ears se

ts “an SnatPai mintwe

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atta Ape Tbe ete ant mia fhep mae Re Re ttn ie Spe oe areas 8 . . : ra oy 7 - Mee : fo Ny eae . nt be Sh ge ha tesa 4
6 te NST ik 2 WRAL yA EES BOR LN SAD Fm oh Sw ma win gine - ¥ ao ede 23 tn vue iy 4 cotie ipe e ! _ " ae perpen aaycg

o

PROCEEDINGS

OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCIETY

OF LONDON.

1923, pp. 1-481,

witn 12 Puares AND 160 TEx'-FIGURES.

1 é

2 al 7750 |
* 4A ah, : ‘ ; Prk if

PRINTED FOR THE SOCIETY,
SOLD AT ITS HOUSE IN REGENT’S PARK.
LONDON:
MESSRS. LONGMANS, GREEN, AND CO,
PATERNOSTER ROW,

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

178

178

Leg

iV

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

179

180

180

180

479

479

479

479

480

480

480

480

cl

10.

iE

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

1a

15

19

4]

59

85

oF

125

129

13.
14,

15.

Ge

We

18.

iS),

20

Zalbe

vi

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

tv

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.
PI

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

15

JES)

Si)

155 ©

273

1x
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

85

125

180

178
179

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

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

65

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.

C

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

XV

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.

d

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.

xx

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.

PRINTED BY TAYLOR AND FRANCIS,

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XX1

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“ZOOLOGICAL SOCIETY

PROCEEDINGS

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APRIL 1923.

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aS

03

: 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

ee

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

1*

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
93
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
39
99
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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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.

es

an
oes

Ist antenna.

Text-figure 4.

ae

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

a

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.
Oo

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
NS
NP

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.

1

i)
1
t
t

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

36

“IENGIpuvmodky yysr1 Fo gaed ATquqoud “wx ¢ [vxrodueqeadns “g's Sesoutends “bs £ umpnosedoqns “do-s
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othed
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it clearly is

seum (the Yorkshire

Philosophical Society’s Museum at York); here the finely to

CUS9qQM snioydo..zUaoy ,,

c

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

‘pl sdllsy-gxey,

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but

serration,

show

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

4*

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.

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=

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tiarum Fennice, I’. 41.

1884. The Myology of Amiurus catus. Proc. Canad. Inst.
‘Toronto.

1882. Development of the Skull in Lepidosteus. Phil.
Trans. London, vol. elxxiii.

1922. Some Points in the Cranial Development of Teleo-
stomian Vishes. Acta Zool. Bd. 3.

1921. Triassic Fishes trom Spitzbergen. Partl. Vienna.
1922. Ueber zwei Celacanthidee aus den Oberdevon yon
Wildungen. Palzeontologischen Zeits. Bd. iv. Hft. 2/3.
1907. Ueber einige Besonderheiten am Primordialcranium

von Lepidosteus osseus. Anat. Hette, Bd. xxxiul.

1911. Beitrage zur Kenntnis des Kopfes der Wirbeltiere.
1. Die Hntwicklung des Primordialcranium von Lepid-
osteus osseus. Anat. Hefte, Bd. xliv.

1874, Untersuchungen zur vergleichenden Anatomie der
Kiemen- und Kiefer-muskulatur der Fische. 1. Sela-
chier. Zeits. Jena, Bd. vii.

1919. The Structure, Evolution, and Origin of the
Amphibia. The “Orders” Rachitomi and Stereo-
spondyli. Phil. Trans. London, vol. ccix.

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, ,

UI

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

5*

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.

a
N
i
!
'

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).

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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

a
_

is
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,

J

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

7*

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

Cr

aE
= ae

le

Le

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.

8*

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.

A.
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.

Ss

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...

4
99 eoe aed
s Sie 3
BIO 5 i

me ep e OO
— eo — 69

3 a
1

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

K

Text-figure 6.

—S SS Se eee

A

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

bo
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.

a

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
Co

oe
PRE EEO oo
pa

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

bo
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.

21

Numbers
referring to
Notes.

6
iG

17

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,

5
6. This was in a Caucasian ibex,
7

2
3. Both of these cases were birds of prey which had over-
4,

128
8.

