arise by

a strong, common musculo-aponeurotic origin from the apex of

the petrous temporal bone, the under surface of the Eustachian

tube and the scaphoid fossa. So the separate origins of the

muscles have fused in this animal. The levator palati (L.P)

runs downwards and forwards and spreads out between the

layers of the palato-pharyngeus. The tensor palati (T.P) is

even more horizontal. Its tendon winds round the pterygoid

hamulus and is inserted by several small tendinous and fascial

bundles in the palatal aponeurosis. The complete limits of the

palato-pharyngeus (P.P) could not be accurately made out, and

the palato-glossus hardly exists. The azygos wvule (A.U) ends

posteriorly in membrane as pointed out by Gratiolet (22).

The thyro-hyoid runs from the entire width of the thyroid ala

to the under and outer surfaces of the body and great cornu of

the hyoid bone. Its nerve from the hypoglossal is well marked.

The scalenus anticus arises from the anterior tubercles of the

transverse processes of the third, fourth, and fifth cervical

vertebre, but others have given its origin from 4, 5, and 6.

It is connected by a tendon to the rectus capitis anticus major.

It is inserted as in Man, the tubercle on the first rib being lateral

to the chondro-costal junction. The scalenus mediusand scalenus

posticus arise as in Man. ‘They unite to form a flat sheet which

courses downwards to be attached to the outer surfaces of the

first five ribs. The fusion and extent are greater than that

described by Gratiolet (22) and others. It is crossed posteriorly

by the slips of insertion of the upward continuation of the sacro-

spinalis.

The omo-trachelian runs as usual from the transverse process

ef the atlas to the upper and outer aspect of the acromion.

It has been recorded by some as being not an omo-trachelian, but

as an acromio-basilaris.

Muscles and Fascie of the Back.

The fascia covering the trapezius and latissimus dorsi is of

great strength, especially below. It is attached above to the

occipital crest, mesially to the vertebral spines and below to the

338 DR. C. F, SONNTAG ON THE ANATOMY,

iliac crest. It is continuous with the fascia over the gluteal

muscles. Laterally it is continuous with deep fascia of the neck,

thorax and limbs. : ‘

The trapezius avises from the inner third of the occipital crest,

the external occipital protuberance, all cervical spines, the thirteen

dorsal spines and the supraspinous ligament. There is no

ligamentum nuche, so the origin differs from that in Man.

Various authors have recorded it as arising from the first ten or

twelve dorsal spines. The lower border is not fused with the

‘atissimus dorsi, as described by Champneys (11), Bland Sutton (4),

and others, though some anatomists did observe fusion. Close

to the lower angle of the scapula there is a triangle of auscul-

tation similar to that in Man. The whole origin is muscular,

there being no aponeurosis close to the vertebral spines as there

is in Man. It is inserted into the outer third of the posterior

border of the clavicle, some fibres passing into the deltoid, the

outer border of the acromion and the whole length of the spine

of the scapula. ‘The most lateral part of the spinous insertion is

aponeurotic. There is no differentiation of fibres inserted into

a special area on the root of the spine of the scapula as there is

in Man. The spinal accessory nerve can be traced almost to the

lower border of the muscle, and gives off numerous branches to it.

It communicates with the third and fourth cervical nerves, but

there is no marked sub-trapezial nerve plexus. It divides at the

root of the neck into two marked branches. One of these goes

to the cervical part of the trapezius and the other to the thoracic

art.

i The latissimus dorsi arises from the lower five dorsal spines

and supraspinous ligaments, the posterior lamella of the lumbo-

dorsal fascia and the posterior lip of the iliac crest from the

highest point to the anterior superior spine, where it overlaps

the outer border of the external oblique. It also receives slips

from the ninth, tenth and eleventh ribs, but none from the

inferior angle of the scapula. On the ribs, whence it derives

slips, it fuses with the origin of the external oblique. The

strong tendon is inserted into the floor of the bieipital groove on

the humerus, and is extensively fused with the teres major and

dorso-epitrochlearis. No band runs across the axillary vessels.

Bland-Sutton (4) emphasises the absence of the latter slip.

Champneys (11) gives the costal origin as ribs 10-13, and

mentions a lesser adhesion to the teres major. Hepburn (24)

gives its origin from the anterior half of the outer lip of the iliac

erest, and records three costal slips; he also mentions slight fusion

with the teres major.

The rhomboideus is an undivided sheet arising from the last

two cervical and first four dorsal spines. It is inserted into the

lower three-fourths of the vertebral border of the scapula. The

insertion reaches higher than the root of the spine of the scapula,

and its upper part overlaps the insertion of the levator anguli

_ scapule. Gratiolet (22) described the origin as extending from

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 339

the occiput to the seventh dorsal spine. The sheet is undivided

into major and minor muscles, in which it agrees with that

described by Gratiolet (22), Vrolik (51), Macalister (33), Bland-

Sutton (4), and Wilder (53). Champneys (22) describes major

and minor rhomboids in great detail.

The levator anguli scapule (text-fig. 33, L.A.S) arises by five

slips from the posterior tubercles of the first five cervical vertebra,

the first being the largest. The lower three origins are tendinous

and fused with splenius cervicis. It is inserted into the upper

fourth of the vertebral border of the scapula. It is not adherent

to serratus magnus, nor is it divisible into two parts. Champ-

neys (11) gives the origin as the first two cervical vertebre ;

Gratiolet (22) gives it as the second and third; Hepburn (24)

records origins from three cervical vertebre ; Wilder (58) gives it

as in Man, and Bland-Sutton (4) records it as springing from the

first five.

The serratus posticus superior (text-fig. 33, 8.P.S) arises by a

thin aponeurosis from the spinous processes of the seventh

cervical and first dorsal vertebre, and it is inserted by four

muscular slips into the outer surfaces of the first four ribs at

their angles. Macalister (33) describes it crossing the first two

ribs to be attached to the third and fourth. The serratus posticus

inferior arises from the posterior lamella of the lumbo-dorsal

fascia and sweeps antero-laterally to be inserted into the lower

borders of ribs nine to thirteen just external to their angles.

Bland-Sutton (4) gives its attachments as ribs nine to twelve.

The lumbo-dorsal fascia is arranged asin Man. The thoracic

part is thin and transparent, and it is difficult to separate it off

from the subjacent muscles as a continuous sheet. The lumbar

part is very dense and strong. The posterior lamella is not easily

separated from the latissimus dorsi, to which it gives origin.

Posterior branches of the spinal nerves pass through its deep

surface. In its lower part it gives origin to the serratus posticus

inferior. Between the outer and middle lamelle the erector

spine is present, and the attachments of the middle lamella are as

in Man. The quadratus lumborum lies between the moderately

strong middle lamella and the weaker internal lamella, whose

attachments to the arcuate ligaments of the diaphragm are as

in Man.

In the possession of a serratus posticus inferior the Chimpanzee

resembles Man, and that muscle is one of the three charac-

teristically human muscles. It will be seen later that the

Chimpanzee possesses the plantaris, which is the second human

muscle, but it does not possess the peroneus tertius.

The muscles of the back described above are relatively stronger

than in Man, and they are relatively weaker than the pectorales,

but the total bulk of these groups has probably diminished

during captivity.

The splenius (text-fig. 33) arises from the sides of the tips of

the fifth, sixth, and seventh cervical vertebre, and from the spines

340 DR. C. F. SONNTAG ON THE ANATOMY,

of the first seven dorsal vertebrae. All the origin is muscular

except that from the sixth and seventh cervical vertebre. The

greater part becomes splenius capitis (Spl.Cap.) which is inserted

into the mastoid and .outer part of the superior curved line of

the occipital bone, and a small slip runs into the first head of the

levator scapula. The splenius cervicis (Sple. Cer.) consists of one

digitation which joins the third head of the levator scapule.

Text-figure 33.

Winns

SCAPULA

Muscles of the back. Comp: complexus; R.M: rhomboideus.

Other letters in text.

Other observers give the splenius cervicis insertions into the first

four cervical vertebree.

The sacro-spinalis (erector spine) is divisible into three columns

—an outer ilio-eostalis, a middle longissimus, and an internal

spinalis—but their characters differ from those of the corre-

sponding muscles in Man. The outer and middle columns are

intimately fused at the origin of the muscle; they separate in the

lumbar and greater part of the thoracic region; in the upper

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 341

thoracic and lower cervical regions they are fused; and the com-

bined mass eventually breaks up again into muscular slips.

The inner column separates off in the upper thoracic region.

The ilio-costalis (text-fig. 33, I-C.C) arises with the subjacent

longissimus from the iliac crest between the highest point and

the posterior superior spine, and it derives fibres from the

covering posterior.lamella of the lumbo-dorsal fascia. It gives

two slips to the lower borders of the twelfth and thirteenth ribs,

of which the former is the larger. It receives a large number of

muscular slips from the outer surfaces of all the ribs, the size

of the latter diminishing from below upwards. Many of the small

muscular slips are continuous with the slips given off from the

longissimus. From the outer border of the muscle long, slender

tendons run to the lower borders of the angles of the first nine

ribs. The longissimus (text-fig. 33, Lo.C) arises from the crest

of the ilium from the highest point to the posterior superior

spine, the posterior sacro-iliac ligament, the back of the sacrum,

all sacral and lumbar vertebral spines and the spines of the

twelfth and thirteenth dorsal vertebre. It gives slips to the

lower borders of ribs 4 to 13 between their angles and the

transverse processes. The attachments to the first three ribs come

from the combined longissimus and ilio-costalis, The combined

outer and middle columns divide into slips which are attached to

the posterior tubercles of the transverse processes of the third,

fourth, and fifth cervical vertebre along with the levator angule

scapule. The third cervical vertebra thus receives splenius

cervicis, levator anguli scapule and longissimus. The part of

longissimus attached to these processes corresponds to longissimus

cervicis in Man, but is not so extensive. The spinalis dorsi

(text-fig. 33, S.C) is a narrow muscle arising from the longissimus

dorsi and the eleventh and twelfth dorsal spines. It is inserted

by muscular and tendinous fibres into the first eight dorsal spines.

Between it and the longissimus is a triangular space into which

the complexus passes and fuses with both. The longissimus capitis

(text-fig. 33, L.Cap) is relatively stronger than in Man. It arises

from the first six dorsal transverse processes by tendinous and

muscular slips. It is fused with the longissimus dorsi, complexus

and scalenus posticus. It breaks up into slips which are inserted

into the posterior tubercle of the atlas and the occipital bone

below the crest. On the atlas its insertion is fused with the

levator scapule and omo-trachelian.

The compleaus (text-fig. 34 A), lying under the longissimus

capitis, has a very extensive origin from the articular processes of

the lower five cervical vertebra, the upper six dorsal transverse

processes and the longissimus and spinalis dorsi. It forms

a number of closely-set parallel muscle bundles which are in-

serted into the inner half of the occipital bone below the superior

curved line. It is not separable into a biventer cervicis and

complexus asin Man. No slip springs from the seventh cervical

spine.

Proc. Zoou. Soc.—1923, No. XXIII, 23

342 DR. C. F. SONNTAG ON THE ANATOMY,

The semispinalis (text-fig. 34 B) has an origin from the arti-

cular processes of the lower five cervical vertebrz by broad bands,

and from the transverse processes of all dorsal vertebree, the lower

five origins being tendinous. It is inserted into the spinous

processes of cervical yertebre two to six by muscle fibres

(semispinalis colli) and to the seventh cervical and first three

dorsal spines by tendons (semispinalis dorsi). It is practically

Text-figure 34.

OCCIPITAL

—

DORSAL TRANSVERSE PROCESSES

Muscles of the back. A: complexus; B: semispinalis and suboccipital muscles.

A.A: posterior arch of atlas; C.N2 and C.N3: posterior divisions of the

second and third cervical nerves ; Lo.C: longus colli; 8.C : spinalis; $.0.T:

suboccipital triangle. Other letters in text.

impossible to distinguish accurately the limits between these

muscles in the combined origins.

A well-marked muscular slip runs from the third articular

process to the transverse process of the atlas and forms a triangle

with the inferior oblique and semispinalis (text-fig. 34, A.O.T).

It alters the course of the second and third cervical nerves. The

mass of the transverse process of the atlas separates it from the

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEL. 343

origin of the rectus capitis lateralis, and the latter is the only

muscle whose fibres run in the same direction *.

The maltifidus spine extends from the sacrum to the axis. It

arises from the sacrum, sacro-iliac ligament, mammillary pro-

cesses of the lumbar vertebrz, transverse processes of all dorsal

vertebree, and lower four cervical articular processes.’ It is

inserted into the spines of all vertebre except the atlas. The

semispinalis covers it but its fibres, which run in the same

direction, are longer than those of the multifidus, and extend

to vertebre farther apart. The cervical fibres of the multifidus

do not form broad bands.

The obliquus inferior (1.0.M) and obliquus superior (S.O.M)

(text-fig. 34 B) are as in Man.

The rectus capitis posticus major (text-fig. 34, R.C.P.M) is a

powerful pyramidal muscle quite concealing the minor muscle.

It arises from the spinous process of the axis and is inserted into

the occiput below the entire length of the superior oblique. The

rectus capitis posticus minor arises from the inner three-quarters

of an inch of the posterior arch of the atlas and is inserted into

the occipital bone below the major muscle. It is quite concealed

by the latter, and by the approximation of the walls of the sub-

occipital triangle. Lectus capitis lateralis is as in Man.

Interspinales, intertransversarli and levatores costarum are as

in Man. Rotatores dorsi are twelve pairs of fan-shaped muscles

running from the transverse processes of the dorsal vertebre to

the laminz of the vertebree above them, the first one being in-

serted into the seventh cervical lamina.

_ Muscles of the Thoracic Parietes.

The external intercostal muscles run in the same direction as

those in Man. They extend from the angles of the ribs to the

sternum in the first three and last two spaces. But there are

external intercostal membranes in the other spaces. The internal

intercostals do not differ materially from those in Man. The

triangularis sternt arises as in Man from the back of the ensiform

cartilage, and it is inserted by radiating slips into the sternal

ends of the second, third, fourth, fifth, and sixth ribs. It has

a slightly different relation to the internal mammary artery from

that in Man, the details being given with that vessel on page 386.

The sternalis muscle is absent. Some authors describe it, and

their observations have been collected by Keith (29).

Contrary to the conditions in Man, there is a well-marked

lateral branch of the first intercostal nerve. It runs over the

pectoralis major and fades away among the glands in the axilla.

Prevertebral Muscles.

The longus colli consists, as in Man, of vertical, superior, oblique

and inferior oblique portions. The vertical part arises from the

* ‘This is probably an individual peculiarity.

23%

344 DR. C. F. SONNTAG ON THE ANATOMY,

lower two cervical and upper four dorsal vertebrae, and is inserted

into the bodies of the second, third, and fourth cervical vertebrz.

The lower oblique portion runs from the first four thoracic to the

fifth and sixth cervical vertebree, And the upper oblique part runs

from the third, fourth, and fifth cervical vertebrz to the anterior

arch of the atlas. ‘The rectus capitis anticus major runs from the

third, fourth, fifth, and sixth cervical vertebre to the basi-

occiput. It receives a well-marked slip from the scalenus anticus.

The rectus capitis anticus minor and rectus capitis lateralis are as

in Man.

It is, therefore, evident that the facial muscles, the muscles of

the back, the scaleni, and the prevertebral muscles are relatively

stronger and more intimately united than in Man. This must

necessarily make the muscular movements less numerous and not

so fine asin him. The muscles forming the suboccipital triangle

are crushed together,

Muscles of the Pectoral Hxtremity.

The pectoralis major, which is less powerful than in Man,

consists of clavicular, costo-sternal, and abdominal parts. The

clavicular part consists of superficial and deep portions. The

former arises from the inner half of the front of the clavicle and

the outer side of the tendon of the sterno-mastoid; the latter

springs from the inner part of the lower surface of the clavicle

and fuses with the former, and with the deep surface of the costo-

sternal part. The costo-sternal portion arises from the whole

length of the body of the sternum and the first six costal

cartilages ; it fuses with both the external oblique and the abdo-

minal part of the pectoralis major at the lower borders of the

fifth and sixth costal cartilages. The abdominal part fuses with

the external oblique as far as the linea semilunaris; it also gets

origins from the lower borders of the fifth, sixth, and seventh

costal cartilages. The abdominal portion joins the deep surface

of the sternal portion to form a muscle which joins the deep

surface of the clavicular part. The combined muscle has a

musculo-aponeurotic insertion into the inner border of the pec-

toral crest, the lower part of the capsule of the shoulder joint

and the deep fascia of the arm. It is never fused with the

deltoid.

T agree with Champneys (11) and Macalister (38) that there is

no actual separation between the clavicular and costo-sternal

parts such as occurs in Man. Champneys records a special slip

arising from the fourth and fifth cartilages, but that is not the

case in my specimen.

The delto-pectoral triangle contains the external anterior

thoracic nerve, thoracic axis vessels, and tendon of the pectoralis

minor, It has no lymphatie glands nor the cephalic vein, which

are present in Man. Bland-Sutton (4) states that the groove

between the pectoralis major and deltoid is absent,

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 345

The pectoralis minor appears to vary considerably both in origin

and insertion. In my specimen it arises by three well-marked

slips from the lower borders of the second, third, and fourth

costal cartilages. Its long, but strong, tendon passes through

a fibrous and synovial sheath over the coracoid process, and it is

inserted into the upper and back part of the capsule of the

shoulder joint. The sheath is adherent to the inner and upper

parts of the coracoid process. Champneys (11) gives its origin

from the first four ribs and its insertion into the capsule of the

shoulder joint close to the supra-spinatus tendon, and Humphry

(26) mentions it as extending across to the great tuberosity of

the humerus. Bland-Sutton (4) describes an insertion similar to

that in my specimen, but gives its origin as ribs three, four, and

five. Wilder (53) and Gratiolet (22) record tendons inserted

into both the coracoid process and capsule of the shoulder joint,

and the latter gives the origin as ribs two to five inclusive. So

if several animals are examined the muscle appears to write its

evolutionary history.

The serratus magnus arises by eleven digitations from the first

eleven ribs, the first one being very small, but it has a remark-

ably rich supply of nerves (text-fig. 48). The first nine arise

from the outer surfaces of the ribs, but the tenth and eleventh

arise from the lower borders. The digitations arising from the

fifth to the eleventh ribs interdigitate with the external oblique.

The muscle is thick at its insertion into the deep aspect of the

vertebral border of the scapula. The part arising from the first

four ribs is much thinner than the remaining part of the muscle.

Champneys (11) has given the origin as from the first ten ribs,

and described the muscle as consisting of three parts, which he

describes in great detail, but Wilder (53) describes two parts,

and gives the origin from all the ribs.

The swbclavius arises from the upper border of the first costal

cartilage immediately internal to the rib. It is inserted into

the under surface of the second quarter of the clavicle from

the inner end. It is enveloped as in Man by the costo-coracoid

membrane.

The costo-coracoid membrane is attached to the first costal

eartilage round the tendon of origin of the subclavius, to the

inferior surface of the sterno-clavicular joint, to the under sur-

face of the clavicle by two layers which enclose subclavius as in

Man, and to the anterior. surface of the clavicle lateral to the

subclavius. The costo-coracoid ligament is well marked. Several

authors have recorded the latter. The clavi-pectoral fascia ex-

tending downwards from the costo-coracoid ligament splits to

enclose the pectoralis minor, and at the same time it sends a

process inwards to the neuro-muscular bundle in the axilla.

External to the pectoralis minor the fascia passes to the deep

fascia of the axilla, and it passes mesially to the deep fascia

covering the serratus magnus between the pectoralis major

and latissimus dorsi. The membrane is pierced by the external

346 DR. C, F. SONN'TAG ON 'THE ANATOMY,

anterior thoracic nerve and the thoracico-acromial vessels, but it

is not pierced by the cephalic vein which runs through it in Man

and many other mammals. The fibres run transversely below

the costo-coracoid ligament.

The deltoid, covered by dense fascia, is coarsely fasciculate. It

arises from the front of the outer half of the clavicle and the outer

border of the acvomion process. A second part has an extensive

wponeurotic and fascial origin from the whole length of the lower

border of the spine of the scapula, and from the fascia over the

entive infra-spinatus. At the inferior angle of the scapula

the fascial origin blends with serratus magnus, the rhomboids,

teres major, and latissimus dorsi. It coneeals a bursa which in-

tervenes between the acromion and upper end of the humerus,

but does not communicate with the shoulder joint. One large

and several small branches of the circumflex nerve, and

branches of the circumflex arteries are seen entering its deep

aspect. Humphry found it adherent to the brachialis anticus

(26), but Macalister (33) denied that it adheres to the triceps

and brachialis anticus. Wilder (53) points out that the attach-

ment to the fascia over the infra-spinatus and the axillary border

of the seapula enables the animal to swing the arm far back.

The muscle fibres all converge to be inserted into the usual

deltoid area on the shaft of the humerus. The insertion is

embraced by the brachialis anticus.

Scapular Muscles :—A\l observers are agreed that the infra-

spinatus greatly exceeds the supra-spinatus in size, and both arise

from the whole of the scapular fosse to which they are attached.

They are inserted as in Man into impressions on the great

tuberosity. Correspons ding vessels and nerves pass into them as

in Man. The infra-spinous fossa is deep, being enclosed between

the prominent spine and a thickening of the axillary border of

the scapula. The teres mtnor arises from the lower border of the

lip of the glenoid cavity and lateral half of the axillary border of

the scapula. It is inserted into the lowest part of the great

tuberosity of the humerus and the upper half inch of the shaft

of the bone. Champneys (11) gives its origin as the mid third

of the axillary border and the adjacent part of the infra-spinous

fossa, and Hepburn (24) records its origin from the upper two-

thirds of the axillary border. The teres major arises from the

medial half of the axillary border of the scapula, and is inserted

into the inner lip of the bicipital groove. It is strongly fused

with the latissimus dorsi. The subscapularis arises from the

whole of the subscapular fossa, and tendinous bands run through

between the bundles of fibres to the bone. It has no origin from

fascia over it. ‘The muscular mass converges, and is inserted by

three tendons into the lesser tuberosity ‘and the shaft of the

humerus over a quarter of an inch below it. Some of the deep

fibres are directly inserted into the capsule of the shoulder-joint.

The coraco-brachialis is fleshy throughout. Jt arises along

with the short head of the biceps from the tip of the coracoid

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE, 347

process, and it is inserted into an impression over an inch long

on the inner aspect of the shaft of the humerus. In its upper

part it is separated by a cellular interval into two parts, and the

musculo-cutaneous nerve passes through the gap. No coraco-

brachialis brevis was present. Some fibres go to the internal

intermuscular septum and dorso-epitrochlearis, and Hepburn (24)

suggests that these represent the coraco-brachialis longus. The

part running to the inner surface of the shaft of the humerus

corresponds to the coraco-brachialis medius. No coraco-brachialis

brevis was recorded by Champneys (11), Bland-Sutton (4),

Dwight (18), and Wilder (53), but it was seen by Macalister (33)

and Hepburn (24). Bland-Sutton gives its insertion into the

upper third of the shaft of the humerus and the capsular liga-

ment. Wilder states that it arises from the coracoid process

through the medium of the short head of biceps.

The biceps arises as in Man, but the bellies remain separate till

they reach the junction of the lower and middle thirds of the

arm. In the upper part of the forearm there is a slight bicipital

fascia (lacertus fibrosus). The muscle fibres of the combined

bellies end in a stout, ribbon-like tendon which is inserted into

the posterior part of the radial tuberosity.

The brachialis anticus is connected by a strong fascial band to

the pectoralis major near its insertion. The origin is as in Man

and embraces the insertion of the deltoid. A slip is given to the

fascia of the forearm. The fibres converge to an insertion into

the coronoid process and inner border of the olecranon.

The dorso-epitrochlearis is a thin muscle, a little more than

half an inch wide, springing from the junction of the muscular

and tendinous parts of the latissimus dorsi. Fibres pass into its

upper part from the coraco-brachialis. It passes into the inner

side of the internal intermuscular septum in the lower third of

the arm, and the latter connects it to the internal condyle.

The triceps differs cousiderably from that in Man. The long

head arises from the dorsal aspect of the outer quarter of the

axillary border of the scapula. The outer head arises from the

upper extremity of the shaft of the humerus and lower part of

the capsule of the shoulder joint. The inner head arises from

the proximal third of the shaft of the humerus along a linear

strip. After a course of two inches the long and outer heads

fuse to form a fleshy belly, and this receives the inner head an

inch more distal. The muscle is inserted by muscular and ten-

dinous fibres into the tip and dorsal surface of the olecranon,

a bursa intervening between the muscle and the capsule of the

elbow joint.

A broad bundle of fibres arising from the deep surface of the

distal half-inch of the triceps runs to the capsule of the elbow

joint and represents the subanconeus.

Champneys (11) states that the triceps, anconeus, and sub-

anconeus are as in Man. Hepburn (24) mentions that the

triceps is as in Man except for the outer head, and mentions

that the anconeus is present.

348 1 DR. U. F. SONNTAG ON THE ANATOMY,

The palmaris longus is absent in both arms in this specimen.

Trail (49) only noted it in one of the arms in his example.

Many authors describe it as being similar to that in Man.

The pronator radii teres has both humeral and coronoid heads

of origin, and the median nerve dips between them. The head

from the internal condyle of the humerus is fused with the origin

of the flexor carpi rvadialis, and the coronoid head is fused with

the deep aspect of the flexor sublimis digitorum. It is inserted

into the middle third of the outer border of the shaft of the

radius, and a few fibres at the upper extremity are inserted into

the inner aspect of the supinator brevis. The upper part of the

insertion is tendinous, but the remainder is muscular. Through-

out its whole length it is fused with fibres of the flexor carpi

radialis. No fibres arose from the dorso-epitrochlearis, but.a few

spring from the extreme distal end of the internal intermuscular

septum. Macalister (33) alone states that there is no coronoid

head of origin.

The flexor carpi radialis has an extremely long origin by a

thick muscular belly, which is intimately fused with the pronator

radil teres and flexor sublimis digitorum. In the second fourth

of the forearm it has no bony attachment, but many fibres run

downwards from the pronator teres and flexor sublimis to its

tendon. Below that it is attached by muscle fibres to the radius

on the inner aspect of the insertion of the pronator teres, and

fibres pass downwards from it to the flexor sublimis. The tendon

receives muscle fibres on its deep aspect till it reaches the annular

ligament. It passes through a tube in the hgament and is in-

serted into the palmar aspect of the bases of the second and third

metacarpal bones. It has a well-marked synovial sheath fused

with the anterior annular ligament.

The flexor carpi ulnaris arises by a narrow head from the

internal condyle, by an expanded head from both interna] condyle

and olecranon, and by fascia from the upper fourth of the shaft

of the ulna. It is inserted into the pisiform bone. It is rela-

tively larger than in Man, and the pisiform is very large.

The flexor sublimis digitorum appears to differ considerably.

In my specimen it has a very extensive origin from the humerus,

ulna, and radius; and fibres spring from those of the other flexor

muscles. The humeral head, arising from the internal condyle

of the humerus, fuses with the flexor carpi radialis, and receives

a few fibres from the flexor carpi ulnaris. It is prolonged almost

entirely into the tendons for the ring and little fingers, the

tendon for these digits separating off in the middle of the fore-

arm. The coronoid head is fusiform and quickly ends in a tendon

which forms the slip to the index finger. The second finger gets

its tendon from a muscle which arises from the lower two-thirds

of the shaft of the radius and from the flexor carpi radialis.

Hepburn (24) states that the tendons to the third and fourth

digits come from the radial part of the muscle, while those for

the second and fifth come from the ulnar part; and the tendon

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 349

for the index goes under those for the second and fourth digits.

Champneys (11) states that the middle finger alone has a separate

radial origin, but Rolleston gives the radial origin to the tendon

for the index. Macalister (83) observed no radial origin at all.

Moore (36) states that the annularis receives two tendons, and

the minimus gets none. Finally, Bland-Sutton (4) describes the

flexor sublimis tendons going to the third, fourth, and fifth digits,

but there is a flexor sublimis indicis arising from both radius and

coronoid. Dwight (18) describes a very complex muscle. The

tendons split over the heads of the metacarpal bones, surround

the deep flexor tendons, and are inserted as in Man into the

middle phalanges.

Muscles of the Hypothenar Eminence (text-fig. 35 A & C):—

The abductor minimi digiti (A.M.D) arises by a broad, but thin,

muscular origin from the pisiform bone. It lies along the ulnar

border of the hand and is inserted by a long, slender tendon into

the ulnar aspect of the base of the first phalanx of the little

finger. Its insertion is closely blended with that of the flexor

brevis. The flewor brevis minimi digiti (F.B.M.D) has a single

head of origin from the anterior annular ligament and hook of

the unciform; and the annular ligament appears to be prolonged

into it. It is inserted along with the abductor. The opponens

minimi digiti (O.M.D) has a double origin from the anterior

annular ligament and uncinate process of the unciform, the latter

being blended with the flexor brevis. It is inserted into the ulnar

aspect of the shaft of the fifth metacarpal bone. The palmaris

brevis is very extensive in both hands, but several authors

describe it as in Man.

Muscles of the Thenar Eminence (text-fig. 35 A & C):—The

abductor pollicis brevis (A.P.B) arises from the anterior annular

ligament, the scaphoid and sesamoid bone and the sesamoid bone

of the thumb, and it is divided into two slips. These unite to a

muscular insertion into the radial side of the base of the first

phalanx of the thumb. It conceals the lateral half of the opponens.

Champneys(11)and Macalister (33)saw no splitting into Seemmer-

ing’s slips. The opponens pollicis (O.P) arises from the anterior

annular ligament and ridge of the trapezium. It is inserted into

the distal half of the radiai aspect of the shaft of the metacarpal

of the thumb. Embleton (19) states that the opponens pollicis is

absent. The fleaor brevis pollicis (¥.B.P) consists of superficial and

deep parts. The superficial part arises by three slips from the

anterior annular ligament and trapezium. The deep part arises

from the ulnar side of the first metacarpal and anterior annular

ligament. Both parts unite and are inserted into the ulnar side of

the base of the first phalanx. The whole muscle forms a large

mass between the opponens superficially and the adductores deeply.

The adductor transversus pollicis (A.T,P) and adductor obliquus

pollicis (A.O.P) are as in Man.

The flexor longus pollicis (¥.L.P) arises from the inner surface

of the shaft of the radius over the whole length except the

350 DR. C. F. SONNTAG ON THE ANATOMY,

proximal inch, and from the interosseous membrane. Some

fibres fuse with the flexor sublimis. It has a strong tendon to

the palmar aspect of the terminal phalanx of the index, and a

very fine tendon to the corresponding part of the pollex. This

agrees with the descriptions of Champneys (11), Hepburn (24),

and others.

The fleaor profundus digitorum arises from the inner side of

the olecranon, the inner surface of the upper two-thirds of the

shaft of the ulna and the interosseous membrane; and some of

the fibres fuse with the flexor sublimis and flexor carpi ulnaris.

The strong tendon divides into three tendons running to the

palmar aspects of the terminal phalanges of the third, fourth,

Text-figure 39.

y p) SEMILUNAR

F.D.I.- i q AE VAG

PALMAR 6 Hill ENN nN AT

inTeR- 5h) aN

IH) Han

A.0.P., OSSEI 4 oan Ki Wii

ATP. EBRD. 2— MMT EA

ie Sig STFFN ONAL

F.B.P-wwes \ ) TDR

==>

LP. ~\ a vp |

i YH I ZANT. ANNULAR

LIGAMENT

»R. F.C.U.

AON Hi

suey :

i ff A.M.D.

OP FOR. pgm. y

‘The muscles and joints of the hand. F.B.P.D: deep part of the flexor brevis

pollicis; F.C.R: flexor carpi radialis; F.C.U: flexor carpi ulnaris; F.D.I:

first dorsal interosseous muscle; F.P.D: flexor profundus digitorum ; F.S.D:

flexor sublimis digitorum. Other letters in text.

and fifth digits. There is no continuity with the flexor longus

pollicis. Several authors have described similar conditions.

The pronator quadratus runs downwards and outwards from

the lower inch and a half of the front of the shaft of the ulna

to the lower inch of the front of the shaft of the radius. Fibres

wrap round both bones and extend down to the interosseous

membrane.

Lumbricales:—These arise as in Man, and the general dis-

position is similar, but a well-marked muscular slip connects the

first and second. ‘The first and second muscles have long origins

from the flexor tendons, but the second and third muscles quickly

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE, 351

separate from the tendons, The imsertious are as in Man.

Hepburn (24) and Macalister (33) describe them as in Man.

Wilder (53), Dwight (18), and Champneys (11) state that the

tendon for the minimus arises from the profundus tendon to

the annularis.

The supinator longus arises from the external supracondylar

ridge and from the shaft of the humerus as high up as the in-

sertion of the deltoid. Some fibres come from the external

muscular septum, and some from the brachialis anticus. It has

a long tendon which is inserted into the shaft of the radius half

an inch above the styloid process. SSome authors have recorded

‘slightly less or a little more extensive origin and insertion. This

animal agrees in this respect with Hepburn’s account (24).

The extensor carpi radialis longior arises from the lower part

of the external supracondylar ridge and septum. Its tendon

separates very high up in the forearm, passes under the extensors

of the thumb, and is inserted into the radial side of the dorsal

aspect of the index metacarpal, and along the radial aspect of the

proximal half inch of the bone. This insertion is more extensive

than in some accounts.

The extensor carpi radialis brevior arises from the lateral

epicondyle, the external lateral ligament of the elbow jomt and

the fascia over the extensor communis digitorum. It is slightly

fused with the long extensor. It is inserted into the dorsal

aspect of the base of the third metacarpal by three small

tendons.

The extensor communis digitorum arises from the external

epicondyle, the fascia over it, and the intermuscular septa on

either side. It remains fleshy to the posterior annular ligament.

The origin from the internal septum is particularly strong. It

is quite separate from the subjacent extensors. It separates into

three broad tendons to the index, medius, and annularis, and a

slender tendon goes to the minimus. Close to the heads of the

‘metacarpals there is strong lateral fusion between the tendons to

annularis and minimus. The tendons are inserted into the bases

of the ungual phalanges. Wilder (53), Vrolik (51), Moore (36),

and Macalister (83) deny the i: esence of a tendon to the minimus.

Dwight (18) and Champneys (11) say there is a slip between the

- tendons to annularis and minimus. The tendons have very

powerful thickened expansions into the sides of the inter-

phalangeal joints.

The Cn eieso: minima digitt has a long, slender tues belly

enclosed in a strong fascial tunnel. It arises from the fascia

‘over the anconeus, and from the common extensor origin from

the external epicondyle. A common dorsal expansion unites its

tendon to the innermost communis tendon over the head of the

fifth metacarpal. he expansion 1s very firmly adherent to

the capsule of the metacarpo-phalangeal joint. The insertion is

into the base of the ungual phalanx of the minimus.

~The extensor carpi ulnaris is as in Man. Several authors

deseribe this.

352 : DR. C. F. SONNTAG ON THE ANATOMY,

The extensor indicis arises from the inner surface of the lower

fifth of the radius, and some fibres blend with the extensor longus

pollicis. Its long, and very slender tendon blends with the

dorsal expansion of the communis tendon to the index over the

first phalanx. No slip goes to any other digit, as in Wilder’s

specimen (53). Hepburn (24), Macalister (33), and Humphry (26)

found it supplying the medius too.

The supinator brevis is wrapped round a little more than the

upper third of the radius. It is musculo-tendinous.

The extensor ossis metacarpi pollicis and extensor primi inter-

nodii pollicis have a common origin from the bones of the forearm.

The latter arises from the upper third of the lateral border of the

ulna, and the former from the upper two-thirds of the mesial

border of the radius. The tendons separate from the combined

muscular mass. The broad tendon of the former runs to the

trapezium and thumb sesamoid, and the slender tendon of

the latter goes to the base of the metacarpal of the thumb.

The extensor secundi internodii pollicis (extensor pollicis longus)

arises from the third fourth of the inner surface of the shaft of

the ulna below the extensor primi internodii pollicis and above

the extensor indicis. Its long, ribbon-like tendon is inserted

into the base of the ungual phalanx of the thumb. Hepburn (24)

gives its insertion as the base of the first phalanx, but Humphry

(26), Macalister (33), Vrolik (51), Wilder (53), and Wyman (54)

recorded conditions as in my specimen.

Interossei :—All authors agree that the dorsal interossei are

as in Man, and several have described the six interossei on the

palmar surface of the manus. Hepburn (24) has shown that

three of the six muscles are the true palmar interossei, namely,

those to the ulnar side of the index and the radial sides of the

annularis and minimus. The others to the sides of the medius

and ulnar side of the annularis are abductors, belonging really to

the dorsal series. With his observations I am quite in agreement.

The six palmar muscles form a very thick stratum. The con-

ditions are shown diagrammatically in text-fig. 35 A. The first

dorsal interosseous wraps round the metacarpal of the index.

Taking the deep muscles from within outwards, we find :—

1. Opponens minimi digiti; 2. palmar adductor interosseous

to the minimus; 3. palmar abducting interosseous of the annu-

laris ; 4. palmar adducting interosseous of the annularis ; 95. pal-

mar interosseous deviating the medius to the ulna; 6. palmar

interosseous deviating the medius to the radius; 7. palmar ad-

ducting interosseous of the index; 8. pollical head of the first

dorsal interosseous covering the metacarpal of the index;

9. deep head of the flexor brevis pollicis.

Muscles of the Anterior Abdominal Wall.

The external oblique arises by well-marked digitations from the

outer surfaces and lower borders of ribs 5-11. The mesial

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 353

digitations fuse with the pectoralis major, aud the outer ones

are only covered by the latissimus dorsi. All interdigitate with

the serratus magnus. The lateral fibres descend to the anterior

superior iliac spine and outer third of Poupart’s ligament. The

other fibres end in the aponeurosis which is attached to the

sternum, the last chondro-sternal junction, the pubis, and the

inner two-thirds of Poupart’s ligament. The aponeurotic fibres

of the two sides cross the mid line into one another. Gratiolet

(11) described the aponeurosis in detail, recording the characters

of Poupart’s ligament, the slight Gimbernat’s ligament, the lax

adhesion to the deep fascia of the thigh, and the formation of the

pillars of the external abdominal ring. The aponeurosis fuses

with the internal oblique mesial to the splitting of the apo-

neurosis of the latter. And the crural fascia and Poupart’s

ligament fuse with the aponeurotic origin of the sartorius.

The internal oblique rises from the outer half of the anterior

border of the iliac crest, the anterior superior iliac spine, the

outer third of Poupart’s ligament, and the lower borders of costal

cartilages 10-13. The aponeurosis, which receives the fibres.

has a curved line of splitting and runs from the tenth costal

cartilage to the inner end of Poupart’s ligament.

The transversalis abdominis arises from the deep surfaces of

ribs 10-13, the lumbar fascia, the anterior quarter of the inner

lip of the iliac crest, the inner surface of the anterior superior

iliac spine, and the outer third of Poupart’s ligament. The

aponeurosis is attached to the xiphoid and pubis.

The sheath of the rectus is as in Man, and the semilunar fold of

Douglas is present. The rectus has two origins, as in Man, but

has four inscriptions running right through it to the sheath

Pyramidalis is absent.

