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PROCEEDINGS
OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCIETY

OF LONDON.

1921, pp. 447-887,

. with 18 Phares AND 184 TEX'r-FIGURES. .
ie 2s 723lp

PRINTED FOR THE SOCIETY,
SOLD AT ITS HOUSE IN REGENT’S PARK.
LONDON:

MESSRS. LONGMANS, GREEN, AND CO,
PATERNOSTER ROW,

fT Sab
OF THE

COUNCIL AND OFFICERS

OF THE

ZOOLOGICAL

SOCIETY OF LONDON.

LO ale

Patron.
His Masgestry Tue Kive.
COUNCIL.
His Grace Tar Duke or Beprorp, K.G., F.R.S., President.

AurreD H. Cocks, Esq., M.A.

CaarLtes Drummonp, Ksq.,
Treasurer.

ALFRED Hizra, Esq., O.B.H.

THe Ricutr Hon. 'Tur Viscount
Grey, K.G., P.C.

Sir Srpney Ff. Harmer, K.B.E.,
M.A., Se.D., F.R.S., Vice-
President.

Pror. JAMES P. Hitu, D.S8c.,
H.R.S., Vice-President.

WILLIAM HuNntsMAN, Esq.

Mayor THe Lorp ALASTAIR
Roper? Innes-KeEr.

Pror.

G Cove Ge ik Cali:

Ernest W. MacBripe,
D.Sce., F.R.S., Vice-President.
Con. Sir Henry McManon,

E. G. B. Mrapr-Watpo, Esq.,
Vice-President.

P. CHatmers MircHe.., Esq.,
C.B.H., M.A., D.Se., LL.D.,
F.R.S., Secretary.

THe Haru or Onstow, O.B.E.

Masor Arperr Pam, O.B.E.

THe Lorp QUEENBOROUGH.

His Grace THe DvuKE oF
Rurtanp, K.G.

Masor Ricuarp 8. Taytor.

A. ‘Treyor- Batryz, Esq.,
M.A.

Antuony H. WINGFIELD, EsqQ.,
Vice-President.

A. Smrra Woopwarp, Esq.,
LL.D., F.R.S., Vice-Pre-

sident.

PRINCIPAL OFFICERS.

P. Cuatmers Mircuett, C.B.M.,

Secretary.

M.A., D.8e., LL.D., F.B.S.,

R. I. Pocock, F.B.S., F.L.8., Corator of Mammals and
Resident Superintendent of the Gardens.

D. SETH

-Smiru, Curator of Birds and Inspector of Works.

Epwarp G. BouLencer, Curator of Reptiles.
Miss L, EK. Cuzxsman, F.E.S., Curator of Insects.

Dr. C. F. Sonntag, Ch.B

., Anatomist.

Dr. N. 8. Lucas, M.B., Ch.B., Pathologist.
R. J. Orriterp, M.A.. Hon. Parasitologist.
FE. Martin Duncan, F.R.M. Si Librarian.

F. W. Bonn, Aeoouatant
Ny Jal Coxe, Chief Clerk.

LIST OF CONTENTS.

1921, pp. 447--887.

EXHIBITIONS AND NOTICES.

Sir 8S. F. Harmer, K.B.E., F.R.S. Exhibition of, and
remarks upon,a mounted specimen of a young female
TERMMOCCTOS SOHAMLCWS Vi. ae icchsiawiah «23 vias ak isd sb ols oF eMRaLN Sa,

Mr. Tuos. Gerrarp, F.Z.S. Exhibition of, and remarks
upon, photographs of the skeleton of an old male
- Kiva Gorilla (Gorilla gorilla beringeri Matschie) ......

Lord Roruscuitp, F.R.S., F.Z.8. Exhibition of, and
remarks upon, a mounted specimen of a male Kivu
Gorilla (Gorilla gorilla beringert Matschie) ............

Mr. D. Seru-Smiru, F.Z.8. Exhibition of, and remarks
upon, a series of skins and photographs of polyandrous
EES HEEL 153100) A) oA OI GNOME calc tneche nla d dn datin do meee «

Dr. Caas. F. Sonntag, F.Z.S. Exhibition of, and remarks
upon, some abnorinalities in the Carnivora ............

‘Mr. L. T. Hoasey, F.Z.S., and Mr. E. A. Spauu. Exhibi-
tion of, and remarks upon, effect of Pineal Gland
administration on Frog Tadpoles ...............:cceeeee

The Secretary. Report on the Additions to the Society’s
Menagerie during the month of April, 1921............

Miss L. E. Coresman, F.E.S. Exhibition of, and remarks
upon, lantern-slides of the larval stages of some
Hydropaychides i.) Fay ote riess Gs MOUNT NSUa eiteta:t Sua:clores

Major Srantey S. Fiower, O.B.E., F.Z.8. 9 Exhibition
of, and remarks upon, examples of three species of
Higy phat Serbeas erates linseed ces yas edie eee ees

643

644

G44

Major Sranuey 8. rower, O.B.K., F.Z.8. Exhibition of,
and remarks upon, living specimens of Zestudo leithit
FE 00 i M0) Y7 eC RE nee BRIS HB NCR AE ME ayetea oe Arce Ar AS

Dr, P. Coatmers MircHett, C.B.E., F.R.S., F.Z.S. Exhi-
bition of, and remarks upon, photographs of the
Death-mask of a young Gonilla iy Webs seer ee e-ee

Dr. F. M. Cuarman. An account of the distribution of
Bird-life in the Urubamba Valley of Peru ............

Prof. J. P. Hiuu, F.R.S8., V.P.Z.8.- Exhibition of, and
remarks upon, a series of lantern-slides of Marsupial
CM DR YOS i. cate meri tle rere Seems me am arena

The Secretary. Report on Additions to the Society’s
Menagerie during the months of May, June, July,
Amoust, andyseptembers VO 2N cana enmeancone eee

Mr. M. A. C. Hinton. Exhibition of an embryo African
Mle plants eee 2 eee eek ee iat re eche Hastie ae As

My. G. E. Boutencer, F.Z.S. Exhibition of photographs

of a Common Grass Snake swallowing a Viper......... j

Mr. F. Martin Duncan, F.Z.S. Exhibition of Sauna

graph films of Marine Animals ............. Lelio ies laa Ale

Mr. D. Seru-Suira, F.Z.S. Exhibition of photographs of
the Sacred Ibis nesting in the Society’s Gardens......

645

646

883

885

Prof. G. Eiutor Sirs, F.R.S., F.Z.S. An account of the- -

habits of Zarsius, illustrated by lantern-slides.........

Mr. Encar P. Caance, M.A., M.B.O.U. Exhibition of
cinematograph films illustrating the laying-habits of,
the Cuckoo (Cuculus canorus) Sand the life of the
Mioume, © uclcooye same maa rence cere Ris tice shen aa Renee

The Secretary. Report on the Additions to the Society’s
Menagerie during the ronth of October, tee seoueane:

886

Mr. A. S. Le Sovir, C.M.Z.S. Exhibition of sistent ;

illustrating the early life-history of Ornithorhynchus.

Dr. A. Smira Woovwarp, F.R.S., V.P. Z.S. Exhibition ob
a Human Skull and other emia from Broken Hl,

North Rhodesia .......... Pe ee aun DS cvs gisele moretmuntnn te an

887

26.

30.

PAPERS.

. The Geographical Distribution of Orthopterous Insects

in the Caucasus and in Western Asia. By B. P.
PLO astern Aud etn: et pete Meee atc tneis dye ecrna pro anshehe eich moe aa

. The Auditory Bulla and other Cranial Characters in

the Mustelide. By R. I. Pocock, F.R.S., F.Z8.
(Memmi umes MAS Sy Baia toes. sacsecns foetsonedec Genrenals el

. On the Venous System of the Lizard Varanus bengal-

ensis (Daud.). By Goxsinp Sineu Tuapar, M.Sc.
(WexGeireranes Ay) ae AP aN ja luau wart « dete e

Abnormalities in the Common Frog (Lana temporaria).
By J. H. Luoyp, M.Se., F.Z.8. (Text-figure 1.)......

. Comparative Anatomy of the Tongues of the Mam-

malia,—IV. Families 3 and 4. Cebide and Hapalide.
By C. F. Sonnrac, M.D., F.Z.8., Anatomist to the
mociety: (Rexti Heures 002.) ticsicseutiesctbieecuotes ee

. Note on the Skull of Dinotherium gigantewm in the

British Museum. By C. W. Anprews, D.Sc., F.B.S.,
EA Sat ( Verb mommecml an) ne. x00). ee eee ae shidog cubed

. On the External Characters of some Species of Lutrinz

(Otters). By R. I. Pococn, F.R.S., F.Z.8. (Text-
HVC AUN ES ILS AL teh bis «oan at aa ane ee Ree Se

The Comparative Anatomy of the Koala (Phascolarectos
cinereus) and Vulpine Phalanger (TVrichosurus vul-
pecula). By Cuas. F. Sonnrac, M.D., F.Z.S. (Plates
Vie Vp ext etgesy ot Obs) oak jundena same ease. sien Jase

. New Indian Drilid Beetles. By S. Mauuix, F.Z8.

(Rextfie ures: 1S. icici scccescsrdnssarscsevsveresenns rie,

Page

A87

493

497

BAT

579

32. On some Abnormalities in-the Carnivora. By CHARLES
F. Sonntac, M.D., F.Z.S. (Text-figures 64-€5.)......

38. The External Characters of the Koala (Phascolarctos).
By R.1. Pocock, F.RS8., F.Z.5. (Text-figures 22-26.)

34, The Nestling Feathers of the Mallard, with Observa-
tions on the Composition, Origin, ond History of
Feathers. By J. Cossar Hwart, ML. 1D Althel eee Aree
(Plates I.-X.; Text-figures 1-14.) NE 2 ss are aa

35. On some new Genera and Species of Anomodont Rep-
tiles from the Karroo Beds of South Africa. By R.
Broom, E.RS., C.M.Z.S8. (Text-figures 28-45.) .

36. The Cichlid Fishes of Lake Nyassa. By C. Tare Recan,
M.A., F.R.S., F.Z.8., Keeper of Zoology, British
Museum (Na tural History). (Plates I.-VI.; Text-
fiomares = 30.) 00% > traene ofa oor epee ee cee eee eee eee ee

37. Note on the Systematic Position and Distribution of
the Actinian Sagartia lucie. By J. PLAYFAIR
McMurreicn, C.M.Z.8. (Text-figures 1-4.)............

38. The Comparative Anatomy of the Tongues of the
Mammalia.—V. Lemuroidea and Tarsioidea. By
Cuartes F. Sonnvrac, M.D., F.Z.S., Anatomist to
the Society. (Text-figures 66-69.) .......... eee:

39. The Comparative Anatomy of the Tongues of the
Mammalia.—VI. Summary and Classification of the
Tongues of the Primates. By Cuarves F. Sonnvac,
M. D,, E\Z.8., Anatomastito the Sociebyaerseas-se: ne

40. On some new Parasitic Mites. By Sranney Hirst,
F.Z.S. (Text-figures 16-48.).......-...54. sees eee

41. On the External Characters and Classification of the
Mustelide. By R. I. Pocock, F.R.S., F.2Z.8. (Text-
fipumes 2L—O,) oo vaisn steele ae ane stele dale ce/feitelle Angier «eee

42. Megalohyraz Andrews and Titanohyrax, g. n.—A
Revision of the Genera of Hyracoids from the
Fayim, Egypt. By H. Marsumoro. (Text-figures
DG), ese, «creat leteareiaeis cee rele) Se Ula eateatc a eee BSE

\ Page

587

59]

609

647

675

769

803

839

vil

. Page
43. Contributions to the Visceral Anatomy and Myology
of the Marsupialia. By Cuarues F. Sonnrac, M.D.,
F.Z.8., Anatomist to the Society. (Text-figures
ETI a 5 <a Bice «ins a er EAR eR Mee MS in ene te Malate ae 851
Aiphabeticall List-of Combributors: co. v.c/.cesaces. \deacee e- vee eee vill
Hrdlexiok Milwstratwonsss Pot ce$. 2g2. cd igs dees ee ss aeestee sia Xi
Xvll

ALPHA BED C Ate)

CONTRI UM ORs;

With References to the several Articles contributed by each.

(1921, pp. 447-887.)

Page

Anprews, C. W., D.Sc., F.R.S., F.Z.S.

Note on the Skull of Dinotheriwm giganteum in the

British Museum. (Text-figures 1-4.) ..................... 525

BovunEencer, HE. G., F.Z.8.

Exhibition of photographs of a Common Grass Snake

Swrallilowmanio a, Vater aed ceeun in. eee eee eee mien te OO

Broom, R., F.R.S., C.M.Z.8.
On some new Genera and Species of Anomodont
Reptiles from the Karroo Beds of South Africa. (Text-

fioures 28245) i aeatacia sects etel ic eee eet tat ether 647

Cuance, Enear P., M.A., M.B.O.U.

Exhibition of Cinematograph Films illustrating the
laying-habits of the Cuckoo, (Cuculus canorus) and

Ghepiteot the youmeiCwekoOees.,.:.ccqa-steererem ede stec ae 886

CuapmaNn, Dr. Frank M.

An Account of the Distribution of Bird-life in the
Urubamba Valley of Peru ...,. obi IU ACA coa aC in a A'S ig 645

1
CurrsemaNn, Miss L, E., F.E.S.

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

larval stages of some Hydropsychide .... ............6.006 644

Duncan, F. Marin, F.Z.S. (Librarian to the Society).
Exhibition of Cinematograph Films. of Marine
PNT DLS es ae ce cE a 5)... Cove eas BURA Beit, eres ke tee 886

Ewart, J. Cossar, M.D., F.R.S., F.Z.S.

“The Nestling Feathers of the Mallard, with Obser-
vations on the Composition, Origin, and History of
Feathers. (Plates I.-X.; Text-figures 1-14.) 20.0.0... 609

Frower, Major Sranuzy 8., O.B.E., F.Z.8.

Exhibition of, and remarks upon, examples of three
species, of Heyptian:Jerboas ot eis. die tes see eke 645

Exhibition of, and remarks upon, living specimens of
Testudo leith and T. ibera

GERRARD, T'Hos., F.Z.S.
Exhibition of, and remarks upon, photographs of the
skeleton of an old male Kivu Gorilla (Gorilla gorilla |
GEN OLIGIET OMAN HST OW (SR HR NGS 1h oF SRN eer eS Haye SRP eae 643

Harmer, Sir Srpney F., K.B.E., M.A., F.R.S., V.P.Z.S.

Exhibition of, and remarks upon, a mounted specimen
of a young female Rhinoceros sondaicus ............600 ees ees 643

Hint, Prof. J. P., F.B.S., V.P.Z.S...

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

slides;of, Marsupial embryos). asin.) Mav ibid. a te. seapitt 646

Hinton, M. A. C.
Exhibition of an embryo African Elephant ............ 885

Hirst, Sranuey, F.Z.8.

On some new Parasitic Mites. (Text-figures 16-43.). 769
Proc. Zoou, Soc.—1921, b

Page ~
Hoasen, L. T., F.Z.S., and Spaun, H. A.
Exhibition of, and remarks upon, effect of Pineal
Gland administration on Frog Tadpoles ..................... 644
Le Sovir, A. S., C.M.Z.S.
Exhibition of photographs illustrating the early Life-
history Of Ormithonhiymcn iis: Aon etre serene sees jane 887

Luoyp, J. H., M.Sc., F.Z.8.

Abnormalities in the Common Frog. (Text-figure 1.) 493

Matsumoto, H.

Megalohyrax Andrews and Titunohyrax, g.n.—A Re-
vision of the Genera of Hyracoids from the Faytm,
Heypt. (Textsfisures 1-6.) 2/10. cban Gee ee ee 803

Mauuik, 8., F.Z.S.
New Indian Drilid Beetles. (Text-figures 1-5.) ...... 579

McMuraicu, J. Puayrarr, C.M.Z.S.

Note on the Systematic Position and Distribution of
the Actinian Sagartia lucie. (Text-figures 1-4.) ......... 729

MircHet, Dr. P. Coaummers, C.B.E., F.R.S. (Secretary to
the Society).
Report on the Additions to the Society’s Menagerie
during the month of April, jlO21. ios eee. eee 644

Exhibition of, and remarks upon, photographs of the

Weath mask of ayounge, Gorilla (ewe! Meh ee eee as 645

Report on the Additions to the Society’s Menagerie
during the months of May, June, July, August, and
Septembers WIA T 9 i.e asc kh ccs re occa teen 883

Report on the Additions to the Society’s Menagerie
during the month of October, 1921... oii. ieecster eee: 886

Pocock, R. I., F.R.S., F.Z.S. (Curator of Mammals).
The Auditory Bulla and other Cranial Characters in
the Mustelide. (Text-figures 14-18.)........0.....eee.

On the External Characters of some Species of Lutrinz
(Otters), “(lext*ficures, 19-21.) We Seen? ee
The External Characters of the Koala (Phascolarctos).
(eset RES! 2A AOR we Cort see hicica « soma neaaeemnla: Cailee eters te
On the External Characters and Classification of the
Mustelide. (Text-figures 27-39.) 2.0.0.0... ccc cccecec ee ees

Reean, C. Tate, M.A., F.R.S., F.Z.8., Keeper of Zoology,
British Museum (Natural History).

The Cichlid Fishes of Lake Nyassa. (Plates. I.-VI.;

Perches ammeds bens (4) Uy Seeker. Vee aiin's Wh lng Wide sees Mamas ee cadre ee

RoruscuitD, Lionen Water, Lord, D.Sc., F.R.S., Ph.D.,
E.ZS.

Exhibition of, and remarks upon, a mounted speci-

men of a male Kivu Gorilla (Gorilla gorilla beringeri

1 GRTIRCET NE)? SOE SES RNS P30 Br i oy ee aR a

Suir, D. Sern-, F.Z.S. (Curator of Birds).
Exhibition of, and remarks upon, a series of skins
and photographs of polyandrous Birds .....................

Exhibition of photographs of the Sacred Ibis nesting
imei be Socteby Ss Gardemay nies uh). dh, heme Ny A ve alee,

Suira, Prof. G. Exxiot, F.R.S., F.Z.8.
An Account of the habits of Tarsius, illustrated by

LAMEOETI SIO COs ee a ER et, anita bn RI A ee

Sonntac, Cuartes F., M.D., F.Z.S. (Anatomist to the
Society).
Comparative Anatomy of the Tongues of the Mam-
malia.—IV. Families 3 and 4. Cebide and Hapalide.
(EU Sse es Whois 7-0) | nee

Page

473

535

D9]

803

675

886

497

xil

Page
Sonnrac, CHaries F., M.D., F.Z.S. (continued).

The Comparative Anatomy of the Koala (Phascolarctos
cinereus) and Vulpine Phalanger (Zrichosurus vulpecula).
(Plates V...ViL.; Wext-fisures D303. )e saa eee en eens DAT

On some Abnormalities in the Carnivora. (Text-
figures|64,'6D.)%. .csc detec n Se eeenMe nae SU TN ecg 587

Exhibition of, and Remarks upon, some Abnormalities
In ‘thie Carnivora: : sioner tuteremen amen va! auch etn praca iam emg 644

The Comparative Anatomy of the Tongues of the
Mammalia.—V. Lemuroidea and Tarsoidea. (Text-
fires O69. ec: Cais abe steer ie eee ee Ace ale ais tee TAI

The Comparative Anatomy of the Tongues of the
Mammalia.—VI. Summary and Classification of the

Mongues*ot the Primates 0: oe. 2s0o0. 6) eee eee Been (57

Contributions to the Visceral Anatomy and Myology
of the Marsupialia. (Text-figures 70-79.) ............... 851

THAPAR, GosiInp Srinau, M.Se.
On the Venous System of the Lizard Varanus bengal-
dusts (Waud:).\(Wext=tigures! t= 4.) 2: eee eee ete 487

Uvaroy, B. P.
The Geographical Distribution of Orthopterous Insects
in the Caucasus and in Western Asia. (Map.)............ 447

Woopwarp, Dr. A. Smiru, F.R.S., V.P.Z.8.

Exhibition of a Human Skull and other remains from
Broken Hill, North Rhodesia .......... Be Si 30 WI aes 887

INDEX OF ILLUSTRATIONS.

Apyprymnus rufescens, Hig. 77, p. 871.

Ailurus fulgens, Fig. 65, p. 589.

Alouatia caraya, Fig. 49, p. 516.

Amblonyz cinerea, Figs. 19, 21, pp.
540.

Aonyx capensis, Figs. 19, 21, pp.
540,

Aotus felinus, Fig. 46, p.512.

Ateles ater, Fig. 42, p. 504.

grisescens, Figs. 37, 41, pp.
503.

Aulonocara nyass@, Pl. V., p. 675.

Bainia haughtoni, Fig. 38, p. 662.
peavoti, Big. 37, p. 660.

Cacajao calva, Fig. 48, p. 514.

Callicebus moloch, Figs. 37, 45, pp.
510.

Canis bengalensis, Fig. 64, p. 588,

Cebus albifrons, Fig. 40, p. 501.

fatuellus, Bigs. 38, 39, pp. 499,
500.

Charronia flavigula, Figs. 27,
815,

Chiromys, Fig. 68, p. 745.

Chlamydophorus, Fig. 12, p. 635.

Conepatus, Fig. 18, p. 483.

proteus, Figs, 29, 30, 37,
pp. 810, 812, 826, 829.

Cyrtocara annectens, Fig. 30, p. 723.

31, pp.806,

9
ve,

Dasyurus viverrinus, Fig. 77, p. 871.
Proc. Zoo. Soc.—1921.

498, |

Dendrolagus ursinus, Figs. 70, 74, 78,
77, 78, 79, pp. 855, 861, 862, 871,
873, 874.

Dicynodon andrewsi, Fig. 30, p. 651.

curtus, Fig. 33, p. 654,
ictinops, Fig. 35, p. 657.
macrorhynchus, Big. 36, p. 658.
osborni, Fig. 31, p. 652.
sollast, Figs. 28, 29, p. 649.
watsont, Fig. 32, p. 653.
woodwardi, Fig. 34, p. 65d.
Didelphys azare, Fig. 77, p. 871.
marsuptalis, Fig. 77, p. 871.
Dinotherium giganteum, Figs. 1-4,
pp. 527, 529, 530, 532.

Emydopsis longus, Fig. 41, p. 667.
parvus, Fig. 45, p. 670.

- sciuroides, Fig. 42, p. 66h.
trigoniceps, Fig. 42, p. 666.
Emyduranus platyops, Fig. 44, p. 671.
Endothiodon crassus, Fig. 45, p. 673.
Eosimops newtoni, Fig. 39, p. 663.

Galago crassicaudata, Fig. 69, p. 747.
Gale nivalis, Figs. 27, 30, 33, pp. 806,
812, 819.

Grison, Fig. 14, p. 475.

rary Hin, 25, p: 808,
Grisonella sp., Figs. 18, 32, 39, pp. 488,
817, 829.
Gulo gulo, Figs. 14, 16, pp. 475, 479.
C

XIV

Hapale jacchus, Big. 50, p. 518.

Haplochromis argyrosoma, Fig. 25,
p. 714.

atriteniatus, Fig. 23, p. 711.

auritus, Fig. 14, p. 699.

breviceps, Fig, 9, p. 694.

ericotenia, Fig. 17, p. 704.

eucinostomus, Pl. 1V., p. 675.

—— fuscoteniatus, Fig. 12, p. 697.

—-~ lateristriga, Fig. 18, p. 705.

lepturus, Fig. 22, p. 710,

leuciscus, Fig. 26, p. 715.

— longipes, Pl. V., p. 6795.

macrochir, Big. 24, p. 713.

macrorhynchus, Fig. 28, p. 718.

—— macrostoma, Pl. IV., p. 675.

melanonotus, Fig. 21, p. 708.

—-— melanotenia, Fig. 20, p.

micrentodon, Fig. 27,

microstoma, Fig. 10, p. 694.

—— ornatus, Fig. 6, p. 691.

—— placodon, Figs. 1, 5, pp. 685, 700.

—— plagiotenia, Fig. 19, p. 706.

—— polystigma, Pl, I., p. 675.

preorbitalis, P|, ITI., p. 675.

—— seafasciatus, Fig. 7, p. 692.

—— similis, Fig. 8, p. 693.

—-- simulans, Fig. 5, p. 690.

spherodon, Fig. 16, p. 704.

spilorhynchus, P\. V1., p. 675.

strigatus, Fig. 13, p. 698.

tetrastigmata, Fig. 4, p. 685.

urotenia, Fig. 11, p. 696.

wooat, Pl. II., p. 675.

Helictis cveretti, Figs. 14, 15, pp. 475,
ATT

—-— personata, Figs. 29, 35, 38, pp. 810,
828, 828.

Hemigalago demidoffi, Fig. 69, p. 747.

Hydrictis macuticollis, Figs. 19, 20,
pp. 536, 538.

Ictonyx, Fig. 17, p. 481.
—— capensis, Wigs. 29, 34, pp. 810,
821.

Lagothrix infumatus, Figs, 87, 43, 44,
pp. 498, 506, 507.

INDEX OF ILLUSTRATIONS.

Lamellipalpodes annandalei, Figs. 4, 5,

p. 5895.

Lamellipalpus bombayensis, Fig. 3,
p- 982.

nigripenmis, Figs. 1, 2, pp. 580,
581.

Lasiorhinus latifrons, Fig. 23, p. 597.

Lenur catia, Figs. 66-69, pp. 743-745,
747.

— fulvus, Figs. 68, 69, pp. 745, 747.

—- —— rufifrons, Figs. 66, 67, 69,
pp. 748, 744, 747.

—— macaco, Figs. 67-69, pp. 740-745,
747.

mongoz, Figs. 67, 68, pp. 744, 745.

rubriventer, Fig. 67, p. 744.

Leontocebus, Fig. 51, p. 520.

Lethrinops leptodon, Fig. 29, p. 721.

Liponyssus ethiopicus, Fig. 27, p. 785.

biscutatus, Figs. 21, 22, pp. 778,

779.

blanchardi, Figs.

pp. 785, 787, 788.

ellipticus, Fig. 36, p. 7995.

flavus, Figs. 34, 35, pp. 792, 793.

——- granulosus, Fig. 35, p. 798.

—— longimanus, Fig. 37, p. 796.

-—— macedonicus, Figs. 32, 33, pp. 790,
791.

madagascariensis, Figs.
pp. 780, 781.

—— sciurinus, Figs.
786.

sternalis, Figs.
783.

— viator, Fig. 20, p. 776.

Loris gracilis, Fig. 69, p. 747.

Lutra lutra, Figs. 15, 19, 20, pp. 477,
536, 538.

Lyncodon patagonicus, Figs. 380, 39,
pp. 812, 823.

28, 30, 31,

28, 29,

25. ‘26, pp. 722;

Macropus bennetti, Figs. 70,77, pp. 850,
871.
giganteus, Fig. 71, p. 858.

| Marmosa elegans, Fig. 77, p. 871.

Martes foina, Figs. 16, 27, pp. 479, 806.
—— martes, Fig. 16, p. 479.

INDEX OF ILLUSTRATIONS.

Megalohyrax pygmeus, Fig. 1, p. 845.

Meles meles, Figs. 14, 16, 65, pp. 475,
479, 589.

Mellivora ratel, Figs. 14, 17, pp. 475,
481.

Mephitis sp., Figs. 14, 18, pp. 475, 483.

mephitica, Figs. 29, 36, pp. 810,
824.

Metachirus opossum, Figs. 75, 76, 77,
pp. 862, 864, 871.

Mydaus meliceps, Figs. 29, 37, pp. 810,
826.

Mystax ursulus, Fig. 52, p. 522.

Neonyssus intermedius, Figs. 18, 19
p. 772.

Nestling feathers, Figs. 1-14, pp. 612,
614, 616, 619, 620, 622, 624, 629,
637, 638; Pls. I.-X., p. 609.

Nycticebus tardigradus, Fig. 69, p. 747.

’

Onychogale tunata, Fig. 70, p. 853.
Otopharynex selenurus, Fig. 1, p. 679.

Palemydops platysoma, Fig. 40, p. 665.
Paratetranychus trinitatis, Figs. 42, 43,
pp. 801, 802.

Perameles obesula, Figs. 72, 75, '76, 77,

pp. 859, 862, 864, 871.

Perodicticus potto, Fig. 69, p. 747.

Petaurus breviceps, Figs. 75, 77, pp. 862,
871.

—- —— papuanus, Fig. 77, p. 871.

— sciureus, Figs. 73, 75, 77, pp. 860,
862, 871.

Phalanger maculatus,
pp. 595, 600.

—— orientalis, Fig. 77, p. 871.

Philander laniger, Fig.77, p. 871.

Pithecia satanas, Fig. 47, p. 513.

Phascolarctos cinereus, Figs. 28, 25, 26,
58, 54, 56, 58-63, pp. 548, 549, 551,
556. 558, 561, 566, 573, 574, 597,
603, 605; Pls. V., VI., p. 547.

Figs. 22,

: nae
Phascolomys ursinus, Figs. 23, 25, 26,

pp. 597, 608, 605.

4 |

Plestogale nudipes, Fig. 33, p. 819.

Podapolipus aharonii, Fig. 41, p. 800.

Pecilictis, Fig. 17, p, 481.

Peecilogale albinucha, Figs. 30, 32,
pp. 812, 817.

Pseudoehirus peregrinus, P\. V., p. 577;
Figs. 22, 24, pp. 595, 600.

Pseudotropheus tropheops, Figs. 2, 3,
pp. 681, 684.

Putorius putorius, Figs. 15, 27, 38,
pp. 477, 806, $19.

Lana temporaria, Big. 1, p. 494.
Khamphochromis macrophthalmus,
el Vile pone

Rhinonyssoides trouessarti, Fig. 17,
p. 770.
PRhinonyssus nove-guinee, Fig. 16,

p. 769.

Sagartia lucie, Figs. 1-4, pp. 780, 731,
735, 736.

Sarcophilus harvisi, Figs. 70, 77, pp. 855,
871.

Spilogale putorius, Figs. 29, 36, pp. 810,
824.

Tarsius spectrum, Fig. 69, p. 747.

Tarsonemus translucens, Figs. 38-40
pp. 798, 799.

Taxidea, Fig. 18, p. 483.

Tayra barbara, Figs. 16, 28, 31, 38,
pp- 479, 808, 815, 828.

Titenohyrax andrewsi, Fig. 6, p. 848.

—— ultimus, Figs. 2-5, pp. 845-847.

Trichosurus vulpecula, Figs. 22, 24, 26,
55, 57, 58, 61, pp. 550, 552, 556, 566,
595, 600, 605; Pl. V., p. 547.

Tropheus moorti, Fig. 2, p. 681.

Varanus bengalensis, Figs. 1-4, pp. 488,
490-492.

Zoogeographical Map Caucasus and
Western Asia, Fig. 1, p. 454.

INDEX.

1921.—Pages 447-887, -

[New names in clarendon type. Systematic references in italics.
(z.8.1.) indicates additions to the Society’s Menagerie. |

Aerida nasuta, 459. | Arctonyx, 473, 474, 803, 805, 828, 833.
—-— robusta, 458. Arcyptera flavicosta transcaucasia, 463,
——-- turrita turrita, 464, 469. 467, 471.

Acrobates pyginea (4. 8. L.), 884. —— ——- flavicosta, 469.

|
Acrotylus longipes, 457. | —— fusca, 458, 470.
Actinia chrysosplenium, 734, 737, 738. | Ardea cocot (Z. §. L.), 885.
pallida, 738. Astatotilapia, 680.
Aipyprymnus, 854, 878. | caliiptera, 680,
rufescens, 854, 855, 857, 872, 874. | Astur leucosomus (4, 8. L.), 889.

Aiptasia (Heteractis) lucida, 737. Asturinula monogrammica (Z. 8. L.), 884.
Ailurus, 473, 474. Ateles; 502.
—— fulgens, 587, 589. Siig, HOEY
Alouatta (= Mycetes), 515. grisescens, 502, 757.
caraya, 515. -—— melanochir, 505.

Amblonyx, 543, 544, 811, 814, 830. paniscus, 505.
Ameles abjecta, 451. | Aulonocara, gen. n., 726.
Amphisbena alba (2.8. u.), 888. —— nyasse, sp. n., 727.
Anas boseas, 609. Axis porcinus (Zz, 8. L.), 888.
Ancistrodon contortrix (Z. 8. L.), 884.
_—— piscivorus (Z. 8. L.), 884. Bainia, gen. n., 659.
Anterastes serbicus, 457. —— haughtoni, sp. n., 661.
Anthropopithecus troglodytes, 757, 758, | ——— peavoti, sp. n., 659.

7ol. -—— tigriceps (Owen), sp. n., 659,
Aonyx capensis, 535, 537, 541, 542,543, | Balearica pavonina (2.8. u.), 884.

544, 805, 830. | Bettongia penicillata, 851, 865.
Aotus, 511. | Blepharis mendica, 467.
—— felinus, 511. | Bos caffer (4. 8. t.), 883.
Aprosmictus amboinensis (4.8. .), 885. | Boselaphus tragocumelus (4. 8. L.), 88.

Proc. Zoou. Soc.—1921. d

XVlll

Bradypus tridactylus, 564, 872.
Brunnerella mirabilis, 465, 466.
Bunohyrax, 841.

Cacajao, 514.

-— calva, 514.
Callicebus, 509.

brunnea, 511.

---— moloch, 509.
personata, 501.
Callimenus dilatatus, 451.
Canis bengalensis, 587.
thous (2.8. L.), 883.
Cebide, 497, 759.

Cebus albifrons, 500.

~—- apella, 502.
capucinus, SOL. —
— fatuellus, 498, 758, 762.
hypoleucos, 501.

Celes variabilis carbonaria, 463, 467,

471.

variabilis, 469.

Cercopithecide, 759.

Cercopithecus ethiops, 761.

-—— patas, 757.

preusst, 762.

tantalus, 757.

Cervus eldi (4.8. u.), 883.

Chalcophaps stephani (4. 8. L.), 889.

Charora crassivenosa, 466.

Charronia, 474, 476, 485, 539, 807, 811,
814, 816, 817, 832.

flavigula, 478.

Chilotilapia, 679.

rhoadesii, 680.

Chiromyide, 759.

Chiromys madagascariensis, 751, 758,
760.

Chironectes, 85d.

Chlamydophorus, 634.

Chrysochraon brachypterus, 470.

-—— dispar, 448, 469, 470.

Chrysoela chrysosplenium, 738.

Cobus leche (z. 8. u.), 885.

Cenolestes obscurus, 851-882.

Colius affinis (z. 8. u.), 884.

Concpatus, 474, 476, 485, 803, 811, 813,
814, 825, 827, 829, 857

|
|
|
|
|

INDEX.

Conepatus proteus, 484, 809, 825.
Corematodus, 677.

shiranus, 677

Crotalus horridus (2. 8. L.), 884.
Cuculligera maculinervis, 459.
Cuculus canorus, 886.

Cuscus maculatus, 866.
Cynelurus jubatus (2. S L.), 883.
Cynotilapia afra, 684.
Cyrtocara, 722.

—— annectens, sp. n., 728.
moortt, 724.

—— venusta, 722.

Dasyuridg, 851, 858, 869, 877.

| Dasyurus macrurus, 866.

-—— viverrinus,

871, 874, 877.

851, 854, 857, 866,

_ Decticus verrucivorus schugurovi, 469.

——- —— verrucivorus, 455, 470, 471.

| Dendrolagus, 853, 854,-857, 859, 869,

880.

—— bennetti, 851, 862, 863, 867, 869.

dorianus, 852, 853, 860, 869.
-— inustus, 868, 869.

— wrsinus, 851-882.

Derocorys roseipennis lazurescens, 465,
467.

—_— —— roseipennis, 467.

Dicynodon andrewsi, sp. n., 650.

—— curtus, sp. n., 654.

—— ictinops, sp. n., 656.

—— macrorhynchus, sp. n., 657.

-—- osborni, sp. n., 651.

— sollasi, sp. n., 648.

—— watsoni, sp. n., 653,

——- woodwardi, sp. n., 655,

Didelphyide, 851, 854, 858, 877.

Didelphys, 851, 852, 856, 861, 870,
877.

albiventris, 851.

—— azareé, 851, 852, 854, 857, 870.
874.

—— marsupialis, 851, 854, 857, 870,
874,

—— virginiana, 863, 868.

Dinotherium giganteum, 525.

—— nat, 526.

INDEX,

Diprotodon, 606.
Diprotodontia, 858, 868, 878.
Docimodus, 721.

Johnstoni, 722,
Dolichopoda euxina, 459.
Dorcopsis luctuosa, 857.
Drymadusa eurvicercis, 459.
—-— grisea, 465, 466.
konow?, 459, 466.
——- recticauda, 459.
spectabilis, 459.
Duronia, 452.

Fracta fracta, 467.
kalmyka, 467.

LEimpusa pennicornis, 459, 462.

Emydops parvus, sp. n., 669.

Emydopsis, gen. n., 666.

— longus, sp. n., 667.

—— sciuroides, sp. n., 668.

trigoniceps, sp. n., 666.

Emydura nove-guinee (2. 8. u.), 885.

Emyduranus platyops, gen. et
sp. n., 670.

Endothiodon crassus, sp. n., 672.

Enhydris, 473.

Eosimops newtoni, gen. et sp. n.,
663.

Epacromia tergestina, 469.

Eremiaphila genet, 458.

persica, 464, 467.

Hudocimus ruber (z. 8. t.), 885.

Eulabeornis castaneiventris (z. 8. L.),

885.
Hurystomus crassirostris (Z. 8. L.), 885.
Huxenura maguari (Zz. 8. L.), 885.

Felis bengalensis (z. s. u.), 884.
leo (Z. 8. L.), 644, 883.
—— pardus (2.8. .), 884,
—— serval (z. 8. u.), 884.
—- tigris (4. 8. L.), 883.
Fischeria baltica, 467.

—— betica, 451.

Galagide, 759.
Galago, 763.
—— crassicaudata, 750,

X1x

Galago maholi (4. 8, 1.), 887.

Gale, 807, 811-814, 818, 882.

africana, 817.

nivalis, 807, 817.

Galera, 473, 476, 803.

barbara (z .8. u.), 883.

Gampsoclets recticauda, 457.

schelkounikove, 455..

Gazella subgutturosa (7. s.L.), 884.

Gecko vittatus (2. s. t,.), 885.

Gelastorrhinus, 452.

sagitta, 467.

Genionyus, 841, 850.

Geomantis larvoides, 457.

Gomphocerus caucasicus, 458.

maculatus, 467.

—— sibiricus, 448, 458, 470.

variegatus, 453,

Gorilla gorilla, 757, 758, 760, 765.

——- -—— lberingeri, 643.

—— matschiei, 643.

Gratidia bituberculata, 465.

Grison, 475, 474, 476, 803, 807, 811,
820, 827, 835,

furax, 807, 813-816.

Grisonella, 480, 482, 484, 485, 807, 813,
814, 816, 829, 835,

Grisonine, subfam. n., 835.

Gryllodes odicus, 469.

Gryllus domesticus, 448,

Frontalis, 458.

—— hebreus, 458.

lateralis, 451, 462.

—— tartarus obscurus, 458.

—— (Gryllodes) herkennensis, 451,

Gulo, 473, 474, 476, 478, 480, 485, 803,
805, 811, 818, 814, 820, 822, 827,
828, 833.

gulo, 478, 539.

Gulonine, 833.

Gymnopus, 818.

Gypagus papa (4. 8. L.), 644,

Hapale, 517.

jacchus, 517, 758. |
—— penicillata, 519.
—— pygmea, 519.

| Hapalemur, 763, 766, 767.

Hapalide, 517, 759, 766, 767.
Haplochromis, 685.

—— argyrosoma, sp. n., 713.
—— atriteniatus, sp. n., 711.
—— auritus, sp. n., 699.

—— breviceps, sp. n., 694.

—— brevis, 709.

—— ceruleus. 712.
-_—-.chrysonotus, 702.

---— compressiceps, 717.

— dimidiatus, 698.

——- ericotzenia, sp. n., fos.
— eucinostomus, sp. n., 716.
—— fuscoteniatus, sp. n., 696.
—— guentheri, sp. n., 707.
—— holotzenia, sp. n., 697.
intermedius, 701.

—— johnstoni, 691.

—— kirkit, 698.

—— lateristriga, 705.
—--lepturus, sp. n., 709.

—— leuciscus, sp. n., 714.

—_— livingstonit, 688.

—— longipes, sp. n., 712.
—— macrochir, sp. n., 7
—— macrophthalnus, 714.

--— macrorhynchus, sp. n., 717.

—— macrostoma, sp. n., 719.
- -— maculimanus, sp. n., 689.
—— melanonotus, sp. n., 708.
-—-— melanotznia, sp. n., 706.
—-— micrentodon, sp. n., 715.
--— microstoma, sp. n., 695.
—— modestus. 701.

—— nototenia, 709.

—— ornatus, sp. n., 691.

—— placodon, sp. n., 700.
—-— plagiotzenia, sp. n., 7U6.
—— polystigma, sp. n., 688.
—— preorbitalis, sp. n., 717.

—— quadrimaculatus, sp. n., 703.

—— rhoadesiz, 710.

—— rostratus, 718.

—— sexfasciatus, sp. n., 692.
—-—- similis, sp. n., 693.

—— simulans, sp. n., 689.
~-— sphezerodon, sp. n., 703.

—~—-- spllorhynchus, sp. n., 711.

INDEX.

Haplochromis strigatus, Ep: n., 697.
—— subocularis, 690.

—-— tetrastigma, 699.

—— urotenia, sp. n., 695.
—— woodi, sp. n., 702.
Helictidine, 831..

Helictis, 473, 474, 476, 484, 803, 831.
—— everett, 480.

Heliopterye humeralis,.459, 466.
Helioscirtus moseri mosert, 467.
—— —— tichomirovi, 465, 467.
Hemigalago demidoffi, 750.
Hemitilapia, 677.

—— oxyrhynchus, 677.

Mierodula, 452.

HMierofalco. candicans (2. 8. t.), 884.
Homo,.757, 760, 761.

—— rhodesiensis, sp. n., 887.
Hyena crocuta (%. 8. u.), 884.
Hyalorrhipis clausi, 469.
Hydrictis, 544, 813, 830.
Hylobates lar, 757.
Hypsiprymnodon, 607.

Hyrax capensis (2. § -U), 884.

This ethiopica (a. 8. u.), 884.
Letonychine, 830.
Ictonyx, 474, 476, 482, 485.

820-822, 828, 835.
Indris brevicaudatus, 746.
Isophya adelungi, 462, 468, 467.
—— acuminata, 459, 462.
amplipennis, 462.
—— bivittata, 463.

——— paveli, 457.

—— poltoratskyi, 459.
—— pyrenea, 462, 463.
— rodsjankot, 459,
schmidti, 467.

Jaculus jaculus, 645.

—— yaculus (z. 8. 1), 883.
-— orientalis, 645.

— orientalis (z. 8. L.), 883.

Kolonakus, 832.

' Kurdia nesterovt, 459,

(Zorilla), 803, 807, 811, 813, 814,

INDEX.

Lagothrix, 506.

—— humboldti, 509,

—— infumatus, 506.

Lamellipalpodes, gen. n., 579,
584.

—— annandalei, sp. n., 584.

Lamellipalpus, gen. n., 579.

—— nigripennis, 580.

Lasiorhinus latifrons, 598.

Latax, 831.

Lataxine, nom. nov., 830.

Lemur, 742, 788, 766.

—— catta, 748, 758.

—— fulvus, 743.

——- —— rufifrons, 743.

—— macaco, 7438.

—— melanocephala, 7438.

—— mongoz, 743.

—— rubriventer, 743.

—— varius, 743.

Lemuride, 742, 759.

Leontocebus, 520.

—— rosalia, 520.

Lepidosiren, 495.

Leptophyes albevittata, 467.

—— punctatissima, 448, 4538.

Leptoptilus crumeniferus (2. s. t.), 884.

Leptoternis gracilis, 467.

Lepus canopus (z. 8. u.), 885.

Lethrinops, gen. n., 719.

-_— albus, sp. n., 719.

-_— leptodon, sp. n., 721.

—-— lethrinus, 720.

—— macrorhynchus, sp. n., 720.

Limnodynastes peronti, 493.

Liogrylius bimaculatus, 452.

Liponyssus, 773.

—— ethiopicus, sp. u., 774, 783.

—-— arcuatus, 774.

—-— bacoti, 775.

——- biscutatus, sp. n., 774, 779.

—— blanchardi, 774, 787.

—— bursa, 775.

—— carnifex, 774.

—— chiropteralis, nom. noy., 773,
776.

—-— confucianus, nom. nov., 774,
789,

XX1

Liponyssus creightont, 774.
-—— ellipticus, 774, 798.
—— flavus, 774, 791.
granulosus, 774, 794.

—— javensis, 774.

—— lacertinus, 775.

—-— liberiensis, 774.

—— longimanus, 796.

— macedonicus, sp. n., 774, 789.

——- madagascariensis, sp. n., 774,
780.

—— natricis, (73, 775.

—_— nyassex, sp. n., 773, 777.

--— saurarwm, 775.

—— sciurinus, sp. n., 774, 785.

—— serpentium, 773.

—— silviarum, 775.

—-- sternalis, sp. n., 774, 781.

—— viator, sp. n., 773, 775.

Locusta caudata caudata, 455, 470.

—— —— kolenatii, 458.

Loris gracilis, 748.

Loriside, 747, 759.

Intra, 473, 476, 478, 484, 544, 803, 811,
8138, 827, 828, 830.

—— barang, £79.

—— cinerea, 479, 535, 537, 539, 541.

—-— lutra, 474, 535, 5387, 539, 541.

—— maculicollis, 479, 5385, 587, 541.

Lutreola, 818, 832.

Lutrine, 830.

Lyncodon, 474, 482, 539, 804, 814, 822,
827, 836.

Lyncodontine, subfam. n., 886.

Macacus leoninus (z. 8. u.), 883.

Macropodide, 851, 858, 854, 871, 878.

Maeropus, 851, 853, 854, 857, 865, 869,
878.

bennetti, 851, 856, 865, 866, 867,
872, 874.

-——~— giganteus, 851-882.

— giganteus (2. 8.L.), 884.

—— melanops, 851.

-— rufus, 851, 852, 853, 868.

rufus (ZS. L.), 884.

Marmosa, 851, 877.

XX11 INDEX.

Marmosa elegans, 851, 854, 857, 867, |
870, 877.

Martes, 473, 474, 476, 803, 805, 809, |
811-814, 816, 817, 820, 822, 827, |
832.

Joina, 478, 480, 485, 816.

martes, 478, 480, 485, 816.

Martine, 832.

Mazama rufa (2. 8. u.), 885.

Meconema thalassinum, 448, 471.

Mecostethus grossus, 448, 470.

Medecticus assimilis, 458.

Megalixalus fornasinii (2. 8. u.), 884.

Megalohyrax, 839, 841, 844.

eocenus, 840, 841.

minor, 840, 842.

-—— niloticus, 840, 843.

-— paleotherioides, 844.

—— pygmeus, sp. n., 840, 843.

— suillus, 840, 843.

Meles, 473, 474, 476, 478, 480, 481,
482, 485, 808, 805, 809, 811, 813,
814, 822-829, 833.

—— meles, 478, 587, 590.

Meline, 808, 833.

Mellivora, 473, 474, 476, 481, 482, 485,
803, 805, 809, 814, 825, 827, 828,
835.

Mellivorine, 834.

Melogale, 811, 813, 81%, 823, 824, 831.

personata, 805, 809, 811, 822.

Melursus ursinus (%. 8. u.), 644.

Mephitine, 836.

Mephitis, 473, 474, 476, 480, 484, 485,
803, 809, 811, 818, 814, 825, 827,
829, 837.

Metachirus, 851, 853, 877.

——— opossum, 851-882.

Metridium, 730.

senilis, 737.

Microcebus murinus, 746, 766.

Mitua salvini (2.8. u.), 887.

Mixohyrax, 839, 841, 843.

Meritheriwm, 582, 5338.

Muntiacus muntjack (2. s.t.), 883. |

Mustela, 478, 474, 476, 484, 893, 807, |
812, 814, 816, 821, 822, 827,828, ;
832. : |

Mustela africana, 480. -
erminea, 480. -

—-— hibernica, 480.
—— nivalis, 480.
Mustelide, 473, 803.
Musteline, 803, 832
Mydaine, 834.

‘| Mydaus, 473, 474, 484, 803, 809, 811,

814, 826, 834.
Myrmecophila ochracea, 455. :
Mystax, 521.
mystax, 522.

—— ursulus, 521, 758.

Nemobius heydeni, 462.
tartarus, 455, 463.
Neogale, 820.

Neonyssus, gen. n., 771.
intermedius, sp. n., 771.
Nocarodes rimansone, 463.
—— schelkovnikovi, 465.

—-— serricollis, 451, 455, 459.
woronowt, 465.

Numida meleagris (1. 8. u.), 884.
Nyctea scandiaca (4. 8. 1..), 887,
Nycticebus tardigradus, 747.

Octodon, 852.
(Edaieus mlokosiewitch, 458.
(Hdipoda caucasica, 469.
—— schochi, 451.
schochi, 459.
(Edipomidas, 523.
Edipus, 528.
Olynthoscelis annulipes, 459.
—— chabrieri, 457.
——. distincta, 463.
—— fallax, 463.
grisea, 458.
—— griseoaptera, 470.
indistincta, 459, 467.
—— kurda, 459.

—— prasina, 457.
—— punctifrons. 461.
—— satunini, 452, 465, 466.
—— signata, 457, 459.
—— zebra, 459,

INDEX.

Onychogale lunata, 851, 856, 857, 863,
869.

Ornithorhynchus, 887.

Orphania scutata zacharovi, 455.

Otopharyne, 678.

—— auromarginatus, 678.

——— selenurus, sp. n., 679.

Ourebia montana (z.8.L.), 885

Ovis orientalis (4.8. u.), 884.

Oxythespis wagneri, 467.

Paleomastodon, 538.

Palamydops platysoma, gen. et
sp. n., 665,

Pallasiella bolivari, 466.

——- truchmana, 458.

Pamphagus brunnerianus, 459.

—— yersini, 459.

Papio, ‘762.

Paracaloptenus caloptenoides, 457.

Paradrymadusa beckeri, 455.

bocquiliont, 452, 465.

-—— longipes, 465.

—— pastuchovi, 452, 465.

persa, 452, 465.

satunini, 465.

—— sordida, 451. 459, 462, 467.

Paraonyx, 544, 811.

Parapleurus alliaceus, 458, 464, 470.

Paratetranychus, 801.

—— trinitatis, sp. n., 801.

Paratettix meridionalis, 462.

Peragale lagotis, 855, 864.

Perameles, 853, 856, 877.

— lagotis, 871.

nasuta, 870.

odesula, 851-882.

Peranelide, 851, 854, 855, 869, 877.

Perodicticus potto (2. 8. L.), 887.

-— potto, 749, 758.

Petaurus australis, 851, 866.

breviceps, 851, 862, 865, 866, 869,

872.

papuanus, 851, 154, 857,
865, 869, 872.

selureus, 801-874.

Petroqgale penicillata, 852.

xanthopus, 8)2, 853.

Pezotettix giornat, 453.

rugulosa, 459.

Phacocherus ethiopicus (4. 8. u-), 885.

Phalanger, 594, 596, 599, 601, 604,
606, 879.

—— maculatus, 598.

851, 856, 857, 869,

—— orientalis,
872.

Phalangeride, 851, 853, 878.

Phascogale, 866.

Phascolarctide, 879.

Phascolarctos, 591, 594, 596, 598, 599,
601, 602, 694-606, 879.

| —— cinereus, 547, 553, 559, 560, 563,

565, 567-569, 572, 852-880.

Phascolomys, 596, 598, 599, 601, 602,
604, 606, 854, 855, 870.

— — mitchelli, 565, 866, 870.

—— ursinus, 594, 596.

Philander, 851, 877.

—— laniger, 851, 852, 854, 857, 870,
874.

Phonochorion satunini, 459.

Phrynosoma regale (4. 8.u.), 887.

Pithecia, 512.

—— monachus, 514.

—— satanas, 512, 762.

Platycleis bicolor, 471.

—— hurri, 467.

—-— capitata, 464.

—— daghestanica, 445.

—— decticiformis, 457.

—— escalerat, 458.

— — fusca, 455.

| ——- grisea, 467.

—— iljenshii, 471.

—— montana, 469.

—— nigrosignata, 457.
—— persica, 452, 465, 466.
—— roeseli, 476.

—— schereri, 457.

—— squamiptera, 459, 466.
—— truncata, 457,

—_— vittata, 463, 467.
—— weseli, 471.

| Platypierna tibialis, 467.

Plesiogale, 805, 812-814, 820, 852.
—— qudipes, 805, 813,

XXIV

Plesiogale strigidorsa, $18.

Podapolipus, 800.

aharonii, sp. n., 800.

Podisma koenigt. 463.

—— pedestris, 448, 470.

Pecilictis, 473, 474, 476, 482, 485, 813,
814, 820, 821, 835.

Pecilimon bosphoricus, 462, 467.

—— concinnus, 459.

—— distinctus, 462, 463, 467.

—— kutahiensis, 459.

—— similis, 455.

—— syriacus, 459,

—_— tschorochensis, 459.

Pecilimonella derjugini, 459.

Pecilogale, 474, 480, 813, 814, 820.

Poliopsar nemoricola (4%. 8. L.). 885.

Polyprotodontia, 857, 877.

Potos caudivolvulus (4. 8.u.), 823.

Presbytes, 762.

Procyon cancrivorus (4.8. L.), 889.

Protopterus, 495.

~ Pseudochirus, 594, 599, 601, 602, 604,
606, 854, 870.

—— peregrinus, 554, 557, 596, 851,
857, 868, 872, 874.

Pseudotropheus, gen. n., 681.

—— quratus, 693.

-—— novemfasciatus, sp. n., 683.

-_—- tropheops, sp. n., 683.

—— williamsi, 682.

-—— zebra, 682.

Psophus stridulus, 448, 453, 470.

Ptilopus aurantiifrons (4.8. u.), 885.

Putorius, 480, 484, 803, 807, 811, 812,
814, 827, 832.

—— furo, 480, 817.

—-— nigripes, 817.

———— putorius, 480, 817.

Pyrgodera armata, 458, 466.

Pyrgomorpha, 452.

Rana temporaria, 493.

Rattus vociferans (z.8.u.), 884.
Rhabdogale {= Ictonyx], 478.
Rhamphochromis, gen. n., 724.
—— esox, 726.

—— ferox, sp. n., 725.

INDEX,

Rhamphochromis leptosoma,
sp. n., 726.

longiceps, 724.

macrophthalmus, sp. n.,

725,

woodi, sp. n., 725.

Rhinoceros sondaicus, 643.

Rhinonyssoides, gen. n., 770.

——- trouessarti, sp. n., 770.

Rhinonyssus nove-guinee,
sp. n., 769.

Saga cappadocica, 459.

Sagartia lucie, 729.

—— pustulata, 733.

Saghathertum, 843.

Saimiris, 509.

seiurea, 509.

Sarcophilus, 605, 854, 856, 877.

—— harrist, 851, 852, 854, 857, 859,
874.

Saza pedo, 469.

Searturus tetradactylus, 645.

—— tetradactylus (z. 8. L.), 883.

Schistocerca peregrina, 467.

Scintharista brunneri, 469, 467.

Selenidera maculirostris (4.8. u.), 644.

Serranochromis, 685.

—— thumbergit, 685.

Sialia sialis (z. s.u.), 884.

Simia satyrus, 758, 759, 761, 765, 766.

Simiide, 759.

Sphingonotus batteatus, 458.

Sphodromerus, 452.

— ce@losyriensis, 459.

serapis, 467.

Spilogale, 474, 476, 485, 803, 899, 811,
813, 814, 815, 837.

Spiziaus canifrons (z. 8. .), 885.

Stauronotus anatolicus, 458.

brevicollis, 457, 469.

— hauensteini, 451.

— kraussi, 469.

hurda, 459.

Steganura paradisea vorreauxt (4%. 8. t..),
884.

Stenobothrus apricarius, 458, 470.

INDEX.

Stenobothrus fischeri, 458.
hemorrhoidalis, 458, 469. 470.
lineatus, 465.

macrocerus, 458, 464.

— nigromaculatus, 453. 458, 470.

parallelus, 467.

petreus, AGT.
pulvinatus, 458.

—— scalaris, 453, 458, 470.
—— simplex, 458.

ventralis, 453, 470.
viridulus, 448, 470.
wernert sviridenkot, 471.
Stria flanieu, 635.

Struthio camelus (z. 8..), 884.
Symphatangus syndactylus, 759.

Talegallus fuscirostris (Zz. 8. L.), 885.
Tamandua tetradactyla, 856, 874.
Tapirus indicus (4.8. L.), 884.
Tarstide, 759.

Tarsipes, 607.

Tarsius, 886.

spectrum, 753, 788, 760, 767.
Tarsonemide, 197.

Tarsonemus translucens, 797.

Taxidea, 473, 474, 476, 482, 485, 803,

805, 807, 811, 813, 814, 821, 825, |

827, 829, 835.
Taxidiine, 835.

Tayra (= Galera), 476, 480, 485, 808, |

811, 813, $14, 715, 416, 827, 833.
barbara, 478, 807, 811.
Tayrine, subfam. n., 833.
Testudo ibera, 648.

leithii, 645.

Tetrabelodon angustidens, 633.
Tetranychide, 81.

Tettix bipunctata, 470.
depressa, 462.

—— subulata, 470.
Thalpomena ledereri, 459, 462.
persa, 465.

XXV

Thylacinus, 605, &66.
Tilapia, 676.
melanopleura, 677.

—— jossambica, 677.

—— shirana, 677.

—— squamipinnis, 677.

Titanohyrax, gen. n., 839, 841
844.

—— andrewsi, sp. n., 845, 847.

—— paleotherioides, 845, 847.

— schlosseri, nom. n., 844, 847.

ultimus, sp. n., 844, 845.

Tmethis bilobus, 459.

wastes carinatus, 459, 465.

—— cisti, 459.

escherichi, 459.

holizt, 459.

persa, 465.

—-— muricatus, 499.

zattzevi, 463.

Tragelaphus scriptus (z. 8. L.), 884.

Trichosurus, 596, 602, 694-606, 854,
866, 879.

vulpecula, 547, 549, 558, 554, 557,
559, 563, 564, 567, 569, 852-880.

Tridactylus tartarus, 469.

Trinchus schrenki, 466.

Troglophilus escalerai, 459.

Tropidauchen, 452.

Tropidopola cylindrica, 452.

Urticina felina, 737.

Varanus arenarius, 487.
bengalensis, 487.
exanthematicus, 487.
—— griseus, 487.

niloticus, 417.

Vormela, 474, 484, 803, 832.

Zorilla {= Ictonyx], 803.

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LIST OF CONTENTS.
“1921, Panr III. (pp. 447-646).

EXHIBITIONS AND NOTICES.

Page

Sir 8. F. Harmer, K.B.E., F.B.S.; V.P.Z.S. Exhibition of, and remarks upon, a mounted —

specimen of a young female Rhinoceros sondaicus....sesceecssecesecesccsceee owes

Mr. Txos. Grrrarp, F.Z.8. Exhibition of, and remarks upon, photographs of the
skeleton of an old male Kivu Gorilla (Gorilla gorilla beringeri Matschie) ..........

Lord Rotuscaip, F.R.S., F.Z.8. Exhibition of, and remarks upon, a mounted specimen
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BLE D. Serx-Smitu, F.Z.8. Exhibition of, and pos upon, a series of skins aya
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Dr. Cuas. F. Sonnraa, F.Z.S. Exhibition of, and remarks upon, some abnormalities in
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Mr. L. T. Hoesen, F.Z.8., and Mr. E. A. Spavu. Exhibition of, and remarks upon, effect

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The Secretary. Report on the Additions to the Society’s Menagerie during the month of
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Major Stanuny 8. Fuowme, O.B.E., F.Z.8. Exhibition of, and remarks upon, living
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643
643
644
644
644
oa
64 ihe
645

645

Dr. P. Cuaumurs Mitcnnt, C.B.H., F.R.S., F.Z.8. Exhibition of, and roa upon, —

_ photographs of the Death-mask of a young Gorilla ...2........eese cece ees gdoce
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645

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PAPERS.

23. The Geographical Distribution of Orthopterous Insects
in the Caucasus and in Western Asia. By Val Be

Uvarov *.
(With Map.)
[Received October 25, 1920; Read March 8, 1921.]

Introduction.