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,

ne
© @ @ pa @
@ 3) ie
® g
(S
ve eo eo
@7° ror f
® @ @
@°o o
@ oS.
A. iN 2
k.

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.
QO

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.

AG
j y 24 Pla) \)\ saya
23
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

a
ie
mde

pa teat

hal aarr sta ef

oe
Arh a py )

et
Sepilbns

Saggy te
, 4 Feabe

wives ee

iy wiry

VV NN VY?
Nouv {

yi
lava
SUG ye

v

Felrs pardus.

Felis tigris.
leo.

v
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!
i

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 %, {
4
Trichechus Cystophora \ | Otaria | f Otaria
TOSINEFUS - proboscidea. it calforniana.\ giilesps.

' Halichoerus “
gry phus.

Phocea

Cystophora ‘
vitulina. A eda!

cristata.

iy APS
ss

t RSShy q
AA
: 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.

99

9?

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.

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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.

a
gg
OOD NE

adie ed

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

ne

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.

v.

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.

6

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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

ia

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.

14

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.

15

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).

16
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.

18

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.

1)

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,

22

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. ]

23

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.

&

ch ee

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
ee

11

15

19

41

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.

x

BS
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

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“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,

ZOOLOGICAL SOCIETY OF LONDON,
INSTRUCTIONS TO AUTHORS.

Owine to the cost of printing, the Publication Committee
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ZOOLOGICAL SOCIETY OF LONDON.

Tas Society was founded in 1526 by Sir Sramrorp RaFFiss,
Mr. J. Sasrne, Mr. N. A. Vigors, and other eminent Naturalists,
for the advancement of Zoology and Animal Physiology, and for the
introduction of new and curious subjects of the Animal Kingdom,

and was incorporated by Royal Charter in 1829.

Patron.
HIS MAJESTY THE KING.

COUNCIL.
HIS GRACE THE DUKE OF BEDFORD, K.G., F.R.S., President.
Cou. S. Moncxron Copeman, || Cot. Sir Henry McManon,
M.D., F.R.S. i G-CMLG., K.C.1E:
CHartes Drummonp, KEsa., E. G. B. Meapz-Waxpo, Ese.,
Treasurer. Vice-President.

Aurrep Ezra, Esa., O.B.E.

: P. CHatmers Mircuert, Hse.,

Hues 8. Guapstons, Ese., M.A., C.B.E., M.A., D.8c., LL.D.,
PRAS.E. F.R.S., Secretary.

Tue Rr. Hon. roe Viscount
Grey oF Fatopon, K.G., |
P.C., Vice-President.

Siz Srpney F. Harmer, K.B.E., | Masor Avsurr Paw, O.B.E.
M.A., Sc.D., F.R.S., Vuce- || Lr.-Cot. Sm Daviy Pram,
President. C.M.G., C.I.E., F.R.S.

Pror. James P. Hitt, D-Sc.,
F.R.S., Vice-President.

Wrt1am Huntsman, Esa.

Lr.-Cozr. tHE Lorp ALASTAIR |
Rosurt Innes-Ker, D.S.O. Ricwarp StepHeNs Tayrtor, Esq.

Tue Ear oF Onstow, O.B.E.,
Vice-President.

Tun Lorp QuEBNBOROUGH.

Harotp Joun Hastres
Russet, Esa.

Sir Water Roper Lawrence, || A. Smita Woopwarp, Esa.,
Br, G.C.LE., G.C.V.07 C8: LL.D., F.R.S., Vice-President.

2

The Society consists of Fellows, Imperial Fellows, Honorary,
Foreign, and Corresponding Members, elected according to the
By-Laws. It carries out the objects of its foundation by means
of its collection of living animals, by its Library, and by its
Scientific Publications.

The Office of the Society, Regent’s Park, N.W.8, where all com-
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The Library, under the superintendence of Mr. F. Martin Duncan,
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The Library is closed from Good Friday to Easter Monday, and
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The Meetings for Scientific Business are held in the Meeting
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The Anniversary Meeting is held on the 29th of April, or the
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The Society’s Gardens are open daily from Nine o’clock until
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Mammals and Birds; Mr. Edward G. Boulenger is Curator of
Reptiles and of the Aquarium; Miss L. E. Cheesman, F.E.S.,
is Curator of Insects; Dr. C. F. M. Sonntag is Anatomist; Dr,
N. 8. Lucas, Pathologist; Prof. R. T. Leiper, F.R.S., with the
assistance of a panel of experts, examines and reports on the
Parasites; Prof. G. H. Wooldridge, F.R.C.V.S., is Honorary Con-
sulting Veterinary Surgeon; and Dr. R. W. A. Salmond, O.B.E,,
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or pathological material, or for facilities for work in the Prosectorium
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TERMS FOR THE ADMISSION OF FELLOWS.