The diaphragm has a comparatively small central tendon

receiving muscles arising from ribs 7-13 and interdigitating with

the transversalis abdominis, from the back of the sternum by two

slips, and from the lumbar vertebre by the crura and extra

slips. The right crus arises as low down as the second lumbar

vertebra, and the left one from the first. A slip arises from the

transverse process of the second lumbar vertebra and one from

the side of the body of tke first. The lumbo-costal arches are

as in Man.

The quadratus lumborum is related to the lumbar fascia as in

Man. It arises from the posterior two-thirds of the inner lip of

the iliac crest, where it is continuous with the iliacus, and from

all the lumbar transverse processes. It is inserted into the inner

four-fifths of the last rib and the bodies of the last two dorsal

vertebre.

Muscles of the Pelvic Extremity.

The psoas parvus arises by fleshy fibres from the last dorsal

and first lumbar vertebre, and it is connected by a fascial sheet

over the psoas magnus to the remaining lumbar transverse

354 DR. C. F. SONNTAG ON THE ANATOMY,

processes. Its broad, flat tendon is attached to the ilio-pectineal

line close to the emergence of the femoral vessels.

The tliacus arises as in Man. It is quite continuous with the

quadratus Jumborum, and it soon fuses with the psoas magnus.

{ts fibres envelop the psoas from each side. The psoas magnus

blends more with the iliacus thanin Man. It arises from the

last dorsal vertebra, the inner inch of the last rib and the bodies

and transverse processes of all the Jumbar vertebre. The com-

bined muscle is inserted into the small trochanter and the.

femoral shaft a little below it. Half of the muscle (mesial part)

passes over the ilio-psoas tendon and is inserted into the bone

posterior to the latter.

The saréorius has a large fan-shaped aponeurotic origin from

the anterior edge of the ilium up to a point a finger’s breadth

below the anterior superior spine. Superficial to the aponeurosis,

but connected to it by fascia, is a long, narrow fascial strip

connecting the muscle to the anterior superior iliac spine. The

strip is connected to the crural fascia and fascia over the

abdominal muscles, thus forming a tunnel for the ilio-psoas.

The comparatively slender muscle is inserted into the upper

third of the anterior berder of the tibia from the attachment of

the ligamentum patelle downwards. Strong fascia unites it to

the mner tuberosity of the tibia and the internal lateral ligament

of the knee. Between it and the subjacent gracilis is the

saphenous nerve, but no bursa.

The gracilis. and adductor longus (text-fig. 36, Gra. and App.

Lone) arise by a common aponeurosis from the inner end of

Poupart’s ligament, the entire length of the side of the symphysis,

and upper third of the descending ramus of the pubis. No other

author describes a precisely similar origin. And the origin

conceals the adductor magnus. The adductor longus occupies

the greater part of the aponeurosis, and its fibres approach closer

to the bones; it is inserted into the third quarter of the back of

the shaft of the femur, and it is fused with the magnus.

Hepburn (24) describes it as arising by a rounded tendon, and

Humphry (26) gives its origin as the spine and inner half of the

horizontal ramus of the pubis. The gracilis is inserted into

the inner aspect of the tibia behind the internal lateral ligament.

It is fused with the subjacent semitendinosus and the fascia over

the inner head of the gastrocnemius. Gratiolet (22), Champneys

(11),and Hepburn (24) & give more extensive origins for a separate

gracilis.

The adductor magnus (text-fig. 36, ApD.Mae) arises by three

heads from the pubis and ischium, and it is inserted into the

upper three-quarters of the back of the shaft of the femur. The

upper head arises from the entire length of the body of the

pubis: the middle head from the arch and ischial tuberosity :

and the lower head from the tuberosity. The upper and mid

heads unite to form a thick muscle inserted into the femoral

shaft. The lower head runs separately to the adductor tubercle,

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 355

and the femoral vessels pass between the two parts to the

popliteal space. Champneys (11) reviews the various descriptions.

Dwight (10) describes fusion of all the adductors except the part

of the magnus to the tubercle. Hepburn (24) states that the

latter is really a hamstring, and is supplied by the sciatic

nerve. I did not observe fibres from the main mass reaching

the knee.

Text-figure 36.

ASG

. \_ ADD. BREV.

ER But

LLM

XSAN aN

SBor: Z

SCHIUM

Muscles and joints of the pelvic extremity. A.C.L: anterior cruciate ligament ;

C.T: fibrous connecting slip; E.L.H: extensor longus hallucis; E.S.C:

external semilunar cartilage; JI.C.B: internal cuneiform bone; I.8.C:

internal semilunar cartilage; Met.I: first metatarsal bone; N.B: scaphoid

or navicular; P.C.L: posterior cruciate ligament; PYR: pyriformis; T.A :

tibialis anticus. Other letters in text.

The adductor brevis (text-fig. 36, ADD.BREv) arises by three

prismatic, interlocking bellies from the angle between the body

and horizontal pubic ramus. Its flat tendon is inserted into a

line from the small trochanter to the middle of the back of the

femur.

The pectineus (text-fig. 36, PEcr) arises from the pubic crest on

356 DR. G. F. SONNTAG ON THE ANATOMY,

the three-quarters of an inch internal to the longus. It hasa

curved insertion running from the lesser trochanter to the back

of the shaft of the femur.

Hamstring Muscles :—With the exception of the short head of

the biceps, all the hamstrings arise together from the lower and

back part of the tuber ischii, and all are fused. The semtendi-

nosus is inserted into the anterior tubercle of the tibia, and by a

large expansion to the fascia of the leg. The tendon is not as

long as in Man (Vrolik and Hepburn), and the insertion is lower.

Cuvier showed that this low insertion is incompatible with

an erect attitude, and Rolleston pointed out that it oceurs

in children. The insertion of this muscle, and that of the

semimembranosus move upwards as the body becomes erect.

The semimembranosus is smaller than the last muscle, and its

long, flat tendon sends no fibres to the fascia over the popliteus,

nor to the internal lateral ligament. It is inserted into the tibia

over a small area proximal to the other hamstrings. The bzceps

differs somewhat in my specimen from the accounts of Hepburn

(24) and Champneys (11). Both heads remain separate. The

ischial head has a strong insertion into the outer side of the head

of the tibia, the head of the fibula, and the fascia over the outer

head of the gastrocnemius, The femoral head is inserted into

the head and proximal inch of the shaft of the fibula, and the

fascia over the gastrocnemius. It is, therefore, evident that the

hamstrings and some of the adductor group are connected to

the fascia over the gastrocnemius.

The gluteus maximus is smaller than in Man. It arises from

the side of the sacrum and coccyx, the great sacro-sciatic ligament

and the ischial tuberosity along with the long head of the biceps.

No muscle fibres arise from the iliac crest, as in Phascolarctos,

but it arises from the strong fascia which covers the gluteus

medius, and is attached to the iliac crest. Hepburn (24) saw it

arising from the crest, Humphry (26) observed no fascial origin,

and Champneys (11) described conditions similar to mine. The

insertion is longer than in Man, for it is fixed to the back of

the femur as low down as the external condyle, and to the

shaft below the great trochanter. Its fibres mingle with the

vastus externus. It is fused with the outer head of the gastro-

enemius (Humphry, 26), and with the tensor fasciz femoris

(Wilder, 53).

The gluteus medius has a fleshy origin from the whole of the

dorsum ilii down to the line from the great sciatic notch to

the anterior inferior spine, and by a devse aponeurosis from the

anterior border between the superior and inferior spines. The

aponeurosis gives way to muscle after an inch. Fibres also arise

from the posterior aspect of the aponeurosis. It is inserted into

the top of the outer aspect of the great trochanter. A small slip

runs from the mesial aspect of the tendon to join the tendon of

the pyriformis, thus bringing the two tendons into connection.

Two communicating burse (text-fig. 36, Bur) separate the

PHYSIOLOGY, AND PATHOLOGY OF THU CHIMPANZEE. DDT

tendons of gluteus medius from those of the scansorius and

pyviformis. It is the largest of the glutei. The whole insertion

is by closely-set ribbon-like muscles. None of these forms a

separated part as described by Champneys (11), but the insertion

is split slightly by the vastus externus, as described by

Hepburn (24).

The scansorius arises from the body of the ilium by a thick

muscular origin, and by a thin curved origin from the ilium an

inch behind and parallel to the acetabulum. The most external

fibres come into relation with the gluteus minimus. It passes

over the acetabulum and head of the femur to be inserted into

the capsule of the hip from the head to the top of the great

trochanter, and to the anterior border of the great trochanter

internal to the pyriformis, gluteus medius and vastus externus.

It is continuous above with the obturator internus and the

gemelli, and below with the gluteus minimus.

The gluteus minimus arises from the anterior border of the

ilium from the anterior superior spine to below the anterior

inferior spine. It is on a slightly more anterior plane than the

scansorius. Its insertion is linear and continues that of scansorius

downwards for a centimetre. The insertion is covered by the

vastus externus.

The tensor fascie femoris arises from the back of the iliacus,

and is inserted into the fascia of the leg more’ than half-way

down the leg. Some authors have stated that the scansorius

corresponds to the tensor fasciz, and others describe only the

former. Champneys (11) describes both.

The gemellus superior (text-fig. 36,, GEM.SuP) arises from a

small area on the outer surface of the ischial spine, and from the

attachment of the lesser sacro-spinous ligament above the groove

for the tendon of the obturator internus. The gemellus inferior

(text-fig. 36, Grem.INF) arises within the pelvis from a linear

origin, and from the top of the outer aspect of the tuber ischii

below the groove for the tendon of the obturator internus. The

tendon of the obturator internus (text-fig. 36, Opr.Inr) formed by

a fusion of fascicles, is separated from its groove in the ischium

by a bursa. ~ It is overlapped by the gemelli which fuse over it.

They all have a common insertion into the femur above the

trochanteric pit. It is intimately connected to the capsule

of the hip joint. Between the hip joint and the tendons of

obturator internus and gemelli and the quadratus femoris and

obturator externus there is a pad of fat. Hepburn (24) states

that the gemellus superior is less than the inferior, and it is

difficult to separate the latter from the quadratus femoris, but

that is not the case in this animal. Champneys (11) agrees with

me in the relative sizes of the gemelli.

The quadratus femoris arises from the inferior ramus and

upper edge of the tuber ischii above the origin of the hamstrings.

It has an angular insertion into the lower part of the inter-

trochanteric crest and lesser trochanter, and then horizontally

Proc. Zoou. Soc.—1923, No. XXIV, 24

308 DR. C. F. SONNTAG ON THE ANATOMY,

outwards for an inch into the upper end of the insertion of the

adductor brevis.

The obturator externus (text-fig. 36, Opr.Exr) arises by two

heads, The large head arises from the inferior ramus of the

pubis between the obturator foramen and the origins of the ad-

ductor magnus, and from the obturator fascia. The small head

arises frem the horizontal ramus of the pubis between the

obturator canal and origin of the lateral head of the adductor

brevis. The two heads unite, and the tendon is inserted into the

trochanteric pit and capsule of the hip joint.

There is greater connection between these muscles, and the

capsule of the hip joint is considerable.

The rectus femoris arises by two heads which, however, are not

very distinctly separate. The straight head arises from the

anterior aspect of the ilium between the anterior inferior spine

and the acetabulum. The retlected head forms an arch over the

whole of the upper part of the acetabulum, and the upper fibres

are connected by a dense aponeurosis with the iliacus. The

vastus externus arises from a small area on the antero-lateral

aspect of the great trochanter below the insertion of the scan-

sorius, whose tendon it splits. The origin is continued down

on the back of the femur, anterior to and continuous with the

gluteus maximus, to an inch above the external condyle. A small

part, arising from the anterior part of the bone below the great

trochanter, is at first separated from the main mass by the

gluteus minimus. And an additional slender belly springs from

the upper end of the intertrochanteric line. The vastus internus

arises from the intertrochanteric line except its extreme upper

end. And it springs from the back of the femur down to a point

an inch above the internal condyle. The crwreus arises from the

upper two-thirds of the surface of the femur between the two vasti.

The quadriceps extensor tendon is wide and receives the muscles

an inch above the patella. It is attached to the upper border of

the patella, the capsule of the joint on either side of it, the

internal condyle of the tibia, and the outer femoral and tibial

condyles. The ligamentum patelle is inserted into the front of

the upper end of the tibia. No suberwreus is present.

Tibialis Anticus :—Macalister (33), Hepburn (24), Champneys

(11), and others have described double origins and insertions,

The insertions are into the entocuneiform and first metatarsal.

Wilder (53) points out that the double insertion is in accord with

the use of the hallux asa thumb. In this animal the origin is

from the outer aspect of the outer condyle and the upper half of

the outer aspect to the tibia, from the middle third of the

delicate interosseous membrane, from the fascia between it and

the extensor longus digitorum, and from the fascia over it. The

lateral superficial fibres form the superficial belly of the muscle,

which is inserted into the base of the first metatarsal bone.

The other fibres form the deep belly, which runs to the internal

cuneiform (text-fig 36 B). The superficial belly has a mucous

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 359

sheath almost to its insertion; but the deep belly loses its sheath

on the dorsum of the foot, and is separated from the cuneiform

bone by a bursa.

The evtensor longus digitorum arises from the external condyle

of the tibia internal to the head of the fibula, the anterior border

of the head of the fibula, the antero-medial surface of the fibula

to within an inch of the malleolus, the anterior peroneal inter-

muscular septum to the same level, the fascia between it and the

tibialis anticus and the fascia over its upper half. The belly

passes through the lateral compartment under the annular

ligament, and divides into four slips. The second slip is thready,

but the others are well developed. The first slip divides into

two; the medial one going to the dorsum of the second toe, and

the lateral one joins the second slip and runs to the third toe.

The third and fourth slips run to the fourth and fifth toes. The

slips form dorsal expansions over the metacarpo-phalangeal joints

and proximal phalanges, which are joined by the lumbricales,

interossei, and extensor brevis, except in the case of the fifth toe.

The actual insertions into the bones are as in Man.

The extensor longus hallucis arises from the middle third of the

antero-medial surface of the shaft of the fibula, and from the

outer part of the interosseous membrane, posterior to the extensor

digitorum longus. The belly passes through the middle com-

partment under the anterior annular ligament, and continues as

a tendon round the inner surface of the entocuneiform. It runs

through the naviculo-metatarsal trochlea (text-fig. 836 B) and

reaches the dorsum of the metatarsal of the hallux. A dorsal

expansion is formed over the first phalanx, the proximo-lateral

part of which joins the tendon of insertion of the most medial

tendon of the extensor brevis digitorum. The rest of the tendon

has an expanded insertion into the base of the terminal phalanx

of the hallux and the capsule of the interphalangeal joint.

The peroneus tertius is absent.

- The gastrocnemius has two heads, but they arise more pos-

teriorly and distally than in Man, and from the capsule of the

knee joint instead of from bones. They spring from the capsule

over the articular surface. They are inserted into a median

tendinous raphé, the inner belly slightly overlapping the external,

and extending more laterally. The flattened tendon joins the

tendon of the soleus, half an inch above its insertion into

the calcaneus, forming the tendo Achillis. The edges of the

muscle are firmly connected to the subjacent soleus.

The plantaris arises lower down than in Man, from the postero-

lateral side of the external femoral condyle. The slender belly,

three inches long, passes under the lateral belly of the gastro-

cnemius, and the very fine thread-like tendon has an expanded

insertion into the tendo Achillis close to the calcaneus.

The soleus has no tibial origin, and is smaller than in Man. It

has a fleshy origin from the posterior aspect of the head of the

fibula, and an aponeurotic origin from the upper part of the

24*

360 DR. C. F, SONNTAG ON THE ANATOMY,

peroneal intermuscular septum. The flat belly gradually expands

till it reaches a point an inch above the calcaneus. ‘The superficial

part of the central portion becomes tendinous in the lower third,

and is joined by the gastrocnemius to form the tendo Achillis.

The tendo Achillis is inserted into the middle of the posterior

aspect of the os calcis. It is separated from the upper part of the

bone by a bursa and a pad of fat.

The popliteus is double, but its origins and insertions are all in

contact to form linear attachments. The proximal fibres rise

from the posterior aspect of the capsule of the knee-joint internal

to the outer head of the gastrocnemius. The distal fibres arise as

in Man. The proximal part is inserted into the vertical line on

the posterior aspect of the internal tibial condyle. The distal

part goes to the oblique popliteal line, the posterior border of the

subcutaneous area, the internal condyle and a curved line run-

ning round the internal condyle.

The flexor longus digitorum arises from the popliteal line and

its medial continuation downwards to within half an inch of the

inner malleolus, and from the septum between it and the tibialis

posticus. The tendon passes round the back of the internal

malleolus and under the inner end of the internal annular

ligament. It crosses to the lateral side of the sole of the foot

and is joined by the tendon of the aecessoriusand a slip from the

flexor longus hallucis. A small slip runs to the lateral tendon of

the flexor brevis digitorum. Then it divides into five tendons

for the four inner toes, the fourth receiving two. The first and

second tendons, to the second and third toes, are accompanied

superficially by tendons of the flexor brevis. In the case of the

fourth toe the superficial tendon of the longus replaces the

tendon of the brevis. The tendons are inserted as in Man.

The tendon to the little toe is attached by a broad vinculum to

the body of the proximal phalanx, by a triangular vinculum to

the second phalanx and is inserted into the base of the terminal

phalanx. The flexor sheaths are as in Man.

The four lumbricales pass to the inner parts of the dorsal

expansions of the extensor tendons of tho four inner toes.

The accessorius is very small and has only the external head

which runs from the outer edge of the calcaneus and the long

plantar ligament to the outer aspect of the flexor longus

digitorum.

The flexor longus hallucis arises from the lower part of the

posterior aspect of the head of the fibula, from the posterior

surface of the fibula almost to the malleolus, from the fascia

between it and the tibialis posticus, from the posterior peroneal

intermuscular septum, the lower part of the interosseous mem-

brane and slightly from the tibia near the interosseous membrane.

The tendon is inserted into the base of the terminal phalanx of

the hallux, and the capsule of the interphalangeal joint. In the

sole a large tendon connects it to the flexor longus digitorum.

It has a flexor sheath. The tendon runs through a trochlea

attached to the base of the first metatarsal.

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE, 361

The tibialis posticus arises from the lower part of the head and

upper part of the body of the fibula anterior to the flexor longus

hallucis, from the upper half of the interosseous membrane, the

upper half of the tibia external to the popliteal line and its

downward continuation and from the septum between it and the

long flexors. It is inserted much as in Man, only the scaphoid

here has no definite tubercle.

The peroneus longus arises from the anterior half of the outer

surface of the head of the fibula, the antero-lateral surface of the

fibula to within an inch of the malleolus, the upper half of the

anterior peroneal septum, the posterior peroneal septum and

the fascia over it. The tendon passes behind the outer malleolus

and external to the peroneus brevis, and from there on it is as

in Man.

The peroneus brevis arises from the lower half of the antero-

lateral surface of the fibula down on to the lateral malleolus, the

anterior peroneal septum and the fascia over it. The tendon

goes behind the outer malleolus and is inserted into the projecting

base of the fifth metatarsal. An inch below the malleolus the

tendon gives a slip to the dorsal expansion on the dorsum of the

proximal phalanx of the fifth toe.

The extensor brevis digitorum arises from the outer side of the

upper surface of the os calcis anterior to the posterior facet, the

dorsal caleaneo-cuboid ligament and the lower border of the ex-

ternal annular ligament. Its four fleshy bellies end in slender

tendons to the four inner toes. The most medial tendon has an

expanded insertion into the base of the dorsum of the proximal

phalanx of the hallux. The other three join the dorsal extensor

expansions, thereby being inserted into the bases of the second

and third phalanges.

The flexor brevis digitorum arises as in Man. It divides into

two tendons, the lateral one to the third toe being joined by a

slip from the long flexor. ‘The medial one goes to the second toe.

Insertion as in Man.

The abductor hallucis arises as in Man, and it also receives

fibres from the inner side of the foot (internal annular ligament,

scaphoid, entocuneiform and capsule of the metatarso-phalangeal

joint). It is inserted into the inner aspect of the capsule of the

first metatarso-phalangeal jointand the base of the first phalanx,

with the interposition of a sesamoid bone. A small slip passes

to the shaft of the first phalanx.

The abductor minimi digiti arises as in Man. It is inserted

into the outer side of the projecting base of the fifth metatarsal

(abductor ossis metatarsi quinti of Hepburn), the inner side of

the base of the fifth metatarsal, the outer side of the plantar

aspect of the capsule of the metatarso-phalangeal joint and base

of the proximal phalanx, and the outer side of the shaft and the

dorsal expansion on the first phalanx. Hepburn (24) mentions

the latter in the Orang.

The flexor brevis hallucis has two bellies. The deep inner one

362 DR. C. F. SONNTAG ON THE ANATOMY,

arises from the entocuneiform, the sheath of the peroneus longus

tendon, the capsule of the first metatarso-phalangeal joint, and the

lower half of the outer border of the first metatarsal. The outer

belly arises from the sheath of the peroneus longus and the

external long plantar metatarsal ligament where it is attached to

the outer surface of the base of the fourth toe. No fibres

arise from the cuboid. The two bellies together with the two

adductores hallucis are inserted into the outer side of the base of

the first phalanx of the hallux and the capsule of the joint with

the interposition of a sesamoid. The internal head is really a

first interosseous.

The adductor obliquus hallucis arises from the base of the third

metatarsal and the proximal half of the external long plantar

metatarsal ligament. The adductor transversus hallucis arises

from the upper half of that ligament and from the internal long

plantar metatarsal ligament, and the capsules of the second

and third metatarso-phalangeal joints.

The flexor brevis minimi digiti arises from the plantar aspect of

the fifth metatarsal bone and the sheath of the tendon of the

peroneus longus. It is inserted into the capsule of the fifth

metatarso-phalangeal joint and outer side of the base of the

proximal phalanx.

The opponens minimi digiti arises from the proximal half of

the plantar surface of the fifth metatarsal and the sheath of the

peroneus longus. It is inserted into the inner side of the

capsule of the metatarso-phalangeal joint. There are four

dorsal and three plantar interosset, but Dwight (18) mentions a

large number. The four dorsal interossei arise as in Man; but

their insertions differ in that they abduct from a line drawn

through the middle toe. This resembles the arrangement

in the hand, but not in the human foot where the mid line runs

through the second toe. Hepburn (24) records the line as

resembling that in the human foot, but Champneys (11) and

Cunningham (13) point out that in all Primates except the

Gorilla and Lemur the interossei abduct the toes from a line

through the middle one. ‘The attachments of the dorsal inter-

ussei are as follows :—

No. 1 inserted into the inner side of the second digit.

2 9 99 9 ye 99 2? 99 third 9

3 ” ” a outer ” I) oD ”

A 29 99 9? 39 99 He) 99 fourth 99

The three plantar interossei adduct towards the middle toe

instead of the second asin Man. The first arises in the second

interspace from the whole length of the outer side of the second

metatarsal, and a few fibres arise from the base of the third

metatarsal, as Hepburn (24) finds in the Gorilla. It is inserted

into the outer side of the second toe. The second arises from

the inner side of the fourth metatarsal and the ligamentous

structure covering the bone. It is inserted into the inner side

PHYSIOLOGY, AND PATHOLOGY OF 'HE CHIMPANZEE. 363

of the same toe. The third is placed more superficially than

the others, and lies in the same plane as the opponens minim1

digiti. Its belly crosses the fourth space from its origin on the

external long plantar metatarsal ligament, and the ligaments

over the base of the fourth metatarsal to its insertion on the

inner side of the fifth toe.

The dorsal interossei are inserted more into the proximal

phalanx than into the dorsal expansion, but the plantar inter-

ossel exhibit the reverse.

The important points to be noted from the physiological point

of view are:—1. The middle line of the foot runs through the

middle toe instead ef through the second, as in Man. So the

interossei of the foot of the Chimpanzee act as they do in a hand.

2. There are no additional plantar interossei in the foot as there

are in the hand.

The disposition of the muscles in four layers in the sole of the

foot is as in Man.

The nature of the terminal part of the pelvic extremity is

described on page 423,

Muscles of the Pelvis.

Levator Ani:—In this animal the levator ani consisted of the

same parts as described by Lartschneider (32). The iliac portion

arose from the margin of pelvis minor, and the pubic portion

arose from the back of the symphysis pubis. It is inserted into

the sides of the lower end of the rectum and the ano-coccygeal

raphe.

The ischio-coccygeus is a small muscle running from the side of

the coccygeal vertebrz to the inner aspect of the back part of the

ischial tuberosity.

Lartschneider has described a pelvic diaphragm in a female

Chimpanzee.

The perineal muscles differ from those in the human condition.

The vulva and anus are very close together, so there is no true

central point of the perineum, nor is there a trace of transverse

perineal muscles. The ischio-cavernosi have long origins from

the ascending pubic rami; they run over the crura clitoridis, and

the opposite muscles are blended. The sphincter vagine is a

strong collar surrounding the upper part of the vagina and

extending backwards on to the rectum. It has a narrow anterior

slip which fuses with the meeting point of the two ischio-

eavernosi. The sphincter ant externus blends in front with the

sphincter vagine, and it is attached behind to the ano-coccygeal

raphé, which is very powerful.

The ischio-rectal fossee are well marked and laden with fat.

No well-marked bursa exists over the ischial tuberosities. And

the fascia is not divided as in Man into a superficial fatty layer

and a deep fascia of Colles.

The orbital muscles are described on page 415, and the laryn-

geal muscles on page 398.

364 DR. C. F. SONNTAG ON THE ANATOMY,

THE JOINTS.

Spinal Ligaments : :—The anterior common ligament extends

from the axis to the upper segment of the sacrum, and is smaller

above than below. It is nisin ona to the front of the centra, but

not to the depressions between the vertebre. The supraspinous

ligaments run as in Man, and are connected to interspinous

ligaments. There is no ligamentum nuche. The iterspinous

ligaments are as in Man. The ligamenta subflava vun from the

anterior aspect of the lamina above to the posterior aspect of the

lamina below. They and the interspinous ligaments are very

elastic. The posterior common ligament runs from the axis to

the sacrum, but is not so dentate in appearance as in Man. It is

narrow on the centra, and expanded over the intervertebral

discs.

Oostal Articulations (text-fig. 37 A & B):—Anteriorly the

head of the rib is connected by a fan-shaped ligament, not

divisible into three parts as it is in Man, and the fibres all gain

attachments to the anterior common ligament (A.C.L). The

upper fibres run to the vertebra above, and the lower ones to the

vertebra below, where they are overlapped by the upper fibres of

the next joint. Posteriorly the tubercle of the rib is attached to |

the transverse process of the vertebra by a capsule. The upper

border of the neck of the rib is connected to the transverse

process by a fan-shaped ligament (C-T.L.).

The intertransverse ligaments (1-T.LL) are fan-shaped. ‘They

connect the tip of the transverse process above to the upper

border of the transverse process below.

The lengths of the spinous processes of the cervical vertebree

“URS :—axial 3 inch; 4th vertebra 2 inch; 6th and 7th vertebre

# inch.

Atlanto-axoid Joints :—The posterior atlanto-axoid ligament,

from the posterior arch of the atlas to the upper border of the

axis, corresponds to the ligamenta subflava Sie It is

strengthened by fibrous bands (text-fig. 37, F.B): From the

transverse process of the axis to an elevation on the posterior

arch of the atlas; 6. A central atlanto-axoid band (C.A.B);

c. Several small bands between the others. The anterior utlanto-

axoid ligament is a very strong band continuing the anterior

common ligament from the axis to the atlas.

Joints of the Occipital, Atlas, and Axis (text-fig. 37 D) :—The

posterior occipito-atlantoid ligament is a thin membrane running

between the posterior arch of the atlas and the edge of the

foramen magnum. It is strengthened by lateral Tends attached

close to the condyles. Jt functions as a posterior capsular liga-

ment. An internal capsular ligament runs from the inner

border of the condyle to the superior articular process of the

atlas. And a strengthening band runs from the posterior arch

of the atlas to the occipital bone close to the attachment of the

internal capsular ligament. The anterior occipito-atlantoid

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 365

ligament is a strong membrane continuing the atlanto-axoid

ligament from the anterior arch of the atlas up to the anterior half

of the circumference of the foramen magnum. The membrana

tectoria continues the posterior common ligament up to the

cranial surface of the basi-occiput where it spreads out in a fan-

shaped manner. The posterior occipito-axoid ligament (P.O-

A.L) is a strong band on each side on the deep aspect of the

membrana tectoria. It extends from the antero-lateral part of

the vertebral canal to the basi-occiput. It passes lateral to the

odontoid process and covers the spinal aspect of the lateral part

of the transverse ligament and conceals it from view. The

transverse ligament (T.L) is a strong band running behind the

odontoid process. It is attached by its extremities to the inner

aspect of the inferior atlantic articular processes. The cruciate

ligament has no inferior crus. The superior crus (8.C.C) is

broader and shorter than in Man. It runs from the transverse

ligament below to the basi-occiput below the membrana tectoria

above. The check ligaments (C.Li)-are more horizontal than in

Man. They run from the odontoid to the inner aspect of the

occipital condyles. The middle odontoid ligament runs from

the tip of the odontoid to the anterior edge of the foramen

magnum. It is stronger and more horizontal than in Man.

Capsular ligaments are as usual. As the odontoid runs farther

upwards than in Man the ligaments are modified.

The lumbar vertebre are connected by the usual capsular

ligaments, intervertebral dises and ligaments connecting their

processes.

Ligaments of the Pelvis (text-fig. 37):—The lumbar vertebrz

are included more closely between the ilia than in Man, and the

pelvic ligaments differ in several respects. The lwmbo-inguinal

ligaments (L-I.L) consist of an upper horizontal ligament

running from the third lumbar transverse process to the inner

lip of the crest of the ilium, and a lower fan-shaped one from the

fourth lumbar transverse process to the inner lip and inner

surface of the ilium. The fibres of these ligaments are con-

tinuous; and the lower one is continuous with the anterior

sacro-iliac ligament (A.S-I.L). The latter is fan-shaped, and runs

from the ala sacri to the anterior surface of the ilium, where it

spreads out in a fan-shaped manner. Some fibres run along the

pelvic brim. The short posterior sacro-iliac ligament is as in

Man. It is superficially thickened to form the oblique posterior

sacro-tliac ligament (O.P.S-I.L), which runs from the posterior

tubercles of the sacrum to the two posterior iliac spines and the

bone between. The great sacro-sciatic ligament (G.S-S.L) runs

from the side of the lower sacral and upper coccygeal vertebre.

It narrows and then it expands again on the posterior part of the

inner aspect of the ischial tuberosity. A falciform process runs

forwards to become the sub-pubic ligament. The ischial spine is

slight, but it receives a broad expansion from the great sacro-

sciatic ligament, and a cord-like small sacro-sciatic ligament

366 DR. C. F. SONNTAG ON THE ANATOMY,

(S8.8-S.L) from the side of the lower end of the sacrum. It can

be seen from text-fig. 37, that the pelvic outlet is divided into

seven parts by ligaments. The obturator membrane is dense and

strong. The symphysis pubis has the same ligaments as in

Man. The upper and sub-pubic ligaments are weak, but the

anterior and posterior ones are strong, particularly the latter.

Within the inter-articular cartilage there is a small synovial

cavity.

The Temporo-maxillary Joint :—The capsule is thick and strong,

the upper part of its outer surface giving origin to some of the

deeper fibres of the masseter. The synovial cavity is divided

into two by an articular disc which, however, is not perforated.

Text-figure 37,

TRANS. PROC.

—!

ie. 6S5L F

Ligaments of the spinal and pelvic joints. ART.P: articular process; CAPS:

capsules; P.C.L: posterior capsular ligaments. Other letters in text.

The disc is much thicker behind than in front, and lies closely on

the condyle of the mandible, moving with it in mastication.

It is concavo-convex from before backwards on its upper surface.

Posterior temporo-mandibular and external lateral ligaments are

present, the latter being very strong. The inner aspect of the

capsule is thin. The internal temporo-mandibular ligament is

smal], but definite, and blends with the capsule. The anterior

aspect of the capsule is covered by the insertion of the external

pterygoid. The spheno-mandibular ligament is very poor.

The acromio-clavicular joint is as in Man.

The Shoulder Joint :—The shoulder joint is in many ways as

in Man, but the following points deserve special mention. The

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 367

subscapular bursa opens into the joint cavity above the superior

gleno-humeral ligament. The coraco-humeral ligament blends

with the capsule near the insertion of the subscapularis. The

coraco-acromial arch is a strong band, and there is also a weak

acromio-humeral ligament. The insertions of the supraspinatus

and infraspinatus are most intimately blended with the capsule,

and cannot be separated from it. The posterior part of the

capsule is weak and loose, but the anterior part contains a broad,

powerful anterior gleno-humeral ligament. The capsule is

attached to the bones as in Man. The superior glene-humeral

ligament divides into two parts, which are inserted separately

into the humerus, and the subscapular bursa communicates with

the joint between them. The cotyloid ligament and tendon of

the biceps are as in Man.

The Hlbow Joint :—The posterior part of the capsule is attached

as in Man; and a strong band passes from the tip of the ole-

cranon up to the outer part of the olecranon fossa, which contains

fat. The anterior part of the capsule is relatively strong all over.

On the radial side the fibres run downwards and inwards, but

those on the ulnar side run downwards and outwards. The fibres

of these sides interlace and make the capsule strong. The internal

lateral ligament is very strong. It arises from the internal condyle

and spreads out to be attached to the whole inner margin of the

olecranon, the coronoid and the great sigmoid notch. The thickest

parts are fixed to the olecranon and coronoid, but it has no trans-

verse part. The external lateral ligament is attached as in Man.

It is weaker than the internal ligament, but muscles arise from it.

The Superior Radio-Ulnar Joint:—The capsule of the elbow

joint is prolonged down for ? inch over the head and neck

of the radius. ‘The orbicular fibres are as in Man, but there is

no oblique cord. The interosseous ligament begins at the lower

part of the coronoid process. One set of fibres runs downwards

and outwards to be inserted into the radius below the bicipital

tuberosity. A second group runs from the insertion of the

first to the ulnar interosseous border; it is by far the stronger

group, and lies anteriorly.

The Synovial Cavity of the Hlbow Joint :—This is attached to

the margin of the coronoid fossa, but more superiorly than the

fossa for the head of the radius. The line of attachment then

passes about half-way between the articular surfaces and the

condyles. On the posterior surface the capsule is attached round

the margin of the olecranon fossa. On the radial side the attach-

ment corresponds to a line joining the epicondyles and drawn on

the part of the humerus which articulates with the radius. On

the ulnar side it is attached to the margin of the olecranon fossa,

greater sigmoid notch and coronoid, but it’ jumps the gap

between the margins of the lesser sigmoid notch, where it gives

rise to orbicular fibres.

The Wrist Joint:—The anterior and posterior aspects of the

carpus are covered by bands of fibres running in all directions; but

368 DR. C. F. SONNTAG ON THE ANATOMY,

there is no centre of radiation asin Man. The following thickened.

bands deserve mention:—(1) A strong band from the anterior

aspect of the base of the radial styloid to the trapezium and os

magnum ; (2)a weak external lateral ligament ; (3) a small internal

lateral ligament; (4) a thickened band from the posterior part

of the distal radio-carpal joint to the back of the os magnum ;

(5) a posterior band from the distal part of the radius to the

cuneiform and unciform bones. Both anterior and posterior

ligaments are stronger than in Man. The cavity of the joint

(text: fig. 35 B) is complicated. The distal end of the radius is

divided into two facets, which articulate with the scaphoid and

semilunar bones. No articular fibro-cartilage lies on the distal

end of the ulna, but a very strong ligament runs from the distal

end of the ulna to the proximal énd of the pisiform bone. It

plays over the outer aspect of the ulnar styloid, which is covered

with cartilage. The synovial cavity between the radius and the

scaphoid and semilunar bones is prolonged into the inferior radio-

ulnar joint, over the ulnar styloid and ligament and upon the

surface of the cuneiform bone; it gets in between the cuneiform

and pisiform. A large cavity separates the head of the os

magnum from the concave surfaces of the scaphoid and semi-

lunar bones, and it also separates the cuneiform and unciform.

It is continuous with the cavity between the scaphoid and trape-

zoid, A small ligament connects the scaphoid and os magnum

and divides the cavity partially into inner and outer parts. This

transverse carpal cavity is prolonged distally on either side of the

os magnum. ‘wo interosseous ligaments are also present in

the joint. A sesamoid bone lying on the apex of the pisiform is

separated from it by a synovial cavity and a strong interosseous

ligament.

Carpo-metacarpal Joint:—A continuous cavity extends along

the proximal ends of metacarpals 2-5. It is connected with the

transverse carpal cavity (text-fig. 35). Two interosseous liga-

ments run from the os magnum to the sides of the third meta-

carpal bone. Inthe intermetacarpal joints there are prolongations

of the carpo-metacarpal synovial cavity, and interosseous liga-

ments connect the second to fifth metacarpals to one another.

The metacarpo-phalangeal joints ave as in Man, but the dorsal

expansions of the extensor tendons can easily be separated from

the capsules. The inter-phalangeal joints are asin Man. ‘Their

internal and external lateral ligaments are powerful. °

Pollical Joinis:—A prolongation of the transverse carpal

cavity runs between the scaphoid and trapezium, and between

the trapezium and trapezoid, where there is also an imperfect

interosseous ligament. A strong ligament runs from the trape-

zium to the base of the index metacarpal. The carpo-metacarpal

capsule is complete; it is thick and strong on its outer and pos-

terior aspects, but its inner and anterior aspects are weak. The

metacarpo-phalangeal and interphalangeal joints are as in Man,

and in the other digits of the Chimpanzee.

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE, 369

The Hip Joint:—The capsule is attached as in Man. The

ilio-femoral ligament consists of one band, which arises under

cover of the rectus femoris and is strengthened by its tendon. It

runs as does the anterior band of the Y-shaped ligament in Man.

No other ligaments are formed from the capsule. The gluteus

minimus strengthens the capsule at the proximo-anterior part of

the great trochanter. The ligamentum teres, cotyloid ligament,

and transverse ligament are as in Man. The joint contains a

pad of fat.

The Knee Joint :—The ligamentwm patelle is broad and strong,

and has a more extensive insertion than in Man (see page 358).

The internal lateral ligament is not inserted into the internal

condyle of the tibia, but is fixed to the upper third of the shaft.

The external lateral ligament isas in Man. The oblique popliteal

hgament is absent, but a strong femoral intercondylar ligament

is attached to the posterior aspects of the two condyles. The

ligamentum nvucosum is just as in Man. The anterior cruciate

ligament is attached to the tibia as in Man, but its femoral in-

sertion is into the upper half of the mesial aspect of the external

condyle. It is smaller than the posterior cruciate ligament; the

latter is attached to the tibia farther back than in Man, and it

receives a slip from the external semilunar cartilage (text-

fig. 36 F). Two semilunar cartilages are present. The internal

one is larger than the external, and is crescentic in shape. It is

attached in front of the anterior cruciate ligament. Its posterior

horn is inserted as in Man. The external cartilage forms a

small, complete circle. Internally it is attached by a broad liga-

ment to the external side of the tubercles and spine of the tibia.