In 1913 I undertook the systematic study of the Orthopteran
fauna of the Caucasus and neighbouring countries (Asia Minor,
Armenia, Northern Persia), based on the large collections of the
Caucasian Museum in Tiflis and literary sources, as well as upon
my own investigations in Northern Caucasus (in 1911-1914)
and in Transcaucasia (in 1915-1919). This work offered great
difficulties, as our knowledge concerning the systematic and
geographical distribution of the Orthoptera, and those of Western
Asia in particular, is as yet very incomplete. Besides, the war
deprived me of the possibility of referring to Western European
scientific centres and of obtaining from them the necessary
literature and information. The materials which I have had
before me, though not very extensive, give different impressions
concerning the faunas of different districts, and some of the latter
are still awaiting more careful investigation. I think, however,
that it would be of some use for these future investigations if I
gave a short summary of the chief zoogeographical results I have
gained from my studies, incomplete and insufficient though they

* Communicated by S. A. Nreave, F.Z.S,
Proc. Zoou. Soc.— i921, No. XXXI. 31

448 MR. B. P. UVAROV ON THE GEOGRAPHICAL DISTRIBUTION

may be. I hope these results will be of interest to every z0o-
geographist studying the distribution of animals in the southern
parts of the Palearctic region.

Before investigating more minutely the distribution of
Orthoptera in the territory under consideration, it is necessary
to give some information concerning the zoogeographical division
of the Palearctic region based on my studies of this group, for
this division differs in some respects from those of other authors
founded on the distribution of other animal groups.

The subregions into which the Palearctic region may be divided
are four innumber: Boreal, Steppe, Mediterranean, and Eremian.
The chief characteristics of these are as follows :—

The Boreal subregion includes the whole zone of the forests of
Northern, Europe and Asia, but some representatives of its
characteristic Orthopteran fauna.penetrate farther on north-
wards—into the zone of the Arctic tundras which has not its own
Orthopteran fauna. The Orthopteran fauna of the Boreal sub-
region is very poor, both in species and in specimens; the
suborders Mantodea and Phasmodea are entirely absent from it,
and of the Gryllodea we only find the Gryllus domesticus here.
As leading characteristics of Boreal fauna should be regarded :
Chrysochraon dispar, Stenobothrus viridulus, Gomphocerus sibiricus,
Mecostethus grossus, Psophus stridulus, and Podisma pedestris from
Acridiodea, and Leptophyes punctatissima, Meconema thalassinum,
and Olynthoscelis griseoaptera from Locustodea.

The great distance between, our country and the Boreal sub-
region causes the Boreal fauna to be of very little importance in
the composition of our fauna. Nevertheless, in some districts
of the Caucasus, as we shall see later on, the influence of the
Boreal fauna is rather striking: in some mountainous districts
we may find the typical boreal species, which are, at the same
time. absent from the intermediate areas between the Caucasus
and the Boreal subregion. These species with such discontinuous
range of distribution are of great importance to the history of
the fauna of the Caucasian mountains, as they give us a hint con-
-cerning the former contact of this fauna with that of the Boreal
subregion ; later on this contact was interrupted, but the cause
of this interruption is still unknown to us with certainty; we
can only suppose that this immediate connection between the
Orthopteran faunas of the Caucasus and of the Boreal subregion
took place during the Glacial period, and ceased after this period
had given place to a warmer and drier one, when the Boreal
elements of the fauna retreated to the North, leaving a few
relics in the high mountainous districts of the Caucasus.

The Orthopteran fauna of the Steppe subregion is rather rich
and includes many typical forms. The most important character
of this fauna is the presence of a large number of species of the
genus Stenobothrus. The Steppe fauna in Europe shows dis-
tinctly marked affinities with the Siberian, and we should suppose
that its representatives have migrated into Europe from Asia.

OF ORTHOPTERA IN THE CAUGASUS AND WESTERN ASIA, 449

The Steppe fanna penetrates into our country through the
steppes of North-western Caucasus, which belong to the Steppe
subregion, and through Daghestan some of its elements reach
Transcaucasia, as well as the mountainous districts of Caucasus
Minor, Armenia, Kurdistan, and Anatolia as far as the northern
limits of the Hremian subregion. On the other hand, the
Steppe fauna also penetrates into Anatolia through the Balkan
Peninsula. It is evident that the influence of this fauna on the
fauna of our country must be very great, and so it is, as we shall
see later on.

The Mediterranean subregion comprises the north-western
mountainous extremity of Africa (Morocco, Algeria, and Tunis—
but the mountains only and not the plains, which belong to the
Kremian subregion), Spain, the southern coast of France, Italy,
the shores of the Adriatic Sea, the Balkan Peninsula south of
the Balkan Mountains, the islands of the Mediterranean Sea, and
Anatolia. The Mediterranean fauna of Orthoptera is extremely
rich, including about 16 Mantedea, 7 Phasmodea, 162 Acridiodea,
317 Locustodea, and 62 Gryllodea. esides its richness this
fauna is remarkable for a large number of peculiar species: out
of 564 species which are known of it, 424 or 75 per cent. are
endemic. Some large families of Locustodea have their centre
of development and distribution here, for~example Sagide,
Decticide, Bradyporide, and, partly, Phaneropteride.

The more detailed study of the Mediterranean fauna allows us
to conclude that it may be divided into two very distinct groups
of species: Western and Eastern. The first has its centre in
Spain and North-western Africa, from where its representatives
disperse to the East and North-east; while the second flourishes
in the southern part of the Balkan Peninsula and in adjacent
parts of Anatolia, sending its migrants into north-western,
northern, north-eastern (into the Steppe subregion), and eastern
directions. According to this distinction of the faunas we may
divide the Mediterranean subregion into two zoogeographical
provinces—Western or Tyrrhenian and Kastern or Balkano-
Anatolian. The Tyrrhenian fauna is of no importance to us, as
it cannot influence the composition of the fauna of the Caucasus,
but of much greater value is the Balkano-Anatolian fauna. This
fauna is very rich and has many characteristic points: here
we find exceedingly rich Gee us of endemic Sagide, of
apterous Phaneropteridee (Jsophya, Pwcilimon), of the genera
Platycleis, Olynthoscelis, Drymadusa, Dolichopoda; some species
of Stenobothrus and the genera Wocarodes (with six species) and
Oallimenus (with two species) are peculiar to it.

The Balkano-Anatolian fauna occupies the greatest part of the
country which we are now studying. Through Asia Minor,
which belongs entirely to this zoogeographical province, through
Armenia, and along the southern aa aes n shores ot the Black
Sea, the Balkano- ‘Anatolian fauna penetrates into the forest
districts of Transcaucasia, giving place to the Eremian fauna in

31*

450 MR. B. P. UVAROY ON THE GHOGRAPHICAL DISTRIBUTION

the desert plains of Eastern Transcaucasia and in the dry rocky |

districts of Persia; we meet it again on the southern shore of the
. Caspian Sea—in the district of Talysh, which is remarkable for
its wet subtropical climate. Some typical Balkano-Anatolian
faunistic elements also reach the Caucasus from the north,
wandering from their native home along the western and northern
shores of the Black Sea through the adjacent steppes of South
Russia.

To the south of the Mediterranean subregion lies the vast:
Eremian subregion, which includes all the deserts of North
Africa (Sahara, Libyan desert, Egypt), Sinai peninsula, Northern
Arabia, Mesopotamia, Persia, the whole of the Aralo-Caspian
impression; perhaps, also, the great deserts of Chinese Turkestan
and Mongolia, the Orthopteran fauna of which is but little
investigated as yet, belong to it. The Orthopteran fauna of the
Eremian subregion has many peculiarities if compared with the
Mediterranean, and I cannot agree with most of the zoogeo-
graphists * who usually unite them together. In support of my
opinion I give the following table :—

E =
. a S .

| s 3 3 = 2

Ss 2 re) 9 cS:

g a S iz £ 1h

2 iS E SN Gea (oi oe

q a S) 5 Ss Sy
| = ay < 4q o a
|
| Mediterranean fauna.
| |
| Total number of species..................| 16 7 162 | 317 62 564 |
| Number of species which do not pene- |
| trate into the Hremian subregion ... 9 | 7 130 | 299 ol 496 |
bind evmnncShscccets cae select etter eetate eects salami an Mead, 108 | 259 | 4 424,
| Percentage of endemism.................: 56 % | 100 Jp) 67 8/9 | 2% | 66% | 75 Oo |
' Common with the Eremian subregion.| 7 | 0 2 leds 12 69 |

[Fig RO STOMRY hs ve Gomtece Crea hae mace eRney Ace 9 105 Be) \ ae) 206
IMIS TY GLETANI GC Srtess step Wyataen GAL ae en rma eee ean (aes 9 99 24 | 16 190
| Percentage of endemism ........ ......| 79 %/o | 100%) 72% | 63%) | 51% | 70%

It is evident from this table that the affinity between the
Eremian and the Mediterranean faunas is restricted to 69 species
only. Out of these we must, however, not take into consider-
ation 48 very widely distributed species (such as Varatettix
meridionalis, Acrotylus insubricus, Conocephalus nitidulus, etc.) and
10 species with great power of flying, which may have wandered
from one subregion to another in recent times ; there are, there-
fore, only 11 species common to the EKremian and Mediterranean

* W. L. Sclater was the first who recognised the difference between the Eremian
and his “Europasian” (Boreal+Steppe+ Mediterranean) subregions, but he

included in it the whole of Northern Africa, the western part of which (Morocco,
Algeria, and Tunis) must be united with the Mediterranean subregion.

————

OF ORTHOPYTERA IN THE CAUCASUS AND WESTERN ASIA, Abd]

faunas, 7. e. but 4 per cent. of the second and less than 2 per cent.
-of the first of these. These eleven species are as follows:
Fischeria beetica, Ameles abjecta, Stauronotus hauensteini, Wdipoda
schochi, Nocarodes serricollis, Callimenus dilatatus, Paradrymadusa
sordida, Olynthocelis punctifrons, Isophya triangularis, Gryllus
(Gryllodes) kerkennensis, and Gr. lateralis. Only two ot them (both
species of Gryllus) may be considered as having originated in the
Kremian subregion, and they are both to be found in the
Mediterranean subregion (in Spain, in Transcaucasia) in prox~-
imity to its southern boundaries and on spots with clearly defined
desert soil and vegetation ; we have the right to believe them to
be comparatively recent invaders from the deserts of the Eremian
subregion. ‘Theremaining nine species are of Balkano-Anatolian
origin and do not penetrate deeply into the Eremian subregion,
being restricted to its northern mountainous parts with mixed
fauna. The relationship of the Mediterranean and of the
Eremian faunas is, consequently, practically absent. On the
contrary, the same table shows us that the difference between
them is a very striking one; 496 species (out of the whole
number 564) of Mediterranean Orthoptera do not reach the
Eremian subregion, and 206 Eremian species (out of 275) do not
go through the northern boundary of this subregion into the
Mediterranean. ‘This difference is not numerical only; t the family
Orthoderide is peculiar to the Eremian subregion*, where
there are 35 species belonging to it; the family Pamphagide is
represented in the Mediterranean subregion by 52 species, of
which only five penetrate into the neighbouring parts of the
Eremian subregion; the family Phaneropteride has move than
80 Mediterranean representatives, and only three of them are to
be found among the Eremian fauna; two families of Locustodea
Ephippigeridz (85 species) and Meconematide (4 species)—and
three of Gryllodea—Gryllomorphide (7 species), Myrmecophilide
(4 species), and Mogisoplistide (6 species), which are very im-
portant in characterising the Mediterranean fauna, do not extend
into the Eremian subregion at all. The generic and specific
differences between these two faunas are yet more considerable,
but I shall not go into details here, as I suppose the above
mentioned facts are sufficient to support my statement that the,
Eremian subregion is of the same zoogeographical value as the
Mediterranean ‘.

There are only two provinces of the large Eremian subregion
which are particularly interesting to us: the Ivanian and the

* With but one exception—Geomantis larvoides—which is Mediterranean
endemic.

+ I even suppose that, when studying the distribution of Orthoptera, we are right
in considering the Eremian subregion of the same value as the whole Palearctic
region: this problem i is, however, too great a one to be discussed here, and I hope
to return to it at some other time ; I am supported in my supposition by the state-
ments of Mr. A. Birula, who, after his studies of the distribution of scorpions,
made an Africano- Asiatic region nearly with the same limits as my Eremian
subregion (see A. A. Bialynicki-Birula, Ayvachnoidea Arthrogastra Caucasica, Pars I.
Scorpiones.—Mémoires du Musée du Caucase, sér, A, N. 5, 1917).

452 MR. B. P. UVAROV ON THE GEOGRAPHICAL DISTRIBUTION

Turanian. The first of these has some peculiarities in its fauna
which are of the greatest interest and value: there are among
the rather numerous. endemics of the Iranian fauna some very
ancient forms (Parudrymadusa bocquilloni, P. persa, P. pastuchovt,
Tropidauchen, Platycleis persica, Olynthoscelis satunini, etc.),
which belong to the groups characteristic of the Balkano-
Anatolian province of the Mediterranean subregion. ‘The presence
of these species in the Iranian fauna allows us to conclude that
this fauna was in some ancient time in close connection with the
Balkano-Anatolian fauna, but afterwards -(from the beginning
of the dry climatic period in Iranian table-lands) this connection
was interrupted, and the further development of the Balkano-
Anatolian and of the Iranian fauna went in different directions :
the remnants of the ‘“‘ Ancient Mediterranean” fauna in Iran
partially died out, partially adapted themselves to the new
conditions of life (the ‘“‘ desert” coloration of Paradrymadusa
bocquilloni and P. persa, etc.); and the recent faunas of the
Iranian and the Balkano-Anatolian provinces, being of the same
origin, are entirely different and belong to the different subregions.

The Iranian fauna occupies a rather large part of Transcaucasia
reaching along the western shore of the Caspian Sea as far as the
neighbourhood of Petirovsk, as we shall see later on.

The Turanian province of the Eremian subregion comprises:
Transcaspia and the southern steppes of Kir ghizes and Kalmyks
along the northern and north-western shores of the Caspian
Sea; its Orthopteran fauna is a rather recent derivate of the
Tvanian which has migrated in northern and north-western
directions, invading the parts of land from which the Caspian
Sea has recently withdrawn. West of the Caspian Sea (in the:
Ciscaucasia) the Turanian fauna spreads southwards, coming im
contact with direct Iranian migrants somewhere near Petrovsk.

To the south of the Eremian subregion lies the Indo-Ethiopian
region, the fauna of which is of some importance for the com-
position of the fauna of the country we are studying now, where
we may find a rather large number of species of undoubtedly
Indo-Ethiopian origin, for example: genera Gelastorrhinus,
Hierodula, Duronia, Pyrgomorpha, Sphodromerus, and species—
Tropidopola cylindrica, Liogryllus bimaculatus, ete. All these
Indo-Hthiopian elements came into the country under consider-
ation through the Eremian subregion, of which fauna they are
very characteristic.

The Zoogeographical Divisions of the Caucasus and
neighbouring countries.

Before continuing the study of zoogeographical districts into:
which the country in question may be divided, I ought to point out
that all my conclusions are based exclusively on the study of the
geographical distribution of Orthoptera, though I have also taken
into consideration the conclusions drawn by the late K. A.
Satunin from his study of Caucasian mammals; by Nikolsky—

OF ORTHOPTERA IN THE CAUCASUS AND WESTERN ASIA. 453

reptiles and amphibians; Birula (/. c.)—scorpions ; as well as by
other zoologists.

A full list of the Orthoptera, which are known to me from the
Caucasus and the neighbouring countries, will be given by me
elsewhere.

1. South Russian Steppe district (R.M.)*.

The Orthopteran fauna of the open grassy steppes lying to the
east of Azoy Sea has not yet been sutliciently investigated, and
we know but 69 species (3 M7 + 32 A 4+ 27L+4+7G) from it.
Notwithstanding, it is evident that this fauna cannot be
separated from that of the steppes north of the said sea and
of the Black Sea—the steppes of Southern Russia, and it has
nothing to do with the fauna of the Caucasus proper. In fact,
there is only one species in this fauna which is unknown from the
South Russian steppes; it is Pezotetiix giornai, a Mediterranean
species which I think has quite recently penetrated here from
the neighbouring Novorossiisk district, and is only restricted to
the south-western part of the Azov steppes. We are right,
therefore, in regarding the latter as simply being the southern
part of the vast South Russian Steppe district. The boundaries
of this district, as far as they are lying within the limits of
country we are interested in, are not quite defined yet; the
southern boundary of it coincides with the northern mit of
the forests growing on northern slopes of the Caucasus; its
eastern limit is not so sharply marked and is dependent on the
westward progression of the Avalo-Caspian (Turanian) flora and
fauna along the Valley of Manytsh (see below).

2. The Kuban-Terek district (K.-T.).

As I have had the opportunity of studying this district for a
rather long time (1911-1914, 7.e. four years), its fauna is well
known to me~t. The number of species known from this district
is77 (2M 4-33 A+33 L+9G) and may be regarded as being very
nearly exact. The bulk of this fauna—68 species—is common to
it and to the foregoing district, which leads us to the conclusion
thatthe Kuban-Terek district ought to be regarded as belonging
to the Steppe subregion. The distinction between the South-
Russian fauna and the fauna of this district is based on 13 species.
Out of these seven are of well-defined boreal origin: Stenobothrus
nigromaculatus, St. ventralis, St. scalaris, Gomphocerus variegatus,
Psophus stridulus, Leptophyes punctatissima, and Olynthoscelis
griseoaptera: they form, no doubt, a rearguard of the relic group
of boreal species which retreated from South-Russian steppes,

* The letters after the name of each district are the initials of their latin names
as adopted in my zoogeographical map.

+ M=Mantodea, P=Phasmodea, A=Acridiodea, L=Locustodea, G=Gryllodea.

+ See my publication concerning the Orthopteran fauna of the province Stavropol
(Bull. du Musée du Caucase, 1915),

454 MR. B. P. UVAROV ON THE GEOGRAPHICAL DISTRIBUTION

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OF ORTHOPTERA IN THE CAUCASUS AND WESTERN ASIA. A455

after the end of the Ice age, southwards into the mountains of
the Caucasus (see p. 448). Of the remaining six species, four
are of Mediterranean (resp. Balkano-Anatolian) origin; these
are: Pecilimon similis, Paradrymadusa beckeri (peculiar to the
Kuban-Terek district, but belonging to a Balkano-Anatolian
genus), Platycleis fusca, and Myrmecophila ochracea, which are to
be regarded as immigrants from Transcaucasia. The ways of this
migration, doubtless, do not lead across the chain of the Great
Caucasus, but I think they go on the west along the shores of
the Black Sea, and on the east through the Somkheto-Kakhetian
and Daghestan districts (see below). The remaining two species
are: firstly, Ganypsocleis schelkovnikove, which has just been
described and the zoogeographical physiognomy of which is as
yet uncertain; secondly, Vemobius tartarus, which has recently
penetrated into the Kuban-Yerek district from the neighbouring
deserts of the Caspian Ciscaucasia.

Thus the Orthopteran fauna of the Kuban-Terek district
may be characterised as derived from the South-Russian Steppe
fauna, with well-marked indicaticns of southern influences—from
Caucasus (forms of boreal origin) and from Anatolia (Baikano-
Anatolian species).

The Kuban-Terek district comprises all the northern slopes
of the Caucasian mountains, including the adjacent hilly country
with insular forests; its northern boundary coincides with the
northern limit of these forests on the west and of the grass steppe
on the east; while the southern is formed by the upper lmnit
of the forests on the main chain of the Caucasus.

3. The Daghestan district (D.).

The fauna of this interesting district has as yet been very little
investigated: the whole number of species known from it is
only 43 (2M +23 A+12L+6G), which is, certainly, not
more than one third of the real number. Notwithstanding,
a eareful analysis of this fauna allows us to draw some very
interesting conclusions as to 1ts composition and origin.

The Daghestanian Orthopteran fauna is in direct contact with
the faunas of four districts: Kuban-Terek, Somkheto-Kakhetian,
Caspian Transcaucasia, and the Eastern Caucasus. But we find
the closest resemblances between our fauna in the two first named
districts, and as they belong to distinct subregions (Steppe and
Mediterranean respectively), it is an interesting problem to be
solved—in which of them the Daghestan should be included.

Out of 39 Daghestanian Orthoptera 34 are common to
Daghestan and to the Kuban-Terek district, and only five are
distinct, as follows: Vocarodes serricollis, Orphania scutata
zacharovi, Locusta caudata caudata, Decticus verrucivorus
verrucivorus, and Platycleis daghestanica. The last named species
is peculiar to the Daghestan and of no interest to us; Vocarodes
serricollis is not to be considered as being characteristic of

456 MR. B. P. UVAROV ON THE GEOGRAPHICAL DISTRIBUTION

Daghestan, since it is not distributed all over this district but
confined to its eastern parts; finally, Locusta caudata caudata,
Decticus verrucivorus verrucivorus, and Orphania seutata zacharovi
are the typical mountain forms and, doubtless, came to Daghestan
from the adjacent mountainous district of Eastern Caucasus. It
is evident, therefore, that there exists but very little difference
between the Daghestanian fauna and that of the Kuban-Terek
district, and that this difference is of a recent date and of an
accidental origin. Onthe contrary, the resemblances between them
are far deeper, for nearly all the chief characteristic steppe forms
(such es Arcyptera flavicosta flavicosta, Celes variabilis variabilis,
Stauronotus brevicollis) range into Daghestan, but not farther
southwards where (in the Somkheto-Kakhetian district) they are
either entirely absent or replaced by other subspecies. ‘Thus, the
steppe Arcyptera flavicosta flavicosta gives place in the Somkheto-
Kakhetian district to the distinct race 4. flavicosta transcaucasica ;
Celes variabilis variabilis is represented there by the subspecies
C. variabilis carbonaria and so on. But the most striking
difference of the Daghestanian fauna from the Somkheto-
Kakhetian (resp. Balkano-Anatolian and even Mediterranean) is
in the negative features of the first: the numerous non-flying
Phaneropteride, which are very characteristic of the Somkheto-
Kakhetian fauna, are strange to the Daghestan, where only three
of them exist: one Orphania, one Pecilimon, and Leptophyes
albovitiata — all three not characteristic of the Somkheto -
Kakhetian district ; the numerous endemies of the latter do not
range into Daghestan at all.

All the above-mentioned facts lead us to the conclusion that
the recent Orthopteran fauna of the Daghestan is in more
intimate relation to the fauna of the Kuban-Terek distvict than to
that of the Somkheto-Kakhetian. I think, therefore, it should
be right to regard Daghestan as an independent zoogeographical
district of the Steppe subregion, characterised by the purely steppe
Orthopteran fauna with but slight admixture of mountainous
forms and of endemicsas well as of some ‘‘ancient-Mediterranean”’
species {see p. 452), like WVocarodes serricollis, or an undescribed
species of Paradrymadusa, known to me from Daghestan only in
females. I think that further investieations of this interesting
district may clear up some details concerning the composition and
origin of its fauna but will not change the views expressed above.

Turning to the establishment of the boundaries of this district,
we can only definitely state the north-western and south-western
ones, which coincide with the lower limits of the alpine district
of the Hastern Caucasus. As for the eastern boundary of the
Daghestan it should be presumed to go along the extreme eastern
chains nearly parallel to the Caspian Sea shore, leaving a narrow
strip along the shore itself bearing quite a different Iranian
fauna of the district of Caspian Transcaucasia. The most obscure
are the boundaries between Daghestan and the districts of
Kuban-Terek and of Somkheto-Kakhetia.

ira
‘

OF ORTHOPTERA IN THE CAUCASUS AND WESTERN ASIA. 45

4, The Western Anatolian district (A.M.).

There are known to us from this district, little investigated
though it is, as many as 103 forms of Orthoptera (7M + 387A +
48 L.+ 11G), which gives evidence that its fauna is a very rich
one. As for the composition of this fauna, it may be regarded
as the purest expression of the Balkano-Anatolian fauna, which
has here its original home.

The most characteristic families of Locustodea for this fauna
are the Decticide and the Phaneropteride : here we find 23 species
belonging to the first named family, and 19 to the second.
Among the Phaneropteridx the flightless species are 15 in
number; 16 species of Decticide are also flightless. It is not
surprising, therefore, that we find many endemies in this district :
13 species are peculiar to it. Among these endemics there are
no fewer than 6 species of Pecilimon, all belonging to the group
with non-denticulate cerci, which group presents one of the most
characteristic features of the Balkano-Anatolian fauna having
its centre of origin and distribution in this and in the neigh-
bouring Armenian district. One species of sophya (J. paveli) i 1s
also peculiar to Western Anatolia. Of Decticida two species of
Platycleis (P. iruncata and P. scherert), two Olynthoscelis (Ol.
signata and Ol. prasina), Drymadusa spectabilis, and Gampsocleis
recticauda are also Western Anatolian endemics. Considering
all these endemiecs, 1t 1s noticeable that their specific features are
very well marked anduer y constant, which gives us the evidence
that these species are ancient and undoubtedly autochthonous
forms. Thus we come to the conelusion that the Locustodean
fauna of the Western Anatolia bears some peculiar features and
is of great age. The composition of the other suborders of this
fauna is of a rather mixed and indeterminate character.

As for the more recent elements of the Western Anatolian
Orthopteran fauna, we may distinguish amongst them the forms
of the Steppe fauna (Stenobothrus spp., Stauronotus brevicollis,
ete.) which came here through the Balkan peninsula, and, what
is more interesting, some species characteristic for the Western
Mediterranean (Tyrrhenian) province, which are six in number,
as follows: Geomantis larvoides, Acrotylus longipes, Paracaloptenus
caloptenoides, Platycleis nigrosignata, Olynthoscelis chabrieri, and
Anterastes serbicus. All of them, except Aciotylus longipes, are
wingless and doubtless very ancient in their origin; the careful
study of their distribution shows that it is discontinuous, which
allows us to think that a connection between the Tyrrhenian and
Balkano-Anatolian has been ancient also and ceased long ago.

The limits of this district are not yet sufficiently known to us.
It is certain, however, that the north-western limit does not
coincide with the recent natural limit of the Anatolia—with the
Bosphorus and the Sea of Marmora, but it is to be looked for
somewhere in the Balkanian peninsula. The Western Anatolian
district occupies, conclusively, the more southern part of the last

458 MR. B. P. UVAROV ON THE GEOGRAPHICAL DISTRIBUTION

named peninsula, the western part of Anatolia from the Mediter-
ranean Sea to the western borders of the interior Anatolian
plateau, extending along the southern shore of the Black Sea
as far eastwards as the neighbourhood of Trebizond.

5. The Armenian district (At.).

The whole number of species known from this district is larger
than that of any other, being 134 (8 M + 62 A+ 54 L+ 10G),
which indicates the exceptional richness of its fauna.

The analysis of this fauna shows its affinity with the fauna of
the preceding district, since 62 species are in common with the
latter; it is of interest that many species are peculiar to both
these districts, being unknown beyond their limits. This affinity
is certainly due to the fact that they both belong to the same
(Balkano-Anatolian) province. Far more interesting is the
difference between them: out of 134 Armenian Orthoptera no
fewer than 74 do not penetrate into Western Anatolia. This
group is composed of very different elements. First of all we can
distinguish in it an admixture of the more northern (steppe and
boreal) forms, which are 12 in number, as follows:

Parapleurus alliaceus. Stenobothrus pulvinatus.
Stenobothrus fischeri. . scalaris.
e nigromaculatus. Gomphocerus sibiricus caucasicus.
a apricarius. Arcyptera fusca.
bd macrocerus, Locusta caudata kolenatii.
ie hemorrhoidalis, Olynthoscelis grisea.

The absence of these forms from Western Anatolia may indicate
that they did not come to Armenia through the Balkanian
peninsula but across the Caucasian isthmus. Some of them, for
example Arcyptera and Gomphocerus, ave absent from Western
Anatolia simply because in this latter district there are no places
(high mountains) suitable for their habitation. The same cause
explains to us why the following four alpine species: Orphania
scutata zacharovi, Psorodonotus brunnert, Ps. fieberi, and Ps.
specsularis, do not penetrate into Western Anatolia.

A very characteristic group of the Armenian Orthoptera is
formed by the 13 species belonging to the true Eremian fauna.
Their list is as follows:

Eremiaphila genei. Sphingonotus balteatus.
Acrida robusta. Sphodromerus ccelosyriensis.
Stenobothrus simplex. Platycleis escalerai.
Stauronotus anatolicus. Medecticus assimilis.
Pallasiella truchmana. Gryllus tartarus obscurus.
Pyrgodera armata. >» hebraus.

(Edaleus mlokosiewitchi.

Since all these species are good flyers it is evident that they
came into Armenia in recent times from the neighbouring
deserts of Persia and Mesopotamia; this view is supported by

:
:

OF ORTHOPTERA IN THE CAUCASUS AND WESTERN ASIA. 459

the fact that most of them are restricted to the eastern parts of
the Armenian district.

But the majority in the group of Armenian Orthoptera which
do not reach Western Anatolia belong to species peculiar to

Armenia (or to both Armenia and Syrian Anatolia). They are
23 in number, as follows :

Stauronotus hauensteini kurda. Isophya rodsjankoi.
*Cuculligera maculinervis. Lone poltoratsky1.
*Pamphagus yersini. Saga cappadocica.

% is brunnerianus. *Drymadusa curvicercis.
*Kunothrotes derjugini. * 33 recticauda.
Peecilimonella armeniaca. : % re konowi.
Pecilimon tschorochensis. *Olynthoscelis annulipes.
rs kutahiensis. * 3 signata.
3 syriacus. * ye zebra.
a concinnus. on kurda.
*Kurdia nesterovi. *Troglophilus escalerai.

*Phonochorion satunini.

No fewer than one half of them should be regarded as the relics
of the ‘‘ancient Mediterranean” fauna (these are marked with an
asterisk), to which also belong the following 13 species whose
area of distribution extends also beyond Armenian limits, as they
are to be found in the neighbouring districts of Aderbeidzhan,
Caspian Transcaucasia, and Somkheto-Kakhetia :

(Edipoda schochi schochi. Tmethis escherichi.
Thalpomena ledereri. py) lnlliivat
Heliopteryx humeratis. Nocarodes serricollis.
Tmethis saussurel. Pezotettix rugulosa.
» carinatus. Platycleis squamiptera.
> CiSti. Paradrymadusa sordida.

» bilobus.

We find, thus, in Armenia 27 relics of the ‘‘ ancient Mediter-
ranean ” fauna which find here their western limit of distribution
and do not reach Western Anatolia.

The remaining 7 species of Armenian Orthoptera which do not
range farther westwards (Zmpusa pennicornis, Acrida nasuta,
Isophya triangularis, [. acuminata, Olynthoscelis indistincta,
Dolichopoda euxina, and Gryllus frontalis) are partly of indeter-
minate zoogeographical value, or their absence trom Western
Anatolia may be explained as a result of insufficient investi-
gations.

Summarizing the results of our analysis of the Armenian fauna
we may conclude that it is a Balkano-Anatolian fauna in its chief
characters but well distinguished from it by (1) the well expressed
influence of the Eremian fauna, (2) the great number of endemics
and relics of the “ancient Mediterranean ” fauna, and (3) some ad-
mixture of boreal and steppe forms.

1 think the frontiers of the Armenian district should be drawn
in the following manner.

460 MR. B. P. UVAROV ON THE GEOGRAPHICAL DISTRIBUTION

The northern boundary coincides with the chain of Pontus,
coming on the east very near to Batoum and embracing a narrow
strip of the Adzharian chain; from here it runs southwards along
the Arsian chain as far as Arax, where the contact of Armenian
and Aderbaidzhan fauna takes place; farther on the boundary
turns eastwards along the chain of Aghridagh as far as Ararat,
from where it goes southwards along the watershed of the basins
of Urmiah and Tigris. The western boundary is presented by
the margin of the central plateau of Anatolia, and the southern
one goes in its western part along the Cilician Taurus and
Antitaurus, not yet being satisfactorily known farther eastwards
owing to the lack of investigation. I suppose it coincides with
the Armenian Taurus.

As the territory of this district is far more extensive than
that of any other, it causes us to suppose that it should be divided
into two or more separate districts. Some modifications of the
Orthopteran fauna of different parts of Armenia support this
conclusion, but I cannot offer any satisfactory division, as our
knowledge of the fauna of southern and south-eastern parts of
Armenia is extremely limited. Further investigations of this
district should be, therefore, of the greatest zoogeographical
interest.

6. The Syrian Anatolia (A.8.)

This district belongs to those which have been less investigated,
as is evident from the small number of species known of it, this
being only 106 (9M+50 A+38 L+9G), though its southern
position and dry climate offer the best conditions for the develop-
ment of the richest Orthopteran fauna.

As for the composition of its fauna, it may be regarded as very
closely related to the fauna of Syria proper, all differences being
of an accidental nature. On the contrary, the difference existing
between this fauna and that of Armenia is rather well defined in
the lack of the most characteristic Armenian endemics and in the
evident influence of the Hremian fauna which isa typical feature
of the fauna of Syria.

The boundary between this district and the Armenian one
coincides with the southern limit of the latter district following
the chain of the Cilician Taurus; along the Euphrat valley this
district penetrates into Armenia, as is to be seen on the map.
All the other boundaries of this district lie beyond the limits of
the country which we are studying now and are entirely unknown

as yet.

7. The Pontian district (P.).

The fauna of this district includes 59 species of Orthoptera
(1M+27 A+20 L4+11G); having been well investigated it
cannot be considered very rich.

The analysis of the Pontian fauna shows us that it is very
closely related to the fauna of Western Anatolia, including only

OF ORTHOPTERA IN THE CAUCASUS AND WESTERN ASIA. A461

15 species which do not range into the latter district. Of these
15 species no fewer than three or four (Gryllus frontalis, Steno-
bothrus macrocerus, Arachnocephalus vestitus and, perhaps, Doli-
chopoda ewxina) should be regarded as not having been found as
yet in Western Anatolia owing to the lack of investigations only ;
one—Wdaleus mlokosiewitchi, being a very strong flyer, doubtless
came to the Pontian district from the east in recent times; the
presence of dipoda schochi schochi may be satisfactorily explained
by the influence of the neighbourmg Armenia; and, finally,
three species are peculiar to the Pontian district—Podisma
koenigi (Pontian endemic ranging also into the adjacent western
portion of the Somkheto-Kakhetian district), P. satunini, and
Olynthoscelis kerketa. The remaining six species are of great
interest’: three of them are definitely boreal in their origin and
inhabit the alpine district of Western Caucasus, from whence
they come into the Pontian district; this descendance of the
representatives of alpine fauna to the sea-level is due to the
great humidity of the Pontian climate, this fact being very
characteristic for the fauna of this district. The last three species
are Pecilimon schmidti, Isophya pyrencea, and Olynthoscelis fallax
all northern Baikanian in their origin and sylvan in their
habitation, which leads us to the conclusion that they come to the
Pontian district from the north—through the Crimea.

Thus, we may consider the Pontian fauna as an unpoverished
Balkano-Anatolian one, with the admixture of peculiar and
boreal forms and species of northern Balkanian origin which came
here from the north.

The eastern limit of the Pontian district is formed by the chain
of Suram; the southern boundary goes along the Adzharo-
Imeretian chain, approaching the Black Sea near Kobulety and
turning from here westwards along the chain of Pontus; the
western boundary is rather obscure and is to be looked for some-
where near Trebizond; the north-eastern boundary coincides with
the upper limit of the forests on the Western Caucasus; and
the north-western separating the Pontian district from that of
Novorossiisk is indefinite, as we shall see later on.

8. Zhe Novorossiisk district (N.).

This district is one of the less investigated ones, the whole
number of Orthoptera known from it being 50(4M-+21 A+
18 L+7G).

Analysing its fauna we observe the very close resemblance of
it to that of the South Russian steppe fauna and, on the other
hand, to that of the Pontian district. This intermediate character
of the Novorossiisk fauna is to be explained by the geographical
position of the Novorossiisk district between the steppe of South
Russia and the district of Pontus. The differences of the
Novorossiisk fauna from that of the South Russian steppes are
expressed in seven species, three of them being boreal in their
origin (Stenobothrus scalaris, Psophus siridulus, and Olynthoscelis

462 MR. B. P. UVAROV ON THE GEOGRAPHICAL DISTRIBUTION

griseouptera) and penetrating hence from the mountains of
Western Caucasus; one (Platycleis sepiunr) is a Balkano-Anatolian
species coming from the Pontian district ; and three remaining
ones (Parameles taurica, Olynthoscelis pontica, and Pezotettix
giornai) are of special interest. The first two of them are peculiar
to the southern part of the Crimea, and the Pezotettix is a
characteristic Mediterranean species, unknown as yet in the
Crimea, though doubtless present there. The presence in the
Novorossiisk district of these three species, which are absent from
all other districts of the Caucasus, indicates that this district was
once in a direct connection with the south of the Crimean
peninsula, the time and place of this connection being at present
unknown to us.

As to the boundaries of the Novorossisk district, its inter-
mediate position and the transitional character of its fauna render
them very obscure; I think they are not very markedly defined ;
its southern boundary separating it from the Pontian district is
to be looked for somewhere between 'Tuapse and Sotchi.

9. The Somkheto-Kakhetian district (S.-K.).

The Orthopteran fauna of this extensive district, though fairl
well investigated, is not yet fully known, as is evidenced by the
fact of the recent description of some new species and subspecies.
inhabiting it. The whole number of the known Somkheto-
Kakhetian Orthoptera reaches 79(7 M+ 35 A + 241 4 138 G),
which should be less than the real number by some 10-12 forms.

The first problem to be solved is whether this district belongs
to the Steppe or to the Balkano-Anatolian province. Let us
consider its affinities to both of them.

This district hasin common with the Steppe province (districts
of Southern Russia and of Kuban-Terek) 69 species, the difference
being 21. Temporarily setting aside eight Somkhete-Kakhetian
endemics, the remaining 13 are as follows:

*Empusa pennicornis. *Tsophya pyrenza.
*Tettix depressa. * =, amplipennis.
*Paratettix meridionalis. +) ee acumimahas
*Thalpomena ledereri. *Paradrymadusa sordida.
*Pcilimon distinctus. Nemobius heydeni.

* 5 bosphoricus. Gryllus lateralis.

*Tsophya adelungi.

No fewer than eleven of these (marked with an asterisk) are
Balkano-Anatolian species or belonging to the characteristic
Balkano-Anatolian genera, and two Gryllids only are recent in-
vaders from the adjacent deserts of the Caspian Transcaucasia.
Thus it is evident that the difference between the Somkheto-
Kakhetian and the Steppe fauna is very well expressed and in-
dicates the entirely different sources of their origin.

Turning to the relation of the Somkheto-Kakhetian fauna to
the Balkano-Anatolian one we see, that only 16 species inhabiting

OF ORTHOPTERA IN THE CAUCASUS AND WESTERN ASIA, 463

this district are foreign to other districts of the Balkano-
Anatolian province, eight of them being peculiar Somkheto-
Kakhetian forms. The remaining eight species are:

Stenobothrus lineatus. Platycleis vittata.

Pecilimon distinctus, Olynthoscelis fallax.

Tsophya adelungi. Nemobius tartarus.
» pyrenza,. Gryllus lateralis.

Out of these only Stenobothrus lineatus and Platycleis vittata
may be regarded as proofs of the influence of the steppe fauna;
two Gryllids are of desert origin; and all others belong to the
Balkano-Anatolian genera.

The above is sufficient to enable us to come to the conclusion
that this district may be regarded as a part of the Balkano-
Anatolian province, a conclusion confirmed by the study of the
Somkheto-Kakhetian endemics. They are eight in number, as
follows :

Arcyptera’‘flavicosta transcaucasica.

Celes variabilis carbonaria.

Tmethis zaitzevi.

Nocarodes rimansone.

Podisma koenigi (peculiar to this and Pontian district).
Tsophya bivittata.

Leptophyes nigrovittata.

Olynthoscelis distincta.

All these peculiar forms, with the exception of Arcyptera,
Podisma, and Celes, belong to the Balkano-Anatolian genera, and
are doubtless not recent invaders. This strongly supports my
opinion as to the affinities of the Somkheto-Kakhetian district.
Podisma koeinigi presents an evidence of the influence of the
Pontian fauna, and two peculiar characteristic steppe species,
Arcyptera flavicosta and Celes variabilis (as well as Stenobothrus
lineatus and Platycleis vittata), are doubtless immigrants from the
Kuban-Terek district (through the Daghestan), the somewhat
different natural conditions of Transcaucasia causing the sub-
specific differences between the Transcaucasian and the primary
steppe forms.

We ought, therefore, to consider the Somkheto-Kakhetia as
a district of the Balkano-Anatolian province, bearing in its fauna
some hints of an influence of the South Russian steppe fauna,
migrating from the north around the eastern end of the
Caucasus through the Daghestan.

The outlines of the Somkheto-Kakhetian district are very
complicated and circuitous. I include in it the southern
forest-clad slopes and hills of the Great Caucasus from Svanetia
on the west to the south-eastern extremity of this chain: here
the district branches around this end on the northern s!opes, thus
coming in contact with the Daghestanian district; through the
Suram meridional chain the northern half of the Somkheto-

Proc. Zoon, Soc,—1921, No, XXXII. 32

464 MR. BP. P. UVAROV ON THE GEOGRAPHICAL DISTRIBUTION

Kakhetian district is connected with the southern part, the district
thus ranging all over the northern slopes of Minor Caucasus
as far eastwards as Karabagh, where its fauna, becoming gradually
poorer, comes in contact with the fauna of Aderbaidzhan.

10. Zhe Talysh district (T.).

The fact that we only know 45 species of Orthoptera (3 M+
24A+8L+10G) from this district is certainly due not to its
poverty but to incomplete investigations.

As regards the composition of the Talysh fauna, it gives the
impression of bearing resemblance to that of the adjacent Caspian
Transcaucasia, being distinguished from it by the presence of
- five forms only, as follows:

Acrida turrita turrita.
Parapleurus alliaceus.
Stenobothrus macrocerus.
Epacromia strepens strepens.
Platycleis capitata.

The latter of these is an endemic species, while the presence of
the four remaining ones indicates the close affinity of the Talysh
fauna to that of the Balkano-Anatolian province. The most
remarkable fact is that dAcrida turrita and Hpacromia strepens
are represented in the Talysh district, not by the desert sub-
species inhabiting the Caspian Transcaucasia and Aderbaidzhan,
but by the same races that are met with in the districts helong-
ing to the Balkano-Anatolian province, the range of these sub-
species being discontinuous ‘The affinity of the Talysh fauna
with the Balkano-Anatolian one is even more defined by the fact
that its difference from the latter can be based upon a single
subspecies (leaving the endemic Platycleis capitata aside)— Decticus
verrucsivorus boldyrevi, which no doubt came here recently from
the neighbouring deserts, The influence of the Eremian fauna on
the fauna of Talysh is, generally speaking, very well marked,
resulting in the presence of such forms as Thisoecetrus dorsatus,
Platycleis escalerai, Liogryllus bimaculatus, ete., but it should be
regarded as of secondary nature. The immediate connection of
the Talysh with the other districts of the Balkano-Anatolian
province is now absent, but it no doubt existed formerly ; I think
it should be looked for in the south-eastern (Karabaghian) branch
of the Somkheto-Kakhetian district which formerly used to
reach the Talysh.

In the district of Talysh I include only the rather narrow strip
along the southern shore of the Caspian Sea, the southern bound-
ary of this district being the upper limit of the forests on the
northern slopes of the Talysh mountains. The north-western
boundary delimitating Talysh from the adjoining deserts of
Caspian Transcaucasia is rather indefinite; the north-eastern one
is completely unknown,

OF ORTHOPTERA IN THE CAUCASUS AND WESTERN ASIA. 465

ll. The district.of Aderbaidzhan (Ad.).

This district possesses a very rich Orthopteran fauna, the
number of species known being 125(9M -+1P+76A 4 24L
+ 15G); the real number should be considerably larger, not less,
I believe, than 150-160.

The most interesting features of this fauna are as follows.
First of all, the presence of a representative of the suborder
Phasmodea ( Gratidia bituberculata)—which is not to be met with
in any other district, except Caspian Transcaucasia—clearly
indicates that the Aderbaidzhan fauna belongs to a quite distinct
zoogeographical division. This is supported by the large number
of Mantodea, Acridiodea, and Gryllodea, while the number of
Locustodea is comparatively small. Among the Acridiodea the
large number of species belonging to the family Gidipodide is very
conspicuous; there are thirty Cidipodids here against ten, for
example, inhabiting the neighbouring Somkheto-Kakhetian
district. But the fauna of Aderbaidzhan is as remarkable for
what it lacks as for what it possesses. Out of these negative
features the total absence of the genera Chrysochraon, Gompho-
cerus, Arcyptera, Psophus, Celes, and Podisma is very remarkable ;
they are all of northern (boreal or steppe) origin, and do not
reach this district. Yet more interesting is the composition of
the fauna of Locustodea: out of the whole family Phaneropteride,
so well represented in the districts belonging to the Balkano-
Anatolian province, we only find two here—Phaneroptera falcata
and Tylopsis thynufolia, both very strong flyers and doubtless
recent immigrants; the highly characteristic for the Balkano-
Anatolian fauna family Sagide is represented in Aderbaidzhan
by one species only, the most widely distributed Saga ephippi-
gera; the majority of Locustodean fauna being thus formed by
the Decticide, which are sixteen in number, mostly species of
distinct ‘ancient Mediterranean” origin.

The originality of the Aderbaidzhan fauna is most clearly
demonstrated by the large number of peculiar species (some of
them also ranging into Caspian Transcaucasia); nearly one third
of them are not to be found in any other district of the country,
being distributed beyond its limits, while eighteen are true
endemics, as follows:

Eremiaphila persica. Derocorys roseipennis lazurescens.

Brunnerella mirabilis. Drymadusa grisea.

Scintharista brunneri. Paradrymadusa pastuchovi.

Thalpomena persa. =) satunini.

Helioscirtus moseri tichomirovi. ae persa.

Tmethis persa. cs longipes.
ecaninanliss % boecquilloni.

Nocarodes woronowi. Platycleis persica.

schelkovnikovi. Olynthoscelis satunini.

33
Amongst these endemics a great percentage of the ‘ancient
Mediterranean ” forms is evident, as, for example, all Paradry-

madusa species, Drymadusa grisea, two species of MVocarodes, etc,
32*

466 MR. B. P. UVAROV ON THE GEOGRAPHICAL DISTRIBUTION

Very characteristic of the Aderbaidzhan fauna are also nu-
merous Hremian genera and species, for example: Hremiaphila,
Oxythespis, LHelioscirtus, Sphingonotus, Derocorys, Sphodro-
merus, ete.

Summarising the results of our analysis, we may characterise
the fauna of the Aderbaidzhan district as a true EKremian one,
with a well-marked admixture of ‘‘ancient Mediterranean” forms
and with very slight indications of the steppe fauna, which pene-
trates here through the Somkheto-Kakhetian and Armenian
districts.

T include in the Aderbaidzhan district the Persian province
bearing this name (but not the recently formed republic of
Azerbaidzhan in Transcaucasia), as well as the valley of middle
Arax from Kaghyzman as far as Migry. Thus, the northern
boundary is to be drawn along the southern slopes of the Minor
Caucasus at a somewhat considerable height, about 6000-7000 feet
above the sea-level. ‘To this district also belongs (I am not yet
sure whether partly or wholly) the Karabagh, where the Ader-
baidzhan fauna comes in direct contact with the here already
impoverished Somkheto-Kakhetian one. Farther eastwards the
northern boundary of Aderbaidzhan goes along the eastern
boundary of the Zangezur chain and near Migry ¢ goes over the
Arax, turning eastwards nearly parallel with the latter river along
the chain of Karadagh as far as the Talysh chain; farther on
the boundary of Aderbaidzhan coincides with the latter chain.
The western boundary is the same as the eastern limit of
Armenia, which we have already considered above. <As for the
southern limit it isas yet unknown, but there are some indications
that it lies rather far southwards in Central Persia.

12. The district of Caspian Transcaucasia (T.C.).
5 ; )

The Orthopteran fauna of the deserts of the eastern or Caspian
Transcaucasia as compared with other districts has been more
fully investigated. The amount of its known species is 98
(6M+1P450A +4 24L4 17 7G), which number is, I suppose,
very near to the real one.

If we take into consideration the uniformity of this district, its
fauna may be regarded as a rather rich one, though poorer than
that of Aderbaidzhan. The difference between the fauna of the
latter district and that of Caspian Transcaucasia appears to be a
very marked one, since as many as 45 Aderbaidzhanian forms do
not reach Caspian Transcaucasia. Out of them 23, that is nearly
half, are species of ‘ancient Mediterranean ” origin, as follows

Pallasiella bolivari. 4. species of Nocarodes.
Pyrgodera armata. Drymadusa grisea.
Brunnerella mirabilis. a3 konowi.
Heliopteryx humeralis. 4 species of Paradrymadusa.
Charora crassivenosa. Platycleis persica.

Trinchus schrenki. a4 squamiptera.

4 species of Tmethis. Olynthoscelis satunini,

OF ORTHOPTERA IN THE CAUCASUS AND WESTERN ASIA. 467

The typical Eremian forms which do not penetrate into
Caspian Transcaucasia from Aderbaidzhan are 19 in number:

Eremiaphila persica. Helioscirtus moseri moseri.
Oxythespis wagneri. Fe » tichomirovi.
Fischeria baltica. 6 species of Sphingonotus.
Blepharis mendica. Leptoternis gracilis.

Duronia fracta fracta. Derocorys roseipennis lazurescens.
Platypterna tibialis. Schistocerca peregrina.
Scintharista brunneri. Sphodromerus serapis.

This list includes a rather large percentage of forms of more
southern origin (7. e. Indo-Ethopian), such as Osythespis,
Fischeria, Blepharis, Schistocerea, Sphodromerus, which are to be
regarded as recent invaders into the Eremian subregion and do
not reach its northern parts, which accounts for their absence
from the Caspian Transcaucasia. Besides, two species are
common to the latter district but represented by distinct sub-
species (Duronia fracta and Derocorys roseipennis), and nearly
all the remaining species are remarkable for their sporadic distri-
bution, being bound to certain habitations which are not to be
met with in Caspian Transcaucasia; their absence from the latter
district is thus easy to understand.

The remainder is formed of three species: Stenobothrus
zubowskyi, Callimenus dilatatus, aud Orphania scutata zacharovi.
The first of them is too little known as regards its geographical
distribution, and the two others are Balkano-Anatolian species
confined to the alpine pastures which are absent from Caspian
Transcaucasia.

The negative features which distinguish the fauna of Caspian
Transcaucasia from that of Aderbaidzhan are thus very numerous.
Nevertheless, they are but of little zoogeographical value, being due
to the comparative youth of the fauna of Caspian Transcaucasia,
it being the cause that ‘‘ancient Mediterranean” species (mostly
flightless or bad flyers) and the HKremian ones of more southern
origin have not had time enough to extend their range of dis-
tribution into this district but recently left by the waters of the
Caspian Sea.

Let us see now what is the positive distinction of the fauna of
Caspian Transcaucasia from that of Aderbaidzhan. The forms of
Caspian Transcaucasia which are not found in Aderbaidzhan are
19 in number, as follows:

Gelastorrhinus sagitta. *<Tsophya adelungi.
Duronia fracta kalmyka. kee SCHMITG ti,
Stenobothrus petreus. **Leptophyes albovittata.
* 55 parallelus. **Paradrymadusa sordida.
*Gomphocerus maculatus. Platycleis burvi.
*Arcyptera flavicosta transcaucasica. 3 decticiformis.
*Celes variabilis carbonaria, * a vittata,
Derocorys roseipennis roseipennis. *% i grisea.
**Pecilimon distinctus. **Olynthoscelis indistincta,

ie 3 bosphoricus.

468 MR. B. P. UVAROV ON THE GEOGRAPHICAL DISTRIBUTION

Two of them belong to the species represented in Aderbaidzhan
by other races (Duronia and Derocorys), and four are of no value,
their zoogeographical character being unknown (elastorrhinus,
Stenobothrus petreus, Platycleis burri, and Pi. decticiformis).
Thus, the difference is based on 13 species only, out of which six
fected: in list with an asterisk) belong to the steppe fauna and
seven (marked with two asterisks) are Balkano-Anatolian ; they
all inhabit the Somkheto-Kakhetian district as well, which
explains their appearance in Caspian Transcaucasia through
recent Immigration from the latter district.

This connection with the Somkheto- Kakhetian fauna is, never-
theless, but of little importance, the differences between it and
that of Caspian Transcaucasia being too numerous.

Summarizing these facts, we may conclude that the Ortho-

-pteran fauna of the Caspian Transcaucasia is undoubtedly in
close affinity with the Aderbaidzhan fauna, being nothing more
than the northern vanguard of the latter, distinguished by some
impoverishment and by slight marks of an influence from the
Somkheto-Kakhetian fauna. The past history of the fauna of
Caspian Transcaucasia should be rather short and simple: the
vast plains of eastern Transcaucasia dried up after the retreat of
the Caspian Sea were populated by the most mobile and progressive
elements of the Aderbaidzhan fauna, while Balkano-Anatolian
and steppe forms proved mostly to be unadaptable to the rough
conditions of life in the newly formed deserts.

The southern boundary of this district, delimitating it from
Aderbaidzhan, is not clearly enough defined, the still continuing
northward migration of Eremian elements being the cause of its
indistinctness; the approximate direction of this boundary is to
be seen above in the section dealing with the Aderbaidzhan
district (p. 466).

The boundaries between the Caspian Transcaucasia and the
Somkheto-Kakhetian district are also mentioned above. The
district of Caspian Transcaucasia gives a very long and narrow
branch northwards, along the western shore of the Caspian Sea,
but it is still uncertain where the northern boundary is to be
drawn, for the deserts along this shore are as yet unexplored.

13. The district of Caspian Ciscaucasia (C.C.).

This district is inhabited by 70 species of Orthoptera (4 M+
39 A+13 L4+14G) and, as its fauna may be regarded as
having been thoroughly investigated, this number cannot be ex-
pected to increase in a marked degree after further explorations.

A glance at the composition of this fauna enables us to conclude
that it belongs to the Eremian subregion. This is evident from
the fact of the relative abundance of Acridiodea and Gryllodea in
comparison with the poverty of Locustodea ; it is even still more
supported by the study of families: Cidipodide being well
represented, the Decticidze also, while of the Phaneropteride there

OF ORTHOPTERA IN THE CAUCASUS AND WESTERN ASIA. 469

is here one species only—the very well-flying Phaneroptera
Jaleatu, penetrating into this district from the neighbouring
district of Kuban-Terek along the valley of Kuma.

The fauna of Caspian Ciscaucasia shows the greatest resemblance
to that of the district of Caspian Transcaucasia, the difference
between them being based upon the negative features of the first,
while a rather large number (46, 7.e. 2M+4+1P+421A+
17 L+5 G) of Transcaucasian Orthoptera do not range into
Ciscaucasia ; a careful examination of this group shows that it is
composed of species of distinct southern origin, except Arcyplera
flavicosta and Celes variabilis, which ave members of the steppe
fauna represented in Transcaucasia and Ciscaucasia as well though
by different geographical races. As for the positive differences
of the Ciscaucasian fauna from that of Caspian Transcaucasia
they are 16 in number, as follows:

Acrida turrita turrita. (Edipoda schochi caucasica.
*Chrysochraon dispar. **Hyalorrhipis clausi.
*Stenobothrus hemorrhoidalis. Tmethis muricatus.
*Stauronotus brevicollis. *Saga pedo.

* ‘5 kraussi. *Platycleis montana.

Arcyptera flavicosta flavicosta. Decticus verrucivorus schugurovi.
*Hpacromia tergestina. **(ryllus odicus.

Celes variabilis variabilis. **Tridactylus tartarus.

Five of them are but geographical races (subspecies) of the
species represented in Transcaucasia as well; one, Z'methis
muricatus, is very closely related to the southern 7. bilobus, aud
is, perhaps, also but a race of the latter. The remaining group of
ten species is composed of seven steppe forms (marked in above
list with an asterisk) and three are desert species originating from
the Turanian province of the Eremian subregion (two asterisks).
The presence of these latter is very interesting, as it gives an
evidence of the affinity of the Ciscaucasian fauna to that of the
Kirghizian district of the Turanian province adjacent to it
on the north-east. Itisa very curious fact that the affinity of the
Ciscaucasian fauna to the Turanian province is far closer than it
is to the Ivanian one; a comparison of the fauna of Caspian
the Ciscaucasia with that of the Kirghizian deserts shows us that
the first contains one species only which is not represented in the
second: this is Wdipoda schochi caucasica, an evidently new
intruder into the Ciscaucasian plains from the dry stony hills of
Transcaucasia. This leads us to the conclusion that the desert
plains of Caspian Ciscaucasia, quite recently left by the retreated
Caspian Sea, got their Orthopteran fauna mostly from the north,
being populated by the most progressive elements of the steppe
and Turanian fauna (the greater part of which are also proper to
the Iranian province of the same Eremian subregion), while the
migration from Transcaucasian deserts was prevented by some
unknown factors. As the eastern Ciscaucasian plains were at
first separated from the Kirghizian deserts by the Strait of

470 MR. B. P. UVAROV ON THE GEOGRAPHICAL DISTRIBUTION

Manytch (which joined the Black Sea to the Aralo-Caspian basin),
it is necessary to conclude that the process of populating these
plains began after the drying up of the above-named strait, and
the whole fauna of the Caspian Ciscaucasia should be regarded
as being of quite recent origin, which explains the absence of
some characteristic Kirghizian forms, as, for example, drmene
alata, Oxythespis turcomanie, Pyrgodera armata, etc. All above
considerations support the idea that the deserts of the Caspian
Ciscaucasia form a distinct zoogeographical district of the
Turanian province.

As for the boundaries of this district, they are all well marked
except the southern one. This district occupies the clay and
sandy deserts adjacent to the lower currents of the rivers Kuma
and Terek, as well as the whole valley of the Manytch. I con-
sider the latter valley as the northern limit of this district,
while the western and south-western are determined by the
corresponding boundaries of the South-Russian and the Kuban-
Terek districts. The southern boundary, delimitating this
district from the Caspian Transcaucasia, is yet unknown, and I
suppose it is not very sharply defined.

14. The district of Western Caucasus (C.0c.).

The Orthopteran fauna of the subalpine and alpine zones of
the western part of the main Caucasian chain includes 40 species
(24A 4+ 1314 3G). Amongst them a very striking group is
formed by 20 species of evident boreal origin, as follows:

Tettix bipunctata. Stenobothrus apricarius.
» subulata. *Gompheocercus sibiricus caucasicus.
Parapleurus alliaceus. Arcyptera fusca.
Chrysochraon dispar. *Mecostethus grossus.
i brachypterus. _ *Psophus stridulus.
Stenobothrus nigromaculatus. *Podisma pedestris.
* x viridulus. *Locusta caudata caudata.
* heemorrhoidalis. *Platycleis roeseli.
* $5 ventralis. *Decticus verrucivorus verrucivorus.
*% " scalaris. *Olynthoscelis griseoaptera.

The presence of these boreal species as well as the total absence
of representatives of Mantodea, the small number of Gryllodea,
the poverty of Gidipodide and Decticide—this all gives to this
fauna a rather northern character. This character is further
strengthened by the remarkable fact that eleven, 7. e. more than.
half of the above-named boreal species (marked by an asterisk),
show a discontinuous range of distribution, their main (northern)
area being separated from the Caucasian one by a large space of
South-Russian steppes from which these species are totally
absent. As regards the way by which these boreal elements
came to the Caucasus there may be two different suggestions :
either they migrated via Balkania and Asia Minor, or reached
the Caucasus direct from the north at some remote time when the

OF ORTHOPTERA IN THE CAUCASUS AND WESTERN ASIA. A7\

climate of South Russia was colder and damper, which might
occur during the Glacial period. Asa good many of the boreal
Species, as for example, Mecostethus, Psophus, Podisma pedesiris,
are doubtless absent from Asia Minor, the latter supposition
should be by far the more correct.

The remaining 22 Orthoptera of this district show rather
mixed affinities. The most marked affinity is that to the
‘Balkano-Anatolian fauna as expressed by the presence of Woca-
rodes cyanipes, four species of ecilimon, four Lsophya and two
Psorodonotus, while the direct influence of the steppe fauna is
evidenced by the presence of such forms as Stenobothrus macro-
cerus, St. parallelus, and Celes variabilis variabilis.

The endemic forms of the Western Caucasus are only four in
number: Podisma satunini, P. rufipes, Isophya caucasica, and
I. kalischevskyi, their small number being a rather characteristic
feature of this fauna.

The northern and southern boundaries of this district coincide
with the upper limits of the forests on the corresponding slopes
of the Caucasian chain; while the eastern boundary, delimitating
Western Caucasus from the district of Eastern Caucasus, is as
yet insufficiently known; I suppose it is somewhere near the
sources of the Terek and the Aragva.

15. The district of Hastern Caucasus (C.Or.).
Only 17 species of Orthoptera (11 A+5 L+i1G) are known in

the eastern part of the Caucasian chain ; its fauna being thus
far poorer in comparison with that of the preceding district.
The most interesting features of this fauna are: the small
number of boreal and, in general, northern species and, further-
more, the presence of two exceedingly well characterized and,
therefore, very ancient endemics—Podisma lezgina and Phlocerus
menetriesi, the latter being the single representative of its genus.
The boundaries of this district are easy to understand.

16. The district of the Caucasus Minor (C.M.).

The Orthopteran fauna of this district, being rather well
investigated, includes but 52 species (30 A+18 L+4G), which
indicates its poverty.