Frtitows pay an Admission Fee of £5, and an Annual Contri-
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4

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Any Frriow, having paid all fees due to the Society, is at liberty
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Ladies or Gentlemen wishing to become Fellows of the Society
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P. CHALMERS MITCHELL,

Secretary.
Regent's Park, London, N.W. 8.
June, 1923.
MEETINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON
FOR

SCIENTIFIC BUSINESS.

1923.
TUESDAY, OCTOBER ......... 23.
i NOVEMBER ...... 6 and 20.

The Chair will be taken at half-past Five o'clock precisely.

ZOOLOGICAL SOCIETY OF LONDON.

LIST OF PUBLICATIONS.

‘THE scientific publications of the Zoological Society of London
are of two kinds—“ Proceedings, # published in an octavo
form, and ‘‘ Transactions,” in quarto.

According to the present arrangements, the “ Proceedings”
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tific meetings, but also all the papers read at such meetings
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“ Proceedings, to illustrate the new or otherwise remark-
able species of animals described therein. Amongst such
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The “ Proceedings” for each year are issued in four parts,
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Fellows also have the privilege of subscribing to the
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The following is a complete list of the publications of the

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TRANSACTIONS OF THE ZOOLOGICAL SOCIETY OF LONDON.

Vols. LIV. (out of print).

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xe met a 910)
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99

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eo eee ee eo eee

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XXI.—Part 1. (7 Plates & 12 Text-figs.)

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Since 1916 (Vol. XXI. Part I.) publication of the Society’s
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the high cost of production.

Wj now able to offer for sale, at the reduced price of £30, sets of Vols. V._XVI. inclusive, and

g In consequence of a re-arrangement of the stock of the ‘Transactions,’ the Society is
,

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LISTS OF THE ANIMALS IN THE SOCIETY'S GARDENS.

List of the Vertebrated Animals now or lately Living in the Gardens
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ZOOLOGICAL RECORD.—Vol. 57, containing literature relating
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price £2. Vol. 58, for the year 1921, is being prepared as
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P. CHALMERS MITCHELL,
Secretary.
‘Regent's Park, London, N.W. 8.
June, 1923.

These publications may be obtained at the Socrery’s Orriczu
or through any bookseller.

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
Tmtrod uctiomyy: ahve caul. Weeks ene cee ee ee eteeny aT
OPP OUIS, 2POS DOTHAN hancriscensg oogsoseoerennont a seocesace » Alley
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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.

ae
ASATW ION;
WU My
NARA
x

AA
) 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.

B
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 —
As

Sn
\
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.

G

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.

B
C. Apex of glans penis of Cervus (Sika) hortulorum, from the front.
D
HE

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.

be ibe tiled ne
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suit

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dA a”

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

2

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.

i)
to
oO

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

16

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.

C

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—

bo

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

Or
bo

. 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

y)
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.

i
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.

9
10
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

S

ce

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.

e

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.

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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.

S

F
=== TEMPORAL -
Z.Mi. FASCIA

Hi) y, Z.Nia. \\ \\
Yy thy, > W\\W i)

LEV. LAB.SUP_<¢ Uf 4) AUR.ANT.
g 2 AUR.POST.7"
ICLE:"} £

pagoTip URICLE: )

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)
x

! \ wD
Wey Ved a
MUA We SSy
\ WAS S
\

\

6

ni

Mi

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/

2
>

WY

S|
STN)

1 Wf, 4] 4 LM.A.
/

i
LDN. ADA.) FF: A

WO
IOOy
SN

S.C.G.

B.

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\\
A
\\

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

Se

—

—

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
IN

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 ”

99
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.
—!

A>

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

KS
se

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
(xi) and hypoglossal (xii) from C1 or ©2. There are no separate
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,

oT

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.