Postero-mesially it is united by a ligament to the outer surface

of the internal condyle of the femur. It is also connected to the

posterior cruciate ligament. The joint differs in many ways

. from that in Man, and its construction is such that it is one of

the factors which prevent the animal assuming a firm, erect

attitude. Humphry (26), who has made a thorough study of this

joint, points out that the femur in Man is broad and compara-

tively flat on the distal end of the external condyle; ard the

attachments of the lateral ligaments are nearer the posterior

parts of the bone. So the joint is firm and locked when it is

fully extended. In the Chimpanzee, on the other hand, the

distal end of the external condyle is rounded, and the lateral

ligament is not attached far back. He also shows that the

lateral, cruciate, and posterior ligaments are all tight when the

human knee is fully extended, but they never become simul-

taneously tight in the Chimpanzee; to obtain tightness of each

ligament it is necessary to divide all the others. Finally, the

attachment of the gracilis and hamstring muscles to the fascia

of the leg, and the laxity of the ligaments of the joint, are con-

tributory factors which prevent the animal assuming the erect

attitude *.

* See the observations on living specimens recorded on page 420.

370 DR. C. F. SONNTAG ON THE ANATOMY,

The Ankle Joint :—The capsule is attached above to the distal

ends of the tibia and fibula and below to the astragalus. A small

deltoid ligament runs from the medial and distal border of the

tibia to the sustentaculum tali and talus. Posteriorly a very

strong fibular calcanean ligament is present; and it is streng-

thened by an accessory ligament from the lateral aspect of the

lower end of the fibula. A ébial calcanean ligament lies on the

inner aspect of the joint. The talo-fibular ligaments, both

anterior and posterior, are present, but the former is ill-defined.

A ligament runs from the middle of the anterior distal border of

the tibia, under the talus, to the medial aspect of the lateral

distal tubercle of the calcaneus. The tibia and fibula are held

together by thin membrane. The ¢éalo-calcanean ligaments are

well-developed. The dorsal, posterior, and two lateral ones

are strong, but the medial one is weak. The posterior ligament

takes part in forming an interosseous ligament. The talus

is further held in position by a ligament running from the

plantar navicular-calcanean ligament to a small impression on

the head of the talus.

Ligaments connected with the Calcaneus:—The plantar calcaneo-

navicular ligament runs from the sustentaculum tali to the

navicular; and there is practically no internal ligament between

these bones. A well-marked ligament connects the calcaneus

and cuboid.

The long plantar ligament is attached proximally to the cuboid,

and distally to the bases of the second, third, and fourth meta-

tarsal bones. No short plantar ligament exists.

A fine ligament attaches the cuboid to the lateral extremity of

the base of the fifth metatarsal, and another connects it to the

third cuneiform.

Navicular Ligaments:—A dorsal ligament runs from the

navicular bone to the base of the second metatarsal, and a medial

ligament connects it to the first cuneiform.

An interosseous ligament holds the cuboid and cuneiforms in

position.

Plantar Metatarsal Ligaments :—The external ligament is an

aponeurotic septum attached obliquely to the fourth metatarsal

from the outer side of the base across the plantar aspect of the

shaft to the head. The adjacent parts of the bases of the third

and fourth metatarsals are covered by ligamentous fibres from

the sheath of the tendon of the peroneus longus, and representing

the termination of the long plantar ligament. This divides into

two strands, the external one joining the external ligament, and

the internal one passing along the whole length of the third

metatarsal as the internal long plantar metatarsal ligament.

Humphry (26) has gone very fully into the shapes of the bones

of the foot and the part played by the bones, ligaments, and

muscles in its mechanics. He draws attention to the following

points :—(1) The shape of the talus throws the weight on the outer

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 371

border of the foot ; (2) the talus and calcaneus are more for pro-

gression than support; (3) the calcaneus easily rolls outwards on

its lower surface, so does not act as a bearing surface; (4) the

calcaneus is thrown out of the plane of gravity, and it is reduced

like its homotype, the pisiform, to a lever for muscles; (5) the

talus, navicular, and calcaneo-navicular ligament transmit weight ;

(6) the posterior surface of the talus slopes downwards and in-

wards; (7) the action of the calf-muscles on the foot is unfavour-

able for lifting weight or propelling the body; (8) there is no

plantar arch, so the navicular bone with the calcaneo-navicular

ligament rest on the ground and do not transmit weight along

the arch to the hallux, which is not adapted for support; (9) the

mobility of the hallux is obtained by articular, osseous, and

muscular arrangements similar to those of the human pollex;

(10) there is more power of flexing the digits towards the sole

than in Man.

It is, therefore, evident that the joints of the neck, pelvis,

wrist, and lower extremity, particularly the latter, differ con-

siderably from those in Man, and all are specialisations in

accordance with the mode of locomotion.

THe ORGANS OF DIGESTION.

The loose, thick, fleshy dips are projected forwards over the

maxille. No philtrum is present, and in the living animal only

a small part of the red margin of the lower lip is visible when

the mouth is closed. Their inner surfaces are studded with the

openings of numerous labial salivary glands. The labial frenula

are long and narrow. The gape of the mouth is wide. The

cheeks are loose and mobile, but no pouches exist. The vestibule

is semilunar, and receives the secretions of Stensen’s ducts, which

open on papillae placed as in Man. And a row of papille lies

beside the salivary papilla on each side (text-fig. 28 C). Rex (41)

has described the histology of the lips, and Ehlers (59) described

folds of mucous membrane connecting the gums and cheeks. The

cavum oris is thrice as long as broad according to Gratiolet (22).

In my specimen the measurements are :—

Length of hard palate...... 2°7 inches.

Length of soft palate ...... AAs Total=3°8 inches,

Greatest width of palate... 14 ,,

Much, however, depends on the age of the animal. Keith has

shown that the breadth of the palate is greater than the length

in new-born animals. The cavity of the mouth has also been

mentioned by Symington (48) and Tyson (50).

The ruge of the hard palate have been figured or described by

Beddard (3), Bischoff (60), Ehlers (59), Gratiolet (22), Symington

(48), and Waldeyer (52). In my specimen there are no complete

ridges crossing the palate, and no incisive papilla. Eleven pairs

372 DR, C. F. SONNTAG ON THE ANATOMY,

of ridges radiate from the median raphé, which is thicker an-

teriorly than posteriorly. The sofé palate has the same histological

structure as in Man, and its glands are very numerous. Using

C. for complete ridges, I. for imcomplete ridges, P. for incisive

papilla, R. for median raphé, and U. for uvula, the formula is C0,

T11-11, PO, R+, U+.

In a former paper (46) I described the tongues of several

specimens, and I collected the literature. In this animal there

are eight papille arranged in a Y.

The pharynx is as in Man. Faucial and pharyngeal tonsils

are both present, and are nourished from the vascular circle

formed by the branches of the ascending pharyngeal artery

(text-fig. 29). The former is covered by fenestrated mucous

membrane. Although lingual, faucial and pharyngeal tonsils

are present I was unable to detect Waldeyer’s lymphatic ring.

Seesel’s pocket is absent. The constrictor muscles are as in Man.

The sinus of Morgagni is large, and the levatores and tensores

palati are more horizontal owing to the prognathism of the skull

than mn Man.

The wsophagus is entirely behind the trachea in the neck. It

has similar relations in the neck and thorax to those in Man.

The mucous membrane is thrown into prominent longitudinal

folds in the cervical and thoracic parts, but the former are more

numerous and closer together. The wells are thinner and more

dilated in the lower part of the cesophagus. In the upper part

the inferior constrictor joins the outer longitudinal muscular coat

of the cesophagus, which increases in thickness from above down-

wards, and becomes continuous with the outer coat of the gastric

musculature. The circular muscle coat thickens from above

downwards and also becomes continuous with the circular fibres

of the stomach. At the lower end of the esophagus it forms the

sphincter of the cardia, which is two inches long. The inner

* longitifdinal fibres in the upper part consist of a few strands, and

the submucosa bulges between them; they are entirely absent in

the lower part of the esophagus. Man has only two muscular

coats.

Cunningham (14) has given a fine illustration of the topo-

graphical relations of the abdominal viscera; and anatomical

details have been given by a large number of authors, whose

works have been collected and grouped by Keith (29). So no

very full account is given below.

The Stomach.

The cesophagus passes through the diaphragm at the level of

the ninth dorsal vertebra, and opens into the stomach after an

intra-abdominal course of half an inch. The stomach is divided

into fundus, body, antrum, and pylorus. The fundus is well

marked and projects up into the left cupola of the diaphragm.

The long axis of the stomach is cresecntic, and is more horizontal

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEK. 373

than in Man. The great curvature reaches a point half an inch

below the fourth lumbar vertebra, and the lowest point of the

lesser curvature is level with the first lumbar vertebra, Con-

sequently the stomach is U-shaped, and the pylorus is not far

from the csophagus. The pyloric antrum is one and a half

inches long, and the pylorus, whose walls are thick, is of the

same length. Between the antrum and pylorus there is a slight

incisura, and there is a sudden transition from the pylorus to the

duodenum. ‘The pylorus does not project like a knob into the

duodenum as it does in Man.

The serous coat is a uniform covering, united in the usual

manner to the omenta. The muscles are thin, but three kinds

are present. The external longitudinal layer is a complete

covering, but it is thicker along the curvatures than on the

intermediate parts of the body. The circular fibres are thickest ;

they are found in the body and pylorus, but only a few run

from the csophagus into the fundus. The oblique fibres are

restricted to the fundus and part of the body to the left side of

a downward prolongation of the long axis of the csophagus, and

they form rings as in Man. Consequently there are only two

muscular layers—an outer one of longitudinal fibres, and an

inner one of oblique fibres to the left and circular ones to the

right. The subserous and submucous coats are thin. The mucous

membrane is thin, and some of the gastric arteries are seen

ramifying in the wall of the stomach. So thin is it that the red

injection matter in the vessels shines up prominently against the

pale mucosa. It is succulent and its surface shows the areas and

glandular orifices as in Man. Only a few longitndinal ruge are

present.

Blood Supply -—The coronary artery (text-fig. 40 A) runs along

the lesser curvature along with branches of the left vagus nerve ;

it gives off tortuous gastric arteries to both surfaces (a.g.a and

p.g.a), and cesophageal arteries which pass up through the

cesophageal opening in the diaphragm. The parent artery anas-

tomoses with the right gastric branch of the hepatic artery. The

splenic artery sends branches to the greater curvature (g.c.b) and

the left gastro-epiploic artery, which anastomoses in the great

omentum with the right vessel from the hepatic artery. The

hepatic artery, in addition to the branches, sends superior pyloro-

duodenal arteries downwards, and these meet with inferior

pyloro-duodenal branches of the cceliac axis; from both the

stomach receives branches. The gastric veins open into the

portal system (text-fig. 43), Barkow (2) has given an illustra-

tion of the stomach and its vessels.

Nerves :—The left vagus sends branches along the lesser cur-

vature as far as the pylorus, and some other twigs form a plexus

over the lower end of the esophagus. The right vagus sends a

rich supply of nerves to the stomach through the splenic plexus

and coronary plexus, and directly.

Proc. Zoou. Soc.—1923, No. XXV. 25

374 DR. C. F. SONNTAG ON THE ANATOMY,

Intestinal Tract.

The duodenum begins opposite the first lumbar vertebra, and

it is throughout entirely behind the peritoneum. It 1s divided .

as in Man into horizontal, descending and ascending parts which

measure 11, 2, and 24 inches long respectively. There is a well.

marked duodeno-jejunal flexure at the level of the first lumbar

vertebra. The common bile-duct and pancreatic ducts have a:

common opening in the descending part, but there is no papilla.

No valvule conniventes are present, and the villi are small.

The ileum and jejunum are 11 feet 5 inches long. Their villiare

small. At intervals there are groups of small longitudinal folds

of the mucosa. Four Peyer’s patches are present at wide intervals

in the lower half of the ileum; the lowest, which is also the

largest, is 2 inches long and # inch broad.

The vermiform appendix is 44 inches long, and the cwcwm is

3 inches. No appendix valve is present, but the ileo-cecal orifice

is guarded by a shelf valve. _

The colon is 4 feet long. It is sacculated as usual by two

longitudinal muscle bands, and there are many appendices epi-

ploic. No Peyer’s patches are present. The most capacious

part is the sigmoid colon. . :

The rectum and anal canal are 54 inches long. In the rectum

there are eight circular folds, of which the fifth is very promi-

nent, and below it there is a deep pocket on the left side. The

anal canal shows numerous strongly-developed vertieal folds of

mucosa, representing the columns of Morgagni, but there are no

traces of the valves of Ball. The entire rectum and anal canal

form a straight tube without any trace of the flexures present in

Man. Herrmann (74) has described the anal mucosa, in detail.

The Peritoneum.

The great omentwm is heavily laden with fat and reaches the

symphysis pubis. All four layers are fused and can only be

separated at the stomach and transverse colon. The anterior

layers are attached to the greater gastric curvature, the first part

of the duodenum and spleen, and bands connect it to the lateral

abdominal parietes. Between the layers are the usual vessels

and lymphatic glands.

The lesser omentum is attached as in Man, and the foramen of

Winslow is large. It is bulged forwards above the stomach.

Between its layers are numerous vessels, sympathetic nerves and

lymphatic glands. hoa

No gastro-pancreatic folds are present. The gastro-phrenic,

gastro-splenic and lieno-renal ligaments are well-marked, A

peritoneal ligament connects the lower pole of the spleen to the

transverse colon, and a small accessory spleen is connected to the

colon at the same point. A well-marked ligament connects

the pylorus to the right ribs, but no suspensory duodenal muscle

exists.

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 375

The root of the mesentery of the small intestine runs from the

left side of the body of the first lumbar vertebra to the right

iliac fossa. In the large intestine there is no mesentery to the

cecum, ascending, descending and pelvic colons. The appendix

has a mesentery, but no other part has such a long mesentery as

the iliac colon. In the rectum the peritoneum is disposed as in

Man, being first surrounded by it, and then the peritoneum

gradually leaves it till the lower part and the anal canal are quite

devoid of a serous investment.

The Salivary Glands.

Many labial salivary glands are present.

The parotid gland (text-fig. 26) is pyramidal in shape, with the

base upwards, immediately below the concha auris. The apex

curves round the angle of the mandible and touches the sub-

maxillary gland. The upper part is composed of small lobules,

but the lower part is coarse. ‘The capsule is well marked, but no

lymphatic glands are included within it. Stensen’s duct emerges

from the upper part of the gland and its course and relations are

asin Man. The relations of the gland to the large vessels and

nerves are also asin Man.

The submaaillary gland (text-figs. 30 & 31) is flat and

triangular and measures one and a half inches in diameter. It is

partly under the horizontal ramus of the mandible and partly on

the interramal muscles. No strong capsule exists. It is com-

posed of superficial and deep parts, each of which is coarsely

lobulated. The duct emerges from the deep surface, runs as in

Man, and opens on a frenal lamella beneath the tongue.

The sublingual gland (text-figs. 31 and 32) is pyramidal with the

apex anterior. Its relations and the course of its duct are as

in Man.

Brief accounts of the glands have been given by Tyson (50)

and Gratiolet (22).

The Pancreas.

Bischoff (60), Cavanna (61), Flower (20), Gratiolet (22), and

Tyson (50) have given details of the pancreas. In my specimen

it is flat, thin, dark in colour and coarsely lobulated. It has the

usual head! body and tail, and it crosses the first lumbar vertebra.

An additional process runs up along the portal vein for a short

distance; and the pancreatic arteries, which are branches of the

splenic, are accompanied by sympathetic nerves. The duct unites

with the common bile-duct.

The Liver.

The liver much resembles that in Man. The umbilical fissure

is much bridged over, and the fissure of the vena cava is also

enclosed. Indications of lateral fissures exist. Near the umbilical

region of the right lobe there is a small lobule directed ventrally.

25*

276 DR. C. F. SONNTAG ON THE ANATOMY,

The gall-bladder is superficial and extends beyond the ventral

margin of the lobe and it is flexed on itself. In occasional

specimens the gall-bladder is deeply embedded in the liver sub-

stance. The Spigelian lobe is subquadrate, and the caudate iobe

is a triangular cone directed to the right. In some specimens

the apex of the caudate lobe reaches the right margin of the

liver, but in others it does not do so. The relative sizes of the

hepatic lobes may be expressed by Garrod’s method thus :—

R2>L>Sp=C.

Figures or descriptions of the livers of other specimens, which

agree with the above, are given by Bischoff (60), Flower (20),

Barkow (2), Gratiolet (22), Cavanna (61), Symington (48),

Traill (49), Tyson (50), and Sperino (47).

The relations of the abdominal organs to the vertebre are

different from those in Man, because there are thirteen dorsal

vertebre. The first lumbar vertebra in the Chimpanzee corre-

sponds to the second lumbar vertebra in Man.

ORGANS OF CIRCULATION.

‘The venous side of the circulation is larger, relatively to the

arterial side, than in Man.

The pericardium adheres strongly to the central part of the

diaphragm. When it is slit open the finger can explore the

aortic arch up to and including the root of the innominate artery,

but the reflection of the serous pericardium prevents one touching

the left subclavian artery. Only a small part of the pulmonary

artery is palpable.

The heart is small, measuring 3:2 inches long, 2°3 inches wide,

and 1:7 inches antero-posteriorly. The upper border is level

with the second costal cartilage, and the apex lies in the fifth inter-

costal space. Fat is present on the base and apex. Its internal

structure is very similar to that in Man. ‘The position and

relations have been recorded or figured by Cunninghani (18) and

Ruge (48), and details of its construction have been given

by Bischoff (60), Cavanna (61), Ehlers (59), Dwight (18),

Gratiolet (22), Tyson (50), and Traill (49). The apex is entirely

formed by the thick, muscular left ventricle, and this differs

entirely from the condition which I have already described and

figured in Mandrillus (62).

The pulse of a young male Chimpanzee, whose age would make

it correspond to the young child, was 150 per minute. It was

regular in rate and rhythm; it was full, and the rise and fall

were moderately rapid. No dicrotism was present. The apex

was not very sustained. It could easily be felt on the radial

aspeet of the lower end of the forearm because the radial artery

is very superficial. Owing to the inability to listen to the hearts

of the larger specimens in the Gardens I am unable to describe

the relation between age and heart rate. The heart sounds were

asin Man, The blood pressure was not obtainable.

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 377

The aortic arch (text-fig. 38, A.A) deseribes a small curve and

gives way to the descending aorta at the sixth dorsal vertebra.

Its relations are as in Man. It gives off the innominate and left

subclavian arteries from its convexity, and the main bronchial

artery to the left lung (L.B. A) arises from its concavity. The

innominate artery (1.A), a large vessel about 2 cm. long, gives off

the left common carotid (L.0.C.A) and divides behind the right

half of the manubrium sterni into the right common carotid

(R.C.C.A) and right subclavian (R.S.A) arteries. The left

common carotid gives off the thyroidea ima (T.1.A) close to its

origin. The intrathoracic parts of the left common carotid and

left subclavian arteries are mostly as in Man, but the latter is

relatively larger and not so vertical.

Text-figure 38.

The main arteries of the thorax and pectoral extremity. Ax.A: axillary artery ;

O.A : occipital artery ; SU.A: subscapular arteries. Other letters in text.

The pulmonary artery (text-fig. 38, P.A) is much more capa-

cious than in Man, and its left branch is united to the aortic

arch by a wide, open, ductus arteriosus (D.A). The presence of

the latter, and the origin of the left bronchial artery are of

interest from the embryological point of view, but the foramen

ovale is closed and the vene cave are normal.

The ductus arteriosus is the sixth embryonic arterial arch, and

the bronchial artery coming from the concavity of the aortic

arch possibly represents the remains of one of the vessels con-

necting the outer extremities of the embryonic arches. So we

have in this animal a combination of interesting embryological

conditions persisting.

378 DR, C. F. SONNTAG ON THE ANATOMY,

As the pulmonary artery is much wider than that in Man, the

velocity and pressure of the blood in it must be relatively less

than in him, as can be shown by applying the laws of velocity

and pressure.

The descending thoracic aorta extends from the sixth to

eleventh dorsal vertebre after which it passes into the abdomen.

It gives off the intercostal arteries to the lower ten intercostal

spaces, a small bronchial artery to the left lung, a large bronchial

artery to the right lung, and several branches to the thoracic

esophagus, which anastomose with cesophageal branches of the

cceliac axis. The lower intercostal arteries supply the diaphragm.

The abdominal aorta extends from the twelfth dorsal vertebra

to the lower border of the fourth lumbar vertebra, where it

divides into the two common iliac arteries. It is relatively

smaller, and its branches are fewer than in Man. The following

is the order of the branches from above downwards :—

1. Phrenic artery.

. Coeliac axis.

. Superior mesenteric artery.

. Renal arteries.

. Right ovarian artery.

. Inferior mesenteric artery.

. Four lumbar arteries arising at different levels from the

back of the aorta.

It does not give off any suprarenal arteries, nor is it continued

as a middle sacral artery.

The phrenic artery is a large vessel arising from the left side

of the beginning of the abdominal aorta. It gives a small branch

to the left crus and left half of the diaphragm, and it is continued

over the right crus as a large vessel which sends branches to the

central tendon, the muscle fibres, the right crus, the process

arising from the second lumbar transverse process and the right

suprarenal capsule,

Blood Supply to the Suprarenal Capsules:—Kach capsule

receives a vessel from the phrenic artery and the corresponding

renal artery, but none from the aorta.

The Celiac Axis (text-fig. 40, C.:Ax) arises immediately above

the upper border of the pancreas, and it quickly divides into

hepatic, coronary, splenic, and inferior pyloro-duodenal arteries.

Of these the hepatic is by far the largest.

The Hepatic Artery (H.A) first runs to the right and then

turns upwards to the liver. Between the layers of the gastro-

hepatic omentum it divides into two branches; and it lies in

front of the portal vein, and to the left side of the common bile-

duct. One of the two terminal branches (a) runs straight to

the liver and divides into two arteries which enter the portal

fissure. The other branch (d) runs in a convoluted course to the

right, gives off. the cystic artery (ca) to the gall-bladder and

divides into two arteries which sink into the right and left lobes

of the liver.

“IO OH CO bD

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 379

' . The artery gives off a trunk which divides into :—

l. Right gastric artery (r.g.a), which anastomoses with the left

gastric artery on the lesser gastric curvature.

2. Pylorc-duodenal vessels (p.d.v), which anastomose with the

inferior pyloro-duodenal artery.

3. Right gastro-epiploic ar tery (r.g.e.a), which anastomoses with

‘branches of the splenic artery in the great omentum.

The Coronary Artery (C.A) is continued through the Pacpingts

as the cesophageal artery (o.a).. Branches arising from its lateral

side are :—

1. Anterior gastric arteries (a.g.a) to the anterior wall of the

stomach in the fundus and body.

2. Posterior gastric arteries (p.g.a) to the posterior wall of the

stomach in the fundus and ae

3. Left gastric artery (l.g.a), in the lesser omentum, anas-

tomoses with the right gastric artery.

The Inferior Pyloro-duodenal Artery (1.P.D.A) anastomoses

with branches of the hepatic and superior mesenteric arteries.

The Splenic Artery (S.A) runs infero-laterally and divides into

two terminal splenic vessels. It gives off branches to the stomach

(g.b) and several vessels which form the left gastro-epiploic

artery (J.g.e.¢) which anastomoses with the right artery. It is

much smaller than the hepatic artery. .

The Superior Mesenteric Artery (text-fig. 40 B) sweeps down-

wards into the right iliac fossa. It supplies the duodenum,

jejunum, ileum, and large intestine as far as the right third of the

transverse colon. It divides into two main branches. The

trunk of the artery and left branch (1.5) supply the small

intestine from the duodenum to the junction of the middle and

lower thirds of the ileum, the vessels for these parts coming off

close together, The highest branch anastomoses with the inferior

pancreatico-duodenal branch of the celiac axis, and the lowest

one anastomoses with the highest branch of the other half of the

superior mesenteric artery. The trunk of the artery gives off a

branch (6) which bifurcates; the one half anastomoses with the

‘lower branch of the parent stem, and its other half anastomoses

in the transverse meso-colon with the middle colic branch of the

inferior mesenteric artery. The arterial arcades in the mesentery

are not numerous. Many glands and sympathetic nerves are

mixed with the vessels.

~ The Inferior Mesenteric Artery (text-fig. 40 C) arises from the

front of the abdominal aorta about three-quarters of an inch

above its bifurcation. It runs downwards for half an inch and

divides into two vessels which subdivide into large bundles of

vessels for the iliac and pelvic colons. These vessels anastomose

with one another. The parent artery gives off a large vessel,

which divides into middle and left colic arteries, and the latter

divides into ascending and descending branches. The mid colic

389 DR. C. F. SONNTAG ON THE ANATOMY.

artery (M.C.A) supplies the left part of the transverse colon and

the hepatic flexure. It anastomoses with the right colic branch

of the superior mesenteric artery, and the ascending branch of

the left colic artery. The left colic artery (L.C.A), supplying the

descending colon, anastomoses with the mid colie artery and

branches to the iliac colon. The superior hemorrhoidal artery

(S.H.A), from the right division of the parent artery, goes down

to the pelvis to supply the rectum and anus. It is relatively

The arterial supply to the pectoral extremity. A: arteries of the forearm and

hand; B: arteries of the arm; A.I.O and-A.J.A: anterior interosseous

artery; D.C: descending branch of the circumflex arteries; M.A; median

artery. Other letters in text.

larger than in Man, and replaces branches of the hypogastric

artery. It divides into two terminal branches; the anterior

supplies the rectum and anus, and the posterior one is limited

to the rectum.

Renal Arteries :—The left vessel (text-fig. 40 D) gives off supra-

renal arteries (s.7.a), branches to the renal capsule (7.c.a), two

small arteries which anastomose with the lumbar arteries (/.a.a)

and the left ovarian artery (/.0.a4). It ends by dividing into two

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE, 381

vessels to the kidney. The right renal artery gives off renal,

suprarenal and capsular vessels, but no parietal nor ovarian

arteries,

Ovarian Arteries:—The left one comes from the left renal

artery, but the right one springs from the aorta. Both run as in

Man, and their terminal parts are convoluted.

Lumbar Arteries :—F¥our single arteries arise from the back of

the aorta, The highest is level with the cceliac axis, the second

is just below the renal arteries, the third is level with the third

Text-figure 40.

Arteries of the abdomen and pelvis. A: celiac axis; B: superior mesenteric artery ;

C: inferior mesenteric artery; D: left renalartery; E: iliac arteries; C.I.A:

common iliac artery; H.I.A: external iliac artery; I.1.A: internal iliac or

hypogastric artery; 7.6: arterial branches to the kidneys. Other letters

in text.

lumbar vertebra, and the fourth is at the fourth vertebra.

These bifurcate, and the halves run like the lumbar arteries

in Man.

The common iliac arteries diverge for one and a half inches

from the aorta, along the pelvic brim, and each gives off the

hypogastric artery and is continued as the external iliac artery.

The left vessel is the more vertical.

The external iliac arteries give no branches, but those which

382 DR, CG. F. SONNTAG ON THE ANATOMY,

arise from them in Man are here replaced by branches of the

last lumbar and femoral arteries. The relations are as in Man.

The hypogastric artery (text-fig. 40 E) on each side divides as

in Man into anterior and posterior divisions. The anterior

division gives off :—

1. A trunk which divides into superior vesical (8.V.A) and

uteri-vaginal (U-V.A) arteries. The former is a small vessel,

which reaches the side of the bladder up which it runs to the

fundus. The latter is larger and breaks up into vessels supplying

the uterus and vagina; details are given on p. 401.

2. The Pudendal Artery (P.A) gives off the inferior vesical

artery (I.V.A), passes through the great sciatic notch at the

lower border of the pyriformis and, after giving off the inferior

gluteal artery (1.G.A), it breaks up into branches which pass

through the ischio-rectal fossa to the rectum and anus, the! vagina,

the levator ani and sphincter vagine.

The posterior division gives off :—

1. A bundle of arteries which enter the anterior sacral

foramina (S.A.).

2. Lateral sacral artery (L.S.A).

3. Superior gluteal artery (8.G.A).

The superior gluteal artery emerges at the upper border of the

pyriformis and supplies it. It divides into two main branches.

The upper one supplies the gluteus medius and gives the nutrient

artery to the ilium. The lower one descends to supply the

gluteus medius, gluteus minimus, and scansorius.

The inferior gluteal artery emerges with the pudendal artery

at the lower border of the pyriformis. It gives branches to the

gluteus maximus, gemellus superior, obturator internus, scam-

sorius, and acetabular part of the ilium. It anastomoses with the

lateral circumflex artery.

Arteries of the Head and Neck.

The common carotid arteries (text-fig. 41,C.C.A) extend from

_the sterno-clavicular articulations to the upper border of the

lateral aspects of the thyroid cartilage, where they divide into

external and internal carotids. They are concealed by the large

external jugular veins, and they line in front of the vagus and

sympathetic nerves. But no internal jugular veins are present

to form lateral relations. No carotid sheath exists. The other

relations are as in Man. It gives off tortuous inferior and

middle thyroid arteries (text- fig. 41, LT.A. and M.T.A), which

replace the inferior thyroid branch of the subclavian arteries.

This may be an individual peculiarity.

The external carotid artery (E..C.A) first ascends almost verti-

cally till it reaches the level of the hyoid bone, where it inclines

posteriorly and upwards, being continued as the temporo-

maxillary artery. Within the parotid gland it divides into

PHYSIOLOGY, AND PATHOLOGY OF THE. CHIMPANZEE. 383

internal maxillary, superficial temporal, and transverse facial

arteries. ‘The branches run in different directions. Coursing

mesially are the superior thyroid (S.T.A), a combined lingual

and external maxillary trunk (L.F.T), transverse facial ‘and

internal maxillary arteries. Running laterally is the occipital

artery (O.A.), and vertically the superficial temporal, ascending

pharyngeal and parotid arteries.

The superior thyroid artery (text-fig. 41, 8.T.A) arises almost

at the beginning of the external carotid. It describes the usual

Text-figure 41.

| [ s BIN ERS oe

The larynx, thyroid gland and vessels of the neck. H.B: hyoid bone; O-H.M:

omo-hyoid muscle; S-H.M: sterno-hyoid muscle; §.T.M: sterno-thyroid

muscle; T.G: thyroid gland; T.H.M: thyro-hyoid muscles; T-M.A:

temporo-maxillary artery; T-P.B: thymus and parathyroid; XII: hypo-

glossal nerve. Other letters in text.

curve, with its convexity upwards, and then descends along the

greater part of the mesial border of the lateral thyroid lobe. It

terminates by anastomosing with the thyroidea ima (T.I.A).

It gives off mesial branches to the omo-hyoid, sterno-hyoid,

sterno-thyroid, crico-thyroid and thyro-hyoid muscles; and a

branch enters the larynx through the thyro-hyoid interval. The

lateral branches supply the thyroid gland and anastomose with

branches of the middle thyroid artery. The corresponding

superior thyroid vein enters the anterior facial vein, The

384 DR. C. F. SONNTAG ON THE ANATOMY,

combined linguo-facial artery (L.F.T) is given off at the point

where the external carotid changes its direction. After an

upward and forward course of half an inch it divides into

lingual and external maxillary arteries.

The lingual artery runs horizontally and disappears under

cover of the hyo-glossus muscle after giving off a large branch to

the submaxillary gland. It then courses between the hyo-glossus

and middle constrictor of the pharynx; and in this situation it

gives off supra-hyoid and muscular arteries. Emerging from

under the anterior border of the hyo-glossus, it passes forwards

and dips downwards between the stylo-glossus and sublingual

gland laterally and the genio-glossus mesially. It sinks into

the latter, and can be traced to a communication with its neigh-

bour below the apex of the tongue.

The external maxillary artery (text-fig. 41, H.M.A) runs first

forwards and upwards on the upper surface of the submaxillary

gland, then between the gland and the mandible. At the

anterior border of the masseter it crosses on to the outer surface

of the mandible and gets into the face (text-fig. 26). There it

runs in a curved, but not convoluted, course to the angle of the

mouth, whereit becomes more vertical ; and it ends in the levator

anguli oris. It is concealed by the platysma and zygomatic mass,

and it lies on the surface of quadratus labii superioris and

buccinator. It gives off several branches to the outer surface of

the submaxillary gland (S.M.G). In the face (text-fig. 26) it

gives off masseteric (M.A.S), inferior labial (I.L.A), inferior

coronary (1.C.A), and superior labial (S.L.A) vessels, whose dis-

tributions are shown in the figure. The submaxillary branches

send vessels to the mylohyoid.

The transverse facial artery (text-fig. 28) runs forwards between

the parotid and masseter, supplying both, and then along the

zygoma.

The internal maxillary artery (text-fig. 29 A) has the same

course as in Man, but it lies on the surface of the external ptery-

goid. It gives off the following branches :—

A. In the Pterygoid Region :—

1. Numerous pterygoid branches to the muscles, especially to

the insertion of the external pterygoid.

2. Inferior dental artery (I.D.A) which runs as in Man along

with the inferior dental nerve.

3. Meningeal artery (M.A) which passes deep to the external

pterygoid, and divides into middle and accessory arteries, which

enter the skull as in Man.

4. Buccal artery (B.A) accompanying the long buccal nerve.

5. Fine arteries to the suctorial pad of fat (S.P.F).

6. Posterior superior dental (P.S.D) to the gums round the

molar teeth.

7. Two large, deep temporal arteries (D.T.A) which run up in

the substance of the temporal muscle and anastomose freely.

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 385

B. In the Pterygo-maxillary Region :—

1. Infra-orbital artery, which passes forwards to the face. It

supplies the incisor and canine teeth, some of the muscles of the

face, and the upper lip.

2. Descending palatine artery to the soft palate, gums and

mucous membrane of the mouth.

3. Pterygo-palatine artery to the roof of the pharynx,

sphenoidal sinus, roof of the nose, and the Eustachian tube.

4. Spheno-palatine to the roof and outer wall of the nose, the

ethmoidal cells, sphenoidal sinus, and pharnyx.

The Ophthalmic Artery, which continues the internal carotid

artery beyond the carotid canal is, except for its size, similar to

that in Man in every way.

The superficial temporal artery (text-fig. 26, S.Te.A) is the

apparent continuation of the external carotid artery. It runs

upwards accompanied by the corresponding vein and the auriculo-

temporal nerve. It divides into two branches which supply the

scalp from the supra-orbital crest anteriorly to the occipital crest

posteriorly,

The transverse facial artery (text-fig. 28, T.F.A) runs forwards

between the parotid gland superficially, and the masseter deeply,

supplying both by large branches. It is continued by a small

artery along the surface of the zygoma.

The occipital artery arises from the lateral aspect of the

external carotid soon after its origin. It passes upwards and

backwards, and under the cleido-mastoid it gives off the posterior

auricular artery, which supplies the parotid gland and back of

the auricle. The parent stem then curves downwards and

disappears under the lateral border of the splenius capitis.

Under the splenius it gives off a descending branch which

passes downwards among the muscles of the neck and supplies

them by small twigs. The parent vessel then passes onwards

under the complexus, and supplies it and the muscles bounding

the sub-occipital triangle. It does not end in the scalp, nor

is the terminal part crossed by the sub-occipital nerve as in

Man.

The branches, with the exception of the posterior auricular

artery, are distributed entirely to the muscles. No meningeal

arteries run from it through the anterior condyloid foramen as

in Man.

The ascending pharyngeal artery (text-fig. 29 B), from the back

of the beginning of the external carotid runs upwards and

supplies the pharynx, levator palati, tensor palati, and pre-

vertebral muscles. It passes deep to the common carotid artery,

and enters the jugular foramen behind the nerves. It gives off

branches which form an arterial circle supplying the tonsils and

pharynx.

The internal carotid artery is as in Man. It is accompanied

by several twigs from the superior cervical sympathetic ganglion.

386 DR. C. F. SONNTAG ON THE ANATOMY,

Arteries of the Pectoral Extremity.

Subclavian Arteries (text-fig. 38. R.S.A. and L.S.A) :— The.

branches of the extra-thoracic parts differ from those in Man,

but the parent vessels are similar, though relatively larger. The

branches are :—

1. Vertebral Artery (V.A) which is very large. It enters the

foramen in the sixth cervical vertebra along with sympathetic

nerves from the inferior cervical ganglion of the sympathetic.

2. A large trunk whose branches correspond to separate

branches of the human subclavian. Its first set of branches,

which arise together, are :—

a. Spinal Arteries (S.A) entering the lower four intervertebral

foramina hehind the corresponding nerves.

b. Profunda cervicis (P.C.A) which passes between the trans-

verse process of the seventh cervical vertebra and the neck of the

first rib. It ascends among the muscles of the back of the neck,

supplies them, and anastomoses with the occipital artery.

c. Muscular branches to the prevertebral muscles.

The trunk then runs outwards, gives off an ascending branch

(A.B) to the muscles in the floor of the posterior triangle and

the upper four spinal foramina and the long thoracic artery

(L.T.A). Finally it divides into the suprascapular artery (S-S.A),

and a branch corresponding to the descending branch of the trans-

verse cervical artery in Man (D.T.A). These terminal branches

course much as in Man.

No inferior thyroid artery is present, its place being taken by

a branch of the common carotid artery.

3. Prevertebral. muscular artery.

4. Superior intercostal artery (S.1.A) is large. It runs over

the neck of the first rib, gives very small branches to the first

two interspaces, and is continued as a very large artery into the

first thoracie intervertebral foramen.

5. The internal mammary artery (I.M.A.) arises close to the

inner border of the scalenus anticus along with the superior

intercostal artery, and its relations are asin Man. It divides at

the fifth interspace into superior epigastric and musculo-phrenic.

It gives off an artery whieh breaks up into branches to the

thymus, pericardium, and mediastinum, and anastomoses with

the thyroidea ima. Muscular branches run to the triangularis

sterni. But the intercostal arteries are not as regularly disposed

asin Man. The phrenic artery divides into two at the seventh

chondro-costal junction; one branch turns inwards and enters

the diaphragmatic musculature, and the other continues along

the origin of the diaphragm to the mid-axillary line where it

enters the diaphragm. It anastomoses with phrenic branches

of the lower intercostal arteries.

Branches of the suprascapular and descending branch of the

transverse cervical arteries take the place of the superior thoracic

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 387

branch of the axillary artery in Man. No branches come from

the second and third parts of the subclavian.

The Awillary Artery has the same course and relations as in

Man. It has the additional deep relation to the axillary

prolongation of the air sac. It gives off the following

branches :—

1. Thoracico-acromial axis (T.A.A), which is not so complex

asin Man. It supplies the pectoralis major, deltoid, and the fat

and lymph glands in the axilla.

2. A muscular artery supplying twigs to the subscapularis.

3. The subscapular artery (S.U.A) gives off branches which

arise separately in Man. ‘hese are :-—a, A branch which runs

to the inferior angle of the seapula between the teres major and

latissimus dorsi, and supplies the latter; 6. A humeral trunk,

which divides into the anterior and posterior circumflex arteries.

These form an anastomosis round the neck of the humerus, and

the posterior circumflex gives a descending branch, which anasto-

moses with the profunda branch of the brachial artery; c. artery

to the teres major; the main stem then runs down the axillary

border of the subscapularis, and ends in the infraspinatus at

the inferior angle of the scapula. Its circumflex branch supplies

the infraspinatus, passes through the great scapular notch, and

ends in the supraspinatus. There is no marked anastomosis

round the scapula as there is in Man.

The Brachial Artery (text-fig. 39 B) differs from that in Man.

It becomes the ulnar artery three inches below the internal

humeral condyle, It lies superficial to the median nerve through-

out, as in the Cercopithecide. Its branches are :—

1. Superior Profunda (S.P.A) divides into twe branches,

which embrace the musculo-spiral nerve. One branch follows

the nerve, anastomoses with the descending branch of the pos-

terior circumflex artery, supplies the triceps and ends in it. An

ascending twig ends in the latissimus dorsi. The other branch

curves mesially round the humerus, and supplies the deltoid and

triceps.