The analysis of this fauna indicates its close aftinity to that of
Armenia, only twelve species being strange to the latter district,
as follows :—-

*Chrysochraon dispar. Pcecilimon similis.
Stenobothrus werneri sviridenkoi. *Meconema thalassinum.
Gomphocerus variegatus. *Platycleis bicolor.

*Arcyptera flavicosta transcaucasica. of . weseli.

*Psophus stridulus. Ms iljinskit.

*Celes variabilis carbonaria. *Decticus verrucivorus verrucivorus.

The bulk of this group evidently belongs to species of boreal
origin (marked in the list with an asterisk), some of them being

472 ON THE GEOGRAPHICAL DISTRIBUTION OF ORTHOPTERA.

the leading forms of the Boreal fauna (Chrysochraon, Psophus,
Meconema); their presence here, together with their absence from
the mountains of Armenia, allows us to conclude that they came
here somehow from the Great Caucasus, perhaps by the transverse
chain of Suram which joins the mountains of the Minor Caucasus
to the main chain. In this characteristic admixture of boreal
forms, as well as in the presence of two endemics (Stenobothrus
wernert svwidenkoi and Platycleis iljinshii), I see the sufficient
cause for separating this district from Armenia.

The district of Caucasus Minor occupies the high table-lands
of Akhalkalaki, Kars, and Alexandropol, sending a narrow and
long branch along the shores of the Goktcha Sea and, farther
south-eastwards, along the chain of Zangezur. The northern
boundary coincides with the upper limit of Sonkheto-Kakhetian
forests; the western goes along the Arsian chain; and the
southern is exceedingly circuitous, being not yet satisfactorily
explored.

The difficult task of drawing the accompanying map of
zoogeographical districts has been undertaken in a most friendly
way by P. I. Nagorny, and I avail myself of the opportunity of
once more expressing my sincere gratitude to him.

EXPLANATION OF THE MAP.

Zoogeographical division of the Caucasus and Western Asia.
(Text-fig. 1, p. 454.)

PALAARCTIC REGION.
I. Steppe subregion.
South-Russian Steppe province.
R.M. (Rossia meridionalis).—South-Russian district.
K.-T.— Kuban-Terek district.
D.—Daghestan district.

Il. Mediterranean subregion.
Balkano-Anatolian province.
W.—Novorossiisk district.
P.—Pontian district.
S.-K.—Somkheto-Kakhetian district.
C.M.—District of the Caucasus Minor.
T.—Talysh district.
Ar.—Armenian district.
A.M. (Anatolia mediterranea).—Western Anatolian district.
A.S.— District of the Syrian Anatolia.
Ill. Eremian subregion.
1. Iranian province.

Ad.—Aderbaidzhan district.
T.C.—District of the Eastern (Caspian) Transcaucasia.

2. Turanian province.
C.C.—District of the Eastern (Caspian) Ciscaucasia.

IV. Caucasian subregion (?).

C.Oc.—(Caucasus occidentalis).—District of the Western Caucasus.
C.Or.—(Caucasus orientalis).—District of the Eastern Caucasus.

ON THE AUDITORY BULLA IN THE MUSTELIDA. 473

24. The Auditory Bulla and other Cranial Characters in

the Mustelide. By R. I. Pococr, F.R.S., F.Z.8.
[Received March 22, 1921: Read May 10, 1921.]

(Text-figures 14-18.)

ConTENTS.
Page
Introduction ........ eee Lao
The Foramina in the Base Be ihe Skull. Beeiseraseaticat ATA,
The Internal Structure of the Auditory Bulla ...... 478
The Classification of the Genera by the Bulle ...... 484

In 1869 (Proc. Zool. Soc. pp. 4-37) Flower described certain
features in the base of the skull of several genera of Mustelide,
especially those connected with the auditory bulla and the
adjacent foramina, recording the position of the glenoid, carotid,
and condyloid foramina, and noting the shape and relative
prominence of the mastoid and paroccipital processes. In many
instances he opened the bulle and deseribed the internal struc-
ture. The genera examined were Lutra, Enhydris, Meles, Taxidea,
Mephitis, Helictis, Arctonyx, Mydaus, Gulo, Mellivora, Galera,
Martes, Mustela, and khabdogale |=TIctonyx|. He summarised
the results of his examination of the skulls of the Urside, Pro-
eyonide, and Mustelidee as follows:—(1) The cavity of the buila
is simple.... That is, although there are frequently trabecule
or partial septa passing mostly transversely across the lower part,
and generally connected with the tympanic ring, there is no
distinct and definite septum dividing it into a separate outer and
inner character [misprint for chamber. (2) The paroccipital
process is more or less triangular, and directed backwards, out-
wards, and downwards, standing quite aloof from the bulla. This
relation depends chiefly on the want of development of the
posterior portion of the bulla; and is absent, or obscure, in
Mustela alone. (3) The alisphenoid canal is present in the
true Bears and Ailurus; absent in all the others fits ef imal
Mustelide ].

Although these statements have been copied over and over
again in works on the classification of the Carnivora, further
investigation shows that not one of them is exact. The bulla
of Helictis, for example, is as completely divided as in the
Felidz, and there is a very distinct partition in Grison, G'ulo,
Mellivora, and other genera, although it is not so complete as in
the typical Afluroidea. As regards the paroccipital process,
although its prominence and remoteness from the bulla are well
marked in such genera as Lutra, Mephitis, Mellivora, Meles,
Taxidea, and others, it is in Helictis and Girison as closely
applied to the posterior end of the bulla as in typical Felidz, and
it is even more confluent with the bulla in Peceilictis than in

ATA MR. R. 1. POCOCK ON THE

Mustela. Flower’s statement about the absence of the alisphenoid
canal may stand with the qualification that the canal may be
retained as an abnormality.

In 1885, Mivart (Proc. Zool. Soc. pp. 363-389) supplemented
Flower’s observations with a few additional facts and included
Conepatus, Pacilogale, and Lyncodon in the list of genera.

Since that date it does not appear that the facts have been
re examined. The observations contained in the following pages,
based mainly on specimens that have died in the Society’s
Gardens, may, therefore, be interesting and useful.

The Foramina in the Base of the Skull.

The Alisphenoid Canal.—This tube, as is well known, is
typically absent in the Mustelide. But in one example of
Lutra lutra, I find the remnants of it on the left side of the skull
although the right side is normal. A little distance in front of
the foramen ovale and higher up there is a small orifice through
which a bristle can be passed so as to appear in the aperture
common to the foramen lacerwn anticum and the foramen
rotundum. ‘The passage is no doubt the homologue of the
alisphenoid canal in the Canide, Urside, Ailurus, Mungotide, ete.
Except as an item of evidence supporting the view that the canal
in question is a primitive Carnivore feature, the discovery of it
in a skull of Lutra is of no great moment. As I have elsewhere
pointed out, it may be present or absent in the Hyzenide (Proc.
Zool. Soc. 1916, p. 444, fig. 2).

The Posterior palatine foramina.—The normal position of
these is approximately opposite the inner lobe of the upper
carnassial tooth (pm.*) on the maxillo-palatine suture, along
which they move to a certain extent backwards or forwards.
They hold this position in Martes, Charronia, Gulo, Tayra,
Mellivora, Meles, Arctonyw, Mydaus, and Helictis. In Vaxidea
they may be as far back even as the anterior edge of the molar.
In Lyncodon, Grison, Ictonyx, Pecilictis, Lutra, Pecilogale,
Vormela, and Mustela, with its related generic and subgeneric
forms, Gale, Putorius, Lutreola, they are on the maxilla
approximately opposite pm. 3 and removed from the maxillo-
palatine suture. In Mephitis, Spilogale, and Conepatus they may
be even nearer still to the anterior palatine foramina, opposite
pm. 2.

It may be noticed that in all the genera with long jaws like
Meles, Arctonyx, Mydaus, Helictis, and Martes they are set back
on the suture, whereas in some genera with short jaws and often
reduced dentition, like Mustela, Vormela, Grison, and Lutra,
they are set forwards, wholly on the maxilla. Nevertheless this
position is not always correlated with the length of the jaws and
teeth, since they occupy the backward position in J/ellivora and
Taxidea.

Their variation in position within the Mustelide is interesting
in view of their constancy in position in other families of

AUDITORY BULLA IN DHE MUSTELID®. 475

Carnivora. In the Ursids, Procyonide, Canide, and Felide they
are always plaeed back upon the suture, whereas in the Hynide,

Text-figure 14.

Palates of Gulo gulo (A), Helictis everetti (B), Meles meles (C), Mephitis sp.
(D), Grison (FE), Mellivora ratel (F), showing the position of the post-
palatine foramina.

Mungotide, and the large number of genera usually assigned to
the Viverride they open on the maxilla in front of the suture ;

476 MR. R. I. POCOCK ON THE

and in connection with the Ailuroid (or Mungotoid) families
cited, it may be noted that the Felidse, which, like Mustela,
Grison, etc., have shorter jaws and the teeth more reduced
numerically than in any other Carnivora, they are set back,
whereas in such relatively long-jawed forms as the typical
Viverride they are wholly maxillary in position.

The Foramen rotundum.—As in the Canide, this foramen is
visibly separated from the foramen lacerum anticum in profile
view of the skull in: Martes, Charronia, Gulo, Helictis, and
Taxidea; but in Mustela, Putorius, Grison, Lyncodon, Tayra,
Tctonyx, Pecilogale, Mellivora, Meles, Mephitis, and Lutra it is
at most visible as a separate orifice when the orbit is looked into
from the front, lying quite close to the foramen lacerum anticum
in a depression common to the two. Not uncommonly the two
orifices are confluent owing to the failure of the partition to
ossify, but within the limits of the genus J/ctonya there is
individual or racial variation in this particular, and no doubt the
same is true of other genera. :

The Foramen ovale.—The position of this foramen with regard
to the orifice of the eustachian tube varies in accordance with the
length of the back of the skull and with the inflation of the
anterior part of the bulla. Typically it is separated by a con-
siderable space from that orifice as in Mustela, Putorius, Martes,
Charronia, Gulo, Tayra, Grison, Mellivora, Meles, Helictis, and
Lutra; but in Ictonyx, Pecilictis, Taxidea, Mephitis, Spilogale,
and Oonepatus the foramen is only separated from the eustachian
orifice by a thin plate of bone.

The Glenoid foramen.—The typical position of this in profile view
of the skull is in front of the external auditory meatus either in
line with it or slightly higher or lower as the case may be. It is
lowest in V'aawidea, where it is almost below the lower edge of the
meatus. In Meles it is, on the contrary, almost above the upper
edge of the meatus. In Mustela, Martes, Charronia, Gulo, Tayra,
Mellivora, Ictonyx, Grison, and Lutra it holds intermediate
positions. In Mephitis, Spilogale, and Conepatus it is, however,
set right over the middle of the upper rim of the meatus; and in
Helictis it occupies almost the same position—farther back even
than in the Canide.

When the skull is viewed from below, the position of the
foramen also varies with respect to the auditory meatus. In
Helictis, Mephitis, Spilogale, and Conepatus ib 1s quite concealed
behind the rim of the orifice. In MJartes, Charronia, and Meles
it is just concealed by its anterior rim. In Mustela, Putorius,
Tayra, Ictonyx, Grison, Taaidea, and Mellivora it is just visible
in front of the anterior rim of the orifice. In Gulo and Lutra it
is exposed in front of the tubular meatus, nearly as far inwards
as in the Urside.

The Stylomastoid foramen.——Typically this foramen lies on the
inner side of the mastoid prominence and between it and the
inflated portion of the bulla, It is approximately in line with

AUDITORY BULLA IN THE MUSTELID. ATT

Text-figure 15,

A. Part of the skull of Lutra lutra showing the abnormal occurrence of the
alisphenoid canal indicated by arrow.

B. Bulla of Helictis everetti, the posterior chamber opened from behind showing
the partition (pt.) descending to the petrous.

C. The same from below cut open to show the partition (pé.) running obliquely
between the anterior chamber (a) and the posterior chamber (0d).

D. Bulla of Putorius putorius opened from behind, showing the thickened spongy
tissue of the walls and the undivided cavity.

KE. Bulla of Lutra lutra showing the partial division of the cavity by rafters.

F. The same from below.

Lettering :—a, 4, anterior and posterior chambers of cavity (in B, C); g, glenoid
foramen ; 0, foramen ovale; y, petrous portion of periotic; po, paroccipital
process ; pt, partition (in B, C) ; 7, rafter; ¢, edge of tympanic annulus,

478 MR. R. I. POCOCK ON THE

the glenoid foramen and some distance from the foramen lacerwm
posticum. Its position varies in accordance with the inflation of
the bulla, and every gradation can be traced from Mustela, where
it is thrust far out, to Gulo, Lutra, and Mephitis, where it lies far
in towards the foramen lacerum posticum. It is relatively closer
to the latter orifice in the Skunks than in other genera.

The Internal Structure of the Auditory Bulla.

The bulla of AMeles meles may be described as representing a
common type in the family *.

The tympanic ring projects far into the cavity of the bulla as a
large semicircular lamina. The space between it and the roof of
the bulla is divided into chambers by a few vertical, arched, bony
plates, arising from the lamina and the superjacent roof and
extending transversely inwards on to the inner wall. These may
be called the rafters. There is also a tolerably large plate rising
from the upper side of the tympanic ring in front and extending
horizontally forwards to the bottom of the anterior wall of the
bulla, terminating internally at a point where the bulla touches
the foramen lacerum medium. Behind this point the floor of the
cavity is subdivided by low anastomosing ridges into a number of
pockets.

The posterior portion of the tympanic ring lies just above the
fenestra rotunda on the periotic bone; and the posterior rafter,
rising about one-fourth of the distance from the posterior end of
the tympanic ring, arches high above the periotic, and terminates
about half-way down the inner wall of the cavity. This rafter,
supplemented by the posterior end of the tympanic ring, partially
divides the cavity of the bulla into an anterior larger and a
posterior smaller chamber. Nevertheless there is a wide passage
between them above the periotic bone. The posterior part of the
cavity of the bulla is closed behind, and does not extend beyond
the limits of the bulla as defined externally.

The bulla of JJartes martes resembles that of deles in all
egsential respects, the posterior rafter being well developed ; but
in WM. foina all the rafters are less well developed, the posterior
being quite short.

In Charronia flavigula there is only a single rafter arising
from about the middle of the tympanic ring, arching with a
strong concave curve to the roof, then descending to the floor
along the inner wall of the bulla.

Tayra barbara resembles Charroma except that the rafter is
less elevated and less arcuate.

Gulo gulo is very like Meles and Martes martes, but the pos-
terior rafter has the lower edge, not lightly concave as in those

* In this paper the bulla is described as seen when the skull is inverted. Hence
the periotic forms part of the floor of its cavity; and the cavity is bounded by the
roof above and by the anterior, immer, and posterior walls. Flower described it as

seen when the skull is in its natural position, and I followed that course in the case
of the Felidae (Ann. Mag. Nat. Hist. (8) xviii. pp, 326-334, 1916).

AUDITORY BULLA IN THE MUSTELIDA. 479

forms, but horizontal or inclined slightly downwards towards the
inner wall of the cavity. It is thus nearer the periotic bone, and
diminishes the height of the passage between the anterior and
posterior portions of the bulla.

Text-figure 16.

A. Bulla of Tayra barbara opened from behind.
B. The same of Galo gulo.
C. The same of Martes martes.
D. The same of Martes foina.
i. The same of Meles meles opened from below in front and behind.
F. The same from behind.
Lettering as in Text-fig. 15.

Lutra lutra, cinerea, barang, and maculicollis have a very simple
type of bulla, differing from those described above in that the
tympanic ring, in conformity with the comparative flatness of

Proc. Zoou. Soc —-1921, No. XX XIII. 33

480 MR. R. I, POCOCK ON 'tHE

the bulla, is almost in contact with the superjacent portion of the
roof, to which it is attached by two or three very shallow rafters.
The posterior rafter, present in Meles, Gulo, and Martes martes,
is undeveloped, the only trace of subdivision of the cavity being
supplied by a rafter which extends transversely from the ring
along the roof and inner wall of the bulla to a point just behind
the orifice of the carotid canal; and, as indicated on the outside
of the skull, the cavity of the bulla posteriorly is far in advance
of the paroccipital process. Except for its flatness the bulla is
very like that of Tayra or Martes foina.

In Pecilogale, judging from a skull with a partially fractured
bulla in the British Museum, it seems that the walls of the bulla
are thin, not spongy, and that the cavity, reaching posteriorly to
the paroccipital, is simple and undivided except by incomplete
rafters, and does not communicate with a chamber hollowed out
in the mastoid, which is quite small. This genus therefore has
a simpler, more primitive bulla than is seen in Mustela or in
Ictonyzx.

Another type of bullaisfound in Gale nivalis, africana, Mustela
erminea, hibernica, and Putorius putorius and furo. ‘The cavity
ef the bulla is continuous from end to end, being undivided by
septa; but it is greatly reduced in size by the thickness of its
walls, which, instead of being composed of solid bone, thicker or
thinner, as the case may be, in the genera previously described,
is composed of spongy bone permeated with air-cells. These fill
the entire space above the tympanic ring, and almost obliterate
the portion of the cavity behind the petrous portion of the
periotic and also the anterior part of the cavity. There is no
definite hollow space in the external portion of the periotic; but
this bone is spongy and porous like the walls of the bulla itself,
and its spaces communicate with the cavity of the bulla.

The bulla of Helictis (H. everettiz) is quite unique. The tympanic
ring is fused to the roof but not to the same extent as in
Grisonella and Mephitis, its free edge projecting slightly, even in
the middle of its curve, and more so in front and behind. The
anterior rafter resembles that of Grisonella; but the posterior
rafter, rising low down on the tympanic ring behind, runs
obliquely forwards and inwards to terminate on the floor of the
bulla just inside the point where the orifice of the carotid canal

opens. Its inferior edge is closely clamped on to the petrous
portion of the periotic except at one point situated externally
towards the stylomastoid, where there is a notch in the partition
rafter close to the fenestra rotunda. Thus the bulla is completely
divided into two subequal chambers, except where the notch in
question affords a passage between them—a_ precisely similar
formation to that which is well-known in many Adluroidea. The
posterior chamber has a few low marginal ridges on its inner and
posterior walls, and the anterior end of the anterior chamber has
a few ridges in addition to the anterior rafter. The external
portion of the periotic is not hollowed.

|
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ome

AUDITORY BULLA IN THE MUSTELID®. 481

The bulla of MJellivora resembles internally that of Meles except
that the cavity is posteriorly in communication by means of a
tolerably large orifice in the anterior part of the hinder chamber
with a smooth-walled cavity hollowed out in the periotic behind
the stylomastoid foramen. ‘The posterior rafter is deeper and

Text-figure 17,

A. Bulla of Ictonyax opened from below, the arrow showing the passage between the
cavity and the hollow in the external portion of the periotic.

B. The same of Pecilictis opened from behind, showing the extension of the cavity
into the occipital and mastoid.

C. The same of Mellivora opened from below, the arrow showing the passage
between the cavity and the small hollow in the external portion of the
periotic.

D. The same opened from behind.

Lettering as in Text-figs. 15 and 16.

more arcuate than in JJeles, and forms a half-partition between
the anterior and posterior portions of the cavity of the bulla
and behind it on the inner side a vertical crest of bone juts out
from the inner wall of the bulla.

33*

482 MR, R. I, POCOCK UN THE

Tetonyx is like Mellivora except that there is no crest behind
the partition on the inner wall of the bulla; and the related
genus Pecilictis only differs in the greater size of the supple-
mentary chamber in the periotic and of the space above the
tympanic ring.

Taxidea is like Pecilictis. In conformity with the great
inflation of the bulla the space between its roof and the tympanic
ring is very high and septate. The posterior rafter is continuous
with the posterior edge of the tympanic ring, and descends so low
that only a small passage is left between its inferior arcuate edge
and the periotic. A posterior chamber is thus cut off almost
as completely as in some Ailuroids. This posterior chamber is
extended anteriorly above the tympanic ring on the inner side of
the stylomastoid foramen, and it communicates by a wide passage
with a large cavity, larger than in J/ellivora, hollowed out in the
periotic between the mastoid and paroccipital processes.

Thus the bulla of Taaidea is very different from that of Weles,
and not essentially like it as Flower stated. It is much more
like that of Mellivora, although the partition is better developed
and the periotic hollow much larger. In Jellivora, too, the
posterior chamber is not extended forwards above the tympanic
ring on the inner side of the stylomastoid.

In Grisonella (sp. from Cordova in the Argentine) the roof of
the bulla is depressed on to the tympanic ring, and fused with it
practically throughout its extent except for the presence of one

two very small cellular spaces in front and behind. The
anterior rafter is very large, and rising from the summit of the
anterior part of the tympanic ring, descends obliquely downwards
and inwards, dividing the anterior part of the cavity, which is
further broken up by. anastomosing ridges, into an upper and a
lower Pore Just behind the posterior end of the ring but
confluent with it arises a comparatively large arched rafter which,
crossing the roof, descends along the inner wall of the bulla,
dividing the cavity into an anterior and posterior chamber
which communicate by a comparatively narrow passage above the
peviotic. The posterior chamber has septa on the inner wall and
anastomosing ridges on the floor, and anteriorly just behind the
main transverse septum formed by the posterior rafter there is a
passage by which the posterior chamber opens into the hollowed

external portion of the periotic. The floor of this hollow is
uneven with ridges and shallow pits.

It is interesting to note that the bulla of Grisonella is widely
different from that of Yayra, although the two genera were
formerly regarded as identical. In possessing a hollowed periotic
chamber communicating with the cavity of the bulla, Grisonella
resembles Mellivora, Ictonyx, and Taaidea, and is further Jike the
latter in the origin of the posterior rafter from the posterior end
of the tympanic ring. But in none of these genera is the ring
confluent with the roof of the bulla as it 1s in Grisonella.

A skull of Zyncodon, with a broken bulla, in the British

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AUDITORY BULLA IN 'THE MUSTELIDA. 483

Text-figure 18.

‘

A. Bulla of Tawidea opened from behind, showing the nearly complete partition
: between the posterior and anterior chambers, and the extension of the
posterior into the external portion of the periotic.

j B. The same of Giisonella sp. opened trom below, showing the extension of the
' posterior chamber into the external portion of the periotic.

C. The same opened from behind, showing the complete partition and the extension
A of the posterior chamber into the external portion of the periotic.

D. Bulla of Mephitis opened from helow showing the absence of partition, the
i arrow indicating the passage from the cavity into the hollow of the external
j portion of the periotic.

E. Hollow in external portion of periotic of Mephitis opened from the outside.

F. The same of Conepatus.

; Lettering as in Text-fig. 15, with the addition of o, the orifice by which

the cavity of the bulla opens into the periotic hollow,

484 MR. R. tT. POCOCK ON THE

Museum shows that the cavity is broken up by trabecule into
spaces which extend into the mastoid. It is not possible, how-
ever, to ascertain from this specimen whether the cavity, before
fracture, was divided by a partition into two chambers as in
Grisonella or not.

In Mephitis, Spilogale, and Conepatus the bulla is still more
peculiar. As in Grisonella, the tympanic ring is fused with the
roof of the bulla and the cavity of the latter is a hollow space,
with at most a few septa or bony ridges, which is closed just
behind the periotic by a thickish wall of bone lying between the
stylomastoid foramen and the foramen lacerum posticum; but
opposite the outer side of the periotic there is a tolerably large
oval orifice by which the cavity of the bulla communicates with
a large hollow space occupying the expanded area of the skull
between. the auditory orifice and a point on the occipital crest
above the paroccipital process. Behind the bulla externally there
is a tolerably long, flat area running back to the paroccipital
process, which, as in Lutra, is placed some distance behind the
bulla.

In Mephitis and Spilogale the cavity in the periotic runs inwards
by means of a narrow passage up to the foramen lacerwm pos-
ticum, and the posterior margin of the orifice, by which the cavity
communicates with the pale is produced into a crescentically
curved ridge of bone. This ridge and the narrow passage running
to the foramen lacerum posticwn are absent in Conepatus proteus,
the walls of the cavity being undifferentiated.

In the British Museum there is a skull of Mydaus with a
broken bulla, which seems to show that in essential points the
bulla of this genus resembles that of Mephitis. The cavity of the
bulla, for instance, comes to an end posteriorly just behind the
petrous portion of the periotic, and on the outer side of this bone
there is an orifice by means of which the cavity of the bulla
communicates with the hollowed mastoid portion of the periotic.

The following table summarises the principal variations in the
structure of the bulla :—

a. Cavity of bulla closed behind, not communicating with a hollow
space inthe external portion of the periotic, the latter being
solid or merely permeated with air-cells.

6. Bulla distmetly divided into two subequal chambers by a
bony partition running obliquely inwards and forwards from
posterior end of tympanic ring to carotid foramen, the two
chambers communicating by means of a notch in the lower
edge of the partition behind poreriane end of petrous ae as

of ‘periatic TUR eer . Helictis.

. At most an incomes partition passing Pe ranteeeing ae
posterior portion of tympanic ring to the foramen lacerum

posticum.
c. Walls of bulla exceedingly thick, permeated everywhere
with air-cells. No partition .............. Mustela, Putorius, | Vormela].

ce’. Walls of bulla thin, without noticeable air-cells.

d. Tympanic ring closely applied to roof of bulla, the pos-
terior end of which is far in advance of the paroccipital. Dutra.

AUDITORY BULLA IN THE MUSTELIDA, 485

a Tympanic ring projecting as a strong semicircular ridge
mto cavity of bulla and attached to its roof by septa
which, vatter-like, pass inwards and downwards along
root and inner wall of the cavity.
e. Cavity very incompletely divided by a median rafter
arching high above the tympanic ring, leaving a wide
space between its inferior edge and petrous portion
COLE OUST NCE neeereee tet PC ia Martes foina; Charronia; Tayra.
e'. Cavity much more completely divided by an additional
rafter, the inferior edge of which extends nearly hori-
zontally inwards and approaches much nearer the
petrous portion of the periotic ......... Martes martes; Gulo; Meles.
a', Cavity of bulla opening behind into a larger or smaller hollow
space in external part of periotic.
Jf. Cavity undivided, communicating with periotic hollow by
means of a well-defined orifice opposite outer side of petrous
portion of periotic (tympanic ring fused with roof of bulla).

g- Walls of periotic hollow undifferentiated ..................... Conepatus.
g'- Walls of periotic hollow with crescentic ridge and a passage
reaching inwards to foramen lacerum posticum ...... Mephitis, Spilogale.

f'. Cavity more or less divided by rafter-like partition into an
anterior and a posterior chamber, the latter opening into
hollow of periotic.
h. Tympanic ring fused with roof of bulla; a very narrow
space, nearly filled by the periotic between the two
chambers; the chambers much broken up by trabecule... Grisonella.
h’, Tympanic ring projecting into bulla, the structure of which
closely resembles that described under heading d’.
7. Periotic hollow smaller, only reaching to a point half-
way between paroccipital and tip of mastoid ............... Mellivora.
z'. Periotic hollow larger, extending to tip of hollow
mastoid.

k. A bony septum separating posterior chambers of the
bulla from periotic hollow behind and leaving a
comparatively narrow passage of communication in
front ; inferior edge of partition between chambers
ol bullleeyevenlyeconcavertancs. scnses nce <coennees osteo eer ee Ictonyx.

k'. Posterior chamber of bulla and periotic hollow
confluent throughout their extent; inferior edge
of partition between chambers angularly arcuate
over petrous.

7. Bulla abutting behind against nearly obsolete par-
occipital and confluent in front with hamular

process of pterygoid, as in Ictonya ..........:.000 0 Pecilictis.
U'. Bulla not reaching the stout, projecting parocci-
pital, and ceasing in front far behind hamular ... Tawidea.

The three genera Pecilogale, Lyncodon, and Mydaus have been
omitted from the table because from want of material my exami-
nation of the bulle was imperfect. So far, however, as the
available evidence goes they would fall provisionally as follows :—
Pecilogale under d alongside Lutra because the roof of the
bulla is thin and close to the tympanic ring, and its cavity is
undivided and not in communication with a hollow in the
periotic. The shape of the bulla is, however, very different from
that of Zuéra. It is long, abuts against the paroccipital behind,
and has a short auditory tube as in Jfustela. Its likeness to the
bulla of Zutra is no evidence of affinity between the two genera,

486 ON THE AUDITORY BULLA IN THE MUSTELIDA,

It merely means, in my opinion, that they have inherited bulle
of a primitive kind, derivable from the Martes-Tayra type, the
flattening in the two cases having been independently acquired.
Lyncodon must be provisionally placed alongside Grisonella, and
Mydaus with Mephitis, Spilogale, and Conepatus.

Since such otherwise dissimilar genera as MJeles and G'ulo or
Martes have similar bulle, it is probable that that type of bulla
is the most primitive in the Mustelide. The rest of the bull
may be derived from: it by modifications in different directions.
The flattening down of the roof on to the tympanic ring and the
shortening of the posterior portion of the cavity gave rise to the
bulla of Zutra; similar flattening of the roof and the development
of a complete oblique partition to that of Helictis; thickening of
the walls with spongy bone filling the spaces between the main
septa to that of J/ustela; and the opening of the posterior
chamber into the hollow of the periotie to that of Mellivora.
From the latter may be derived the bulle of Zctonyx, Pwcilictis,
and Taxidea by extension of the periotic space, and in the case of
Grisonella this was accompanied by the flattening of the bulla and
the confluence of its roof with the tympanic ring and reduction
of the posterior chamber. The bulla of Mephitis and other
Skunks may also be derived from the type seen in Mellivora by
the disappearance of the partition behind the petrous portion of
the periotic, shortening of the portion of the cavity behind it,
and the forward movement of the orifice by which the cavity
communicates with the greatly enlarged periotic hollow.

ON THE VENOUS SYSYEM OF VARANUS. 487

25. On the Venous System of the Lizard Varanus bengal-
ensis (Daud.). By Goptmnp Srycn Tuapar, M.Sc.,
Professor of Zoology, Canning College, Lucknow*.

[Received April 5, 1921: Read May 10, 1921.]
(Text-figures 1-4.)

The only data we possess concerning the venous system of the
genus Varanus appear to be contained in the work of Corti,
‘De Systemate Vasorum Psammosauri grisei,’ 1847 (which I have
not seen; the observations are reproduced in Bronn’s Thierreich,
vol. vi. Abt. i11. Reptilien, by C. K. Hoffmann, p. 1010); Hoch-
stetter, “ Beitrage zur Entwickelungsgeschichte des Venensystems
der Amnioten, II. Reptilien (Lacerta, Tropidonotus),” Morph.
Jahrb. xix. 1893, p. 464 (mainly the renal and hepatic portal
systems of V. arenarius, according to Beddard a synonym of
V. griseus); Beddard, ‘‘On the Venous System in certain Lizards,”
P. Z.8. 1905, 1. p. 447 (V. griseus), and Beddard, ‘On the Vas-
cular System of Heloderma, with Notes on that of the Monitors
and Crocodiles,” P.Z.S. 1906, 11. p. 610 (V. griseus, niloticus,
and exanthematicus).
These descriptions, however, do not completely apply to the
present species, V. bengalensis, common in and near Lucknow,
and it seems desirable, therefore, to record the results of my
investigation.
Besides ordinary dissections, injections were made through

the anterior abdominal vein, using a thick solution of gum arabic
to which a little colouring matter—carmine or methyl blue—
was added. The fluid passes completely into the system, and it
is easy then to dissect out even the finer vessels. I have used
this fluid in preference to gelatine, as it can be employed cold,
and sets satisfactorily after being placed for a short time in
alcohol, swelling the vessels. I have to thank Prof. H. C. Ahuja,
of the Central Hindu College, Benares, for supplying me with
the necessary literature; and my grateful acknowledgments are
due to my friend Mr. 8. K. Zibbu, of the Lucknow Christian
College, for the assistance he has given me in the course of my

work,

I. The System of the Anterior Vena Cava (text-fig. 1).

(1) The Haternal Jugular Vein (¢.j.). It is stated that this
vein is ordinarily absent on the left side in Lacertilia ; in the
present species, however, it occurs on both sides, the vessels
running parallel on each side of the trachea. It joins the internal
jugular at the base of the neck. The external jugulars com-
municate with each other by four transverse connections, the
plexuses. ‘he first plexus is situated at the level of the thyroid
gland, close to the place of union of the external with the internal

* Communicated by Dr, J. StepHEnson, F.ZS.

488 PROF. G. 8. THAPAR ON THE

jugular vein. The second plexus occurs a little higher up, near
the hyoid, from which it receives branches (A.). The third and
fourth plexuses are near each other, on the floor of the buccal
cavity. The formation of plexuses between the external jugular
veins is essentially an avian character, and I am not aware that
it has hitherto been described in any Lacertilian.

Text-figure 1.
a

The Anterior Vena Cava and its branches.
a.v.c., anterior vena cava; an., antebrachial; 67., brachial; e.j., external
jugular; 4., hyoidean; ,7., internal jugular; 7., lateral vein; p., pectoral ;
pr.br., prebrachial; s., subclavian; ss., subscapular.

(2) The Internal jugular Vein (1. 7.) receives a short vessel, the
lateral vein (/.), from the hinder region of the neck.

(3) The Prebrachial (pr.br.) joins the vena cava at the junction
of the neck with the body. It is a vessel of fair size, running
along the preaxial border of the arm.

(4) The Subdclavian Vein (s.) is a very large vessel running
along the postaxial side of the arm; it receives many branches
from the arm, shoulder, and chest, the chief of which are

(a) Subscapular (ss.), from the muscles of the back,
(6) Pectoral (p.), from the pectoral muscles,

(d) Antebrachial (an.), from the forearm.

(5) The Azygos Vein (text-fig. 3,az.), which enters the right
anterior vena cava, isa fairly large vessel formed by the union of
two branches, a vertebral (v.) from the vertebral column, and a
parietal (pa.) from the body-wall. Beddard states that in
V. griseus the parietal branch of the azygos is connected with

VENOUS SYSTEM OF VARANUS. 489

the suprarenal portal; there is, however, no such connection in
V. bengalensis. The lateral parietal vein (l.pa.), ranning laterally
along the body-wall, communicates with it; behind, this vein
forms an anastomosis with the dorsal parieto-hepatie vein. The
vertebral branch of the azygos disappears from view at the side
of the vertebral column.

Il. The Posterior Vena Cava (text-fig. 4)—This vessel begins at
» the hinder end of the body a little behind the kidneys, and bifur-
cates into two branches, each of which runs in a groove along the
ventral surface of the kidney. Leaving the kidneys (‘“ posterior
cardinals,’ Beddard) they traverse the suprarenal bodies, beyond
which they again unite. The vessel passes through the appendage
of the right lobe of the liver and discharges into the sinus venosus.
The anatomy thus indicates the origin of the posterior vena cava
from two originally distinct vessels, the fusion of which to form
a single trunk is here still incomplete.

III. The Hepatic Portal System (text-fig. 2) is peculiar in the
present species. The femorai vein (f.) is formed by the union of
several small vessels from the muscles of the leg and the pelvis;
on entering the trunk it divides into two branches, the pelvic, and
one which I propose to call the ischio-mesenteric. The pelvic
vein (p.) runs forward along the inner border of the corpus
adiposum of its side, and after receiving the sciatic (sc.) from the
posterior region of the leg and a varying number (4-5) of veins
from the fat-bodies (f,-f,), unites with its fellow of the other
side to form a median abdominal vein; the pelvic vein also
receives two or three dorso-lumbar veins from the body-wall.

The epigastric veins (ep.v.) are two in number, and are described
by Beddard as originating in V. griseus from the sciatic veins.
In the present species I find that they arise from the pelvic;
each is a fairly long narrow vessel running dorsally over the fat-
body, from which it receives a few branches. It then curves
backwards, and again forwards, assuming a position ventral to
the fat-body, and, receiving many branches from the parietes,
proceeds forwards to open into the corresponding lobe of the
liver at its anterior ventral margin. Before entering the liver
each is joined by a ventral parieto-hepatic vein from the ventral

body-wall.
~The anterior abdominal ventral vein (a.a.), formed by the union
of the pelvic veins, is a single vessel throughout its course, and
joins the hepatic portal vein.

The other branch of the femoral, which I have called the
ischio-mesenieric (r.), is an altogether new vessel, not so far
described by any observer. It receives a small offshoot (x.) from
the hypogastric, and runs along the rectum, receiving minor
branches from the rectum and cecum. In front of the cecum
the two ischio-mesenterics unite to form one of the main roots
of the hepatic portal vein.

490 PROF. G. §. THAPAR ON THE

The hepatic portal vein (h.p.v.) forms two complete loops in the
mesentery of the small intestine (text-fig. 2); im the course of the
second loop it receives splenic, pancreatic, and gastric veins. It
runs forwards to the hinder end of the median sulcus of the liver,
and after receiving the anterior abdominal vein divides into two
branches, one entering each lobe of the liver towards its dorsal
side.

Text-figure 2.

The Hepatic Portal Vein and its connections.
aa., anterior abdominal vein; c¢., caudal; ep.v., epigastric; f., femoral ;
*,-f veins from the fat-body ; h., hypogastric; h.p.v., hepatic portal vein ;
p., pelvie; se., sciatic; 1., ischiomesenteric; I, 2, 3, 4, 5, vems forming
the hepatic portal; 6, gastric; x., connection between hypogastric and
ischiomesenteric veins.

It would thus appear that all the blood from the limbs and
alimentary canal passes through the liver before entering the
posterior vena cava. A large part of the blood from the hinder
region passes along the ischio- mesenteric vein into the liver, while
a small part enters the kidneys by way of the hypogastric veins.
Thus while the hepatic portal system is increasing in complexity
the renal portal is reduced. The short connection between the
hypogastric and ischio-mesenteric veins would allow the blood to
flow from the ischio-mesenteric to the hypogastric or vice versa ;
but the possibility of a flow into the hypogastric is perhaps
diminished by reason of the direction of the connecting vessel,

VENOUS SYSTEM OF VARANUS. 49]

taken in conjunction with the forward stream into the hypogastric
from the tail. phi

The dorsal pariecto-hepatic vein (d.p.v., text-fig. 3). The right
lobe of the liver has a small elongated appendage on the dorsal
side towards its hinder end, which encloses in its whole Jength the

posterior vena cava. Into this appendage opens a vessel from the

Text-figure 3.

The Veins in connection with the Liver.

d.p.v., dorsal parieto-hepatic vein; ep., epigastric; g.h., gastro-hepatic ;
h., hepatic vein; p.v., posterior vena cava; v.p.h., ventral parietohepatic
veins.

dorsal body-wall, the dorsal parieto-hepatic vein, supported by a
pocket-shaped fold of peritoneum which, as in /gwana and
Heloderma, attaches the appendage to the body-wall. This vein
is described by Hochstetter, and by Beddard, who finds it in other
species of Varanus, in the same position. The dorsal parieto-
hepatic is formed by the union of the dorso-lateral vein from the
body-wall with a vertebral vein from the vertebral column ; the
dorso-lateral forms an anastomosis with the parietal vein, as
previously stated.

Another vessel opens into the left lobe of the liver at its

extreme anterior tip—the gastro-hepatic (g.h.), coming directly

492, ON THE VENOUS SYSTEM OF VARANUS.

from the stomach. Hochstetter noted its position in V. griseus,
and Beddard in J. niloticus and exanthematicus. I find exactly
the same state of affairs in V. bengalensis.

The ventral parieto-hepatic veins (v.p.h.) are three in number.
Two of these are laterally situated, and are connected with the
corresponding epigastric vein close to its entrance into the liver.
The third is a median vessel from the hinder end of the body,
which receives smaller branches from the sides and enters the
liver at the median sulcus near its anterior end.

IV. The Suprarenal Veins (text-fig. 4, 1, 2, 3).—The suprarenal
bodies are situated in front of the kidneys, and each encloses the

Text-figure 4.

The Posterior Vena Cava and its connections.

ce. caudal vein; h., hynogastric; &., kidneys; p.v., posterior vena cava;
s.p., suprarenal body; 1, 2, 3, veins forming the suprarenal portal system.

corresponding branch of the posterior vena cava. Hach has its
own blood-supply ; the right suprarenal portal system consists of
three veins: one formed by the union of three vessels of the
lateral body-wall; the second, from the body-wall near the
vertebral column; and the third, from the omentum. The left
suprarenal portal is formed by two veins only, the vein which on
the right side runs close to the vertebral column being absent.

The chief peculiarities of the venous system of Varanus bengal-
ensis are thus :—

(1) The presence of jugular plexuses like those of birds, formed
by the external jugulars.

(2) The azygos vein joins the right anterior vena cava, and
one of its branches forms an anastomosis with the dorsal parieto-
hepatic veins.

(3) Presence of an ischio-mesenteric vein, and consequent
complication of the hepatic portal system.

(4) The origin of the epigastric veins from the pelvic and not
from the sciatic,

——

ls ees ee a

— a ee ee

SS EE Se ee ee

ABNORMALITIES IN THE COMMON FROG. 493

26. Abnormalitiesin the Common Frog (Rana temporaria).
By J. H. Luoyp, M.Se., F.Z.8., Zoological Depart-

ment, University College, Cardiff.
(Text-figure 1.)
[Received January 13, 1921: Read April 5, 1921.]

Both specimens described in the following notes were dis-
covered on dissecting a number of frogs for'class purposes at the
beginning of the present session.

Specimen A. Persistence of the right Posterior Cardinal
vein in an adult male Rana temporaria.

The occasional persistence of the posterior cardinal vein was
first noted by Howes (2) in 1888, in an adult female Rana
temporaria. In this specimen the postcaval vein was normally
developed and the persistent left posterior cardinal functioned
as an azygos vein. The cardinal was continuous posteriorly with
the renal portal vein. According to Parker (6), Howes afterwards
came across another frog in which an almost similar arrangement
occurred.

In the following year Parker (6) described a male frog with
a persistent left posterior cardinal vein, in which the hepatic
veins opened direct into the sinus venosus.

In 1905 Woodland (7) described a male. Rana temporaria in
which a large posterior cardinal vein persisted which was
continuous posteriorly with the enlarged right renal portal vein.
The venous blood was supplied to both kidneys by the left renal
portal only.

O’Donoghue (4 & 5) has described eight cases of the persistence
of posterior cardinal veins in adult frogs. Seven of his specimens
were males (six Rana temporaria and one Limnodynastes peronit),
and the other was a female Rana temporaria. The majority of
his specimens in addition exhibited abnormalities of the renal
portal veins.

Description of Specimen A.

This specimen exhibits a persistent right posterior cardinal
vein which runs from the anterior of the interrenal portion of
the posteaval vein, and opens anteriorly into the subclavian vein
about halfway along its length. The interrenal vein is normally
developed and receives the renal and spermatic veins from the
kidneys and spermaries of right and left sides. There is abso-
lutely no trace of the left posterior cardinal vein or of the
postcaval vein anterior to the kidneys. The renal portal veins
exhibit no abnormalities.

494 MR. J: H: LLOYD ON

The liver in this specimen is also peculiar. The left lobe is
normally developed and the left hepatic vein opens directly into
the sinus venosus. The right lobe is an elongated, slightly
dorso-ventrally flattened, cylindrical structure which is anteriorly
directed and terminates near the posterior edge of the mylohyoid
muscle. This lobe is divided into two portions by a transverse
constriction. About halfway along its length on the mesial
side of the dorsal surface, a hepatic vein runs out and opens into
the right precaval vein. ‘The internal and external jugular veins
enter the precaval together dorsal to the abnormal lobe of the
liver.

As far as I can ascertain this is the only case yet recorded of a
hepatic vein opening into a precaval. This irregularity is
obviously due to the abnormal condition of the liver.

Text-figure 1.

Sketch of specimen A from ventral surface, shewing persistent right posterior
cardinal vein and abnormal liver. B.V. brachial vein; G.B. gall-bladder ;
I.R. interrenal vein; L.H.V. left hepatic vein; L.L. left lobe of liver;
P.C.V. right posterior -cardinal vein; R.H.V. right hepatic vein; R.L. right
lobe of liver ; S.V. subclavian vein.

According to Hochstetter (1) the posterior portions of both
posterior cardinal veins fuse in Amphibia to form the interrenal
“Urniere abschnitte’) portion of the postcaval, the anterior
portion (‘ Leberabschnitt”’) of the latter beg formed by a venous
connection between the anterior end of the interrenal vein and
the tip of the liver. It is suggested that in the specimen
described above the formation of this venous connection was

ABNORMALITIES IN THE COMMON FROG. 495

interferred with by the growth of the liver towards the anterior
end of the animal. (Kerr (3) has pointed out “that in
Lepidosiren and Protopterus, the tip of the liver is in contact,
and fused, with the tip of the right kidney.”) This necessitated |
the persistence of one of the posterior cardinal veins to carry on
the functions of the postcaval in returniug blood from the
posterior end of the body.

it is interesting to note that this specimen being a male
supports O’Donoghue’s (4) suggestion that the absence of the
postcaval and persistence of a posterior cardinal vein appears to
be correlated with the sex of the animal.

SpecIMEN B. Abnormal Genital Organs in a male
Rana temporaria,

This specimen possessed a greatly hypertrophied testis, which
completely covered the left kidney, when viewed from the ventral
surface. There was absolutely no trace of a testis on the right
side, but the fat-body on both sides was well developed, that on
the right side being attached to the kidney.

The right kidney was about one-eighth of an inch shorter than
the left, and was distorted in such a way as to give it a
cylindrical appearance, and to cause the ureter to run on the
dorsal surface instead of on the outer edge. Both kidneys were
so closely apposed that it was difficult to ascertain whether there
was any actual fusion. There was a well developed seminal
vesicle attached to each ureter, and from the fact that both
seminal vesicles were apparently filled with sperms it seems
probable that there was some degree of fusion.

The left renal portal vein was normal but the right renal
portal was much reduced in size.

LITERATURE.

1. Hocusrerrer, F.—Beitriige zur vergleichenden Anatomie und
Entwicklungsgeschichte des Venensystems der Amphibien
und Fische. Morphol. Jahrb. Bd. 13, 1888.

2. Howes, G. B.—Note on the Azygos Vein in the Anurous
Amphibia. Proc. Zool. Soc. 1888, p. 122.

3. Kerr, J. Granam.—Note on the Posterior Vena Cava in
Polypterus. Proc. Roy. Physiol. Soc. Edinburgh, vol.
xviii. 1910.

4. O’Donocuusr, C. H.—The Persistence of Posterior Cardinal
Veins in the Frog, together with some Remarks on the
Significance of the Renal PortalSystem. Anat. Anz. Bd.
a0, LOLO,

5. O'Donoeuusn, C. H.-—Further Instance of the Persistence of
Posterior Cardinal Veins inthe Frog. Anat, Anz. Bd. 43,
LOTS:

Proc. Zoou. Soc.—1921, No. XXXIV. 34

496 ABNORMALITIES IN THE COMMON FROG.

6. Parker, W. N.—On the Occasional Persistence of the left
Posterior Cardinal Vein inthe Frog, with Remarks on the
Homologies of the Veins in the Dipnoi. Proc. Zool. Soc.
1889, p. 145.

4%. WoopLiaNnp, W.—On a New Mode of Persistence of the
Posterior Cardinal Vein in the Frog: with a Suggestion
as to the Phylogenetic Origin of the Post-caval Vein.
Zool. Anz. vol. xxviii. 1905.

THE ANATOMY OF THE TONGUES OF THE MAMMALIA. 47

27. The Comparative Anatomy of the Tongues of the
Mammalia.—IV. Families 3 and 4. Cebidee and Hapa-
lide. By Cuartes F. Sonntag, M.D., F.ZS.,
Anatomist to the Society.

[Received April 15, 1921: Read May 24, 1921.]

(Text-figures 37-52.)*

ConTENTs.

“Page Page

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Mannily @ebideeere trees | 4070) | Genus Alowatia.. ...me.0-..%5... Sid
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Genus @ebrrstps.n ns. yeh sete: AGS) Hannay, Hiapalidaey 2). 5..ceee.. eee ey.
GemiseAtelesm & -.. ssa teeter seis 502m “Genus Hapale see. 20
Genus Lagothrix ...............55 506 Genus Leontocebus ............... 520
Genus Saimiris .............0....... 6509 | Genus Wystqa 4. .5.-.8. 2... 823
Genus Callicebus .............0605 509 | Genus Gidipomidas ............... 523
(Creraitiss AGG ese Sanaa a eteeentl le GaLLy eek aan PRCT au ay cette, ee WP a elated Webuaen 2/5733
GleimS TERVAGCUG) so toopsenceeconcasne » DIAS) TBM biorewy Dh Coscanodsnanespboleon manson (a2!

INTRODUCTION.

The literature of the Cebide and Hapalide contains several
papers on the tongues of Cebus, Ateles, and Callithrix (= Hapale),
but few on those of other genera. The objects of this paper,
which is based on fifty-five specimens, are to supplement existing
accounts, and describe several tongues for the first time.

The tongues of the Cebidee and Hapalide resemble those of the
Cercopithecide in the characters of the apex and lateral borders ;
and they agree with them in that the foramen cecum, lytta, plice
fimbriatee, and apical gland of Nuhn or Blandin are absent.

I have given the measurements of fresh specimens only, for a
list of the dimensions of preserved ones is worthless.

I have employed Pocock’s nomenclature (12) for the Hapalidee.

Family CeBpip2&.

Pigmentation.—Many of the tongues of the Cebide are
pigmented, and the colour is evenly distributed over the entire
dorsum, or arranged in a definite pattern; on the inferior surface
it covers the parts around the frenal lamella which, as a rule, is
colourless. The vallate and fungiform papille may be colourless
or pigmented.

If several specimens of each pigmented species are examined it
will be seen how the distribution of the pigment varies greatly,
so the colour is of no value for purposes of classification.

* The explanation of the lettering of the Text-figures is given in the text.
DA*K
34

498 DR. C. F. SONNTAG ON THE ANATOMY

Several forms of pigmentation are shown diagrammatically im
text-fig. 37, but the descriptions are given with each species.

Text-figure 37.

Ateles Colicebus Lagothrix
QFiSeESCENS. rma/oct

watumzte

Patterns of pigmentation of the tongue: A. dorsum; B. inferior surface.

Genus CEBUS.

Tue Brown Capucuin (C. fatuellus).

The largest of three specimens exhibited the following measure-
ments :-—total length 5-45 em.; length of the oral part 3:9 em.;
length of the pharyngeal part 1:55 cm.; width between the
lingual attachments of the palato-glossal folds 2-1 em.

The Cirewmvallate Papille.—Specimen No. 1 (text-fig. 38 A, EB,
& F).—Three papille form an isosceles triangle with the apex
behind. Each lies in the centre of a low, flat, circular elevation
which is crowded with small cylindrical conical papille. Only a
small part of each papilla projects beyond the vallum. The basal
papilla is level with the lamine of the posterior half of its corre-
sponding lateral organ. Specimen No. 2 (text-fig. 39 A).—Five
vallate papille form a VY. The anterior papille are circular and
large, the posterior papilla is large and oval, and the mesial
papillae are small and circular. All are recessed within their
well-marked fosse.

The Fungiform Papille (text-fig. 38 A, B, & H).—-The papille,
which have a pearly appearance, form a dorsal bounding zone on
which they have the usual arvangement in rows and clusters,
but the transverse rows extend far forwards at the expense of
the apical cluster. On the inferior surface there are two rows— .
an inner one of closely- set large papille, and an outer row of small
discrete ones (text-fig. 38 B, b).

They are hemispherical, flat, or provided with a central eleva-
tion, and none are overlapped by conical papille.

OF THE TONGUES OF THE MAMMALIA. 499

The Conical Papille (text-fig. 38 A, B, & G).—Although they
have the usual distribution according to size, and their points
run in the usual directions, they are irregular in arrangement on
the oral part of the dorsum. They are disposed in oblique chains
on the base of the tongue. This lack of definite arrangement in
clusters and rows on the oral part of the dorsum is very common
on the tongues of the Cebide, and rare in the Cercopithecide.
On the lateral borders and inferior surface they are very closely
ageregated, but have no definite arrangement.

They are filiform or cylindrical, and haye one or more points.

Lymphoid Tissue and Glands (text-fig. 38 A, «).—The base of
the tongue is covered with nodules, or is delimited by a crescentic
zone of elevations. A few orifices of ducts and pits are prosent.

Text-figure 38.

The tongue of Cebus fatuellus.

The Lateral Organs (text-figs. 38 D and 39 B & C).—The
appearances vary greatly in different examples. They may form
concave bodies on the sides and dorsum (text-fig. 38 D), convex
bodies on the sides and dorsum (text-fig. 39 B), or long rows of
lamine and sulci on the inferior surface (text-fig. 39 C); the
concavities or convexities face the lateral vallate papillee (L.v.P.).
The lamine are rounded or pointed, and many are traversed by

500 DR. C. F. SONNTAG ON THE ANATOMY

secondary sulci. In two specimens the following measurements,
etc., were observed :—
Length. Laminee. Sulei.
Specimen No. 1.

Right organ...... 1-1 cm. tte) 10

Left organ......... "9 cm. 10 11
Specimen No. 2.

Right organ ...... enn: 9 10

Left organ......... *8 cm. 12 13

The Frenal Lamella (text-figs. 38 B, a, and 39 D).—The tri-
angular lamella has a bifid apex, and the edges, which extend
postero-laterally as far as the middle of the lateral organs, may be
plain or have long sharp processes. The free part, which is
triangular on section (text-fig. 38 1), has a ridge on its upper
surface.

The Ventral Papillary Border (text-fig. 38 B, 5) is narrow, and
the arrangement of its conical and fungiform papille has already
been described.

Text-figure 39.

@iue

The tongue of Cebus fatuellus.

The Ventral Mesial Sulcus (text-fig. 38 B,c¢ and C) is narrow
and deep anterior to the frenal lamella (text-fig. 38 A, c).
Posteriorly it is wide and triangular (text-fig. 38 C) and lodges
the ridge on the upper surface of the lamella.

The frenwm is short and thick. It passes along with the
lamella into the wide part of the ventral median sulcus.

THe WHITE-FRONTED CapucHtn (C. albifrons).

The following measurements refer to the larger of two
specimens :—total length 3°6cm.; length of the oral part 3-1 em.;
length of the pharyngeal part °5 em.; width between the lingual
attachments of the palato-glossal folds 1°6 em.

The Circumvallate Papille (text-fig. 40 A).—Three papille
form a triangle, but none are situated on elevations as in
C. fatuellus. The basal papille are circular, glistening, and
slightly retracted, but the apical papilla is deeply recessed (qa).
The fosse are all well-marked and the vallums are nedulated.
The right basal papilla (6) is compound, but all the others are

OF THE TONGUES OF THE MAMMALIA. 50}

simple (¢). The vallate area is crowded with conical papille, and
a fungiform papilla bisects the base.

The Fungiform Papille have a similar arrangement to those of
C. fatuellus, and exhibit the same characters.

The Conical Papille have no definite arrangement in clusters
and rows, but they increase in size in the usual manner. None
overlap the vallate and fungiform papille.

Lymphoid Tissue and Glands.—The base of the tongue is
covered with nodules, but no duct orifices are visible through the
lens. They are intermingled with conical papille.

The Lateral Organs lie on the dorsum and lateral borders of
the tongue, and are curved with the convexities forwards. Their
surfaces are smooth, and some are traversed by secondary sulci.
The inner ends of the laminz are pointed. Both organs have
nine lamine and ten sulci.

The Inferior Surface.—The ventral papillary border is narrow
round the apex, and widens on the edges of the inferior surface.

Text-figure 40.

° 900
cv) "a, fon

The vallate papillee of Cebus albifrons.

Its fungiform papille are arranged in two rows as in C. fatuellus.
The ventral mesial sulcws is short, narrow, and deep. It opens
posteriorly into a triangular fossa, which also receives the frenum.
The frenal lamella is triangular, with a bifid apex and serrated
edges which extend postero-laterally as far as the middle of the
lateral organs.

THe WeEEPER CAPUCHIN (C. capucinus).

Several authors have described the tongue of C. capucinus, and
the following arrangements of the vallate papille have been
recorded :—

(1) Three papille in a triangle..................... Mayer (4).
(2) Four papille as a double-pair ............... Mayer (4).
(3) Whree papille- in a triangle >) 0h. 0. 7.012... Flower (3).
(4) Three papille in a triangle (in 5 cases) ... Mtinch (5).

THe WHITE-THROATED CapucHIN (C. hypoleucos).

Tuckerman (9) described the macroscopic and microscopic
appearances, and recorded three papillz in the form of a triangle.

502 DR. C. F. SONNTAG ON THE ANATOMY

THe Hornep Carucuin (C. apella).

F, J. C. Mayer (4) pointed out that the tongue has four vallate.
papillz, and nine sulci in the lateral organs.

Podwisotsky (7) described six papille arranged in the form
ofa V. He also described the lingual glands which are well-
developed.

Boulart and Pilliet (10), Cuvier (2), and Minch (5) state that
the triangular pattern is the usual arrangement of the vallate
papille in the genus Cebus. It is present in the majority of my
specimens, but the V-arrangement and double-pair type appear
in the remainder.

Genus ATELEs.

THe GRizzLED Sprper Monkey (A. grisescens).

Measurements :—total length 3°6 em.; length of the oral part
' 25 em.; length of the pharyngeal part 1-1 cm.; width between
the lingual attachments of the palato-glossal folds 1-7 em.

Pigmentation.—The apex and a small area of the dorsum
behind it are unpigmented. Behind the latter the dorsum is
yellowish- brown, with a white cross in the centre. The long
limb of the cross is continuous with the colourless area behind
the apex, and each of the short limbs lodges a vallate papilla
(text-fig. 37). The frenal lamella is colourless, but the rest of the
inferior surface is yellowish-brown.

The Circumvallate Papille (text-fig. 41 A).—Three large
vallate papille form an equilateral triangle with the apex behind.
The left basal papilla is compound, and consists of two elements ;
the right basal papilla is oval and excavated; and the ,posterior
papilla is circular. The fosse are well-marked, and the vallums
form clear zones.

A fungiform papilla occupies the centre of the vallate area.

The Fungiform Papille (text-fig. 41 A) form a dorsal bounding
zone on which they have the usual arfangement. Those lying
beneath the apex are large and closely aggregated, but the
transverse rows extend far forwards on the dorsum.

The Conical Papille (text-fig. 41 A) have the usual arrange-
ment in clusters and rows; they increase in size in the usual
manner, and their points run in the usual directions. On the
pharyngeal part of the dorsum they ave small and irregular.
On the lateral borders and inferior surface they have the usual
arrangements. Ail the papille are small or of medium size.

The Lateral Organs (text-fig. 41 C) extend from the dorsum to
the lower parts of the edges of the tongue. Hach consists of
elongated fusiform lamine running forwards and upwards, and
the separating sulci are narrow. Each organ is surrounded by a
diamond-shaped unpigmented area. The right organ is ‘8 em.
long, and has 5 lamine and 6 sulci. The left organ is °7 cm

OF THE TONGUES OF THE MAMMALIA, 503

long, and has 4 Jamine and 5 sulci. The dotted line in text-
fig. 41 C represents the margin of the unpigmented area round
the lateral organs.

Lymphoid Tissue and Glands—The base of the tongue is
delimited by a zone of large nodules with a concave anterior
border. No duct orifices are visible even under a lens.

The Frenal Lamella (text-fig. 41 B, a)—The large, flat,
triangular lamella covers a large area of the inferior surface of
the tongue, and is devoid of a ridge on its upper surface. The
apex is divided into small sharp processes, and the plain edges
extend back to the middle of the lateral organs.

The Median Ventral Sulcews (text-fig. 41 B, b) lodges a well-
marked crest. It opens posteriorly into a flat, shallow, unpig-
mented triangular area.

Text-figure 41.

The tongue of Ateles grisescens ; ¢.t. tonsils.

The frenwm is very short. ;
The ventral papillary border is narrow, and its papille have
already been described.

Tur Buack-FAceD SpipER Monkey (4. ater).

Tuckerman (9) described the tongue, and the following
description is supplementary to his account. The measwrements
of the larger of my two specimens are :—total length 4:9 cm.;
length of the oral part 4:3 cm.; length of pharyngeal part °6 cm. ;
width between the lingual attachments of the palato-glossal folds
2 cm.

The Circwmvallate Papille (text-fig. 42 A & C).—Four large
papille form a double-pair, and two small papille lie on the.

504 DR. C. F. SONNTAG ON THE ANATOMY

lateral sides of the resulting rhomboid. Tuckerman states that
there are three papillz within the vallate area, but believes that
they are intermediate between the fungiform and vallate groups.
All the papille are circular, the fosse are well-marked, and the
vallums appear as clear zones. They vary in prominence, but
the posterior papille (C, 5) project to a greater extent than the
others. All are smooth and glistening,

Only conical papille are seen within the vallate area.

The papille of the anterior pair are level with the most anterior
Jamine and sulci of the lateral organs. They may be excavated
(text-fig. 42 C, a),

Text-figure 42.

Ye
Dd.

@.ve

The tongue of Ateles ater.

The Fungiform Papille (text-fig. 42 A & B), which are larger
than in A. grisescens, form a narrow dorsal bounding zone on
which they have the usual arrangement. They are numerous,
but small. On the inferior surface they are large; round the
under surface of the apex they are closely aggregated, but farther
‘back they are discrete (text-fig. 42 B).