The brachial plexus. ©5-D1: lower cervical and first dorsal nerves.
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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. Sonnraa, C. F.—Proc. Zool. Soc. London, 1921, pp. 1-29.
. SPERINO, G.—‘‘ Anatomia del Cimpanzé.” Torino, 1897.

. Symmneton, J.—Proc. Phys. Soc. Hdin. 1890, pp. 297-312.

. Trartt, Dr.—Mem. Wern. Nat. Hist. Soc. Edin. 1817,
pp. 1-49.

. Tyson, E.—“ Orang-outang, sive Homo sylvestris.” London,
1699.

. Vrouik, W.—‘‘ Recherches d’Anatomie Comparée sur le
Chimpanse.” Amsterdam, 1841.

. WALDEYER, W.—Monit. Zool. Ital. No. 4, pp. 73-74.

. Wiper, B.—Boston Jour. Nat. Hist. 1862, pp. 352-382.
. Wyman, J.—Proc. Boston Soc. Nat. Hist. 1856, pp. 274-275.
. ZUCKERKANDL, H.—‘“‘ Das Periphere Geruchsorgane der Siuge-

tiere.” Stuttgart, 1887.

. Kipp, W.—Proc. Zool. Soc. London, 1904, p. 263.

. Reape, W.—‘‘ Savage Africa.” London, 1864.

. Watuis, H. M.—Proc. Zool. Soc. London, 1897, pp. 298-310.
. Esturs, E.—Abh. Phys. Cl. Ges. Wiss. Gottingen, 1881,

Beli
. Biscuorr, T. L. W.—Mitth. K. Zool. Mus. Dresden, 1877,

pp. 251-260.

. Cavanna, G.—Arch. per l Antropologia, 1876, pp. 211-215.

. Sonntaa, C. F.—Proe. Zool. Soc. London, 1922, p. 441.

. Pautia, 8.—Morph. Jahrb. xxvii. (several papers).

. Kerra, A.—‘* Human Embryology and Morphology.”

. Harrmann, R.—Arch. f. Anat. und Physiol. 1872 (several

articles).

. Giacomini, C.—Quoted by Sperino (see no. 47).

. Duvat, M.—Quoted by Sperino (see no. 47).

. Buumenpacs, J. F.—Quoted by Humphry (see no. 26).

. Hermes, O.—Veth. Berl. Gesell. f. Anthrop. 1876, p. 88.

. Jounstong, G. L.—~—Proc. Zool. Soc. London, 18$7, p. 183.

. Doran, A.—-Trans. Linn. Soe. 1897, pp. 371-497.

. Hatrorp, G. B.—“ Not like Man bimanous and biped, not yet
quadrumanous, but cheiropodous.”” Melbourne, 1863.

. GruBer, W.—Arch. f. Anat. 1870, p. 501.

. Herrmann, G.—Journ. de lAnat. et de la Physiol. norm. et
path. 1880, pp. 484, 451, 472.

. Owen, R.—Proc. Zool. Soc. London, 1836, pp. 41-42.

a Proc. Zool. Soe. London, 1846, pp. 2-3.

. Scumipt, M.—‘‘ Die Krankheiten der Affen.” Zool. Clinik.
Berlin, 1870.

. Rotietr, E.—C. R. Ac. Paris, 1891, pp. 1476-1478.

. Mrver, A. B.—Mitth. Zool. Mus. Dresden, 1877, pp. 223-247.

. Pocock, R. I.—Proe. Zool. Soc. London, 1905, p. 169.

PHYSIOLOGY, AND PATHOLOGY OF THE CHIMPANZEE.

Fig.

Fig.

Qn

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.

4

9

PZ. S. 1923, RING, Pl. 1.

a

4 John Bale.Sons & Danielsson, Lt?

ELEPHANT-SEALS OF KERGUELEN LAND.

ae

ere

o

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

BO

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
ke
~~ 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
\¥
WG

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 ;

(c) in both types the motive power used on land or in water is
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,

26

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.

27

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.

SPERM

z

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.

30

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.

31

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,

o4

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 :—

©. Tare Recan, M.A., F.R.S., F.Z.S.

' The Skeleton of Lepidosteus, with Remarks on the Origin
- and Evolution of the Lower Neopterygian Fishes.

39

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
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