2. A large artery (M.A) to the biceps, braehialis anticus, and

triceps.

3. Three muscular arteries to the brachialis anticus (Br.A).

4, 'T'wo muscular arteries to the biceps (Bi).

5. Articular to the elbow joint (Art.A).

6. Radial artery (R.A).

7. A muscular artery to the triceps and muscles arising from

the external supracondylar ridge and external epicondyle

Ge A).

8. Articular artery to the elbow joint.

9, Muscular arteries to the deep flexors and extensors (D. F).

10. Radial recurrent artery (R.R.A).

11. Muscular to the superficial flexor muscles (S.F. ):

12. Anterior interosseous artery (A.I.A).

388 DR. C. F. SONNLAG ON THE ANATOMY,

There is no anastomosis round the elbow joint as there is

in Man.

The Radial Artery (text-fig. 39 A), which is the largest vessel

in the forearm, runs down the forearm very superficially, curves

round the back of the wrist and over the trapezium, and passes

into the first interosseous space, so it is divisible into three parts

as in Man. The first part, lying in the forearm, gives off the

following branches:—1. A large muscular artery to the supi-

nator brevis (S.B); 2. Numerous fine twigs to the superficial

flexor muscles; 3. Anterior radial carpal artery (A.R.C.A),

which arises in the upper third of the forearm, runs down

parallel to the radial artery, and crosses behind it in the lower

part of the forearm. It ends by a series of arteries over the

palmar ligaments of the inferior radio-ulnar, radio-carpal, inter-

carpal, and carpo-metacarpal jomts. It also supplies the flexor

museles; 4. Superficialis vole (S.V.A) runs downwards and in-

wards across the thenar eminence and supplies its muscles. And

it is continued along the inner border of the pollex. It gives off

a fine twig which curves inwards and helps to form the irregular

superficial palmar arch. The second part of the artery, lying on

the trapezium, gives off twigs to the dorsal aspect of the inter-

carpal joints (D.C.B), and a muscular artery to the first dorsal

interosseous muscle (D.I.M). From the dorsal carpal branch

there rises a vessel to the adjacent sides of the dorsal aspect of

the index and medius. The third part of the artery hes in the

interval between the palmar aspects of the index and _ pollex.

It gives off a thenar artery (T.A) to the thenar muscles, a

muscular artery to the first dorsal interosseous muscle (D.I.M),

several adductor twigs and branches to the lumbricales. Over

the heads of the metacarpal bones it gives a branch to the radial

side of the index finger.

The Ulnar Artery (text-fig. 39 B, U.A) runs downwards as

in Man, curves round the mesial aspect of the pisiform bone and

enters the palm. It bifurcates about the middle of the palm.

One branch runs to the inner border of the minimus; and the

second divides into two branches which supply respectively, the

adjacent sides of the minimus and annularis, and annularis and

medius, From the latter branch, two arteries pass to joi with

branches of the radial artery and form the superficial and deep

palmar arches. As the artery turns round the pisiform it gives

off a dorsal branch which curves round the ulnar border of the

manus to supply the tissues on the back of the ulnar border of

the carpus.

The anterior interosseous artery (text-fig. 39 B, A.I.A) is as

in Man.

Three palmar arterial arches are present (text-fig. 39 A) :—

(1) The deep arch (D.A), lying in front of the carpus, is formed

by a branch of the ulnar artery, the superficialis vole, and the

branch of the radial artery to the thenar eminence; (2) the

superficial arch (S.A), lying in the front of the deep arch, is

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 389

formed by the superficialis vole and a branch of the ulnar artery;

(3) a digital arch (Dig.A) over the metacarpo-phalangeal joints

of the index and medius is formed by a branch of the ulnar

artery with the continuation of the radial artery. ‘The deep and

digital arches are connected by a thick vessel.

The wrist joint is supplied by dorsal branches of the radial and

ulnar arteries, the anterior radial carpal artery, the anterior

interosseous artery, and the deep palmar arch. There is no

anastomosis round the elbow joint.

The arrangements of the arteries in the pectoral extremity

favour a relatively slower circulation than in Man. The pro-

funda arteries break up into a much larger number of branches,

and the brachial artery terminates in a large number of vessels

which run distally in long, parallel trunks. Consequently the

frictional resistance resulting from more numerous branches,

combined with the relatively smaller and more uniform brachial

artery slow the circulation much more. The addition of a third,

or digital, arterial arch is an additional factor. As there are no

anastomoses round the joints and scapula, the connections must

be more numerous in the muscles, which will consequently play

an important part in maintaining the circulation. And finally,

the vascular arrangements are such that the head, neck, and arm

get a relatively greater supply of blood than do the thorax,

abdomen, and legs.

Arteries of the Pelvic Extremity.

The femoral artery begins about the middle of Poupart’s

ligament and courses downwards for an inch and a half. Then

it gives off the profunda and is continued as the superficial

femoral artery. The latter passes between the two parts of the

adductor magnus and becomes the popliteal artery. There is no

adductor canal. The common femoral artery gives off a trunk

which divides into an abdominal artery and the mesial circum-

flex artery; and the former, after giving a nutrient aitery to

the ilium and the deep epigastric, is continued as the obturator

artery. From the common femoral artery the deep circumflex

iliac artery also arises. The profunda gives off the lateral

femoral circumflex artery, and the superficial femoral artery

gives off the saphenous artery, which goes down to the foot.

The deep epigastric artery runs up in the sheath of the rectus,

but does not anastomose with a superficial epigastric branch of

the internal mammary artery.

The obturator artery passes through the obturator foramen

after running down over the horizontal ramus and back of the

pubis. It supplies the symphysis pubis and muscles attached to

the bone around the foramen.

The mesial femoral circumflex artery runs down over the head

of the femur under the adductor muscles, and supplies the

capsule of the hip joint, psoas, obturator internus, and adductor

Proc. Zoot. Soc.—1923, No XX VI. 26

390 DR. C, F. SONNTAG ON THE ANATOMY,

magnus. It then passes round to the back of the leg and gives

branches to the adductor magnus, quadratus femoris, biceps, and

gluteus maximus: it also gives off the arteria comes nervi

ischiadict.

The deep circumflex ihae artery runs up to the ilium. It

supplies the sartorius and ilio-psoas, and ends between the

internal oblique and transversalis abdominis.

The profunda femoris gives off the lateral circumflex and a

branch passing back under the rectus femoris to the gluteus

medius.. It then passes through the middle head of the adductor

magnus, supplies the adductor longus and vasti, and ends in the

biceps. There is no series of perforating arteries as in Man.

The lateral femoral circumflex artery gives off :—1. an ascend-

ing branch to the glutei, rectus femoris, and hip joint; 2. a

transverse artery to the gluteus maximus, vastus externus, and

hip joint; 3. a descending artery to the rectus femoris, vastus

externus, crureus, and hip joint.

The popliteal artery gives off muscular twigs to the heads of

the gastrocnemius, an articular artery to the knee and a geni-

cular trunk, the latter dividing into three branches:—l. a

Jateral geniculate artery which gives a nutrient artery to the

femur, a branch to the back of the joint and one which passes

round to the front of the capsule; 2. a mesial genicular artery

which supplies the capsule tn the popliteal space; 3. a descending

artery which supplies the popliteus and passes through between

the tibia and fibula to the anterior tibial muscles.

The posterior tibial artery continues the popliteal. It gives

off a recurrent branch which anastomoses with the geniculars.

A long branch, corresponding to the human anterior tibial

artery, descends to the lower end of the posterior tibial region

and curves forwards to the anterior tibial region ; it gives off

the following branches:—1. an artery which anastomoses with

the saphenous artery and helps to form the arterial arcade on the

dorsum of the foot; 2. muscular arteries; 3. malleolar arteries ;

4. nutrient artery to the fibula; 5. articular arteries to the

ankle. The arterial arch on the dorsum gives digital arteries to

all toes except the hallux. The saphenous artery, after forming

the arterial arcade, supplies the tarsal joints and gives an artery

which replaces the dorsalis pedisin Man. This dips in between

the hallux and first toe and gives a branch to the lateral side

of the hallux and several muscular arteries. It then passes

between the heads of the adductor hallucis and anastomoses with

the deep branch of the lateral plantar artery to form the plantar

arch. The latter vessel is the terminal branch of the posterior

tibial artery.

The posterior tibial artery divides under the laciniate ligament

into medial and lateral plantar arteries. The lateral plantar

artery gives off the lateral caleanean artery to the skin of the

heel and branches to the flexor brevis digitorum, accessorius,

and abductor minimi digiti. It then divides into superficial

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 391

and deep divisions. The former continues as digital branches

to the outer two digits; the latter passes between the heads of

adductor hallucis and supplies the interossei, tarso-metatarsal

joints, and anastomoses with the vessel corresponding to the

dorsalis pedis to form the plantar arterial arch. The medial

plantar artery gives off medial caleanean branches to the skin

of the inner side of the sole of the foot and muscular branches

to the abductor hallucis and flexor brevis hallucis and digital

arteries to the inner three toes.

Veins of the Thorax.

Innominate Veins (text-fig. 42 B):—The left vein runs as in

Man and unites with the more vertical right one to form the

superior vena cava, It receives inferior thyroid (I.T.V), thymic

(T.V), internal mammary (I.M.V) and superior intercostal

(L.I.V) veins. The short right vein only receives the forma-

tive vessels.

The superior vena cava (S.V.C) is large, vertical, and enters

the upper part of the right auricular appendix. It receives the

vena azygos major asin Man. The thoracic part of the inferior

vena cava is 2°6 cm. long.

The azygos veins drain the lower nine spaces, the first three

being drained by the superior intercostal vein. All are small.

No ascending lumbar veins were found. The vena azygos

major enters the superior vena cava at the level of the fifth

dorsal vertebra.

Veins of the Head and Neck (text-fig. 42 A).

The intra-cranial blood sinuses have the same general arrange-

ment asin Man. ‘The chief difference lies in the union of the

inferior petrosal and lateral sinuses within the jugular foramen

to form the internal jugular vein. The groove in the skull for

the right lateral sinus is much larger than that for the left, and

a very shallow bony groove connects the two.

The anterior facial vein (A.F.V) begins by the confluence of

palpebral and lateral nasal veins. It runs downwards and back-

wards and crosses the mandible at the anterior border of the

masseter. It crosses the levator anguli oris and buccinator, and

it is covered by the zygomaticus, risorius, and platysma. It runs

between the mandible and sub-maxillary gland, and then under

the stylo-hyoid and posterior belly of the digastric. Finally it

unites with the temporo-maxillary vein to form the external

jugular vein. It receives the following tributaries :—(1) Pal-

pebral veins (P.V) from both eyelids. (2) Lateral nasal veins

(L.N.V), (3) Masseteric veins (M.V). (4) Deep facial vein

(D.F.V), which runs under the malar bone and buccal pad of

fat to the pterygoid region. (5) Submaawillary glandular vein

(S.M.G). (6) Lingual vein (L.V). (7) Laryngeal and superior

26*

392 DR. C. F. SONNTAG ON THE ANATOMY,

thyroid venous trunk (L-T.V). (8) Pharyngeal veins (Ph.V).

No vena transversa exists.

The temporo-maxillary vein (T-M.V) is formed within the

parotid gland by the union of the internal maxillary (I.M.V)

and superficial temporal (8.T.V) veins. It drains the side of

the head and pterygoid region and parotid gland. It receives

the very short internal jugular vein (1.J.V), passes under the

Text-figure 42.

The cephalic veins (A) and thoracic thymus gland (B). L.I.V: superior inter-

costal vein ascending from hehind the heart to enter the innominate vein ;

P.B.D: posterior belly of the digastric muscle; T.G: thymus gland (the

small veins and arteries on its surface are the thymic vessels, described in

the text as T.V.). Other letters in text.

posterior belly of the digastric and stylo-hyoid muscles and

unites with the anterior facial vein to form the external jugular

vein. The vein shows no trace of a division. No jugular bulb

is present on the internal jugular vein.

It has been shown on p. 330 that there is no close pterygoid

plexus, and the pharyngeal veins do not form a rich plexus.

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE, 393

The external jugular vein (H.J.V) does not lie on the surface

of the sterno-mastoid. It dips down and lies on the surface of

the common carotid artery, vagus and sympathetic. At the

outer border of the first rib it unites with the subclavian vein to

form the innominate vein.

Veins of the Abdomen.

The Portal System (text-fig. 43).

The general arrangement of the tributaries of the portal system

is the same as in Man, but there are differences in detail. The

vein formed by the confluence of gastric, splenic and inferior

mesenteric veins unites with a large trunk formed by pyloro-

duodenal, superior mesenteric veins and the veins from the

Text-figure 43.

LESSER CURVE

JEJUNUM

The portal vein. Letters in text.

transverse colon to form the portal vein. And the veins from

the transverse colon form a connecting loop between the two

systems. The portal vein begins behind the pancreas and ascends

to the portal fissure in the liver behind the hepatic artery and in

front of the foramen of Winslow. It divides into two large

branches which enter the liver. The main vein is two and a

half inches long. It is behind the pancreas, but it is later accom-

panied by a process of pancreatic tissue. It is surrounded and

394 DR. C. F. SONNTAG ON THE ANATOMY,

accompanied by numerous sympathetic nerves and lymphatics.

The arrangement of the system facilitates a slow and even flow

of blood from the digestive organs.

Tributaries :-—

1. Splenie vein (S.V) formed by several veins from the hilus

of the spleen. It passes through the lieno-renal ligament and

unites behind the pancreas with the inferior mesenteric vein.

Tt receives (a) vasa brevia (V.B) from the body of the stomach ;

(b) left gastro-epiploie vein (L.G.E.V) which runs along the

greater curvature of the stomach, receiving veins from it, and

connects the spienic and duodenal veins; (¢} pancreatic veins.

2. Inferior mesenteric vein (1.M.V), which communicates with

the superior mesenteric vein by a vessel whieh runs through

the transverse meso-colon and supplies the transverse -colon

(T.M.C.V). Tt drains the large bowel from the splenic flexure

to the beginning of the rectum, and it unites with the splenic

vein.

3. Superior mesenteric vein (S.M.V), which drains the ileum

and jejunum, and the large intestine from the appendix to the

hepatic flexure, communicates with the venous arch in the trans-

verse colon. At the point where it participates in the formation

of the portal vein it receives the venous arch of the transverse

colon, duodenal veins (d.v) and gastric veins (g.v).

4. The cystic vein (e.v) enters the portal trunk itself.

The Inferior Caval System.

The vena cava inferior is formed by the union of the two

common iliac veins deep to the right common iliac artery. It

ascends on the right side of the abdominal aorta. In the upper

part of the abdomen it bends to the right and passes through

a tunnel in the liver. Its relations are much as in the human

body, but the right ovarian artery passes behind it. It

receives :-—

1. Four single lumbar veins, the first or lowest entering the

left side of the vein, and the others pass into its dorsal surface.

2. Right ovarian vein.

3. Two renal veins.

4. Right suprarenal vein.

5. Gastric vein which serves as a link between the systemic

and. portal circulations.

6. Hepatic veins.

Veins of the Pelvie Extremity.

Superjicial Veins ;—The venous arch on the dorsum of the foot

receives veins from both sides of the digits. The inner extremity

is continued upwards by two internal saphena veins, which are

united by cross branches. They pass upwards, dip under the

sartorius and end in the femoral vein. The large external

saphena vein runs up the back of the leg, dips through the fat

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 385

in the popliteal space and enters the femoral vein. There is no

upward vein running through a saphenous opening, and that

opening is a human characteristic.

Two venz comites accompany aJl the branches of the posterior

tibial artery. They unite to form one popliteal vein which

accompanies the artery and becomes the femoral vein. The

venous circulation closely follows the arterial supply, but no

epigastric vein enters the femoral. The saphenous veins open,

as described above, into the popliteal and femoral veins.

The veins of the pelvis follow the branches of the hypogastric

artery, and the hypogastric vein joins with the external iliac

vein to join the common iliac vein. The two common iliac veins

unite to form the vena cava inferior. These veins have rela-

tions similar to those in Man.

Veins of the Pectoral Extremity.

The venous circulation differs in several points from that in

Man. The veins of the hand unite to form the cephalic vein

which only extends up as far as the antecubital fossa. There

it dips inwards and unites with vene comites following the

branches of the brachial artery to form the brachial vein. No

basilic vein is present. The brachial vein runs upwards, re-

ceiving tributaries corresponding to the branches of the artery.

It is successively followed by the axillary and subclavian veins

which receive tributaries corresponding to the branches of the

arteries. The subclavian veins unite with the external jugular

veins to form the innominate veins. The venous circulation

differs from that of Man in the shortness of the cephalic vein,

the absence of the basilic vein, the presence of a brachial vein

instead of venz comites, and the absence of an internal jugular

vein uniting with the innominate vein.

Tue Ducriess GLANDS.

The thyroid gland (text-fig. 41) is long, narrow, and thin.

The lateral lobes are bent on themselves at the upper ends,

which lie against the criccid and lower end of the thyroid

cartilage. The thicker isthmus crosses the fourth and fifth

tracheal rings. There is no strong capsule and no pyramidal

lobe. It receives a complicated series of arterial anastomoses

from the superior (S.T.A), middle (M.1T.A), and inferior (1.T.A)

branches of the external and common carotids, and the thyroidea

ima (T.I.A) from the left common carotid. No subclavian

branches pass to the gland. The superior thyroid vein (8.T.V)

opens into the anterior facial vein, and the inferior thyroid vein

L.T.V) goes to the innominate vein.

At the lower border of the isthmus there is, on each side, an

oval body, the size of a pea, consisting of the parathyroid gland

and a piece of thymus. No other parathyroid tissue was present.

396 DR. C. F. SONNTAG ON THE ANATOMY,

The thymic constituent consisted mainly of concentric corpuscles

and little lymphoid tissue. These conditions are, however,

individual peculiarities.

The thymus (text-fig. 42), lying in the thorax, consisted of a

large left part reaching the level of the third costal cartilage,

and a small right part reaching the second cartilage. These

parts touched over the pericardium and the left part sent a

process up under the great veins. Both parts have coarse

lobules, and no cavity is present in either. It is supplied by

the internal mammary artery, and the veins enter the left

innominate vein.

The spleen is small, measuring 3°8 ins. long, 2°4 ins. wide and

1:2 ins. thick. It has the same shape as in Man. The hilum

is elongated. A small, oval accessory spleen is present. The

artery is smaller than the hepatic artery. Other examples have

larger spleens; but the form and size depend on the stages in

digestion.

The suprarenal capsules are elongated bodies, with rounded

ends, lying in the usual positions. They receive their arteries

from the phrenic and renal arteries, but none from the abdominal

aorta. The suprarenal plexuses are well marked.

THE Broop.

Gulliver (23) pointed out that the red blood corpuscles have a

diameter of 1/3412 inch, whereas those of Man are 1/3200 inch

wide. The precipitin reactions have been described by Nuttall

(38) who found that the blood of the Chimpanzee gives strong,

positive reactions with those of Homo and Simia, but he does

not mention its reaction with those of Gorilla and Hylobates.

Tse LyMpHaAtic System.

The thoracic duct arises from a receptaculum chyli of con-

siderable size by two vessels, which unite later. One vessel runs

upwards on the right side of the thoracic aorta, and the other

courses on the posterior surface of the esophagus. At the level

of the sixth dorsal vertebra the two vessels unite to form a

trunk, which runs up between the cesophagus and vertebral

column, and to the left side of the former. It then passes into

the neck where it opens into the junction of the left subclavian

and jugular veins.

No right lymphatic duct was detected.

The lymphatic glands are fewer than in Man, and the following

groups were isolated :—

A. Glands in the Head and Neck:—1. A row of small glands

lying on the surface of each submaxillary gland; 2. A group of

both large and small glands between the cleido-masteid and the

larynx; 3. No glands were found on the surface of the parotid

or along the great vessels; 4. A group of glands over the sub-

occipital region.

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 397

B. Glands in the Pectoral Hxtremity:—1. Two small glands

on the axillary surface of the teres major receiving vessels

along the axillary vessels; 2. A row of glands, both large and

small, along the course of the long thoracic artery. It is

divisible into an upper group draining the glands on the teres

major, and a lower group draining the side and back of the

thoracic parietes; 3. No delto-pectoral nor cubital glands were

found.

C. Thoracic Glands :—1. Several glands in the pulmonary

roots; 2. Three small glands among the cardiac plexuses; 3. No

retro-sternal nor vertebral glands were found.

D. Abdominal Glands:—1. Several small glands along the

lesser gastric curvature; 2. Several small and large glands on

the greater gastric curvature; 3. Numerous glands between the

layers of the mesentery; 4. A chain of glands along the common

iliae vessels.

E. Glands in the Pelvic Hxtremity:—1. A group of glands

close to the mid point of Poupart’s ligament.

It is, therefore, evident that the groups of lymphatic glands

are fewer than in Man.

RESPIRATORY ORGANS.

The external nose is small, fat, and has no lateral cartilages.

It is surrounded by a groove in the upper lip. Its muscles and

nerves have already been described (see p. 328). The vestibule

is well marked, and has numerous vibrissee. The mucosa lining

the nose has the orifices of numerous glands, and the upper

fourths of the septum and lateral wall have striations produced

by the olfactory nerves. The septum is as in Man, but I could

not detect any pit corresponding to Jacobson’s organ. The

inferior turbinate bone (Pl. II. A, 1.2.8) is long and almost

horizontal; it is prolonged backwards by a fold of mucous

membrane. The inferior meatus receives the naso-lachrymal

duct (N.L.D) in its middle part. The middle turbinate bone

(M.T.B) is bifid posteriorly, and is shorter than the lower one.

On elevating it, a movable mucosa-covered bony crest is revealed,

and between them lies the opening of the frontal sinus (F.S)

in the middle meatus. But there is no actual bulla similar to

that in Man. Above the middle turbinate bone there are three

turbinal crests with four grooves. The longest is the swperior

turbinate bone (S.T.B). The sphenoidal sinus (8.8) is large and

opens into the upper turbinal region. It is undivided and

excavates the alisphenoids. The frontal sinus is narrow. The

antrum of Highmore (Pl. II. B) is large, strengthened by

buttresses, and has elevations produced by the roots of the

canine, premolar and molar teeth. The turbinal region has been

mentioned by Zuckerkandl (55), Keith (64), and Paulli (63). It

has several air cells in its walls.

398 DR. C. F. SONNTAG ON THE ANATOMY,

Larynx * :—The thyroid cartilage is shaped somewhat differently

from that in Man, for it has median notches both above and below.

Theangle between itsalez is about 90°. Its superior and inferior

cornua articulate, as in Man, with the hyoid bone and cricoid

cartilage. The cricoid cartilage is as in Man. The arytenoid

cartilage is shaped as in Man, and the cartilages of Santorini and

Wrisberg are very small. The epiglotiis has the same shape as in

Man. It is small and does not rise freely above the level of the

aryteno-epiglottidean folds. The true vocal cords are attached to

the thyroid ale and vocal processes of the arytenoid cartilages.

They are soft and flaccid, consisting almost entirely of mucous

membrane, and a little elastic tissue. The false cords are

likewise soft, and between them there are well-marked ventricles.

The latter extend upwards behind the false cords to the

aryteno-epiglottidean folds, and they are prolonged upwards by

well-marked diverticula to the air-sac, which begins above at the

excavated hyoid bone and extends downwards even into the

axille. The cervical part of the pouch has already been

described. ‘This pouch is much larger than that described in

many other animals, and resembles that in Simia in its extent.

The communications between the sac and ventricles pierce the

thyro-hyoid membrane, which is large. ‘The crico-thyroid

membrane is as in Man. The laryngeal joints (crico-thyroid,

thyro-hyoid, crico-arytenoid) are as in Man.

Laryngeal Muscles:—The thyro-hyoid muscle runs from the

lower half of the thyroid ala to the lower border of the hyoid

bone. The crico-thyroid muscle runs from the anterior two-thirds

of the lower border and outer surface of the cricoid cartilage to

the lower border of the thyroid cartilage anterior to the inferior

cornu. It is not fan-shaped as in Man. The posterior crico-

arytenoid muscle arises from the posterior cricoid lamina, and is

inserted into the processus muscularis of the arytenoid. It is

more vertical and not so fan-shaped as in Man, and a small

branch of the superior thyroid artery runs on its posterior

surface. The lateral crico-aryienoid muscle runs from the upper

border of the lateral part of the anterior cricoid arch to the

processus muscularis. ‘The thyro-arytenoid muscle does not divide

asin Man. It runs from the inner surface of the thyroid ala in

its lower half, near the mid plane, to the outer border of the

arytenoid cartilage above the crico-arytenoid. The oblique and

transverse arytenoid muscles are as in Man, but are diminutive,

as is the ary-epiglottidean muscle.

Interior of the Larynu:—The ary-epiglottic folds are almost

absent. Posteriorly the cuneiform tubercles and tubercles of

Santorini are close together. The pyriform sinus is more

marked than in Man. The upper division of the larynx is

shallow, and the cushion of the epiglottis is well marked. The

middle division is relatively larger than in Man. ‘the false cords

are 4mm. apart. The lower part of the eavity is as in Man.

* Excellent illustrations have been published by Gratiolet (22).

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 399

The trachea has nineteen rings, all of which have cartilaginous

hoops, with the gaps behind. ‘The first is very wide.

Lungs :—The left lung has upper and lower lobes, and the

right one has upper, middle, and lower lobes. No azygos lobe is

present. ‘The right lung receives a large bronchial artery from

the descending thoracic aorta, but the left one receives a large

artery from the concavity of the aortic arch, and a fine thread-

like vessel from the descending aorta. Mayer (84) opserved

three lobes in the left lung, and two in the right. But Tyson

(50), Vrolik (51), Gratiolet (22), Chapman (12), Hartmann (65),

Sperino (47), and Symington (48) observed conditions similar to

those in my specimen. Bischoff (60) observed two lobes in the

left lung, and four in the right in an old animal. The bronchial

glands are of moderate size, and adherent to the bronchi.

The limits of the pleura were described by Ruge (43). They

presented nothing remarkable in my specimen.

UROGENITAL ORGANS (text-fig. 44).

Kidneys :—It is frequently stated that the Primates, with the

exception of Man and Aéeles, have only one renal papilla. But

in this specimen, end in former animals examined by me there

were respectively four and five. Ehlers (59), and Bischoff (60)

noted one papilla, Symington (48) found the pyramids fused to

form one papilla, and Sperino (47) noted three papille. The

right kidney reaches lower down than the left one, and the

measurements are :—

Left kidney :—5:5 em. long; 3°3 em. wide; 1°6 cm. thick.

Right 7 66 ,, cy) 3°38, re) 18 ,, ”

There is much peri-renal fat, but no fascial shelf supports the

kidney ; and there is no fat in the pelves, although Sperino (47)

observed some. The capsule, which strips easily, is well

vascularised. The cortex is thick, and sends prolongations in

between the pyramids, which are finely striated. The blunt

apices of the pyramids do not project much, and they are not

embraced by large calyces. The relative positions of the

structures in the hilum are as in Man, and the suprarenal

capsules are similarly placed. The shape of the kidneys alters

with the movements of the viscera apposed to them.

The wreters* have the same course, relations, and terminations

as in Man, but I was unable to detect lymphatics running along

them between the kidneys and bladder.

The bladder is thick and muscular, but no urachus nor anterior

ligaments are present. Lateral ligaments are well marked; and

the thick round ligaments of the uterus are connected to them by

peritoneal folds, so the utero-vesical pouch is crescentic. The

mucosa is thick and corrugated, and the ureteric papille he at

the ends of prominent ridges. The patulous urethral orifice is

close to the ridge, so the trigone is small. The vesical

musculature consists of two layers. There is an external

400 DR. C. F. SONNTAG ON THE ANATOMY,

longitudinal layer, half an inch wide,in the middle line. Lateral

to this band the fibres run obliquely, both upwards and

downwards, and interlace with one another. The submucous

coat is thick, and composed of more elastic areolar tissue than in

Man. The mucous membrane is loosely attached to it. The

serous coat exhibits nothing peculiar.

Text-figure 44.

The internal organs of generation. A: anterior aspect; B: posterior aspect. B.L:

broad ligament; C.U: cervix uteri; F: fimbrie; F.T: Fallopian tube;

M.U: meatus urinarius; O.A., O.L., O.F.,O.Vn: ovarian artery, ligament,

fimbria, and vein; R.L: round ligament; V.C: vaginal columns. Other

letters in text. The arrows in figure A show the natural inward curvature

of the Fallopian tubes.

Ovaries (text-fig. 44, O):—The left ovary is long, narrow and

very thin, measuring 2°5x°5x-2cm. The right ovary is flat and

rounded, measuring 155x1:5x°2cm. Neither has any super-

ficial scars nor rugs. Each lies vertically behind the broad

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 401

ligament at a higher Jevel than the uterus. The histology has

been described by Giacomini (66), Duval (67), and Sperino (47).

Ligaments connect it to the utero-tubal junction, and to the

tube below and behind the fimbriz. The ovarian fimbria is well

marked. Sperino describes triangular ovaria. The primordial

ova are innumerable, and are similar to those in the human

species. And Graafian follicles can be seen in various stages of

development according to Sperino.

Fallopian Tubes :— Both are 65cm. long when drawn straight.

They hardly increase in calibre from their uterine to their

ovarian ends. The fimbria form a dense cluster, the ovarian

fimbria is well marked, and uterine and abdominal orifices are

plain; but one cannot easily pass a bristle through the tube.

Each tube curves uver the anterior border and upper pole of the

corresponding ovary. The hydatid (text-fig. 44, H.M) is well

marked on the right side. The epoophoron and paroophoron are

present.

Uterus (text-fig. 44) :—The uterus is isolated from the bladder

and rectum by peritoneal fosse, and its summit lies 1°5 em. above

the floor of each. There is no marked fundus, the body is

triangular and the cervix is fusiform. The body is 1°5 cm. long,

and its base is 1°5 cm. across. The cervix is 1:2 cm. long, and

1-1 cm. across at its widest part. It has very infantile propor-

tions. The round ligaments are large and run directly upwards

and forwards from the utero-tubal junction. The interior of the

body of the uterus is smooth between the tubes, but lower down

it has an upward continuation of the median dorsal crest and

transverse ridges which occupy the cervix. The musculature in

the upper part of the uterus is thinner than in the lower part of

the body and the cervix. The external os uteri is oval, with

nodulated continuous lips. Both lips are of equal length. This

account differs in several respects from the accounts of Sperino

(47), and others. Gratiolet (22) described a bicornuate uterus.

The vagina is 5 cm. long, and expands from above downwards.

Anterior and posterior fornices are both present, but the latter is

much the larger. In its upper part there is a median dorsal

cushion, and the mueosa has transverse folds. Below that it has

longitudinal folds. In its lower part it has fine longitudinal

strie and several pockets (text-fig. 44). The urethra opens on

its anterior wall about the middle.

The uterine artery (U.A) supplies the vagina, uterus, tubes,

ovaries, epoophoron, etc. It anastomoses with the very small

ovarian artery. Its complexity is shown in text-fig. 44.

The external generative organs (Plate I. B) are built on the same

plan as, but differ from those of the human female. The mons

veneris (M.V) is slight, and has a few hairs. The labia majora

(L.M) are represented by slight elevations of skin over thick-

enings of the subcutaneous fat. The labia minora (L.Mi) are

large and folded, and divide to surround the large clitoris (CL)

the latter having two crura covered by well-developed ischio-

cavernosi muscles. A small fourchette exists, but there is no

402 DR. C. F. SONNTAG ON THE ANATOMY,

hymen. The meatus urinarius is within the vagina, so no

prominent vestibule is seen as in the human condition. The

glands of Bartholin lie between the vagina and rectum. Sperino

(47), Bischoff (60), Chapman (12), Gratiolet (22), Hartmann (65),

Barkow (2), Hoffmann (68), Symington (48), and Traill (49)

have described the external genitalia; and many of these ob-

servers have described the internal organs.

Winwoode Reade (57), Garner (21), and Mohrike (35) describe

a sexual season, and Bolau (5), Ehlers (59), Hermes (69), and

Keith (30) describe either the periodicity or characters of men-

struation. Pocock (80) contrasts menstruation in the Chim-

panzee and Hainan Gibbon.

Tue NERyous System *.,

The olfactory nerve terminates by marked branches on the

upper thirds of the turbinate regions and nasal septum.

The optic nerve is large and surrounded by a sheath of dura

mater. No arteria retine centralis was detected in it, but the

injection material may not have traversed it.

The oculo-motor nerve has superior and inferior divisions. The

superior division does not pierce, but runs to the inner side of,

the superior rectus. It supplies the superior and internal rectus

muscles and ends in the levator palpebre superioris. The inferior

division runs downwards and outwards on the outer side of the

rectus inferior, gives a motor branch to the ciliary ganglion,

supplies the inferior rectus and ends in the inferior oblique. The

branch of the superior division to the internal rectus is very large.

The trochlear nerve ends by three divisions to the superior

oblique muscle.

The trigeminal nerve has three divisions as in Man, radiating

from the Gasserian ganglion. The ophthalmic division courses as

in Man, and breaks up into:—1. Lachrymal nerve, lying between

the orbital wall and upper border of the external rectus. It

supplies the lachrymal gland, conjunctiva and skin of the eyelids.

2. Frontal nerve resembles that in Man. It breaks up into

supra-orbital and supra-trochlear branches. 3. Wasal nerve.

This is distributed as in Man, but the lateral terminal branch,

which is very large, comes out of the nasal cavity direct, and not

between bone and cartilage, asin Man. The ciliary ganglion is

larger than in Man. It lies on the lateral side of the oph-

thalmic artery and receives filaments from both divisions of the

third nerve, a twig from the naso-ciliary nerve, and sympathetic

filaments from the carotid plexus. It gives off short ciliary

nerves: one large one, lying on the outer side of the optic nerve,

divides into upper and lower divisions on reaching the eyeball.

The superior and inferior maxillary divisions of the trigeminal

are similar to those in Man, but I was unable to detect as many

branches radiating from Meckel’s ganglion. The chorda tympani

* The brain will be described in a separate paper by Professor G. Elliot Smith,

E.RBS.

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 403

joins the inferior maxillary division before the latter separates

into its lingual and inferior dental nerves. The submaxillary

ganglion is not separate as in Man, but is fused with the

hypoglossal nerve. The otic ganglion was not recognised with

certainty.

The abducens emerges between the two heads of the external

rectus and sinks into the ocular surface of the muscle.

The facial nerve emerges from the stylo-mastoid foramen. Its

intra-petrous course was not traced. It divides in the parotid

gland into temporal. zygomatic, maxillary, buccal, mandibular, and

cervical divisions. The temporal branches run upwards and are

distributed as in Man. The zygomatic and maxillary divisions

eventually unite and give off from their combined trunk a

number of branches to the muscles of the face. The mandibular

and cervical divisions are as in Man. The union of the chorda

tympani and trigeminal nerves has already been deseribed.

The auditory nerve was not traced.

The glosso-pharyngeal nerve emerges from the inner part of the

jugular foramen and communicates with the other nerves at the

upper part of the neck. It passes between the external and

internal carotid arteries, curves round the stylo-pharyngeus

muscle and disappears under the free outer edge of the hyoglossus.

Finally it breaks up into branches to the tongue, pharynx, and

tonsil. It supplies the stylo-pharyngeus. The tympanic and

petrosal branches were not traced.

The Vagus Nerve (text-figs. 45 & 46) emerges from the jugular

foramen wherein it is lateral to the glosso-pharyngeal nerve,

posterior to the internal jugular vein and mesial to the accessory

nerve, to which it is closely adherent. Immediately below the

base of the skull it develops the ganglion nodosum (G.N.) on its

lateral aspect. The nerve separates from the ganglion again at

the level of the posterior border of the hard palate. At the root

of the neck it runs on to the posterior aspect of the common

carotid artery and then it enters the thorax on the left side.

The right one disappears under cover of the innominate artery

where the latter bifurcates into right common carotid and sub-

clavian arteries. The left vagus (text-fig. 45 A) only communicates

with the sympathetic, but the right one (text-fig. 45 B) is ex-

tensively used with the sympathetic.

Branches in the Neck :—

1. Communicating nerves to the glosso-pharyngeal (c. ix),

hypoglossal (c. xii), superior cervical ganglion of the sympathetic

(S.C.G) and cervical plexus (¢.C.P).

2, Pharyngeal nerve (a).

3. Superior laryngeal nerve (6).

A, Right recurrent laryngeal nerve (d).

5, Cardiac branch of the left vagus ( f).

6. Plexus of carotid, tracheal and cardiac branches of the

right vagus.

404 DR. C. F. SONNTAG ON THE ANATOMY,

The left thoracic vagus (text-fig. 45 B) has the same relations

and course till it reaches the posterior aspect of the pulmonary

root as in Man. It gives off recurrent (e), cardiac (/), and

anterior pulmonary nerves. Behind the root of the left lung it

gives off posterior pulmonary nerves. It does not break up into

the posterior pulmonary plexus as it does in Man. Leaving the

back of the root of the lung it gains the front of the esophagus,

which position it maintains into the abdomen. It supplies the

cesophagus and pleura and communicates with the right vagus.

The right thoracic vagus (text-fig. 45 B) gets into the thorax

after crossing the right subclavian artery, at which point it gives

off its recurrent branch. It crosses the right side of the trachea

Text-figure 45.

The vagus and syrapathetic nerves. A: cervical parts of the left vagus and sympa- _

thetic; B: thoracic parts of the left vagus and the cervical and thoracic

parts of the right vagus and sympathetic. Letters in text.

from before backwards and it is crossed by the vena azygos major.

It passes down the back of the right pulmonary root and inclines

downwards and backwards to reach the dorsal aspect of the

esophagus. There is no plexus gule. It gives off three cardiac

nerves (f), three anterior pulmonary (g'), two tracheal (T.B),

two posterior pulmonary (/), and several cesophageal nerves.

Abdominal Paris of the Vagi (text-fig. 46):—The left vagus

passes through the anterior part of the cesophageal opening in

the diaphragm, and divides into two. One branch divides into

twigs which run along the lesser curvature as far as the pyloric

antrum. The other branch gives twigs along the lesser curvature,

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 405

twigs running along the cesophageal branch of the celiac axis,

coronary plexus, and cesophageal nerves, which ramify over the

lower end of the esophagus. It anastomoses with branches of

the right vagus. The right vagus (R.V) passes through the

posterior part of the cesophageal opening in the diaphragm,

and it ends in the left semilunar ganglion. The branches

are very numerous and extend widely in the abdomen. They

are :—

1. Gsophageal nerves (O.N) to the back of the lower end of

the cesophagus.

Text-figure 46.

Abdominal parts of the vagus and sympathetic nerves. Letters in text.

2. Gastric nerves (L.C.N) running along the lesser curvature

of the stomach.

3. Twigs to the hepatic (H.P), coronary (G.P), splenic (8),

superior mesenteric (S.M.P), inferior mesenteric (I.M.P), and

aortic (A.G.C) plexuses. The twigs can be traced far into the

plexus, some in fact being so well marked that they can be

followed to the organs. I did not trace twigs as far as the

cecum nor could I trace them to the sigmoid and rectum.