Tuckerman showed how the fungiform papille on the under
surface of the apex are richly supplied with taste-buds.

The Conical Papille (text-fig. 42 A) are all very small, those
at the posterior part of the tongue, which are the largest, being

OF THE TONGUES OF THE MAMMALIA. 505

just visible to the naked eye. They have the usual arrangement,
and their distribution according to size takes the usual form.
Their characters are shown in text-fig. 42 D.

They resemble the papille of A. grisescens in that those within
and behind the vallate area are not regularly arranged in oblique
chains.

Lymphoid Tissue and Glands.—The base of the tongue is
delimited by a zone of prominent elevations, but no orifices of
glands are visible. These nodules have not been mentioned by
Tuckerman, who merely states that the base of the tongue is
wrinkled. He describes the mucous glands, which are arranged
in a small central and two lateral groups.

The Lateral Organs (text-fig. 42 K).—The prominent lateral
organs form crescentic bodies on the sides of the tongue, the
concavities facing the lateral vallate papille, which are level with
the anterior laminze and sulci. ‘The lamine and sulci are
numerous, and are irregular in position for they have been ac-
commodated withina small space. The right organ is ‘9 cm. long,
and has 9 lamine and 10 sulci. The left organ is 1 cm. long,
and has 10 lamine and 11 sulci. Many of the lamine have
secondary sulci, and some of the primary sulci are very wide.

Tuckerman described the histology of the lateral organs.

The Frenal Lameila (text-fig. 42 B)—The triangular lamella
forms a stout, thick plate with a notched apex and nodular sides.
Tuckerman described a lamella, with a fimbriated edge, which
possessed sensory (tactile?) end organs on its non-medullated
nerves, which form a network. Gegenbaur (11) criticises Tucker-
mau’s observations and thinks that the end organs are tactile in
function.

The long, narrow, deep ventral mesial sulcus opens posteriorly
into a small deep, triangular fossa. It lodges a small, median,
triangular crest derived from the short frenum.

THe BLACK-HANDED SpipER Monkey (4. melanochir).

Flower (3) describes the tongue as follows:—‘‘ The tongue is
rather lancet-shaped and pointed at the apex. The lateral group
of linear follicles at the base of the palato-glossal fold is very
distinct and large. The papille generally are soft and small.
Of cireumvallate papille there are only two, situated side by side,
close to the middle line opposite the attachment of the palato-
glossal folds. The frenum is attached very near (three-tenths of
an inch) to the apex of the tongue, which has a stong median
groove below. The sublingual papilla is rather small, soft, thick
and pointed, with a slightly bifid apex.”

THe Rep-raceD Sprper Monkey (4A, paniscus).

Mayer (4) described three vallate papilla forming a triangle.
Miinch (5) described the vallate papille of two specimens of
A. vellerosus. In one there were seven papille on each limb, but

506 DR. CG. F. SONNTAG ON THE ANATOMY

the central papilla was absent. The central papilla was present
in his second example.

Genus LAGoTHRIX.

THE Smoky Wooutty Monkey (L. infumatus).

IT examined three specimens, and the following measurements
refer to the largest:—total length 4°6 em.; length of the oral

Yext-figure 43.

The tongue of Lagothrix infumatus.

part 3°8 cm.; length of the pharyngeal part °8 em.; width
between the lingual attachments of the palato-glossal folds 1-9 em.
Pigmentation (text-fig. 37).—The three specimens are bluish-
black in colour, and the colourless fungiform and conical papille,
frenal lamella, and mesial part of the inferior surface stand out
prominently from the pigmental parts. The shape is variable in
resh specimens, being conical or oval.

OF THE TONGUES OF THE MAMMALIA. 5O7

The Circumvallate Papille (text-figs. 43 & 44 A & B).—Specimen
No. 1 (text-fig. 43 A).—Five papille form a V with long arms.
The two anterior papille are of medium size, the mesial papillee
are small, and the posterior papilla is large. All are prominent,
smooth and colourless, the fosse are well-marked, and the vallums
appear as clear zones. The apex of the V is far from the
epiglottis, and the anterior papille are close to the edges of the
tongue.

Text-figure 44.

Om eo @®

A B.
LEX ADS (SI
(Ss

DOCS NG Wale
/ 4 9 ‘i

Sa OV Gir
at(D0o-o OMe

E. iy

Dh dees 12
Ma AD
G.
The tongue of Lagothria infumatus.

Within the vallate area there are many conical papille, and
the posterior rows of fungiform papille enter the mouth of the V.

Specimen No. 2 (text-fig. 44 A).—Three papille form an
isosceles triangle whose equal sides are long (2 cm.), and the
posterior papilla is a considerable distance from the epiglottis.
The anterior papille are small and circular, the fosse are well-
marked, and the vallums are divided into lobes by fissures from
the fosse. The posterior papilla is oval and surrounded by a
prominent furrowed vallum.

508 DR, C. F. SONNTAG ON THE ANATOMY

Specimen No. 3 (text-fig. 44 B).—Four vallate papille are
arranged in the form of a Y, and all have well-marked fosse and
clear vallums. I do not consider that the Y-form in this case
corresponds to that of the Simiid, but I believe that the papille
of the mesial limb are members of a double-pair put into line
with the long axis of the tongue.

The Fungiform Papille (text-figs. 43 A and 44 C).-— The
papillae, which are of small and medium sizes, form a dorsal
bounding zone, but the transverse rows extend as far as the apex,
thereby eliminating an apical cluster. The rows contain many
papillee.

On the lateral borders and inferior surface the papille are
small, discrete, and arranged in single chains.

All are glistening and hemispherical, and are closely sur-
rounded by shaggy conical papille.

The Conical Papille (text-figs. 43. A and 44D) are closely-
aggregated and increase in size in the usual manner, but have no
definite arrangement on the oral part of the dorsum. They are
arranged in chains on the pharyngeal part of the dorsum and on
the inferior surface. All are filiform, and stand up round the
fungiform papille when the tongue is dried.

Glands and L ymphord Tissue,—The baseot the tongue Is occupied
by a large number of nodules, but no orifices of ‘ducts and pits
are visible. The nodules, however, are not aggregated together
on a crescentic zone in front of the epiglottis.

The submaxillary and sublingual glands are enormous (text-
fig. 43 B), and play the greatest part in the lubrication and
softening of the food in the buccal area,

The Lateral Or ‘gans.—Specimen No. 1 (text-fig. 44 E’—The
lamine and sulei vary greatly in size, and form irregular rows
on the lateral borders. The spacing between them is also very
variable, for the lamine are closely-packed posteriorly, and discrete
anteriorly. Many of the lamine are oval and traversed by wide
secondary sulci.

The right organ is 1 em. long, and has 13 lamine and 14 sulci.
‘The left organ is ae em. long, and has 12 lamine and 13 sulci.

Specimen No. 2 (text-fig. 44 F).—Both organs consist of long,
pointed laminze running upwards and on wards towards the
lateral vallate papillee (1. v.P), which are level with the anterior
lamine. The right organ is 1:1 cm. long,and has 11 lamine and
12 sulci. The left organ is 1-1 cm. long, and has 12 lamine and
13 sulci.

Specimen No. 3 (text-fig. 44 G)—The lamine and sulci all
run from behind forwards and upwards, and their white colour
makes them stand out prominently against the bluish-black
background of the tongue. The anterior amine are level with
the antero-lateral vallate papille, and the posterior laminz are
level with the antero-mesial vallate papilla. The right organ is
1 em. long, and has 12 lamine and 13 sulci. The left organ is
1:1 em. long, and has 11 lamine and 12 sulci.

OF THE TONGUES OF THE MAMMALIA. 509

The Frenal Lamella (text-fig. 43 Ba, Ca, & Da).—tThe tri-
angular lamella may have an entire or bifid apex. In the latter
the halves may be long and narrow, or wide and short. The
edges are plain or have processes.

The Ventral Papillary Border (text-fig. 43 Bb, Cb, & D b)
is wider than in Cebus and Ateles. Its conical papille are in hori-
zontal rows, but the fungiform papille form a single chain.

The Ventral Mesial Sulcus (text-fig. 43 Be, Cc, De) may be
narrow and deep, or wide and shallow. It has no mesial crest.

Humporpr’s Wootty Monkey (ZL. humboldti),

The tongue has three vallate papilie forming an isosceles
triangle; of these the apical one is large. The fungiform
papillz are large and conspicuous, and have the usual arrange-
ment in clusters and rows. The triangular frenal lamella has a
bifid apex.

Genus SAImMIRIS (=CuHRYSOTHRIX),

THE SqurrreL Monkey (S. sciwrea).

Measurements :—total length 2:1 em.; length of the oral part
1-9 cm.; length of the pharyngeal part °4 em.; width between
the lingual attachments of the palato-glossal fold 1:1 em.

The Circumvallate Papille.—Three papille form an isosceles
triangle with the apex behind. The right basal papilla is
strongly retracted within its fossa, but the others are prominent,
granular, and surrounded by nodulated vallums.

The Conical and Fungiform Papille have the usual arrange-
ment, and the latter form a dorsal bounding zone.

Glands and Lymphoid Tissue ave present as usual on the base
of the tomgue, but no duct orifices are visible.

Both lateral organs are 5 cm. long, and have five primary sulci
separating four lamine; the latter are traversed by secondary
sulci. Each organ appears asa straight line of lamine and sulci.

The frenal lamella, as shown by Pocock, is long, narrow, and
has a slightly divided apex. The frenwm is long.

Genus CALLICEBUS.
THe Moxocw Treree (C. moloch),

Three specimens were examined, and the measurements were
very similar in all. The following figures refer to my largest
example :—total length 2°55 cm.; length of the oral part
2°15 em.; length of the pharyngeal part -4 cm.; width between
the lingual attachments of the palato-glossal folds 1-2 em.

The tongue is bluish-black in colour on the entire oral part
and on most of the pharyngeal part of the dorsum. The inferior
surface has a bluish-black bounding zone. The frenal lamella,
mesial part of the inferior surface, and cuneiform areas on the

510 DR. C. F. SONNTAG ON THE ANATOMY

base of the tongue are unpigmented, and the vallate and fungi-
form papille may or may not be colourless; in the former case
the appearance of the dorsum resembles that of Lagothrix
infumata (text-fig. 37).

The Circumvallate Papille (text-fig. 45 A).—Three papille
form an isosceles triangle with long sides, and an acute
vertical angle. The posterior papilla is granular and _ projects
considerably beyond the coarsely-nodulated vallum. The right
basal papilla is granular, and the vallum is overlapped by conical
papillz. The left basal papilla is round, granular, and pigmented.

The Fungiform Papille (text-fig. 45 A & C) form a narrow
dorsal bounding zone on which they have the usual arrangement,

Text-figure 45.

The tougue of Callicebus moloch.

but the transverse rows extend far forwards. Those on the
inferior surface are minute and form a single row. Some are
overlapped by conical papille.

The Oonical Papille (text-fig. 45 A & D) are all shaggy, and
stand up around the fungiform papille when the tongue is dried.
The rows are mostly transverse, but the distribution according to
size is not according to the typical plan. All the papille are of
the filiform types.

Lymphoid Nodules and Glands.—The base of the tongue“is
occupied by nodules, but no orifices of ducts of glands are visible.

The Lateral Organs (text-fig. 45 E)—The wide primary sulci
appear as incisions on the lateral borders, for the intervening
lamin are flat. They form aslightly concave row whose posterior
members are level with the lateral vallate papilla (1.y.P.).

ee ee ee ee eS ee

"a7

OF THE TONGUES OF THE MAMMALIA. 5y ial

The entire organ is surrounded by an oval colourless area of
the tongue, so they offer a marked contrast to the bluish-black
lateral borders.

The right organ is °6 em. long, and has 10 lamine and 11 sulci.
The left organ is °5 em. long, and has 9 lamine and 10 sulci.

The Ventral Papillary Zone is bluish-black and narrow. It
has many small irregularly-placed conical papille, and a few
fungiform papille in a single row posteriorly,

The Ventral Mesial Sulcus is narrow and deep, and the frenal
lamella is triangular and entire with lobulated edges,

THE Brown TEEres (C. brunnea).

In a hitherto unpublished paper Garrod states: ‘There are
three circumvallate papillz in the form of a triangle. The fungi-
form papille ere most numerous at the tip of the tongue, and
there are a few in front of the circumvallate papilla and Mayer's
Organ. There is a good bifid sublingua” (=frenal lamella).

Owen (6) described a vallate triangle in C. personata,

Genus Aorus (=NycriprrHects).
Tue Fevine Dovrovucoeut (A. felinus).

The two specimens which I examined were preserved, so no
measurements were made, One of them has a single vallate
papilla.

The apex and lateral borders have the usual characters, and a
well-marked mesial dorsal sulcus runs along the base of the
tongue.

The Circumvallate Papille (text-fig. 46 A & C).—Two pro-
minent vallate papille lie near the lateral borders. Each is
granular, and has a prominent nodular vallum. The fossa are.
well-marked. Both are round on plan and on conical elevation.

The Fungiform Papille (text-fig. 46 A & D) form a dorsal
bounding zone on which they have the usual arrangement. Some
are flush with the general surface of the tongue (0), but others
stand up prominently (a & 6). They form a “single row on the
ventral surface of the tongue, and some are overlapped by conical
papille (d).

The Conical Papille (text-fig. 46 A & E) have the usual dis-
tribution according to size, and the usual direction of their
points. They do not have the usual arrangement, however; on
the oral part of the dorsum they are irregular, but on the base
they form oblique chains. They are cylindrical or filiform, and
may have central depressions.

The Lateral Organs (text-fig. 46 F).—The lamine and deep
narrow primary sulti lie on the edges and the inferior surface
of the tongue. They run forwards and upwards, and the laminz
are traversed by secondary sulei. The right organ is ‘9 cm. long,

Proc, Zoou. Soc,— 1921, No, XX XV, 35

512 DR. C F. SONNTAG ON THE ANATOMY

and has 5 lamine and 6 sulci. The left organ is ‘8 cm. long, and
has 4 lamin and 5 sulci.

The Inferior Surface (text-fig. 46 B)—The narrow ventral
papillary zone has irregularly disposed conical papillee and a single
row of discrete prominent fungiform papille; the latter are most
numerous and closest together round the apex.

Text-figure 46.
Vad hs

Elevation.

Leve/ of
surface of 2&- 6 SS a
tongue.

Kee) COA

[Ey
The tongue of Aotus felinus.

The frenal lamella is triangular. Tt has a bifid apex and plain
sides. The ventral mesial sulcus is shallow and deep, and the
frenum is of moderate length.

Genus PIPHECIA.
THe Buack Saki (LP. satanas).

The specimen which I examined was preserved, so no measure-
ments were made.

The tongue has almost parallel sides, and the apex and lateral
borders have the usual characters. The thickness rapidly
diminishes from frenal lamella to apex.

The Circumvallate Papille (text-fig. 47 A & C).—The vallate

apillae form an isosceles triangle with the apex behind. All are
slightly retracted, with granular surfaces, the fosse are well-
marked, and the vallums are nodulated,

2 a ee

ee ee ae ee

OF THE TONGUES OF THE MAMMALIA. 513

The Fungiform Papille (text-fig. 47 A & B) form a dorsal
bounding zone on which they have the usual arrangement, and
the apical cluster is very well-marked. They form a single row
on the inferior surface. They are all prominent, smooth, and
hemispherical, and none are overlapped by conical papille.

The Conical Papille (text-fig. 47 A & D) have the usual
arrangement in clusters and rows of varying degrees of obliquity.
The vertical direction of the rows on the lateral borders can only
be made out by very careful examination, however. All belong
to the filiform series.

Text-figure 47.

- The tongue of Pithecia satanas.

The Lateral Organs (text-fig. 47 E) form triangular or concave
bodies on the sides and inferior surface, and the lateral vallate
papille are level with the apex. Several of the pointed laminz
have secondary sulci. The right organ is ‘9 cm. long, and has
5 lamine and 6 sulci. The left organ is ‘8 cm. long, and has
4 lamine and 5 sulci.

The Inferior Surface (text-fig. 47 B).—The broad bounding
papillary zone is wide, and has papille disposed in the usual
manner.

The frenal lamella is a long plate with rounded apex. and the
orifices of the Wharton’s Ducts open on the ventral surface near

the base (text-fig. 47 B, a).
The ventral mesial sulcus is wide and deep, and the frenwm is

short.
Mayer deseribed a vallate triangle in P. satanas (4).
Bie

514 DR. C. F. SONNTAG ON THE ANATOMY

Humpoupt’s Saxt (P. monachus).

Flower (3) describes the tongue as follows :—‘“ The length of
the tongue, from the base of the epiglottis to the tip, is 1-4 inch;
its breadth *5 inch; its sides are parallel, the end square, with
the corners shghtiy rounded. The sublingual organ is fleshy,
except towards the end, which is sharp-pointed and cleft at the
extreme tip; it is free to the extent of -25 inch, and its apex is
‘3 from the tip of the tongue.”

In a hitherto unpublished note Garrod states that the tongue
of Pithecia albinasa has a triangular vallate papillary area.

Genus CAcagao.
THe Batp OvaKari (C. calva).

The specimen had been preserved, so no measurements were
made.

The apex and lateral borders have the usual characters, but fine
lateral striz, only visible on high magnification, run in from them
across the dorsum.

Text-figure 48.

The tongue of Cacajao calva.

The Circumvallate Papille (text-fig. 48 A).— Three vallate
papille form an isosceles triangle whose apical papilla is far from
the epiglottis; the basal papillae are close to the edges of the
tongue. The apical papilla is small and the basal ones large, the
fosce are all well-marked and the vallums are nodular.

The Fungiform Papille (text-fig. 48 A & B) form a dorsal
bounding zone on which they have the usual arrangement in

OF THH TONGUES OF THE MAMMALIA. 515

clusters and rows, but the apical cluster is small. Nowhere are
they numerous, however. ‘They are absent from the inferior
surface.

The Conical Papille are irregularly arranged, but they increase
in size in the usual manner and their points run in the usual
directions. They form a narrow papillary zone on the inferior
surface, but this is widest round the apex.

Glands and Lymphoid Tisswe—The base of the tongue is finely
nodular, and a few orifices are visible when it is highly magnified.

The Lateral Organs (text-fig. 48 C) form concave bodies on
the sides of the tongue, and the pointed or round Jamine have
fungiform papille below their bases. The lateral vallate papillee
are level with the centres of the organs. All rounded lamine
have wide secondary sulci.

The right organ is 1 cm. long, and has 7 lamine and 8 sulci.
The left organ is -9 em. long, and has 6 lamine and 7 sulci.

The Inferior Surface (text-fig. 48 B).—The ventral papillary
zone is narrow, the ventral median sulcus is shallow, and the
frenum is small.

The frenal lamella is long and narrow, and terminates in two
small points (text-fig. 48 B, a).

Genus ALouarra (= Mycerss).
Tue Brack Howrter Monkey (A. caraya).

The specimen had been preserved, so no measurements were
made.

The Circwnvallate Papille (text-fig. 49 A & C).— Three
papille form an isosceles triangle with long sides. The apical
papilla is far from the epiglottis, and the basal papille are close
to the lateral borders of the tongue. The length of the vallate

triangle is one-third the length of the dorsum.
’ The apical papilla is large, oval, granular, and excavated in the
centre, the fossa is well-marked, and the vallum striated.

The Fungiform Papille (text fig. 49 A & B) are not numerous,
but have the usual arrangement on the dorsum; and a few lie at
the sides of the vallate triangle. They appear to stand on ridges
of the mucosa when the tongue is highly magnified. Those on
the inferior surface are very small.

The Conical Papille (text-fig. 49 A, B, & D), which are closely-
set, are irregularly disposed on the dorsum, but they increase in
size in the usual manner, and their points run in the usual
direction. They are filiform, cylindrical, or fusiform, and the
number of points to each papilla varies. Some stand up so
prominently round the fungiform papille that they conceal them
entirely.

Glands and Lymphoid Tissue (text fig. 49 A).—The base of the
tongue is covered with nodules of various sizes, and many orifices
of ducts are visible to the naked eye. The lens reveals the
existence of many minute openings, and the base of the tongue is

516 DR. C. F. SONNTAG ON THE ANATOMY

more pitted than in other species of the Cebide. Many nodules
with central depressions are also present on the anterior surface
of the epiglottis.

The Lateral Organs (text-fig. 49 E).—On each side they form a
straight row of lamine and “sulci. All the sulci and lamine
are short ; the posterior ones are straight, and the anterior ones,
which are level with the anterior vallate papille, are convex
forwards.

The right organ is ‘9 em. Jong, and has 10 lamine and 11 sulci.
The left organ is ‘7 em. long, and has 8 lamine and 9 sulci.

Text-figure 49.

MP1!) yy )))) ))
| I

The tongue of Alouatta caraya; t.t. tonsils.

The Inferior Surface (text-fig. 49 B).—The ventral papillary

zone is narrow, but has papilla disposed i in the usual manner.

The ventral mesial sulcus is short, but does not open up much
when traced posteriorly.

The frenum is short, and the frenal lamella is long, narrow,
deeply incised at the apex, and has plain sides (text- fig. "49 B, a).

Carus and Otto (1) described a vallate triangle (A. fuscus)
closely resembling that of A. caraya, described above.

——

~~ ee ee ee

SS, ee

CE ew 5

OF THE TONGUES OF THE MAMMALIA. 517

Summary of the Tongues of the Cebide.

1. All tongues, except those of the Sakis, are conical, and some
are pigmented.

2. The conical papille are irregularly disposed on the oral part
of the dorsum in most of the Cebide, but they may be arranged
in chains on the base of the tongue.

3. The fungiform papille always form a dorsal bounding zone.
They may be absent from the inferior surface.

4, The vallate papillary patterns are—a single papilla, a pair,
a triangle, a V, or a double-pair.

5. The vallate triangle is far from the epiglottis in many
species, and its median may be very long (the median is the line
drawn from the apex to the mid-point of the base).

6. The orifices of the ducts of glands are fewer than in the
Cercopithecide. They are most marked in Alowatta.

7. The lateral organs are most frequently concave towards the
epiglottis.

8. The frenal lamella is usually bifid at the apex.

9. The lingual characters are not sufliciently distinctive to be
of value for the classification of species.

Family HAPALID&.
Genus HAPpALe.
Tue Common Marmoszer (7. jacchus).

‘The comparatively long, slender tongue has the following
measurements :—total length 2°1 em.; length of the oral part
1-9 cm.; length of the pharyngeal part *2 cm. ; width between
the lingual attachments of the palato-glossal folds °65 cm.
These figures were found to be the average in twelve specimens.

The apex is flat and tuberculated by fungiform papilla; it is
devoid of a notch. Running back for 1 cm. is a median dorsal
sulcus from which transverse sulci pass to the sides of the tongue,
but these are the result of preservation and are not shown in
text-fig. 50. he lateral borders have the usual characters.

The Circumvallate Papille (text-fig. 50 A).—Three vallate
papill form an isosceles triangle with the apex behind, and the
two basal papille are close to the edges of the tongue. ‘The
vallums are covered by conical papillae which obscure the fossee,
and the surfaces of the papillary bodies are finely granular.
This pattern was present in all twelve specimens. Tuckerman (9)
also described this as the type.

The Fungiform Papille (text-fig. 50 A & B) form a dorsal
bounding zone on which they have the usual arrangement, in
clusters and rows, and they are surrounded by prominent
upstanding conical papilla. They are few in number, however,
on the dorsum, and are absent from the ventral bounding
papillary zone.

518 DR. C. F. SONNTAG ON THE ANATOMY

The Conical Papille (text-fig. 50 A, B, & C) have an irregular
disposition, but they increase in size in the usual manner, and
their points run in the usual directions. They appear as imbri-
cated scales, and some stand up prominently round the fungiform
and vallate papille, concealing the vallums of the latter
(text-fig. 50 C, e). .

Lymphoid Tissue and Glands—A few nodules are present

among the conical papille on the base of the tongue, but no

duct orifices are visible.

Text-figure 50.

Wy ints, *
vty, a)

ue 4 V tm

Cc.

The tongue of Hapale jacchus.

The Lateral Organs (text-fig. 50 D).—These consist of a few
fine lamine and sulci on the sides of the tongue. The right
one has 6 lamine and 7 sulci, and the left one has 5 lamine and
6 sulci. Some of the lamine have small secondary sulci.

The Inferior Surface (text-fig. 50 B).—A narrow ventral
papillary border, possessing conical papille ‘only, is present, and
the narrow, deep ventral mesial sulcus opens posteriorly into a
triangular fossa.

_ The frenal lamella is triangular, with a bifid apex and plain or
nodulated sides,

—=—;

eT eee ee

a ae

—_ =. ee eee

OF THE TONGUES OF DHE MAMMALIA. 519

Tse Buack-wARED Marmoser (ZZ. penicillata).

The conical tongue, which is green in colour, has the following
measurements :—total length 2°35 em.; length of the oral part
2-1 em.; length of the pharyngeal part -25 em.; width between
the lingual attachments of the palato-glossal folds -9 em.

The apex is pointed and tuberculated ; running backwards
from 1:4 cm. is a median ridge, whence transverse ridges, which
are convex forwards, run out to the lateral borders.

The Circwmvallate Papille.—Three papille form an isosceles
triangle. The apical papilla is close to the epiglottis, and the
lateral papille are close to the edges of the tongue. The lateral
papille are small, and their fossee and vallums are concealed by
conical papille (text-fig. 51 A, 6). The posterior papilla, which
is eval and excavated, has a nodulated vallum composed of two
rows of lobules (text-fig. 51 A, c).

The Fungiform Papilic form a wide dorsal bounding zone, and
the area of the centre without papille is small. They have the
usual arrangement, but the posterior rows form thick clusters in
front of the vallate papille. They are hemisphericai or flat, and
the surfaces are smooth and glistening. As in 1. jacchus, there
are none on the inferior surface of the tongue.

The Conical Papille have the usual arrangement, but their size
remains the same from apex to base. They are scale-like and
imbricating, as in H. jacchus. They form a narrow zone on the
inferior surface of the tongue (text-fig. 51 A, a).

Lymphoid Tissue and Glands—The area occupied by the
lymphoid nodules is small, and no duct orifices are visible.

The lateral organs consist of a few very fine laminz and sulci.
It is difficult to detect these, even on high magnification, by a
hand lens, so microscopical examination is necessary in order to
settle their characters.

The Inferior Surface (text-fig. 51 A, a).—A narrow ventral
papillary border is present; the narrow, deep ventral mesial sulcus
opens posteriorly into a triangular fossa; and the frenal lamella 1s
triangular, with an entire apex and notched sides.

Tur Pyemy Marmoser (41. pygmea).

The tongue has the following characters :—

1. It is conical with a rounded apex.

2. The vallate papille form an isosceles triangle.

3. The fungiform papille resemble those of H. penicillata.
. The conical papillae form imbricating scales.

. The lateral organs are well-marked.

. The ventral papillary zone is well-marked.

. The ventral mesial suleus is long and deep.

. The triangular frenal lamella has a bifid apex.

OID Oe

520 DR. C. F. SONNTAG ON THE ANATOMY

Genus LEONTOCEBUS.
Tue Lion Marmoser (L. rosalia).

The two specimens examined were preserved, so no measure-
ments were taken.

The tongue rapidly tapers from base to apex ; and the latter is
pointed and covered with a dense cluster of small fungiform
papille. The middle of the dorsum is occupied by a strip of
mucosa covered by closely-aggregated conical papille.

The Circumvallate Papille (text-fig. 51 B, 6)—Three papille
form an isosceles triangle with two short sides and a long base.

Text-figure 51.

fi
sf ANN

a.
(Ge
® @@ 6
ae
c Wy © tGewe
é. a. las
4,
A 4 (e) Dd.
Cc.
B.

The inferior surfaces of the tongue and yallate papillae of some Marmosets.

The apical papilla is a short distance anterior to the epiglottis,
and the basal papille are close to the edges of the tongue.

All the papille are large and nodulated, the fosse are well-
marked, and the vallums are clear zones or lobulated bands.

The Fun giform Papille, which are prominent, form a well-
marked dorsal bounding zone on which they have the usual
arrangement in clusters and rows, and they increase in size in the
usual manner. ‘T'wo rows of small elements connect the basal
vallate papille. None are present on the posterior part of the
ventral papillary zone.

All are prominent, hemispherical, and smooth.

The Conical Papille (text-fig. 51 B, c).—No apical cluster is
present, for the transverse rows extend right forward to the apex.

eS."

OF THE TONGUES OF THE MAMMALIA. 521

Hach is leaf-shaped, and has several hair-like processes along its
free edge, and all the points are directed backwards. Some of
the posteriar papille may be umbilicated. They increase in size
in the usual manner.

Lymphoid Tissue and Glands.—The nodules form a narrow
zone in front of the epiglottis, but no duct orifices are visible,
even through the lens.

The Lateral Organs.—The lateral organs consist of well-marked,
but small, backwardly-directed Jamin separated by deep primary
sulci. The right organ has 3 lamine and 4 sulci, and the left
organ has 4 lamine and 5 sulci.

The Inferior Surface (text-fig. 51 B, a & a'').—-No ventral
papillary zone is present, but the outer part of the inferior
surface incised by small fissures.

The ventral mesial sulcus, which is narrow and deep, opens
posteriorly into a large triangular space.

The frenum is short, and the frenal lamella is triangular, with
a bifid apex, or a deep notched lateral border.

Genus Mystax.

Tae Necro Tamarin (J. wrsulus).

My fresh specimen has the following measwrements :—total
length 2°8 em.; length of the oral part 2°4 em. ; length of the
pharyngeal part ‘4 cm.; width between the lingual attachments
of the palato-glossal folds 1:2 em.

The apex is rounded, and a dorsal median sulcus, which is more
marked than in any specimen belonging to the Cercopithecide,
Cebide, or Hapalide, runs back to the apex of the vallate triangle.
No transverse ridges or sulci are present.

The Circumvallate Papille (text-fig. 52 A & C).—Four
papille are arranged in the form of a V. Including the apical
papilla the right limb has three papille and the left one has two.
The posterior papilla (A, C a) is oval, with a well-marked fossa
and nodular vallum. The left lateral papilla (A, C 3) lies in a
wide fossa with three small ones. The right lateral papilla
(A, C c) lies in a wide fossa with two small ones, and the right
mesial papilla is circular.

The Fungiform Papille (text-fig. 52 A & B).—Small, but
well-marked papille form a dorsal bounding zone, but the trans-
verse rows are maintained as far forwards as the apex. They
are absent from the inferior surface.

The Conical Papille,—No apical cluster is present, for the
papille maintain their arrangement in rows right forward to the
apex, and there is no great increase in size from before-backwards,
or without-inwards. They have the usual arrangement on the
ventral papillary zone. All are cylindrical and surrounded by
zones of interpapillary dorsum. ;

Lymphoid Tissue and Glands.—On each side of the epiglottis
there is a large nodular mass, and the two are connected by a

522 DR. C. F. SONNTAG ON THE ANATOMY

band of small nodules which also ‘passes towards the posterior
vallate papilla. Some of the large nodules have small central
pits.

The Lateral Organs.—The lamine, which increase in size from
before backwards, are separated by wide primary sulci. The right
organ has 3 lamine and 4 sulci, and the left one has 4 lamine and
5 sulci. e

The Inferior Surface (text-fig. 52 B).—The ventral papillary
border, which is narrow, possesses conical papille only.

Text-figure 62.

¢
2pAt wx 27450
NS bo Al)

t i Ya
/|
I

0 23 Voy
0 eS AS
10,4 (CA) \—w

The tongue of Mystax ursulus.

The ventral mesial sulcws, which is long, narrow, and deep,
opens posteriorly into a Jarge triangular depression containing a
well-marked median crest.

The frenal lamella is a long, flat body with its anterior

extremity deeply cleft into several long, thin club-like processes.

Tue Movusracup Tamarin (I. mystaz).

The tongue, whose form resembles those of the preceding
species, has the following measurements :—total length 2-1 em. ;
length of the oral part 1:8 cm.; length of the pharyngeal part
-3 cm.; width between the lingual attachments of the palato-
glossal folds °6 cm.

A deep median dorsal sulews runs back for 1:1 em. from the
apex, and transverse furrows run towards the lateral borders ;
these separate broad ridges which are convex forwards and con-
cave backwards.

The Circumvallate Papille.—Three papille form an isosceles
triangle with two short sides and a long base ; the apical papilla
is close to the epiglottis, and the basal papille are close to the
lateral borders of the tongue.

eI ee eee eS ee eee

ee ee eee

Tie

le ea a ett

ene ee se es AS a

OF THE TONGUES OF THE MAMMALIA, a8

All are round and finely-granular, the fosse are well-marked,
and the vallums are lobulated. .

The Fungiform Papille, which are all small but prominent,
form a dorsal bounding zone on which they have the usual
arrangement in clusters and rows, the usual number on each row
being two. Most of them are equal in size, and are smooth and
hemispherical.

The Conical Papille, which are all closely-ageregated, have the
usual arrangement in rows and clusters; they increase in size
from without-inwards, but their increase in size from before-
backwards is slight. They take the form of overlapping scales
with hair-like processes.

Lymphoid Tissue and Glands.—The nodular zone on the base
of the tongue is narrow, and no orifices of ducts and pits are -
visible, even when the tongue is highly magnified.

The Lateral Organs ave small and oval: their centres are level
with the lateral vallate papille. They consist of well-marked
primary sulci separating comparatively broad lamine with pointed
or rounded upper extremities. The secondary sulciare very thin.

The right organ is *1 cm. long, and has 3 lamine and 4 sulci.
The left organ is ‘1 cm. long, and has 4 lamine and 5 sulci.

The Inferior Surface.—The narrow ventral papillary zone has
only conical papillx, but is incised by transverse sulci running in
from the dorsum.

The ventral mesial sulcws, which is narrow and deep, opens
into a large triangular space, which also receives the short
frenum.

The frenal lamella is triangular, with a bifid apex and fissured
lateral borders.

Genus CipIpomipas.
Tue Pincnt Monkey (@. edipus).

Flower (3) described the tongue and gave the following data :—

1. The tongue tapers gradually.

2. There is a median dorsal sulcus posteriorly.

3. The ventral median sulcus is deep.

4. The fungiform papille are regular, and cover the entire oral
part of the dorsum.

5. Three vallate papilla form a triangle.

6. The lateral organs ave well-marked.

7. The frenal lamella has a dorsal ridge and cleft apex.

Summary of the Tongues of the Hapalide.

1. The tongue is either long, narrow, and tapering slowly, or
short, conical, and tapering rapidly from a wide base.

2. Mesial dorsal ridges are frequently present.

3. The tongue of Mystax ursulus has a better-marked mesial
dorsal sulcus than those of all other Primates,

54 THE ANATOMY OF THE TONGUES OF THE MAMMALIA.

4, The vallate papille always form a triangle.

5. The fungiform papille always form a dorsal bounding zone.

6. Seale-like conieal papillz, with or without filamentous pro-
cesses, are exceedingly common.

7. The ventral papillary zone is small or absent.

8. The frenal lamella is usually subdivided at the apex, but
undivided lamelle occasionally appear.

9. The lateral organs are always small.

10. The lytta, foramen cecum, plice, fimbriate, and apical

gland of Nuhn are absent.

BIBLIOGRAPHY.

1. Carus and Orro.—Erliuterungstafeln zur verg]. Anatomie.
Heft 4. Leipzig, 1835.
. Cuvrer.—Legons d’Anatomie comparée. Paris: Crochard et
Cie., 1835.
. Frower, W. H.—Medical Times and Gazette, 1872, vol. i.
pp. 507-509; and Proc. Zool. Soc. London, 1862, p. 331.
. Mayer, F. J. C._—See Bibliography to paper No. 8.
Mincu.—See Bibliography to paper No. 8.
Owen, R.—The Comparative Anatomy of the Vertebrata,
vol. 111.2 199:
”. Popwisorsky, V.—‘‘ Anat. Untersuch. iiber die Zungendriisen
des Menschen und der Siiugethiere.” Diss. Dorpat, 1878.
8. Sonnrac, C. F.—“The Tongues of the Cercopithecide.”
Proc. Zool. Soc. London, 1921, pp. 277-322.
9. TuckerMAN, F.—Journal of Anatomy and Physiology, 1892,
pp. 391-392.
10. Bovunarr and Piniier.—See Bibliography to paper No. 8.
11. Gecensaur, C.—Morphol. Jahrbuch. 1894, pp. 1-18.
~ 12. Pocock, R. I.—Ann. Mag. Nat. Hist. (8) xx. pp. 247-258.

Oop w »w

ON THE SKULL OF DINOTHERIUM GIGANTEUM. 525

28. Note on the Skull of Dinotherium giganteun in the
British Museum. By C. W. Anprews, D.Sc., F.R.S.,
F.Z.S.

(Submitted for Publication by permission of the Trustees of the British Museum.)
[ Received March 5, 1921: Read May 24, 1921.]
(Text-figures 1-4.)

One of the most important specimens in the Geological Depart-
ment of the British Museum is the fine skull of Dinotheriwm
giganteum, which seems to be the only even approximately
complete example at present known. The history of this spe-
cimen is of considerable interest. It was discovered in 1835 by
Klipstein, about eighteen feet deep in the well-known ossiferous
sands of Eppelsheim in Hesse-Darmstadt. An account of this
discovery, including a description of the specimen and of the
deposits in which it occurred, was published by Klipstein and

‘Kaup in 1836 under the title ‘“‘ Beschreibung und Abbildungen

von dem in Rheinhessen gefundenen colossalen Schiidel des
Dinothervi giganter.” : in the following year a French translation
of this paper appeared. Kaup, in his part of the memoir, not
only gives a description of the skull itself, but also'a very
interesting account of the method of collecting the specimen.

_It appears that it was found lying on its dorsal surface in a bed

of clay and sand. This was partly removed, leaving the skull
resting on six pillars of matrix. Next, iron bars were passed
through the openings thus made, which were then filled up with
plaster of Paris. Next, the remaining portions of the matrix
were removed and their place filled with more plaster, so that
finally the skull rested on a solid base of plaster strengthened by
iron bars. A stout board was inserted beneath the whole mass,
which was then lifted from the excavation by twenty-four men
with ropes. A quaint picture of this operation is given on the
cover of Klipstein and Kaup’s memoir. In this figure there
is also shown a huge proboscidean femur, which is perhaps the
original from which the cast of a Dinotherium femur now in the
British Museum was made. In 1837 the skull was sent to be
exhibited in Paris, part of the expenses of transport being borne
by the French Academy. While there it was examined by
de Blainville, whose description of the specimen is the best yet
published ; but even he was not allowed any opportunity of
examining the dorsal surface. On the same occasion Isidore
Geoffroy, Straus, and others took the opportunity of giving their
views as to the structure and affinities of Dinotheriwm. It was
then intended to send the specimen to London, but whether this
was done on this occasion is uncertain. Some years later, in
1849, it was certainly in London, and was offered to the British

526 DR. C. W. ANDREWS ON THE

Museum for purchase ; it was then examined and reported on by
Buckland and Owen. ‘The purchase was not completed, and
the skull seems to have been sent back to Darmstadt. About
1866, Klipstein sold his collection to Dr. Oldham, the Director
of the Geological Survey of India, and from him the British
Museum acquired not only this, but also some other valuable
specimens, including the skull of Dorcatheriwm nawi and the front
of the skull of Z'apirus priscus.

Dr. W. D. Mathew, when visiting the Museum recently,
pointed out to me that there is a widely-spread idea on the
Continent that the skull now under discussion was broken up
on its journey to London. This mistake has even appeared in

vint: thus Weinsheimer in his memoir on Dinotheriwm (Pal.
Abhandl. Bd. i. 1883, p. 244), in speaking of the cast of this
specimen, states that ‘“ The original of this cast no longer exists,
since on its journey to London it was irretrievably broken into
fragments.” Examination of the skuil shows that, in fact, at some
time it has been much broken, but has been, on the whole, skilfully
mended, the figures and cast no doubt having been followed
in making the restoration.

In addition to Kaup’s original descriptions and _ figures,
de Blainville has given a very good account of the skull so far as
he was able to examine it, and numerous other writers have
referred to it. The most complete summary of the various views
that have been held as to the affinities of Dinotherium is given by
Stefanescu in his paper on Dinotherium gigantissimum (Anuarulu
Museulni de Geologia, ete., Bukarest, 1894, p. 126). It is now

voposed to give a short description of the skull as it now is,
followed by some discussion of the reasons for its peculiarities
and of the relationships of Dinotherium to the other Proboscidea.

The occipital condyles (0.c.) are very large and prominent ;
ventrally and laterally they are limited by a deep groove, but
dorsally they seem to pass uninterruptedly into the occipital surface
(oc.s.). The articular surface of the condyles from above down-
wards makes rather more than a semicircle, so that the range of
movement of the head up and down was extensive. Ventrally the
condyles are separated by a broad, deep notch, in front of which
the basioccipital runs downwards and forwards as a broad ridge,
narrowing a little towards its junction with the basisphenoid, at
which point. however, there is a prominence. The basisphenoid
is not quite in the same straight lineas the basioccipital, but seems
to have been directed a little more upwards. The basis cranii as
a whole slopes rather steeply downwards, making an angle of about
135 degrees with the plane of the teeth and the posterior part of
the palate, which is regarded as being horizontal. From the outer
angle of each condyle a ridge (7.) runs outwards and a little
upwards, terminating in the outer angle of the squamosal (a.s.),
which projects outwards far beyond the level of the zygomatic
processes. These ridges, which towards their outer ends rise into
thick and prominent crests, separate the upper occipital surface

SKULL OF DINOTHERIUM GIGANTEUM, 527

from the large post-tympanic flanges (pé.fl.), which appear to be
formed partly by the exoccipitals and partly by the squamosal,
though no suture can be seen; the angle between the post-
tympanic flanges and the upper occipital surface is about 130
degrees. The ventral border of the flanges is gently convex, and
is separated from the condyles on the inner side by a deep notch,
which is partly occupied by a tongue-like process of bone,
apparently the paroccipital process (p.p.) of the exoccipital. The
anterior face of the flanges is concave from side to side, and is
separated superiorly from the greatly elongated glenoid surface
(gl.) for the mandible by a shallow transverse channel. Above

Text-figure 1.

i
1

a @

fl {|
fi n )

Vien)
( y

GAN) )y¢f\*
rong | wey

Hig

——=

Skull of Dinotherium giganteum.
From right side: about js nat. size.
af., antorbital foramen; a/.c., posterior opening of alisphenoid canal; ea.n., external
nares; /.c., lambdoidal crest; m.s., depression for muscle attachment on
preorbital prominence; o.c., occipital condyle; oc.s., occipital surface; o7b.,
orbit ; or.r., orbital ridge; pmx., premaxilla; p.orb., post-orbital process of
frontal; pt.fl., post-tympanic flange; zyg., zygomatic process of squamosal.

the transverse ridges just described the occipital surface slopes
forwards to a remarkable degree, giving the skull a curious
resemblance, in this region, to the skull of some types of Whales.
-e. g. Balenopiera. The angle between the occipital surface and
the basis cranii is about 535 degrees, so that the occiput is nearly
parallel to the palatal surface. This extraordinary forward
inclination of the occiput evidently limits the size of the cranial
cavity, and de Blainville suggested that possibly the occipital
surface had been crushed down towards the floor of the skull.
Careful examination of the specimen, however, does not seem to
support this idea, and it seems probable that, remarkable as the

Proc. Zoou. Soc.—1921, No. XXXVI, 36

528 DR. C, W. ANDREWS ON THE

structure of this region of the skull may appear, it represents
nearly the condition in the living animal, In other Proboscidea
the occiput slopes strongly forwards, but this inclination is
masked, in the adult at least, by the enormous development of
cellular bone that takes place in this region, except over the area
of the insertion of the ligamentum nuche, which occupies in
consequence a deep depression.

In young individuals in which this development of the bone
has not taken place, it can be seen that the occiput is inclined to
the basis cranil at an angle about equal to that occurring in
Dinotherium. Thus, in a figure of a median longitudinal section
of the skull of a young African Elephant given by Flower
(‘ Osteology of the Mammalia,’ ed. 2, p. 181, fig. 59), the somewhat
convex occipital surface is inclined to the basis cranii at an angle
of about 40 degrees—that is, at a more acute angle than in Dino-
therium. In the skulls of two very young Indian Elephants this
angle was about 50 degrees, and in Paleomastodon, in which in the
adult the spongy bone is relatively little developed, the angle is
only about 35-40 degrees in the middle line. Thus it appears
that the forward slope of the occiput relatively to the basis cranii
is not really greater in Dinotheriwm than in the later Proboscidea,
but only appears so because, when the palate and tooth series in
Dinotherium is placed horizontally, the basis crani slopes steeply
downwards instead of being nearly horizontal.

In Dinotherium no development of cellular bone seems to have
taken place, the necessary surface for the attachment of the
muscles supporting the heavy head being in this case supplied by
the great widening of the occipital surface, which is carried out
laterally on to the lateral extensions of the squamosals (a.s.). The
pit for the nuchal ligament (7./.) is both wide and deep: behind
it the occiput is convex from before backwards, while at the sides
it is gently convex in all directions. Anteriorly the occiput is
bounded by the transverse portion of the lambdoidal crest (U.c.),
which is slightly convex forwards: in front of this the parieto-
frontal surface makes an angle cf about 140 degrees with the
occiput. Laterally the lambdoidal crest reaches the upper border
of the temporal fossa at about its middle point, and thence is
continued downwards and backwards along its border to the
lateral processes of the squamosal. The lateral portion of the
occiput thus bounded, projects far over the enormous temporal
fossa, forming a kind of roof to its posterior portion. The fronto-
parietal region of the roof is short from before backwards and in |
its middle portion is flat or slightly concave. It first narrows a
little, and then widens out and slopes down towards the post-
orbital (p.orb.) processes, which are presumably borne by the
frontals though no sutures are visible. A little in front of the
level of the post-orbital processes the fronto-parietal region of the
roof is bounded by a thickened border, which is concave forwards
and forms the upper edge of the remarkably wide nasal opening
(er.n.), There is no trace of the nasal bones; they may have been

SKULL OF DINOTHERIUM GIGANTEUM. 529

lost, but in any case do not seem to have formed any median pro-
jection over the nasal opening. The maxilla (ma.), which formed
the sides of the nasal opening at least in part, is a very massive
and greatly developed bone. Externally to its tooth-bearing
and palatal region it is produced laterally into a shelf of bone
the hinder border of which is opposite the hinder lobe of the
first molar. Posteriorly it forms the floor of the orbit, and no

Text-figure 2,

~
SS
SS
SSS
= ———————

\ \ S

Skull of Dinotherinm giganteum.
From above: about 7s nat. size.

a.s., outer angle of squamosal; ev.n., external nares; jug., jugal; U.c., lambdoidal
crest; 2.f., nuchal fossa: 0.¢., occipital condyles; oc.s:, occipital surface;
p.orb., postorbital process of frontal; pt,fl., post-tympanie flange; 7., ridge
from outer angle of squamosal.

doubt joined the jugal, on which the lower post-orbital process

is borne. In front of the orbit the maxilla forms a great mass of

bone, terminating laterally im a rugose somewhat concave surface

for the attachment of a muscle (m.s.), probably the maxillo-
36*

530 DR. C. W. ANDREWS ON THE

labialis superior. The Jachrymal bone perhaps took part in
the formation of this pre-orbital mass, but its limits cannot
be determined. The very large antorbital foramen (a,f.) opens
on the lower surface of the maxilla, about mid-way between its
prominent pre- orbital boss aud the premolar teeth. The point of
union of the maxille with the premaxillz is probably marked by

Text-figure 3.

Cie Wed te i
i

tip. Phe Oe

Skull of Dinotherium giganteun.
From below: about 7's nat. size.

a.f., anturbital foramen; ad.c., posterior opening of alisphenoid canal; a.s., outer
angle of squamosal; aw. external auditory meatus; f.l.p., ? foramen
lacerum posterius; j., jugal; g/., glenoid surface for mandible; m.8, third
molar; m«., maxilla: oe.c., occipital condyles; pima., premaxilla ; pé.fl., post-
tympanic flange; ¢.6., Ptympanie bulla; zyg., zygomatic process of the
squamosal.

the sudden narrowing of the snout about 40 centimetres in front
of the orbit. The suture hetween the two bones can be traced
for a short distance on the palatal surface.

SKULL OF DINOTHERIUM GIGANTEUM. Dol

The premaxille (pme.), though narrower than the maxille, ave
still very massive and widely expanded. ‘Their upper surface is
concave from side to side, the lower convex in the same direction.
In the mid-ventral line they are separated in the actual skull by
a cleft, probably the result of distortion through crushing: in the
east, which seems to have been made before the skull had been
broken, there is only a deep groove. From the posterior angles of
this groove a pair of ridges run back on the palatal surface to the
anterior end of the tooth series.

The anterior ends of the premaxille are thickened, and terminate
in a nearly flat surface in which there are some irregular asym-
metrically arranged pits but no real trace of any incisor alveoli,
though it seems not unlikely that incisors will be found to have
been present in the young animal. The anterior portion of the
skull in front of the tooth series is curved downwards, its palatal
surface approximately following the curve of the downturned
mandibular symphysis.

The structure of the basal region of the skull is not very clear,
probably in consequence of the crushing and fracturing it has
undergone. On either side of the basioccipital there is a fairly
weil-developed auditory bulla (¢.b.). External to this is a strong
erest of bone, terminating posteriorly at the process described
above as the paroccipital (p.p.):; Anteriorly it runs inwards to
the sides of the basisphenoid. This crest, which seems—at least in
front—to be formed by the pterygoid, turns downwards anteriorly,
and forms the border of the narrow opening of the posterior
nares (p.n.): ventrally it terminates just behind the tooth series
in a boss of bone which forms the posterior angle of the hard
palate. Dorsal to this boss there is a deep fossa, presumably
in the palatine, passing upwards to the hinder opening of the ali-
sphenoid canal (al.c.). Behind and a little to the outer side of this
opening is the inner end of the extraordinarily elongated glenoid
surface (g/.) for the mandible. This surface is very narrow
from before backwards and gently convex in the same direction:
posteriorly it is bounded by the depression separating it from the
post-tympanic flange. Its outer end is immediately behind the
base of the zygomatic process (zyg.) of the squamosal, and its
anterior border, at least in its outer half, abuts on the temporal
fossa. The auditory opening (aw.o.) is opposite the outer border
of the zygomatic process: the external auditory meatus is greatly
elongated, and its floor is formed by the roof of the channel
behind the glenoid surface. Above the auditory opening the
massive lateral angle of the squamosal (a.s.) projects outwards some
fourteen or fifteen centimetres. The structure of a bony laby-
rinth supposed to belong to Dinotheriwm has been described and
figured by Claudius (“Das Gehorlabyrinth von Dinotheriwm
giganteum,” Paleontographica, vol. xiii. (1864-66) p. 65).

The opening of the optic foramen is indicated by a depression,
from the upper edge of which a downwardly deflected ridge (o07.7".)
runs upwards and forwards across the side wall of the skull and

532 DR. C. W. ANDREWS ON THE

terminates in front in the post-orbital process (p.orb.). Towards its
anterior end it becomes very prominent, sharply delimiting the
eye-socket above. This ridge seems to be very characteristic of
the Proboscidea: it occurs in Meritheriwm and probably in all
the members of the group.

The skull of Dinotheriwm is, in many respects, one of the most
remarkable known. Although fundamentally its characters are
clearly Proboscidean, nevertheless it differs widely from the skulls
of the other members of the group and, indeed, in some respects
from that of any other mammal. In the true Elephants and

Text-figure 4.

Skull of Dinotherium giganteum.
From behind: about 33 nat. size.
a.s., outer angle of squamosal; fm., foramen magnum; 2.f., nuchal fossa; 0.¢.,
occipital condyle; p.., posterior nares; p.p., paroccipital process; ptf.
post-tympanic flange; 7., ridge from outer angle of squamosal.

Mastodens the peculiar form of the skull is mainly due to the
enormous development of cellular bone in the occipital region,
increasing the area available for the attachment of the muscles
necessary for the support of the heavy trunk and tusks. In
Dinotherium, although the upper tusks are wanting, the trunk,
judging from the large size of the nasal opening, must have been
enormous, and the weight of the head was further increased by
the great deflected mandibular symphysis with its large tusks.
In this case, however, the area for the attachment of the support+
ing muscles was supplied by the widening out of the occipital
surface, which was further increased laterally by the extension
outwards of the squamosals. Little or no cellular bone seems to
have been developed, the occipital surface above the post-tympanic
flanges being nearly flat except for the depression for the nuchal

SKULL OF DINOTHERIUM GIGANTEUM. 533

ligament. This flattening of the occiput, combined with its
forward inclination, must have made it possible for the animal to
move its head up and down through a large arc, a movement
perhaps connected with the use of the downwardly directed lower
tusks. The great width of the proboscis, probably rendered
possible by the absence of upper tusks, led to the widening out
of the skull in the orbital region, producing the shelf-like
projection of the maxiliz above noticed, The great width of the
glenoid surface for the mandible is a peculiarity for which it is
dificult to account, unless it is correlated with the general
widening of this region of the skull.

The skeleton of Dinotheriwm is still very imperfectly known,
but such bones as have been described show that the animal must
have been quite Hlephantine in structure and appearance except
as to its head, the legs being pillar-like and the neck short. A
femur probably associated with the skull above described measured
150cm.in length. The numerous speculations as to the appearance
and habits:of Dinotheriwm have been summarized by de Blainville
and Stefanescu in their works referred to above. Most writers
seem to suppose that the animal was chiefly fluviatile and
aquatic in its habits, but there appears to be no good reason for
believing that it was more so than the Elephants.

The origin of Dinotheriwm is a question of much obscurity.
The genus must have branched off from the main Proboscidean
stem at a very early date, probably originating from an animal in
much the same stage of evolution as Meritherium, the teeth being
still simply bilophodont. The premolar series is more highly
modified than in Meritheriwm, the series being reduced to pm. 3
and pm. 4, the latter being bilophodont like the posterior molars.
This modification of the premolars, like the acquisition of the third
ridge in m.1 and the reduction of the heel of m.3, may have
arisen after the divergence from the majn stock had taken place.
It is interesting to note that the last lower milk molar of
Meritherium shows a strong tendency to trilophodonty, the
posterior ridge being fairly distinct: in Dinotheriwm also this
tooth is trilophodont. If, as seems almost certain, Dinotheriwm
originated from some small comparatively generalized type, it is
interesting to note how, after its separation from the main stock,
the direction of evolution is almost the same as in the latter.
Thus there was a_ general increase in stature, which, being
accompanied by a shortening of the neck, necessitated, as in the
Elephants and Mastodons, the development of an elongated
snout ; so that it may be regarded as certain that Dinothervum
passed through a longirostrine stage comparable to that of
which the beginning is seen in Palwomastodon and the culmina-
tion, so far as the elongation of the lower jaw is concerned, in
Tetrabelodon angustidens. Subsequently the symphysis became
somewhat shortened and was deflected, the lower incisors at the
same time becoming enlarged. It is interesting to notice that a
tendency to a similar type of modification of the mandible occurs

534 ON THE SKULL OF DINOTHERIUM GIGANTEUM.

in that group of T'etrabelodonts to which Professor Osborn gives
the name Rhynchorostrine, the most extreme case being
L. dinotherioides. This deflexion of the mandibular symphysis
led to the development of the flexible free trunk, just as the
shortening of the symphysis did in the main stem, but while in
this the upper incisors tended to increase in size, in the
Dinotheres they disappeared.

The dimensions of the skull above described (taken on the cast)
are i—

. cm.
Length from occipital condyle to tip of premaxille. 121
Length from posterior border of palate to tip of

prem axal les). eee! a OEE ea ie See 84
Width between outer angles of the squamosals ... 90
Width between outer borders of the zygomatic

GOCESSOS sac CUT a tates Arua ei eave Ce ae oa, 74
Width between the openings of the ear ............ 68
Length of glenoid surface from before backwards. 6°3
Width of glenoid surface from side to side......... 24
Width of the occipital COMAVIES Ck A ee ee see 31
Width of skull just in front of the ant-orbital

ROWAN Gini): Pare ps mle ol ioeule Ur asta ae aac 57
Width of the anterior end of the premaxille ...... 3]
Length of molar-premolar series .........0.0,.020000+ AT

Length of molar series

ON THE EXTERNAL CHARACTERS OF LUTRIN A. 535

29. On the External Characters of some Species of Lutrinze
(Otters). By R. I. Pocock, F.R.S., F.Z.8.
[Received April 19, 1921: Read May 24, 1921.]

(Text-figures 19-21.)

CONTENTS.

Page
pis en EU na ani uunrieat, Metis we Pee, ac namareiy cena atelier Ey
Tn JOB vere Wailea SE) gee nee bannices WecmabieGsae tuaanl Gna eo aL
SMe Ce tReet ame te hhh Niels Mieniiane. setnmne ara
The Anus and External Genitalia....................... 542
The Genera of Otters of the Old World............... 542
The Descent of the Lutring .............0.......000000... 544

Since the different species of Otters are remarkably alike in
general appearance, and the published notes on their external
characters, to which Gray in particular paid great attention more
than half a century ago, were taken apparently from dried skins,
the following illustrated descriptions ot the three species, Lutra
lutra, L. maculicollis, and L. cinerea, exhibited during the past
few years in the Zoological Gardens, supplemented by an
examination of the softened skin of a stuffed example of donya
capensis, and followed by some general conclusions with regard to
the systematic status of the species examined and the descent of
the Lutrine, may prove of interest.

The fhinarvwm.—vhe extent to which the rhinarium is covered
with hair has frequently been used since Gray’s time as one of
the best external characters for distinguishing the species of
Otters.

In Lutra lutra it is entirely naked and a little wider than
long ; but judging from descriptions is variable in shape. Gray,
for example, described the lower edges as straight and the upper
as “rather produced and angular.” Blanford corroborates this,
saying that the ‘‘ upper edge is not straight but projects in the
middle and is coneave on each side, running up considerably to
the hinder edge of the nostril.” Miller, on the contrary, says the
upper margin is “strongly convex at middle, the lower slightly
so.” In the specimen here figured the upper edge agrees precisely
with Blanford’s description, and the lower margin is angularly
produced. The nostrils are widely separated and there is no
philtrum, the upper lip being continuously hairy and about as
deep as the height of the rhinarium. This species has a com-
paratively long muzzle, as shown by the skull, and the plane of
the rhinarium is inclined upwards and forwards at an angle of
about 60° or less.

In the Asiatic small-clawed Otter (Lutra cinerea), a short-

536

AHMaQeaeoanrPr

MR. R. I. POCOCK ON THE EXTERNAL

Text-figure 19.

Nan WEN
ff MM dN AN INS
TASS

. Head of Amblonyz cinerea.
. Khinarium and upper lip of the same from the front.

Rhinarium of the same trom above.

. Head of Hydrictis maculicoliis.

. Rhinarium and lip of the same from the front.

. Rhinarium of the same from above.

- Rhinarium of Aonya capersis from the front (skin dried and softened),
. Rhinarium of Lutra lutra from the front.

. Rhinarium of Lutra lutra trom the side.

. Har of Lutra lutra.

All figures, except K, x4.

CHARACTERS OF SOME SPECIES OF LUTRINA. 537

jawed form, the plane of the rhinarium is almost vertical.
Otherwise it closely resembles that of the European species,
except that its upper edge is straight transversely between the
posterior edges of the nostrils and its inferior edge less produced.

So far asit is possible to judge on the dried skin, the rhinarium
of the African clawless Otter (Aonyx capensis) resembles that of
the Indian form Z. cinerea, which is often associated with it.

The other well-known African Otter (Lutra maculicollis) has a
very different rhinarium, although as in JZ. cinerea its plane is
vertical. It is very much reduced in size, being overgrown with
hair to such an extent that little more than the edges of the
nostrils are naked. The inferior border is quite straight and the
infranarial portion on each side is shallow, while its upper edge
is sinuously concave backwards, owing to the advancement of the
hair in a curved line on each side from the posterior margin of
the nostril to the middle line, where it nearly divides the rhinarium
into two portions. From this it results that the middle of the
rhinarium is less than one-fourth of the depth of the upper lip.

The Facial Vibrisse.—The facial vibrisse in Otters are, as a
rule, exceedingly coarse and numerous as in most predatory
aquatic mammals. Two points to be noted, however, are that
the mystacials, as compared with those of normal terrestrial
mamnials, are relatively short, not longer that is to say than
the genals, and that the gena! tufts, especially the inferior, are
exceptionally well developed, the vibrissee being numerous and
long. The material examined does not warrant the opinion that
Lutra lutra, L. cinerea, and L. maculicollis differ in any way
with respect to their vibrisse ; but the vibrissx of donyx capensis
are decidedly less stout and stiff, especially perhaps those forming
the upper genal tuft *.

The Har.—The external ear in Otters is always small and oval,
and comparatively simple in structure, the bursa being absent.
In the European species (Zatra lutra) the tragus and antitragus
are tolerably well developed, with a well-marked notch between
them, and just inside the antitragus there is a strong ridge-like
thickening forming the posterior rim of the cavity. The supra-
tragus is a shelf-like ridge with well-developed fleshy lobe.