Proc, Zoou, Soc.—19238, No. XX VII, 27

406 DR. C. F. SONNTAG ON THE ANATOMY,

The Spinal Accessory Nerve emerges as in Man from the

jugular foramen, pierces the cleido-mastoid, runs deep to the

sterno-mastoid and gains the deep surface of the trapezius, where

it has already been described. It supplies the cleido-mastoid,

sterno-mastoid and trapezius, and it communicates with the

cervical plexus, but not with the sympathetic.

The Hypoglossal Nerve emerges as in Man from the skull, and

has a similar disposition till it reaches the hyo-glossus muscle.

At the anterior border of that muscle it forms a loop and exhibits

a swelling slightly anterior to it. This swelling receives filaments

from the lingual nerve, and there is no separate submaxillary

ganglion. Finally it divides into twigs for the stylo-glossus and

genio-glossus. Branches:—(1) On the left nerve there is a

strong descendens hypoglossi, but it is replaced by two branches

on the right side. (2) Nerve to the thyro-hyoid muscle. (3)

Nerves to genio-hyoid. (4) Communicating to the lingual nerve.

(5) Nerves to genio-glossus. (6) Nerves to stylo-glossus.

The Cervical Plexus (text-fig. 47).

The cervical plexus is formed from the first four cervical nerves,

and its relations are similar to those in Man; but there are

differences in the branches. The first and second nerves form a

loop. Branches of the second and third nerves form cords; a

mesial cord forms the nerve to the sterno-hyoid (S-H.M) and

a lateral cord forms the transverse cervical (T.C.N) and occipital

nerves (O.N). Branches of the third and fourth nerves form the

descending supraclavicular nerves (S-C.N). The fourth nerve

communicates with the fifth.

Branches :—

I. Superficial Cutaneous Nerves:—Small occipital (O.N) and

transverse cervical (1.C.N) from C2 and C3; Descending

branches (acromial, sternal, and clavicular) from C3 and C 4.

Il. Deep Muscular Branches to sterno-mastoid (S-M. from C 2),

trapezius (Tra. from C3 and C4), levator anguli scapule (L.A.S.

from C3), scalenus medius (Seal. Med. from C4), omo-hyoid

(O-H.M. from C€ 2), sterno-thyroid (S-T.M. from C 2), sterno-

hyoid (S-H.M. from C2 and C3), and diaphragm (by phrenic

(P.N) from C 2, C3, 04, C5).

TIT. Deep Communicating Branches to vagus (G.N), accessory

branches to the sympathetic on the left side, but the ganglion

nodosum and superior cervical sympathetic ganglion are con-

nected close to the spot where the vagus communicates with the

cervical plexus. On the right side communications go from the

sympathetic ganglion to the first and second cervical nerves.

There is no marked ansa hypoglossi.

The Phrenie Nerve (P.N) is mainly derived from the fourth

cervical nerve, but it receives fine fibres from C3, C2, and C5.

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE, 407

It passes downwards through the neck on the scalenus anticus

and it enters the thorax between the subclavian artery and vein.

Its general relations to the aortic arch, vagus, heart, and root of

the lung are the same as in Man. Close to the diaphragm it

divides into five branches which subdivide. Some of these supply

the thoracic surface of the muscle, but others pass through it to

Text-figure 47.

s.c.c. GN. xq,

Scal.Med. §

PN.

The cervical plexus. C1-C5: cervical nerves. Other letters in text.

supply the abdominal surface. It is accompanied by an artery

and a vein. It gives branches to the pleura and pericardium,

and communicates with the phrenic sympathetic plexus, but I

did not trace branches of this anastomosis to the inferior vena

cava, hepatic, and suprarenal plexuses. No arteria comes nervi

phrenici was seen.

The Brachial Plexus (text-fig. 48).

The plexus is formed by the lower four cervical and first dorsal

nerves as in Man, but the arrangements differ. Before they form

the plexus C5, C6, and C7 give off the three roots of the long

thoracic nerve (L.T.N), and a well-marked branch runs from the

upper root to the first digitation of the serratus magnus (Serr.

Mag). C7 and C08 also give twigs to the scalenus anticus (Scal.

Ant.).

C5 unites with the anterior division of C6 after giving off :—

1, A nerve to the levator anguli scapule (L.A.8S), the rhomboidei

(R.M) and the first digitation of serratus magnus (8.A.M);

2. the suprascapular nerve (S-S.N). As C5 joins a division of

C6 there is no upper trunk as in Man.

C6 divides into anterior and posterior divisions,

408 . DR. C. F, SONNTAG ON THE ANATOMY,

C7 gives off the external anterior thoracic nerve (H.A.T) and

forms a trunk which divides into anterior and posterior divisions

as in Man.

C8 and D1 unite to form a short trunk which divides into

anterior and posterior divisions. At the point of division the

internal anterior thoracic nerve (1.A.T) is given off. The two

anterior thoracic nerves are connected by a loop as in Man.

C5 and the anterior divisions of C6 and C7 unite to form a

thick cord which, after giving off the masculo-cutaneous nerve

(M-C.N), unites with the anterior division of the combined C 8 and

D1 to form a cord, which divides into median (M.N) and ulnar

(U.N) nerves. There is no separation into outer and inner cords

as in Man, but the musculo-cutaneous nerve represents the

former.

Text-figure 48.

a Ca,

Scal.Ant.

Letters in text.

The posterior divisions of C6, C7, and the combined C 8 and

D1 unite to form a posterior cord, which gives off four swbscap-

ular nerves (U.S.N., L.8., Lo.S) and divides into the musculo-

spiral (M-S.N) and circumflex (C.N) nerves.

The internal cutaneous (1.C.N), lesser internal cutaneous

(L.1.C), and a coraco-brachial (C-B.N) twigs come from the

representatives of the outer and inner cords.

The suprascapular nerve from C5, passes through the supra-

scapular notch, supplies supra-spinatus and turns through the

great scapular notch to supply infra-spinatus. As it passes

through the greater notch it gives a second branch to the supra-

spinatus.

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 409

_ The musculo-cutaneous nerve from C5, C6, C7, gives a branch

to the coraco-brachialis and then pierces the muscle. It then

gives a large branch to the biceps. Finally it divides into a

muscular.trunk to the brachialis anticus, and a cutaneous trunk,

which gives a small nerve to the spinator longus.

Four subscapular. nerves are present. The two upper ones go

to the upper and lower parts of subscapularis. The long sub-

scapular communicates with the musculo-spiral nerve and supplies

the latissimus dorsi. The lowest nerve supplies the subscapularis

and teres major.

The median nerve arises from tne anterior divisions of all the

nerves forming the plexus. It almost immediately after its

formation gives a small branch to the coraco-brachialis. No

branches arise in the arm. Just below the bend of the elbow it

supplies the flexor carpi radialis, flexor sublimis, and both heads

of the pronator radii teres. Then it communicates with the ulnar

nerve by a thick branch: In the middle of the forearm it supplies

the radial fibres of the flexor sublimis digitorum. About an

inch distal to the radio-carpal joint it bifurcates. The outer

division supplies the thenar muscles, first lumbrical, and the skin

of the radial side of the index and ulnar side of the thumb. The

inner division gives a small twig to the third and fourth lum-

bricals. Then it divides to supply adjacent sides of the second

and third and fourth digits. The nerve to the second and third

digits also supplies the second lumbrical. All the branches pass

deep to the superficial palmar arch.

The circumflex nerve gives the nerve to the teres minor before

it passes through the quadrilateral space. No definite anterior

and posterior divisions are present. After giving off the large

lateral cutaneous nerve of the arm it breaks up into deltoid

branches.

The war nerve arises in common with the median. Its course

is much asin Man. In the forearm it supplies the flexor carpi

ulnaris and flexor profundus digitorum and communicates with

the median nerve. Two inches proximal to the wrist it divides

into anterior and posterior divisions. ‘The former supplies the

hypothenar muscles, the skin of the adjacent sides of the annu-

laris and minimus and the inner side of the minimus; and the

latter goes deeply to supply the palmar interossei. A dorsal

branch leaves the main trunk at the level of the pisiform bone

and supplies interossei.

The musculospiral nerve from the posterior divisions of C 6, C7,

C8, gives off a slender, but long, nerve to the dorso-epitrochlearis.

Its course is asin Man. In the arm it gives off branches to the

triceps and skin as in Man. Im the lower part of the arm it

supplies the supinator longus and extensor carpi radialis longior.

At the bend of the elbow it divides into radial and posterior

interosseous nerves. The former runs down to the skin of the

wrist. The latter perforates the supinator brevis. It supplies

the extensores carpi radialis longior and brevior, and the muscles

410 DR. C. F. SONNTAG ON THE ANATOMY,

on the extensor aspect of the forearm. Finally it sends a long,

fine nerve to the wrist joint.

The internal cutaneous and lesser internal cutaneous nerves arise

from the conjoined median and ulnar nerves in the brachial’

plexus. They are distributed as in Man.

The intercosto-humeral nerve is as in Man.

The Lumbar and Sacral Plexuses (text-fig. 49).

The lumbar plexus is formed by the anterior primary divisions

of the four lumbar nerves, the sacral plexus is formed by the

Text-figure 49.

The lumbo-sacral piexus. D XIII-S2: nerves forming the plexus proper; S3 and

$4, not shown in the diagram, supply pelvic muscles; Pyr: nerves to pyri-

formis; H.M. and Q.F: nerves tu the hamstring muscles and quadratus

femoris. Other letters in text.

anterior primary divisions of the four sacral nerves, The two

plexuses are connected by the lumbo-sacral cord from the third

and fourth lumbar nerves. Hepburn (24) gives the cord as coming

from the fourth lumbar, Champneys (11) records a_ totally

different formation of the plexuses, but does not mention: the

cord. Bolk (7) showed that there are slight variations in

different animals.

The following table shows the origins of the nerves as observed

by myself and others :—

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE,

Nerve Boux. HEPBURN. CHAMPNEYS. Self. |

Ilio-hypogastric ...| D xiii. D xiii. | Not recorded. D xiii. D xiii.

Tlio-inguinal......... Dxui. Dxii. L1 Lil D xiii. Lil

Genito-crurals esa esse Li D xii. Ll

Anterior crural...... Ll, 2,3 Wal, WAP, ys | 10) ee IWS, |) Ving es WB

Lateral cutaneous... Li, 2 L2, L3 D xii., L1 L2,L3

Obturator ............ WZ BS INO SM Ibe We bil |) Deshi, Wii, Wye) |p I tls Wee bs}

Superior gluteal ... Tai. 9 1 L-S.C, $1 Liu., Liv., Si L-S.C, $1

Inferior gluteal Liv.,S1,S2 | Small sciatic. | Not recorded. Sup. gluteal.

Great sciatic ......... WS! CrSils S12) ease Sone L-S.C, $1, $2

Sinallisciatichweses ir orn teacc. Not recorded. | Liu., Liv., S1 $2

Pudendal ............ $2 Not recorded. | Not recorded. $2

N. Obturator Int....|_ Tibial nerve. $1,82 Not recorded. $2

N. Pyriformis ...... Peroneal nerve. S82 $2 $2

N. Quadratus Fem.| ‘Tibial nerve. $1, $2 Not recorded. Great sciatic.

N.to Gemelli ......) 9... $1, 82 Not recorded. $2

of Man.

lt is also evident that the branches of the plexus are

liable to considerable variation in different animals, and I found

some differences on both sides in my own.

For example, the

obturator nerve came from L2 and L3 on the right side, but

from L1, L2, L3 on the left.

The ilio-hypogastric nerve (I-H.N) is the continuation of

Dxiui., and it is distributed as in Man.

The «aio-inguinal

(1-I.N) and genito-crural (G-C.N) nerves are also as in Man;

the former is large.

The anterior crural nerve (A.C.N) divides into anterior and

posterior divisions in the upper part of Scarpa’s triangle. The

former gives off a cutaneous patellar branch, the cutaneous

saphenous nerve and muscular branches to the sartorius, gracilis,

and pectineus. The posterior division supplies the quadriceps

extensor and the hamstring muscles *. _

The obturator nerve (O.N) divides into inner and outer parts

between the inner and outer heads of the adductor brevis. ‘The

former emerges to the outer side of the superficial head of the

muscle and supplies the pectineus, gracilis, adductor longus, and

adductor brevis. ‘Che latter emerges to the inner side of the

superficial head and supplies the adductor magnus, adductor

brevis, and obturator externus. No branch accompanies the

popliteal artery.

The lateral cutaneous nerve (H.C.N) branches off from the

anterior crural nerve, and is distributed as in Man.

The great sciatic nerve (G.S.N) at first supplies the obturator

* This distribution to the hamstrings was not observed in another Chimpanzee.

412 DR. C. F. SONNTAG ON THE ANATOMY, .

internus, gemelli, quadratus femoris, biceps, and gluteus maxi-

mus. As it courses round the tuber ischii and down the thigh

it gives branches to the hamstrings. In the popliteal space

it divides into external and internal popliteal nerves. The

external popliteal nerve passes under the biceps and through the

extensor longus digitorum and supplies both. It is continued as

the anterior tibial nerve. The latter supplies the anterior

tibial muscles at the top, the ankle joint, the flexor brevis

digitorum, the tarso-metatarsal joints and the skin of the

adjacent sides of the hallux and index. It gives off the

musculo-cutaneous nerve which, however, only supplies the skin

of the adjacent sides of the index, medius, annularis, and

minimus. No nervus suralis exists. ‘The internal. popliteal

nerve becomes the posterior tibial nerve. This passes between

the heads of the gastrocnemius and supplies them. As it passes

down the leg it gives a branch to the upper part of the anterior

tibial muscles and branches to the posterior tibial muscles,

peronei, ankle joint, and flexor brevis digitorum. It divides

into three terminal branches. A muscular branch runs to the

abductor minimi digiti. The internal plantar nerve, or second

terminal branch supplies the abductor hallucis, lumbricales,

flexor brevis hallucis, adductor hallucis, joints of the foot, and

the skin of the inner four toes. The nerve to the last digit

communicates with the lateral plantar nerve. The lateral plantar

nerve, or third terminal branch divides into superficial and deep

parts. The former supplies the abductor and flexor and skin en

the outer side of the fifth toe. The latter supplies the adductor

hallucis, interossei, and tarso-metatarsal joints.

The superior gluteal nerve (S.G.N) emerges above the

pyriformis, and divides into two branches which follow those of

the artery. A special branch runs to the gluteus minimus. but

the scansorius is supplied by the sciatic nerve.

The «inferior gluteal nerve (1.G.N) accompanies. the corre-

sponding artery to the gluteus maximus.

The pudendal nerve (Pud.N), after emerging through the

sciatic notch, forms a prominent cord lying alongside the

pudendal vessels. It lies in the outer wall of the ischio-rectal

fossa, but no well-marked Alcock’s canal exists. It gives

several twigs to the rectum, external sphincter ani, levator ani,

sphincter vagine, and ischio-cavernosus. It also supplies the skin

of the perineal region. It differs from that in Man in that it

does not pierce any triangular ligament, and it has no branches

to the transverse perineal muscles, for the latter are absent. It

does not divide into two terminal branches of large size.

The small sciatic nerve (S.S.N) courses much as in Man.

I agree with Bolk (7) that the Chimpanzee, like the other

Anthropoids, differs from Man in the absence of a nervus

suralis.

The lumbar and sacral nerves receive grey rami communicantes

from the gangliated cords of the sympathetic nerves (text-fig. 46).

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. A13

The Sympathetic Nervous System (text-figs. 45 & 46).

The long, oval superior cervical ganglion (S.C.G) extends from

the level of the hard palate to the hyoid bone. It is connected

by communicating branches to the ninth (IX) and twelfth

cranial nerves, and to the ganglion nodosum (G.N) and its

superior laryngeal (b) branch. On the left side it sends no

twigs direct to the cervical plexus, but it is connected to the

first and second cervical nerves on the right (text-fig. 459A.) It

gives off pharyngeal nerves and the external carotid plexus, but

no cardiac nerve arises from it. The internal carotid branch

(I.C.N) breaks up into a plexus before it enters the skull.

The left sympathetic runs separate from the vagus and ends

in the middle cervical ganglion (M.C.G) whence the following

branches radiate :—(1) A stout cord which divides into branches

accompanying the thyroidea ima artery to the thyroid gland

(T.B.S), tracheal nerves and cardiac nerves (C.B.S) to the deep

part of the cardiac plexus and plexus round branches of the

aortic arch. (2) Nerves to the cardiac and aortic plexus (C.B.S)

(3) Continuation of the cord to the inferior cervical ganglion

(1.C.G). This also communicates with the vertebral plexus

(V.A.P), brachial plexus (¢.B.P), and cardiac plexus.

The right sympathetic fuses with the right vagus, but

separates from it lower down again, and a rich plexus of nerves

comes from it, both above and below, and accompanies the

common carotid artery to the plexus on the branches of the

aortic arch. The middle ganglion does not send off many

radiations as on the left side.

The inferior cervical ganglion (1.C.G) and first thoracic

ganglia are fused. It gives off rami communicantes to the

brachial plexus (c.B.P), a thick plexus which accompanies the

vertebral artery (V.A.P), a nerve to the cardiac plexus, and

the thoracic sympathetic cord (T.C.5).

The Thoracic Cords have fewer ganglia than the number of

intercostal nerves. The left one gives off the great splanchnic

nerve (G.S.N) at the level of the fifth and sixth thoracic nerves.

At the level of the diaphragm it divides into the small

splanchnic nerve (S.S.N) and abdominal sympathetic cord

(S.C). In addition to these it gives off rami communicantes to

the intercostal nerves and some of these are long. Aortic nerves

accompany the intercostal arteries to the plexus around the

aorta, and some of these reach the root of the lung, but were

very delicate at that region.

Abdominal Cords (text-fig. 46):—The left cord runs down and

passes under the left renal artery. It possesses four ganglia.

The first (G.1) lies at the level of the superior mesenteric artery.

The cord which emerges from it gives off nerves to the inferior

mesenteric plexus (I.M.P) and divides into two. The halves are

collected again into the second ganglion (G.2); this gives off

rami communicantes (R.C) to the first two lumbar nerves, a

414 DR. C. F. SONNTAG ON THE ANATOMY,

branch to the inferior mesenteric plexus, and the ovarian plexus

(O.P). The cord cornecting the second and third ganglia gives

twigs to the aortic plexus (A.P). The third ganglion (G.3)

lies at the beginning of the common iliac artery. It gives

off rami communicantes to the lower two lumbar nerves,

hypogastric nerves (H.N) and the external iliac nerves. The

fourth ganglion (G.4), situated within the pelvis, gives off

strong rami communicantes to the sacral nerves, and a nerve to

the hemorrhoidal plexus. The right cord has two abdominal

and one pelvic ganglia. ‘lhe first ganglion gives rami communi-

cantes to the lumbar nerves from its lateral aspect. From its

mesial aspect a stout cord comes and divides into an upper

branch to the left renal plexus (L.R.P) and a lower bunch of

three nerves to the inferior mesenteric plexus (I1.M.P). The

second ganglion is at the level of the common iliac artery.

Vagus and Sympathetic Plecuses.

A. Pharyngeal Plewus:—This is formed by branches of the

glossopharyngeal nerve and sympathetic, and the pharyngeal

branch of the vagus.

B. Cardiac Plexus (text-fig. 45 .B):—The cardiac plexus lies

chiefly between the aortic arch and heart anteriorly, and the

trachea posteriorly. It receives two cardiac branches of the

left vagus. One rises in the neck and divides into four branches

on the front of the aortic arch; two of the branches pass under

the arch to the deep part of the plexus, the third runs to the

surface of the arch, and the fourth supplies the pulmonary

artery. No sympathetic filaments run over the arch to the

superficial part of the plexus (S.C.P). The deep part of the

‘plexus (D.C.P) communicates with the superficial part and

receives :—(1) Many filaments from the left sympathetic, a

thoracic cardiac branch of the left vagus, the cervical cardiac

branch of the right vagus, three thoracic cardiac branches of the

right vagus and filaments from the right cervical sympathetic.

The plexus contains two clusters of ganglia, one behind the

beginning of the innominate artery, and the other between the

aortic arch and bifurcation of the trachea. The large vessels

arising from the arch haye associated plexuses or sympathetic

nerve cords.

C. Anterior pulmonary plexuses derived from the vagi. No

separate sympathetic filaments are seen.

D. Posterior pulmonary pleauses derived from the vagi and

upper thoracic ganglia of the sympathetic.

K. Solar Plexus (text-fig. 46):—The gangliated ring, which

acts as a centre, lies in front and at the sides of the cceliac axis.

It receives the greater part of the right vagus (R.V) and the

great splanchnic nerve of the left side (G.S.N). It gives off

a trunk at its lower end which runs into the gangliated cord of

the sympathetic of the left side (A.G.C). It gives off the cceliac

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. A15

plexus, which breaks up into gastric (G.P), splenic (S.P), and

hepatic (H.P) plexuses. It sends off the superior mesenteric

(S.M.P), left renal (L.R.P) and left suprarenal (L.S.R.P)

plexuses.

EF. Inferior Mesenteric Plewus (1.M.P):— A well-marked

ganglion (I.M.G) is present. It supplies the descending colon

and rectum and communicates with the left abdominal sympa-

thetic cord.

The right celiac ganglion (R.S.G) receives the great

splanchnic nerve. It is connected to the left ganglion by

several communicating nerves. Offshoots go into the coeliac,

right renal and superior mesenteric plexuses.

G. The right and left renal plewuses (L.R.P. and R.R.P) are

offshoots of the corresponding halves of the solar plexus. ‘They

receive splanchnic nerves and branches from the abdominal

syiapathetic cords.

H. The ovarian plexuses (O.P) are offshoots of the abdominal

sympathetic ganglia.

The Hye and its Appendages (text-fig. 27).

The skin over the supraorbital margin has a few long hairs

running in different directions, but there are no pronounced

eyebrows. The upper lid is longer than the lower, and has

longer cilia. The Meibomian glands form projections on the

back of the lid, but a strip of darkly-coloured conjunctiva prevents

them from forming ridges on the ocular surface of the lid. The

capacious lacus lachrymalis is lined by black conjunctiva. No

caruncula is present, but the plica semilunaris is well marked.

The bulbar conjunctiva is dark in colour, but only the marginal

part of the palpebral conjunctiva is pigmented. The lower

lachrymal papilla is larger than the upper one, and the internal

tarsal ligament is larger than the lateral tarsal raphe.

The lachrymal gland is small and flat, and consists of two

parts asin Man. The ducts open into the superior conjunctival

fornix. And the naso-lachrymal duct opens below the inferior

turbinate bone into the middle of the inferior nasal meatus

(Pl. II. fig. A). The gland is deeply embedded in thick fat.

The fascia is very strong, and is attached as in Man to the

tarsal ligaments.

Orbital Muscles:—The levator palpebre superioris arises as in

Man. But it has only two insertions—into the tarsus and

conjunctiva—instead of three. It is supplied by the third nerve

as in Man. The frontal nerve is far internal to it. The

superior oblique arises as in Man, and the trochlea is well

developed. Its long, fan-shaped tendon is inserted into the

eyeball distinctly to the outer side. It passes under the superior

rectus. ‘The fourth nerve supplies it by three branches. The

rectus swperior arises, and is inserted, as in Man. As _ its

insertion the ocular surface plays upon the anterior border of the

416 DR. C. F. SONNTAG ON THE ANATOMY,

superior oblique. It passes through an arch formed by the

capsule of Tenon. The rectus externus arises by two heads

and ig inserted asin Man. It is broad and moderately thick.

The third nerve crosses both heads instead of passing. between

them. The fourth nerve passes over both heads as in Man.

And the sixth nerve comes out between them before sinking

into their ocular surface. The naso-ciliary nerve also crosses

both heads. The rectus internus is broad and thick, and its

attachments are as in Man. Its nerve, from the superior

division of the oculomotor nerve supplies it by several twigs.

The inferior oblique arises by fleshy and tendinous fibres from

the floor of the orbit a quarter of an inch external to the naso-

lachrymal duct. It is not spread out as in Man, but remains as

a thin belly, which is inserted farther back into the sclera close

to the entrance of the optic nerve on the postero-lateral aspect

of the ball (text-fig. 27). The rectus inferior isas in Man. It

is, therefore evident that the recti are almost as in Man, but the

obliques and Jevator palpebre differ.

The nerves and vessels are described in other sections of this

paper.

The capsule of Tenon is very strong.

The ophthalmic veins are as in Man.

On pulling the eye forwards it was seen that the fascia lying

next to the eyeball was seen to be well developed, and almost

free from fat. The globe itself is relatively smaller than in

Man, but the ophthalmoscopic appearances are very similar in

both, as pointed out by Lindsay Johnstone (70).

Auditory Apparatus.

It is well-known that the auricle is less degenerate in the

Chimpanzee than in Man and the other Anthropoids. And

from the numerous accounts which have been published it

appears that the auricle is one of the most variable parts of the

external anatomy of the Chimpanzee. Its very complete form

in my specimen is shown in Plate I. fig. A. It has few hairs, and

Wallis (58) pointed out that it has this feature in all examples.

Darwin (16) noted that neither the Orang nor the Chimpanzee

move their auricles, and I was unable to detect any movements

on any occasion when I made observations in the Ape House in

the Gardens. In Plate IT. fig. B it is shown how the auricular

cartilage is very complete, and it has a wide, thin peripheral

rim. But the human auricular cartilage is a totally different

thing. I was unable to detect intrinsic muscles in the cartilage.

The tympanic membrane cannot be seen through the ordinary

aural specula, for it lies at the end of a long, bony external

auditory meatus.

The Eustachian tnbe has no well-marked torus round its

pharyngeal end, and I did not detect a salpingo-pharyngeus

muscle.

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. Al7

Doran (71) pointed out that the auditory ossicles, taken as a

whole, resemble those of Man more than do those of the Gorilla

and Orang. But in these Anthropoids the ossicles resemble

those of Man more than do those of the Chimpanzee in a few

points. In the Chimpanzee the malleus is more human than

those of the Gorilla and Orang. ‘In the shape of its head,

which projects markedly forwards, and in the nature of its

articular surface, of which the outer segment is much the widest,

it approaches 7’. gorilla more than Homo or Simia; but in the

neck and manubrium it is very human, the only difference being

that the latter, in this ape, is narrower at the base, and more

eurved than in Man, and its well-developed processus brevis is

directed upwards, and hardly outwards. In length the handle

does not exceed that of our species—another prominent distine-

tion from the other two apes. The body of the incus resembles

that in Homo: the processus brevis is more slender, and ends in

a sharp point, with no trace of any depression on it. The

processus longus is rather stouter and shorter than in Man; it

forms with the posterior crus a right angle. The stapes is

smaller than in Man. The crura are almost equally curved;

they are shorter and more slender than in Man, but wider apart

at their insertion. They are well grooved towards the aperture

which is wide. ‘The base resembles that of Homo, though less

distinctly reniform, and equally rounded off at both extremities.”

The Skin and Tegumentary Organs.

As the Chimpanzee uses the extensor surfaces of his fingers in

progression the skin has become modified. On the penultimate

phalanges it exhibits long, oval callosities; and it has papillary

ridges on its terminal ones. These ridges appear to increase

during the period of growth, and Kidd (56) after describing

their longitudinal direction states: “their long axes are at right

angles to the line of progression of the animal. There is no

correlation between the act of prehension and the direction of

the ridges, though it agrees closely with the general rule which

obtains in so many regions, that the ridges lie at right angles to

the line of incidence of the predominating pressure on the part.”

The mamme are two in number, and pectoral in position.

The umbilicus was very faint in this specimen.

The following account of the comparative histology of the

hairs of the Anthropoid Apes has been written by Mr. F. Martin

Duncan, F.R.M.S., F.Z.8.:—

The hair of the Chimpanzee is lank, coarse in texture, and

jet black in hue. Microscopically it presents certain interesting

features. The cuticular scales are well marked, narrow, and of

the imbricate-ovate type characteristic of the Anthropoid Apes,

“and in contour bear a closer resemblance to Gorilla than to

Simia. In the cortex, between the cuticular scales and the

medulla, the pigment granules are very numerous, opaque, and

418 DR. C. F. SONNTAG ON THE ANATOMY,

tend to coalesce in short, regular lines. The medulla is con-

tinuous, homogeneous, and densely pigmented. The hair shaft

is cylindrical. (PI. ITI. fig. A.)

A transverse section of the skin, passing across a hair-

follicle, shows the thick outer and inner root-sheath, with the

layers of Henle, and of Huxley, both well developed; while the

mass of elastic tissue closely surrounding the. hair-follicles

presents a very striking appearance (PI. aes figs. B and C).

PATHOLOGY.

Nothing is known of the diseases to which the Chimpanzee

is subject in its native surroundings, In captivity in Europe it

usually succumbs to diseases of the respiratory or digestive

organs. Some animals die from generalised tuberculosis, or from

osseous and arthritic changes after many years in confinement.

The following table, compiled from the death reports, shows the

causes of death and duration of life of animals which have been

in the Bacighy! s Gardens since 1882.

No. Date of Tage atihe Cause of Death.

1, 7. 5.1882 1 month. Ulcers of tongue. Viscera healthy.

2. 8. 6.1883 16 days. Pneumonia.

3. 29. 10. 1883 Se Typhoid fever *.

4, 22. 6.1884 1 month. Bronchopneumonia. Ascites.

5. 4.11. 1886 43 months. Acute bronchitis.

6. 20. 3.1889 11 days. Pneumonia.

Uo 1. 6.1889 1 year, 7 days. Bronchitis.

8. 15. 4.1891 |2 years, 4 months. | Pneumonia.

9. PRE fey dlsiMe 7 WO Pneumonia, peritonitis (TB.).

| 10 15. 11.1891 43 months. Pneumonia.

11 23. 9.1895 1 year, 5 months. | Hypertrophied liver. Ascites.

12 17.11. 1895 8 months. Bronchitis. Pneumonia.

13 12.12. 1896 5 months, 9 days. ie os

14 13.12.1896 | 1 year, 2 months. £p A

15 1. 6.1897 4 days. Not opened.

16 27. 9.1898 3) 45 Debility.

17 8. 10. 1898 1 year, 5 months. | Bronchopneumonia.

18 3. 12. 1899 114 months. Not examined.

19. 6. 12. 1899 7 months. 3 3

20. 24. 3.1900 | 4 years, 1 month. | Chronic pneumonia.

21. 16.10.1901 | 10 months, 1 week.| Pneumonia. Hepatic congestion.

22. 9. 1.1903 7 months. Prolapsus ani. Killed by order.

23. 4.12.1904 | 1 year, 10 months. | Bronchitis.

* Authority J. B. Sutton.

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 419

No. ern | tee Bes Cause of Death. |

24. 20. 3.1905 | 1 month. Generalised tuberculosis.

25. 16. 1.1906 3 months,3 days. | Colitis.

26. 24, 4.1907 | 4 years, 0 months. | Trauma.

oy 7. 9.1907 |2 ,, 5 4, | Ulcerative colitis.

28. 19.11.1907 | 3 years, 12 days. | Colitis. Enteritis.

29. 6. 7.1908 1 year, 2 months. | Bronchopneumonia.

30. 8. 7.1908 | 3 years, 2 months. 3 55

31. AY, BUSY) NR ee WE ce Fracture of skull.

32. cS ale, 5 AS SS al i eine ts oh Pneumonia.

33. | 1. 8.1919 | 10 ,, 10 ,, | Chronic arthritis. |

3d. 7.10. 1922 Pas id 3) es Pneumonia.

It has been shown by Metschnikoff, Roux, Neisser, and Lassar

that the Chimpanzee is more susceptible to the virus of syphilis

than any other Ape or Monkey. The primary lesions appear

in thirty days after inoculation; the secondary symptoms develop

after a further period of more than thirty days; but tertiary

signs have never been observed.

All experimental inoculations with the gonococcus have failed

to produce a result.

Keith has collected papers by Owen (75, 76), Schmidt (77),

Rollet (78), and Meyer (79) on the pathology of the Chimpanzee.

And the works of Ehrlich and Hata give accounts of the trans-

missibility of yaws to Apes; but the actual Apes employed have

not been mentioned.

I desire to express my thanks to Dr. Doreen Stranger, Dr.

J. H. James, Miss Kahan and Messrs. Aurounin, Henderson,

Meneces, and McCormick, students in the Anatomy Department

of University College, for their assistance in the dissection of

the animal described above.

CoMPARISONS witH Man.

The Chimpanzee resembles Man in a general way in form and

structure, but it differs from him in many respects. Some of the

differences are associated with habits and diet; others are

dependent on differences in the size and complexity of the brain ;

and others again are the outcome of different developmental

processes.

At a certain stage the fetuses of the Chimpanzee and Man

have several features in common, but the subsequent develop-

mental changes—both intrauterine and extrauterine—proceed in

different directions. In the Chimpanzee they are marked by a

progressive increase in certain parts, such as the hair and facial

skeleton. In Man, on the other hand, they are characterised

by suppression ; but the power to develop farther lies dormant.

A420 DR. C. F. SONNTAG ON THE ANATOMY,

The suppressive agents are the various ductless glands. When

they are diseased the suppressive power is removed, the latent

power reasserts itself, and Man assumes certain ape-like

characters. Man, in fact, retains more fetal characters than the

Chimpanzee. The most distinctive character of the human

foetus is the foot, for it has never been seen with the hallux

projecting from the postero-mesial aspect of the sole.

The Chimpanzee differs from the white races of Man in its

pigmented, hairy skin, its thick lips, and its overgrown facial

skeleton, which exhibits large supra-orbital crests, prominent

zygomata and malar bones, prognathism and large mandible.

But diseases of the ductless glands cause Man to assume one or

more of these charaeters, for they remove the suppressive

agencies. In Addison’s disease of the supra-renal capsules the

skin becomes pigmented; and in the various disorders of the

pituitary body, so beautifully monographed by Cushing (15), the

lips thicken, the skull exhibits large crests, zygomata and malar

bones, maxillary or mandibular prognathism occurs, and there is

a variable amount of hirsuties. The extremities also become

large and clumsy. Many of these conditious are present as the

normal characteristics in the lower races of Man; and one of the

most prominent features in the skull of Homo rhodesiensis is the

enormous development of the supra-orbital crests.

At a certain stage in development the foetuses of all Primates

have external genital folds. In the human fetus they continue

to develop and form the labia majora and mons veneris, and they

bury the labia minora and clitoris. In the lower Primates they

disappear and the clitoris is exposed on the surface. But the

Chimpanzee exhibits an intermediate condition. The mons

veneris is slight, and the labia majora are represented by two

slight elevations of the skin over thickenings of the subcutaneous

tissue (Pl. I. fig. C). The chief difference between the Chim-

panzee and Man is the absence of the hymen. In diseases of the

ductless glands the organs atrophy in Man.

The biochemical reactions of the blood show that Man is

related to the Chimpanzee and other Anthropoids, and it is

evident from the above that the actions of the ductless glands

have altered the appearances of these relatives in a pronounced

manner. Bolk (7) has shown that the suppressive action has not

only influenced the somatic features of Man, but it has retarded

his development and succeeding life phases. He believes that the

ancestor of Manchanged his diet from frugivorous to omnivorous,

and the change may have been the factor which evoked the

suppressive action of the endocrine organs.

The compressed head appears sunk between the shoulders, for

the neck is short. It is also more rigid than in Man. This

arrangement throws no obstacle in the way of the long arms, and

the shortness of the neck may be designed to give the powerful

levator anguli scapule and levator clavicule a very strong fixed

origin.

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 421

If some object is held above the animal’s head one can see that

there is a considerable upward movement of the eyeballs, but the

head does not move much. And the greater upward movement

of the eyes compared with that in Man is effected by a more

posterior attachment of the inferior oblique muscle.

The Chimpanzee uses its arms as hook-like suspenders, but the

diminutive thumb is of no great use for suspension. The new-

born child can, it is well known, support the weight of its body

for a half to two minutes in a similar manner. Its fingers

reflexly assume this position if one places his index finger in its

alm.

; Much has been written about the attitude of the Chimpanzee,

but the conclusions, in several instances, have been drawn from

the study of dead material, or from the observation of sluggish

animals moving clumsily across the floors of their cages. Those

who have observed Chimpanzees in their natural haunts testify

to their activity and agility ; and I have been fortunate in being

able to examine a male Uganda Chimpanzee, lately arrived at

the Gardens, which still exhibits much of its originai activity.

It runs about actively, using its arms and legs almost equally ; it

occasionally uses its foot as a spring-board; and it swings about

on the branches in its cage very actively.

Anatomical descriptions state that the Chimpanzee keeps its

knees semi-flexed and give that as one of the factors which

prevent the animal from assuming the erect attitude. And

Humphry (26) states that one cannot fully extend the knee

without doing violence to the muscles. If, however, the living

animal is examined a different state of affairs can be observed ;

but the observations must be long and frequent. I observed the

active animal mentioned above extending its joints fully,

both during active progression and while standing up and

holding on to the bars of its cage. Two young animals were then

examined during their active movements, and the same conditions

were observed. After studying the active range of movement I

examined the passive movements in two other young animals,

and I found that I could easily extend the knees; but the

curvature of the upper end of the tibia gave the leg an apparent

slight flexion even when the knee is lightly extended. It is,

therefore, evident, from the results obtained on these five living

animals that the knee-joint can be fully extended. The position

of semi-flexion is, however, more comfortable in the Chimpanzee,

as it is in Man, and an animal which becomes sluggish in

captivity will develop stiff joints, so full extension of the knee

will then become impossible, either actively or passively. And I

believe that some anatomical accounts have been based on the

examination of limbs so affected.

If the animal were deprived. of its arms it could not stand

upright like Man, but it can under momentum be erect for a

short period; I have observed the active animal mentioned above

Proc. Zoou. Soc,— 1923, No. XX VITI. 28

422 DR. C. F. SONN'LAG ON THE ANATOMY,

run for a few paces in the erect posture. ‘he maintenance of

the erect attitude in Man is effected by a very compiex and

beautifully adjusted nervous, muscular, and osseous mechanism ;

but many factors co-operate in the Chimpanzee to make it

quadrupedal when on the ground. Jn the first place, the centre

of gravity is high, for the greatest weight of the body is nearest

the arms. ‘he animal will naturally fall to the ground unless

it uses its arms as supports. In Man, on the contrary, the

centre of gravity is low down near the supporting legs. As

distension of the abdomen by food-and pregnancy throws the

line of gravity farther forwards in Man, the effect of similar

conditions on the Chimpanzee will make the arms all the more

necessary as supports. In the second place, the muscles of the

back are more rigid in the Chimpanzee, so they are not employed

as in Man for adjusting the Lalance to suit awkward positions.

Thirdly, the arrangement of the bones and joints of the pelvis

and lower limbs in the Chimpanzee is such that the lower limbs

cannot be converted into strong supporting pillars. Finally, the

muscles are not so subdivided as in Went so the movements are

more massive. ‘There is not the fine co-ordination of movements

which Man obtains through a highly organised brain, a delicate

and complex nervous mechanism, and a subdivided muscular

system, whose elements can group themselves to produce complex

actions.

The. Chimpanzee experiences joy and anger, and young

ones manifest jealousy if their companions are petted. It

expresses these emotions by grimaces instead of fine facial

expressions. The lips and cheeks exhibit gross movements, and

many teeth are exposed. ‘The reasons for this are the coarseness

of the platysma and its very intimate union with the labial

muscles; and the latter ave coarse, fused, and devoid of fine

subdivisions.

The muscles of mastication are built on the same plan as in

Man, but they are more powerful. And the prognathism makes

the levatores and tensores palati more horizontal than in Man.

The columns of the erector spine are coarser than in Man, and

pass farther up into the neck. And the shortness of the neck

almost obliterates the sub-occipital triangle.

The muscles attached to the shoulder girdle are so arranged

that the arm can be moved far backwards. In addition to the

usual elevators, which are more powerful than in Man, there is a

levator clavicule. The nerve supply to the rhomboideus, levator

seapule, and first part of the serratus magnus is very rich. The

Chimpanzee has also a dorso- epitrochlearis.

If several animals are examined it is seen how the pectoralis

minor writes its evolutionary history

There is considerable fusion between the muscular bellies of

the flexors and extensors of the wrist and fingers. The flexor

carpi ulnaris is more bulky than in Man, and it is inserted into a

PHYSIOLOGV, AND PATHOLOGY OF THE CHIMPANZEE. 423

very large pisiform bone. The trapezium, on the other hand,

is small, and there is no os centrale as in Simia. The minute

flexor longus pollicis is an offshoot of the tendon of the flexor

profundus to the index. The palmaris brevis is large, the

first and second lumbricales are connected by a muscular slip,

and there are six palmar interossei. Still those anatomical

differences are not sutlicient to show that the hand of the

Chimpanzee is not such a marvellous mechanism as that of

Man.

The muscles of the abdominal parietes are very strong, for

they act as flexors of the trunk, and they support the abdominal

viscera when the animal is walking. A pelvie floor is present,

but there is no true central point of the perineum, which plays

such an important part in supporting the uterus in woman.

The vulva and anus are behind a line connecting the anterior

extremities of the ischial tuberosities, but a corresponding line in

woman passes between the vulva and anus,

The quadratus lumborum is shorter than in Man, and is strongly

fused with the iliacus, and the latter is longer than in Man.

The glutei are less than those in Man, but the maximus has a

longer insertion. The glutei and other thigh muscles exhibit a

considerable degree of adhesion, and some of the thigh muscles

are inserted into the fascia over the muscles of the leg. The

scansorius is absent in Man. The adductors form a powerful

mass, aud they help to keep the inverted foot against a tree in

climbing; from the backward projection of the ischium the

adductor magnus is a powerful extensor of the leg in leaping.

In the muscles of the leg it is interesting to note the doubling

of the tibialis anticus and the absence of a tibial head of the

soleus.

Anatomical literature contains many speculations as to the

nature of the foot of the Chimpanzee. Cuvier, Blumenbach, and

Owen, and in later years Huxley (27), Humphry (26), and

Embleton (19) have proposed different views. <A survey of their

writings shows that it has been regarded as a foot, a hand, or a

compromise between them, that is a chiropodous structure. To

settle this question it is necessary to consider the extremities

from the anatomical and physiological points of view, and it is

necessary to examine them in several animals. Using the human

hand as a standard, it is, in the first place, necessary to see how

it differs from a fore-foot. Humphry (26) points out that the

elongation of the phalanges and the shortness and opposability

of the pollex are the characters which transform a fore-foot into

a hand. In the case of the terminal part of the lower extremity

many of the myological and osseous features are characteristic,

not of a hand or a foot, but of a hind limb. So we must look to

the characters of the digits and hallux to determine the nature

of the part in the Chimpanzee. Comparing the measurements in

28*

424 DR. C. F. SONNTAG ON THE ANATOMY,

the animal whose anatomy is described in this paper with those

in Man the following results are obtained :—

Chimpanzee. Man.

End of heel to tip of middle digit... 7-4 ins. 8-4 ins

Hallux and its metatarsal bone ...... 40 ,, ATOM.

Hallux without metatarsal bone ... 2°2 , DO hos

Length of metatarsals 2-5 ............ 20, PAS bans

Length of phalanges :—

Secan@ydwott este... reo ZONE iL Siers

LOUD EO gs eh tg Dena CALs SARL DrSt he

Fomnbl hie i Be en AAD at

fifth BSS SY Ue 203) Ne

Total length of second digit ......... AD ,, Aes.

It is thus seen that in the Chimpanzee the phalanges of digits

9-4 are more than a third of the length of the foot, but the fifth

digit is less than a third; but in Man they are less than a

quarter of the length of the foot. The hallux im the Chimpanzee

is shorter than the other digits, but it is slightly longer than

them in Man. In the Chimpanzee the relative lengths of the

digits are as in a hand, and the digits act towards or from a line

passing through the middle digit; in a foot the basal line passes

through the second digit. All these data show that the pelvic

extremity in the Chimpanzee is terminated by a hand, And if

the grasping action of the human hand be taken as a pattern it is

even more effective than the hand which terminates the pectoral

extremity. The hind hand is not employed as an exploring organ,

and its grasping action is chiefly an aid to progression, so

Humphry (26) thinks that the term “‘ chiropod ” best describes it.

And he makes the following important statement :—‘‘ Whichever

term is used it must not be forgotten that the configuration on

which it is based is not peculiar to the monkeys, but is common

to them with some other tree-roving animals, such as Iguanas

and Opossums. In human beings, who are born without arms,

the foot can be educated to take the place of the missing bands.

And medical literature contains records of such persons who

could grasp objects and carry out complex movements, such as

painting, with their hallux and foot digits. These cases have

been adduced by some writers to prove that the leg of the ape

is also terminated by a foot. They do not really disprove the

views stated above for:—1. The proportions of the hallux and

digits to each other and to the whole foot are normally foot-like ;

2. The middle line, or basal line, runs through the second digit ;

3. The hallux can be abducted, but it is not really opposable;

4. The prime work is to act in conjunction with the other

characters of the leg in forming a strong supporting basis for

the body.

The joints between the occipital bone, atlas, and axis differ

from those in Man. The inferior crus of the cruciate ligament

is absent, and there are additional strengthening bands. No

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE. 425

ligamentum nuche is present. Owing to the greater forward

projection of the odontoid process, the check and middle odontoid

ligaments are more horizontal than in Man. ‘The articular

processes cause more locking of the atlas and axis. The head has

a much more limited movement, and this is accompanied by a

greater upward mobility of the eyeballs. The carpal ligaments

are more complex than in Man. And the construction of the

joints of the pelvis and leg are such that the anima] cannot stand

stiffly in a fully erect attitude without holding on to branches

above it.

The mouth differs from that in Man in the greater number of

palatal rugs and in the characters of the tongue, although

Wood-Jones states that the tongue is like that in Man. The

pharyngeal musculature is similar in both, but the Chimpanzee

has the larger sinus of Morgagni. Man has only two muscular

coats in his esophagus, but the Chimpanzee has an additional

inner longitudinal coat in its upper part. Man has three

muscular layers in his stomach, but the Chimpanzee has only

two; and the inner coat is composed of oblique or circular fibres

in different parts. As regards the intestinal tract the most

striking differences are found in the rectum The Chimpanzee

has five enormous valvule conniventes running completely round

it, and the columns of Morgagni are very prominent longitudinal

folds running down to the lower end of the anal canal. When

the anus is dilated these columns are seen at once. No valves of

Bull connect the columns asin Man. The anus is prominent as

the gluteal regions are small, whereas the reverse is the case in

Man.

The amount of lymphoid tissue in the intestinal mucosa is less

in the Chimpanzee than in Man.

The liver and pancreas are practically the same in both.

The heart is small in the Chimpanzee, but its structure is as in

Man. ‘The branches of the aortic arch take one of two forms,

and one of these varieties isas in Man. The subclavian arteries

give off well-marked spinal branches, and the branches of the

axillary artery come off as in the Cercopithecide. No arterial

anastomoses exist round the scapula or elbow joint as they are in

Man. The vessels in the forearm form long parallel branches,

much as in the Lorises, and there are three palmar arterial

arches, whereas Man has only two. The branches of the

abdominal aorta are fewer than in Man; and in my specimen

the femoral artery gave off branches which combine those of the

external iliac and femoral arteries in Man. The femoral artery

gives off a saphenous artery to the foot, and the anterior tibial

artery courses differently from that in Man.

In the venous system the chief differences from the condition

in Man are the freer intercommunications between the parts of

the portal circulation, the absence of a saphenous opening, the

ending of the cephalic vein in the lower end of the brachial veins.

and the presence of a large brachial vein instead of two vene

comites of the brachial artery.

426 DR. C. F. SONNTAG ON THE ANATOMY,

The lymphatic system is characterised by a doubling of the

thoracic duct; and the groups of glands are much fewer than in

Man.

- The ductless glands generally resemble those in Man, But

the thyroid is a long, narrow U-shaped organ instead of being

composed of an isthmus and two lateral lobes. The vascular

supply is interesting in my specimen, but I am unwilling to

regard it as the normal condition till other material becomes

available for examination.

The lungs are divided into the same number of lobes as in

Man, but they ave subdivided differently. The trachea and

bronchi are as in Man, but the larynx differs in some points of

structure, and in the possession of a large air-sac. Vrolik has

shown that they are largest in the aged, and it is possible that

this fact is correlated with weaker muscles at that period of

life,

The external generative organs have already been alluded to.

The internal organs are built on the same plan; but the round

ligaments are relatively much thicker than in the human

condition. They may play a more important part in fixing the

uterus than they do in woman. ‘The vestibule is very small, and

the meatus urinarius opens within the vagina.

It is sometimes stated that Man and Ateles are the only

Primates which possess more than one renal papilla. But that is

not the case, for I have seen kidneys of Chimpanzees with three

to six papille. The left renal artery is peculiar in my specimen

for it anastomoses with lumbar arteries, whereas the renal

arteries are end-arteries in Man.

In its origin from the femoral artery, and its course up

through the femoral sheath the obturator artery im my specimen

courses as in one form of abnormality in Man.

Man differs from the Chimpanzee in being bimanous and

bipedal, and in the possession of those higher mental powers

which we designate by the name of the Soul.

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PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE.

Fig.

EXPLANATION OF THE PLATES.

Prats I.

. The external ear.

. The auricular cartilage.

. The external generative organs. Letters in text.

Prate II.

. The turbinate region. Letters im text.

. The antrum of Highmore. Description in text.

Prate III.

Fig. A. The hair shaft.

Figs. B & C. Sections through the skin and the root of a hair.

PZ. S. 1923, RING, Pl. 1.

4 John Bale.Sons & Danielsson, Lt?

ELEPHANT-SEALS OF KERGUELEN LAND.

ere

Sie recs

P. Z.S. 1923, RING, Pl. II.

fe. John Bale.Sons & Danielsson, Lt?

ELEPHANT-SEALS OF KERGUELEN LAND.

ELEPHAN?-SEALS OF KERGUELEN LAND. 431

23. The Hlephant-Seals of Kerguelen Land.

By T. P.. A. Rine *.

[Received November 21, 1922: Read March 6, 1923. ]

(Plates I. & 11.7)

The hunting of the Elephant-Seals of the southern hemisphere,

on a scale which threatens these interesting animals with

extinction, began at the South Georgia group of islands in the

South Atlantic, which are under the jurisdiction of Great

Britain, and form part of the Falkland Islands dependencies.

A factory for whaling was established in the year 1905-6, by a

Norwegian, Captain C. A. Larsen, of Antarctic exploration fame.

This enterprise proved so successful that other establishments for

utilizing the larger cetacea weresoon erected. Asa ‘“ side-issue,”

the capture of the Elephant-Seals was also included, and in con-

sequence their numbers must have been so reduced that, unless

the British Government introduce legislation forbidding the

capture of these animals for, say, 10 years, they will become

extinct. Such legislation ought also to embrace the protection

of the Fur-Seals.

At the Crozet Islands, situated in the South Indian Ocean,

and which are under the jurisdiction of France, a Norwegian

steam sealing-factory made a great haul of Hlephant-Seals in

1907, and three years later a French floating-factory cleared the

beaches of the remnant left by the Norwegian vessel. In the

year 1908 a Norwegian company established a whaling factory at

Kerguelen Land in the South Indian Ocean. This also is a

possession of France, and in the course of four years (1909-1913)

Klephant-Seals were killed in such great numbers that, unless

the Government of France takes steps for protecting them from

destruction by enforcing strict protection, say for. 10 years, their

extermination is practically certain. The Hlephant-Seals and

Fur-Seals of Marion, Prince Edward, and the Crozet Islands

should be similarly protected.

The transport vessel of the Kerguelen Whaling Company,

the 8.8. ‘Jeanne d’Arec,’ arrived at the Island with all the

paraphernalia for establishing a whaling factory, on the 29th of

October, 1908, coming to an anchor in Gazelle Basin, a fine

harbour situated centrally on Kerguelen Land. During the

fortnight following, the vessel visited a great many bays looking

for a suitable site for putting up the factory, which finally was

established to the west of Long Island in Royal Sound, thus

named by Captain James Cook in 1776. On every sandy beach

* Communicated by Sir S. F. Harmer, K.B.E., F.R.S., F.Z.S.

+ For explanation of the Plates, see p. 443.

432, MR. T. P. A. RING ON THE

of any extent a huge bull Elephant-Seal, the beachmaster of the

old-time sealers, was found to be in possession of a herd com-

prised of females and youngsters—dams and pups—whilst on

smaller beaches, and lolling about on the grass-covered plains

above the beaches, were flocks of male Elephant-Seals of various

ages and dimensions, termed “‘ pods” by the sealers.

The dams were all brown in colour, measuring from seven to

about eleven feet in length, and rather emaciated after suckling

their pups, who were in splendid condition, almost bursting with

fat, and covered with a pretty silver-grey coat of short hairs,

The beachmasters were most imposing looking, compared with

their offspring, among which the young bulls measured as much

as seven feet in length, whilst the females were no more than five

feet long. The guardians, or “sultans,” of the harems were, in

most cases, dark brown, like the dams; though some were dark

grey in colour. On some beaches a solitary, dun-coloured bull

was seen, who, like the beachmasters, was scarred from many a

fight; and among the “pods” of variously sized grey bulls,

fighting was the order of the day, combats being waged at all

hours if the sun was not out, and for the mere pleasure of

battling ; the hoarse bellowings of struggling bulls being audible

at night as well.

In the course of a week several anchorages on Kerguelen Land

were visited by the 8.8.‘ Jeanne d’Are,’ and about the 6th of

November the beachmasters were observed to be very much on

the alert to prevent the dams from abandoning the pups: placing

themselves between their harems and the water. Whenever a

dam tried to pass, the bull simply crushed her down, and he

would even administer a bite; but whilst he was thus engaged

another dam would watch her chance, and slip past him into the

water. With angry bellowings the beachmaster would try to

intercept the runaway ; but no sooner did the dam kept down by

the bull feel the pressure diminish, than she, in her turn, slipped

away, and there the baffled Sultan stood, bellowing forth his rage

at the escape of his odalisques.

The beachmasters were found to measure from about 18 to 21

feet in length: and had always a well-developed proboscis, which

is an enlargement of the snout more nearly resembling that of

the tapir than the trunk of an elephant; whence, however, is

derived the name “ Sea-Hlephant.” As long asa bull is undis-

turbed this “‘ proboscis ” hangs flabby and limp to one side of the

jaw ; but when excited, or roused to anger, it becomes rapidly

inflated with air and enormously enlarged, the process of inflation

being accompanied by short snorts. The bulls, when fighting

each other, are very careful to prevent the trunk from being

seized, contracting and raising them, as far as possible, out of

harm’s way. The dams are entirely devoid of this enlargement

of the nose, which is a sign of sexual maturity in the males, and

is most pronounced during the rutting period; while it is per-

ceptible in the pups when only seven to eight weeks old.

ELEPHANT-SEALS OF KERGUELEN LAND. 433

By the 10th of November only a few cows were left, and it

had been noticed that each had but one pup to suckle. In several

harems there were a few without youngsters; and these were

assumed by us to have been unfecundated. As late as the 21st

of the month a dam was seen to have come up to the beach to

suckle her pups, and as the youngsters were increasing in fatness

for some time after the disappearance of the mother seals to

recoup their strength after the exhausting strain of suckling, the

dams had evidently returned, now and then, to the beaches

before finally abandoning the pups. After Christmas the cows

began to reappear, all being in fine condition after their voyage

of recuperation, some arriving even earlier, and they now went

far up on land to loll about among the bulls changing their coats,

and growing slim again, until the month of April when the eveat

exodus of the Elephant- “Seals took place.

In the month of November the “rene was fairly fine; and

the pups which, on the departure, or rather the disappearance of

the dams, had withdrawn from the beaches and collected on the

green slopes above, were seen by the end of the month to

approach the water again. The beachmasters had remained to

guard their youngsters, and were very prompt to come to their

assistance when teased by us. Indeed those bulls, which had

been harassed and troubled by the men, had at last become quite

furious, so that the sight of a man was suflicient to drive them

into a towering rage.

Whilst lying on the slopes, the pups were seen to charge each

other in clumsy hops, and express their feelings in a kind of

hooting when catching each other up. The male pups also tried

to fight one another ; but on raising their fat, round bodies they

generally tumbled over; and when teased the nose would be

puffed up twice its original size. The quivering of the muscles

of the face gave the otherwise docile physiognomy of these

youngsters quite a ferocious aspect.

They had decreased considerably in fatness, though grown in

length when mustering on the beaches about the 3rd_ of

December: and by the 10th the beachmasters, who were very

much reduced in bulk, and who oceasionally would disappear for

a couple of days, doubtless to seek some food, took their departure

with their offspring. In the month of February the pups

reappeared, the males having grown to a length of about nine

feet, whilst the females nerdy measured more than seven feet in

length.

The « pods” of bulls or bachelor seals, which on our arrival

were seen on the beaches and the flats above, disappeared at

various times to feed themselves up, and on their return, after

about two months’ absence, they resorted far up on land, to spend

the remainder of the summer basking in the sunshine on the

slopes and plains above the beaches, shedding their hair and

growing thin again; many were seen in the miry troughs or oval

hollows made by generations after generations of seals, and others

434 MR. I. P. A. RING ON THE

in bog-holes, chiefly above Cape Digby on the lowland, where the

animals were wallowing about like so many crocodiles. Fighting

would be the usual pastime in rainy weather; but with the sun

out, they were chiefly engaged in throwing up mud and sand on

to their backs with a backward sweep of the fore-flippers, so

that the body was entirely covered. It being too hot to fight,

they would also chase each other about and play at pairing.

Instead of going up the slopes some seals seemed to prefer lying

among the boulders forming the talus, at certain parts of the

coast-line, and the grey, clumsy shapes of the animals were diffi-

cult to distinguish from the boulders.

By the end of March, and in the beginning of April 1909, the

Elephant-Seals were coming down to the beaches again: travel-

ling leisurely, and having bouts of contests as shown by their

tracks and the downtrodden mud.

The gathering of the clans took place on the beaches, the seals

departing in companies on their four to five months’ tour of

migration, but unfortunately it was impossible to ascertain if the

males and females travelled together or separately. By the Ist

of June the beaches had become deserted; but some seals, males

and females, were seen to remain behind and winter over, the

animais probably suffering from some disease or other. Snow

did not affect them much and they were lying on the beaches as

well as in the shallows, covered with snow, and looking like so

many logs of timber. When asleep in the water the heads of

the seals were sometimes visible, and at other times submerged,

and for hours they would remain asleep without stirring. When

a snap of frost set in the beaches were deserted, the temperature

of the water being more congenial than that of the air.

It has been mentioned that after their tour of recuperation

the seals were in splendid condition: and that they travel far

has been proved ; the late Captain Robert Falcon Scott having

observed Elephant-Seals at South Victoria Land in latitude

77° 50' South, during the Antarctic summer. The animals fed

themselves well when travelling, as shown by their sleekness

when hauling on land again ; and when returning after their five

months’ voyage of migration, the seals were in even a better

condition. Their route is unknown, as no congregations of seals

have ever been met with by vessels “‘ running the easting down ”

in the latitudes of the roaring forties, and the probability is that

these animals resort to the regions of the pack-ice during the

winter, unless they perform a round voyage, like the hump-back

whales, or follow the route of migrating fishes: They may, of

course, visit unknown feeding-banks; but the pack-ice seems a

more likely region where an abundance of food may be found,

and the seals are safer from their chief enemy, the prowling

Orea gladiator. Several of these ferocious “ Dolphins” were

seen cruising in the fjords of Kerguelen Land, and many a seal

had deep cuts which only could have been inflicted by a powerful

beast like the ‘‘ Killer- Whale.”

ELEPHANT-SEALS OF KERGUELEN LAND. 435

Several Elephant-Seals were seen during the winter in the

various anchorages resorted to by the whalers in bad weather, a

few Crab-Haters and one Ross-Seal, as well as a small number

of Sea-Leopards, also having been observed, and specimens

secured for Museum purposes.

On the 28th of August 1909 three large, grey bulls hauled up

at Swain’s Haulover, a neck of land separating Swain’s Bay from

the fjord to the west of Long Island, in Royal Sound, where the

whaling factory had been established, and about 24 miles distant

from the Jatter. The weather had been, and still was, abomin-

able, and the seals disappeared again. Tracks in the snow

showed that they, or some others, had been upa day or two

earlier, but none were seen again until the 3rd of September,

when several “hauled up” at Swain’s Haulover, as well as in

Royal Sound, and at Greenland Harbour Haulover, which

separates this harbour from an arm of Royal Sound. The new

arrivals drove the bulls that had wintered over, and which were

very thin, compared with the others, away to the corners of the

beaches, or compelled them to seek another beach, and they then

commenced to battle between each other for the supremacy,

which meant the possession of the beach. The weather was most

uncongenial however, the thermometer only registering 304° F.,

though the sea-water was much warmer. On this account the

fights were rather half-hearted, and the water was preferred.

In the course of a week an increasing number of large bulls

hauled up: and on the 11th of September a few dams reappeared.

The battles between the rival males now began in real earnest,

the arrival of the females adding a zest to what previously

seemed a mere pastime. Thecombats now assumed a sanguinary

and decisive character. It was always bull to bull, and whenever

a large bull without provocation attacked a weaker one, others

would come to the assistance of the latter, and hurl themselves

at the bully. The duels were fought in the shallow water until

the bulls were, so to speak, sorted out in pods of equal strength

and dimensions, and at last only one or two remained, the others

taking themselves off, and away from the victorious bull, who

took up his position among the dams. At Swain’s Haulover two

fine bulls lingered in the shallow water without ascending the

beach. Both had come out as victors over their rivals, and both

had had several trials of strength. ;

What may. be called real water-pantomimes had been witnessed

by us: the pelting of thesundry fighting bulls with stones having

nonplussed the animals to such an extent that, when a stone hit

one, the beast would thrust his head into the water after the

stone, and on finding what had given him the stinging blow, he

looked round in a dazed way and next furiously went for the

nearest of his compeers.

It was rather puzzling to see that neither of the two bulls

took possession of the beach, although an increasing number

436 MR. T. P. A. RING ON THE

of dams were hauling up, but on the 14th of September the

riddle was solved—our friend of last year, the Sultan, made

his appearance.

This special bull, and also one in Seelhorst Harbour, near

Gazelle Basin, on the east coast, had been the objects of much

harassing by our people, both being fine brown specimens about

20 feet long, and both possessing a mutilated proboscis, which

distinguished them from their confréres. No sooner did the

Sultan of Swain’s Haulover, as he had been named, perceive his

tormentors of the previous season than he made for us in eager

strides, and a scene followed similar to others that had taken

place, and probably remembered by the bull, as no provocation

whatsoever had been offered by us, the remembrance of last year’s

harryings rankling in his memory no doubt.

Hurling his ponderous body along, the dark brown, eager bull,

without uttering a single snort, evidently had the intention to

run the object of his attack over and crush it, and great was

therefore his surprise when it disappeared, the man in front

having quickly slipped aside. The discomfiture of the beast was

apparent as it came to a sudden stop, the inflated proboscis

dropping, and a loud snort being emitted. Raising himself on

his fore-flippers the bull lifted his head with the snout pointed sky-

wards, and, looking over his back and turning his head slowly from

one side to the other, a searching glance was directed backwards

and sideways. The astonishment of the bull was great when he

saw his enemy standing behind him, so to speak, and on getting

a smart blow from a stick on his hind-flippers his surprise and

disappointment gave place to a most sudden outburst of fury.

With bulging, blood-red eyes, and inflated proboscis trembling,

a succession of stentorian snorts and gurgling roars came from

the vibrating snout and open mouth of the bull; and on getting

a second stinging cut he flopped down and curved his back, whilst

at the same time the hind-flippers were thrown up, opened out,

and swung to one side, imparting amomentum to the body which,

oivoting on the fore-flippers, was turned with a grand sweep in

nlmost a semicircle until he was again facing his adversary. He

was undecided what to do it seemed; but a blow on the head

roused him to action, and with a magnificent motion he rose from

the ground, towering up to a height of 10 feet, and looking most

imposing as he thundered forth his fury, which was genuine

enough, and with which his whole body was shaking.

It isin this magnificent posture that the battling bulls pose,

for some seconds, when challenging each other to combat; and

they next hurl themselves at one another with allthe momentum

they can possibly impart, clashing together with a curiously soft

thud, and with such force that one expects the oil to squirt out

of the pores of the skin. Seizing hold of each other with their

fore-flippers, the antagonists sway about, attempting to gouge

out an eye, or seize the proboscis, both being careful to contract

this appendage, and turn it upwards out of reach, When trying

ELEPHANT-SEALS OF KERGUELEN LAND, 437

to bite each other on the sides of the neck, the animals incline

their heads towards the threatened side, and the foldings pro-

duced by this movement in the skin make a good bite difficult.

Nevertheless, deep bleeding gashes are inflicted, which, however,

heal quickly, and rarely fester. The fight ends, as a rule, in

mutual exhaustion : the bodies of the combatants finally cannoning

off each other, and dropping to the ground,

At times the combat would be renewed until the weaker one

gave up the fight and took himself off, gashed and bleeding, but

mostly the bulls were seen to separate, unless fighting for the

supremacy of the beach.

The temper of the Elephant-Seals is normally docile; and by

slowly approaching a sleeping animal, and waking it up without

any undue noise, one could come quite close up to a seal, and

even sit on it, and scratch it with one’s cane; the soft eyes, after

an almost friendly glance, closing again. If frightened the pale

green pupil of the eye, set in a brown iris, would become blood-

red however, a mist of tiny drops of blood gradually filling the

pupil, a hue which was also assumed when a seal was angry.

Tears came into the eyes when, on waking an animal suddenly,

the abrupt opening of the lids exposed the eyes to the light ; but

when harassed a seal would also shed tears, and the pupil then

changed from pale green into blood-red,

By the 20th of September the dams hauled up in force, some

arriving singly, but generally they came in companies, and on

the 24th the first birth took place. The weather was cold and

snowy however, and only a few dams were delivered during the

two following days. On the 27th and 28th the wind went

northerly, with the temperature of the air rising from 33° to

44° and 52°F., and the greater part of the pups were then pro-

duced, but as late as October 7th what appeared to be new-born

pups were seen in a bay on the South Coast of Kerguelen Land,

though only a few, the arrival of the dams being belated,

The delivery of the pups took place close to the sea, and the

travail of the dams did not last longer than about five minutes, the

heads of the youngsters appearing first, and each dam only pro-

ducing one pup. ‘The loss of blood was slight, but the delivery

was attended with pain, and cries of distress were uttered at

certain moments, the dams facilitating the event by movements

of the body. With a sweep of the hind flippers the umbilical

cord, which remained attached to the youngster, was severed.

It gradually dried up, and had a length of from two to three feet.

The female pups were, at birth, about three feet long, whilst

the males measured as much as five feet in length; all being

covered with jet-black, curly hair, and presenting more the

appearance of a chameleon than that of a seal.

The patresfamilias, as well as the other bulls, were quite dis-

interested in what was taking place, although they must have

been cognizant of it, as their moustaches and nostrils were

moving.

Proc, Zoon, Soc.—1923, No, X XIX, 29

438 . oo MER. T, B.A. RING ON Dre

The growth of the pups was astonishing: day by day one

seemed to notice the progress, and in about 10 days they had

more than doubled their bulk and weight. No attempt was made

to seize a youngster and weigh it, but on comparing the size of

some of them with that of a man, the estimate was arrived at

that the pups in question weighed 180 to 200 Ibs.; and at birth

from 65 to 80 Ibs.; the largest male pups 85 lbs. There was a

preponderance of females in nearly all harems, but no certain

ratio could be determined. When suckling the youngster, the

mother-seal reclined to one side, offering the nipples protruding

through two holes—-one on each side of and equidistant from

the navel, as well as in line with it—to the pup, who, after

having sucked his fill, painfully dragged himself, with the beavy

head bobbing up and down, into the water; the dam following

up, and placing herself alongside of the youngster, who rested

with his fore-flippers on the shelving bottom, and both animals

facing landwards. When, as often happened, the slight. rough of

the sea drew the pup out, and he lost the support of his flippers,

the head became submerged. Immediately the dam swept the

youngster up, so that he again recovered his footing. If

prevented from getting into the water the pups vomited the milk.

According to the American authority, Mr. N. W. Elliott, in

regard to the Pribyloff-Islands seals, they become land-sick ;

digestion, which must take phe while the pup is in the water,

having been prevented.

In the third week after their birth the jet-black pelts of the

pups changed into brown, the black curly hairs dropping out, and

brown, straight hairs taking their place, the coats being thick

and rough. Gradually these were changed into the pretty silver-

grey coats, which the pups had when leaving the beaches on

about the 10th of December, but which, on their return in

February, had assumed a darker hue.

On the 17th of October the pairing of the Elephant-Seals was

observed, this event taking place when the first cireumnavigation

of Kerguelen Land was made by us. In Thunder Harbour, near

Sadole Island, on the West Coast, where a party of men had been

landed to capture the Fur-Seal,—which was believed to visit this

part of our domain, but which proved not to be the case, only a

couple of these seals being seen, and a small one captured—a

curious commotion was noticed on the 16th of October among

the hundreds of Hlephant-Seals that were lying on the plain

above the beach. Large bulls were seen to descend, or rather to

tumble down, the steeply shelving beach from the plain, gashed

and bleeding, to dive into the sea and disappear; whilst other

powerful beasts were coming out of the water to ascend the

beach and making their way up to the plain, whence a con-

tinuous roaring of bulls was hear d, and the sharp agonized yelps

of pups and dams. From an island, and at a height overlooking

the harbour, the plain was dimly visible through the mist; and

the huge shapes of careering bulls were discernible, the animals

ELEPHANT-SEALS OF KERGUELEN LAND. 439

coming’ to a sudden stop amidst the harems. Fierce fights

between-the bulls lying outside:the herd were taking place, ‘and

occasionally an immense bull was seen making his way through

the seals. Waning daylight, and the necessity for finishing a

rough survey of Thunder Harbour, and taking soundings, pre-

vented a visit to the plains, but next day, in Christmas Harbour,

the riddle was solved.

The 8.8. ‘Jeanne d’Are’ had called here on the 24th of

December, 1908, in order to takea couple of Sea-Hlephants to the

Durban Zoological Gardens, and two pups were most unexpectedly

secured, a smail bull being too tough a customer to tackle. As

the migration of beachmasters, with their pups, had been noticed

to happen on the 10th of December, it became evident that the

season's youngsters did not abandon the beaches simultaneously

with their sires, but that the latter left for the most part when

it suited them, to a certain extent, together with their pups.

In Captain Cook’s Christmas Harbour, two fine bulls with

their harems occupied the beach, separated by a space of about

500 feet. The same commotion, but ona much smaller scale than

what dimly was observed in Thunder Harbour, and which up to

the day before had never been seen anywhere since the arrival

of the 8.8. ‘Jeanne d’Arc’ at Kerguelen Land, was also raging

among the seals lying on the beach in this anchorage, and, on

getting ashore, the reason was discovered—the cows were “in

heat.” Fierce, sanguinary battles were fought between the

bachelor bulls lying in the shallows, the water shining crimson-

red in some places, while rolling bellowings mixing with the

savage hootings of the obsessed ‘beasts resounded between the

hillsides. On the beach, the two Sultans were standing alert and

threatening, ready to repulse an aggressor; both mutually

concurring not to poach on each other’s preserves, and watching

the approach of the boats with evident concern. Suddenly one

of them was seen to face about, pivoting his body in a magnificent

sweep on his fore-flippers, and gallop madly into his harem, where

he disappeared from sight, and on gaining the beach and coming

through the crowd of pups and dams we found the bull serving

one of the latter.

The commotion among the “water-bulls” became pande-

monium. The frenzied creatures went blindly for each other,

digging their sharp incisor teeth into each other, and careering

about; the sound of their roars and bellowings filling the whole

harbour. One, more venturesome than the others, ascended the

beach and made for the nearest cow, which he tried to ravish ;

the unwilling harem beauty, with Tend cries of distress, making

her lord aware of the situation, and appealing to qa for

assistance. Having satisfied the call of the other dam, the harem

master came bounding along, running the offender ‘down, and

erushing him with his weight so that the onder lying bull

trdcehod with terror or agony, whilst making frantic eftorts to

440 . MR, T. P. A, RING ON THE

wriggle away, in which he finally succeeded, and wobbled into

the water again, so much wiser, if not better, after his futile

raid,

The Sultan of the other seraglio had in the meantime driven

off a slightly smaller bull after a fairly severe trial of strength,

and was found lying on the ground, recovering his breath,

when he spied the two-legged intruders coming along. Raising

himself on his fore-flippers the bull lifted his head, and with

the snout pointing to the zenith, turned his head slowly from

side to side, taking a scowling look at us, and after what must

have been a satisfactory scrutiny, lying down again.

On walking among the inmates of the harem, a black dead pup

was found and later two more, which the prancing bulls must

have crushed in their mad careering through their harems. The

Sultan, probably being aware of our heavy footfall on the hard

sand, would occasionally rise to have a look round, and suddenly

he became alert, the proboscis, which had been hanging limp and

flabby, being inflated to a size which exceeded anything previously

seen. Up went the head and with the moustache bristling, and

eyes turning blood-red, the bull slowly moved his head about, the

proboscis vibrating, and saliva running from the open, pink,

cavern-like mouth. The posture of the seal was one of tense

concentration, and all at once he threw up his hind-flippers, as

usual with the animals when making a grand swerve, and

bounded off through his harem, the dams and pups scuttling out

of the way of the prancing bull with terrified yelpings. Coming

to a stop before a dam whose youngster had scrambled well out

of the way, and who seemed to be waiting for her master, and

inviting him by twisting her body about, the bull with a loud

snort seized the dam with a fore-flipper and served her.

The bulls in the water, who fought fiercely with each other for

the privilege of being onlookers, as the men said, must also have

received the silent call, as pandemonium again reigned among

them, until the bull had satisfied the demand of nature. Within

about 45 minutes the two beachmasters served each three dams,

and every time the call came from different cows. Had all six

made their demand on the services of the bulls at the same time,

those in the water, which were equally possessed by the pairing

instinct, would have ascended the beach and satisfied the calls of

the dams.

The following day the pairing of the seals in Christmas Harbour

continued, and in all the bays which were visited on our return

to the whaling station—when large harems were found,—the

bulls were engaged in satisfying the calls of their dams, of which

none were seen to be served twice. On beaches where only a few

dams formed the harem no pairing was observed, this act, no

doubt, having taken place before our arrival.

In Royal Sound a beachmaster had been vanquished by a bull,

which had successfully contested his supremacy. The fight had

HLEPHANTESEALS OF KERGUELEN LAND. 44]

been savage, bout after bout having been fought, according to

the testimony of the men from the whaling station, who had been

deputed to make tours in one of the motor boats of the Kerguelen

Whaling Company, to observe the seals. The two rivals had

kept up the contest for nearly 40 minutes, and the beachmaster

was at last vanquished. He was almost unable to move, and one

of the incisor teeth had gone, the jaws of the combatants having

become interlocked, and only released by the snapping of the

incisor of the beachmaster. Gashed and streaming with blood,

the defeated Sultan then dragged himself painfully to the beach,

and dived into the water, the wake of the animal as he swam

away being crimson-red. The victorious bull, who looked only a

degree less frightful than his late opponent, had taken possession

of the harem, that is, he simply dragged himself in among the

dams, being almost spent, and went to sleep at once. Some

hours later, the motor boat having in the meantime visited other

harems, the bull was observed to have recovered his mettle, and

was on the gui vive when the boat appeared, though no pairing

took place as the men passed the beach on returning to the

whaling factory.

The Sultan of Swain’s Haulover had also had his hands full in

coping with the exigencies of the situation; but had come out

victorious in all contests, although one was almost a draw. Had

he been obliged to wage a similar combat soon after, the men

opined, he would hardly have come out of the battle with the

palm of victory.

What may be termed the rutting period of the Elephant-Seals

began on the 16th of October, when the inexplicable commotion

among the seals in Thunder Harbour was noticed, though it

may have commenced two to three days earlier, and it lasted

until all the dams in each harem had been satisfied. The mature

bachelor bulls, compelled by the pairing instinct, came to contest

the supremacy of the beachmasters, who by right of superior

strength and size, and by right of conquest of course, had

occupied the beaches and were harem-masters. The dams in each

harem requested the service of their masters at different times,

and not simultaneously, every dam being served only once.

They evidently roused the pairing instinct of their respective

masters by emitting a distinctive odour, imperceptible to the

human sense of smell, and a call, which also reached the water-

bulls, equally possessed with the pairing instinct, and which

drove them to frenzy.

During the rutting period it was seen that the beachmasters

received the call not only from mature cows, with or without a

pup, but also from what appeared to us to be immature females.

These females were of an almost uniform size, not more than

eight feet in length, of a slender shape, and born during the last

-week of September of the previous year. These female Elephant-

Seals when slightly more than a year old are then in heat and

A49, MR. T. P. A. RING ON THE

paired. The bulls of the samé age were naturally the smallest in

the herds, but had outstripped the females in growth by as much

as three feet in length. Like their older confréres they were

excited by the pairing instinct, fighting furiously with one

another when the call of the dams was sensed by them. When

only five months old these young males had been seen playing at

pairing, on their return to the beaches after their first voyage of

recuperation under the guardianship of their sires. Their seniors,

however, also indulged in this habit, but seldom troubled a

female seal, attempted overtures being promptly and indignantly

repulsed.

The dams had produced their pups about the 27th of September;

their fecundation took place about the 16th of October, and they

would again give birth to their youngsters about thé 27th of

September the following year. The period of gestation is thus

about 49 to 50 weeks.

On the 2nd of Nevember the slaughter of the seals was ordered

to begin, those at Swain’s Haulover and in Seelhorst Harbour to

be spared for further observation purposes, and the men were

also ordered to spare all dams. The pups were almost bursting

with fat, and most of them had changed into silver-grey coats ;

whilst the mother-seals. were emaciated from suckling their

youngsters ; but the bulls were in first-class condition. By the

8th of the month the dams began to abandon their pups, having

produced them about the 27th of September. Thus the period

of lactation lasts about six weeks.

In the season of 1909-10 the Elephant-Seals were observed to

leave the beaches, and return about the same dates as during the

season of 1908-9; and be it noted, as in the two following whaling

seasons, only 82 Cetaceans were captured in 1910, and about

the same number in 1911-12, the Kerguelen Whaling Co. would

have been obliged to go into liquidation but for the “successful ”

sealing of those seasons. But as a consequence, in 1913, there

would only be a small remnant of Elephant-Seals left to repro-

duce themselves. How far this remnant has succeeded in

recovering—if it has been able to recover—during the decennium

which has passed since the slaughter of the animals ceased, is

difficult to conjecture. It is by no means improbable that the

existence of these seals has been jeopardised through the strain

which the stock has suffered during three seasons’ intense hunting,

as all the large and virile bulls would have been killed on account

of the greater yield of blubber from such individuals. Con-

sequently only young bulls would be left, and the offspring of

such animals, deficient in virility, would be of an inferior stamina

compared with the old, extinct stock. It was seen by us that as

much as 35 per cent. of the pups in a harem succumbed to the

dangers of the first voyage of migration, and the percentage will

be much greater with seals of inferior stamina and vitality.