In the Indian small-clawed Otter (Z. cinerea) the ear is very
like that of Z. lutra, but relatively smaller, the cavity not
extending so high above the supratragus; and the tragus and
antitragus are inconspicuous. On the dried skin I could find
nothing distinctive about the ear of Aonyx capensis. In L, macu-
licollis the ear is simpler than in L. cinerea. The inferior edge of
the cavity shows no trace of a notch, the tragal and antitragal
thickenings being altogether suppressed, and the supratragus is
represented merely by a rounded tubercle corresponding to the
fleshy lobe of this thickening seen in L. lutra and cinerea.

* Hinton (Ann. Mag. Nat. Hist. (9) vii. p. 195, 1921) has recently proposed the
new generic name Paraonyx for two clawless Otters from the Congo which differ,
amongst other things, from donyx in the complete suppression of the superciliary and
upper genal tufts.

538

MR. BR. I, POCOCK ON THE EXTERNAL

Text-figure 20.

Be aaah
NESGNAID Ween
we Nate

Nac

a 1
aa \
DA ee)

Ni Olas walle V39,\ 4,
Ny Sahel Pap 1 [ts

% Gali (ee rhe yi
ME ES

A. Right fore foot of Dutra lutra.
B. Right hind foot of Dutra lutra.
C. Right fore foot of Hydrictis maculicollis.
D, Right hind foot of Hydrictis maculicollis.

CHARACTERS OF SOME SPECIES OF LUTRINA. 539

The Feet *-—The feet of the Lutrine or Otters are so vari-
able that it is hardly possible to mention a single character in
which they differ from those of the other Mustelidz collectively,
except that the fore paw is always much smaller than the hind.
In some cases the digits are less fully webbed than in any other
genus of the family except Lyncodon ; but in the typical forms
the feet are more fully webbed because the digits are relatively
longer and more widely separable, owing to the greater width
and less emargination of the webs, especially of those joining the
first and second, second and third, and fourth and fifth digits.

In Lutra lutra, for example, the third and fourth digits of the
fore foot are not more widely separated nor more fully webbed
than in Charronia or Gulo; but the webs between the fourth
and fifth, and third and second, and second and first digits extend
with a nearly straight edge between points near the middle of
the margins of the digital pads. The claws are well developed
and project well beyond the tips of the digital pads, and the sole
of the foot is naked back to the proximal margin of the single
large, sometimes nearly hemispherical, sometimes transversely
elongated carpal pad. The radial or internal element of the
carpal pad is not separately distinguishable. There is a short,
naked area between the carpal and the plantar pad; and the
main part of the latter is well developed but narrow, that is to
say about as wide as long, and distinctly three-lobed ; the two
lateral lobes nearly meet behind the median, and the pollical
element is small angl indistinctly defined from the posterior end
of the adjoining lobe. The hair of the wrist extends right down
to the carpal pad. Tt hind foot is markedly larger than the
fore foot owing to the greater elongation of the digits, which
have consequently deeper and wider webs. The plantar pad is
remarkably modified. * Its median lobe is elongated and some-
what heart-shaped; the external 1be, defined by a deep groove,
extends far back behind the median. in contact throughout its
leneth with the inner lobe, which is almost as long but has its
free edge emarginate, the proximal or pollical element being
large. The three main lobes just described are defined from
each other by a Y-shaped groove. The hair of the metatarsus
extends down to the proximal end of the plantar pad, and there
is no trace of metatarsal pad.

In Lutra maculicollis the digits are relatively longer than in
L. lutra and the webs are better developed, extending to the
distal end of the digital pads on their inner sides. The plantar
pad of the fore foot is more nor mally shaped, being considerably
wider than long and four-lobed., The median lobe is not narrowed
behind, the _ posterior ends of the main lateral lobes are widely
separated, and the pollical lobe is clearly defined. The single
ulnar carpal pad is much smaller than in L. lutra; and apart
from the pads which are naked, the underside of the foot is
covered with fine, short scattered hairs. The hind foot differs

* The carpal vibrisse are present in all the specimens described.

MR. R. I, POCOCK ON THE EXTERNAL

Text-figure 21,

A. Right fore foot of Aonyx capensis.

B. Right hind foot of Aonyx capensis.

C. Tip of digit with minute claw of hind foot of same.
D. Right fore foot of Amblonya cinereus.

E. Right hind foot of Amblonyx cinereus.

F. External aspect of fore foot of same, showing claws.

CHARACTERS OF SOME SPECIES OF LUTRINA. 54]

from that of ZL. lutra in similar respects, except that the plantar
pad is very poorly defined, its three lobes gradually fading away
into the scantily hairy area of the metatarsus at the proximal
end of the digits. The median lobe is tolerably broad in front
and tapering behind, the external lateral lobe is scarcely differen-
tiated from the base of the fifth digit, the internal lateral is a
mere thickened ridge of skin covering the base of the second
digit, and there is no trace of the internal or hallucal lobe. The
skin of the metatarsus close to the proximal ends of the digits
and of the plantar pad is scantily hairy, like the lower side of the
webs and digits; but above it is thickly haired as in Z. lutra.

In Lutra cinerea the feet differ from those of Lutra lutra and
I. maculicollis in being narrower owing to the shortness of the
digits and the comparative shallowness and narrowness of the
webs, which reach only to the proximal end of the digital pads,
those joining the second and third, and fourth and fifth digits
being decidedly emarginate, although not quite to the same
extent as in cursorial and fossorial Mustelide. The claws, more-
over, are very short, almost erect, and quite concealed when the
feet are examined from below. The plantar pads also are better
developed and more normal in shape, being longer than wide,
subsymmetrical and four-lobed, the inner main lobe being pro-
longed by the pollical and hallucal elements. In the hind foot
the outer main lobe is distinctly divided into an anterior and
posterior portion by a transverse groove and almost in contact in
the middle line with the internal lateral lobe. Above this pad
the metatarsal area is mostly covered with naked wrinkled skin,
the heel only being hairy. In the fore foot the carpal pad is
relatively as large as in L. lutra, but it is defined above by a
naked strip of skin over which the hair of the wrist does not
extend. The lower sides of the webs and the grooves between
the pads are beset with small hairs, but they are shorter and fewer
than in L. maculicollis.

In Aonyx capensis *, the African clawless Otter, the hind foot
is very like that of L. cinerea in the development of the digits
and webs; but the hair on the metatarsal area extends farther
towards the plantar pad. The fore foot also is tolerably similar
to that of ZL. cinerea apart from the great difference that the
webs between the digits reach only to the distal end of the first
phalange, the ends of the digits being free. This terminal
freedom of the digits from webbing is not found in any of the
cursorial or fossorial Mustelide. A. further peculiarity of this
Otter is the suppression of the claws on the fore foot and on the
first, second, and fifth digits of the hind foot. On the third and
fourth digits of the hind foot they are merely retained as small
flattish nails.

* Tam greatly indebted to Mr. Fitzsimons, of the Port Elizabeth Museum, for very
kindly sending me at my special request an example of this Otter from the collection
under his charge. It was a mounted specimen, and the method of preservation made
it impossible for me to ascertain the precise shape of the plantar and carpal pads.

542 MR. R. I. POCOCK ON THE EXTERNAL

A further stage in the disappearance of the webs is shown in
Paraonyz, which is described by Hinton as having unwebbed
digits on the fore foot and the digits of the hind foot webbed
to the base of the second phalange, apparently as in the fore
foot of Aonyx.

Anus and Haternal Genitalia.—About the anus and external
genitalia there is very little to be said, since they closely resemble
those of such typical Mustelide as Martes in having the anus
encircled and the perinzum covered with hair and the two normal
anal glands present, with the scent of the secretion compara-
tively inoffensive. The penis of Lutra lutra is moderately long,
and is supported by a stout, nearly straight baculum* about
60 mm. long, convex above, flatter and sometimes faintly grooved
below, and gradually narrowed from the base up to the apex,
where it expands into two short, deeply cleft, thick condyle-like
branches, curved downwards. The branches are not alike, the
right being a little thinner and Jess curved than the left. The
urethra runs along the underside of the baculum, which is in-
vested in soft vascular distensible tissue, and opens in the middle
line below, just behind the condyle-like thickenings of the
baculum.

The baculum of Z. cinerea is very like that of L. lutra, but is
of more even width throughout and the two terminal branches
are not curved downwards.

The Genera of Otters of the Old World.

Apart from the incidental mention of Paraonya, the Otters
discussed in the foregoing pages have been referred to the two
genera, Lutra and Aonyx. Butif the nomenclature of this group
is to be brought into line with that of other groups of Mammals,
as the publication of Paraonyx seems to demand, it is certain
that several generic and subgeneric terms long ago proposed but
never as yet adopted will have to be restored to use. I have
neither the material nor the time to go into the question beyond
the limits of the species exhibited in the Zoological Gardens, of
which skulls or skins, or both, are in the Society’s possession.

Otters unfortunately, although so w.dely distributed, are very
seldom in the live animal market, and it does not appear that
more than the following species have been exhibited in the
Gardens :—Lutra lutra, L. barang (entered as nair), L. cinerea,
L. maculicollis, and L. platensis.

Each of the four exotic species has been severed from Lutra,
and received.a generic or subgeneric name. About platensis,
named Nutria by Gray, I have nothing to say, the specimen
having been sent unexamined to the British Museum. The

Oriental species determined by Thomas as barang was referred «

by Gray to the subgenus Lutrogale under the name monticola.
I have no personal knowledge of the external characters ; but the

* Described by Blainville, Pohl, and other authors.

CHARACTERS OF SOME SPECIES OF LUTRINA. 543

skull appears to me to differ from that.of Zutra in characters
which by the modern standard must be regarded as of generic
value. It is much higher; the much larger orbit is thrust so far
forwards that its anterior rim is above the front edge of pm. 3,
thus greatly reducing the length of the muzzle, lengthening the
floor of the orbit, bringing the frontal postorbital process over
the middle of the upper carnassial, and lengthening the area of
the cranium between those processes and the “ waist.” Added
to this, the anterior nares are much less sloped, the infraorbital
foramen is almost concealed in profile view by its upper bar, and
the teeth are out of all proportion larger, ete.

The long-clawed African Otter (maculicollis), the type of
Gray’s genus Hydrogale, difters from typical Lutra in the reduction
of the rhinarium and the simplification of the ear; in the rela-
tively large, more fully webbed and hairy-soled feet with reduced
plantar pads; and in many cranial characters, especially the
shortness of the muzzle, length of the orbital floor, and the
generally immature aspect of the skull owing to the feeble
development of constrictions, crests, and prominences.

Since the sum of the differences appear to me to be of generic
value, I propose the name Hydrictis to replace Gray’s twice
preoccupied name Hydrogale. !

In defence of the adoption of the name Aonyx for the African
small-clawed Otter I need say nothing, since the genus appears to
be admitted as valid.

The Oriental small-clawed Otter (cinerea) was named Amblonya
by Rafinesque in 1832. By modern writers it has been referred
either to donyx or to Lutra, despite its differences from both.
Tt resembles Lutra in the rhinarium, facial vibrisse, and the ear,
but differs in the structure of the feet, which are narrower
owing to the shorter digits being joimed by narrower webs,
especially between the second and third and fourth and fifth
digits, none of the webs extending beyond the proximal ends of
the digital pads. The claws also are short and nearly erect or
sometimes, according to Blanford, absent*; and only the
calcaneal area of the hind foot is hairy. The skull also is very
different from that of Zutra, and much resembles that of Zutro-
gale barang in the character of the muzzle and the size of the
teeth, but is altogether much shorter from the frontal postorbital
processes and the molars back to the occipital crest and condyles ;
the interorbital region is wider, the waist much shorter, and the
orbital floor less extensive owing to the shortening of the tooth
row.

In external characters Amblonyx differs from Aonya, at all
events, in the extension of the webbing of the fore feet to the
digital pads and in the normal coarseness of the facial vibrisse.

* The abbreviation of the claws of the front foot is accompanied in this Otter by
extreme delicacy of the sense of touch in this extremity. I have seen one of these
animals manipulating and playing with a marble in a manner recalling that of a
conjuror juggling with a cricket-ball.

Proc, Zoou. Soc.—1921, No, XXX VIT, 37

544 MR. R. I. POCOCK ON THE EXTERNAL

Thus four genera of Otters occur in Africa: Latra lutra,
north of the Sahara, Hydrogale maculicollis and Aonyx capensis,
south of the Sahara, and Paraonyx congicus, etc., in the Congo
area; and three in India, where they seem to be widely dis-
tributed: Lutra lutra or allied species; Lutrogale barang; and
Amblonyx cinerea *.

Omitting Lutrogale barang, whose external characters are only
known to me from descriptions which supply no evidence of
difference from Lutra lutra, these Otters may be distinguished as
foliows :—

a. Fore feet with digits unwebbed ; hind feet webbed to proximal end
of second peaeteys pete aal and nD EEE genal vibrisse sup-
pressed (Hinton) . y: FI REEE CEMA e ECO CON is

a’. Fore feet with ahete Hee leant Halearentede ond feet webbed at
least to proximal end of third phalange; ‘superciliary and upper
genal vibrisse retained.

b. Fore feet webbed to proximal end of second phalange; facial
vibrisse comparatively slender and soft . bapeanes Aonyx.

b’. Fore feet webbed at least to proximal oil of weal ‘Gielianees
facial vibrissee comparatively stout and stiff.

c. Claws small or vestigial; digits short, webs narrow, reaching to
proximal end of digital pads; hairs on metatarsus restricted

to calcaneum ; skull short, especially behind zygomatic arches. Amblonya.
ce’. Claws long, digits longer, webs wider, reaching at least half-
way along digital pads; metatarsus hairy to plantar pad ;

skull long behind zygomatic arch.

d. Digits longer, more fully webbed, plantar pads voorly
developed, especially on hind foot, carpal pad small, webs
hairy below; rhinarium nearly covered with hair; ear
simplified; skull with muzzle and Hebert fossa

JNORH “ahheacoser . Hydrictis.

. Digits short, hes fay, Ww seueat inte a ae Gell Pas Hopedi
carpal pad large, webs naked below; rhinarium naked ;
ear with well-developed tragus, antitragus, and supra-
tragus ; skull with muzzle and mesopterygoid fossa long... Dutra.

The Descent of the Lutrine.

So far as I am aware, Miller is the only author who has
attempted to affiliate the Lutrine definitely with one of the other
subfamilies of Mustelide. ‘This is embodied in the suggestion
that they “appear to be essentially badgers modified for semi-
aquatic life.” Even granting the widest possible meaning to the
word “badgers,” I am quite unable to agree with this view.
Miller, I think, placed too much reliance upon the teeth; but
it does not appear to me that the teeth of Lutra lutra, perhaps
the least modified of all the Otters, are more like those of Meles
than of Martes. Again, in the Otters the shape of the skull, with
its flat or slightly arched crown, its short muzzle, long back, and
forwardly set posterior palatine foramina, forcibly recalls that of
Mustela or Grison, and is quite unlike that of J/eles, or indeed of
any of the genera usually referred to the Meline, in all of which
the skull has a long jaw and short back and, except in the

* T have not examined the type of Gray’s genus Barangia, namely sumatrana,
which is, I believe, the only other admitted species of Old World Otter,

ee ee ee

Se a

Se Ss ee ee ee a ee

CHARACTERS OF SOME SPECIES OF LUTRIN®. 545

Skunks, the palatine foramina in question placed on the maxillo-
palatine suture.

Both Mustela and Grison are predatory terrestrial forms; and
on @ priort grounds it seems to be much more probable that the
Otters are the descendants, modified for aquatic life, of some
group of active, predatory terrestrial Mustelide than of com-
paratively inactive, specialized diggers like the Badgers. This
view is supported by the long, slender, flexible body, the long
tail, the broadly webbed, well spread digits, small claws and small
front paws. These characters are found associated only in the
genera currently referred to the Musteline, but not in any of
the so-called Melinz, which have a short thick body, short tail,
closely tied toes, powerful claws, especially on the fore feet, and
the latter are larger than the hind feet.

Another point worth considering is the presence in a few
unrelated species—i.e. Lutra canadensis, according to Gray, and in
Hydrictis maculicollis and Amblonyx cinerea—of hairs upon the
lower side of the webs of the feet. It seems to me to be difficult
to explain their occurrence on the hypothesis of the descent of
the Otters from naked footed Meline burrowers. The hairs seem
to be quite functionless, and they have all the appearance of
harmless vestigial structures suggesting the descent of Otters
from hairy-footed forms. And since the Martens, Wolverenes,
and Weasels, in the broad sense of the word, are the only members
of the Mustelide, apart from the Otters just mentioned, which
have hair upon the webs below, these hairs may be regarded as
an additional item of evidence pointing to those active predatory
Mustelide as standing nearest to the extinct ancestors of the
Otters.

Tt is not of course suggested that the Otters are the descendants
of any of the Martens or Weasels known to us, but that they are
an offshoot from that line of descent; and I visualize their ancestor
as an active terrestrial, predatory Mustelid with a well-developed
tail and feet with the longish digits provided with short, sharp
claws and united by wide webs, hairy below but not so hairy as
in Martes or Mustela, and with the pads probably as large as in
Charronia, with the skull shaped like that of Mustela or G'rison,
the auditory bulla like that of MJartes and also the dentition,
except that the upper carnassial had a larger inner lobe such as
is seen in Grison.

The enlargement of the hind feet as compared with the fore
feet, the thickening of the facial vibrisse, with the correlated
expansion of the upper lip and enlargement of the suborbital
foramen, the reduction and simplification of the ear, the texture
of the fur, and the muscular development of the tail, are special
adaptions to aquatic life. Similarly, the enlargement of the back
teeth, recalling in a measure that of the Melinz, may be looked
upon as an adaptation for breaking up fish-bones and rendering
them harmless to swallow.

The explanation of the great difference in dentition between

37*

546 ON THE EXTERNAL CHARACTERS OF LUTRINA.

the Otters and the Seals may be explained in this way. Otters
masticate their fish and require broad, flat-crowned teeth for the

purpose. Seals swallow them entire, and the teeth are modified
for catching and holding the slippery prey. ;

[Nore.] After this paper was written, Mr. Thomas drew my
attention to the elaborate monograph of the Lutrinz published
by Pohle in November 1920 (Arch. f. Naturg. 1919, pt. 9, pp. 1-
247). So far as the species I have discussed are concerned, Pohle’s
systematic results, based upon skulls, teeth, and dried skins, differ
im two points only from mine. While admitting the genera
Amblony« and Aonyax* as distinct from Lutra, he granted Latro-
gale barang only subgeneric rank under Lutra, and dismissed
maculicollis and two related species as ‘‘ Die maculicollis-Gruppe”
of Lutrat. Since the properly constituted subgenera and
“oyoups” of species of one generation of workers almost
invariably become the genera of the next, the fulfilment of their
destiny in that respect by Lutrogale and the maculicollis-group
may, I think, be confidently predicted.

I see no reason, therefore, for making any change in the text
of this communication.

* Pohle missed Paraonyx although he had the description of congica before him
and apparently plenty of material of the genus in the skins and skulls of the species
he described as Aonya microdon. This remark, however, is not intended as any
disparagement of Pohle’s work, which appears to me to be exceptionally good. In
addition to the genera and species, it deals with geographical distribution, cranial
changes with growth, milk-dentition, and so forth.

+ Similarly, Pohle established “ Die Swmatrana-Gruppe” [= Barangia Gray} and
“Die platensis-Gruppe” for the American Otters canadensis, annectens, enudris,
platensis, provocax and felina—a group tor which the name Lontra Gray appears
to be the oldest, with Latawina and Nutria as synonyms.

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COMPARATIVE ANATOMY OF THE KOALA
(PHASCOLARCTOS)

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COMPARATIVE ANATOMY OF THE KOALA
(PHASCOLARCTOS) :

ON THE ANATOMY OF THE KOALA AND VULPINE PHALANGER. 547

30. The Comparative Anatomy of the Koala (Phascolarctos
cinereus) and Vulpine Phalanger (Zrichosurus vulpe-
cula), By CuHartes F. Sonntag, M.D., F.Z.58.,
Anatomist to the Society.

[Received May 7, 1921: Read June 7, 1921.]
(Plates V., VI.; Text-figures 53-63.)

The observations recorded here are based on the examination
of an adult male Zrichosurus vulpecula which died in the Society’s
Gardens, and a young female Phascolarctos cinereus which
Mr. A. F. Richardson, of Hull, sent to us for examination after
death.

The measurements of the Koala are as follows :—

Centimetres.
Length from tip of nose to root of tail......... 50
i CUNT AVELICG) po AU" cet pre aa i aR Pee Sa he RO Ae 12°5
Oly tami a Pans er AM kas se ee alts 2°5
Greatest, widtheotimead, 22, e204. 0Gasecceeeacec es 75

Length of fore limb (arm, 9°5; forearm, 13:5;

TAVATATES AO) acs eee seed ashe Wri kee HSER Sa. e 5:0 275
Length of hind limb (thigh, 11°5; leg, 12;
[OSIRIS ids. d Sus BO ROSE AnH Oey cnr ar ries ee eee 28:5

The external characters presented nothing new.

Sir Everard Home, who wrote the first account of the internal
anatomy of the Koala, in 1808, described the character of the
glandular patch of the lesser curvature of the stomach, and the
prostate gland which resembles that of the Kangaroo; he also
recorded the absence of the patella (5). Highteen years later
Dr. Robert’ Knox, of Edinburgh, described the long cecum for
the first time, and named the Koala the ‘‘ Wombat of Flinders ”
in honour of its discoverer. He also criticised the writings of
Home, Illiger, Cuvier, Desmarest, and De Blainville. Since
then accounts of the anatomy of the entire animal have been
written by Martin (8), Forbes (4), and Young (18), and indi-
vidual organs have been described by Flower (3), Owen (11),
Macalister (7), and Chalmers Mitchell (9). The histology of
different parts has been described by Oppel (10) and Braus (1).

THe ANATOMY OF THE Heap AND NECK.
Phascolarclos cinereus.

When the skin of the head is removed it is seen how the
platysma myoides, muscles of mastication, auricular muscles, and
those round the orbit and mouth are strongly developed.

The platysma myoides avises posteriorly from a strong panni-
culus carnosus. It has a clearly-defined line of attachment to

548 DR. C. F. SONNTAG ON THE COMPARATIVE ANATOMY

the lips, and some of its fibres fuse with those of the muscles
of the angle of the mouth and orbit. It covers the anterior part
of the masseter muscle and the facial vessels. It has been divided
along the lines 7.7. in text-fig. 53.

The masseter muscle (text-fig. 53) has been described by
Macalister. It has a tendinous lower border (6), from under
cover of which the facial artery (a) emerges and runs antero-
superiorly ; it gives off an external nasal branch (c) which runs
along the antero-syperior border of the cheek-pouch (CP). The
masseter is crossed by the anterior facial vein (d), transverse
facial artery (f), and Stensen’s Duct (e).

Text-figure 53.

The side of the head of Phascolarctos cinereus.
j.k. posterior auricular vessels; 7. parotid gland. Other letters in text.

Macalister has described the attolens, attrahens, and retrahens
muscles which act on the large auricles.

The anterior facial vein has the usual origin, course, relations,
and branches. It unites with the posterior facial vein on the
surface of the exoccipital process (g) to form the external jugular
vein. No part of it is seen in the interramal space of the
mandible, and the vena transversa (text-fig. 54, 12), which is
long and V-shaped, connects the external jugular veins (11)
instead of the anterior facials; 10 receives lingual, muscular (3,
18), sub-maxillary (21), and a single submental vein (2), but no
anterior jugular vein connects it to the internal jugular vein.

The Parotid Gland is large, and forms a prominent mass in the
face and neck. Running on its surface are the posterior auricular
vessels.

The neck is divided, as usual, into mesial and lateral triangles
by the sterno-mastoid muscle (text-fig. 54, 10), and these have
the usual boundaries. The base of the lateral triangle is formed

OF THE KOALA AND VULPINE PHALANGER. 5A9

by the clavicle (17) as in all Marsupials except the Peramelide,
which have no clavicles.

In the space between the mesial borders of the sternomastoids
there is a thin muscular sheet formed by the mylo-hyoid (1)
anteriorly, the digastrics (9) antero-laterally, and the sterno-
hyoids (13) posteriorly ; the sterno-hyoids have become separated
from the hyoid bone in this specimen, but the whole sheet plays
over it. Anterior to the digastrics are the submaxillary glands (4).

Text-figure 54.

The superficial anatomy of the neck of Phascolarctos cinereus.
5 and 6. anterior and posterior facial veins; 7, 20. exoccipital process; 8. posterior
auricular vein; 11. external jugular vein; 16. supraclavicular nerves;
17. clavicles; 19. masseter. Other numbers in text.

The space between the sheet and the sterno-mastoids is bridged
over by a triangular fascia (14) which covers the common carotid
artery, vago-sympathetic nerve-trunk, and the nerve to the
pretracheal muscles. It does not cover the internal jugular vein,
which is under cover of the sterno-mastoid.

Trichosurus vulpecula (text-fig. 55).

The superficial anatomy differs from that of Phascolarctos in
the following respects :—

1. The platysma is not so well developed. 2. The external
nasal artery does not follow a cheek-pouch. 3. The auricular

550 DR. C. F. SONNTAG ON THE COMPARATIVE ANATOMY

muscles are not so powerful. 4. The vena transversa is horizontal,
and runs between the anterior facial veins. It is connected to
the internal jugular veins (#) and receives two submental veins ;
a small part of it loops into the interramal space. ‘5. The
sterno-hyoid muscles are attached to the hyoid bone. 6. The
superficial sheet between the sternomastoids of Phascolarctos
is absent. Neither in Phascolarctos nor in Trichosurus does the
vena transversa receive laryngeal veins.

Text-figure 59.

The superficial anatomy of the neck of Trichosurus vulpecula.
Numbers as in previous figure.

The deep anatomy of the neck differs greatly in the two
species :—
Phascolarctos cinereus.

When the muscular sheet described above is removed the
conditions shown in text-fig. 56 make their appearance:

The walls of the mouth (/) and pharynx (g) lie immediately
under cover of the muscular sheet, and the following structures
are seen to lie on their external surface :—

1. Two long, narrow strap-like genio-hyoid muscles (a, @’)
running from the symphysis menti to be attached to the hyoid
bone (m) and larynx (/). ‘They cross the lingual arteries (c).
2. The lingual arteries. 3. The hypoglossal nerves (d). 4. The
pharyngeal branches of the vagi (e), which are very tortuous.

OF THE KOALA AND VULPINE PHALANGER. 551

A short interval separates the insertions of the genio-hyoid
and sterno-thyroid muscles (7), which arise from the back of the
sternum and third costal cartilages. At the posterior part of

Text-figure 56.

The deep anatomy of the neck of Phascolarctos cinereus.
J. level of opening of nasal tube into the pharynx; &. thyro-hyal capping
the shaded basi-hyal (m). Other letters in text.

the neck the lateral lobes of the thyroid gland (A. A) project

from under cover of the sterno-thyroids, and extend for a short
distance dorsal to the clavicles,

552 DR. C. F. SONNTAG ON THE COMPARATIVE ANATOMY

Macalister (7) described the styloid group of Biaees as
follows :—“ The styloid group form a single sheet, the hind
fibres of which pass to the pharynx, the middle to the ceratohyal
under the stylohyoid ligament; the anterior pass to the side of
the tongue.”

The digastrics have the usual attachment, and they are united
by muscular fibres to other muscles around.

The stylo-hyoid ligament is loose (text-fig. 56, h). The genio-
hyoids are connected to the digastrics by Hinsemlnn slips (b).

Text-figure 57.

The deep anatomy of the neck of Trichosurus vulpecula.
D. sterno-mastoid; E. digastric; k. hyoid bone; m. common carotid artery;
m. external jugular vein; o,, vena transversa; qg. anterior jugular vein ;
7. s. thyroid and laryngeal arteries. Other letters in text.

The cervical lymphatic glands form two groups. The first,
consisting of a few large giands, is a little posterior to the
prominent alisphenoid bullae The small glands of the second
group are around the great veins at the root of the neck.

The distance from the symphysis menti to the level of the
angles of the mandible is 6 em., from the angles of the mandible
to the hyoid bone 1:2 cm., and from the hyoid bone to the
sternum 6 cm,

OF THE KOALA AND VULPINE PHALANGER. 553

Trichosurus vulpecula (text-fig. 57).

The deep anatomy of the neck differs from that of Phasco-
larctos in the following respects :—1. The walls of the mouth
and pharynx are more deeply placed under muscles. 2. True
hyoglossus muscles are present (B), so the lingual artery (¢) and
hypoglossal nerve (¢) are more deeply placed. 3. The thyroid
gland is applied to the posterior part of the larynx and is far
anterior; it does not pass dorsal to the clavicles. 4. The
alisphenoid bulle are not so prominent. The exoccipital pro-
cesses are not visible. 5. The external jugular vein is formed on
the surface of the sternomastoid muscle.

THe RESPIRATORY ORGANS.

In Phascolarctos the epiglottis is broad and entire, with an
emarginate apex, and each lung contains two lobes; Forbes (4)
described two left and three right lobes. No azygos lobe is
present. In Zvrichoswrus the epiglottis is notched ; the left lung
has two lobes and the right cne has four, of which one is azygos.
The tracheal rings are wide, but deficient dorsally.

The nasal tube of Phascolarctos has no median dorsal ridge,
and does not grip the larynx, whose epiglottis is so large that it
could only be accommodated by considerable stretching of the
orifice of the tube.

In Trichosurus the nasal tube has a median dorsal ridge, and
it gripped the larynx firmly in my specimen.

Sir Everard Home states that the Koala utters a shrill yell
when hungry or hunted, but Owen says that he never heard a
note of any kind from the Phalangers.

THe Moura.

Phascolarctos cinereus.

The Vestibule.—Fleshy lips are connected to the gums by lax
frenums, of which the upper is ‘6 and the lower is 1:2 em. long.
The cheeks are attached to the gums opposite the canine tooth
and posterior border of the premolar of the upper jaw on each
side; between these lie the mouths of the cheek-pouches
(Plate V.,A) already described by Owen (11), Young (13), and
Forbes (4).

Forbes states that “the mucosa of the cheeks is smooth
throughout”; but in my specimen it is covered with rounded
papille, which are all visible to the naked eye. They extend on
to the fauces, and for a short distance along the outer walls of
the cheek-pouches, but they are absent from the lips whose
mucosa is quite smooth. They are not regularly arranged like
the papille of the cheeks of the Ungulata.

554 DR. C. F. SONNTAG ON THE COMPARATIVE ANATOMY

The vestibule and cavum oris communicate through the
diastema anteriorly, and behind the last pairs of molar teeth
posteriorly ; the former of these is wide and the latter narrow.

The Palate (Plate V., A).—The upper incisor and canine teeth
surround a small area on which there are six small tubercles, —
but these do not cover prominences on the premaxille. Imme-
diately behind these lies the first of the nine palatal ridges, which
are most marked anteriorly. Forbes also described nine ridges,
and Young recorded eight. Martin described eight furrows. ‘The
anterior ridges are transverse, and are complete or incomplete,
but some of the posterior ridges are curved and bear small
nodules on their anterior borders.

When the mucous membrane is removed the bony palate is
seen to have a sinuous outline when viewed laterally (text-
fig. 59, HK).

The soft palate is long, smooth, and devoid of a uvula.

The isthinus faucium (text-fig. 59, f) is a circular orifice which
admits the tip of the index finger ; and the fauces form a septum
without the tonsils, which lie far back in the pharynx at a
distance of 6 cm. from the incisor teeth.

Trichosurus vulpecula (Plate V., B).

Cheek-pouches are absent, and the vestibule and mouth com-
municate as in the above. On the palatal area surrounded by
the incisor and canine teeth, which form a more perfect arcade
than in Phascolarctos, there is a median ridge and several small,
hard papillee.

The palatal ridges, of which there are six, are sharper than in
Phascolarcios, and the anterior ones are convex forwards, with
the free edges directed backwards. Between several of the
ridges the mucosa is covered with papille.

The tonsils have the usual situation in the fauces.

Pseudochirus peregrimus (Plate V., C).

There are eight palatal ridges, most prominent anteriorly, and
many papille crowd the mucosa between them. Cheek-pouches
are absent.

Tue Toncve (text-fig. 58).

Martin (8) states that the tongue of Phascolarctos rises abruptly
from a furrow SuLWe wag its base, and the distance from
its root to the epiglottis is $ of an inch. I found, however, that
it is much farther removed, the distance being slightly more than
13 inches, and the mucosa of its dorsum is continuous with that
of the ventral wall of the pharynx (text-fig. 59, A,@).

Measurements :—Total length 5:5 em.; length of the oral

OF THE KOALA AND VULPINE PHALANGER. 555

part 5 em.; length of the pharyngeal part ‘5 cm.; greatest
width 15cm. The slightly spatulate narrow tongue does not
narrow greatly from base to apex.

The Papille.—A single prominent vallate papilla (text-fig. 58,
A) lies in the mid line; it is circular on plan and conical on
elevation, with the base of the cone projecting beyond the
well-marked fossa ; the vallum is rough, granular, and not easily
distinguished from the rough surface of the tongue. Martin,
Owen, and Forbes each describe a specimen with one papilla.

The fungiform papille ave small, and form a dorsal bounding
zone on which they have the usual arrangement in clusters and
rows; none are overlapped by the conical papille, which are
small, pointed, and irregular in distribution. On the ventral
papillary zone only conical papille are found.

Although there are no lateral organs similar to those of the
Primates, the posterior extremities of the corrugated lateral
borders have circular nodules with well-marked central orifices
(text-fig. 58, I); some of these are single and isolated, but
others are in small chains. Poulton (16) has described similar
bodies in Halmaturus walabatus, Macropus melanops, and Petro-
gale wanthops, but points out that their arrangement varies ;
they may be irregular or arranged in curves. He draws attention
to their appearances being similar to those of gland-ducts, and
sugeests that the lateral organs arise from glands; “ this view,”
he states, ‘is confirmed by a study of minute structure.”

No Apical Gland of Nuhn is present, but the glandular mass,
as described and figured by Oppel (10), sends two long processes
forward almost as far as the apex of the tongue. He believes
that the Gland of Nuhn, which is present in the tongue of Man
and .the Orang-Outan, are pieces of the basal glandular mass
which have become separated. Perhaps they have become
separated off from a prolongation similar to those of the Koala
and other Marsupials.

The trferior surface is wrinkled and fissured. In the mid line
in front there is a well-marked rough area, bounded laterally by
fissures. Posterior and lateral to the central area are crenated,
transversely-furrowed plice fimbriate (text-fig. 58, B), which
increase in width from before backwards.

Many Marsupialia have a sublingua which differs from that of
the Lemuroidea in being firmly fixed by its apex to the under
surface of the tongue. It varies in size in different species, but
that of the Koala is the smallest; it has been reduced to the
condition of plice fimbriat in the same way as the sublingna of
the Lemuroidea gives way to the plice of the Simiide. A few
illustrations of the degrees of development in the Marsupialia
are shown in text-fig. 58, B~-H. The sublingua also becomes
softer as well as reduced in size, as in the Primates. Thus there
is the complete horny sublingua as in Lemur, the softer and
smaller sublingua as in Varsiws, and the plice fimbriate as in
-Anthropopithecus.

556 DR. C. F. SONNTAG ON THE COMPARATIVE ANATOMY

Text-figure 58.

Trichosurus Pseudochirus
vulpecu/a. PETegrinUs.
i Dd.

Didelphys Dasyurus Perameles. Dendrolagus.
wirgimana. hellucatus. gunn. inustus.
(Ei is G H,
P2060 ea 200 02°
anc
J. A.
/.

The tongues of some Marsupials.

A. and B. Phascolarctos cinereus; C-H. inferior surfaces of the tongues; I-K.
lateral organs of Phascolarctos, Trichosurus, and Pseudochirus respectively ;
L-N. vertical sections of the tongues of Phaseolaretos, Trichosurus, and
Pseudochirus showing degrees of freedom of the sublingua.

OF THE KOALA AND VULPINE PHALANGER, 557

Trichosurus vulpecula.

Poulton (16) described the tongue under the old name of
Phalangista vulpina, so I shall only mention the differences
between it and that of Phascolarctos :—1. The epiglottis overlaps
the base. 2. Three papille form a vallate triangle. 3. The
lateral organs consist of fissures and lamine. 4. The sublingua
is a flat horny plate with a strong median ridge and a denticu-
lated apex bound down to the under surface of the tongue
(text-fig. 58, C). 5. The two forwardly-directed glandular
processes are stouter and relatively shorter.

Pseudcchirus peregrinus.

The pavrallel-sided tongue has the following dimensions :—
Total length 4:2 em.; length of the oral part 3:7 em.; length
of the pharyngeal part °5 em.; width between the lingual attach-
ments of the palato-glossal folds ‘4 cm.

The Papille.—Three circular vallate papille form an equilateral
triangle with the apex behind; the basal papille are prominent,
but the apical one is strongly retracted.

The fungiform papille, which are small and few in number,
form a dorsal bounding zone, but they have no definite arrange-
ment in transverse rows and clusters; they are surrounded by
sharp conical papille, which form transverse rows almost up to
the apex of the tongue. Only conical papille are present on the
ventral bounding papillary zone.

No orvfices of ducts and glands are seen on the base of the
tongue.

The Lateral Organs (text-fig. 58, K).—On each lateral border
of the tongue there is a row of circular elevations with central
slit-like orifices, so they are not quite similar in appearance
to those of Phascolarctos. The slits run upwards and backwards.
Each organ consists of six areas. On the surrounding mucosa
there are many nodules and papille.

The Sublingua (text-fig. 58, D).—The most pronounced feature
of the tongue is the large, soft sublingua. It is firmly bound
to the under surface of the apex, as in all Marsupialia in which
it is complete, but its under surface is deeply fissured and
separated by a deep gap from the tongue (text-fig. 58, D and N).
Running backwards in the mid line is a strongly-developed
median crest, which is bounded behind by a fissure.

There are great differences, therefore, between the lateral
organs, ventral surfaces, and vertical sections of the tongues of
these three species, as seen in text-fig. 58, B-D and I-N. The
essential differences between the sublingue of the Marsupialia
and Primates lie in the apices—in the former the apex is fixed
to, but in the latter it is free from, the under surface of the
tongue,

558 DR. C. F. SONNTAG ON THE COMPARATIVE ANATOMY

THe PHARYNX.

Phascolarctos cinereus.

The distance from the fauces to the beginning of the cesophagus
is 4°5 cm., and the greatest width of the pharynx, which narrows
from before backwards, 1s 2°2 em. The mucosa is smooth, and
two ridges run along the sides of the mid-dorsal line (text-

fig.) OO He):

Text-figure 59.

Mouth

Fauces.

Pharyn x.

| Ocesophagus

The pharynx of Phascolarctos cinereus.
Explanation of letters in text.

The éonsils (text-fig. 59, c) are small oval bodies in the dorso-
lateral aspect of the pharynx, a little distance anterior to the
hyoid bone. They appear as pits on the interior of the pharynx,
and the right one is surrounded by a prominent ridge.

The naso-pharynx.—The nasal fosse are prolonged backwards
by a conical tube, which opens on the mid-dorsal wall of the
pharynx by a small circular orifice guarded by two fleshy cushions
(text-fig. 59, Ad and C). It has no mesial dorsal ridge as in

OF THE KOALA AND VULPINE PHALANGER. . 559

Trichosurus and Pseudechirus. The opening lies within the
hyoid ring about 5 mm. anterior to the beginning of the
cesophagus, so the naso-pharynx is exceedingly small. his is a
remnant of the condition present in the new-born animal.
Owen (11) describes how the larynx is gripped by the nasal tube
in the mammary foetus of the Kangaroo, and the milk stream is
divided by the larynx, which crosses the pharynx; he gives
illustrations.

Sections through the head and neck at different levels show
how the shape of the nasal tube alters from before backwards
(text-fig. 59, F and G).

The Hustachian Tubes (text-fig. 59, b).—The lower orifices of
the tubes are slit-like, and lie on the infero-lateral aspects of the
nasal tube at a distance of 2:9 cm. posterior to the hard palate,
and at a point about 1 em. posterior to the fauces. In Plate V.
pins are inserted into the inferior orifices of the Eustachian
tubes,

Trichosurus vulpecula.

The pharynx differs from that of Phascolarctos as follows :—
1. The tonsils are in the fauces.
2. There is a ridge on the dorsal wall of the nasal tube, but
none in the pharynx.
3. The larynx crosses the pharynx and enters the nasal tube.
The nasal tube of Pseudochirws has a median dorsal ridge
(Plate V., C).

THE CisopHaaus.
Phascolarctos cinereus.

The cervical part (4:3 cm. long) is entirely concealed by the
larynx and trachea, and its mucosa is smooth throughout. The
thoracic part (7:8 em. long) has small longitudinal folds on its
inner surface. The abdominal part (1:8 cm. iy is lined by mucosa
thrown into prominent longitudinal folds. At the point where
it emerges from the posterior surface of the liver it is in contact
dorsally with the right suprarenal capsule.

Trichosurus vulpecula.

The esophagus projects beyond the left border of the trachea
in the neck, and it has a longer course (=3 cm.) in the abdomen
than has that of Phascolarctos. It is not in contact with the
right suprarenal capsule. In some examples its abdominal part
is much shorter, however.

Tur PERITONEUM.

Phascolarctos cinereus.

The great omentum is devoid of fat, but contains a well-marked
arterial arcade formed by a branch of the splenic artery and a

Proc, Zoot, Soc,—1921, No, XX XVIII, 38

560 DR. C. F. SONNTAG ON THE COMPARATIVE ANATOMY

twig from the arterial ring round the pyloro-duodenal junction
(text-fig. 60, B, a’). There is nothing new to record about its
attachments.

The lesser omentnm has a T-shaped attachment to liver and
stomach (text-fig. 60, A, @). The horizontal limb lies along the
lesser curvature, and the vertical limb runs back along the
ventral surface of the stomach.

The great omentum of Trichosurus vulpecula is well laden
with fat.

THE STOMACH.
Phascolarctos cinereus.

The observations recorded here are supplementary to the
descriptions by Martin (8), Young (18), Forbes (4), Edelmann (2),
Flower (3), and Home (8).

The long axis of the sac-like stomach, which lies in the
epigastrium and left hypochondrium. runs posteriorly, ventrally,
and to the right, and the well-marked fundus is directed forwards
and to the left. Projecting from the greater curvature imme-
diately below the pyloric cylinder is a sac whose walls are thinner
than those of any other part of the stomach.

Relations :—

Anteriorly—iiver and right suprarenal capsule.

Posteriorly—spleen, intestines, solar plexus, left kidney, and
suprarenal capsule.

Right—duodenum, gall-bladder.

Left and ventral—abdominal parietes.

Dorsal—crura of the diaphragm, aorta, pancreas, right kidney,
inferior vena cava.

Measurements :—

Nobtal lengthy. 255%. trades tees aeeenere a
eng th sot fumals: ficctseteseus Aen ee 2
Width between curvatures ............ Be
2
1

Arteries (text-fig. 60).—The stomach receives its blood supply
from the three branches—gastric (@’), hepatic (6), and splenic (c)
—of the celiac division of the cceliaco-mesenteric artery (see
page 570), and each vessel is accompanied by fine nerves derived
from the vagus and solar plexus (text-fig. 60, A).

The gastric artery divides into a large ventral branch (d) and
two dorsal branches (e)—large and small. The former, after a
straight course towards the stomach, breaks up into many
tortuous vessels ramifying over the fundus, and some can be
traced into the left side of the gland-patch. The latter make a
similar vascular arrangement on the dorsal aspect of the f undus,
and supply the patch as well.

OF THE KOALA AND VULPINE PHALANGER, 561

The hepatic artery gives off a large gastric branch (G. A.)
which divides into four—two dorsal (g.g) and two ventral—
branches (h. 2). Each pair supplies one surface of the stomach,

Text-figure 60.

Ovesophagus.

Saccu/us,

CA.

Oesophagus.

The ventral (A) and dorsal (B) surfaces of the stomach of
Phascolarctos cinereus with arteries.

38"

562 DR. C. F. SONNTAG ON THE COMPARATIVE ANATOMY

and forms half of a vascular ring encircling the pyloro- duodenal
junction. They supply the gastric wall and gland-patch. The
pyloro-duodenal ring communicates with the mesenteric arteries,
and a twig forms a long arcade (777) in the great omentum with a
branch of the splenic artery ; this arch gives off vasa brevia,
which enter the stomach-wall vertically (text-fig. 60, B).

There is no continuous vessel or arcade on the lesser curvature,
for the gland-patch interrupts it.

The veins enter the portal and splenic veins.

The gastric nerves (text-fig. 63, V.G.P.) form a complex plexus
on the left side of the gland-patch, and are intermingled with
branches of the gastric artery, and fine nerves reach the stomach
along branches of the hepatic artery. They will be described
later along with the solar plexus and vagus nerves. The nerves
reach the interior along the course of the arteries.

The interior of the stomach (Plate VI., A) was briefly described
by Martin (8). Apart from the enumeration of the orifices of
the gland-patch by several authors, no other description of the
interior has been published.

The long axis of the oval gland-patch, which lies along the
lesser curvature of the stomach, is 2 cm. long, and the width is
1:7 em. Its inner surface has thirty orifices of different sizes,
arranged in rows at right angles to the long axis of the patch.
In addition to these the lens reveals how the surface of the patch
is finely-pitted.

The remainder of the inner surface is traversed by ruge
radiating from the edges of the gland-patch like the spokes of a
wheel. Those running towards the lower end of the cesophagus
are short and Y-shaped; those passing into the fundus and body
of the stomach are at first straight but undulating later ; none
are present in the sacculus on the greater curvature ; and those
passing to the pylorus are few in number, but stronger than in
other parts.

The musculature of the pylorus thins out on the walls of the
sacculus, and that of the ceesophagus thins out on the fundus and
lesser curvature.

The gastrie contents consisted of a viscid creamy chyme, and
they resembled the contents of the stomachs of two Langurs
which I examined. In the Sloth, on the other hand,as I showed
in a previous paper, the stomach contained many hard, almost
entire leaves.

Special gastric glandular apparatus is also present in the
Wombat, Beaver, Dormouse, Manatee, and Dugong. It was also
present in the now exterminated Steller’s Sea-Cow (Rhytina
stelleri). Knox thought that the apparatus in the Sirenia was
an electric organ, and traced the two vagus nerves into it.

The pylorus projects into the duodenum (Plate VI., B), as in
Man. The human form has been likened to the projection
of the cervix uteri into the vagina, The free surface is rough
and villous, ‘

OF THE KOALA AND VULPINE PHALANGER. 563

Trichosurus vulpecula.

The stomach differs from that of Phascolarctos in the following
respects :—

1, It is more globular.

2. There is no sacculus on the greater curvature, but the
stomach projects to the right of the pylorus.

3. The gland-patch is not present on the lesser curvature.

4, The pylorus is not such a well-defined cylinder, and does
not project into the duodenum as in Man and Phascolarctos.

5. The walls of the stomach are relatively thinner.

6. The mucous membrane is rough, but has no ruge.

In a hitherto unpublished note Garrod states that the fundus
of Cuscus maculatus has a reticulum of ruge.

Tur INTESTINES.
Phascolarctos cinereus.

The external and internal characters have been fully described
by Forbes and Chalmers Mitchell (9), and I have nothing to add
to their accounts. My measurements differ greatly, however,
from those recorded by various authors. The figures in the
following table represent inches, and the asterisks denote that
fresh specimens were examined. Lonnberg’s specimen was a
mammary fetus*. It due allowance is made for shrinking
induced by preserving fluids, it will be seen that the lengths of
the different parts of the intestinal tube vary greatly in indi-
viduals, and have no relation to the length of the animal :—

ache Ae z es ae bith Seen es. SET
* Knox (1826) ...... 23 92 128 17
Martin (1836) 2. not given not given 76 50
Owen (1837)...... 23 92 125 a
*Horbes (188). oF 115 93°25 46°75
Forbes (1881) 9. 20°5 st) 160°8 66
Lonnberg (1902). 3°5 14°55 11°45 34
*Sonntag (1921)... 21 54:5 (5) 36

Lonnberg (17) compared the measurements of the different
parts of the intestinal tract of Phascolurctos and several other
Marsupials, and he also compared those of the feetal and adult
intestines, taking Forbes’s figures as the standard for the latter.
He came to the following conclusions :—“. ... the difference
is quite striking with regard to the cecum and the large intestine.
The former is proportionately only about a third as long in the
foetus as in the full-grown animal, and the latter less than half

* The mammary foetus is the new-born animal fixed to the nipples in the pouch,

564 DR. C. F, SONNTAG ON THE COMPARATIVE ANATOMY

as large in the fetus as in the adult. The difference of the
small intestine in both stages is not so great, that of the foetus
being about four-fifths of the same in the adult. It is also to be
remarked that in the feetus the small intestine is considerably
longer than the colon, but in the adult the reverse is the case.
The differences can of course be ascribed to the difference of the
diet of both stages. The milk food of the mammary fetus is
chiefly or completely digested in the small intestine, but the
vegetable diet of the adult needs a greatly developed cecum and
colon, The longitudinal folds of the cecum and colon are,
however, already developed in the feetus.”

Tt is possible that the growth of the intestinal tract in my
specimen has been considerably retarded.

Ellenberger, Tullberg, and Lonnberg believe that the large
size of the cecum in many animals, such as the Horse, Koala,
and some Rodents, is brought about by the presence of a large
amount of cellulose in the diet; and when the small intestine
deals with all the elements of the food, there is no specialisation
in the cecum and colon. Lonnberg also speculates about the
great length of the small intestine in phyllophagous Mammals.
He states that the leaves consist of cellulose and certam proto-
plasmic, amylaceous, and other substances contained within
resistant envelopes. Some time must elapse, therefore, before
these are broken down and their contents become available for
digestion and absorption; the small intestine is, consequently,
long.

Lonnberg’s views, however, will not account for the conditions
present in the intestinal tract of the phyllophagous Three-toed
Sloth (Bradypus tridactylus). In my paper dealing with its
anatomy (18), I showed that the intestinal tract is short, the
cecal pouch is rudimentary, and the interior of the colon and
rectum offers a large absorptive area, due to the excavation and
folding of the mucous membrane; it possesses sphincters which
are similar in character-to, but larger than, those of the cecum of
Trichosurus vulpecula, and its muscular coat, which thickens
from before backwards, may have a similar function to that of
the cecum of 7’. vulpecula.

Although the stomach of 86. tridactylus contains many hard,
almost entire leaves, the intestinal tract contains none, so the
cellulose and hard materials have been completely comminuted
and reduced to the state of chyme; Rapp’s analogies are interest-
ing in this connection, for he likens the paunch and pylorus to a
crop and gizzard. This thorough digestive process must throw
less work on the small intestines, which, in consequence, are short.

The chyme yields a number of substances for absorption in the
small intestines, and the remainder is absorbed from the large
intestines, whose pockets and sphincters retain it in the colon
and rectum, and the muscular coat moves it backwards and
forwards; and the sphincter ani has ‘a retaining function similar
to that of the septa higher up in the colon.

OF THE KOALA AND VULPINE PHALANGER. 56d

The differences between these two phyllophagous Mammals
must therefore be due to complexity of the stomach and the
different varieties of specialisation of the interior of the colon
and rectum, but the increased internal surface of the colon and
rectum of the Sloth is compensated for by the large size and
great length of those of the Koala. The Langurs are inter-
mediate in the complexity of their intestinal tract between the
Sloth and Koala.

In Phascolomys mitchelli the cecum is considerably reduced,
and Lénnberg believes that the colon has become considerably
enlarged to take on the cellulose-digesting function of the ceecum.
The stomach is simple as in the Koala, so it has not had any
modifying influence on the complexity of the intestinal tract
as in the Sloth. The Wombat has external sacculi, but the
Sloth has internal cavities, so there is a greater similarity
between their colons than exists between the colon of either of
these animals and that of the Koala.

I have already shown how the complexity of the stomach is
also influenced by the degree of development of the buccal organs
(19). One must take a wide view of the whole alimentary canal
from mouth to anus in framing an explanation of the physiology
of any one part.

THe Urinary Orcans (text-fig. 61).

Phascolarctos cinereus.

The right kidney is 3:2 cm. long, 1:8 cm. wide, and 1:6 em.
thick ; it lies entirely anterior to the left kidney, whose corre-
sponding measurements are 3°7, 2°4, and 15 cm.; ts hilum is
ventral in position, whereas that of the left kidney is mesial.
The structures in the hilus have the usual disposition in regard
to one another.

On section each kidney is seen to possess a thin cortex, and a
single large papilla.

The surface of each renal pelvis and hilus is covered by a plexus
of small veins. These communicate posteriorly with the veins
ascending along the ureter, and they open anteriorly into the
renal vein.

The right ureter lies parallel to, but 1 em. to the right of, the
vena cava inferior. It passes ventral to the iliac vessels and
runs posteriorly along the dorsal pelvic wall. It opens into the
lower and back part of the bladder.

It is accompanied by small veins from the pelvis; the latter
are joined by veins from the ovaries, after which a Jadder-like
arrangement of veins runs up to the venous plexus on the ventral
surface of the renal pelvis (text-fig. 61 C, wv).

The left wreter runs along the left border of the abdominal
aorta, crosses the iliac véssels, and courses through the pelvis as
does the right one. The veins accompanying it are numerous

566 DR. C. F. SONNTAG ON THE COMPARATIVE ANATOMY

and some of them ramify; so dense is the venous plexus that
the ureter is concealed by them. At first sight it looks like a
cord of veins.

John Hunter (22) showed that the relative positions of the
kidneys of Petaurus taguanoides are similar to the above.

Text-figure 61.

A. heart and aorta of Phascolarctos cinereus with origins of right subclavian,
right common carotid, left common carotid, and left subclavian arteries, from»
right to left, and described in text as a, b, c, d: B. heart and aorta of

Trichosurus vulpecula: C. kidneys and some of the abdominal veins: ~

L.S.R.C. and R.S.R.C., suprarenal capsules; P.L., peritoneal ligament: D and
E, origins of vertebral arteries in Phascolarctos cinereus and Trichosurus
vulpecula respectively: F and G. spleens of Trichosurus vulpecula and
Phascolarctos cinereus respectively.

——-

OF THE KOALA AND VULPINE PHALANGER. 567

The bladder is pyriform and firm, but no urachus is seen.
Lateral peritoneal folds connect it to the pelvic wall, but no
other ligaments are present.

Trichosurus vulpecula.

The anterior half of the right kidney. is received into a deep
recess in the liver, and its posterior pole is level with the hilus
of the left kidney. The bladder is transversely corrugated on
the surface and no urachus is present. Only one wide renal
papilla is present in each kidney.

GENERATIVE ORGANS.

I have nothing to add to existing accounts of the generative
organs of either species.

Tur PANCREAS.

The duct is dead-white in Phascolarctos and shines through
the translucent brown glandular tissue. The latter in my speci-
men has the same appearances as those described by Martin (8).

Tue Liver.

. r
Phascolarctos cinereus.

Forbes (4) described and figured the liver, and Braus (1)
described its histology.

The hepatic artery is very convoluted, and divides into two
terminal convoluted branches which diverge at the sides of the
gall-bladder. From the left branch a straight, slender cystic
artery runs in the free edge of the vesical mesentery to the gall-
bladder, which it reaches about its middle; then it runs poste-
riorly along the surface of the gall-bladder as far as the fundus.

The postcaval vein is never deeply embedded in the hepatic
substance, and receives three hepatic veins at its emergence from
the anterior surface of the liver. A fourth vein enters it within
the organ.

The common bile-duct does not exhibit cysts of its mucous
membrane similar to those of some Marsupials described by

Owen (11).

Trichosurus vulpecula.

_ The liver differs from that of Phascolarctos as follows :—1. It
is not so compressed antero-posteriorly. 2. Its edges are not
notched by many small fissures. 3. The hepatic tissue is not so
subdivided. 4. The right lobe recesses the anterior half of the
right kidney. 5. The gall-bladder is shorter and broader, and
only projects for a short distance into the abdominal cavity

568 DR. C, F, SONNTAG ON THE COMPARATIVE ANATOMY

beyond the posterior edge of the right lobe when fully distended
with bile.

THe Ductuess GLANDS.

Phascolaretos cinereus.

The Thyroid Gland (text-fig. 56, A, A).—The two lateral lobes
are unconnected, and vary in shape and extent. The lefé lobe is
pyriform, with the broad extremity posterior and slightly within
the thorax. It is 2 em. long, *8 em. wide, and °3 cm. thick. It
is separated from the larynx by the first three tracheal
rings. The right lobe is elongated and ovoid, and measures
27x'‘5x:3 em. Its anterior extremity overlaps the lower end
of the larynx, and its posterior pole stops anterior to the thorax.
The arterial supply comes from the thyroid and laryngeal branches
of the common carotid artery (m), and the veins run to the
internal jugular vein (7).

I did not observe any structures which could be definitely
described as parathyroid bodies, but they may have been lying
among the small glands which are numerous at the root of the
neck.

The Thymus Gland has been described hy Symington (15).

The Suprarenal Capsules (text-fig. 61, s..c.). —The left capsule
is a round body, of the size of a large pea, lying on the mesial
aspect of the anterior pole of the left kidney, to which it 1s
attached by a fold of peritoneum. The right capsule is oval, and
has a peculiar position; it lies in contact with the posterior
surface of the liver, and is situated between the inferior vena
cava to the right, the right crus of the diaphragm to the left,
and the esophagus ventrally.

The greater part of each capsule is composed of brown
medulla.

The spleen has been described by Martin (8).

My specimen is triangular and measures 7°4 cm. long, but
that of Zrichosurus vulpecula has a lateral process (text-fig. 61,
Fand G). In Phascolomys it is an equilateral triangle.

Trichosurus vulpecula.

The lateral thyroid lobes (text-fig. 57 A) measure 1:3 cm. long
and *3 cm. wide. ‘They lie at the sides of the posterior extremity
of the larynx and anterior seven tracheal rings. The connecting
isthmus crosses rings 5, 6, and 7.

The thyroid artery describes a curve, with the convexity
forwards, and runs along the anterior five-sixths of the mesial
border of its corresponding lateral lobe. The thyroid vein runs
to the internal jugular vein(p). Thisarrangement of the thyroid
vessels is seen in animals belonging to several of the Mammalian
orders.

OF THE KOALA AND VULPINE PHALANGER. 569

Tur ORGANS OF CIRCULATION.
Phascolarctos cinereus.

The Heart (text-fig. 61, A).—The fibrous pericardium is firmly
bound to the dorsal Surface of the sternum and left ribs by
connective tissue. It narrows out and is lost on the great vessels.
These conditions are also present in J'richosurus vulpecula.
The area of adhesion is greater than that of Dendrolagus ursinus.

The long axis of the heart is antero-posterior and parallel to
the left side of the sternum, and the apex is level with the
posterior border of the fourth left costal cartilage. Its apex is
more obtuse than that of 7'richosurus vulpecula *

Measurements :—

Phascolarctos. Trichosurus.

Meme this. 2 uay hehe 5: 4-6 cm. ae em.
Greatest width ...... DWART os Dahan
Greatest thickness ... 1:9 ,, Toes.

The characters agree with those already described for the
Marsupial heart.

The Ascending Aorta (text-fig. 61 A) is embraced by the two
parts of the right auricular appendix, and is parallel to the
right precaval vein. It quickly gives way to the aortic arch, whose
most anterior point is level with the anterior end of the first
intercostal space. At the level of the fourth dorsal vertebra it
gives way to the descending aorta; this is the commonest level
for the transition in the Mammalia. The relations of the arch
are :—

Ventral.—Left lung, left precaval vein, left azygos vein, left

vagus nerve, left phrenic nerve.
‘Dor sal.—Trachea, cesophagus.
Posterior.— Root of left lung.
_ Anterior. — Great vessels of head, neck, and fore limbs.
Branches :—The right subclavian (a), right common carotid (4),
and left common carotid (c) arise directly, and close to one
another, from the aortic arch, so there is no innominate artery.
After a short interval the left subclavian artery is given off (d).
Martin (8) and Owen (11) describe an arrangement similar to
that of Man, but Forbes (4) states that the innominate artery
gives off three vessels close together from a common trunk. In
Trichosurus vulpecula the aortic arch gives off the innominate
and left subclavian arteries, and the former, after giving off the
left common carotid, divides into right common carotid and right
subclavian arteries; the left common carotid arises almost at the
beginning of the innominate ar tery.

The Common Carotid Arteries give off fine entocarotids at the

level of the anterior border of the hyoid bone, and are continued

* Some specimens of Trichosurus vulpecula have a more obtuse cardiac apex than
the above.

570 DR. C. F. SONNTAG ON THE COMPARATIVE ANATOMY

as stout ectocarotids. The former give off no branches in the
neck, and enter the carotid foramen in the basisphenoid. The
latter pass antero-laterally between the exoccipital process and
the mandible, and the former separates it from the external
jugular vein. In Trichosurus vulpecula the exoccipital process
does not separate these vessels.

The Descending Thoracic Aorta lies first on the right half of
the body of the fifth dorsal vertebra, but passes more to the
right as it runs backwards through the thorax. It is connected
to the esophagus by a triangular fold of pleura, whose base is
attached to the diaphragmatic pleura. It has the usual relations
and gives off the usual branches.

The vertebral arteries come off from the subclavians, but in my
specimen of Trichosurus they arise from axes (text-fig. 61, E).

The Abdominal Aorta differs from the common Marsupial
condition in that it is entirely to the left of the postcaval vein.
Hochstetter (20) examined several species, and found that the
postcaval vein concealed the lower end of the aorta in all but
Petaurus taguanoides, but he did not include Phascolarctos
cinereus in his list. Beddard (21) also found the posteaval vein
covering the aorta in all his species. My observations on 7'richo-
surus vulpecula and Dendrolagus ursinus confirm those of Hoch-
stetter and Beddard.

Relations.—At the point where the abdominal aorta reaches
the level of the posterior surface of the liver, it is separated from
the right suprarenal capsule by the right crus of the diaphragm.
These structures separate it from the postcaval vein.

A little posterior to the capsule the solar plexus lies between
the aorta on the left, the postcaval vein on the right, and the
coeliaco-mesenteric artery ventrally.

Posterior to the solar plexus the postcaval vein lies immediately
to the right side of the aorta till the latter divides into the
internal iliac and mid-sacral arteries, and then the vein crosses
the surface of the aorta. The whole trunk of the aorta is
visible.

The relations to other structures show nothing unusual.

Branches.—With the exception of the four pairs of lumbar
arteries which arise at regular intervals from the sides, the
following is the order of the branches from before backwards :—

Phrenics, cceliaco-mesenteric, suprarenals, renals, ovarians,
external iliacs, internal iliacs, and middle sacral (caudal).

After giving off the external iliac arteries, the trunk of the
aorta diminishes in calibre to a little more than half of that of
either of the external iliac arteries.

The Celiaco-mesenteric Artery divides into celiac axis and
anterior mesenteric artery, and a branch of the latter replaces
the posterior mesenteric branch of the abdominal aorta of the
higher Mammalia.

The celiac axis divides into gastric, hepatic, and splenic
arteries, whose distribution has already been described.

OF THE KOALA AND VULPINE PHALANGER. 571

The anterior mesenteric artery describes a curve, convex
forwards and to the right, and terminates in the cecum; it
occupies the free edge of the cecal mesentery in the latter part
of its course.

The branches supplying the large intestines arise from the
convexity of the curve, and those destined for the small intestines
are given off from the concavity. The highest branch anasto-
moses with the duodenal branches of the hepatic artery, and the
lowest one anastomoses in the pelvis with the hemorrhoidal
vessels. The lowest branch to the ileum anastomoses with the
branches to the cecum.

The mesenteric areas between the branches of the mesenteric
artery contain rich networks of fine lymphatic vessels, and fine
branches of the solar plexus accompany the arteries. The
mesenteric veins have been figured already by Chalmers
Mitchell (9).

In Trichosurus vulpecula the celiac axis arises separately.

Tur VEINS.

The Anterior and Posterior Facial Veins have the usual forma-
tion, course, and tributaries. They unite to form the external
jugular vein on the surface of the exoccipital process. The latter
runs postero-mesially across the sternomastoid to reach its lateral
border in the posterior sixth of the neck. It passes dorsal to
the posterior end of the sterno-mastoid and clavicle, and, anterior
to the neck of the first rib, it unites with the axillary vein to
form the precaval vein. Both veins are united by a long
V-shaped vena transversa. In Trichosurus vulpecula the vein is
more transverse (text-fig. 55).

Tributaries.— Auriculo-tem poral, acromial, transverse scapular,
muscular, ascending cervical and internal jugular veins. ‘hese
vessels have the usual distribution. No anterior jugular vein
is present. _

The vena transversa veceives veins from the tongue, sub-
maxillary glands, and muscles of the mylo-hyoid region.

At the point where the internal and external jugular veins
unite there are several small glands.

The innominate and precaval veins follow the usual course of
the Marsupial type, and the azygos veins in my specimen are
similar to those described by Forbes (4). The intrathoracic part
of the postcaval vein measures 1-5 em.

The postcaval vein is formed in the usual manner, but the
only part lying ventrally superficial to the abdominal aorta
crosses the termination of the latter. This arrangement is an
exception to Beddard’s statement that “the Marsupials show a
very constant condition in that the postcava lies medianly
ventral to the aorta, so that on dissection the aorta is not seen,
being completely covered by the postcava ” (21).

It lies to the right side of the aorta, and close to it till it

yf DR. C. F. SONNTAG ON THE COMPARATIVE ANATOMY

reaches the right renal vein. After that it bends to the right
and enters the liver. At the point where it disappears from
view it is in contact with the right suprarenal capsule, and is
accompanied by twigs from the right half of the solar plexus.

The left ovarian vein (text-fig. 61 C, ov.) joins the thick venous
plexus running along the ureter; therein it is united to veins
ascending from the pelvis. The right ovarian vein communicates
with the right ureteric veins by a stout trunk, and then runs
forwards to join the postcaval vein. The ureteric veins (uv. and
wv.) form a plexus on the ventral surfaces of the renal pelves and
open into the renal veins (rv.).

Tue Neryous System.

The brain has been described by Owen (11), Forbes (4), Elliot
Smith (12), and Ziehen (14), and I have nothing to add to their
accounts,

Tur Vacus Nerves (text-figs. 62 and 63).
Phascolarctos cinereus.

A short distance posterior to the base of the skull the vagus
nerve on each side receives the hypoglossal (XIT.) nerve, and their
fibres intermingle in a very thorough manner. Soon they
separate again, and the vagus runs postero-mesially as far as the
anterior extremity of the thyroid cartilage, where it unites with
the cervical sympathetic cord(C.5.) to form a vago-sympathetie
(V-S.). A short distance anterior to the clavicle it separates again
into vagus (V.) and sympathetic (8.).

This arrangement is present in most of the Carnivora. It
differs from that of Dendrolagus, in which the vagi and sympa-
thetics remain separate.

There is no trace of the ganglion of the trunk of the left
vagus (ganglion nodosum) in the neck. The right nerve has
one (G.N.).

Branches in the neck.—Pharyngeal (a), anterior laryngeal (6),
communicating to the first cervical nerve (c), and recurrent
branch of the right vagus (d).

The relations of the vagiin the thorax differ. The left nerve
erosses superficial to the left subclavian artery and aortic arch,
lying on the inner side of the left precaval vein and left phrenic
nerve, and on the outer side of the trachea, cesophagus, and left
common carotid artery. It passes dorsal to the root of the left
lung and gains the dorsal aspect of the cesophagus. The right
nerve crosses the right subclavian artery under cover of the
right precaval vein, runs down the side of the right main
bronchus, and gains the dorsal aspect of the root of the right
lung; lastly it passes to the dorsal aspect of the cesophagus,
where it fuses with the left nerve to form the combined trunk of
the vagi (C.T.V.).

OF THE KOALA AND VULPINE PHALANGER, 573

Branches of the separate nerves in the thorax.—Left recurrent
laryngeal nerve (e), cardiac (f), pulmonary (g), and a branch
from each nerve which unites with its neighbour to form a fine
cord, which runs posteriorly along the ventral surface of the
cesophagus (/).

The combined trunk of the vagi enters the abdomen dorsal to
the cesophagus, runs dorsal to the stomach, and ends in the left

Text-figure 62.

a--2- + se e-

Qs te ee

b.=)= =i

‘

‘ :

b= 22
ine

The vagus nerves of Phascolarctos cinereus in neck and thorax.

half of the solar plexus. It gives off ventral gastric nerves in
the thorax (7), and stout ventral (j), and strong dorsal (4), gastric
nerves in the abdomen.

It is possible that sympathetic filaments run anteriorly from
the solar plexus to the stomach in the combined trunk of the
vagus, for it has been shown that sympathetic filaments run in

other nerves (¢. g..the sciatic).

574 DR. C. F. SONNTAG ON THE COMPARATIVE ANATOMY

The two halves of the solar plexus (text-fig. 63) he between
the abdominal aorta and postcaval vein under cover of the
origin of the cceliaco-mesenteric artery. The right half (A)
is a fusiform ganglion, but the left half (B) consists of a nerve

Text-figure 63.

The vagus nerves and abdominal sympathetic of Phascolarctos cinereus.
t=} J

plexus with a small round ganglion. The combined trunk of
the vagi (C) enters the left half of the plexus by a brushwork
of nerves. ‘The right and left sympathetic nerves (D and K) end

in the right half of the plexus, but the left sympathetic gives off

OF THE KOALA AND VULPINE PHALANGER. 575

a fine nerve (F), which enters into the formation of the aortic
plexus farther back (G). Fine fibres (H) connect the halves of
the plexus.

Branches of distribution :—

1. Three nerves which course along the posteaval vein and
embrace the right suprarenal capsule (1).

2. Hepatic plexus (J).

3. Splenic plexus (K).

4, Right renal plexus (L).

5. Two cords (M and N) which emerge from the posterior end
of the right half of the plexus and form a loop lying dorsal to the
postcaval vein. The left one breaks up into a number of fibres
which reunite later with the right one, and it gives off the left
renal plexus (QO).

The loop unites with the left sympathetic by a T-shaped
junction (P), and fine filaments (Q, Q, Q, Q) also run between
them.

Only one cervical sympathetic ganglion
one—exists,

namely the superior.

SUMMARY.

The differences between Phascolarctos and T'richosuruws may be
summarised as follows :—

Phascolarctos. ' Trichosurus.

1. Platysma myoides strong. 1, Platysma weaker.

2. No true hyoglossus muscle. 2. Hyoglossus present.
3. Vena transversa A-shaped. 3. Horizontal.
4, A.
5 5

. Absent. . Anterior jugular vein present.
. Lateral organs of tongue look like . Composed of fissures and lamine.
elands.
6. One vallate papilla. 6. A vallate triangle.
7. Plice fimbriatz present. 7. Sublingua well developed.
8. Cheek-pouches ‘present. 8. Absent.
9. All anterior palatal ridges trans- 9. Anterior palatal ridges convex for-
verse. wards.
10. Tongue far from epiglottis. 10. Close to epiglottis.
11. Epiglottis broad and entire. 11. Narrow and notched.
12. Gland-patch in stomach. 12. Absent.
13. Pylorus projects into the duodenum. | 13. Pylorus does not project.
14, Tonsils far back in the pharynx. 14. Tonsils in fauces.
15. Pharynx has two dorsal ridges. 15. Absent.
16. Absent. 16. Dorsal ridge in nasal tube.
17. Cardiac apex obtuse. 17. More acute.
18. Absent in my specimen. 18. Innominate artery present.
19. Postcava crosses end of abdominal | 19. Postcava conceals posterior half of
aorta. aorta.
20. Liver more subdivided.
21. Gall-bladder long and narrow. 21. Short and wide.
22. Right kidney posterior to liver, and | 22. Sunk in liver, and each kidney has
each kidney has one conical papilla. one broad, flat papilla.
23. Thyroid gland in posterior part of | 23. In anterior part.
neck.

Proc, Zoot, Soc.—1921, No. XX XIX, 39

576 DR..C. F. SONNTAG ON THE COMPARATIVE ANATOMY

Phascolarctos. Trichosurus.
24. No thyroid isthmus. 24. Isthmus present.
25. Patella absent. . Patella thin and scale-like.

25
26. Absent. 26. Right lung has an azygos lobe.
27. Two lobes in each lung. 27. Left lung has two, and right has
four lobes.
28. Spleen triangular, and has no lateral | 28. Spleen has a lateral piece.

process.

29. Vertebral arteries arise from sub- | 29. Vertebral arteries arise from an
clavian arteries. axis on each side.

30. Intestinal measurements much
greater.

Owen stated that the Koala is the Marsupial which has the
greatest claim to typical pre-eminence. MacAlister, on the other
hand, showed that it has more individual peculiarities in its
myology than has any other Didelphian; and the remarks made
in this paper show how many of the appearances of its internal
organs are peculiar to itself, for several of those described for
Prichosurus vulpecula are similar to the conditions present in
many Marsupialia. There are also many striking differences
between the external and skeletal characters of Phascolarctos and
those of Tvichosurus, but I have nothing new to add to existing
descriptions.

BIBLIOGRAPHY.

1. Braus, H.—Jen. Denkschr. Bd. 5 (Semon, Zool. Forschungs-
reisen, Bd. 2). Jena. G. Fischer, 1896.

2. Epe~Mann, T.—Deut. Zeitschr. f. Thiermed. Bd. xv.
pp. 165-214. 1899.

3. Frower, W. H.—‘ The Comparative Anatomy of the Organs
of Digestion of the Mammalia.” Medical Times and
Gazette, 1872, p. 645.

4. Forpes, W. A.—‘‘On some Points in the Anatomy of the
Koala.” Proce. Zool. Soc. London, 1881, pp. 180-195.

5. Homs, E.—‘‘ An Account of some Peculiarities in the
Anatomical Structure of the Wombat.” Phil. Trans. Roy.
Soc. 1808, pp. 304-312.

6. Knox, R.—‘‘ On the Wombat of Flinders.” Edinburgh New
Philosophical Journal, vol. i. p. 104. 1826. .

7. MacAutstrer, A._—‘‘ The Muscular Anatomy of the Koala.”
Annals and Magazine of Natural History, 1872, pp. 127-
134.

8. Martin, W.—“ Notes on the Anatomy of the Koala.” Proc.
Zool. Soc. London, 1836, pp. 109-113.

9. MircuEett, P. CHoatmers.—‘ Further Observations on the
Intestinal Tract of Mammals.” P.Z.S8. 1916, pp. 195-
19%.

10. Orret, A.—Text-book of Comparative Microscopic Anatomy,
vol. i. p. 291, and vol. i. p. 308.

11. Owen, R. Art. « Marsupialia” in Todd’s ‘ Cyclopedia of
Anatomy and Physiology,’ vol. i.

‘
!
q
:
:
;

12.
13.

14.
15.

16.
re
18.
19:

20.
21.

22.

OF THE KOALA AND VULPINE PHALANGER. iy

Suirn, G. Exxror.—Journal of Anatomy and Physiology,
EXxili. 1898, p. 30.

Youne, A. H.—Journal of Anatomy and Physiology, xii.
1880, pp. 305-317.

ZienEN, T'.—Jenaische Denkschr., Bd. vi. 1897, p. 98.

Symineron, J.—Journal of Anatomy and Physiology, xxxiv.
p. 226.

Pouttron, E. P.—‘On the Tongues of the Marsupialia.”
Proc. Zool. Soc. London, 1883, pp. 599-628.

Lonnperc, E.—‘‘On Digestive Adaptations in the Marsu-
plata. Pawns. 1902.1. p. 12.

Sonnrac, C. F.—‘‘ A Contribution to the Anatomy of the
Three-toed Sloth.” P.Z.S8. 1921, pp. 157-177.

Sonnrac, C. F.—‘‘The Tongues of the Cercopithecide.”
P.Z.S. 1921, pp. 280-281.

Hocusretrer, F.—Morphol. Jahrb. xx. 1893, p. 626.

BepparbD, F. E.—‘“‘ On the Postcaval Vein and its Branches
in certain Mammals.” P.Z.S. 1909, pp. 496-526.

Hunter, Joun.—‘ Essays and Observations. Edited by
Richard Owen. 1861, p. 257.

EXPLANATION OF THE PLATES.
Prate VY.

The Palate of Phascolarctos cinereus with the cheek-pouches packed with wool
and pins in the Eustachian Tubes.

. The Palate of Trichosurus vulpecula.
. The Palate of Pseudochirus peregrinus.

Prate VI.

The Stomach of Phascolarctos cinereus.

. The interior of the stomach.
. The projection of the pylorus into the duodenum.

cr
“I
Je}

ON NEW INDIAN DRILID BEETLES.

31. New Indian Drilid Beetles. By 8. Mavuix, F.Z.S.,
Professor of Zoology in the University of Calcutta.

[Received May 11, 1921: Read June 7, 1921. |
(Text-figures 1-5.)

In the present paper four beetles from India of the Family
Drilide are described. They are all characterized by the pos-
session of extraordinarily developed lamellate terminal joints of
both the maxillary and labial palpi. These joints, of both pairs
of palpi, are similar in structure, and are of almost equal size.
They naturally fall into two genera, and can be distinguished
thus :—

Head almost as long as the pronotum, so that the eyes

are distant from the front edge of the pronotum.
Antennee distant ............ecsssececeecsesetereeesasess, Lamellipalpus, gen. 0.

Head short, the eyes almost touching the front edge of
the pronotum. Antennze more or less approxi-
THU C Meee sneer aee ven aa eee Re EBs esac LOMeELlipalpodes, Senn Ml:

LAMELLIPALPUS, gen. n.

The type of this genus is Hugeusis nigripennis Pascoe, from
Burma. At the end of his description of this insect, (Ann. Mag.
Nat. Hist. xx. 1887, p. 10, t.i.f.7) the author remarks: ‘“ The
description is from a female; the male has probably flabellate
antenne, as in #7. palpator Westwood” (the type of the genus
Hugeusis). Ihave examined Pascoe’s type; it is not a female,
therefore his conclusion that the male has flabellate antenne is
not correct. Under this assumption he included his nigripennis in
Westwood’s genus Hugeusis, of which I have seen a specimen in
the collection of the British Museum, and with which it is not
congeneric. It is necessary, therefore, to erect a new genus for
the reception of this species as well as two others in the collection
of the British Museum.

Generic characters.—The antenne are 11-jointed, filiform;
sometimes each joint is slightly expanded on the inner side. They
are separated from each other by the whole breadth of the head,
and are situated in front of the eyes. The eyes are convex and
prominent. The mandibles are large, curved, and pointed; in
repose they lie crossed. The maxillary and labial palpi are 4-
and 3-jointed respectively, the terminal joint in each being very
large, elongate, and compressed. The front coxe are large and
exserted, having a large trochanter ; the intermediate and hind
coxe are contiguous, but not so large as the front ones. The
tarsi are 5-jointed, the fourth being bilobed. The elytra are not
of a membranous texture and the suture is closely fitting.

580 PROF. S. MAULIK ON

LAMELLIPALPUS NIGRIPENNIS Pascoe.

The following description and illustrations are from a fresh
specimen taken “by Mrs. Kemp in Assam :—

Colour testaceous brown; underside paler; elytra blackish,
except an ill-defined basal area, the suture and the Perel
margins; the antenne except the two basal joints blackish ; eyes
deep black. The upper side of the head and prothorax is shining,
the rest of the body being subnitid. The whole body of the
insect is covered with fine brown pubescence.

Text-figure 1.

Lamellipalpus nigripennis Pascoe.

Head almost as broad as long, slightly narrowed towards the
neck, shallowly depressed in front; upper surface smooth, shining,
spar sely covered with fine brown hei Viewed from aber e, the
extraordinarily developed terminal joints of the palpi and the
large mandibles are conspicuous. The joints of the antenne are
not at all expanded on the inner side. Pronotwm broader than
long, front margin straight, lateral and posterior margins slightly
sinuate, anterior lateral angles rounded, posterior ones very
acute. The upper surface is gently convex, shining, covered with
fine brown pubescence, more so on the sides than on the middle
area, deeply depressed at the base on either side of the longi-

|
:

NEW INDIAN DRILID BEETLES. 581

tudinal middle line. Zlytra as broad at the base as the pro-
thorax; the humeral angles are rounded. Corresponding to the
basal depressions of the pronotum, the opposite elytral areas at
the base are also depressed. The surface is punctate, covered
with brown pubescence. Lach elytron has four feeble coste,
the two inner ones—z. ¢e., those nearer the suture—being more
prominent than the outer ones; the two inner ones commence
at the base and terminate before reaching the apex; the third,
commencing posterior to the humerus, terminates about the
middle; the fourth, commencing below the humerus, terminates

Text-figure 2.

Underside of Lamellipalpus nigripennis Pascoe.

just beyond the middle. Of the four cost the third is the
feeblest ; to a certain extent they converge towards the apex.
Scutellum triangular, light brown, pubescent, impunctate, sub-
nitid. Underside brown, but lighter than the colour of the
upperside; the abdominal segments have a slight suffusion of
black. The surface is sparsely covered with fine brown hairs,
their roots give the surface an appearance of being finely punc-
tate. The abdominal segments are at a lower level than the
metathoracic segment; each is convex in the middle and de-
pressed at the sides, these lateral depressions forming an oblique
line on either side. The underside of the femora and tibie is
not suleate.

Length 10 mm., breadth 43 mm. Length of head 2 mm.,

589 PROF. Ss MAULIK ON

breadth of head across the eyes 3 mm., without the eyes 23 mm.
Length of pronotum 23 mm.

Assam, Garo Hills; Tura, 1400 ft.; Oct. 1917 (Mrs. Kemp)
(Indian Museum).

The type specimen (Brit. Mus.) is from Burma. The brown
parts in the type are lighter than Mrs. Kemp’s specimen ; also
in the type the lighter colour of the suture and the lateral
margins of the elytra is not so well marked.

LAMELLIPALPUS BOMBAYENSIS, Sp. n.

Upperside shining, dark brown, in one specimen of the two
lighter brown; the antennal joints from the third joint to apex

Text-figure 3.

Lametlipalpus bombayensis Maulik.

brownish-black, and the apical area of the elytra obliquely con-
tinuing to the middle along the side smoky black; eyes deep
black. Underside shining light to dark brown. The whole insect,
both on the upper and lower sides, is covered with light brown
pubescence.

Head almost as long as broad, depressed in the middle in front
but convex round the roots of the antenne ; surface punctate,
each puncture having a stiff hair. The eyes strongly convex.

NEW INDIAN DRILID BEETLES. 583

The first joint of the antenne club-shaped, the second very short,
the third to the tenth almost equal in size and slightly expanded
triangularly towards the inner side, the last joimt bluntly
rounded. In the expanded character of the several joints of the
antenne it bears a certain amount of resemblance to Hugeusis
palpator Westwood, in which these points are extraordinarily
expanded into rami. Prothorax about as broad as long, slightly
narrowed towards the front; the posterior lateral angles pro-
duced to acute points; surface convex, with a faint longitudinal
impression along the centre and deeply sloped on each side ; this,
with a similar slope on the elytra in front of the humerus, forms
a deep depression ; closely punctate. Scutellum triangular, punc-
tate. Hlytra as broad at the base as the prothorax. The lateral
margins and suture costate. On each elytron there are three
more or less prominent coste; one runs close to the suture, the
second along the longitudinal middle line, and the one nearer
the margin starts from a point below the humerus, a feeble
one, corresponding to the third in Lamellipalpus nigripennis
Pascoe, is observable in certain lights. These coste approach
one another towards the apical area, where they disappear. The
surface is irregularly punctate. Underside shining, smoother
than the upperside. :

Length 9-10 mm.

Bombay.

Described from two examples, the larger of which I have made
the type specimen. It has the apical black patch on the elytra
diffused, while in the other it is more sharply defined. In the
type specimen the lamellate joints of the palpi are broader, with
apex slightly narrowed, while in the smaller specimen they are
uniformly narrower.

Type in the British Museum.

LAMELLIPALPUS MANIPURENSIS, Sp. n.

Upperside subnitid, thorax and head more shining than the
elytra ; elytra and eyes black, the four lamellate terminal joints
of the maxillary and labial palpi, and the antennal joints except a
part of the two basal joints, smoky black ; the rest of the body,
including the underside, brown. The lamellate joints of the
palpi are narrower and more elongate in this species than in the
two others of this genus.

Head almost as broad as long; interantennal space shallowly
depressed, covered with light brown pubescence. The first joint of
the antennz is almost as long as the third joint but very slightly
thicker, second joint small and rounded, third joint slightly club-
- shaped, fourth to tenth almost equal to each other in length and
thickness, the eleventh slightly lorger than each of the preceding
ones and rounded at the extremity; the joints of the antenne
are not at all expanded. Prothorax slightly broader than long,
sides and front margin straight, basal margin sinuate; the

584 PROF. 8. MAULIK ON

. posterior exterior angles acutely produced, the extreme edges of
the produced part sharply depressed. The pronotum is covered
with light brown erect pubescence. Scutellwm brown, triangular,
covered with light brown pubescence. lytra almost as broad at
the base as the prothorax. The surface is confusedly punctate
and covered with light brown pubescence. The three cost on
each elytron are very feeble though distinctly recognizable.
Underside: the elytra project to a certain extent beyond the
abdomen, shining, covered with stiff brown hairs.

Length 7:5 mm.

Manipur (Doherty).

Deseribed from one example.

Type in the British Museum.

LAMELLIPALPODES, gen. n.

The genus differs from Lamellipalpus in the length of the
head and in the approximation of the antenne, as has been
indicated in the key given above. The type of the genus is the
following species.

Although I include the two new genera described here in the
family Drilide, I am not sure about the position of Lamelli-
palpodes, gen.n. When I examined Lamellipalpus nigripennis
Pascoe (Garo Hills, Mrs. Kemp) and Lamellipalpodes annandalet
Maulik at Calcutta, I was impressed by the fact that they both
possessed the remarkable lamellate palpal joints but did not
appear to. belong to the same family. Dr. C. J. Gahan thinks
that Lamellipalpodes Maulik does not belong to the Drilide,
but is more allied to the Rhagophthalmide, a neighbouring
family of about three genera, including nine species. Without
further research it is‘not possible to decide this point.

LAMELLIPALPODES ANNANDALEI, sp. n.

Upperside yellow-brown, underside lighter; antennz except

the two basal joints, and the elytra except an ill-defined basal.

portion (which shares the colour of the pronotum) smoky black.
Eyes deep black. The whole body is covered with brownish
pubescence.

Head narrower than prothorax, the interocular space piceous.
The first joint of antenne stout and large, second joint very
small and rounded, the third to the tenth similar in structure but
of gradually diminishing length, the eleventh bluntly rounded.
Prothorax broader than long, with a longitudinal median sulca-
tion; posterior angles acute, pointed, anterior ones rounded ;
basal area depressed on either side of the middle longitudinal
line, basal margin bisinuate. Scutellwm triangular, with the apex
rounded, brown, smooth. Zlyira as broad at the base as the
prothorax. Corresponding to the prothoracie depressions, the
elytra are also depressed at the base. The surface neither

|
|
:

NEW INDIAN DRILID BEETLES. 585

smooth nor regularly punctate-striate. Each elytron is feebly
tricostate; the costz are more visible if the insect is held at
certain angles; the first—that which is nearest to the suture—
proceeds from the apex of the basal declivity of the elytra, the

_Text-figure 4.

Lamellipalpodes annandelei Maulik.

Text-figure 5.

5 Bagche:
Underside of Lamellipalpodes annandelei Maulik.

second commences from the edge of the elytral black area, the
next commences about the middle of the elytron, each of these
terminating on the apical area of the elytron-

Length 53 mm.

586 ON NEW INDIAN DRILID BEETLES.

' Described from five examples.

Type in the British Museum.

- Cotype in the Indian Museum.

Of the five examples four belong to the British Museum, and
these have only ‘‘ Bengal” on the locality labels; the specimen
belonging to the Indian Museum was taken by Dr. N. Annandaie
from the following locality :—

Barkuda Island, Chilka Lake, Ganjam district, Madras Pre-
sidency, 22. vii. 1920; found on bath-room window.

In this specimen the cost of the elytra are more pronounced,

In preparing this paper I have had the advantage of seeing
specimens in the British Museum collection. I wish to thank
Dr. C. J. Gahan and Mr. K. G. Blair for their courtesy, valuable
suggestions, and criticisms. I wish to thank also Dr. N. Annan-
dale and Dr. and Mrs. Kemp for kindly letting me examine
their respective insects. The drawings of two insects were made
by Mr. 8. C. Bagchi, of the Indian Museum, Calcutta, and of one
by Mr. A. J. E. Terzi; to these artists my thanks are also due.

The preliminary study of these interesting beetles was made
in the Indian Museum, Calcutta, while I held charge of its
Entomological Section.

ON SOME ABNORMALITIES IN THE CARNIVORA, 587 |

32. On some Abnormalities in the Carnivora. By CHarEs
F. Sonntac, M.D., F.Z.8., Anatomist to the Society.

[Received May 10, 1921: Read May 10, 1921.]
(Text-figures 64-65.)

The following abnormalities were seen in animals dissected in
the Society’s Prosectorium during the past two years, They
consist of :—

1. Elongation and contortion of vessels and nerves in the neck
of a Bengal Fox (Canis bengalensis). :

2. The occurrence of two separate precaval veins in a Panda
(Ailurus fulgens).

3. Abnormal modes of termination of the vertebral veins of a
Common Badger (J/eles meles).

These conditions were compared with the normal state of the
vessels, which is identical in all three animals.

The Normal Blood-vessels.

The aortic arch gives off the innominate and left subclavian
arteries, and the former divides into the right subclavian and
right common carotid arteries after giving off the left common
carotid. The common carotid arteries run antero-laterally, and
then straight forwards. The internal jugular veins are on their
lateral aspects, and the vago-sympathetic nerve-cords lie on their
dorsal surfaces.

The external jugular veins unite with the subclavian veins to
form the innominate veins, of which the left one runs obliquely
across the anterior part of the thorax and joins the more vertical
right vein to form the precaval vein, The vertebral veins open
by single trunks into the innominate veins.

The vena azygos major opens into the right side of the precaval
vein immediately anterior to the root of the right lung.

Abnormalities in a Bengal Fou (text-fig. 64).

The animal, which was young, poorly-nourished, and rachitic,
had a symmetrical and uniform enlargement of both lateral
thyroid lobes (text-fig. 64, A.A), and the thymus gland was well-
developed. The former were about twice the size of the lobes in
a normal adult male. The isthmus of the thyroid gland was
normal,

The innominate artery (B) divided in the normal manner, but
the right subclavian artery was concealed by the right innominate
vein (C).

The left common carotid artery (D) has a more horizontal
course than the right one (EK), and describes a very acute flexure
with the convexity to the left. The right common carotid artery
is not only flexed, but recurved, with the result that there is
an S-shaped flexure. The left vago-sympathetic cord runs a

588 Dh. G. F, SONNTAG ON SOME

straight course on the dorsal surface of the left artery, but the
right cord (F) follows the right carotid in its wanderings, and
appears on the surface for a short distance.

No growths or adhesions are present in the root of the neck
which could be responsible for distortion of the vessels and

Text-figure 64.

aa
st

RUNEUEEH

SESS

—s

The vessels of the neck and thorax of Canis bengalensis.
For explanation of letters see Text-figure 65.

nerves. All other vessels and nerves are normal as regards their
course.

The precaval vein (1) receives internal mammary (N), peri-
cardiac (J), and azygos veins (M), and a vein from the diaphragm
running along the surface of the left phrenic nerve (1). No
phrenic vein accompanied the right phrenic nerve.

1
;

ABNORMALITIES IN THE CARNIVORA,. 589

The Occurrence of T'wo Precaval Veins in a Panda (text-
fig. 65 A).

The innominate veins were formed as usual, but soon gave way
to two precaval veins (L.L). The latter at first ran posteriorly
and slightly mesially to points level with the anterior limit of the
heart, when they turned directly inwards and ran horizontally ;
in this part of their course each vein receives an azygos vein (M).
The orifices of the two precaval veins and the wide postcaval
vein (‘I’) form the angles of a triangle with the apex posterior.

This arrangement of the precaval and postcaval veins is the
rule in Marsupials, but it occasionally occurs in higher. Mammals.
The chief difference between the condition described above and

Text-figure 65.

The veins of Ailurus fulgens (A) and Meles meles (B).

Explanation of Letters.

G. subclavian veins; H. intercostal vein; Ic.1 and Ic.2. intercostal veins;
O. esophagus: P. trachea; Q. right recurrent laryngeal nerve; R. internal
jugular vein; 8. inferior thyroid vein; U. internal jugular vein; V. vertebral
vein; W. externa] jugular vein. Other letters in text.

that of the Marsupials lies in the nature of the terminal orifices.
In the latter the openings are closer together, and the postcaval
and right precaval veins are only separated by a crescentic fold.
In the former they are wider apart, and the crescentic fold is not
present.

In the process of development the anterior cardinal veins from
the head and the posterior cardinal veins from the body unite to
form the Ducts of Cuvier, which open into the sinus venosus.
The anterior cardinal veins become united by a transverse vessel,
which takes a gradually increasing share in returning the blood
from the left side of the head. Kventually the part of the left
anterior cardinal vein between this vein and the sinus venosus

590 ON SOME ABNORMALITIES IN THE CARNIVORA.

atrophies, and the blood is conveyed entirely to the vein of the
right side. The part of the right anterior cardinal vein between
the cross vein and the sinus venosus becomes the precaval vein of
the normal animal.

I believe that the persistence of two precaval veins in this
animal is due to the non-formation of the cross vein.

No abnormality was found in the postcaval vein.

The veins of the father of this animal were quite normal.

Abnormalities in the Vertebral Veins of a Badger (text-
fig. 65 B).

The left vertebral vein bifurcates in the lateral triangle of the
neck; one of the divisions opens into the left external jugular
vein, and the other one opens into the left innominate vein.

The right vertebral vein runs posteriorly dorsal to the right
subclavian vessels, collects the veins from the first and second
right intercostal spaces, and opens into the precaval vein.

ON THE EXTERNAL CHARACTERS OF THE KOALA. 597

33. The External Characters of the Koala (Phascolarctos)
and some related. Marsupials. By R. I. Pocock,
FLR.S., F.Z.8:

[Received May 10, 1921: Read June 7, 1921.]
(Text-figures 22-26.)

CoNnTENTS. Page
MtVOOUCTIONM Nee Cor arnt er Seen we ar OL
AHR HTMar MING A eet Lae een ee Cee eta Dee
rep hacialeValbnisscocecs det cee eee ee eee OG
AHeMMiouibligye eco ete een ean oa eats L1G
Teh B aie ae ee SS POP LCE Moh Se er OG
RhepMore/ Hootie sass eas ee eas Sa LOS
MhegtHan dub oO ten 4-8 tyes sees Bat danke Pere ie ee (GOL
hewRouchvwas eh. cies Dettat nes LOO:
The Classification of crete. eratgdants, ees 606

Introduction.

The body of the female Phascolarctos described in the following
paper was very kindly sent to me by Mr. A. F. Richardson when
I explained to him the zoological importance of the animal and
the infrequeney with which fresh examples come into the hands
of anatomists in this country.

The need for renewed examination of Phascolarctos is especially
shown by the disagreement amongst modern zoologists regarding
the position and status to be assigned to it in the classification of
Diprotodont Marsupials. Thomas, for instance (Uat. Marsupialia
and Monotremata in Brit. Mus. 1888), divided the suborder into
three families: (1) Macropodide with the subfamilies Macro-
podinz, Potoroine, Hypsiprymnodontine ; (2) Phalangeride with
the subfamilies Tarsepedine (Varsipes), Phalangerine (Plelane ger,
Trichosurus, Pseudochirus, Petauroides, etc.), and Phascolarctine
(Phascolar hos); and (3) Phascolomyid (Phascolonys).

In connection with the Macropodide and Phalangeride, Thomas
pointed out that the characters of Hypsiprymnodon are inter-
mediate between those of the two families, and render the division
between them by no means so sharp and well-defined as it has
been usually considered to be. Nevertheless, his arrangement
suggests that Phascolarctos is an aberrant member of the
Phalangeridez more nearly related to Phalanger than the latter is
to Macropus.

Thomas's classification has been adopted by English zoologists
and by Trouessart in his Catalogue (1912).

Bensley (Amer. Nat, xxxv. pp. 117-138 and 245-269, 1901;
and Tr. Linn. Soc. Lond. (2) ix. pt. iii. pp. 83-214, 1903) followed
Thomas in admitting the family Phascolomyide and in regarding
Phascolarctos as the representative of a subfamily of the Phalan-
geride; but he made the important modification of transferring

Proc. Zoou. Soc.—1921, No. XL. 40

592 MR. R. I. POCOCK ON THE

Pseudochirus and Petawroides from the Phalangerine to the
Phascolarctinz because of the crescentic (subselenodont) pattern
of the molar teeth in the three genera. Bensley’s classification
was’ adopted, presumably with approval, by W. K. Gregory
(Bull. Amer. Mus. Nat. Hist. xxvii. pp._215-216, 1910), who was
acquainted with but rejected the classification by Winge quoted
below.

Thomas, of course, was quite familiar with the dental character
to which Bensley and Gregory attached so much importance.
On p. 167 of his Catalogue he pointed out that by the compli-
cated subselenodont character of its molars, Psewdochirus, with
its close ally Petawroides, stands somewhat apart from most of
the other Phalangers, and approaches Phascolarctos, in which a
similar but simpler modification is observable. But his reasons
for attaching to it subordinate systematic value lay apparently in
the circumstance that in young examples of Phalanger a tendency
towards the same structure is visible, but the crests on the molars
- goon wear off, leaving little difference between them and the
simple quadricuspid molars characteristic of typical genera of
Phalangeride. The obvious, but not on that account necessarily
true, inference to be drawn from this fact is that the tendency
towards the subselenodont molar pattern exhibited by Phalanger
is a primitive character of the family Phalangeridee, which is tem-
porarily retained in Phalanger, lost in Trichoswrus, Dactylopsila,
and others, and elaborated in Pseudochirus and Phascolarctos.

Winge (E Museo Lundii, viii. pt. 1, 1893) held very different
views. He adopted two families: (1) Phalangistide |= Phalan-
geride] with the subfamilies Pseudochirini for Psewdochirus
and Petauroides, and the Phalangistine [= Phalangerine; for
the two groups Phalangistee (Phulanger, Trichosurus, Petaurus,
Tarsipes, etc.), and Macropodes (Macropus, Hypsiprymnodon,
ete.); (2) Phascolarctide with the subfamilies Phascolarctini
(Phascolarctos) and Phascolomyini (Phascolomys) oe

Winge took as the basis for his classification the degree of
extension of the tympanic process of the alisphenoid, which in
the Phascolarctidee (Phascolomys, Phascolarctos) is small, does not
envelope the tympanic cavity, and fails to reach the paroccipital
process, whereas in the Phalangeridw, comprising the rest of the
genera of Diprotodont Marsupials, the bone in question is large,
envelops the tympanic cavity, and reaches the paroccipital process.
As accessory characters, the vestigial tail, the presence of a cardiac
gland in the stemach, and the loss of one of the two normal
pairs of teats further serve to distinguish the Phascolarctide
from the Phalangeride 7.

%* To these Winge added Thylacoleontini (Thylacoleo) related to Phascolarctini
and Diprotodontini (Diprotodon, Nototherium) related to Phascolomyini.

+ Winge’s valuable paper is unfortunately written im Danish. The statements
about the extension of the alisphenoid is taken from Max Weber’s work. I have
not been able to verify it im the case of the two skulls of Phascolarctos available
for examination.

EXTERNAL CHARACTERS OF THE KOALA. 593

Attention may also be drawn to Winge’s severance of the
erescent-toothed or subselenodont Phalangers (Pseudochirus and
Petauroides) from the rest as representing a special subfamily
Pseudochirini equivalent to the’ Phalangerine containing not
_ only the rest of the genera Thomas assigned to that subfamily,
but all of those constituting the family Macropodide of that
author.

Max Weber (Die Siug. p. 348, 1904) followed Winge in
admitting the two families Phalangeride and Phascolarctide as
he defined them; but he did not admit the subfamily groups
Phascolarctine and Phascolomyine for Phascolarctos and Phasco-
Jomys respectively, and in the case of the Phalangeride he made
a compromise between Thomas’s and Winge’s systems by dividing
the family into three subfamilies : se Mlk n peas, Hypsiprymno-
dontine, and Macropodine.

It will be noticed that neither Winge nor Max Weber attaches
any particular importance to the structural peculiarities of
Tarsipes.

‘The divergence of opinion between the authors quoted may be
briefly and more clearly expressed by the following tabulation of
their classifications :—

Thomas. Family MACROPODID.
Subfamily Macroropin# (Macropus, Dendrolagus, etc.).
Potoroinz (Potorous, Bettongia, etc.).
55 HyYPsiPRYMNODONTIN® (Hypsiprymnodon)-

Family PHALANGERID A.

Subfamily Tarsrpepinm (Tarsipes).
PHALANGERIN® (Phalanger, Trichosurus, Pseudo-

chirus, Petauroides, etc.).
x PHASCOLARCTIN® (Phascolarctos).

Family PHASCOLOMYID (Phascolomys).

Bensley. Family MACROPODID (Macropus, Hypsiprymnodon, etc.).
>» PHALANGERIDA.
Subfamily Tarsrpepinz (Tarsipes).
PHALANGERINE (Phalanger, Trichosurus, etc.).
PHASCOLARCTINE (Pseudochirus, Petauroides,
Phascolarctos).

Family PHASCOLOMYID (Phascolomys).

Winge. Family PHALANGERIDE*.
Subfamily PHALANGERINE.
a. Macropodes (Macropus, Hypsiprynnodon, etc.).
b. Phalangeri (Larsipes, Phalanger, Trichosurus).
Subfamily PspeupocurIrinw (Pseudochirus, Petauroides).
Family PHASCOLARCTID As.
Subfamily PuascoLarcrinz (Phascolarctos).
PHAscoLoMYinm® (Phascolomys).

* Por the sake of clearness in comparison, I have here altered the name Phalan-
gista and its derivatives used by Winge to Phalanger and its derivatives.

40*

59A MR. R. I. POCOCK ON THE

Weber. Family PHALANGERIDA. C
Subfamily Macropopinm (Macropus, Dendrolagus, Potovous,
Bettongia, etc.).

_ HypsIPRYMNODONTIN® (Hypsiprymnodon).
5 PHALANGERINE (TLarsipes, Phalanger, Trichosurus,

Pseudochirus, Petauroides).
Family PHASCOLARCTID A (Phascolarctos, Phascolomys).

Kuternal Characters.

The Rhinarium.—In Trichosurus the rhinarium is large, naked,
and convex above; the infranarial portions are complete laterally
and narrow; there is a median groove extending between the
nostrils on to the philtruam, which is divided into two narrow
strips, with a little process, overlying the gum between the median
incisors, at the apex of the angular excision above. The nostrils
are of the typical form, consisting of a rounded orifice in front,
moderately widely separated iran its fellow of the opposite side,
and of a lateral aud posterior narrow slit.

In Phalanger the rhinarium is in a general way similar to that
of Trichosurus, but the nostrils ave more widely separated, the
infranarial portion is deeper in front, and the philtrum 1S eX-
ceedingly wide, and, although gradually narrowed below, is quite
wide where it terminates on the edge of the upper lip. There is,
moreover, no angular excision such as is seen in Z'richosurus, but
the median groove broadens below, its floor terminating in a
median process which projects at least as low as the lateral
portions of the philtrum.

In Pseudochirus the rhinarium is more like that of T'richo-
surus, but the infranarial portions are wider in front, with the
inferior edge more steeply inclined ; the median groove is con-
tinued between the nostrils to the summit of the rhinarium; and
the inferior portion of the philtrum is quite narrow, with a small
median notch.

In Phascolarctos the nose is widely different. The muzzle is
abruptly sloped downwards about an inch above the nostrils, and
the whole of this inclined area is covered with very fine, short,
scattered hairs, leaving a narrow, naked, thickened 11m round the
nostrils. Thus there is no true rhinarium. The nostrils them-
selves are also peculiar. Hach consists of an ovally elongated
valvular orifice, with a thickened superior and lateral rim, but.
with the inferior rim hardly defined from the upper lip; all
trace of differentiation into lateral slit and anterior orifice has
disappeared ; the septum is very narrow, and is continued in-
feriorly between the two halves of the upper lip, ending below in
a process over the gum between the median incisors.

In Phascolomys ursinus* the rhinavium is quite unlike that of
Phascolarctos, and resembles, broadly speaking, the rhinarium

* The rhinarium of Lasiorhinus latifrons, the hairy-nosed Wombat, which
should, I think, rank as a distinct genus, is unknown to me except from descriptions

and a dried skin. It appears to differ solely from that of Phascolomys in being
covered with short hair, the nostrils being normally formed.

EXTERNAL CHARACTERS OF THE KOALA. 5G5

of the three genera of Phalangers above described. It extends
dorsally, however, further backwards in the middle line, and the
hair of the muzzle encroaches over the well-developed slit of the
nostrils, the expanded portions of which are widely separated in
the middle line. The infranarial portions are moderately deep,

and reach almost to the posterior end of the nostril-slits.

Text-figure 22.

. Head of Pseudochirus peregrinus, with pocket of ear (a) seen from above.
. Rhinarium of the same.

. Ear of Trichosurus vulpecula.

. Rhinarium of the same, from the front.

. Rhinarium of the same, from the side.

. Head of Phalanger maculatus, the ear stripped of hair.

Rhinarium of the same, from the front.

Qertoawe

Their inferior edge slopes obliquely downwards and inwards to
the gum of the incisors, the two lobes of the upper lip being

596 MR. R. I. POCOCK ON THE

tolerably widely separable at this point. There is no median
excision in the inferior edge of the rhinarium above the teeth,
and there is no median groove, but a groove runs on each side
from the nostril downwards and inwards to the gum, defining
the infranarial portion from the rest of the rhinarium.

Facial Vibrisse.—The facial vibrissee are well and normally
developed in Z'richosurus, Phalanger, Pseudochirus, and Phasco-
lomys, being represented by numerous long mystacials, super-
ciliaries, genals—one tuft,—submentals, and Unterramals. In the
examples of Phalanger and Pseudochirus examined, however, the
interramals were few in number and slender. In Phascolarctos
the vibrisse are comparatively poorly developed, the mystacials.
especially being few and short and the interramal tuft wanting.

The Mouth—There are only two points for me to notice in
connection with the mouth. The first is the presence, with which
everyone is familiar, of cheek-pouches in Phascolarctos. These lie
alongside the gum of the upper jaw in front, the orifice looking
downwards iit the cavity extending upwards and backwards.
towards the eye. No such pouches occur in 7 richosurus, Pha-
langer, Pesnialoaiar us, or Phascolomys. The second is the presence
in Phascolomys of a flap of naked skin extending into the mouth
from the inner surface of the lips laterally. This 1s analogous to-
the similar ingrowths well known in the Rodentia, where they
serve to close the throat during gnawing. Their function is no
doubt the same in Phascolomys, Ghich has rodent’ incisor teeth.
These oral flaps of skin are not developed in the other genera
here discussed, nor so far as I am aware in any other Marsupial.

a. A flap of skin jutting inwards from the lower ip and lying

alongside the tongue on each side .............. veseseeueees. Phascolomys.
a’. No flap of skin jutting into mouth from the ie
6. A cheek-pouch on each side of the upper jaw .................. Phascolarctos.
bP Niovcheek-pouchessers: 15 seer eee eek eeeees ee .... Pseudochirus,

~ Phalanger, Dr PenGR cee f

The Ear.—In Trichosurus the ear is high, but tolerably narrow
and nearly naked internally. The supratragus (metatragus) is.
a well-developed thick, semicircular, slightly upturned disk.
Above it there is a low curved ridge. In front of it the cavity
of the ear is defined by a strong oblique ridge passing upwards.
slightly above the low ridge above described and inferiorly
curving backwards beneath the supratragus and passing thence
downwards on the inner side of the well-developed tragal
thickening forming the anterior rim of the inferior notch (aditus
inferior), the posterior border of which is defined by a strong
thick antitragal ridge, capable of being turned outwards and
backwards to a certain extent; but it exhibits no pocket-like
depression.

In Pseudochirus peregrinus the ear is nearly as naked intern-

ally, and relatively nearly as high as in Jr ichosurus but somewhat
broader. ‘The supratragus is as well developed, but forms a more
distinct roof to the deep lower portion of the cavity. The low

EXTERNAL CHARACTERS OF THE KOALA. 597

ridge seen in Zvrichosurus is represented by a lobate thickening,
and there is a similar thickening lower down behind, above the
posterior end of the supratragus. The anterior ridge forms a
lobate expansion above the anterior end of the supratragus,
and the tragus itself is less well defined. The ridge behind the

Text-figure 23.

A. Head of Phascolarctos cinereus, with position of cheek-pouch (p) dotted in and
pocket of ear (a) seen from above.

B. Nose and part of palate of the same, showing the nostrils and the orifices of the
cheek-pouches (p).

C. Nose and mouth of Phascolomys ursinus, with lower incisors cut short and
flap of skin (f) projecting into mouth.

D. Side view of rhinarium of the same.

E. Ear of Lasiorhinus latifrons, from dried skin softened in water.

598 MR. R. I. POCOCK ON THE

inferior notch turns inwards above towards the posterior end of
the supratragus, and it is provided with a shallow pouch defined
externally by a definite thickened ridge.

The ear of Phalanger maculatus is greatly reduced, thickly
covered with hair externally, and scarcely projects above the level
of the head. The supratragus is large and valvular, and the
cavity of the ear above it is deep and roofed over by a ridge
resulting possibly from the confluence of lobate thickenings
homologous to the two seen in Phalanger. The anterior ridge
does not extend so high above the supratragus as in the two
genera already discussed, and there is a trace of the pouch on
the posterior ridge, although it is not so well defined as in
Pseudochirus.

In Phascolarctos the ear is more expanded even than in
Pseudochirus, but it differs from the ear of that genus, 7'richo-
surus, and Phalanger in having the supratragus reduced to a low
curved ridge without any trace of the lobe*. There is a low
ridge above it as in Z'richosurus, and the anterior ridge bounding
the cavity of the ear in front is as in that genus; the tragus,
however, is much smaller. The pocket on the posterior ridge is
present as in Pseudochirus, but is deeper and has a more sharply
defined posterior rim.

In Phascolomys the antero-internal ridge, continuous above
with the slightly overfolded anterior margin of the pinna, is
produced inferiorly into a large compressed lamina, jutting back-
wards into the cavity of the ear and descending to the auditory
orifice. Anteriorly and externally this lamina is marked by a
deep groove which descends and passes into a low ridge defining
the anterior border of the inferior notch, but there is no definite
tragal thickening. The antitragal ridge is also simple. It
ascends and forms the posterior edge of the cavity of the pinna,
and is itself marked externally and posteriorly by a groove.
The supratragus appears to be represented by a low ridge visible
above the laminate expansion of the antero-internal ridge.

The ear of Lasiorhinus latifrons is very like that of Phascolomys,
but is longer. On askin softened in water I could find no trace
of the supratragus.

The ear of Phascolomys resembles that of Phascolarctos in the
complete or almost complete disappearance of the supratragus—
an important character considering the constancy in the deve-
lopment of this ridge in many Mammals. But in -the simple
structure of the posterior antitragal ridge it is more like that of
Trichosurus.

The differential characters of the ear may be used to define and
classify the five genera as follows :—

a. Supratragus represented by a strong ridge with well-developed
lobate thickening.

6. No pocket above the antitragal ridge ..........seccceuseee wees Trichosurus.

* Thomas mentioned this peculiarity and wrote ‘‘metatragus almost obsolete.”

EXTERNAL CHARACTERS OF THE KOALA. 599

b’. A pocket above the antitragal ridge.
c. Pinna greatly reduced; pocket small ...............0.05. Phalanger.
ce’, Pinna large; pocket better developed ............ 06000000000 Pseudochirus.

a’. Supratragus at most represented by a low curved ridge;
without trace of lebate thickening.

d. Antero-internal ridge normally developed; a large pocket
abovethe antiinaral rid cert comes cer slavaeneitntentnse os eietan le) Phascolarctos.

d'. Antero-internal ridge large and laminate; no pocket above

amtifvagall tGee wes cia sud sdoves cocclugeer Gandy ced tatebcrahadcss sun mR ASCOLOMYS.

The Fore Foot.—In Trichosurus, Phalanger, and Pseudochirus
there is a tuft of long carpal vibrisse. In Phascolarctos and
Phascolomys these tactile bristles are absent.

In Trichosurus the fore foot presents no special modifications.
The five digits are free of webbing down to the plantar pads and
are nearly evenly spaced, the third and fourth being the longest
and subequal, the second and fifth a little shorter but subequal
and rising at approximately the same level, and the first or
pollex much the shortest, about two-thirds the length of the third
or fourth, and not opposable but closing obliquely backwards and
inwards across the sole (palm). The claws of all the digits are
compressed, curved, and pointed, and the integument of the
lower side of the digits is transversely grooved. The uniformly
granular plantar pad is four-lobed, the three main lobes being in
contact. The inner or pollical lobe, sometimes separated from
the main lobes, is fused with the inner element of the carpal pad
into a longitudinal mass, longer than wide. The external element
of the carpal pad is elliptical, narrowly separated in front from
the external lobe of the plantar pad, and, like the internal element,
passes imperceptibly proximally into a narrow naked area of skin
between them and the hair of the wrist. The centre of the sole
is depressed and granular like the pads and the depressions
between them.

In Phalunger the fore foot is considerably modified from the
type seen in Zrichosurus. The third, fourth, and fifth digits are
evenly spaced; but the second is capable of being separated from
the third by a much greater space than that between the third
and fourth. With the pollex it is capable of being extended
almost at right angles to the axis of the foot and of being closed
transversely upon the sole. The pads are transversely striate,
with the spaces between them granular, the space or groove
between the second and third being deeper than that between the
third and fourth. The first, the internal or pollical lobe, forms a
continuous subtriangular mass, directed obliquely backwards and
inwards when the pollex is drawn back, and the external moiety
of the carpal pad is larger than in Zrichosurus.

The capacity for co-operative movement of the first and second
digits in a plane nearly at right angles to that of the third,
fourth, and fifth seems to have been overlooked in Phalanger
maculatus. Bensley, at all events, records the similar phe-
nomenon in Pseudochirus and Phascolarctos as peculiar to these
two genera. Nevertheless, the modification in question may be

600

MR. R. I. POCOCK ON THE ;

Text-figure 24,

. Right fore and hind foot of Trichosurus vulpecula.
. aie % Phalanger maculatus.
5 oy i) Pseudochirus peregrinus.

x F-

EXTERNAL CHARACTERS OF THE KOALA. 601

clearly seen even on dried skins; and it is shown in the figures
of the Spotted Cuscus in the Royal Natural History, iii. p. 257,
1894, and Harmsworth’s Natural History, ii. p. 888, 1910.
These figures bear the impress of having been drawn from life by
Mitzel and Kiihnert respectively.

The fore foot of Pseudochirus resembles tolerably closely that
of Phalanger, except that the third digit is not lengthened and
the lobes of the plantar and carpal pads are separated and smaller,
the fused pollical element of the plantar pad and the inner moiety
of the carpal pad forming an oblique transverse mass about twice
as wide as long.

The fore foot of Phascclarctos is an extreme exaggeration of
the type seen in Phalanger and Pseudochirus, although the pads
and intervening spaces are areolated or granular, not striated.
The sole is much longer as compared with its width: the first
and second digits are completely isolated from the rest, and rise
close together from the postero-internal angle of the foot, at right
angles to its long axis and close transversely across the proximal
half of the sole, the posterior border of the pollex when drawn
back being approximately in the same transverse line as the
posterior border of the sole. There is a large three-lobed plantar
pad at the base of the third, fourth, and fifth digits, and there is
a similar but smaller lobe upon the base of each of the second
and first digits. The one on the pollex probably represents the
inner moiety of the carpal pad, and the one on the base of the
second digit the first or external element of the plantar pad,
widely severed from the lobe in front of it and altogether discon-
nected from the pollex behind it. The external moiety of the
carpal pad is small, restricted to the postero-external angle of
the foot and widely separated from the plantar pad.

In Phascolomys the fore foot, modified for terrestrial progres-
sion and digging, is very different from that of the preceding
genera, especially from that of Phascotarctos. It may be derived
from the type seen in Zrichosurus by the shortening and widening
of the digits and sole, the obliteration of nearly all trace of the
individual elements of the pads, and by the straightening,
lengthening, and blunting of the claws.

The Hind Foot.—In Trichosurus*, Phalanger, and Pseudochirus
there are one or two tactile vibrisse, similar to the carpal vibrisse,
on the inner side of the heel behind the hallux. These are
absent in Phascolarctos and Phascolomys.

The hind feet of the four arboreal genera—Trichosurus,
Phalanger, Pseudochirus, and Phascolarctos—difter from each
other much less than the fore feet. Their general structure is
well known. Hence only the comparatively minor points of
difference need be noticed.

* Represented in the figure of the hind foot of Psewdochirus cooki (pl. 20. fig. 4,
of Waterhouse’s ‘ Marsupiata’), but apparently unnoticed in the text.

602 MR. R. I. POCOCK ON THE

In Trichosurus the conjoined second and third digits* are
much shorter than the fourth. The three lobes of the plantar
pad at the base of the terminal digits are in contact, the outer
lobe being larger than the other two taken together. The large
lobe of the hallux shows no trace of division, and there is a
tolerably well-defined, long external metatarsal pad reaching
from the heel almost to the plantar pad.

The digits of Phalanger are approximately as in Trichosurus;
but the external lobe of the plantar pad is separated to a certain
extent in front from the other lobes, which are fused although
defined by a groove, and are together larger than the outer lobe.
The entire hallux is relatively larger than in T’richosurus, and
its great pad is indistinctly divided; there is no distinct external
metatarsal pad and the heel is shorter than in 7’richosurus.

In Pseudochirus the digits are as in the foregoing genera, but
all the elements of the striated pads on the sole are better defined
even than in 7richosurus. The outer lobe of the plantar pad is
completely isolated from the two conjoined inner lobes and about
half their size. The great pad of the hallux is distinctly divided,
into a distal and a proximal portion, the latter representing the
inner metatarsal pad; the outer metatarsal pad is well developed,
sharply defined, and about twice as long as wide.

In the foregoing genera the hind foot is larger in every way
than the fore foot.

In Phascolarctos the hind foot is not larger than the fore foot.
It differs from the hind foot of the other genera in having the
united second and third digits relatively much longer and only a
little shorter than the fourth; the Jobes of the plantar pad are
feebly differentiated, being fused to form a transverse cushion,
with irregularly convex anterior and irregularly concave posterior
border. The great pad of the hallux is undivided, and there is
no distinctly defined external metatarsal pad. Thus in the
development of the pads the foot of this genus differs more from
that of Pseudochirus than from that of Trichosurus.

The hind foot of Phascolomys, as has often been pointed out,
is a fossorial modification of the Phalangerine scansorial hind
foot, resulting from the conversion of the sharp, curved claws of
the latter into longer, stronger, and straighter claws, from the
approximate equality in length between the fourth and the united
second and third digits, the reduction in bulk of the great pad
of the hallux so that it projects only slightly beyond the inner
margin of the foot, with the terminal segment of the digit planted
like a hemispherical tubercle upon it, and from the practical

* These two united digits act as a fur-comb in Marsupials. Possibly they were
primarily modified for that: function. Generally they are too small in arboreal
forms to be of much use fer grasping. But this cannot be maintained in the case
of Phascolarctos; and in Phascolomys they are large enough to be subservient to
digging. But in the Kangaroos they appear to be retained solely for the purpose
mentioned, and may frequently be seen to be so employed. Moreover, it is signifi-
cant that these are the only digits in Tarsipes which have other than rudimentary
claws.

EXTERNAL CHARACTERS OF THE KOALA, 603

obliteration of the pads with expansion of the heel; but, as in
the Phalangerine foot, the fifth digit is more widely separated
from the fourth than the latter is from the third, although the
digit in question is relatively shorter and the space above
mentioned somewhat wider.

Text-figure 25.

\
\
\

B. Right fore and hind foot of Phascolarctos cinereus.
D. 53 Ss Phascolomys ursinus.

604 MR. R. I. POCOCK ON THE

By their feet the genera may be distinguished as follows :—

a. Feet fossorial. with digits short, claws long, blunt and
slightly curved, and the granular pads but itttle differen-
tiated ; hallux short, its distal phalange reduced to a button-
like excrescence on the shortened lobe, which, however, still
retains the capacity for movement in the opposable plane ... Phascolomys.
a'. Feet scansorial and preensile, with longer digits and sharp,
curved claws, and differentiated pads; hallux very large and
opposable, with well-developed terminal phalange.
b. Fore foot as large as hind foot, with comparatively long and
narrow sole; the inner Icbe of the plantar pad small aud
attached to the base cf the second digit, which is separated
by a long space from the third; the imuer lobe of the
carpal pad small, isolated from the outer and attached to
the base of the first digit (pollex) ; second and thivd digits
of hind foot comparatively large, strongly prekensile ..... Phascolarctos.
b', Fore foot smaller than hind foot, with shorter, broader sole ;
the inner lobe of the plantar pad large, in contact, or
nearly so, with the rest of the pad and fused into one mass
with the inner lobe of the carpal pad, there being no differ-
tiated pad at the base of the first digit (pollex) ; second
and third digits of hind foot short, weakly prehensile.
c. A comparatively long space between the third and second
digits of the fore foot, the second actimg in unison with
the firzt like a double opposable pollex; pads striated ... Phalanger and

Pseudochirus.
c/. The five digits of the fore foot evenly spaced, the second
in no respect opposable to the next, and the first or pollex
only slightly so; pads granular..............:0::-.eee Trichosurus.

The Pouch.—I\ have seen no fresh female examples of Phalanger
and Pseudochirus.