Although the capture of the animals would cease automatically

BLEPHANT-SEALS OF KERGUELEN LAND. 448

with their scarcity which rendered any venture on a large scale

unprofitable, even after a close period of ten years, it is doubtful

whether the Elephant-Seals of Kerguelen Land would be pre-

served from extinction, sickness and the dangers of migration

being important factors threatening the lives of the young seals.

EXPLANATION OF THE PLATES.

Prats I.

. Young Bulls fighting.

. A Bull with inflated proboscis.

. The Challenger.

. Bull with deflated proboscis.

wm Ww be

Pruate II.

Fig. 1. Elephant-Seal calling.

2. Young Elephant-Seals resting on the slopes above the beaches.

ON THE SKELETON OF LEPIDOSTEUS:; 445

24, The Skeleton of Lepidosteus, with remarks on the origin

- and evolution of the lower Neopterygian Fishes. By

C. Tate Ruean, M.A., F.R.S., F.Z.8., Keeper of

Zoology in the British Museum (Natural History).

[Received April 24, 1923: Read May 29, 1923.]

(Text-figures 1-8.)

Although a detailed and well-illustrated account of the skeleton

of Lepidosteus was given by Agassiz in 1833 (Poissons Fossiles,

vol. i1.), the nomenclature makes his description very difficult for

a present-day student to follow, and it has been ignored by most

modern authors. In his memoir on the development of the skull

of Lepidosteus osseus (Phil. Trans. 1882), Parker has given a

description of the skull of the adult fish which is in some respects

less complete and less accurate than that of Agassiz. More

recently Collinge (Proc. Birmingham Phil. Soc. viii. 1893) and

Allis (Internat. Monatsb. Anat. Physiol, xxi. 1905) have studied

Lepidosteus especially in relation to the sensory canal system.

Amia is much better known than Lepidosteus, but a proper

comparison of these two genera has never been made.

I propose, therefore, to describe the skeleton of Lepidosteus, to

compare it with that of Amia, to discuss the systematic position

of these genera, and to put forward my views as to the origin and

evolution of the group to which they belong.

I have examined skeletons of Lepidosteus platystomus, L. osseus,

and ZL. tristoechus; all three species are essentially similar in

structure, differing from each other mainly in the shape of the

bones and the number of segments of the maxillary, due to

the fact that in Z. osseus the snout is longer and narrower and

in L. tristoechus shorter and broader than in ZL. platystomus.

The nature of these differences will be seen on comparing the

accompanying figures (text-fig. 1).

SKELETON OF LEPIDOSTEUS PLATYSTOMUS.

Cranium (text-fig. 2).—The skull is elongate, narrowed forward,

depressed, with the upper surface flattish anteriorly and convex

posteriorly. It is well ossified, but a considerable part of the otic

region remains cartilaginous, and there is a large median ethmoidal

cartilage running the whole length of the rostrum.

On the upper surface a transverse series of 6 dermo-occipital

plates is rigidly united to the parietals, pterotics, and epiotics, and

the small, simple post-temporal is very firmly attached to the

dermo-occipitals, epiotic, and pterotic. Parietals, pterotics, and

446 _ fir. c. TATE REGAN oN WITH

frontals are normally developed. Anteriorly there is a small

dermal ethmoid, and paired nasals and adnasals cover the

olfactory sacs, which lie in concavities of the premaxillaries,

above the dentigerous margin of the latter. The premaxillaries

run backwards to below the frontals, expanding on the upper

surface of the snout to form a pair of rugose plates (ethmo-nasals

Text-figure 1.

A. B.

Upper surface of skull of A, Lepidosteus tristoechus; and B, L. osseus.

Lettering as in text-fig. 2.

of Parker); below this they form a pair of thick-walled tubes

that carry the olfactory nerves; they embrace the slender

anterior portion of the rostral cartilage and are attached below

to the vomers and parasphenoid (text-fig. 3). fey

The vomers are laminar bones attached to the lower surface

of the parasphenoid and premaxillaries ; each sends Lack a long

SKELETON OF LEPIDOSTEUS. 447

process attached to the parasphenoid. In front of the: orbital

region the parasphenoid is VW-shaped in transverse section, the

ascending lamine overlapping the premaxillaries in front and

descending laminze of the frontals behind, with the latter

Text-figure 2.

se SSS

BNO ISS

ER Se et

eosS

ix»

Lepidosteus platystomus. Skull from above and from ‘below; the pterygo-quadrate,

suspensorium, opercles, etc., of one side removed.

eth. mesethmoid ; ». nasal; 2.’ adnasal; pmax. premaxillary ; ma, maxil-

lary; f. frontal; p. parietal; pto. pterotic; spo. sphenotic ; epo. epiotic ;

pro. pro-otic; doc. dermo-occipital; eoc. exoccipital; boc. basioccipital ;

ptte. post-temporal ; psp. parasphenoid ; v. vomer; pal. palatine; ect. ecto-

pterygoid; ent. entopterygoid; m¢. metapterygoid; g. quadrate; corb. cir-

cumorbital; op. operculum ; sop. suboperculum; iop. interoperculum ; pop.

praoperculum.

enclosing a chamber that contains the posterior part of the

rostral cartilage. This does not completely fill the chamber,

but. on each side there is a channel for the passage of the

olfactory nerves (text-fig. 3).

448 MER. C. TATE REGAN ON THE

Behind the orbital region the parasphenoid expands, the

‘anterior edge of each wing forming a transverse condyle for

articulation with the metapterygoid. Parkew’s researches show

that these condyles originate as separate ossifications in cartilage,

and he terms them alisphenoids, but in the adult fish they are

completely ankylosed with the parasphenoid.

The true alisphenoids were not seen by Parker ; they are well

developed and separate ; they meet the pro-otics behind and are

in contact with the frontals above; anteriorly they join the

orbitosphenoids, which converge and ankylose anteriorly to form

a short vertical plate that rests on the parasphenoid ; just behind

this anterior plate the orbitosphenoid is pierced on each side for

the passage of the olfactory nerves (text-fig. 4).

The parasphenoid condyles are continued on to the pro-otics,

which intervene between the parasphenoid and the sphenotics ;

inner lamine of the pro-otics meet in the middle line and form a

roof for the myodome; the pterotics ossify downwards a short

Text-figure 3.

Lepidosteus platystomus. Diagrammatic transverse sections of the snout : A, at

the level of the posterior end of the maxillary; B, at the middle of the length

of the premaxillaries. ~

c. rostral cartilage; f frontal; x. olfactory nerve; py. parasphenoid;

gm. premaxillary ; v. vomer. i

distance in the cartilaginous wall of the otic region, and the

epiotics are well developed ; there are no opisthotics. The ex-

occipitals meet, and behind them. upper wings of the basioccipital

‘almost meet in the middle line, roofing the foramen magnum.

Circumorbitals, etc. (text-fig. 5,A)—The orbit is surrounded

-by a complete series of 12 circumorbital bones; these appear

externally as flat plates, but internally they are strengthened by

‘a stout circular ridge. There are 3 preorbitals, the anterior

overlapping the end of the maxillary. Behind the cireumorbitals

the cheek is covered with a number of irregularly arranged

plates.

Jaws, Suspensorium, etc. (text-fig. 5).—The preemaxillaries have

already been described. The maxillary is segmented into 8,

the anterior piece firmly fastened to the side of the premaxillary,

the others attached to the palatine and ectopterygoid.

In the lower jaw the concave surface for the articulation of the

convex condyle of the quadrate is formed by the posterior face of

SKELETON OF LEPIDOSTEUS. 449

an upper and the superior face of a lower ossification in Meckel’s

cartilage. There are well-developed angular and “ supra-angular”

bones, ‘the latter curving in above the coronoid elevation, On

the inner side there is ‘posteriorly a large laminar prearticular,

which ossifies through the coronoid cartilage below the anterior

part of the supra- ‘angular. Tn front of this a ridge of the dentary,

parallel to its margin, is covered by two elongate dentigerous

lamine, the shorter anterior one entering the symphysis, the

posterior ascending the coronoid elevation for a short distance

external to the przarticular ;-on one side a small bone is inter-

calated at the junction of these two bones, which may be termed

intra-dentaries. The prearticular and intra-dentaries are often

termed collectively ‘‘ splenials,” but it is very improbable that any

of them is the homologue of the true splenial.

The pterygo-quadrate is completely separated from the hyoid

arch, and supported by it only through the articulation of the

quadrate with the preoperculum, The ectopterygoid is a very

long and stout bone; anteriorly it articulates with the pre-

maxillary above the expanded head of the vomer; the posterior

Text-figure 4.

pes

Lepidosteus platystomus. Orbitosphenoid (os.), alisphenoid (as.), and pro-otic

(pre.) bones; pas. parasphenoid, Lateral view.

seven segments of the maxillary are firmly united to its outer

face, which has a groove for their attachment; the long laminar

palatine is attached to its lower surface. The entopterygoid is

small, the metapterygoid bears a process for articulation with

the parasphenoid and pro-ctic, and the quadrate has an anterior

condyle for articulation with the lower jaw and a posterior one

for articulation with the anterior end of the prazoperculum,

The large interoperculum is rigidly attached to the hyo-

mandibular, symplectic, and preoperculum; the last-named is

reduced to a small bone that lies above the anterior part of the

interoperculum and ends in a concave facet for articulation with

the quadrate. . Collinge and Allis transpose the names of the

przoperculum and interoperculum, on the ground that the latter

is attached to the hyomandibular and transmits a branch of the

sensory canal system. It may be pointed out that the. inter-

operculum is quile normal in its relation to the suboperculum

behind and the lower jaw in front, and that where it is in

450 MR. C. TATE REGAN-ON THE

contact with the preoperculum the latter lies above it. Also

the analogous case of Phractolemus may be quoted; in that

genus the very large interoperculum transmits the sensory canal,

but it is the horizontal limb of the preoperculum that is lacking,

whereas in Lepidosteus the vertical limb has gone.

The operculum articulates with a knob on the hyomandibular,

the suboperculum is well developed, and there are 3 branchio-

stegal rays. . oy es

Text-figure 5.

Lepidosteus platystomus.

A. Premaxillary, maxillary, cireumorbital, and cheek bones.

B. Palato-quadrate (outer view).

pal. palatine; ect. ectopterygsid (showing groove for attachment of:

maxillary) ; ent. entopterygoid; m¢. metapterygoid; g. quadrate.

C. Opercular bones, ete.

hm. hyomandibular; sy. symplectic; op. operculum; sop. suboperculum;

iop. interoperculum ; pop. przeoperculum.

DP, E. Lower jaw, outer and inner aspects.

den. dentary; an. angular; san. supra-angular; ar. articulars ; par. pre-

articular; id. intra-dentaries.

The hyoid arch comprises the hyomandibular, which articulates

with the sphenotic and pterotic, the symplectic, which is firmly

attached below to the preoperculum but is remote from the

quadrate, the interhyal, the ceratohyal, the hypohyal, and a

median basihyal. The ceratohyal has two ossifications, the

hypohyal only one. The upper ossification in the ceratohyal is

generally termed “epihyal;” this is certainly wrong, and I

SKELETON OF LEPIDOSTEUS, 451

think that there can be little doubt that the hyomandibular

is the true epihyal, 7. e. the epibranchial of the hyoid arch.

Vertebral Column.—The vertebra are solid and opisthoceelous,

with the neural arches and parapophyses ankylosed to the centra.

The vertebra number66; from the fifty-third the vertebral column

curves upwards and the vertebre decrease in size. The neura-

pophyses of the first two vertebra are short and stout; those

of the succeeding vertebre are prolonged into slender spines; the

spines. of each pair are in contact: distally, but remain separate ;

the last six vertebre have no neural spines and the last three

no distinct neurapophyses. The series of supraneurals extends

backwards from the first vertebra to the dorsal fin, and reappears

above the end of the vertebral column, where 4 bones (epaxial

supports of the caudal fin) are probably to be interpreted as

belonging to it. Hach supraneural is more or less expanded or

even bifid proximally; the first two are short and stout, and

articulate with the neurapophyses of the first two vertebre ; the

rest are slender, attached to the neural spines. The ribs are

expanded proximally for articulation with the distal ends of the

strong transverse pavapophyses. From the forty-first vertebra

backwards the parapophyses become shorter and are directed

downwards ; their ribs also are downwardly directed, and behind

the anal fin unite to form hemal spines; the hemal spines of

the upturned vertebre are expanded as hypurals to support the

caudal fin; the hypurals are only one less in number than

the upturned-vertebra, but in the specimen examined they are

displaced, so that the sixtieth vertebra bears two and some of

the last six vertebre do not bear any.

Fin Skeleton.—The dorsal and anal fins are supported by a

series of pterygiophores, each of which is divided into a long

proximal (basal) and a short distal (radial) segment ; each fin-ray

is articulated to its radial. The pelvic bones are rather long and

flat; they converge and slightly overlap anteriorly; only the

innermost radial is well developed, the fin-rays articulating

directly with the pelvic bones. The pectoral arch includes a

small simple post-temporal, which is firmly united to the skull, a

supra-cleithrum, and a cleithrum. The coracoid cartilage has

only one ossification, the hypercoracoid (or “scapula”); there

are 4 radials, 3 of which articulate with the hypercoracoid and

one with. the metapterygium.

CoMPARISON OF LEPIDOSTEUS WITH AMIA,

- Cranium.—In Amia the orbito-rostral part of the skull is

much shorter and broader than in Lepidostews, and the cavum

cranii extends forward between the orbits to the ethmoid region.

On the upper surface a single pair of loose dermo-occipital plates

represent the three pairs or more of Lepidosteus; the other bones,

parietals, ptevotics, frontals, preemaxillaries, nasals, adnasals, and

dermal ethmoid, have’ precisely the same relationship to each

452, MR. C. TATE REGAN ON THE

other as in Lepidosieus, differing only in form and proportions ;

owing to the shortness of the snout the premaxillaries are

scarcely exposed on the upper surface behind the nasals, and

when the nasals and adnasals are removed it is seen that the long

tubes that carry the olfactory nerves in Lepidosteus are top

sented by mere foramina.

The rostral cartilage contains paired septo-maxillary and eel

ethmoid ossifications ; the vomers and parasphenoid are broad

and flat, and the latter is unconnected with the frontals; the

Text-figure 6.

Amia calva., Skull from above. For comparison with text-fig. 2.

Lettering as before.

wings of the parasphenoid do not bear condyles and do not unite

with the pro-otics, but extend upwards to unite by suture with

the sphenotics and internally with the alisphenoids ; the orbito-

sphenoids are paired and separate. Small paired basisphenoids

are present, narrowly separated and projecting upwards in front

of the pro-otics, which roof the myodome just as in Lepidosteus ;

sphenotics and epiotics are much as in Zepidosteus, but in

addition there are well-developed opisthotics, and above them on

SKELETON OF LEPIDOSTBHUS. 453

each side a posterior temporal fossa roofed by the epiotic and

pterotic. Exoccipitals are as in Lepidosteus, but the upper wings

of the basioccipital in that genus are represented in Amia by two

or three pairs of bones, generally interpreted as the neural arches

of centra incorporated in the basioccipital.

Circumorbiials—In Amia the circumorbital series is incom-

plete above, and the bones are reduced in number; there is a

single preorbital, two large postorbitals cover the cheek, and

above them a smaller one appears at the side of the frontal as a

bone of the cranial roof.

Jaws, Suspensoriwm, etc.— As already noted, the premaxillaries

of Amia have the same relation to the dermal ethmoid, nasals,

adnagals and frontals, and to the olfactory sacs and nerves as in

Lepidosteus, but the maxillaries are quite unlike those of that

genus, for they are unsegmented and free ; each has an inwardly

Text-figure 7.

Amia calva. Pterygo-quadrate, suspensorial, and opercular bones,

Lettering as in text-fig. 5.

directed articular process at the anterior end, and each carries a

supplemental bone (or supramaxillary). The lower jaw of Amia

has two endosteal bones for articulation with the quadrate as in

Lepidosteus, but there is an additional small bone in front of the

upper of these, and above them is another additional bone with

a convex posterior surface for articulation with the concave

anterior end of the symplectic. The dentary, angular, and supra-

angular correspond to those bones in Lepidosteus, and, as in that

genus, there is a large laminar prearticular ; in front of this

Allis shows two intra-dentaries, but in the skeleton I have

examined, the posterior of these is represented on one side by four

bones and on the other by six; also in my example the endosteal

mento-Meckelian figured by Allis appears to be fused with the

anterior intra-dentary.

Proc. Zoou. Soc.—i923, No. XXX. 30

454 MR. C. TATE REGAN ON THE

Except that the palatine is separable into an endosteal

ossification and. two dentigerous lamin, the pterygo-quadrate,

hyoid, and opercular bones are the same as those of Lepidosteus ;

but they are very differently arranged, for the metapterygoid 1s

firmly attached to the hyomandibular and the quadrate to the

symplectic, the well-developed crescentie preeoperculum is united

to the hyomandibular and symplectic, and the interoperculum is

normally developed and movable. Scme authors have described

a process of the metapterygoid as articulating with the pro-

otic, but actually there is no such articulation, and the process,

Text-fieure 8.

Amia calva. Wower jaw, inner and outer aspects.

Lettering as in text-fig. 5.

which is an intermuscular lamina directed upwards, forwards,

and outwards, does not go anywhere near the otic region of

the skull.

Amia has a good series of branchiostegal rays and a median

gular plate. The ceratohyal has two ossifications and the hypo-

hyal one as in ZLepidosteus, and the branchial skeleton shows

no important differences from that genus.

Vertebre: and fins.—The vertebral column of Amia is well

known; the most important differences from Lepidosteus are

that the vertebre are amphicelous, and that in most of the

SKELETON OF LEPIDOSTEUS. A455

caudal vertebrz each centrum is divided into a precentrum

(without arches) and a ‘postcentrum. ‘he terminal vertebra,

hypurals, ete., are arranged much as in Lepidostews. The

skeleton of the fins is as in Lepidosteus, except that the pectoral

radials are more numerous and more than one articulates

with the metapterygium; the pectoral arch differs from that of

Lepidosteus in that the eoracoid cartilage is unossified and the

post-temporal is large and loosely attached, with a lower fork

running to the opisthotie.

Systematic Position of Lepidosteus and Amia.

It is probable that most of the characters in which Lepidosteus

and Amia agree are common to all the Holostean Ganoids (Semio-

notide, Pycnodontide *, Macrosemiide, EKugnathide, Amiide,

Pachycormide 7, Aspidorhynchide). Further, it is evident that

the essential features which distinguish Lepidosteus from Amia,

t.ée. the characters of the jaws, suspensorium, and opercles,

distinguish it also from all the others. Lepidotus is extremely

like Amita in its head skeleton, except that the skull is more

compressed, with the epiotics meeting above the exoccipitals and

the orbitosphenoids united (cf. Smith Woodward, P. Z. 8. 1893,

and Mon. Palwontogr. Soe. 1916-1919). I am fairly certain

that I can make out in this genus the symplectic articulation

with the lower jaw.

The Semionotide, to which family Lepidotus belongs, first

made their appearance in the Upper Permian, and are very

distinct in structure from the Paleoniscide, the only fishes

‘known which can be regarded as their ancestors. The typical

Palzoniscids appear to have been swift-swimming predacious

fishes, with a large mouth and sharp teeth. If we regard them

as giving rise to the Semionotide, probably slow-swimming

bottom-feeding fishes, with a small mouth and styliform or

tritoral teeth, we can interpret many of the differences as related

to a change of habits.

The change trom dorsal and anal fins with numerous rays

forming a close-set series to fins with the rays relatively few and

spaced, each ray with its own pterygiophore, and the reduction

of the muscular lobe at the base of each fin and of the radial

segments of the pterygiophores, are readily understandable if

the use of these fins changed from cleaving the water and with-

standing strains to performing the delicate movements of a fish at

rest or swimming slowly. Correlated with this modification of the

dorsal and anal fins is that of the caudal, the upturned end of the

* These are aberrant in the reduction of the opercular apparatus and the arrange-

ment of the bones of the cranial roof.

+ The preemaxillaries are said to be loose and separated by an ethmoidal rostrum,

but I am not satisfied that this interpretation is correct. I think that the so-called

premaxillaries may be the fractured anterior ends of the maxillaries.

30*

456 MR, C. TATE REGAN ON THE

tail being shortened so that the fin became terminal rather than

ventral. The use of the jaws for crushing shell-fish instead of

seizing fishes may account for the firm union of the premaxillaries

with the frontals. The lower jaw of the Paleoniscids appears

to have consisted of the same four bones as that of Polypterus,

namely dentary, angular, articular, and prearticular; that of

the Semionotids is shorter, deeper, and stronger, and the

additional articular ossifications (as seen in Amia) may simply be

due to the increased size and complexity of its articulation, whilst

the intra-dentaries may be regarded merely as plates developed to

support teeth. The ‘supra-angular” may be an anterior bone of

the outer circumorbital series which has adhered to the coronoid

process, in the same way that a bone of the temporal series has

become attached to the hyomandibular in Polypterus; and I believe

that the supplemental maxillary had a similar origin. The small

size of the mouth involved a change in shape and direction of the

preoperculum and hyomandibular ; the preoperculum, which in

the Palzoniscide appears to have been simply a plate covering

the cheek behind the circumorbitals, now became used to

strengthen the suspensorium, a necessity on account of the

forward position of the articulation of the lower jaw; the lower

end of the elongated hyomandibular ossified as the symplectic,

which may have been originally developed either, as I am inclined

to think, for the articulation of the lower jaw, or perhaps only

to support the quadrate and connect it with the preoperculum ;

the original hyomandibular ossification developed a knob for

_articulation with the operculum; the upper ossification in the

cerato-hyal appears to have been developed for the attachment

of the interhyal, which seems to be an ossified ligament. It is

obvious that the character of the respiration would be greatly

changed, the movements being smaller, and I think that a better

vegulation of these would result from the definite articulation of

the operculum with the hyomandibular ; moreover, the hyomandi-

bular being now but little movable, movements of the operculum

became of more importance. The inter operculum is a new bone

of some interest; it would almost seem as if the suboperculum,

retaining its attachment to the posterior end of the lower jaw,

had peen elongated and then fractured, the posterior part thus

retaining its freedom of movement, whilst the anterior—the

interoperculum—was overlapped by the horizontal limb of the

preoperculum, Finally, one may suspect that the disappearance

of the clavicles is in some way connected with the other read-

justments in this region.

The Semionotide appear to have developed in various direc-

tions, giving rise for example to the remarkable Pycnodontidea,

whieh ‘show! many resemblances to our modern Plectognaths; and

through the Kugnathide to the Amiidz and to the Pachycor mide,

which) parallel “the Scombroids. The development of a large-

mouthed piscivorous type appears to have taken place in two

SKELETON OF LEPIDOSTEUS. 457

ways. In the Eugnathide the cleft of the mouth extended back-

wards and the suspensorium became vertical or directed back-

wards; in these strong-swimming fishes the dorsal and anal rays

have increased in number and become concentrated, and the

caudal has become forked. On the other hand, in the Lepido-

steidee the snout and lower jaw have grown forward, the quadrate

retaining its anterior position, and we may suppose that as the

lower jaw became longer and more powerful the strain on the

suspensorium was relieved by the acquirement of the articulation .

of the metapterygoid with the parasphenoid. Other modifiea-

tions, such as the great development of the interoperculum, seem

to be related to the elongation of the orbital region, which gives

an increased surface for attachment of the strong mandibular

muscles. In these sluggish, lurking fishes the fins remain essen-

tially as in the Semionotide.

The Aspidorhynchide, which have sometimes been associated

with the Lepidosteide, have none of the peculiarities of that

family. An analysis of their characters shows a close agreement

with the Kugnathide, from which they differ in the prolonga-

tion of the premaxillaries to form a beak and the development

of a presymphysial bone in the lower jaw. Especially important

is the resemblance of the symplectic to that of the Hugnathide

and Ama (Smith Woodward, Monogr. Paleont. Soc. 1916-1919,

p- 98, pl. xx. fig. 2).

It is well known that the Teleosteans agree with the Holostean

Ganoids in the structure of the dorsal and anal fins, the absence of

clavicles (infra-clavicles), and the structure of the suspensorium

and opereles. But the most primitive Teleosteans (e. g. the

Elopide) differ from the Holostei in several important respects,

as follows :—The caudal fin is homocercal*. There are three

coracoid bones (hyper-, hypo-, and meso-coracoid), whereas in

Lepidosteus and Amia there is not more than one (hypercoracoid

or “scapula ”) and the mesocoracoid bridge is cartilaginous. The

pectoral radials are all inserted on the hypercoracoid and hypo-

coracoid, the lowest radial probably representing the metaptery-

gium. An endochondral supraoccipital bone is present. The

vomer is unpaired. The lower jaw has no “supra-angular” and

no intra-dentaries, and the prearticular is reduced. The pre-

maxillaries are loosely attached, remote from the frontals, and in

front of the olfactory sacs.

* Regan, Ann. Mag. Nat. Hist. (8) v. 1910, p. 353.

+ The current nomenclature of the Teleostean lower jaw is faulty. ‘The bone

- usually called angular is the homologue of the lower articular of Amia and the single

articular of Polypterus. Ridewood’s “ endosteal articular ” is the homologue of the

upper articular that articulates with the quadrate in Lepidosteus and Amia. His

“ectosteal articular” is the angular, and his “sesamoid articular” in all pro-

bability the przearticular.

According to Ridewood (P.Z.S. 1904) the bone last named serves for the attach-

ment of the tendon of part of the levator muscle, but it may well have persisted

rather than have been developed for this purpose. In the Cyprinodonts the opisthotic

is reduced to a nodule to which the lower fork of the post-temporal is attached.

458 MR. CG, TATE REGAN ON THE

If only the living forms were considered, it might be held that

the Chondrostei, Holostei, and Teleostei were groups of equivalent

rank. But a study of the fossils shows that this conclusion is

wrong. The mesozoic Pholidophoride and Oligopleuridz resemble

the Holostei in the structure of the caudal fin (and in the

Pholidophoride of their scales), but in other characters, such as

the small loose premaxillaries, the maxillary with convex oral

edge and bearing two supramaxillaries*, and the absence of a

“‘supra-angular,” they show clear evidence of their relationship

to the Hlopide.

The Pholidophoride had minute teeth, and were prebably

plankton-feeders like the Herrings, which they resemble in form

and in the shape of the fins. They are evidently another off-

shoot of the Semionotidee, from which they differ especially in the

structure of the mouth, and they lead to the Leptolepide and

Elopide. These are stronger swimmers, with a deeply-forked

caudal fin, in which the upper hypurals are for the first time

supported by uroneurals, paired bones that replace the upturned

end of the vertebral column.

The Holostei and Teleostei, therefore, are one group, for

which it seems better to use the name Neopterygii, rather than

to use Holostei or Teleostei in a new and extended sense;

whilst the name Paleopterygii may be used to designate the

group including the Palzoniscoids, Chondrostei, and ._Belono-

rhynchii.

I have long considered that the Selachians constitute a distinct

class (cf. P. ZS. 1906, p. 724), and if that be admitted, the main

groups of the Pisces? may be termed subclasses. These are six

in number—namely Paleopterygii, Neopterygii, Cladistia, Rhipi-

distia, Actinistia, and Dipneusti, the probable relationships of

which are indicated in the following scheme : —

Actinistia. Dipneusti.

Cladistia. | Rhipidistia.

Neopterygii.

Paleopterygii

The Neopterygii are defined by the dorsal and anal fins with

pterygiophores equal in number to the dermal rays, the absence of

clavicles and of paired gulars, the presence of an interoperculum,

* It is worth noting that one supramaxillary and a supra-angular, and two supra-

maxillaries and no supra-angular are alternative conditions, almost suggesting that

the maxillary has captured the supra-angular from the lower jaw. But it seems more

likely that the supra-angular has gone, and that as the maxillary lengthened another

suborbital has become attached to it.

+ TI exclude from the Pisces a number of Paleozoic groups of uncertain relation-

ships,.e. g. Arthrodira, Asterolepida, Osteostraci, etc.

SKELETON OF LEPIDOSTEUS. 459

and by the structure of the skull (e.g. typically 5 otic bones,

alisphenoids and orbitosphenoids, etc.) and of the’ hyoid arch

(a symplectic, two ossifications in the ceratohyal, etc.). This sub-

class contains the great majority of living fishes, and includes a

large number of orders, of which the first three are :—

Order 1. ProrosponDyYLi.

Caudal fin abbreviate heterocercal. Vertebral column acentrous,

or with centra variously developed*. Premaxillaries fixed,

attached to frontals and pierced for passage of olfactory nerves ;

maxillary free, unsegmented, usually with a supramaxillary.

Lower jaw with ‘supra-angular,” large dentigerous prearticular,

and intra-dentaries. Hyoid arch and preoperculum normally

developed, and symplectic firmly attached to quadrate. Only one

coracoid ossification, or none ; mesocoracoid bridge cartilaginous ;

one or more pectoral radials inserted on metapterygium.

Principal families :—Semionotide, Macrosemiide, Kugnathide,

Amiidee, Pachycormide, Aspidorhynchide, Pyenodontide.

Order 2. GINGLYMODI.

Distinguished from the preceding by the characters of the

jaws, Suspensorium, and opercles. Maxillary segmented, the first

one or two segments attached to preemaxillary, the rest to ecto-

pterygoid ; metapterygoids articulating with transverse facets on

wings of parasphenoid ; hyoid arch free from pterygo-quadrate ;

preoperculum small, its anterior end articulating with a condyle

on quadrate ; interoperculum large, fixed, connecting hyomandibu-

lar with preoperculum. Vertebral centra solid, opisthoccelous.

One family, Lepidosteidee.

Order 3. HALECOSTOMI.

Caudal fin abbreviate heterocercal. Pramaxillaries small,

loosely attached ; maxillary typically with two supramaxillaries ;

lower jaw without ‘“ supra-angular ” or intra-dentaries. Vertebral

centra annular or amphiccelous.

Two families, Pholidophoride and Oligopleuride.

* The various conditions are :—(1) No centra; bases of arches’ not expanded.

(This is found in all families except Amiidx and Aspidorhynchide.) (2) No centra ;

bases of arches may form laminar expansions over the notochord (some Pycno-

donts). (3) Annular or amphicclous centra of uncertain origin; no pre- and

post-centra (Aspidorhynchide, Neorhombolepis). (4) Solid outgrowths of arches

may enclose notochord, forming annular centi'a, with pra- and post-centra in caudal

region. (Some Macrosemiidze, Hugnathide, and Pachycormide.) (5) Centra disc-

“like, formed by ossification of tissues surrounding notochord, enclosing bases of

arches; prae- and post-centra in caudal region (Amiidz).

460 MR. C. TATE REGAN ON THE

The fourth Order, the IsosponDYL1, is distinguished from the

Halecostomi by the homocercal structure of the caudal fin; it is

followed by the Orders Ostartopuys1, APODES, etc.

The relationships of the families dealt with above may be

expressed diagrammatically as follows :—

Lepidosteidee.

Hlopide.

Aspidorbynchide. ;

| Leptolepide. /

\ jp

Pachycormide. |

Ayiidee ; 4

Pin ut | | | Oligopleuride.

Eugnathide. | Wy

Pholidophoride.

| i

Pyenodontide. | ve

~~ Macrosemiide. we

aoa | ca

Semionotide.

Paleoniscide.

GENERAL CONSIDERATIONS.

In dealing with the origin and evolution of the lower Neoptery-

gian Fishes I have tried to emphasize the conclusion at which

I have arrived, that their differentiation has been adaptative,

changes of structure being especially related to the nature of

the food and the method of procuring it.

This is well illustrated by the following example :—The Semi-

onotide, probably slow-swimming fishes feeding on molluscs,

crustaceans, etc., appear to have been derived from the Palzo-

niscidee and have themselves given rise to the Hugnathide,

SKELETON OF LEPIDOSTEUS. 461

which resemble the Palzoniscidx in form, shape of the fins, size

of the mouth, and structure of the teeth.

In comparing Hugnathus with Palwoniscus, both strong-

swimming predacious fishes, it is not evident that any of the

differences between them are adaptative. But these differ-

ences are due to the Semionotid ancestry of the Hugnathide,

and there is good reason for believing that they were adapta-

tive in origin,

The later evolution of the Neopterygian Fishes offers numbers

of parallel examples: modifications persist and become the basis

for further modifications.

ON THE ORIGIN OF FLIGHT IN BIRDS. 463

25. On the Origin of Flight in Birds.

By Baron Francis Noposa.

[Received April 24, 1923: Read April 24, 1923. ]

(Text-figures 1-7.)

In 1907 { brought forward the idea that birds originated from

bipedal Dinosaur-like running reptiles in which the anterior ex-

tremities, on account of flapping movements, had gradually turned

to wings without thereby affecting terrestrial locomotion (14).

This idea has been on several occasions criticised by various

authors (1, 3, 9, 12, 16).

In consequence of new material having turned up that has some

bearing on this subject, the time seems to have arrived when

the different criticisms to which the hypothesis of a ‘‘ running

Proavis”’ has been subjected can be reviewed.

The first criticism was brought forward by Hay (9), who dis-

agreed with my idea on account of my having drawn Proavis

with too short arms and with a reduced and rotated haliux.

According to Hay’s view, birds never could have originated from

such a form unless considerable rejuvenation might be presumed.

The Dinosaurs Hay supposes to have been originally quadru-

edal.

: Hay’s view was endorsed by Abel (1), who, on account of

the rotation of the hallux and the development of a prehensile

manus in Archeopteryx and Dinosaurs, in 1912 supported the

idea that both birds and Dinosaurs were derived from arboreal

forms. Steiner tried to derive arguments against my hypo-

thesis from the diastaxic arrangement of the feathers, the origin

of which he traced back to somearboreal Agamid (16). Ina later

paper (17) he likewise emphasised the prehensile structure of the

manus in primitive birds, but at the same time he believed. that

Abel’s arguments, based on hallux and pollex, were devoid of

foundation.

Beebe (3) and Lucas (12) fully appreciate the difficulty that

arises from the fact that, while in all arboricolous passive fliers the

centre of the parachute surface coincides, as it must, with the

centre of gravity of the animal, this could not have been the case

in a primitive arboricolous bird having feathers only on its tail and

arms. They think to overcome this difficulty by assuming that

the most primitive arboreal birds had quills also on their legs.

This stage in avian evolution Beebe terms the Tetrapteryx

stage; and in support of his hypothesis, he falls back on the

observation that temporarily quills develop on the legs of several

kinds of birds, and are later replaced by down.

464 BARON FRANCIS NOPCSA ON THE

In 1922 Abel (2) dropped the idea that birds and Dinosaurs

originated from quadrupedal arboreal forms, and suggested that

birds might be the offspring of a bipedal Varsius-shaped reptile.

Heilman tried (11) to reconcile my views with those held by

Hay and Abel, etc.

It is apparent from this review, that anatomical arguments

against the hypothesis of a running Proavis are mainly based

on the structure of the manus and the pes, and partly on the

occurrence of certain quills. None of the authors cited has

considered either the shoulder-girdle or the pelvis of birds, in

spite of these parts being correlated with the feet.

Since Hay and Abel’s argument from the occurrence of a

functional hallux in different birdsis of great anatomical weight,

it has to be dealt with first. The functional hallux in many

birds must either be an old structure inherited from an arboreal

Proavis, or it must be a newly-acquired character that was

missing in the running Proavis.

Although we know comparatively little about the Triassic

bipedal reptiles among which the running Proavis might be

looked for, nevertheless by the study of fossil footprints some

points of the structure of the feet of bipedal Triassic reptiles can

be elucidated (13). In my argument, four types of Triassic

footprints are of special interest, for they throw some light on

the evolution of the hallux in these otherwise nearly unknown

types. These four types can be grouped around the footprints

called Plectoterna, Ammopus, Anchisauripus, and Gigandipus.

Believing that I have proved all Dinosaurs to have developed

from bipedal lightly-built animals (15), I consider these tracks

to be Dinosaurian.

Plectoterna has long and slender toes, and a very long metatarsus

resting always on the ground and indicating a plantigrade animal

like Hallopus (text-fig. 1). Ammopus has also slender toes, but

it shows, instead of the metatarsal impression only, the impres-

sion of a rounded pad, proving that this animal was digitigrade,

as is Procompsognathus (text-fig. 2). Anchisawripus has thicker

toes than either of these two types, and was frequently planti-

grade. It is the track of Anchisawrus (10) (text-fig. 3). Gigan-

dipus is of still heavier build, but always digitigrade, and the

track of some unknown Megalosaurian.

Around each of these four types several other Triassic foot-

prints can be grouped that differ mainly in the development of

the hallux. Tothe Plectoterna type belong Palamopus, Haocampe,

Herpedactylus, Xiphopeza, and Corvipes; to the Ammopus type

are allied Hupalamopus, Polemarchus, and Platypterna; the

Anchisauripus type is suggested by Anomepus, Apatichnus, and

Grallator; and the Gigandipus type is approached by Sawropus

and Hyphepus.

Considering the hallux of these different tracks, it may be

remarked that in some the hallux articulates very low down and

is not rotated (Palamopus, Hxocampe, Anomepus) ; in others the

ORIGIN OF FLIGHT IN BIRDS. 465

rotation is more or less marked, but the hallux is still always well

in touch with the ground (Herpedactylus, Xiphopeza, Platy-

pterna, Palamopus, Polemarchus) ; yet in other tracks (Ammopus,

Apatichnus, Anchisauripus, Sauropus) the rotation is still more

accentuated, and the articulation is well raised above the ground,

so that only the tip of the hallux makes an impression ; and finally

in some of the tracks (Corvipes, Platypterna, Grallator) the hallux

can be seen no more.

Thus in all four types of tracks a simultaneous and indepen-

dently acquired rotation and abbreviation of the hallux can be

detected. Since it is quite impossible to assume that all four

types of tracks were exclusively made by animals becoming

arboreal, it is evident that the curious rotation of the hallux

in all these types was not due to arboreal adaptation, but to

some other hitherto unknown factor *.

The hallux that is attained in all these types recalls that

Text-figure 1.

Cc D E

Footprints of plantigrade Triassic reptiles.

(A) Herpedactylus; (B) Xiphopeza; (C) Plectopterna ; (D) Plectopterna ;

(E) Corvipes. ‘

of those birds in which it is a non-functioning digit. If it

can be proved that the functioning hallux in birds developed

from a non-functioning hallux, this observation is a strong

argument for a Dinosaur-like terrestrial bipedal running Proavis.

If the functioning hallux in birds developed from a non-

functioning but rotated and abbreviated hallux, this change

must have left some traces. In the functioning hallux of birds

two distinct features seem actually to point to such a change.

First, one may mention that even in birds with a strong func-

tioning hallux the first metatarsal is reduced. This shows that

in the history of these birds some stage of evolution was passed

in which the hallux was but feebly attached to the leg, and when

consequently the hallux was not much used for grasping.

%* The reason why the rotated hallux is more reduced in some Dinosaurs than in

others will be discussed on another occasion.