In Trichosurus the pouch, as in J/acropus, is deeper than
wide, its orifice has a well-developed lateral and posterior rim
but no overhanging anterior rim; it therefore leoks forwards * ;
and in the specimen examined there were only two teats,
not four, as stated by Winge to be characteristic of the
Phalangeride.

In Phascolomys the pouch is a little wider than long, being
deeper laterally, especially anteriorly, than behind or mesially in
front, and a little deeper behind than mesially in front, the
muscular rim overhanging the cavity all round; and there is a
single pair of teats.

In Phascolarctos the pouch, as recorded by Forbes, is much
wider than long, being extended, gradually narrowing as it goes,
along the depression between the muscles of the thigh and
abdomen nearly as far as the edge of the flap of integument
joining the hind leg to the body, its width being about three
times its length. The orifice looks slightly backwards, the
muscular rim overhanging the cavity laterally and to a slight
extent in the middle line in front, but ceasing on each side close
to the position of the single teat. The orifice therefore looks

backwards.

% In his volume on Marsupials (Allen’s Naturalists’ Library, p. 76, 1894),
Lydekker wrongly states that the orifice of the pouch is directed backwards in
Phalangeride.

EXTERNAL CHARACTERS OF THE KOALA. 605

From the description of these three pouches it is evident that
the pouch of Phascolomys is structurally intermediate in many
respects between those of Vrichoswrus and Phascolarctos. The
differences between the pouches of the last two genera, consider-
ing that both are arboreal forms, is remarkable, especially in view
of the theory that has been advanced that the anterior upward
aspect of the orifice is an adaptation to arboreal life or to the

Text figure 26.

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Way

Nes

ig "| oy ae WNL
CY a CH iohss NHS

§ th TR sant .

(<— m %. Rijs F at
— CI mee enon ‘ ( =
Sen ere ar ry QJ NY)? W)C ———— DAS =
RSs Raia y XS HS } ~7r :~
ger 7 a te (PA QIOR LN? hea Se 2 ia ‘o
“2G ec a ol wo 4/)) wy 4 Fe al Gat wea & (Ss
thd et b t A 7 = Se SS =
AAA) (7
Sis
ZS k roan
JE NN Wesnes
SAS7 ae
Ss VU WAY 528%
(277 i IN <i Fan
Yai Zz, All| WP aie Nas

A. Pouch of Phascolarctos cinereus.
B. 5 Phascolomys ursinus.
C. i Trichosurus vulpecula.

The extension of the pouch beneath the integument with the mamme (m)
at its posterior end shown diagrammatically with dotted lines.

more or less upright attitude assumed by such terrestrial bipedal
forms as Macropus, and conversely that the posterior aspect of
the orifice is an adaptation to the terrestrial life and quadru-
pedal gait of such genera as Thylacinus and Sarcophilus. This
theory entirely breaks down in the case of Phascolarctos, per-
haps the most specialized scansorial genus in the entire order,

606 MR. R. I. POCOCK ON THE

The deep, backwardly directed pouch of Trichosurus seems so
well adapted for the safe retention and carrying of the young up
in the trees that its replacement by a laterally directed pouch
with the orifice opening towards the tail, such as is seen in
Phascolarctos, seems most unlikely. The use of the lateral
extension for lodging the young carried by a mother who doubt-
less climbs at times head-downwards is easy to imagine, but the
reason for the reversion of the orifice demanded by the theory of
the descent of Phascolarctos from the Phalangerine stock is
puzzling. The structure of the pouch alone supplies evidence
that Phascolarctos is not closely related to any genus of the
Phalangeride.

The characters of the three types of pouches above described
may be summarized as follows :—

a. The orifice, not encircled behind, opening backwards and

downwards; cavity of pouch extended laterally on each
side alone the depression between the thigh and the

Enhoys Kovaaycsulamvepieet: carlo ees teye bee YR AC RIS Ra aIAce MMA cia est oe al Phascolarctos.
}. The orifice subcircular, surrounded by a flap of skin all

round; cavity subcireular, a little deeper laterally than

elsewhere ............ Phascolomys.

ce. The orifice opening forwards, at the anterior end of the

cavity, which is longer than wide Pseudochirus,

Phalanger, Trichosurus.

The Classification of existing Diprotodonis.

My opinion on the classification of the Diprotodonts is as
follows :—

1. Winge’s removal of Phascolarctos from the Phalangeride
must be accepted. But although that genus shows points of
resemblance to Phascolomys not shared by other existing Dipro-
todonts, the differences between them are teo many and too
important to admit of their ascription to the same family.
Moreover, if we adopt as criteria of family rank such characters
as those distinguishing the Kangaroos from the Phalangers, the
characters separating the Koala and the Wombat should, I think,
be given superfamily rank indicated by the titles Phascolarc-
toidea and Phascolommyoidea. Assuming Winge to have correctly
placed the extinct forms above referred to, the Phascolarctoidea
will contain the two families Phascolarctidee (Phascolarctos) and
Thylacoleonide (Thylacoleo); and the Phascolomyoidea the two
families Phascolomyide (Phascolomys, Lasiorhinus, Phascolonus)
and Diprotodontide (Diprotodon, Nototheriwm).

2. The two above-mentioned superfamily groups are individu-
ally equivalent to a group of that rank, containing the rest of
the Diprotodonts, for which two names are available, Hypsi-
prymnoidea and Phalangeroidea. I prefer the latter on account
of its greater familarity and its derivation from the name of a
more primitive genus. Accepting for this group Thomas’s families
Macropodide and Phalangeride, it seems clear in the case of the
former that if the characters of Potorous and its allies entitle

EXTERNAL CHARACTERS OF THE KOALA, 607

them to rank as a subfamily—and I see no reason for dissenting
from that view,—the characters of Hypsiprymnodon must be
given the higher rank of a family, the Hypsiprymnodontide
equivalent to, and standing between, the Macropodide and the
Phalangeride, but not definitely assignable to either.

From the typical Phalangeride constituting the subfamily
Phalangerine, the genera Psewdochirus and Petauroides, character-
ized by the crescentic pattern of the molar teeth, may be separated
as the Pseudochirine as proposed by Winge. But it does not
appear to me that proper appreciation has been shown for the
characters of the very highly specialized genus Tarsipes *, which
differs profoundly from the Phalangeride in the structure of the
skull, teeth, tongue, snout, and alimentary canal, as Thomas re-
corded. The genus seems to he at least as widely divergent from
the Phalangeridew as are the Macropodide. These two families,
indeed, are linked by the Hypsiprymnodontide, and the
evolutionary stages by which the Kangaroos have been derived
from the Phalangerine stock may be traced with reasonable
certainty through living forms. But all the hypothetical inter-
mediate genera between Tarsipes and the Phalangers have died
out, leaving that genus isolated. And since, in my opinion, it
differs more from the typical Phalangers than do the Pseudo-
chirines, it may be raised at least to the rank of a family—
Tarsipedide.

Briefly, the existing genera of Diprotodonts may be classified
as follows :—

Phalangeroidea.
‘Fam. MACROPODIDE.
Subfam. MAcRoPoDIN&.
3 PoToROINA.
Fam. HyPpsSIPRYMNODONTIDA.
Fam. PHALANGERIDZ.
Subfam, PHALANGERINA,
PSEUDOCHIRINA,
Fam. TARSIPEDIDZ.

Phascolarctoidea.
Fam, PHASCOLARCTIDA,

Phascolomyoidea.
Fam. PHASCOLOMYIDA,

* Winge and Bensley even considered the characters of this genus to be of less
systematic value than the concentric molars of Pseudochirus.

Proc. Zoou. Soc.—1921, No. XLI. Al

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EWART, Pl. I.

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INPESIOOONG IVE/MUSVIRS Ol Wists MUA IC Ms |B),

P. Z. S. 1921, EWART, Pl. II.

NESTLING FEATHERS OF THE MALLARD,

NESTLING FEATHERS

2 4, So 12, EW AIRIT,

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NESTLING FEATHERS OF Wish. IMUNEIL/NE ID),

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NESTLING FEATHERS OF THE MALLARD.

VII.

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NESTLING FEATHERS OF THE MALLARD,

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NESTLING FEATHERS OF THE MALLARD.

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NESTLING FEATHERS OF THE MALLARD.

ON THE NESTLING FEATHERS OF THE MALLARD. 609

34, The Nestling Feathers of the Mallard, with Observations
on the Composition, Origin, and History of Feathers.
By J. Cossar Ewart, M.D., F.R.S., F.Z.8., Regius
Professor of Natural History, University of Edinburgh.

[Received March 7, 1921; Read March 22, 1921. |

(Plates I.-X.; Text-figures 1-14.)

I. Tue Nestiinc FEATHERS OF THE MALLARD (ANAS BOSCAS).

In a recent paper in ‘ The Ibis’ it is pointed out that “ there
is no branch of ornithology that has remained so long neglected
as the study of nestling birds, nor is there one in which so many
problems await solution” (1). That the study of nestling feathers
has been neglected will be admitted when it is mentioned that up
to 1906 it was not realized that the true feathers (pennz) may be
preceded by two generations of nestling feathers (prepenne), that
ornithologists have not yet made up their minds whether the
familiar coat worn by newly-hatched chicks and ducklings corre-
sponds to the first (protoptile) or to the second (mesoptile)
nestling coat of Penguins, and that morphologists have not yet
ascertained whether nestling and other feathers originally con-
sisted of one shaft or of two complete shafts like the nestling and,
adult feathers of the Emu.

True or definitive feathers vary greatly in size and structure ;
nevertheless, they may be said to belong either to a plumose
(metaptile) or to a pennaceous (teleoptile) type. The plumose
true feathers are characterized by barbules which may possess
cilia but never have hooklets (hamuli); in pennaceous feathers
some of the barbules are armed with hooklets, by means of which
the barbs are ‘‘ woven” into a web—in the wing-quills of flying
birds we have highly specialized teleoptiles; in the plumose
feather from a Mallard represented in PI. I. fig. 1 we have an
example of a simple metaptile, bearing a protoptile.

In Penguins all the true feathers are probably preceded by
both protoptiles and mesoptiles (text-fig. 3), but in many birds
there is only one nestling coat—the mesoptiles have either been
completely suppressed or are represented by inconspicuous and
barely recognizable vestiges.

In most birds there are numerous hair-like feathers associated
with, and intimately related to, the true feathers. As these”
hair-like feathers are preceded by minute prepenne—are, in fact,
degenerate pennz,—they might be known as filopenne.

Further, in many birds there appear between the true or
contour feathers (penne) true down feathers (plumule). In all
the Ducks, Geese, and Penguins examined the plumule are
preceded by preplumule,

41*

610 PROF. J. COSSAR EWART ON THE

The relation of the various kinds of feathers of mature birds to
nestling feathers is indicated in the following table :—

Papille.
Filamenta
(Cryptoptiles).
Prefiloplumee I. Prepenne Preplumule.
(Prefilopenne). (Protoptiles).
IJ. Prepennee
(Mesoptiles).
Filoplume Plumose Pennee Pennaceous Penne Plumule.
(Filopenne). (Metaptiles). (Teleoptiles).

1. The Prepenne.

The most concise and authoritative statement I have come
across about nestling feathers occurs in ‘ Newton’s Dictionary of
Birds.’ It is there stated that neossoptiles (2. e. protoptiles) ‘ are
characterized by (1) a very short calamus, (2) an insignificant or
ill-defined rhachis—if there be one at all, (3) the almost universal
absence of cilia, (4) long and slender rami (barbs), and (5) the
absence of an aftershaft except in Dromzeus”’; in another para-
graph it is stated that the barbules (radii) of the aftershaft have
no cilia (2). A more recent statement about neossoptiles is that
the feathers forming the familiar coat of newly hatched ducklings
and chicks are not protoptiles but mesoptiles, and hence corre-
spond to the feathers forming the long fur-like second coat of
Penguins (3).

T may at once state that in Mallard ducklings the outer
nestling coat consists of typical protoptiles characterized by
(1) a well-developed calamus which may contain over twenty
“eones,” (2) a well-developed rhachis, (3) well-developed and in
some cases hook-like cilia, (4) short stiff, as well as long slender
barbs, and (5) by a well-developed aftershaft, the barbules of
which bear cilia.

Tn ducklings the protoptiles vary in size—some are over 30 mm.
in length and resemble plumose (metaptile) feathers, others are
under 10 mm. in length and resemble preplumule. But the
small as well as the large protoptiles consist of a calamus, a shaft,
and an aftershaft. In addition to assisting the preplumule, and
later the plumule, in diminishing the flow of heat from the skin,
the protoptiles, as a rule, prevent water reaching the under coat

NESTLING FEATHERS OF THE MALLARD. 611

of true down; by the pigment they contain they help to make
the duckling inconspicuous.

(1) Zhe Wing-Quall Protoptiles.

Though Ducks have been living under domestication for well-
nigh two thousand years, their nestling feathers have apparently
never been systematically examined. ‘That no serious attempt
has been made to study the nestling feathers of ducks and other
Anseres, and of fowls and other Galli, is made evident by a
perusal of Mr. Pycraft’s important monograph on the Emperor
and Adélie Penguins. In an interesting chapter on the com-
position and sequences of the neossoptiles, after pointing out that,
“the penguin must certainly be regarded as having preserved
what must be looked upon as a somewhat, perhaps very, ancient
succession of plumages,” Mr. Pycraft states that “in the light of
my recent discovery” it is clear that the ‘‘down” feathers of the
Galli and Anseres ‘do not, 1s I imagined, represent a primitive
type of down feather homologous with the woolly, and so pre-
sumably degenerate down of say the Alcide, but answer to
mesoptyles. The protoptyle or first generation of feathers would
seem to be wanting in these birds, but I had the good fortune to
discover small tufts of down adhering to the tips of the meso-
ptyles of a young Chloéphaga rubidiceps. Thus we may assume
that this first generation, since it has not yet been traced, has
been lost in all the Galli, and probably all the Anseres save
perhaps this species and one or two allied genera” (3).

As Pl. I. figs. 2 & 3 show, I have succeeded in tracing two
generations of nestling feathers in the Indian Runner Duck and
in the common Domestic Goose, and thereby have proved that
the coat worn by newly-hatched Ducks and Geese consists of
protoptiles corresponding to the first or protoptile nestling coat
of Penguins. In the case of the King Penguin the first or
protoptile coat is apparently in the act of disappearing; but in
all the Anseres and Galli I have examined it is the second coat
that has been lost or is in the act of disappearing. Though in
the wings of some Ducks and Geese well-developed mesoptiles
still make their appearance, the tail-quill mesoptiles are either
vestigial or have been completely suppressed. That the pro-
toptile is in the act of disappearing in the tail of the Penguin is
suggested by Pl. I. fig. 4, while Pl. II. fig. 5 clearly indicates
that as the mesoptile was suppressed in the tail of the Mallard
the protoptile was enlarged.

With the help of a prematurely developed ‘nestling tail” the
Mallard duckling begins to dive at the end of the second week,
but, owing to the development of the wings being retarded, flying
only becomes possible at the end of the eighth week. That the
development of the wing is from the outset retarded is suggested
by Pl. II. fig. 6, a ten days’ duck embryo. In this embryo the
papille which develop into the tail-quill protoptiles are large,

612 PROF, J. COSSAR EWART ON THE

but the wing-quill papille are only beginning to make their
appearance*. :

In a five weeks’ duckling some of the tail-quills measure
50 mm., but the wing-quill rudiments at the end of the fifth
week only measure 4 mm. The wing of a twenty-five days’
Mallard is represented in Pl. II. fig. 7, naturai size, and the
protoptile of a secondary wing-quill of a thirty-eight days’
Mallard in Pl. II. fig. 8. Though the wing-papille appear late
they soon develop into filaments—filaments of a thirteen days’
Mallard embryo are represented in Pl. III. fig. 9. With the
exception of the calamus the development of the protoptile is, as

Text-figure 1.

oo a | ———
p PPE SS

OLLIE ET Fo
Ss TIT a we == Pea : :
SASS SSS

=. SEN =
—— AN i N

zt Sh

—_

Part of a barb from the shaft of a protoptile of a 16 days’ duckling showing
barbules spirally twisted at their origin. X 26 dia.

Text-figure 2.

hot SS oe

Distal part of a barbule of a protoptile of a 23 days’ duckling
showing cilia. X 250 dia.

a rule, completed before the end of the twenty-eighth day of
incubation, so that the vascular pulp—which made the develop-
ment of the nestling feather possible—may be converted into
bloodless pith before hatching. Hven when the duckling escapes
from the shell the nestling coat consists of bristle-like filaments.
But as the duckling dries, the thin friable sheaths which pro-
tected the protoptiles during development disintegrate, with the
result that the protoptiles expand and provide the duckling
with a wonderfully complete outer nestling coat. A wing-quill

* It is conceivable that there is some relation between the slow development of
the wing and the persistence of wing-quill mesoptiles.

™ =

NESTLING FEATHERS OF THE MALLARD. 613

protoptile of a twenty-six days’ duck embryo artificially removed
from a filament (like the long filaments in Pl, III. fig. 9) is
represented in Pl. III. fig. 10.

The wing-quill protoptiles are complete feathers: 7. ¢., each
consists of a calamus, a shaft, and an aftershaft. The calamus,
though only 3 or 4mm. in length, is well formed, contains
several ‘ cones,” and remains intact until the protoptile is shed—
in Penguins the part of the epidermic tube which represents a
“‘calamus” usually soon splits, with the result that the individual
protoptile barbs are directly continuous with mesoptile barbs
(text-fig. 14). Though the growth of the shaft and aftershaft is
arrested before hatching, the calamus may continue to grow alter
hatching.

The shaft is made up of a rhachis, continuous with the outer
segment of the calamus, and usually of seven or eight pairs of
barbs, which vary in length and in the number of barbules they
possess. The two terminal barbs (which by uniting formed the
distal part of the rhachis) end in long slender processes destitute
of barbules. The aftershaft (PI. IT. fig. 11), continuous with the
inner segment of the calamus, usually consists of eight barbs
bearing barbules. Sometimes the four mesial barbs of the after-
shafts unite to form a short rhachis. The barbules of the shaft
are spirally twisted at their origin (text-fig. 1), but the aftershaft-
barbules only curve slightly on leaving the barb. It has frequently
been stated that aftershaft-barbules of true as well as nestling
feathers have no cilia, but cilia are invariably present on the
barbules of the protoptile aftershaft of Mallard ducklings. Part
of a barbule with cilia is given in text-fig. 2.

The wing-quill protoptiles may all be present at the end of the
sixth week (PI. III. fig. 12), but some of them have usually been
shed before the end of the seventh week. Though Mallards
begin to fly during the ninth week, the development of some of
the wing-quills is only completed during the tenth week.

(2) The Wing-Quill Mesoptiles.

Up to 1906, as already mentioned, it had not occurred to
ornithologists that true feathers (penne) might be preceded by
two generations of nestling feathers (prepenne), and mesoptiles
have apparently not yet been recorded in connection with wing-

uills.
‘ Early in the century two National Antarctic Expeditions were
fortunate enough to collect material which made possible a fairly
exhaustive study of the development and history of the nestling
feathers of Penguins. The penguin material brought home by
the English National Antarctic Expedition was reported on by
Mr. Pyeraft ; that collected by the Scottish National Antarctic
Expedition by Dr. Eagle Clarke. In a paper published in 1906
in ‘The Ibis,’ Dr. Clarke announced that he had discovered two
coats of nestling feathers in the Ringed and Gentoo Penguins,

614 PROF. J. COSSAR EWART ON THE

Text-figures 3 & 4.

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Sor

af SSS MARA M4 SCO
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BY TATAY YAN Z
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UO MINAIN og
MAA (NN
RN NY
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BOUT
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ie:
af RNA
a Sete to ANN
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=
Ae
2
1
1
1
1
1
1
'
7
1
'
i
os)
(mr

ees

eB.
ies

Diagrams indicating the relation of the mesoptile to the protoptile and to the first
true feather of a Ringed Penguin :—pro., the protoptile, consisting of four
outer barbs representing a shaft and of four inner barbs representing an after-
shaft. sh.mes., shaft of mesoptile continuous distally with protoptile and
proximally with the shaft of the true feather. afmes., mesoptile aftershaft also
continuous with the protoptile and the true feather. cé., connection with pro-
-toptile. c¢., connection with shaft of true feather. ea., connection with after-
shaft of true feather. cal., calamus.

NESTLING FEATHERS OF THE MALLARD. 615:

and in the following year an important paper by Mr. Pycraft on
the Emperor and Adélie Penguins was published which deals
with the composition and sequences as well as with the structure
of nestling feathers.

Dr. @lanke refrained from stating whether the silky-white
protoptile coat, or the grey fur-like mesoptile coat, of the Ringed
Penguin, corr esponds to the nestling coat worn by newly -hatched
chicks and ducklings. But Mr. Pycraft, in discussing the
sequences of nestling-feathers, states that there ave good reasons
for assuming that in most of our common birds the first or
protoptile coat has been lost, and adds that the feathers forming
the adult coat of the Emu might possibly consist of mesoptiles.

Whether ornithologists have, as a rule, adopted Mr. Pycraft’s
views it is impossible to say, but ib may be mentioned that
Mr. Ingram, in the ‘Ibis’ paper already referred to, points out
that a study of the nestling plumage of the Raptores suggests.
that in some cases, at any abe, it is the second and not the first
generation of nestling ‘‘ down ” that has been suppressed (1).

When, some years ago, I was led to study the feathers of
Penguins, I assumed that ‘birds with only one nestling coat had
lost the first or protoptile coat; but the examination of a
number of nestlings soon made it evident that when in the Galli
and Anseres there is only one generation of nestling feathers,
it is owing, not to the absence of the protoptiles, but to the
suppression of the mesoptiles.

In the account of the nestling feathers of the Emperor
Penguin, Pycraft states that the mesoptiles are umbelliform, and
that in the Adélie Penguin the mesoptile is in part attached to
the main shaft, but mainly to the aftershaft. I have not had an
opportunity of studying the nestling feathers of either the
Emperor or Adélie Penguins, but from material placed at my
disposal by Dr. Eagle Clarke I have worked out the structure
of the mesoptiles of the Ringed Penguin. In this species the
mesoptile is extremely complex: it consists (1) of an outer small
series of barbs, which extends between the protoptile and the tip
of the true feather, and hence occupies the position of, and
doubtless represents, a shaft; and (2) of an inner series of barbs
(text-fig. 3), arranged to form two or three bundles, which as
obviously represent an aftershaft. ‘The chief connections of the
mesoptile in the Ringed Penguin are diagrammatically repre-
sented in text-fig. 4, which indicates that the mesoptile consists
of a simple shaft extending between the protoptile and the tip of
the true feather, and a complex aftershaft connected with the
shaft as well as with the aftershaft of the true feather.

In a Mallard duckling at the end of the fifth week the
mesoptile looks as if it would develop into a simple umbel not
unlike the umbelliform protoptiles of Penguins. But this
appearance is due to the fact that nestling feathers, like true
feathers, are developed, not out of a scale- like plate, but out of an
epidermic tube surrounding a highty-vascular dermic pulp and

Text-figure 5.

+
te}.

Ady)
Yy

yy
ps

Diagram indicating the structure and relationships of the mesoptile with the pro-
teptile and with the true feather in the Chinese Goose. The mesoptile shaft
(sh.mes.), which lies between the protoptile shaft (sk.pro.) and the tip of a true
feather, consists of numerous barbs and of a band representing a rhachis. Con-
tinuous with the rhachis is a perforated band from which slender cords (c.)
proceed to the barbs forming the tip of the true feather (¢eZ.). The mesoptile
aftershaft is also continuous with a perforated band which ends in a vestige of
an aftershaft of the true feather (aft.). From the perforated band slender
cords (¢1) proceed to barbs near the tip of the true feather. 1¢ will be ob-
served that the aftershaft of the protoptile consists of twelve well-developed
barbs armed with numerous barbules.

NESTLING FEATHERS OF THE MALLARD. 617

protected by a thin epidermic sheath. When the development
is carried further, it becomes evident that in the Mallard the
mesoptile consists of two sets of fibres, an outer set representing
a shaft, and an inner, an aftershaft, each provided with a more
or less distinct rhachis. Though the mesoptiles in Ducks may
reach a considerable size, it is easier to make out their develop-
ment and structure in Geese. The wing-quill mesoptiles, though
well-developed in the Mallard, are not present in the Emden
goose but as Pl. I. fig. 3 shows, the major wing-covert mesoptiles
may reach a considerable size in Geese.

To start with, the mesoptile of a Chinese gosling looks as if it
possessed a calamus (Pl. IV. fig. 13); the mesoptile barbs seem
to proceed from a tube lying immediately in front of the tip of
the true feather. That this tubular part represents a calamus
is supported by the fact that it contains several ‘“ cones,” but
ugainst the view that it is a true calamus is the fact that it is in
the act of splitting longitudinally into numerous slender cords.
In mesoptiles from older goslings part of the tubular portion is
represented by two perforated bands, from which proceed
numerous slender cords towards the wing-covert. Sometimes
the mesoptile in goslings splits into two portions (Pl. IV. fig. 14,
& text-fig. 5), an outer (the shaft) extending between the shaft
of the protoptile and the tip of the wing-quill, and an inner (the
aftershaft) extending between the protoptile aftershaft and the
vestigial aftershaft of the wing-quill.

It may be mentioned that in the Anseres the mesoptiles vary
greatly in structure—the proximal part of the aftershaft may be
perforated and assume the form of a section of a relatively
large tube (PI. IV, fig. 14), or, like the rhachis of the shaft, be
represented by a simple narrow band. Though in the Mallard
and Chinese goslings the aftershaft, as well as the shaft of the
mesoptile, has usually a distinct rhachis, in Emden goslings the
whole of the mesoptile, as Pl. 1. fig. 3 suggests, usually consists
of simple barbs, one or two of which are continuous with the
rhachis of the wing-covert shaft, while two or more end in a
vestigial aftershaft. The barbules of the mesoptile, like those of
tlie protoptile aftershaft, are narrow and provided with well-
developed cilia. In the Ringed Penguin the mesoptiles closely
agree in structure, but in the Ducks and Geese examined no two
mesoptiles were alike in structure. The mesoptile coat is probably
as useful now to Penguin chicks hatched within or near the
Antarctic Circle as it was when originally acquired, during perhaps
a cold phase of a glacial epoch. In the Anseres, though the
protoptile coat still plays an important part, the mesoptile coat
probably no longer counts in the struggle for existence. If this
is the case there is no difficulty in accounting for its being well-
developed in some parts but entirely absent in others, and for
the marked variation amongst the mesoptiles which still persist.

The Mallard has wing-covert, as well as wing-quill, mesoptiles.
The mesoptiles which precede the wing-coverts in the Mallard

618 _, PROF. J. COSSAR EWARY ON THE

are especially interesting, because they indicate how the suppres-
sion of the mesoptiles was gradually accomplished. In the case
of the disappearing protoptiles of the King Penguin (PI. I.
fie. 4) the size is gradually reduced until they are smaller than
the minute bunch of barbs which precede the filoplumes of
ducklings. In the case of the vanishing mesoptiles two sets.
of factors are evidently at work ; one set arrests the development
of the mesoptile barbs, another set accelerates the development of
the barbs of the true feather, with the result that in course
of time the tip of the true feather is found projecting into the
base of the protoptile calamus.

Though in most cases the mesoptiles are gradually reduced in
size, in some eases the length is maintained and either a few
barbs with barbules are developed, or the simple epidermic tube
splits into two or more bands, which for a time connect the
protoptile to the tip of the true feather.

It may be mentioned that though all the wing-quill protoptiles
and mesoptiles may be present in the Mallard duckling at the
end of the sixth week (PI. III. fig. 12), some of the mesoptiles.
are usually shed before the end of the seventh week, and all are
usually lost before the middle of the eighth week. But though
the wing-quills lose all their nestling feathers during the eighth
week, some of the feathers of the humeral track may retain
protoptiles and mesoptiles to the end of the eleventh week. The
existence of the wing-quill mesoptiles in the Mallard proves
conclusively that the nestling coat worn by newly-hatched Ducks
and Geese corresponds, not as Pyeraft suggested to the second
generation of prepenne in Penguins, but to the first or protoptile
generation.

(3) The Tail-Quill Protoptiles.

The tail varies greatly in young aquatic birds. In Penguins
the tail protoptiles have almost disappeared and the mesoptiles
are only represented by a few simple barbs (Pl. I. fig. 4). In the
Mallard the mesoptiles of sixteen of the eighteen tail-quills have
completely disappeared, but the protoptiles of all the eighteen
quills are larger and more complex than in the plumose (meta-
ptile) feather represented in P]. I. fig. 1. In Penguins steps are
soon taken to develop the tail-quills with a view to their forming
with the hind limbs a tripod useful in maintaining the erect
attitude. In the Mallard the protoptiles form a ‘nestling tail”
which plays an important part during the earlier portion of the
period that Mallard ducklings behave like diving ducks™.

In a ten days’ duck embryo the tail-quill papille ? are easily

* The adult Mallard, Mr. G. J. Millais informs me, “hardly ever dives except
when in play during the love chase, or to escape when wounded.”

+ There ave twenty tail-quill papilla, but only eighteen of them develop into
feathers large enough to rank as tail-quills. A paper “On the development of the
Feathers of the Duck during the Incubation Period” was recently communicated
to the Royal Society of Edinburgh by Augusta Lamont, B.Sc.

NESTLING FEATHERS OF THE. MALLARD. 619

identified (Pl. IL. fig. 6), and by the twenty-fifth day of incubation
they have grown into long filaments out of which escape imme-
diately after hatching highly specialized protoptiles. During
the first week the nestling tail consists only of protoptiles (PI.IV.
fig. 15), but during the second week the protoptiles are pushed
from the skin by the tail-quills, or by vestigial mesoptiles. At

Text-figure 6.

, HA
SS
if vy ve

7 his ast mK
Dy

ends in a calamus containing “cones,” that this calamus is directly continuous
with the expanded distal portion of the tail-quill rhachis, and that the connection
of the protoptile with the teleoptile is strengthened by barbs at the tip of the
tail-quill ending in the proxinilal portion of the protoptile calamus.

the end of the fourth week the tail-quills may be nearly an inch
in length, and they are still longer at the middle of the fifth
week (Pl. [V. fig. 16). During the latter part of the fifth week
the protoptiles begin to break off, and with the exception of the
ninth from the middle line at each side, they are usually all shed
by the middle of the sixth week.

620 PROF. J. COSSAR EWART ON THE

The calamus of the protoptiles is strong and rigid and firmly
fixed to the expanded tip of the tail-quill rhachis. By way of
strengthening the connection between the protoptile and the tail-
quill, four or more pairs of the terminal barbs of the tail-quill
are attached to the protoptile calamus (text-fig. 6). At the end
of the second week the “* nestling ” tail and the other structures.
concerned are sufficiently developed to admit of young Mallards.

Text-figure 7.

The proximal portion of the calamus of a tail-quill protoptile containing “ cones.’”
As the calamus approaches the tip of the tail-quill it splits into an outer
‘portion continuous with the expanded rhachis of the quill and a more slender
inner portion continuous with a vestigial tail-quill aftershaft. c., protoptile:
calamus with “‘cones.” sf., tail-quill shaft. af., vestige of tail-quill after-.
shaft.

obtaining the greater part of their food by diving*. Notwith-
standing the loss of the protoptiles during the sixth week, young
Mallards usually continue to behave like diving ducks up to the
end of the eighth week. From what has been said it is evident
that the “ nestling tail” of the Mallard duckling is not from first.
to last entirely composed of nestling feathers. It is a true
nestling tail up to the middle of the second week, but from the

* Two of nine Mallard ducklings I had under observation began to dive on the-
thirteenth day: they were all diving on the fourteenth day.

NESTLING FEATHERS OF THE MALLARD. 621

beginning of the third to the end of the fifth week the true tail-
quills take an ever increasing part in forming the “ nestling tail.”
Hence nestling feathers only function during three of the six
weeks that Mallard ducklings feed and disport themselves like-
diving ducks. When the protoptiles are shed, the tail-quills,
owing to the rhachis being short and incomplete, look unfinished.
Perhaps for this reason the first set of tail-quills is ere long
superseded by larger and more perfect quills in which the rhachis,
as in adult ducks, ends in a fine point. Usually the first set of
tail-quills begins to drop out during the fourteenth week, and a
new tail consisting of feathers of the adult type is eventually
developed.

The Structure of the Tail Protoptiles—The calamus of the tail--
quill, like that of the wing-quill, protoptile, continues to grow
after hatching. In the case of the protoptiles at each side of the:
middle line the calamus only reaches a length of 4 or 5 mm. and
may have only three or four “cones”; but the calamus of the:
outer tail protoptiles may eventually measure 15 mm. and contain
over twenty ‘‘cones.” I expected the entire protoptile calamus:
to be directly continuous with the expanded tip of the tail-quill
rhachis, but I found that, as text-fig. 7 shows, the calamus splits.
into an outer segment continuous with the tail-quill rhachis and
an inner segment continuous with the rhachis of a vestigial
aftershaft, the presence of this unexpected vestige of an after-
shaft indicates that even the highly specialized tail quills.
originally consisted of two shafts, and hence were constructed on
the same plan as the double feathers of the Emu.

The shaft of the tail-quill protoptiles of a Mallard duckling
as a rule consists of fourteen or more pairs of barbs, each pro-
vided with barbules. As already mentioned, the majority of the-
barbs of the wing-quill protoptiles are long and pointed and have
only a limited number of slender barbules (PI. II. fig. 8), but some
of the barbs of the tail-quill protoptiles are ribbon-shaped
(text-fig. 9), and they all have barbules along their whole length..
The majority of the barbules have simple cilia along both
margins; but the six or seven pairs of ribbon-shaped barbules at
the tip of the protoptile have large curved cilia along one margin,
which differ but little from the hooklets on the barbules forming
the tip of the true tail-quills. The difference between the
specialized barbules with hook-like cilia at the tip of the pro-
toptile and the slender proximal barbules with small cilia will be-
evident if text-fig. 8 is compared with text-fig.9. When the
broad barbules with hook-like cilia on one edge cross each other
the approximation to a true teleoptile is especially marked..
What Mallard ducklings gain by having the tips of the tail-quill
protoptiles highly specialized is not very obvious. The after-
shaft of the Mallard’s tail quill consists of eight long barbs
(Plate III. fig. 11) bearing slender almost straight barbules, each
with two rows of small cilia.

It may be mentioned that in the case of the goslings of Chinese-

622 PROF. J. COSSAR EWART ON THE>

and Emden geese the tail-quill protoptiles—though possessing a
long, strong calamus firmly connected with the rhachis of the tail-
quill, a well-developed shaft, and an aftershaft made up of twelve
barbs—are not provided with broad distal barbules armed with
hook-like cilia. In goslings, as in ducklings, the inner segment
of the calamus of the tail-quill protoptile is continuous with the
rhachis of a vestigial tail-quill aftershaft.

Text-figure 8.

Two of the proximal barbules of a tail-quill protoptile; they are narrow
and armed with small cilia. x 64.

Text-figure 9,

!
'
'
'
1
'
1
'

‘Three of the broad distal tail-quill barbules armed along one edge with tooth-like
cilia (hamuli) of true feathers. X 64. The difference between the simple
‘proximal and the specialized distal barbules is also indicated in text-fig. 6

(4) The Tail- Quall ehante

In sixteen of the tail-quill protoptiles of the Mallard the
mesoptiles are completely suppressed, but in the remaining two—
the ninth from the middle line at each side—the mesoptile is
represented by a simple tube from 10 to 12 mm. in length ex-
tending between the protoptile calamus and the tip of the tail-
quill. This tube, though complete to start with, sooner or
later splits into two ov more bands, the outer of which is con-
tinuous with the rhachis of the tail- quill, while the inner ends in
the rhachis of a vestigial tail-quill aftershaft. As these bands
have no barbules they evidently do not represent barbs. Though
there are only vestiges of two tail-quill mesoptiles in the Mallard,
there are sometimes vestiges of all the mesoptiles in cross- bred

NESTLING FEATHERS OF THE MALLARD, 628

domestic ducks. In cross-bred, as in Mallard ducklings, the
feathers at each side of the true tail-quills approach in structure
the wing-quill mesoptiles. In Hider ducklings the wing-quill, as
well as the tail-quill, mesoptiles are either absent or represented
by minute vestiges, hence the Hider supports the view that in
the Anseres the mesoptiles no longer count in the struggle for
existence.
(5) The Prepenne of the Trunk.

About the prepennz which precede the feathers of the head,
neck, trunk, and hind limbs, very little need be said. In many
cases the protoptiles closely resemble, and may be as large as,
the wing-quill protoptiles, but they are sometimes so small and
poorly developed that it is difficult to distinguish them from pre-
plumule. ‘The protoptile from the dorsal track of a thirty-eight
days’ Mallard duckling represented in PI. I. fig. 1 is only half the
size of the wing-quill protoptile in Pl. IT. fig. 8; but some of the
protoptiles of the wing-coverts are smaller than the majority of
the preplumule. Usually the protoptiles of the trunk are shed
about the same time as the wing-quill protoptiles, but in a small
chequered area at each side of the body near the tail well-
developed protoptiles are still present at the end of the eleventh
week. Though mesoptiles precede some of the trunk penne,
they are never as well developed as the wing-quill mesoptiles.

The examination of the mesoptiles of ducks and penguins lends
support to the view that the mesoptile coat was specially acquired
to meet a marked change of climate—a change perhaps from cold
dry to cold damp conditions—and that in most cases as the
climate improved the fur-like second coat was more or less com-
pletely suppressed, and a new coat, made up of feathers con-
structed on the protoptile rather than the mesoptile plan, was
subsequently evolved. It is conceivable that some of our modern
birds, e. g., the Emu and Cassowary, are descended from ancestors
which never acquired a mesoptile plumage.

2. The Preplumule and Plumule.
(1) Zhe Preplumule.

In many birds the nestling coat sooner or later consists of pre-
plumule as well as prepenne. In the Gannet the preplumule
appear before the prepenne, and grow so rapidly that even when
the prepenne are well developed they are difficult to find; but
in the Mallard in most regions the preplumule are hidden by a
relatively dense coat of protoptiles. Though the preplumule
papille in the Mallard appear three or four days later than the
protoptile papille, the majority of the prepiumule are ready to
escape from their filaments as soon as the duckling is hatched.

The relative size of the protoptile and preplumule filaments of
the Mallard is indicated in Pl. IIT. fig. 9 and in text-fig. 10. The
difference in size of the protoptiles, preplumule, and prefiloplume

Proc. Zoou. Soc.—1$21, No. XLII. 42

624 PROF. J. COSSAR EWART ON THE

is made evident by Pl. V. fig. 17, which shows a protoptile, three
plumule, each with a preplumula on its tip, and, on the left
of the protoptile, a prefilopluma on the tip of a developing
filoplume.

The preplumula consists of a short conical calamus, a shaft
often consisting of a short rhachis and four barbs, and an after-
shaft also as a rule consisting of four barbs. It is sometimes
difficult to distinguish the small protoptiles which precede small
true feathers from preplumule, but when in ducklings the

Text-figure 10.

Filaments from a Mallard embryo.
P, proximal part of a protoptile filament. P.1, a preplumula filament.
P.2, a prefiloplume filament. S., skin.

barbules are characterized by nodular swellings and very few
cilia it may be safely assumed that the nestling feather under
examination is a preplumula.

(2) The Plumule.

At the beginning of the Jurassic Age the coat of birds may
have consisted only of protoptiles and preplumule; but in
many recent birds the plumule form a very important part of
the nestling plumage. From Pl. V. fig. 17 it may be inferred
that during the fifth week the plumule form an inner coat as
effective in preventing a loss of heat as the woolly under coat of
wild sheep.

,
;
4

SS ee ee

eee

NESTLING FEATHERS OF THE MALLARD, 625

Like plumose cr metaptile feathers, the plumule vary in size,
colour, and structure, but they, asa rule, consist of a calamus,
a shaft, and an aftershaft. ‘lhe calamus, especially well developed
in the Hider Duck, is barvel-shaped and contains relatively large
cones. As it leaves the skin it divides into two portions, an
outer which forms the short rhachis of the shaft, and an inner
which forms the still shorter rhachis of the aftershaft( Pl. V. fig. 18).
The shaft consists of a dozen or more pairs of very long slender
barbs from which spring numerous slender barbules, the proximal
portions of which are characterized by pyramidal swellings,
while the distal portions like the preplumule have nodular
swellings. The aftershaft (Pl. V. fig. 18)is always well developed,
and like the shaft is largely made up of long barbules having the
characteristic pyramidal swellings. When a plumula is shed the
tip of the sheath of the growing new plumula is often found
attached to the calamus. Evidence of this we have in Pl. VY.
fig. 19.

(3) Prefiloplume and Filoplume.

Nitzsch, in his account of filoplume, says they are strikingly
distinguished from the other three kinds of feathers (contour,
down, and semiplume feathers) by their peculiar position, their
extraordinary slenderness, and by the entire deficiency or very
small size of their vane; but he states subsequently that he
believes the narrow white downy feathers of the Cormorant must
be regarded as filoplumes, even though they are furnished with
perfect vanes. Nitzsch further mentions that filoplumes are
“associated with the contour feathers, one or even two filoplumes
standing quite close to every contour feather of the head, neck,
and trank, apparently issuing almost out of the same pouch of
the skin.” Fuiloplumes are usually regarded as degenerate
feathers because during development they have barbs arranged
after the fashion of a down (plumula) feather.

In the Mallard during development there are usually two or
more minute filaments at each side of the filaments containing
the wing-quill protoptiles. These minute filaments contain
prefiloplume. In text-fig. 10 the two short filaments (P. 2)
lying in contact with the large protoptile filament are prefilo-
plume filaments. ;

In the newly-hatched duckling the minute prefiloplume and
the large wing-quill protoptiles escape simultaneously from their
sheaths—being barely visible to the naked eye, the prefiloplume
have apparently hitherto escaped the notice of ornithologists.
In course of time, as the protoptiles are pushed from the skin by
mesoptiles, the prefiloplume are pushed from the skin by
filoplume. The wing-quill protoptile of a thirty-eight days’
duckling represented in PI. II. fig. 8 is attached to a developing
mesoptile. At the base of the mesoptile are three young filo-
plumes, two of which support prefiloplumes—a young filoplume

with a prefiloplume attached to its tip is also seen in. Pl. V. fig. 17.
42*

626 PROF. J. COSSAR EWANT ON THE

Notwithstanding their small size, the prefiloplume have the
structure of prepenne. They are hence not dwarfed pre-
plumule, as their name suggests, but dwarfed protoptiles. It is
important to note that, though several developing filoplume may
be intimately related to a protoptile, they are not imbedded in
its sheath; each projects from a separate pit in the skin. At
the end of the seventh week, when the filoplumes have reached a
length of about 20 mm., they are still found in contact with the
developing true feather. A filoplume from a forty-eight days’
duckling, still carrying the minute prefiloplume on its tip, is
represented in-Pl, V. fig. 20. At this stage the filoplume still con-
sists of a calamus, a shaft and an aftershaft; but as the duckling
increases in size the filoplume in connection with the wing-quills
degenerate. ach loses its prefiloplume, its aftershaft, and most
of the barbs of its shaft, with the result that, as a rule, only the
rhachis of the shaft and two or three vestigial terminal barbs are
left.

That the filoplumes are degenerate penne is supported by a
study of the foot-feathers of the Barn Owl (Striw flammea). In
a newly-hatched Barn Owl one large and two small filaments are
seen projecting from under the foot-scales. Soon a well de-
veloped protoptile escapes from the large mesial filament and
a minute protoptile from each of the two small filaments. In
course of time a simple plumose feather (metaptile), with a large
aftershaft, but with a rhachis which suggests a filoplume, succeeds
the large protoptile, and typical hair-like filoplumes succeed the
small protoptiles.

Tf further proof were needed in support of the view that
filoplumes are degenerate penne, it is obtained by the examina-
tion of the white neck feathers which form part of the nuptial
plumage of Cormorants, z. e., the feathers Nitzsch assumed were
tiloplumes. Instead of regarding the narrow neck feathers of
Cormorants as filoplumes, it would be better to regard them as
true feathers (metaptiles) which have undergone degeneration.

It may be added that, notwithstanding the small size of the
prefiloplume, a microscopic examination clearly proves that they
are more intimately related to protoptiles than to preplumule.
Seeing that the filoplumes are degenerate penne, it might be
better if the name filopenna were substituted for filopluma.

Ti. THe Compostrion oF FEATHERS.

The feathers forming the coat of the adult Emu consist of a
calamus and two blades, an outer, the shaft, and an inner,
generally known as the aftershaft; the wing- and tail-quills of
ducks consist of a calamus and a well-developed shaft, but there
is only a minute vestige of an aftershaft. Hitherto it has been
taken for granted that a feather with only a vestige of an after-
shaft is more primitive than a feather with an aftershaft as long
and as complex as the shaft, that, in fact, the aftershaft is not a

a ee eed

NESYLING FEATHERS OF THE MALLARD. 627

primitive but a secondarily acquired feature. The chief reasons
given for regarding the aftershaft as an accessory and secondarily
acquired structure are: (1) that the aftershaft is developed from
a forward elongation of the calamus, and (2) that the tip of the
aftershaft of the new feather is never attached to the calamus of
the feather about to be shed—that, for example, in the Emu the
tip of the aftershaft of the feather of the second generation is
from first to last free.

If I succeed in showing that the aftershaft, like the shaft,
grows from apex to base and is completed before the calamus 1s
formed and that, during development, the tip of the aftershaft
is connected with the calamus of the feather about to be shed
is, in other words, for a time attached to the old feather—it
may be safely assumed that the aftershaft is a primitive feature,
and that a complete true feather (penna) like a down feather
(plumula) consists of two shafts or blades.

Owen evidently assumed that a true feather was made up of a
calamus and a single shaft, for he states that “‘ besides the parts
which constitute the perfect feather there is an appendage
attached to the upper umbilicus called the accessory plume” (4).
According to Nitzsch, the aftershaft ‘‘ originates from the under-
side of the umbiliciform pit” (5). Gadow, in discussing the
aftershaft, states that when present it is developed as ‘“‘a ventral
elongation of the calamus,” and points out that if we consider the
various types of nestling feathers ‘‘ with reference to the presence
or absence of an aftershaft in the Teleoptiles, we are led to
conclude that this appendage and eonsequently also the double
feathers of certain Ratite are secondarily acquired not primitive
features” (6).

That the aftershaft is a primary feature was suggested in 1903
by Degen in a paper on Ecdysis. In the chapter on “ Adven-
titious Plumage,” Degen writes: ‘‘I may mention here that
owing to the yet ‘more primitive conditions of the feathers of the
trunk in some Ratite, the extreme tips of the aftershafts, which
in the Cassowary and the Emu attain a length equal to that of
their main shafts, jointly support the new-growth feather with
the latter” (7). But Degen, following Gadow, seems to adopt
the view that the calamus is concerned with the formation of the
aftershaft, for he assumes that while the one half of the calamus
“deposits substantia rhachidis evterna for the main shaft, the
other half deposits” the same substance required in the building
up of the other shaft! Obviously the best way to throw light on
the status of the aftershaft is to study its development.

1. The Development of the Aftershaft of Plumule.—tIn the
Mallard a preplumula consists of a small conical calamus, a shaft
usually made up of four barbs, two of which coalesce to form a
short rhachis, and of an aftershaft also made up of four barbs.
Soon after hatching the preplumule are pushed from the skin by
plumule. Some of the plumule at the end of the fifth week,
though still bearing preplumule on their tips, are 15mm. in

628 PROF. J. COSSAR EWART ON ‘tHE

length and consist of two distinct bundles of barbs—a large outer
bundle in a line with the shaft of the preplumula and a smaller
bundle in a line with the aftershaft.

In the plumula from a five weeks’ duckling represented in
IBIS AY Ihe fig. 21, the outer bundle already consists of a rhachis and ~
eight pairs of barbs, but the inner or aftershaft bundle only
consists of five pairs of barbs. By the end of the sixth week the
development of both the shaft and aftershaft is complete, and
the unsplit portion of the epidermic tube is giving rise to a
calamus. In the completed plumula (Pl. V. fig. 18) the shaft
consists of a rhachis and over twenty pairs of barbs; and the
aftershaft of a rhachis with about a dozen pairs of barbs.

From Pl. VI. fig. 21 it is evident that the aftershaft of a
plumula in its development and structure agrees with the shaft ;
as it is developed before the calamus it cannot be regarded
as a secondarily acquired appendage produced from a forward
elongation of the calamus.

2. The Development of the Aftershaft of Emu Feathers.—It
has hitherto been often assumed that in the Emu the proto-
ptile has ‘no distinct aftershaft.” That there is anaftershaft in
the first, as wellas in the second and third Emu feathers, is indi-
cated by the diagram, text-fig. 11. In the young Emus I have
examined the protoptile aftershaft has, with rare exceptions,
consisted of seven or more barbs longer than, or at least as long
as, the barbs of the shaft. In the Emu protoptile given in
Pl. VI. fig. 22, the aftershaft has a short rhachis. It is especially
noteworthy that in the Emu the rhachis of the protoptile shaft,
instead of ending in a hollow calamus containing ‘‘ cones,” is
connected with the relatively thick rhachis of the shaft of the
feather of the second generation by a solid band (Pl. VI. fig. 23).

In a like manner the rhachis of the protoptile aftershaft is
connected with the aftershaft rhachis of the second feather.
When, as sometimes happens, the connecting bands occupying
the position of a calamus separate from each other (Pl. VI. fig. 23),
it becomes obvious that the tip of the aftershaft of the second
feather is not free as generally assumed, but from the outset
continuous with the protoptile aftershaft.

Though as in plumule the rhachis of the aftershaft is formed
by the fusion of two barbs, the distal portion of the rhachis of the
shaft is simply.a continuation of the band proceeding backwards
from the protoptile shaft. That the aftershaft of the feathers of
the second generation is not ‘‘ developed as a ventral elongation
of the calamus,” but develops from apex to base like the shaft,
is made clear by Pls. VI. and VII. figs. 23, 24, & 25, which show
the second feather at various stages of growth.

As the destined length of the second-generation feather is
reached the rhachis of the shaft expands, gives off six or more
pairs of relatively long barbs, and thereafter becomes continuous
with the rhachis of the shaft of the feather of the third generation
(Pl. VII. fig. 26). Im a like manner the proximal end of the

NESTLING FEATHERS OF THE MALLARD, 629

rhachis of the aftershaft, after giving off several pairs of barbs
but without appreciably increasing in width, becomes continuous
with the rhachis of the aftershaft of the feather of the third
generation (P]. VII. fig. 26). When (as was the case in the
specimen represented in Pl. VII. fig. 27) the two connecting bands
naturally separate from each other, it is impossible to escape

Text-figure 11.

Eee

In this diagram the structure and relations of the feathers forming the first three
generations in the Emu are indicated. sh.1, shaft, and af. 1, aftershaft of pro-
toptile. sh.2, shaft, and af.2, aftershaft of a feather of the second generation.
sh. 3, shaft, and af. 3, after shaft of a feather of the third generation. The
aftershaft of the protoptile i is from the outset free, but the tip of the aftershaft
of the feather of the second generation is, to start with, continuous with the
protoptile aftershaft, and the aftershaft of the feather of the third generation is
also for a time continuous with the aftershaft of the second generation feather—
according to Gadow the tip of the aftershaft of the second generation feather is
a forward prolongation from the calamus which is never during development

connected by its tip with the “calamus” of the protoptile or first generation
feather.

from the conclusion that, just as the aftershaft of the second
feather of the Emu is connected at the outset with the proto-
ptile, the aftershaft of the feather of the third generation is
connected with the feather of the second generation.

The feathers represented in Pl. VII. figs. 26 & 27 (from a seven

630 _ bRor. J, COSSAR EWART ON THH

months’ Emu bred in the Gardens of the Zoological Society of
London) prove conclusively that in the Emu the aftershaft in
origin and structure agrees with the shaft, while the feathers
represented in Pl. VII. figs. 24 & 25 (from a much younger Emu
in the Museum of the Philosophical and Literary Institute, Leeds)
as clearly indicate that as the aftershaft is developed before the
calamus it cannot be an outgrowth from the calamus. The first
three generations of the feathers of an Emu are diagrammatically
represented in text-fig. 11.

Before dealing with the aftershaft in the Carinate, attention
may be directed to the remarkable Emu feather of the second
generation represented in P]. VIII. fig.28. In this growing feather
the aftershaft, instead of resembling the shaft, simply consists of
two barbs bearing barbules. An Emu feather with an aftershaft
represented by two simple barbs is interesting, because in true
feathers near the oil-gland of ducks two long barbs sometimes
occupy the position of an aftershaft. That the two long barbs in
this Emu feather represent a phase in the degeneration of the
aftershaft is suggested by Pl. VIII. fig. 29, a developing true
feather with the protoptile intact from the rump of an eighteen
days’ Mallard duckling. Connected with the protoptile calamus
are two bundles of barbs—a large bundle with a distinct rhachis
which is obviously a developing shaft and a smaller bundle repre-
senting an aftershaft. The smaller or aftershaft bundle is con-
nected to the inner aspect of the protoptile calamus, as in the
Emu feather, by two long barbs. In course of time the large
bundle in Pl. VIII. fig. 29 would have developed into a shaft like
the one represented in Pl. I. fig. 1, and the small bundle into
an aftershaft with a short rhachis. Possibly long barbs which
represent an aftershaft will be met with in the feathers of other
Ratite.

The two long barbs in the Emu feather in Pl. VIII. fig. 28
support the view that the coat of the adult Emu consists of true
feathers and not, as Pycraft suggested, of mesoptiles.

3. The Development of the Aftershaft of the True Feathers of
Penguins.—In the Ringed Penguin the protoptiles though um-
belliform may be said to consist of two sets of barbs—an outer
representing a shaft and an inner occupying the position of an
aftershaft. Though in ducks and geese there is a well-developed
protoptile calamus containing “cones,” in penguins as in the
Emu a true protoptile calamus is never developed. Soon after
the protoptile escapes from its sheath the part of the epidermic
tube representing a calamus splits into slender cords, which
connect the barbs of the protoptile with barbs of the developing
mesoptile. The mesoptile consists of a shaft made up of seven or
more simple barbs which end in the tip of the true feather, and
of a complex aftershaft connected with the aftershaft of the true
feather. In text-figs. 3 & 4 the mesoptile aftershaft is con-
nected to the aftershaft of the true feather by a narrow band
formed by the fusion of the distal portions of the barbs of the

NESTLING FEATHERS OF Tite MALLARD. 631

aftershaft of the true feather. Pycraft considered the “ ribbon-
shaped stem” connecting the mesoptile to the aftershaft of the
true feathers as somewhat remarkable, because ‘‘in all other
instances yet known the nestling-down when attached to contour
feathers is attached to the rami (barbs) of the main shaft” (9).
When it is realised that in Penguins the mesoptile may have a
well-developed aftershaft the connection of the mesoptile with
the aftershaft of the true feathers ceases te be remarkable.
Though in the Ringed Penguin the mesoptile aftershaft is de-’
cidedly larger and more complex than the shaft, it is developed at
the same time as the shaft, and is directly continuous with the
aftershaft of the true feathers. That in Penguins the aftershaft
of the true feathers has the same origin as the shaft is made
abundantly evident when the development of the feathers of the
fourth generation is studied. In P]. VIII. figs. 30 & 31 we have
feathers from a moulting King Penguin in the Scottish Zoological
Park. From fig. 30 it is evident the new feather has from the
first been double, consisted of outer barbs representing a shaft and
of inner barbs in the position of an aftershaft. Had the develop-
ment proceeded the tip of the aftershaft would have been set
free when the old feather was shed. In fig. 31, the shaft having
been removed, the aftershaft is seen emerging from the tip of the
sheath which invested the growing feather. These figures show
that in Penguins the aftershaft of a feather of the fourth genera-
tion is for a time as complete as in the Emu. The aftershaft of ,
the mesoptile and of the first true feather of a Ringed Penguin
are diagrammatically represented in text-figs. 3 & 4.

4. The Development of the Aftershaft in the Ptarmigan, the
Common Fowl, and the Guillemot.—As already mentioned, it
has been suggested that in Grouse, Fowls, and other Galli the
feathers forming the first nestling coat may correspond to the
mesoptiles forming the second nestling coat of Penguins.
Against this view it may be pointed out (1) that unlike the
mesoptiles, but like the protoptiles of Penguins and Ducks, the
first nestling feathers of the Galli are developed in filaments; (2)
that in structure they are intermediate between the protoptiles of
Penguins and Ducks; and (3) that they are sometimes succeeded
by vestiges of mesoptiles. Degen mentions that in Grouse the
aftershaft may be two-thirds the length of the shaft but, as
Pl. IX. fig. 32 shows, the aftershaft in Grouse may be as long and
as complex as the shaft—the only difference in the feather
figured apart from its width is that, as in the Emu, the
rhachis of the aftershaft is slightly shorter than the rhachis
of the shaft. In the Ptarmigan the protoptile often con-
sists of fifteen barbs, eight representing a shaft with a short
rhachis, and seven forming an aftershaft also with a short
rhachis. From the protoptile calamus there proceed about
twenty barbs, which in some cases readily separate into three
distinct bundles—a large one destined to form the tip of the shaft,
and two small ones which later unite to form the tip of the

632 PROF. J. COSSAR EWART ON THE

aftershaft. In some metaptiles the rhachis of the shaft extends
to the tip of the feather, in others it ends in two slender barbs
some distance from the tip. Though at the outset in Grouse
both the shaft and the aftershaft of the true feather may consist
of several more or less distinct bundles, sooner or later the outer
barbs give rise to the shaft, while the inner form an aftershaft.
When the destined number of barbs for the shaft and aftershaft
have been produced the splitting of the epidermic tube is
arrested, and steps are taken to form a calamus in which

“cones” make their appearance. Evidently in Grouse, as in
Penguins, the aftershaft is not a forward elongation of the
calamus. An interesting stage in the development of a
Ptarmigan true feather is given in Pl. IX. fig. 33.

It has been asserted once and again that ‘in the great
feathers which form the ‘quill’ or‘ flight’ feathers (remiges)
of the wing and the tail-quills (rectrices) the aftershaft is in-
variably wanting” *, Assuming that in the case of the wing-
quills the aftershaft is invariably absent, it might be argued, that
these all-important feathers differed in their origin and history
from the other true feathers, that while the trunk feathers may
possibly have developed from an epidermic tube, the wing- and
tail-quills were formed by the splitting of elongated scales.
Attention has already been directed to the fact that in the Mallard
during the earlier stagesin the development of the wing- and tail-
quills, there is a vestige of the distal portion of an aftershaft (text-
figs. 5 & 7). That in wing- and’ tail-quills there may also be a
vestige of the proximal portion of an aftershaft is made evident
by Pl. IX. fig. 34, which represents the aftershaft of a wing-quill
from a full-grown Grouse.

Fowls are not supposed to have the aftershaft as well developed
as Grouse, nevertheless, as Pl. 1X. fig. 35 shows, the aftershaft in
the silky breed may be as long as the shaft. In its development
the aftershaft in the silky breed follows the same course as in
Ptarmigan.

_ In the Guillemot the aftershaft is, to start with, sometimes as
long as the shaft. A Guillemot feather from the side of the
trunk is represented in Pl. X. figs. 36 and 37. In this case the
protoptile consists of seventeen barbs. Of the seventeen barbs
nine end in the tip of the shaft of the true feather, four end in
barbs of the shaft which join the rhachis a considerable distance
from the tip, and four, which represent the protoptile aftershaft,
are connected with the aftershaft of the true feather (Pl. X.
fig. 37).

Bvidently the history of the aftershaft in Grouse, Fowls, and
Guillemots affords no support to the view that the aftershaft is a
secondarily acquired appendage, and may be said to fully estab-
lish the view that a complete true feather like a plumula consists
of a calamus and two shafts.

* Pycraft, ‘A History of Birds,’ p. 9.

NESTLING FEATHERS OF THE MALLARD. 633

III. Ortein anp History oF FEATHERS.

It seems to be generally assumed by naturalists and others
interested in birds that feathers are modified scales, and were
originally acquired to enable birds to fly. When evidence of the
scale origin of feathers is asked for, the reply usually is, ‘If
you accept the view that birds were evolved from Reptile-like
ancestors you may safely also accept the view that feathers are
specialized scales.” Having adopted the view that feathers are
modified scales, investigators interested in the coat of birds
proceeded to inquire if each feather corresponds to an entire
scale or to part of a scale, Wiedersheim favoured the view that
each feather represents a complete scale, but Bornstein (13) and
others came to the conclusion that a feather papilla only repre-
sents a fragment of a scale.

How scales were transformed into feathers has been indicated
by various writers. Baron Frances Nopcsa, in a paper “ On the
Origin of Flight,” states that we may quite well suppose “ birds
originated from bipedai long-tailed cursorial reptiles which
during running oared along in the air by flapping their free
anterior extremities,’ and added that the double running and
flapping action would subsequently easily lead to an enlargement
of the posterior marginal scales of the antebrachium, and even-
tually to their development into actual feathers(10). Lull,
another believer in the transmission of acquired somatic charac-
ters, points out that the anterior extremities of birds when in the
making would be more effective if their breadth was increased,
and that an increase of the size of the scales along the arm margin
would be a ready means to this end. Lull then proceeds to say
‘the “scales would extend, lighten, and ultimately evolve into
feathers which would not only subserve the function of flight,
but acting as clothing retain and aid in the increase of tempera-
ture” (11). Similar views are held by W. K. Gregory, who tells
us the Pro-Aves jumped lightly from branch to branch and
finally from tree to tree partly sustained by the long scale-like
feathers of the pectoral and pelvic wings.