466 BARON FRANCIS NOPCSA ON THE

As a second feature pointing in the same direction, the

great variety of the flexor muscles of the hallux can be

brought forward. If the functioning type of hallux were the

primitive type in birds, then the arrangement of the flexor

muscles could be everywhere alike, whereas, if the hallux was

rejuvenated and secondarily but independently strengthened in

different groups of birds, the flexor muscles might vary. As is

known, the latter is actually the case, and the variation has been

used in the classification of birds (7). These detailed observa-

tions show that the functioning hallux in birds is perhaps a

secondary adaptation. A feature worth mentioning is that in

the apparently tridactyle foot of those bipedal animals that passed

through a stage of arboreal specialisation (Kangaroo) the fourth

toe is the longest (5), while it is the third that is the longest in

Dinosaurs, birds, and Alactaga (14).

Text-figure 2.

Footprints of digitigrade Triassic reptiles.

(A) Anomeepus ; (B) Apatichnus ; (C) Anchisauripus; (D) Grrallator.

Turning now from the pes to the manus in primitive birds

(Archeopteryx), it may be remarked that this is a long and

slender organ, adapted to some extent for grasping, but this alone

is again no sign of arboreal specialisation.

The partial use of the anterior limb as a grasping organ

occurs in different terrestrial quadrupeds. The fore leg is used

as a manus in the Kangaroo when taking up food ; it is similarly

used in mice and rabbits and, partly for the same purpose and

partly also as a weapon, in cats and bears. Members of each

of these groups are, it is true, partly also arboreal ; but neverthe-

less the terrestrial representatives in these groups do not descend

from the arboreal forms, for, on the contrary, it was the use

of the anterior limb as a grasping organ that enabled some of the

terrestrial forms to climb trees.

Steiner has proved in a recent highly valuable contribution (17)

ORIGIN OF FLIGHT IN BIRDS. 467

that in most Archosauria the ulnar phalanges are reduced,

while the radial phalanges are retained or sometimes even

strengthened. He takes this to be a sign of the arboreal adapta-

tion of Proavis. The reduction cannot be denied, but again it

does not seem to be a sign of any arboreal specialisation. In

the one group of extinct reptiles that certainly was derived

from arboreal types this veduction does not take place, for in

Pterosaurians the last finger is very strong, and this reduction

does not occur in any of the arboreal mammals (5). It is well

marked only in Crocodilia, Dinosaurs, and birds. The primitive

Dinosaurs are, as is well known, bipedal (15). In the Crocodilia

the reduction of the shoulder-girdie and the curious elongation

of the carpal bones indicate that formerly also in this group

the disproportion between the fore limbs and the hind limbs

was also decidedly greater (10). Considering that all those

mammals that are partly bipedal use the anterior limb for

Text-figure 3.

Footprints of digitigrade Triassic reptiles.

(A) Hupalamopus (somewhat modified); (B) Polemarchus; (C) Ammopus ;

(D) Platypterna.

grasping, and that Dinosaurs and Crocodilia are terrestrial

forms with a strong posterior body, it should be investigated

whether the ulnar reduction is not due to the use of the manus as

a grasping organ during terrestrial and not during arboreal life.

The first change that is attained when both the as yet unmodified

anterior limbs are used together as a primitive grasping organ, is

the inward rotation of both palmar surfaces; the next change

that must be expected is a strengthening of the radial side, for

in two palms that are half-way rotated the radial borders of the

hands come always nearer together than the ulnar borders ;

consequently they are always exposed to much greater strain

than the ulnar sides of the hands. This greater strain will be

especially felt when the food is held with both anterior feet and

pieces are detached with the teeth. Such a manus must of course

differ strongly from the manus of every arboreal type, for in the

468 BARON FRANCIS NOPCSA ON THE

arboreal types it is of importance that the grasping surface of

the whole hand should be as great as possible. his is attained

by the retention of all phalanges, and especially by the retention

of the fourth (5). In animals in which the function of the

manus is different, a reduction of the ulnar phalanges, recailing

somewhat the reduction of the fourth and fifth digits of the

human hand, can take place.

The most reduced manus of this sort attained by a terrestrial

animal is the manus of Struthiomimus, and the resemblance tothe

manus in Archwopteryx is quite easily made out (18). Struthio-

mumus, however, did not live on trees (text-fig.5). This genus is

not only suitable for demonstrating that a hand can be developed

on terra firma, but also that a secondary elongation of the fore

limb can be attained without adaptation to flight (18). Com-

paring the Triassic Procompsognathus, the Jurassic Ornitholestes,

and the Cretaceous Struthiomimus, none of which were arboreal,

Text-figure 4. _

Bones of pes in Dinosaurs.

(A) Massospondylus ; (B) Antrodesmus; (C) Compsognathus.

it may be remarked that the anterior limb is short in Pro-

compsognathus, longer in Ornitholestes, and longest in Struthio-

mimus. ‘This shows a steady increase in length. Since a similar

elongation occurs also in the Sauropoda (for in this group the

long-armed Brachiosauride certainly descended from the other,

but short-armed forms) this elongation may well be compared

with the elongation that can be seen in the anterior limb of the

most primitive bird (Archwopteryx). It is a sort of rejuvenation

that suggests also the rejuvenation of the avian hallux already

discussed. ‘This observation proves to a certain extent that

Hay’s objection to my drawing Proavis with relatively short arms

does not weigh as much as it seems to do at first.

After having discovered in the course of our investigation

that the similarity of hands and feet in birds and Dinosaurs, and

their fundamental difference from these parts in all arboreal

ORIGIN OF FLIGHT IN BIRDS. 469

mammals and lizards, not only do not form an argument for the

arboreal origin of birds and Dinosaurs but on the contrary’ plead

for their origin, from running ancestors, and after: having, as I

believe, refuted the arguments brought forward by Abel, Hay, and

Steiner, I think the next step should. be to mention all those

points in the anatomy of birds which exclude.an arboreal Proavis

from the history of this group. #4 peas OITA

All birds have a fused metatarsal. Where it is but: feebly

fused, as in the Penguins, this isa return toa primitive stage, for

in the fossil Penguins of the Seymour Islands the metatarsals are

more strongly fused than in any recent species (20).

With exception of arboreal birds, not a single arboreal verte-

brate animal is known in which a fusion of the metatarsals is

even indicated. This. fusion shows that Proavis must have

moved differently from all recent and fossil arboreal forms.

Text-figure 5.

a Yy

\¥

Bones of manus in Dinosaurs and birds.

(A) Massospondylus; (B) Antrodesmus ; (C) Struthiomimus; (D) Ornitholestes ;

(E) Archeopteryx.

Proavis cannot have leaped in a more or less frog-like manner like

Tarsius, Galago, or frogs, for in such arboreal animals the

flexible tarsus is modified and its proximal part elongated ; it

cannot have climbed, for the manus is specialised on other lines :

it cannot have jumped like a monkey, for birds show no tuber on

the heel. Even Pedetes, though terrestrial and provided with com-

pact tarsal bones, shows how even the firmly united tarsal bones

are. finally affected by jumping, for in the recent Pedetes these

bones are somewhat elongated, while they are short in a more

primitive fossil type that Prof. Stromer intends to describe.

This elongation of the tarsal bones is necessary in Jumping

animals, for in these itis essential that that part of the pes

should be rigid and long which immediately touches the tibia,

and not the part beyond the flexible tarsal bones. In bipedal

stalking and walking animals the metatarsals can move easily

elongate so as to ensure'a greater stride.

Proc. Zoot. Soc.—1923, No. XXXI. 31

‘470 BARON. FRANCIS NOPCSA ON THE

| Since the fusion of the metatarsals occurs exclusively in terres-

trial: forms.(Alactaga, ete., and Artiodactyla), Proavis must once

have, been terrestrial. Supposing Proavis acquired this marked

specialisation while living in trees, it must have moved about

there in a hitherto unknown fashion ; supposing, however, that

birds learnt to fly, and only then became bipedal, then we have a

quite unique sort of evolution, for all those quadrupeds that learnt

to fly in the quadrupedal state became unable to walk. Conse-

quently birds’ must ‘first’ ELS been bie and only then have

learnt to glide.

Another argument that can also be brought forward against

the hypothesis of an arboreal Proavis can be deduced from the

pelvis of the most primitive bird we know (Archeopteryz).

While the pes in dArcheopterya shows that this type had

Text-figure 6,

Pubes of Dinosaurs and birds.

(A) Celurus; (B) Archeopteryx; (C) Struthio; (D) Dromeus.

already taken to arboreal life, the pelvis, as shown by the pubes,

is of the long and narrow type. This type is only met’ with

in Dinosaurs jand one cursorial bird. In the other birds, and

especially in the arboreal birds, the pelvis is usually broad and

short, and the pubes are wide apart (text-fig. 6).

In accordance with this structure and with the abbreviation of

the vertebral column, the arboreal birds mostly hop, while the

ground. birds walk or run, In the best runners the body is

the longest, and the pelvis always remarkably narrow. Since

a long body and narrow pelvis are seen in Ar cheeopteryx in a

marked manner, its arboreal specialisation can only be called

very slight. It was assuredly derived from'a terrestrial stock.

The shoulder-girdle in birds also indicates that their common’

ancestor must have once passed through: a sea Seage. of

ORIGIN OF FLIGHT IN BIRDS. 47 |

specialisation. Whenever in nearly-related Mammalia terrestrial

and arboreal forms occur, it is in the latter that the clavicles

are certainly present. According to Weber (19), the clavicles are

missing in the terrestrial genus Hystrix, but present in the

arboreal Hrethizon.

Since in cursorial forms in which the humerus moves only in

a plane that runs parallel to the body, no lateral strain is brought

to bear on the glenoidal fossa, of course in such forms clavicles

are useless, but they must be retained in the forms that extend

the arms strongly sidewards, for here such lateral strains occur.

If birds are derived from terrestrial forms in which the avms

and the shoulder-girdle were temporarily reduced, it is in the

shoulder-girdle that this must show.

In the Thecodontia, in which, as shown by the relative length

of the fourth digit, the elbows stood yet somewhat off from the

body, an interclavicle and clavicles are present ; in the Crocodilia,

only the interclavicle remains, but also the fourth toe is abbrevi-

ated, showing that the lateral strain on the leg was already

relatively small ; finally in all the Dinosaurs, as we know, clavicles

and interclavicula are absent. In Dinosaurs, moreover, a well-

ossified sternum is present, arising from two centres of ossification.

If we turn now from these reptiles to modern and extinct birds,

we find always an osseous sternum and further on a bone that

has been considered as corresponding to the clavicle of reptiles,

This bone has been called the furcula, and it is this name that

I adopt, for it has no phylogenetical meaning. In some birds

this bone touches the sternum, in others, as Archwopieryx, it

does not.

Not even the slightest trace of an interclavicle has ever been

found in birds (6), although this bone has been eagerly sought

for. The scapula and coracoid in Archeopteryx recall these

elements in the primitive bipedal Dinosaurs and Struthious

birds (text-fig. 7).

If the present arboreal birds are the direct descendants of a

élavicle-bearing quadrupedal lizard-like reptile that took to

climbing trees, surely a clavicle ought to be present. and perhaps

even an interclavicle might be expected, for it is exactly the

interclavicle that is retained both im Crocodiles and Mosasaurs

when the osseous clavicles are already gone. If features exist

that show that the furcula of birds is a newly-acquired bone and

that both clavicles and interclavicle are gone, then this is an

important argument in favour of a primitive running Proavis.

As can be seen, a great part of the question turns on the nature

of the furcula of birds.

While the clavicles in all animals are always paired bones

attached to the scapula with one end and to the interclavicle

with the other, the furcula of birds adheres to the coracoid with

one énd and fuses, even in Archeopteryx, in the middle with the

furcula from the other side.

31*

472, BARON FRANCIS NOPCSA ON THE

If we wish to identify the furcula of birds with the clavicle of

other vertebrata, we must assume that, first of all, the clavicle

managed to slip along the anterior border of the scapula, then, to

pass the glenoid fossa without, however, affecting the motion

of the humerus by so doing, and finally to attach to the coracoid.

Worse than that, we must also suppose that in the middle of the

body a very curious, almost mysterious, and unique reduction

managed to wipe out even the last ontogenetical trace of the

interclavicle without affecting the clavicles. Surely it is some-

what risky to evoke a hypothesis of this sort.

Supposing now, instead of these changes, that the furcula is an

ossified tendon that was developed when birds began to glide to

strengthen a reduced scapular arch, it is easy to explain its

origin on the coracoid, the primary fusion of the median ends, its

Text-figure 7.

Scapula and coracoid in Dinosaurs and birds,

(A) Gorgosaurus; (B) Archeopteryx ; (C) Struthio; (D) Nothura.

successive fixing on to the sternum, and the absence of clavicles

and interclavicle. Of course it is necessary in this case to fall

back on the hypothesis of a terrestrial and bipedal Proavis in

which the shoulder-girdle had suffered reduction.

In accordance with some old zoologists, I consider the furcula

to be an ossified tendon, and not to be homologous with the

clavicles in reptiles. Since this supposition agrees with the

results hitherto obtained, I think this hypothesis is the right

one.

All evidence brought forward up to the present shows that the

flying state of birds was preceded by a state of bipedal running.

Having established this, of course it becomes necessary to investi-

gate when and where primitive feathers and passive flight began.

if passive flight and feathers arose after cursorial reptiles had

ORIGIN OF FLIGHT IN BIRDS. 473

taken up an arboreal life, they must have developed feathers

on arms and legs, for also in these long-necked and long-tailed

forms the centre of the supporting gliding surface had to coincide

with the centre of gravity of the body. Hence, evidently out of

these reptiles a Vetrapteryx had to develop.

Of course an exclusively bipedal Tetrapteryx would encounter

great difficulties when climbing. During gliding, furthermore,

in spite of new difficulties presenting themselves, the knees of such

a creature had evidently to be turned outward; the feet had to

be stretched away from the bady and had to be brought into the

same plane as the anterior members, else the feathers of the legs

would have been quite ineffective. All this and the difficulties

encountered when climbing without using the anterior limbs

would have been very detrimental to bipedal locomotion. Even

amere glance at Beebe’s hypothetical Tetrapteryx shows that such

a creature could not have been bipedal.

Since Beebe’s Tetrapteryx is a rather short-necked animal,this

point of his drawing is likewise open to criticism. All birds

originated evidently from a bipedal and long-necked ancestor

that carried the head upright. It will, however, always remain

an unsolved problem how Tetrapteryx could have managed to

carry such a long neck and an upright head when gliding:

arboreal animals are rather short-necked.

In consequence of all this, the Vetrapteryx hypothesis, which

is but a revival of Pycraft’s arboreal Proavis, must evidently be

dismissed, for it can in no way account for the persistence of

bipedal locomotion in birds.

The whole Tetrapteryx hypothesis is based exclusively on

the one interesting observation, that in some birds a row of

quills is present on the upper part of the legs.

Without diminishing the interest of this observation, it has to

be remarked that in a long-tailed animal this is the very place

where quills would be ineffective, and that the phylogenetical

value of this observation varies according to the manner in which

one estimates the relation between quills and down.

If the down is considered to be the primitive type of feather

which originated as a body cover to prevent the loss of heat, and

the quills are considered as its modification, of course in such a

case the occurrence of a row of quills on the leg can be taken as

a proof that the legs were once used for flying; but if, on the

other hand, the quills are considered as primitive, then such a

reasoning is fallacious. In this latter case the occurrence of

quills in places that are later covered with down possibly only

shows that formerly a part of the side of the body of the primitive

birds was covered with quills arranged in rows. Such quills can

then gradually have been replaced by down, except in places

where they served for flight or steering.

In such a case it may be assumed that the quills which

evidently first originated on the arms and tail gradually spread

to adjacent places where they were not really wanted,

ATA BARON FRANCIS NOPCSA ON THE

A hypothesis that assumes that organs that develop in one

part should be capable of spreading on to adjacent regions where

they are of little or no value, needs naturally to be proved.

In all species of Rhinoceroses in which a strong nasal rugosity

is present to serve as bage for the dermal horns, the surface of the

jugal arch, the lacrymal, prefrontal, the otic region, and even

the posterior and exterior surface of the mandible, is also rough

(Rh. sumatrensis, Rh. bicornis). Inthose Rhinoceroses, however,

in which the nasal tuberosity is weak or wanting, the jugal and

lacrymal and the other bones mentioned above are also smooth

(Lh. unicornis, Aceratherium).

A similar feature, as in Rhinoceros, can be found also in

Suide. Hylceochwrus does not only show strong rugosities on

the maxillaries and jugals, where the facial warts are developed,

but also on the mandible and the. frontal. Sus verrucosus

shows similar but less marked features. Contrary to this, in

Sus barbatus and Sus cristatus all the skull bones are smooth,

and correspondingly on the cheeks strong warts and rugosities

are wanting.

That the rapidly spreading rugosities in Rhinoceros and Sus,

though in connection with warts, do not exclusively depend on

the presence of these excrescences, can be demonstrated by the

different Giraffes. Also in the latter a spreading of exostosis 1s

traceable until the whole head is covered and even the mandible

is affected: nevertheless, the skin in these animals remains smooth,

and ig quite normally covered with hair. Less clearly than in

these three groups, the spreading of useless exostosis can be

observed in the bovine tribe. Tn Bos bubalis, Bos gaurus, and

the Bison, all of which have strong bony horn-cores, the

rugosities of the frontal spread to the orbital and otic region,

while in Sos depressicornis, in which the horn-cores are weak,

the orbital and otic region is smooth. Bearing all these facts

in mind, and they could easily be multiplied, and considering

furthermore that in Crocodiles the skull bones and the bones of

the dermal armour are of altogether different origin but that

they nevertheless show the same pattern of sculpture, and that

thus also they show the spreading of one feature over the

whole body and that something similar occurs in Lepidosteus (8),

[ think i is not too much to assume that also in the primitive

Proavis the primitive feathers of the tail spread even to regions

where they were not wanted. Their attachment on a patagium

is the best proof that they never functioned. That the primitive

feathers of birds were quills and not, as generally thought, down

was recently proved by Steiner (17).

In this way also Beebe’s chief argument for the reconstruction

of his Tetrapters yx loses a great deal of its value. It only shows

that first in birds only a marginal row of quills was developed

on the arms and on the tail, that later these spread towards the

feet, and that the rest of the quills spring up later.

Summing up, one can safely state that in the skeleton of recent

ORIGIN OF FLIGHT IN BIRDS. 475

and extinct birds not a single character can be detected which

points conclusively towards a primitive arboreal adaptation, but

that several skeletal characters can only be explained by the

hypothesis of a cursorial running ancestor of birds. The other-

wise inexplicable features are: the fused median metatarsals,

the reduction of the first metatarsus, the small tarsal bones,

the position of the knee, the long symphysis of the pubes in

Archeopteryx, the lack of clavicles and interclavicle in all birds,

and the position of the occipital condyle. The latter is vertical

in birds and primitive Dinosaurs, and proves that, “unlike

arboreal forms, in both these long-necked groups the head was

primarily carried upright on a sigmoidal neck.

Apart from all these important points, it is only by means of

the running-Proavis hypothesis that the early occurrence of

numerous flightless birds can be explained. It is evident that

badly flying ground birds will much more readily return to

exclusively cursorial habits than arboreal birds, for these, before

becoming again flightless, must first descend from their trees and

become thoroughly adapted to ground life.. There exists a

great difference between the pelvis of formerly arboreal flightless

birds (for example, Dodo) and that of the Paleognathe.

The hypothesis of a running Proavis is also the single one that

accounts for the primitive structure of the palate, the free dorsal

vertebre, the low body-temperature, and the strong penis of the

greater part of the Paleognathe, for according to this hypo-

thesis these birds are, of course, much nearer related to Proavis

than the later and evidently more specialised aboreal birds. If

the birds were of arboreal origin, one woula expect the most

primitive birds among the arboreal and not the terrestrial birds.

Tn compliance with a wish of Steiner (17), I desist from using in

the course of these comparisons the term Ratite.

That in one primitive arboreal bird more specialised, however,

than an ostrich (i.e. Opisthocomus) the claws of the wing are

more used for climbing than in all the other recent birds, is not

of very great weight, for other nestlings use even their beak

when climbing, and yet nobody will consider the Parrots to be

ancestral to all the other birds. The climbing of Opisthocomus

may quite well be regarded as a secondary adaptation, for the

development of the pinions of the wings in Opisthocomus sutters

a curious retardation that points in this direction (14). The

sternum of Opisthocomus shows likewise a curious modification.

The primitive structure of the nest of ground birds when

compared with the nest of the arboreal birds, and the more

brilliant coloration of the latter, also agree very well with the

hypothesis of the terrestrial origin of all birls. Even the

curious fact that the nestlings of ground birds as a rule leave

their nest comparatively soon after birth seems to some extent

to favour this hypothesis (14).

Up to the present all critics of the “running Proavis” hypothesis

have only tried to find apparent difficulties in that hypothesis,

476 BARON FRANCIS NOPCSA ON THE

and have never considered what it explains; now it is their

turn to explain all the points that have been brought forward

in the course of this paper by means of their hypothesis of an

arboreal Proavis. It may .be that more difficulties will be

encountered than are expected.

Some critics (12) say that the comparison of a running

Proavis with the different flying fishes is no comparison at all,

because fishes do not move their pectoral fins in flight. In spite

of this, I repeat that the comparison is rather a good one

because

(a} both types are vertebrata in which the anterior limbs are

primarily only used for a similar passive gliding ;

(b) in both, during gliding the centre of gravity is behind the

upholding surface, giving it a convenient elevation ;

situated at the posterior end of the body and behind the

gliding-planes ;

(dz) in both, the locomotive power can immediately be brought

again into action as soon as gliding through the air ceases,

and so gliding can soon be regained.

None of these four characters can be found in arboreal passive

fliers.

Zschokke (21) has remarked that some Crustacea develop a

sort of flying like that of flying fishes; so there is no reason why

a terrestrial vertebrate animal should not also have been capable

of developing it. No mechanical difficulties exist. To those who

go on to argue that, with the exception of flying fishes, all other

flying vertebrates descended from arboreal animals, and that

therefore this was presumably also the case in birds, I retort

that, with the exception of flying snakes, all arboreal vertebrata

and naturally all their flying offspring are without exception

quadrupedal, while this is not the case in birds.

Literature referred to.

(1) Apet, O.—Grundziige der Paleobiologie der Wirbeltiere,

Stuttgart, 1912.

(2) Ase, O.—Lehrbuch der Paleozoologie. Jena, 1922.

(3) Berse, C. W.—Tetrapteryx Stage in the Ancestry of Birds.

Zoologica, New York, 1915.

(4) Branca, W.—Fossile Flugtiere und der Erwerb des Flugver-

mogens. Abhandl. Preuss. Akad. d. Wiss., Berlin, 1908

(5) Dotto, L.—Les ancétres des Marsupiaux étaient-ils arbori-

coles. Miscellanées biologiques dediés au Prof. A. Giard.

Paris, 1899.

(6) Firprincer, M.—Vergleichende Anatomie des Brustschul-

tergirtels. Jenaische Zeitschr. f. Naturwiss. 1902. .

(7) Garrop, A. H.—On the Disposition of the deep Plantar

Tendons in Birds, Proe. Zool, Soc, London, 1875,

ORIGIN OF FLIGHT IN BIRDS. ATT

(8) Goopricxu, E.S.—On the Structure of Bone in Fishes. Proc.

Zool. Soe. London, 1913.

(9) Hay, O. P.—On the Manner of Locomotion in the Dinosaurs,

with remarks on the Origin of Birds. Proc. Washington

Acad. of Science, 1910.

(10) Hurnz, Fr.—Beobachtungen iiber die Bewegungsart bei

Krokodilen. Biolog. Centralblatt, xxxii. 1913.

(11) Herrmann, G.—Fuglenes Afstamning. Dansk. Ornithol.

For. Tidskr. Kopenhagen, 1916

(12) Lucas, F. A.—Animals of the Past. 6th Edition. New

York, 1922.

(13) Luni, L.—Triassic Life of the Connecticut Valley. Con-

necticut State Geolog. and Nat. Hist. Survey, Bulletin 24.

(14) Noposa, F.—Ideas on the Origin of Flight. Proc. Zool.

Soc. London, 1907.

(15) Norcsa, F.—Notizen iiber Dinosaurier, Parts 1 and 5.

Centralb. f. Min. Geol. Paleont. 1917-1918.

(16) Sremer, H.—Das Problem des Diastataxie des Vogeltligels.

Jenaische Zeitschr. f. Naturwiss. 1917.

(17) Sverver, H.—Die ontogenetische und phylogenetische Ent-

wicklung des Vogeltliigel Skelettes. Acta Zoologica,

Stockholm, 1922.

(18) Sternmann, G.—Laufvogel und Flugvégel. Anatom.

Auzeiger, 1922.

(19) Wexner, M.—Die Saugetiere.

(20) Wiman, C.—Vorlaufige Mittheilung tiber alttertiare Verte-

braten der Seymourinsel. Bull. Geol. Inst. Universitat

Upsala, 1905.

(21) Zscuoxke, F.—Der Flug der Tiere. Berlin, 1919.

198): ad ha

TAMA ET

THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 479

EXHIBITIONS AND NOTICES.

March 20th, 1923.

Prof. EK. V. MacBrips, 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 February, 1923 :—

The registered additions to the Society's Menagerie during the

month of February were 141 in number. Of these 76 were

acquired by presentation, 27 were deposited, 17 were purchased,

and 21 were born in the Menagerie.

The following may be specially mentioned :—

2 Iliger’s Marmosets (Mystax illigeri), from the Upper

Amazons, new to the Collection, deposited on February 6th.

1 Banded Coati (Nasua vittata), from British Guiana, pre-

sented by The Viscount Dunsford on February 13th.

4 Reindeer (Rangifer tarandus), from Finland, purchased on

February 24th.

2 Hared Vultures (Otogyps auricularis), from South Africa,

purchased on February 16th.

Mr. F, Martin Duncan, F.R.M.S., F.Z.8., exhibited a speci-

men of a caterpillar (¢ Charagria virescens) infected with the

Entomogenous fungus Cordyceps robertsii, which he had recently

received from Tauranga, New Zealand. The specimen showed

the long stroma of the fungus growing out through the tissues

of the joint between the head and first segment of the thorax of

the caterpillar, and bearing at its free end the densely packed

perithecia; the whole aerial stem bearing a strong resemblance

to a miniature bullrush. The native names for this so-called

plant-eaterpillar are “ aweto,” “ pepeaweto,” and “ hotete”; and

the burnt ashes of these fungus-infested caterpillars, powdered

and mixed with water, were used in ceremonial tattooing of rank

and tribe. es

Mr. M. A. C. Hriyton, F.Z.8., exhibited, and made remarks

upon, the skin of a Lioness showing considerable development

of mane.

April 10th, 1923.

Dr. A. Surrn Woopwarp, F.R.S., Vice-President,

in the Chair.

The Secrerary exhibited, and made remarks upon, a series of

photographs of the deep-level reservoirs of the Society’s new

Aquarium.

480 CAPT. R. B. MURRAY ON A GIANT CENTIPEDE,

Mr. G. C. Rosson, M.A., F.Z.S., exhibited, and made remarks

upon, the snail Planorbis dufouri Graells, the intermediate host

of Schistosoma (Bilharzia) hematobium, in Portugal.

Dr. G. M. Vevers, F.Z.S., gave an account of his recent visit

to Zoological Gardens in Holland and Belgium.

April 24th, 1923.

Prof. EK. W. MacBrips, F.R.S., Vice-President,

in the Chair.

The Secrerary read the following Report on the Additions to

the Society’s Menagerie during the month of March, 1923 :—

The registered additions to the Society’s Menagerie during the

month of March were 234 in number. Of these 154 were

acquired by presentation, 21 were deposited, 54 were purchased,

1 was received in exchange, and 4 were born in the Menagerie.

The following may be specially mentioned :—

2 Black Spider-Monkeys (Ateles ater), from South America,

purchased on March 21st.

1 Indian Wild Dog (Cyon dukhuensis), from the Central Pro-

vinces, India, presented by Major T. §. Paterson on March 2nd.

3 Axis Deer (Amis amis), from India, presented by James A.

de Rothschild, Esq., on March 29th.

1 female Llama, presented by A. H. Winefield, Esq., F.Z.8.,

on March 24th.

Capt. R. B. Murray, F.G.S., F.R.G.S., exhibited living and

mounted specimens of a Giant Centipede from Trinidad, and made

the following remarks :—

The two specimens of the Giant Centipede recently presented

to the Zoological Gardens were captured on the small island of

Gasparee on the west coast of Trinidad and in the Gulf of Paria,

which latter separates the island of Trinidad from the Spanish

Main, or Venezuela.

The species is confined to the tropical latitudes of South

America, and in certain localities is by no means uncommon, as,

for instance, amongst the Gulf islands of Trinidad, which enjoy

the reputation of breeding “the largest centipedes in the world.”

Trinidadans are familiar with this little dragon of the islands,

yet remarkably enough it is not’ known to have been found on

the mainland of Trinidad, though the whole group of islands are

closely associated both geographically and geologically. It is,

however, possible that the Gulf islands offer certain peculiar

conditions of environment favouring their existence, and that

their occurrence on the mainland of Trinidad has escaped

observation by reason of their comparative scarcity.

LIEUT.-COL. S: MONCKTON COPEMAN ON A RARE FLY. 481

The Giant Centipede is nocturnal in habits, but it may some-

times be found on the move during the day, and when disturbed

its rustling passage through the undergrowth is distinctly

audible. Occasionally it is found sheltering in the rock fissures

leading to a deep-seated Parasol Ants’ nest, and when the latter

is attacked with a chemical exterminator the bewildered lodger

emerges.

Lieut.-Col. S. Moncxron Copeman, M.D., F.R.S., and Major

EK. EK. Ausren, D.S.0., F.Z.8., exhibited, and made remarks upon,

photographs and specimens of the rare fly Hylephilas obtusa Zett.

The exhibition included the only male of this species captured

in this country in the neighbourhood of the burrows of the bee

Andrena fulva, on which the female of this fly is parasitic. ‘The

specimen was captured on Primrose Hill, London.

No. 238.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

March 20th, 1923.

Prof. E. W. MacBrrps, F.R.S., Vice-President,

in the Chair.

The Secrerary read a Report on the Additions te the Society’s

Menagerie during the month of February 1923.

Mr. F. Martin Duncan, F.R.M.S., F.Z.S., exhibited, and

made remarks upon, a specimen and photographs of a Caterpillar

from New Zealand infected with the Entemegenous fungus

Cordyceps robertii Hooker.

Mr. M. A. C. Hinton, F.Z.S., exhibited, and made remarks

upon, the skin of a Lioness. .

Dr. Puiirpra C. Espairr, F.Z.8., gave a résumé of Mr. SuSHTL

Cu. Sarkar’s paper on “ A Comparative Study of the Buceal

Glands and Teeth of Opisthoglyph Snakes, and a Discussion on

the Evolution of the Order from Aglypha.”

A paper “On the Mammals obtained in Darfur by the Lynes-

Lowe Expedition” by Mi. Oupristp Tuomas, F.R.8., F.Z.5., and

Mr. M. A. C. Hinton, F.Z.8., was communicated. by Mr, Hinton.

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent's Park, N.W., on the Tuesday followmg the date of Meeting to which

it refers.. It wall be issued, along with the ‘ Preceedings,” 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, er, if desired, sent post-free for

the sum ef Sew Shidlings per annum, payable in advance,

Mr. R. I. Pocock, F.R.S., F.Z.8., gave a résumé of his papers

(1) On the External Characters of Hlaphus, Hydropotes, Pudu,

and other Cervide ” and ‘“ (2) The Classification of the Sciuridz.”

The next Meeting of the Society for Scientific Business will be

held on Tuesday, April 10th, 1923, at 5.30 p.m., when the

following communications will be made :—

Dr. G. M. VeEvers, F.Z.S.

Notes on a recent visit to Zoological Gardens in Holland

and Belgium.

Prof. K. Kosranecki, M.D., LL.D.

On a Remnant of the Omphalo-mesenteric Arteries in the

Manatee.

Dr. Cuas. F. Sonntac, F.Z 8.

On the Anatomy, Physiology, and Pathology of the Chim-

panzee.

The following Paper has been received :—

EK. C. Stuart Baker, F.Z.S.

Cuckoos’ Eggs and Evolution.

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,

Rrcent’s Park, Lonpon, N.W. 8.

March 27th, 1923.

No. 239.

ABSTRACT OF THE PROCEEDINGS

ZOOLOGICAL SOCIETY OF LONDON.?

April 10th, 1923.

Dr. A. Smira Woopwarp, F.R.S., Vice-President,

in the Chair,

THE SEcRETARY exhibited, and made remarks upon, a series of

photographs of the deep-level reservoirs of the Society’s new

Aquarium,

Mr. G. C. Rorson, M.A., F.Z.S., exhibited, and made remarks

upon, the snail Planorbis sufourti Graells, the intermediate host

of Schistosoma (bilharzia) hematobiwm, in Portugal.

Dr. G. M. Vevers, F.Z.S., gave an account of his recent visit

to Zoological Gardens in Holland and Belgium.

Dr. Cuas. F. Sonntac, F.Z.8., gave a réswmé of his paper “ On

the Anatomy, Physiology, and Pathology of the Chimpanzee.”

* 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 Stx Shillings per annum, payable in advance.

In the absence of the Author, Prof. K. Kostanecki’s paper

“On a Remnant of the Omphalo-mesenteric Arteries in the

Manatee,” was taken as read. -

The next Meeting of the Society for Scientific Business will be

held on Tuesday, April 24th, 1923, at 5.30 p.m., when the

following communications will be made:—

R. B. Murray, F.G.S., F.R.G.S.

Exhibition of a Giant Centipede from Trinidad, and mounted

skins of Oil-birds.

Lt.-Col. 8. Moncxron Copeman, M.D., F.R.S., and Major E. E.

AustEN, D.S.0O., F.R.S.

Exhibition (with photographs) of an unique British Dip-

teron, taken on Primrose Hill.

Baron F. Nopcsa.

On the origin of flight in Birds.

E. C. Stuart Baxer, F.Z.S.

Cuckoos’ Eggs and Evolution.

The following Paper has been received :—

Harry Burret, C.M.Z.S.

The Transmission of Marsupial Embryos.

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,

Scevetary.

ZOOLOGICAL Society oF Lonpon,

Recent’s Park, Lonpon, N.W. 8.

April 17th, 1923.

No. 240.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

April 24th, 1923.

Prof. E. W. MacBripzg, F.R.S., Vice-President,

in the Chair.

The SecrErary read a Report on the Additions to the Society’s

Menagerie during the month of March 1923.

Capt. R. B. Murray, F.G.S., F.R.G.S., exhibited, and made

remarks upon, mounted and living specimens of a giant Centipede

from Trinidad,

Lt.-Col. S. Moncxron Copeman, M.D., F.R.S., F.Z.S., and

Major E. E. Austen, D.S.O., F.Z.S., exhibited, and made

remarks upon, photographs and specimens of an unique British

Dipteron, taken on Primrose Hill.

Baron F. Norcsa gave a réswmé of his paper “On the Origin

of Flight in Birds.”

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent's Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications ; but it may be obtained on the

day of publication at the price of Stxpence, or, if desired, sent post-free for

the sum of Sia Shillings per annum, payable in advance,

Mr. E. C. Sruarr Baker, F.Z.S., communicated his paper on

* Cuckoos’ Eegs and Evolution.”

The next Meeting of the Society for Scientific Business will be

held on Tuesday, May 8th, at 5.30 p.m., when the following

communications will be made :—

THE SECRETARY.

Exhibition of Photographs of Big Game from Choma,

Northern Rhodesia.

Miss L. E. Coersman, F.E.S., F.Z.S.

_ (1) Exhibition of living specimens of Peripatus from

Trinidad. j

(2) Exhibition of section of a nest of the Stingless Bee

from Australia.

Harry Burnett, 0.M.Z 8.

Note on a hibernating female specimen of the Marsupial

Acrobates pygmeus.

F. Marty Duncan, F.R.M.S., F.ZS.

On the Microscopic Structure of Mammalian Hairs.—I. The

Hairs of the Primates.

The following Paper has been received :—

' The Skeleton of Lepidosteus, with Remarks on the Origin

- and Evolution of the Lower Neopterygian Fishes.

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 LonDon,

Recent’s Park, Lonpon, N.W. 8.

May \sé, 1923.

18.

20.

hey

PAPERS (continued.)

On the Mammals obtained in Darfur by the Lynes-Lowe Expedition. By OnpriELp

Tuomas, F.R.S., F.Z.8., and Marrin A. C. Hinton, F.Z.8, 00.0.0... ..00 cee cece 247

Page

. On a Remnant of the Omphalo-mesenteric Arteries in the Manatee. By K.

Kosranecsi, M.D., LL.D., F.Ac.Sc. Cracow. (Text-figure 1.) ......-...-+..---+ 278

Cuckoos’ Eggs and Evolution. By E. C. Sruarr Baxer, F.Z.S. (Plates 1-IV.).... 277

. A Comparative Study of the Buccal Glands and Teeth of the Opisthoglypha, and a

Discussion on the Evolution of the Order from Aglypha. By Susuin Cu. Sarkar,

Zsa) (Rextehoures ul DOM acer we sitin Sevaidsis Sues clipe iste ot Ne Came Pee hey A

22. On the Anatomy, Physiology, and Pathology of the Chimpanzee. By Cuaruzs F.

Sonnac, M.D., F.Z.S. (Plates I.-III.; Text-figures 25-49.) .........--+...-.. 323

23. The Elephant-Seals of Kerguelen Land. By I. P. A. Rive. (PlatesI.&II.).... . 43]

24, The Skeleton of Lepidosteus, with remarks on the origin and evolution of the lower

Neopterygian Fishes. By O. Tare Reeay, M.A., F.R.S., F.Z.S. (Text-figures 1-8.) 445

25. On the Origin of Flight in Birds. By Baron Francis Norcsa. (Text-figures 1-7.).. 463

Alphabetical List of Contributors .../.... 0.4 2 oe en re mete eh ee

Ingles: OF JUDO: Shea ceentienee 4 cas Go| ageane Pn Men nero n 4.5.0.) (ttn

TE Ve epee 6 25s a oe Soar eee eee ees Pane nee ash A Ua akg: ny aoe eae eae Xvii

LIST OF PLATES.

1923, Parr II. (pp. 181-481).

Page

E. C. Stuart Baxer: Pls. 1.-1V. Cuckoos’ Eggs and Evolution .... 277

Cuas, F. Sonnvac: Pls. I.-IlI. Anatomy of the Chimpanzee .... 323

TT. P. A. Rine: Pls. I.—II. Elephant-Seals of Kerguelen Land 431

NOTICE.

The ‘ Proceedings’ for the year are issued in fowr parts, paged consecutively,

: go that the complete reference is now P. Z. 8. 1922, p.... The Distribution

is usually as follows :— ¢

Part I. issued in March.

Perio (8 I e June.

pice LE aerated September.

DNV 5-4; December,

‘Proceedings, 1923, Part I. (pp. 1-180), was published on April 12th,

1923.

The Abstracts of the ‘ Proceedings,’ Nos. 238-240, are

contained in this Part.

The dates of Publication of ‘Proceedings’ 1880-1858 will be found in the

‘ Proceedings’ for 1893, page 436.

The dates of Publication of ‘ Transactions’ 1882-1869 will be found in the —

‘ Proceedings’ for 1913, page 814. oN Vy

\ woe

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