The view that feathers are modified scales has long been held
by Pyeraft. In a work published in 1910 Pycraft points ont
that, to start with, the body clothing of birds probably consisted
of relatively large scales, ‘those covering the hinder border of
the incipient wing growing longer would still retain their original
overlapping arrangement and along its hinder border would in
their arrangement, appearance, and function simulate the quills
of modern birds; as their length increased they became also
fimbriated and more and more efficient in the work of carrying
the body through space.”

It is now evident that the wing-quills of birds are preceded by
nestling feathers, and may be regarded as modified protoptiles:
hence the question at issue is, Are the protoptiles modified scales?

634 PROF. J. CoSsAR EWART ON THE

It is inconceivable that the small protoptile forerunrers of the
wing-quills of Ducks were evolved from enlarged scales, which as
they increased in length became fimbriated. The more the
history of feathers is studied the more untenable becomes the
belief in their origin from scales. When special attention 1s
directed to the phases through which feathers pass during their
development, it becomes evident that in an attempt to trace their
origin we are not concerned with the wing-quills or other contour
feathers, but with the feather papille, and still more with the
filaments, in which the nestling feathers are developed. It is
conceivable that by the appearance of new and dominant factors
in the germ-plasm, scale papille, instead of developing into scales,
developed into feather filaments, but it is difficult to imagine
how scales by growing longer and splitting were transformed into
double-shafted nestling feathers. There is little chance of the
geological record ever throwing any light on the origin of either
feather papille or feather filaments, hence we must make the
most of the embryologicai record. Thus far the embryological
record has afforded no support to the view that scales in whole or
in part were concerned in the making of feathers.

When birds first acquired feather filaments it is impossible to
say, but we may assume that when the remote ancestors of birds
and mammals became warra-blooded any outgrowths from the
skin which prevented a loss of heat would from the first count in
the struggle for existence.

A hint as to the nature of the coat in the progenitors of
mammals we have from Armadillos, and an indication of the kind
of coat worn by the Pro-Aves is obtained by studying the deve-
lopment of nestling feathers, more especially of the protoptiles of
feather-footed birds, The Six-banded Armadillo has an extensive
scaly covering, but only a sparse coat of hair; birds, with few
exceptions, have during development a more or less complete
coat of filaments*, and probably in Owls the foot-feathers still
bear the same relation to scales as in the primeval birds. In the
developing Armadillo a stage is eventually reached when hairs
are seen projecting from under the developing scales or from the
skin between the scales. ‘The number of hairs developed in con-
nection with scales varies. There are often groups of three hairs
projecting from under the tail scales of mammals; in the Six-
banded Armadillo there are often three or four hairs to a scale,
but sometimes only one; in the small Argentine Armadillo Chlamy-
dophorus, as text-fig. 12 shows, fifteen hairs may be developed in
connection with one foot-scale.

In Armadillos and other mammals having scales as well as
hairs there is no evidence that the hairs are ever developed from
scales.

* All the birds I have examined, with the exception of the House Sparrow, had a
more or less complete coat of protoptiles—in the Rook a nestling coat is said to be
absent, but even in Rooks there are a few well-developed nestling feathers,

it.
-
Wy

i
oY
ih
ott

NESLLING FEATHERS OF THE MALLARD. 635

That in ancient birds feathers were intimately associated with
scales is suggested by the feather papille having approximately
the same arrangement as the scales of reptiles, by feathers
emerging from under scales above the tarsal joints of the Wood-
cock, and by chicks and ducklings having vestiges of scales in
connection with the feather filaments of the legs. In the case of
Grouse and other birds with only one feather emerging from
under a foot-scale, we are tempted to assume that each foot-
feather is developed from part of the scale it is associated with.
But when we find three feathers projecting from under the foot-
scales of the Barn Owl (Séria flammea) and seven or more
feather filaments occupying an area equivalent to a foot-scale, we

Text-figure 12.

Scale from the foot of the small Argentine Armadillo (Chlamydophorus) from under
which proceed a number of hairs developed from the skin underlying the scale.
The three filaments proceeding from under the foot-scales of Barn Owl embryos
like the hairs associated with the scales of mammals are developed from the

skin under the scales and not by a splitting of the scales or from fragments of
the scales.

are less inclined to admit that each of the foot feathers of Grouse
corresponds to part of a scale. When the development cf the
foot feathers of the Barn Owl is studied, one large and two small
filaments are found projecting from many of the scales; the large
filament contains a protoptile, and each small filament contains a
prefilopluma. Similar groups of filaments which occur above the
tarsal jot are overlapped by scale vestiges, but in the three
feather groups on the head scales are conspicuous by their
absence. The well-developed scales in connection with the foot-
feathers and the vestiges of scales in connection with feathers
above the tarsal joints support the view that birds, to staré with,
had a complete scaly covering and a sparse coat of feather filaments.

636 PROF, J. COSSAR EWART ON THE

As in feather-footed birds the feather papille are from the first as
distinct from the scale papille as the hair rudiments are distinct
from scales, there seems no escape from the conclusion that the
foot feathers are not modified scales. Further, when it is realized
that the foot feathers often belong to a simple primitive type,
have the aftershaft as long as the shaft, and that in many birds
there are seven or more feathers occupying the position of an
ancestral scale, we are justified in assuming that feathers, like
hairs, were originally intimately related to, but not derived from,
scales.

As a matter of fact, believers in the scale origin of feathers
admit that the foot feathers of recent birds are not derived from
the scales from under which they project. Just as it has been
assumed that the aftershaft in Emu feathers is a secondarily-
acquired structure, it is assumed the scales on the feet of birds
are secondary structures. That the scales overlapping the feathers
of modern birds were secondarily acquired and that the foot-
feathers were formed from the original foot-scales, are gratuitous
assumptions wholly unsupported by facts.

How or when feather filaments made their appearance it is
impossible to say, but we may safely assume that many centuries
elapsed before there was evolved in each filament a more or less
complex feather; needless to say that unless a filament in which
a feather was evolving was of more use than a simple filament,
natural selection took no part in providing birds with their
original coat of what are now familiarly known as nestling
feathers. On the other hand, when one bears in mind that blood-
vessels extend right to the tip of a feather filament, and not merely
a short distance into its root as in a hair, it is probable that early
specialization of the filaments was well-nigh inevitable. Given a
constant flow of blood through the axis (pulp) of the filament, an
inner epidermic layer capable of rapidly adding new cells to the
middle layer, an outer epidermic layer forming a protecting
sheath and, in addition, an ever increasing demand for a coat
capable of arresting the flow of heat from the skin, changes were
almost bound to take place in the bristle-lke outgrowths pro-
jecting from under or between the scales of the remote ancestors
of birds.

Judging by what takes place to-day during the development of
the simple protoptiles of pigeons, the first step in the making of a
feather probably consisted in the splitting of the middle epidermic
layer to form hair-like barbs. The result of this splitting of the

hollow cone formed from the middle epidermic layer of a pigeon ‘

filament is represented in text-fig. 13. The appearance of a
completed pigeon protoptile after escaping from its protecting
sheath is represented in Pl. X. fig. 38.

The protoptiles of Penguins represent a second stage in the
evolution of true feathers (text-fig. 3). Like the pigeon protoptiles
all the barbs are alike (form a tassel or umbel), but by sprouting
they have acquired barbules,

te tt

\ eae

NESTLING FEATHERS OF THE MALLARD. 637

A series of links connect the relatively simple umbelliform
protoptiles of Penguins with the highly-specialized protoptiles of
Ducks and Emus. There are good reasons for believing that the
warm-blooded progenitors of birds and mammals were evolved
out of reptile-like ancestors during a period when desert con-
ditions prevailed. How long this period lasted, and to what extent
feathers had been specialized when it came to an end it is impos-
sible to say, but judging by what happens during the development
of the wing-quills of the Mallard, it may be assumed the period of
aridity came to an end, aiid a glacial period had fairly set in long
before wing-quills and other true feathers were evolved. In
other words, before protoptiles had been modified to form meta-
ptiles and teleoptiles, progress in the evolution of true feathers was

Text-figure 13.

Section through the filament of a Pigeon in which the middle epidermic layer has
been split longitudinally to form barbs. s., the thin sheath which disintegrates
immediately after hatching. 6., one of the eight barbs formed by the splitting
of the hollow cone formed by the middle epidermic layer. p., the pulp con-
taining blood vessels, 7. e., the dermis or true skin which extends during
development to the tip of the protoptile. After Davies.

arrested owing to the necessity of providing birds with a fur-
like coat as capable of arresting the flow of heat from the skin as
the dense coat now worn by Polar Bears and other Arctic
mammals.

The discovery by Dr. Eagle Clarke and Mr. Pycraft of a
mesoptile coat in penguins has profoundly modified our views
about the plumage of birds. We do not yet know if the remote
ancestors of all our modern birds acquired a mesoptile coat —
probably in the case of the ancestors of the Ratite this was
unnecessary—but the more the nestling feathers are studied the
evidence of the existence of two coats of prepenne in the ancestors
of modern birds is increased. The fur-like mesoptile coat is
probably as well developed to-day in penguins as it was in their
remote ancestors of the Jurassic Age; relatively large mesoptiles
_ still occur in ducks and geese, and I have found recently distinct
vestiges of a mesoptile coat in pigeons, How long birds, or most

638 PROF. J. COSSAR EWART ON THE

of them, in order to survive required a fur-like coat throughout
life we have no means of knowing; but this much is certain, that
in most recent species of birds the mesoptile coat is in the act of
being suppressed, or has already disappeared—even in the King
Penguin chicks it is probably never worn in the Antarctic for
more than 9 or 10 months. From what we know about the
mesoptile coat it seems to have been originally produced partly
by lengthening the protoptiles and partly by the development of

Text-figure 14.

A developing mesoptile of an Adélie Penguin. In penguins the mesoptiles consist,
as a rule, of numerous barbs which in some areas may reach a length of over
three inches. A true protoptile calamus containing “cones” though present
in ducks is seldom mei with in other birds. As the protoptile is pushed from
the skin the inner ends of the proteptile barbs, destitute of barbules, separate
from each other and are subsequently seen to be continuous with mesoptile
barbs. mb., one of the numerous barbs of the developing mesoptile. pé., one
of the nine protoptile barbs, each of which is connected with a mesoptile barb.
b., one of the nine slender cords which together for a time playeu the part of a
protoptile calamus.

new barbs. This is suggested by text-fig. 14, which represents a
developing mesoptile feather from the breast of an Adeélie
Penguin. The nine barbs forming the protoptile are directly
continuous with nine mesoptile barbs; the rest of the mesoptile
consists of barbs having no connection with the protoptile. As
Pyeraft points out, the second coat of the Adélie Penguin is “a

4
;
i
i
‘a

NESTLING FEATHERS OF THE MALLARD. 639

long, dense woolly covering, so dense that it is difficult to expose
the skin.” In course of time the protoptile coat is shed and the
mesoptile coat is pushed from the skin by the true feathers.

The long fur-like mesoptile coat of the King Penguin suggests
that during the Ice Age, which succeeded the period of desic-
cation, the evolution of true feathers was arrested, and that the
mesoptiles cannot be regarded as representing a stage in the
evolution of true feathers. Though to the naked eye the meso-
ptile barbules seem to be identical with the barbules forming the
protoptiles, they are seen to differ when examined under the
microscope—instead of being nearly straight they are spirally
twisted at their origin. In the case of King Penguin chicks the
mesoptiles differ from the protoptiles more than in the Adélie
chicks, and, as already pointed out, the mesoptiles of the Ringed
Penguin decidedly differ from both the protoptiles and the
metaptiles. In all the three Penguins mentioned the first true
feathers differ from the prepennze which precede them. Hence,
though the prepenne and all the true feathers which succeed
them must be regarded as links in a chain, it should be borne in
mind that the first link differs both in function and structure
from the second, and the second from the third. Further, it
should be borne in mind that there are good reasons for believing
that, but for the advent of an Ice Age, the protoptiles at a
comparatively early period would have been succeeded by true
feathers, and that probably long before Archeopteryx made its
appearance on the scene some millions of years ago, steps would
have been taken in most birds to suppress the fur-like coat of
mesoptiles, now apparently only well developed in Penguins.

In conclusion, it may be stated that a study of the develop-
ment of birds suggests: (1) that their coat, to start with, consisted
of various kinds of simple filaments; (2) that in course of time
the hollow epidermic cone in each filament, in some incompre-
hensible way, was converted into a protoptile or preplumula:
(3) that the protoptiles in some cases soon acquired the chief
characteristics of true feathers ; (4) that for a time progress was
arrested in order to provide birds (or most of them) with a fur-
like (mesoptile) coat, by way of giving them a chance of surviving
during the cold phases of an Ice Age: (5) that as the climate
improved the mesoptile coat was in many cases superseded by a
coat of true feathers ; and (6) that by the specialization of feathers
along the posterior margin of the hands and forearms and the sides
of the tail, birds of the Archzopteryx type were eventually evolved
capable of flight, or at least of gliding easily from tree to tree.

Text-figures 1, 2, 8, 9, 12 are from drawings by Miss Augusta
Lamont, B.Se., text-figures 3-7 and 10, 11, 13, 14 are from
drawings by the author, all the other figures are from photographs
by Mr. John Chisholm.

Towards defraying the cost of the drawings and photographs
a grant was obtained from the Moray Fund of the University of

Proc. Zoou. Soc,— 1921, No. X LIII. 43

640 PROF. J. COSSAR EWART ON THE

Edinburgh and a grant has been voted from the Carnegie Trust
for the Universities of Scotland, towards the cost of publishing
this paper.

In addition to material in the form of ducklings, goslings &c.,
bred at Craigybield, Penicuik, it ought to be mentioned that
valuable contributions for the study of nestling feathers were
provided by the Director of the Zoological Society of Scotland,
and that but for the Institution of the Scottish Zoological Park
I would not have been led to undertake research work on the
Structure and History of Feathers. further, it ought to be
mentioned that during the investigation I received valuable
material from the Zoological Society of London, the Royal
Scottish Museum, and the Philosophical and Literary Institute
of Leeds, and that I am especially indebted for specimens that
proved most useful to Colonel Ashley, M.P., Miss Clerk of
Penicuik, William J. Gordon, Esq., of Windhouse, Yell,
Shetland, Professor Meakins, the University of Edinburgh,
and Professor Philip J. White, the University of North Wales.

References to Literature cited in Paper.

1. Iveram, CoLtitinewoop.—‘ The Ibis,’ October 1920, p. 856.
2. Gavow, Hans F.—Newton’s ‘ Dictionary of Birds,’ p. 243.
3. Pycrart, W. P.—‘‘ The Emperor and Adélie Penguins,” Nat.
Ant. Expedition, Zoology, vol. it. p. 12. 1907.
4. Owren, R.—Comparative Anat. vol. il. p. 235.
5. Nirzscu.— Pterylography,’ English Translation, p. 8. 1867.
6. Gapow, Hans F.—Newton’s ‘ Dictionary of Birds,’ p. 245.
7. Decen.—Trans. Zool. Soe. vol. xvi. pp. 362 & 363.
8. Pycrarr, W. P.—-Trans. Zool. Soe. vol. xv. p. 164.
9. an Nat. Ant. Expedition, Zool. vol. ii. pp. 7 & 8.
10. Norcsa.—Proe, Zool. Soc. 1907.
11. Luti.— Organic Evolution,’ pp. 533 & 534, 1917.
12. Pycrarr.— A History of Birds,’ p. 39. 1910.
13. Bornsrey.—Archiv f. Naturgesch., Band I. Supplement,
heft 4.

EXPLANATION OF THE PLATES.

PATE I.

Fig. 1. Protoptile and metaptile from vicinity of preen gland of a 38 days’
Mallard. The mesoptile has been suppressed. T he] long barbs proceeding
from the inner aspect of the protoptile calamus towards the calamus of
the metaptile are vestiges of a metaptile aftershaft. X 2.

2. Protoptile, mesoptile, and tip of a wing-quill of a 5 weeks’ Indian Runner
duckling.

3. Protoptile, mesoptile, and tip of a wing-covert of a 6 weeks’ Emden
gosling.

4, Tip of tail-quill of a young Ringed Penguin (u. s.).

‘Prats II.

.5. Tail-quill and protoptile of a 5 weeks’ Mallard duckling (n. s.).

6. A 10 days’ Mallard embryo. The tail-quill papille are well developed, but
there is tae indication of the wing-quill papille.

7. Wing of a 25 days’ Mallard duckling showing the wing-quill protoptiles (n.s.).

8. Wing- quill protoptile from a five ‘weeks’ Mallard duckling attached to tip
of developing mesoptile,

10.

ie
12.

Fig. 13.

14.

NESTLING FEATHERS OF THE MALLARD. 641

Prats III.

. Filaments from a 13 days’ Mallard embryo; the long filaments contain

developing protoptiles, the small ones preplumule or prefiloplume.

x 2.

Wing-quill protoptile artificially removed from a filament of a 26 days’
Mallard embryo.

Aftershaft of the protoptile of a duckling.

Wing of a 38 days’ Mallard duckling showing mesoptiles between the pro-
toptiles and the developing primary and secondary wing-quills. Lying
between developing wing-quills (each still invested by a sheath) are wing-
coverts in which the mesoptile is vestigial.

Prats IV.

Protoptile, mesoptile, and tip of wing-quill of a Chinese gosling. Though
the mesoptile seems to be umbelliform and to end in a calamus, it really
consists of a shaft and an aftershaft from which slender threads proceed
to the tip of the true feather. :

In this figure the proximal portion of the mesoptile is represented. It con-
sists of a narrow, perforated band representing the rhachis of the meso-
ptile shaft, of a broad perforated band representing the rhachis of the
aftershaft, and of numerous slender threads which connect these bands
with the shaft and the vestigial aftershaft of the true feather.

15. Nestling tail of a 7 days’ Mallard duckling. The protoptiles at each side

of the middle line have beeh pushed from the skin by the developing
true tail-quills.

16. The “nestling tail”’ at the middle of the fifth week. It now consists of

growing true tail-quills as well as of protoptiles, usually with the excep-
tion of the ninth from the middle line at each side the protoptiles are
directly continuous with the tail-quills—in other words, in the tail of the
Mallard duckling nearly all the mesoptiles have been completely

suppressed.
Prats V.

Fig. 17. Nestling feathers from the back of a 33 days’ Mallard duckling. The large

protoptile has a distinct calamus containing “cones.” ‘To the left of
the calamus are: (1) a filoplume with a minute prefiloplume on its: tip;
and (2) two plumule each carrying a preplumula, to the right a plumula
with a preplumula on its tip. X 5.

18. A plumula from a young Aylesbury duck showing a large shaft consisting

of long, slender barbs springing from a relatively short rhachis and a
well-developed aftershaft with a short rhachis.

19. The calamus of a naturally shed plumula with part of the sheath of the

growing plumula attached.

20. A developing filoplume with a prefiloplume at its tip from a 7 weeks’

Aylesbury duckling. X 2°5.

Prate VI.

Fig. 21. Developing plumula from a 5 weeks’ duckling. The aftershaft is developing

from apex to base in exactly the same way as the shaft. X 4.

22. A protoptile from a young Emu showing an aftershaft with a short

rhachis.

23. Emu feathers of the first (protoptile) and second generation. The protoptile

shaft is directly continuous with the shaft of the second feather, and the
protoptile aftershaft is continuous with the aftershaft of the second

feather.
Pirate VII.

Figs. 24 & 25. In 24,we have an early stage in the development of an Emu feather of

the second generation. In 25 the development of the second generation
feather has been nearly completed. In both cases the aftershaft is deve-
loped in the same way as the shaft—the tip of the aftershaft of the
second feather is continuous with the protoptile aftershaft and the tip of
the shaft is continuous with the protoptile shaft,

642 ON THE NESTLING FEATHERS OF THE MALLARD.

Fig. 26. Emu feathers of the first, second, and third generations. The tip of the
aftershaft of the feather of the third generation is continuous with the
rhachis of the aftershaft of the feather of the second generation.

27. In this specimen the tip of the aftershaft of the second generation feather
had lost its connection with the protoptile, and the tip of the aftershaft
ot the third generation feather had lost its connection with the “ ealamus ”
of the feather of the second generation.

Pratt VIII.

Fig. 28. Emu protoptile and a developing feather of the second generation, the
aftershatt of which is represented by two barbs. X 5.

29. Developing feather from rump of 18 days’ Mallard duckling. This feather
consists of two portions, one, the larger, representing a shaft, and one
representing an aftershaft, connected like the Hmu feather given in
fig. 28, to the protoptile hy two barbs.

30. Breast feathers from an adult moulting King Penguin. The growing
feather consists of a shaft and an aftershaft connected by their tips with
the calamus of the feather about to be shed.

31. In this figure the shaft of the new feather has been removed to show the
connection between the developing aftershaft and the old feather.

Prate DX:

Fig. 32. Grouse feather with an aftershaft as long and as complex as the shaft.

33. A protoptile and a developing true feather from a young Ptarmigan. The
part of the developing feather with a distinct rhachis represents the
shaft, the part m which a rhachis has not yet appeared represents an
aftershaft.

34. Calamas and aftershaft of the wing-quill of a Grouse.

35. Feather of a fowl of the Silky breed with an aftershaft as long as the shaft.

Prate X.

Figs. 36 & 37. Feather from a nestling Black Guillemot. 36, represents the shaft,
and thirteen barbs of the protoptile. 87, the four remaining protoptile
barbs connected with the rhachis of the aftershaft which was detached
along with the calamus.

38. Protoptile of a Wood Pigeon connected to the tip of a true feather by a
vestige of a mesoptile. The pulp projects from the tip of the teleoptile.
Note that the barbs resemble hairs, have no barbules.

SS:

ON A MALE KIVU GORILLA. 643

EXHIBITIONS AND NOTICES.
May 10th, 1921.

Dr. A. Smirx Woopwarp, LL.D., F.R.S., Vice-President,
in the Chair.

Sir S. F. Harmer, K.B.E., F.R.S8., exhibited a mounted spe-
cimen of a young female /hinoceros sondaicus, the one-horned
** Javan Rhinoceros,’ presented by T. R. Hubback, who shot it in
Lower Tenasserim, to the British Museum (Natural History).
Attention was called to the absence of a horn in the female, a
character which Mr. Hubback believes to be normal. A young
male of this species, from Java, was formerly exhibited in the
Society’s Gardens, and was described by Dr. P. L. Sclater, F.R.S.
(P. Z. 8. 1874, p. 182; Trans. Z. S.ix. p. 649). Although osteo-
logical material is to be found in various Museums, there appear
to be extremely few mounted specimens in existence. The Javan
Rhinoceros, which formerly ranged from Assam to Lower Bengal,
the Malay Peninsula, and Java, has become increasingly rare
during recent years, and it is in urgent need of protection, Its
occurrence in Sumatra and Borneo has been questioned, but
reference was made to the statements of W. Volz. (‘ Nord-
Sumatra,’ vol. ii. p. 372, Berlin, 1912), as proving its occurrence,
in limited numbers, in North Sumatra.

Mr. THos. Gerrarb, F.Z.8., exhibited, and made remarks
upon, a series of photographs of the skeleton of an old male Kivu
Gorilla (Gorilla gorilla beringert Matschie), the property of Lord
Rothschild, and also a series of photographs showing the milk-
dentition in the skull of a young African Elephant.

On behalf of Lord Rothschild, Mr. Gerrarp exhibited a speci-
men of a male Kivu Gorilla (Gorilla gorilla beringert Matschie)
mounted by Messrs. Rowland Ward, and stated that it differed
from the other races of Gorilla externally by its long, thick
pelage, intense black colouring, and the large fleshy protuber-
ance on the top of the head, which often measured 6 inches by
6 inches (7. ¢. width and height). The skull was much wider at
the occiput even than in G. gorilla matschiet Rothschild. The
specimen was a very aged male, long past the age of repro-
duction, and was found solitary, several miles from the two herds
inhabiting the region. The specimen was of great pathological

644 ON SNARE-BUILDING CADDIS WORMS.

interest, as it was the first instance known of pyorrhea alveolaris
in a Gorilla. Habitat Mikeno Volcano, Kivu District.

Mr. D. Seru-Smiru, F.Z.8., exhibited a series of skins and
photographs of some polyandrous Birds, and described their
mating habits.

Dr. Cuas. F. Sonnrac, F.Z.8., exhibited and made remarks
upon, some abnormalities in the Carnivora.

Mr. L. T. Hoesen, F.Z.8., and Mr. HE. A. Spauu exhibited,
and made remarks upon, the effect of Pineal Gland administra-
tion on the pigment-cells of Frog Tadpoles.

May 24th, 1921.

Prof. E. W. MacBrips, D.Sc., F.R.S., Vice-President,
in the Chair.

The Srecrerary read the following Report on the Additions
to the Society’s Menagerie during the month of April ;—

The registered additions to the Society’s Menagerie during
the month of April were 198 in number. Of these 81 were
acquired by presentation, 27 were deposited, 84 were purchased,
and 6 were born in the Menagerie.

The following may be specially mentioned :—

A pair of Lion Cubs (/elis leo), from Somaliland, presented by
Sir Geoffrey Archer, K.C.M.G., on April 21st.

2 Sloth-Bear Cubs (Melursus ursinus), from the Central Pro-
vinces of India, presented by Capt. R. C. Ansdell on April 5th,

2 King Vultures (Gypagus papa), from Tropical America,
purchased on April 6th.

2 Spot-billed Toucanets (Selenidera maculirostris), from Brazil,
purchased on April 28th.

A collection of Tropical Fish, including several species new to
the Collection, purchased on April 6th.

Miss L. E. Cunssman, F.E.S., exhibited, and made remarks
upon, a series of lantern-slides illustrating the habi's, in the .
larval stage, of the Hydropsychide (Snare-building Caddis
Worms).

DISTRIBUTION OF BIRD-LIFE IN PERU. 645

Major Stanuey 8. Frower, O.B.E., F.Z.S., exhibited examples
of three species of Egyptian Jerboas—Jaculus jaculus, Jaculus
orientalis, and Scarturas tetradactylus, the latter being the first
living specimens ever brought to England; and made remarks
upon the distribution of the three species in North Africa.

June 7th, 1921.

Prof, J. P. Hiuu, F.R.S., Vice-President,
in the Chair.

Major Srantey S. Fiower, O.B.E., F.Z.S., exhibited, and made
remarks upon, some living specimens of Testudo leithit and ar
example of Z'estudo ibera.

Dr. P. Cuatmers Mircueni, C.B.H., F.R.S., exhibited, and
made remarks upon, a photograph of the Death-mask of a young
Gorilla, presented to the Society by Prof. Howard Macgregor, of
Columbia University, New York.

Dr. Franx M. CuapmAn gave an account of the distribution of
Bird-life in the Urubamba Valley of Peru, and of his expedition
down the Urubamba river from its sources in the Puna zone, at
an elevation of 14,200 feet, to the Tropical zone at 3500 feet.
He reviewed the existing climatic conditions in Peru, and com-
mented on the faunal characteristics of the Puna, Temperate,
Subtropical, and Tropical zones encountered in the descent. A
comparison of the Avifauna of the forests of the humid Temperate
zone (alt. 10,000—12,500 ft.) with that of the adjoining treeless,
grass-grown plains and slopes of the Puna zone, showed that that
of the first-named area has evidently been derived from the
tropical forests at the eastern base of the Andes; that of the
latter from the South Temperate zone plains of Argentina and
Patagonia. The environmental change experienced in passing
from the hot, tropical forests, at the foot of the mountains, to
the cold temperate forests near their summits, is much greater
than that existing between the plains of Patagonia and’ those of
the Interandine tableland. This is obviously raflected in the
degree of differentiation exhibited by the birds inhabiting these
elevated forests and plains. Thus 55 per cent. of the genera of
birds inhabiting the Temperate zone forests are endemic, while

646 ON SOME MARSUPIAL EMBRYOS.

only 7 per cent. of the genera found on the plains of the Puna
are endemic. These facts are believed to indicate that the extent
of environmental change, expressed chiefly in climatic conditions,
is a more important evolutionary factor than those exerted by
time of isolation or extent of distance from the presumed ancestral
forms.

Prof. J. P. Hruz, F.R.S., V.P.Z.S., exhibited, and made remarks
upon, a series of lantern-slides of some Marsupial embryos,
dealing especially with the Koala (Phascolarctos) and the Wombat
(Phascolomys).

No. 217.

ABSTRACT OF THE PROCHEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*
May 10th, 1921.

Dr. A. Smita Woopwarp, LL.D., F.R.S., Vice-President,
in the Chair.

Sir 8. F. Harmer, K.B.E., F.R.S., exhibited, and made re-
marks upon, a mounted specimen of Rhinoceros sondaicus.

Mr. THos. GerRRARD, F.Z.S., exhibited, and made remarks
upon, a series of photographs of the skeleton of an old male Kivu
Gorilla (Gorilla gorilla beringeri Matschie), the property of Lord
Rothschild, and also a series of photographs showing the milk-
dentition in the skull of a young African Elephant.

On behalf of Lord Rothschild, Mr. Grerrarp exhibited the
mounted specimen of the Kivu Gorilla (mounted by Messrs.
Rowland Ward), and stated that it differed from the other races
of Gorilla externally by its long thick pelage, intense black
colouring, and large fleshy protuberance on the top of the head.

Mr. D. Sera-Suirn, F.Z.S., exhibited a series of skins and
photographs of some polyandrous Birds, and described their
mating habits.

* 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 Strpence, or, if desired, sent post-free for
the sum of Stix Shillings per annum, payable in advance.

Dr. Cuas. F. Sonwrac, F.Z.S., exhibited, and made remarks
upon, some abnormalities in the Carnivora.

Mr. L. T. Hoesen, F.Z.S., and Mr. E. A. Spaut, exhibited,
and made remarks upon, the effect of Pineal Gland administra-
tion on the pigment-cells of Frog Tadpoles.

My. R. I. Pocock, F.R.S., F.Z.S., communicated his paper on
“The Auditory Bulla and other Cranial Characters in the Mus-
telide (Martens, Badgers, etc.).”

Tn the absence of the Author, Mr. G. 8S. THapar’s paper “ On
the Venous System of the Lizard, Varanus bengalensis Daud.,”
was taken as read.

The next Meeting of the Society for Scientific Business will be

held on Tuesday, May 24th, 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 April, 1921.
Caas. F. M.D., F.Z.8., Anatomist to the Society.

“The Comparative Anatomy of the Tongues of the Mam-
malia.—IV. Families 3 and 4. Cebidee and Hapalide.

R. Broom, F.R.S., C.M.Z.8.

On some new Genera and Species of Anomodont Reptiles
from the Karroo Beds of South Africa.

Rel, Pocock, HRIS:, EIZS:

On the External Characters of some Species of Lutrine

(Otters).
C. W. Anprews, D.Sc., F.R.S., F.Z.S.

Note on the Skull of Dinotheriwm gigantewm in the British
Museum.

j 27
The following Papers have been received :—

Dr. C. F. Sonntac, F.Z.S.

The Comparative Anatomy of the Koala (Phascolarctos
cinereus) and the Vulpine Phalanger (77richosurus vulpeculu).

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

The External Characters of the Koala (Phascolarcios) and
some related Marsupials.

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 Lonbon,
ReEcENtT’s Park, Lonpon, N.W.8.
May \7th, 1921.

No. 218.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

May 24th, 1921.

Prof. E. W. MacBrips, D.Sc., LL.D., F.R.S., Vice-President,
in the Chair.

The Secretary read a Report on the Additions to the Society’s
Menagerie during the month of April, 1921.

Miss L. E. Curssman, F.E.S., exhibited, and made remarks
upon, a series of lantern-slides illustrating the habits in the
larval stage of the Hydropsychide (Snare-building Caddis
Worms).

Major Sranuzy 8S. Frower, F.Z.S., exhibited examples of three
species of Egyptian Jerboas—Jaculus jaculus, Jaculus orientalis,
and Scarturus tetradactylus, the latter being the first living
specimens ever brought to England; and made remarks upon
the distribution of the three species in North Afriea.

Dr. C. W. Anprews, F.R.S., F.Z.S., gave a réswmé of his
paper ‘‘ On the Skull of Dinotherium gigantewm in the British
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 Sex Shillings per annum, payable in advance.

Dr. Cuas. F. Sonntac, F.Z.8., gave a réswmé of his papers
(1) ‘Lhe Comparative Anatomy of the Tongues of the Mam-
malia, Families 3 and 4, Cebide and Hapalide,” and (2) ‘‘ Some
Points in the Anatomy of the Tongues of the Lemureidea.”

Tn the absence of the Author, Prof. R. Broom, F.R.S.,C.M.Z.8.,
his paper, ‘‘On some new Genera and Species of Anomodont
Reptiles from the Karroo Beds of South Africa,” was taken as
read.

Mr. R. I. Pocock, F.R.S., F.Z.8., gave a résumé of his paper

‘““On the External Characters of some Species of Lutrine
(Otters).”

The next Meeting of the Society for Scientific Business will
be held on Tuesday, June 7th, at 5.30 p.m., when the following
communications will be made :—

Dr. Frank M. CHAPMAN.

The Distribution of Bird-life in the Urubamba Valley of
Peru. (Illustrated by lantern-slides. )

S. Mauuik, F.Z.S.
New Indian Drilid Beetles.
IP gots do 126 deliei. Iolaus, Wale As

Exhibition of some Marsupial Embryos, especially the
Koala and the Wombat.

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

The External Characters of the Koala (Phascolarctos) and
some related Marsupials.

Dr. C. F. Sonntaa, F.Z.S.

The Comparative Anatomy of the Koala (Phascolarctos
cinereus) and the Vulpine Phalanger (Zrichosurus vulpecula).

Communications intended for the Scientific Meetings should
be addressed to

P. CHALMERS MITCHELL,

Secretary.
ZooLoGicaL Society oF LonDoN,
ReceEnt’s Park, Lonpon, N.W. 8.
May 31st, 1921.

tee aN

aie aeons mil

No. 219.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

June 7th, 1921

Prof. J. P. Hin, F.R.S., Vice-President,
in the Chair.

Major Stantry S. Frower, O.B.E., '.Z.8., exhibited, and made
remarks upon, some living specimens of Testudo leithit and an
example of Testudo ibera.

Dr. P. CHatmers Mircueto, C.B.E., F.R.S., exhibited, and
made remarks upon, a photograph of the Death-mask of a young
Gorilla, presented to the Society by Prof. Howard Macgregor, of
Columbia University, New York.

Dr. Frank M.CHApPMAN gave an account of the distribution of
Bird-life in the Urubamba Valley of Peru, and illustrated his
remarks with a remarkable series of lantern-slides.

In the absence of the Author, Mr. S8. Mauvtix’s paper on
“ New Indian Drilid Beetles ” was taken as read.

* This Abstract is published by the Society at its offices, Zoological Gardens,
Regent's Park, N.W., on the Tuesday following the date of Meeting to which
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,
to all Fellows who subscribe to the Publications; but it may be obtained on the
day of publication at the price of Sixpence, or, if desired, sent post-free for
the sum of Six Shillings per annum, payable in advance.

Prof. J. P. Hiut, F.RS., V.P.Z.S., exhibited, and made
remarks upon, a series of lantern-slides of some Marsupial
Embryos, dealing especially with the Koala (Phascolarctos) and
the Wombat (Phascolomys).

Mr. R. I. Pocock, F.R.S., F.Z.S., gave a résumé of his paper
on ‘The External Characters of the Koala (Phascolarctos) and
some related Marsupials.”

Owing to the lateness of the hour, Dr. C. F. Sonntae’s paper
on “The Comparative Anatomy of the Koala (Phascolarctos) and
the Vulpine Phalanger (Zrichosurus vulpecula) ” and Mr. C. Tarr
Reean’s paper on “The Cichlid Fishes of Lake Nyassa” were
taken as read.

The next Meeting of the Society for Scientific Business will
be held on Tuesday, October 18th, 1921, at 5.30 P.M.

A notice stating the Agenda for the Meeting will be cireulated
early in October.

The following Paper has been received :—

R. I. Pocock, F.R.S., F.Z.8.
The External Characters and Classification of the Mustelide.

Communications intended for the Scientific Meetings should
be addressed to
P. CHALMERS MITCHELL,

Secretary.
ZooLoGiIcAL Sociery oF Lonpon,

-Recent’s Park, Lonpon, N.W. 8.
June 14th, 1921.

23.

24,

25.

26.

27.

28.

29.

30.

dl.

32.

33.

34,

PAPERS.

The Geographical Distribution of Orthopterous Insects in the Caucasus and in
Meuiera Asia. by bP. UYAROV,; 2 CMlnp.); oe de cctinie viele o's ess Had ce bites re’s

The Auditory Bulla and other Cranial Characters in the Mustelide. By R. I. Pococs,
E-.R.S., F.Z.8. (Text-figures 14-18.) ....cecececsseeseoves taiahe) ct otatel sichniet ole. aie

On the Venous System of the Lizard Varanus bengalensis (Daud.). By Gopinp Sincu
amma, Vise, (Rext-fpuras lA.) sl es oe cs hace 6 eicciesie sa bin's/s\e sie) Ore siete selena s

Abnormalities in the Common Frog (Rana temporaria). By J. H. Luoyp, M.Sc.,
HZ.8.- (Lext-figure 1.):. 5/0... 05. tet eteceare BStcigy oGc osname Ulaos cobs an
Comparative Anatomy of the Tongues of the Mammalia.—IV. Families 3 and 4.
Cebide and Hapalide. By C. F. Sonnraa, M.D., F.Z.8S., Anatomist to the Society.
(Bext=fimuresia i Oas)\ ee iictele sisie's ceisjoisisiee ja) oh ew cies's cles ees ORO staresclevone Stars ya tecs -

Note on the Skull of Dinotherium gigantewm in the British Museum, By OC. W.
Anprews, D.Sc., F.R.S., F.Z.S. (Text-figures 1-4) .......... cece et ee eee eeee

On the External Characters of some Species of Lutrinz (Otters). By R. I. Pococs,
F.R.S., F.Z.S. (Text-figures 19-21.) .....0.. ees. eee eee Sal oi wi Sraptote ce cummhereeterale ae 5

The Oomparative Anatomy of the Koala (Phascolarctos cinereus) and Vulpine
Phalanger (Trichosurus vulpecula). By Ouas. F. Sonntag, M.D.,F.Z.8. (Plates V.,
Vik; Ae ee ENGI) Uke ols manasa elena Cisvels ister stone) a elmtete/n ie eter nfal srosees socstorscere

New Indian Drilid Beetles. By S. Mauuix, F.Z.S. (Text-figures 1-5.).........2- ‘

On some Abnormalities in the Carnivora. By Cuas. F. Sonntac, M. D.; E.Z.8.
(Text-figures 64-65. ) eeeteesese seve Ce @eeereece OF ereoe 8 ons

The External Characters of the Koala (Phascolarctos). By R. I. Pococx, F.R.S.,
EZS. (Text-figures 22-26.) ee seee @eonvere eereseee ee ee ee oe @oertes et eeerseeoe eo

The Nestling Feathers of the Mallard, with Observations on the Composition, Origin,
and History of Feathers. By J. Cossar Ewart, M.D., F.R.S., F.Z.S. (Plates I.-X. ;

Text-figures 1-14.) eee rvee sees Sotelo erelscaansieveyaiose, err eeseeresesesse es sears 000080

Page

447
473
487

493

497

535-

547

579

587 -

609

LIST OF PLATES.

1921, Parr III. (pp. 447-646).

Page
Cuas. F. Sonntag: Pls. V.-VI. Comparative Anatomy of the Koala
(Phascolarctos) 0. /'s ses sie cae es) MOLT

J. Cossan Ewart: Pls. I.-X. | Nestling Feathers of the Mallard .. 609

NOTICE.

The ‘ Proceedings’ for the year are issued in four parts, paged consecutively,
s0 that the complete reference is now P. Z. 8.1921, p.... The Distribution
is usually as follows :— :

Part I, issued in March.
Sey aed an ays June.
ob. i September.
AREA IATA December,

‘ Proceedings,’ 1921, Part II. (pp. 187-446), was published on July 8th,
1921.

The Abstracts of the ‘ Proceedings,’ Nos. 217-219, are
contained in this Part.

The dates of Publication of ‘ Proceedings’ 1830-1858 will be found in the
‘Proceedings’ for 1893, page 436.

The dates of Publication of ‘'T ransactions,’ 1833-1869, will be found in the
-« Proceedings’ for 1913, page 814.

PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF AEE =

ZOOLOGICAL SOCIETY
~ OF LONDON,
1921.

PART. LV.

conrainine Paces 647 To 887, with 6 PiatTEs aND
113 Texr-riGuREs.

ey lal Uist;
easel Slit Y
&

JANUARY 192
Ns,
4tlong) Muse
PRINTED FOR THE SOCIETY,
SOLD AT ITS HOUSE IN REGENT’S PARK.
MESSRS. LONGMANS, GREHN, AND CoO.,
PATERNOSTER ROW.

[Price Twelve Shillings. ]

LIST. OF CONEEANES:

1921, Part IV. (pp. 647-887).

EXHIBITIONS AND NOTICES.

The Szcrerary. Report on Additions to the Society’s Menagerie during the months of
May, June, July, August, and September, 1921......-+.. 1.0... eee ee eee eee ees

Mr. M. A. C. Hiyron. Exhibition of an embryo African Elephant ..-.......-...----

Mr. E. G. Boutrncrr, F.Z.S. Exhibition of photographs of a Common Grass Snake

swallLowimlertey Viletiere rele seals tetris ie PNG COM OE BE OA GOOCH 0020 AC
Mr. F. Marrin Duncan, F.Z.S. Exhibition of cinematograph films of Marine Animals ..

Mr. D, Serm-Smiru, F.Z.S. Exhibition of photographs of the Sacred Ibis nesting in the
Society’s Gardens ................

Prof, G. Exzior Smiru, F.R.S., F.Z.S. An Account of the habits of Zurszus, illustrated
by Tantern-slides 2220 ics7se) 0G hare, vee wl eee tie ei eae eet ae ee oer eee

My. Epear P. Cuancn, M.A., M.B.O.U. Exhibition of cinematograph films illustrating
the laying-habits of the Cuckoo (Cuculus canorus) and the life of the young Cuckoo ,

‘The Secretary. Keport on Additions to the Society's Menagerie during the month of
Oleoloere MeN Oss boso scat so aneobauuer EHide UM SON Akaa Ta aISCE RO OAS Ao Bu ac

Mr. A. &. La Sovir, C.M.Z.S, Exhibition of photographs Cae the early Ilfe-
LAVORO Ol IAU ON (OOM rolesn Gh OA ald op o wan oO FobGdG haan noah oon cat ones

Page

886

887

Dr. A. Smiri Woopwanrp, F.R.S., V.P.Z.S. Exhibition of a Human Skull and other —

TeMaims tromebrolcony Ell sNorbidhe hesitant eieiceiae eters oie ena ea

Contents continued on page 3 of Wrapper.

eT Pa

‘
)

ZOOLOGICAL SOCIETY OF LONDON.

Tars Society was founded in 1826 by Sir Sramrorp Rarruzs,
Mr. J. Sasinz, 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. yo sti
HIS GRACH THE DUKE OF BEDFORD, K.G., F.B.S., President.

t OO D2
AuFrep H. Cocks, Hse., M.A. Cot. diy Hiner 1 ¥enk AifON,

Ceoncns Drummonn, Ksa., E. ree ae i, A ee yey
Treasurer. SAFO AF OR| Neo)

Vice-President:

AurFrep Ezra, Ksa., O.B.E.
2 2 P. CHatmers Mrrcuetr,

C2B.n, MEAy DSc.
E.RAS., Secretary.

Tuer Hart oF Onstow, O.B.E.

Ksq.,
LL.D.,

THe Rient Hon. rae Viscount

Grey, K.G., P.C.
Sir Sipnny F. Harmer, K.B.E.,

M.A., Sc.D., F.RS., Vice-
President.

Pror. Jamuus P. Hirt, D.Sc.,
F.R.S., Vice-President.

Witiiam Huntsman, Ese.

Mason tHE Lorp AtAsSTAIR
Rosset Innes-Ker, D.S.O.

Pror.ErnestW. MacBrips,D.Sc.,
LL.D., F.R.S., Vice-President.

Masor AtBert Pam, O.B.E.

Taz Lorp Quennporoven.

H.G. Tae Doxezor Rurtanp, K.G.

Masor RicHarp S. Taytor.

A. Truvor-Bartys, Hse., M.A.

Awnrnony H. Winertetp, Ese.,
Vice-President.

A. Smrra Woopwarp, Esa.,
LL.D., F.B.S., Vice- President.

2
The Society consists of Fellows, and Honorary, Foreign, and
Corresponding Members, elected according to the By-Laws. It

garries 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-
munications should be sent, addressed to “The Secretary,” is open
from Ten till Five, except on Saturdays, when it closes at OnE P.M.

The Library, under the superintendence of Mr. F. Martin Duncan,
F.Z.S., F.R.M.S. is open daily (except Sunday) from Ten a.m. till
Five p.m.; on Saturdays, Ten a.m. till One p.m.

The Library is closed from Good Friday to Easter Monday, and
upon all other Bank Holidays. It is also closed annually for
cleaning purposes during the whole month of September.

The Meetings of the Society for General Business are held in
the Meeting Room at the Society’s Office on the third Wednesday
of the month at 4.30 p.m. except in September and October.

The Meetings for Scientific Business are held in the Meeting
Room at the Society’s Office-fortnightly on Tuesdays, except in
July, August, September, and December and January, at halfpast
Five o’clock pP.n.

The Anniversary Meeting is held on the 29th of April, or the
nearest convenient day, at Four p.m..

The Society’s Gardens are open daily from Nine o’clock until
Sunset. Mr. R. I. Pocock, F.R.S., F.L.S., is the resident Super-
intendent and Curator of Mammals, Mr. D. Seth-Smith is Curator
of Birds and Inspector of Works, Mr: E. G. Boulenger is Curator of
Reptiles, Miss L. K.Cheesman, F.E.S., is Curator of Insects. Appli-
cations for anatomical material or facilities for work in the
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ON ANOMODONT REPTILES FROM SOUTH AFRICA. 647

PAPERS.

30. On some new Genera and Species of Anomodont Reptiles
from the Karroo Beds of South Africa. By R. Broom,
FRS., CM-Z:8:

[Received April 6, 1921: Read May 24, 1921.
Jy
(Text-figures 28-45.)

INDEX.
Page Page
Dicynodon sollasi, sp. V.... ...... +45 648 | Eosimops newtoni, gen. et sp.n.... 663
7 andrewst, Sp. 0.......... 650 | Palemydops platysoma, gen. et
a OSUOTNI) SP. Ne) Sine oe 6al | SPELT Cees ace ed Nur es ea 665
3 watsoni, sp.n. ......... 653 | Hmydopsis tr Ge ise Broo)
us CLURGUS Se Neier 654 gen. n. as 666
py woodwardi, sp.N. ...... 655 = longus, sp. bans danasaene 667
A ictinops, SP. 0. ......... 656 | a Sciuroides, sp. n....... 668
macrorhynchus, sp.n.. 657 Emydops parvus, sp... ....... 669
Bainia tigriceps (Owen), gen.n.... 659 EHmyduranus platyops, gen. et sp.1 n. 670
» peavott,sp.n. ............... 659 | Hndothiodon crassus, sp. N.......... 672
LAUGH TONDASPS Heese ene OGL:

The genus Dicynodon was established by Owen in 1845 for the
remarkable South African fossil reptiles discovered by Bain and
referred to by him as Bidentals. As the result of the labours of
Owen, supplemented by that of later workers, nearly every detail
in the structure of the skeleton of Dicynodon is now as fully known
as that of the Dog. It was a primitive reptile with a skeleton built
essentially on the same plan as that of the mammal, but with
certain remarkable specializations of the skull, especially in the
complete loss of incisor teeth, the presence of a large permanently
growing canine in the male, and in the fact of the front of the
upper and lower jaws having formed a horny beak as in the
tortoise or bird.

While there remains little to add to our knowledge of the
skeleton, there is still much to be done ere we have “anything
like a satisfactory knowledge of the large number of species
which must be referred to the genus.

Dicynodon first occurs in South Africa at the base of the
Parevasaurus zone, which may be regarded as Middle Permian
in age, and it extends through all the zones to the top of the
Permian, and forms which are very closely allied, if not indeed
referable to Dicynodon, are met with to thé top of the Trias. A
form closely allied to Dicynodon oceurs in the Dwina of Russia,
and Placerias of the Trias of North America is, if not Dicynodon
also, at least very closely allied.

Oudenodon, originally described as a distinct genus differing
from Dicynodon in the absence of the tusk, we now know to be
the female. So long as all the known species of Oudenodon
differed from the known species of Dicynodon in other characters
besides the absence of the tusk, it was felt wiser to keep Oudenodon

Proc. Zoot. Soc.—1921, No. XLIV. A4

648 MR. R. BROOM ON ANOMODONT REPTILES

as a distinct genus. In 1909, in summing up the position at
that time, I stated that the evidence seemed ‘to be getting
stronger in favour of Oudenodon being but the female Dicynodon.”
By 1911 the discovery of tusked Tanid tuskless specimens of
Dicynodon bolorhinus in one locality removed all doubt, and
since then we have discovered tusked and tuskless specimens of
quite a number of species. We do not yet know whether the
females of all species are tuskless, and there is considerable reason
to believe they are not. And it is also possible that even the
males of some species at present referred to Dicynodon may be
tuskless. The genus Lystrosaurus is certainly tusked in both
sexes, and as all known specimens of Cistecephalus ave without
tusks, presumably both sexes are tuskless. All the known
specimens—about a dozen—of Kannemeyeria are tusked. Also
every known specimen of Dicynodon tigriceps or allied species 1s
tusked, while the few known specimens referred to Hocyclops are
tuskless. Species where both sexes are tusked or both tuskless
will bave to be placed in distinct genera or at least subgenera.

When Owen worked at Dicynodon and Oudenodon he made
nearly every skull the type of a distinct species, and there were
doubtless many who considered that he was over liberal in the
making of new species. Lydekker in 1890 reduced Owen’s
20 species to 11 good and 3 doubtful species, but there is now
little doubt, with our much greater knowledge of the genus, that
practically all Owen’s types are good. In my opinion, out of the
20 specific names only two can be regarded as synonymous—
D. pardiceps and D. rectidens, both of which I refer to D.
leoniceps Ow.

It is hardly surprising that a genus which survived for
probably over 2,000,000 years should have over 50 species
already known, and probably very many more to be discovered,
when we consider how many species of such reptiles as Zestwdo
or Lygosoma are at present alive. Until a few years ago nearly
every good specimen discovered seemed to belong to a new
species. Now the corner seems to be turned, and the majority
of Dicynodon skulls now picked up can be veferred to species
already known.

In the present paper I wish to des a iis a number of new forms
T have discovered in the past three years.

DIcYNODON SOLLAS!, sp. n. (Text-figs. 28, 29.)

This new species I have much pleasure in naming after
Prof. W. J. Sollas, who with his daughter has added so consider-
ably to our fuller knowledge of the Puan of the Dicynodon
skull*. It is founded on a series of over 20 good skulls and

* In connection with the work on Dicynodon by Prof. and Miss Sollas, I should
like to point out that the skull examined by them in 1913 cannot possibly be
Dicynodon leoniceps‘Owen. As the specimen is now, of course, completely gone,
and as the postorbitals have manifestly been partly weathered away before the
specimen was ground down, it will never be possible to determine the species with
absolute certainty. In my ‘opinion the specimen was Dicynodon feliceps Owen.

FROM THE KARROO BEDS OF SOUTH AFRICA. 649

many imperfect ones discovered near Biesjespoort on a horizon
which I believe to be at the very top of the Hndothiodon zone,
and it appears to extend a short way into the Cistecephalus
zone. Though many skulls, both males and females, are known,
I take as the type one of the best preserved, which I believe to
be an adult male. Itis about 5 inches in length and a little less
than 4 inches in width.

The beak is about the same length as the orbit and moderately
sharp in front. The nostrils are fairly large, and behind them
the maxillaries meet the nasals, and the septomaxillaries do not
appear on the side of the face.

Immediately behind the plane of the nostrils is a thickening of
the bones which forms a round button-like low boss in the middle
line. Above and behind the canine is a thickened elongated
ridge.

Text-figure 28. Text-figure 29.

Text-fig. 28.—Skull of Dicynodon sollasi Broom. Reduced.
Text-fig. 29.—Preparietal region in Dicynodon sollasi Broom. Nat. size.

The interorbital region is flat and narrow. The orbits look
upwards and outwards.

The parietal region is only very slightly wider than the inter- —
orbital, and the large postorbitals largely overlap the parietals
and almost meet in the middle line.

The preparietal region is not well displayed in the type skull,
but is well seen in a number of others.

The pineal foramen is relatively small, and is situated a few
millimetres behind the plane of the back of the postorbital arch.
The preparietal is small, and is flanked by fairly large anterior
processes of the parietals. The postfrontals are nearly obscured
by the parietals and postorbitals, but form a distinct though

small part of the orbital margins.
44*

650 MR. R. BROOM ON ANOMODONT REPTILES

The following are the principal measurements of the type
skull :——

Greatest antero-posterior length ...... 128 mm.
Width across squamosals ' ............... SB) tees
Intertemporal width eres. ee eeeeeeeee DD eae.
nterorbibaliiwidichy ieee ree tere eee ee Dal pees
Width across maxillary bosses ......... D2 Wes
Width between mostrils 24 27-ecee- «css HRS Pope oe
Width between canine roots ...:........ 32

Dicynodon sollast bears considerable resemblance to D. feliceps,
D. ictidops, and D. testudiceps. From D. feliceps it differs in
being a much more delicately built skui] and with a very much
feebler tusk, the tusk in J. sollasi being about half the thickness
of that in D. feliceps. In D. testudiceps the postorbitals meet
each other in the middle line over the parietals; in D. sollast
the postorbitals do not overlap the parietals so very greatly.
D. ictidops comes fiom a horizon which is probably 500 feet lower
than that which yields JD. sollasi; so that there is a strong
presumption in favour of the two being distinct species, though
they are undoubtedly closely allied. The preparietal in D. sollasi
is smaller than in JD. ictidops, and the general contours of the
bosses differ considerably in the two species.

DICYNODON ANDREWS, sp. n. (Text-fig. 30.)

This species is founded on a nearly perfect skull from the same
horizon and nearly the same spot at Biesjespoort that yielded the
type of D. sollasi. In addition to the type specimen I have two
others which I refer to the same species—one from a locality
between Murraysburg and Biesjespoort, and the other from a
locality 3 miles east of Biesjespoort. As this latter specimen was
found in near association with a specimen of Cistecephalus sp., 1b
gives us the horizon as the base of the Cistecephalus zone.

The species is a very strikingly new type, which may ultimately
have to be placed in a new subgenus.

In general shape the species resembles typical Dicynodons.
The snout is unusually short, the preorbital region being only
about one-fifth the length of the skull. Above and behind each
nostril is a well-marked bony boss, with a fairly broad flat suleus
between. Hach of the two specimens which has the mawxille
preserved is tuskless.

The orbits are large, with supraorbital thickening on the
frontal bones. The frontal region is shghtly concave behind, and
in front a low median elevation leaves each frontal bone slightly
concave.

The parietal region is broad, and unlike the typical species of
Dicynodon, the pavietals are not overlapped by the postorbitals.
The large size of the parietals makes a distinct approach to the
broad-headed Dicynodonts of the type of Dieynodon tigriceps
Owen, and for which I am proposing a new genus. In fact,
D. andrewsi may be the ancestor of D. tigriceps.

FROM THE KARROO BEDS OF SOUTH AFRICA, 651

The preparietal is well developed, and the relations of the
bones in this region are very similar to the typical Dicynodon
arrangement. There is a large postfrontal.

Text-figure 30.

Parietal region of Dicynodon andrewsi Broom.

The following are the principal measurements of the type :—

Greatest antero-posterior length... about 260 mm.

iBasalelemetine | es aa. dees see Rea KOE a8
Greatest: widthieess ja cce aes. coke ao Ours.
Wmbteronloubal waddlay 2 ahs eke. aoe Aes
intertemiporaliwidthapes...8 ss iynses seas sos AGH. 5.

I have much pleasure in naming the species after Dr. C. W.
Andrews, F.R.8., of the British Museum.

DiIcyNODON OSBORNI, sp.n. (Text-fig. 31.)

This very strikingly new species is from Wapatsberg, near
New Bethesda Road, and is from beds which are undoubtedly in
the Lystrosaurus zone. In the seme horizon are found not only
Lystresaurus, but the Therocephalian recently described by me
as Moschorhinus kitchingi. When Moschorhinus was described
the exact locality was unknown to me, and when later I dis-
covered it I made a special expedition to the spot to try and
discover the back portion of the skull. Though I did not succeed
in getting the rest of the type, I got another imperfect skull, two
specimens of Lystrosaurus, and the type skull which I am now
describing.

The present skull is not only of much interest as a new type,
but is of great importance as being one of the very few Dicynodon

852 _MR. R. BROOM ON ANOMODONT REPTILES

skulls known from the Lystrosaurus zone, and the only known
large one.

The skull lacks the lower jaw, and has lost by weathering the
front of the snout, the right jugal arch and part of the left, but
otherwise is fairly complete. Unfortunately, the matrix is hard,
and though most of the bony surface has been displayed, it is
almost impossible to be sure of the sutures.

The whole skull is broad and flat. When perfect it probably
measured about 163 inches in length, and it must have been
fairly wide.

Text-figure 31.

Skull of Dicynodon osborni Broom. Greatly reduced.

The snout is relatively long, and apparently resembled in some
respects that of Kannemeyeria, and as in that genus, there are a
pair of very powerful tusks. The nasal region has been broad
and rounded, without any very striking bony bosses.

The frontal region is wide and moderately flat. Low but well-
developed bosses are above the anterior half of each supraorbital
margin.

The intertemporal region is relatively narrow, and a marked
median crest is formed by the flattened parietal overlapped by
the large postorbitals.

The preparietal region has not been satisfactorily revealed.
The pineal foramen is evidently small, and situated well back
between the approaching postorbitals. Whether a preparietal

FROM THE KARROO BEDS OF SOUTH AFRICA. 653

bone is present or not the evidence does not show. There is
certainly no preparietal boss such as occurs in the species of
Hocyclops.

The orbits look almost directly upwards.

The postorbital arches are long and well developed,

The squamosals must have been extremely broad, judging from
the portions preserved.

The following are the principal measurements of the type :—

Greatest antero-posterior length... about 460 mm.

Greatest) wath. ere oe Ba 40 ules
Distance between tusks .................0085 "SO. 5
Mnteronbitall wid tay oe ces es <5 NGOr ar.
Jnitertemamearalltwrahbhy Yeates is cect ot eens 45

Basallene tii sce desde c sess about 410 ,,

Ihave much pleasure in naming this new type after Prof. H. F.
Osborn.

Text-figure 32.

SESS
asses

SSS
eS

Top of parietal region of skull of Dicynodon watsoni Broom.
The degree to which the postorbitals overlap the parietals is unknown, but most
probably it is not greater than is indicated in the figure.

DicYNoDON WATSONI, sp. n. (Text-fig. 32.)

This new species is founded on a very fragmentary skull, of
which little more than the frontal and parietal regions are pre-
served, found on the mountain to the east of New Bethesda,
about 800 feet above the horizon of the village and probably in
the Lystroswurus zone. It is a near ally of D. osborni though
quite a distinct species, and important in giving details of the
structure of the preparietal region.

654 MR. R. BROOM ON ANOMODONT REPTILES

The frontal region is broad and slightly convex, and the
parietal region is in the same plane as the general frontal plane.
There are no supraorbital bosses as in JD. osborni, the whole
frontal surface being unusually smooth and slightly convex.

The postorbitals are large and form a considerable part of the
upper cranial surface. Posteriorly they clasp the parietals, and
with them form the parietal crest. Curiously enough, each post-
orbital has had its posterior process broken off near the plane of
the pineal foramen, probably as the result of the bite of some
carnivorous enemy ; but there is little doubt that they extended
backwards, though not so far as in typical Dicynodons, as the
grooves for their lodgement are preserved. Probably, however,
they did not quite extend to meet the squamosals.

The preparietal is moderately large and wide, and forms the
margin of anterior third of the pineal foramen.

The postfrontal, though fairly large, is almost entirely hidden
on the surface by the frontals and the postorbitals.

I have much pleasure in naming the species after Mr. D. M.S.
Watson.

The following are the principal measurements :—

DIcYNODON cuRTUS, Sp. n. (Text-fig. 33.)

The type of this new species is a complete but distorted skull
found by me at Biesjespoort. The horizon is probably about

Text-figure 33.

Preparietal region of Dicynodon curtus Broom. Wat. size.

100 feet above that in which the type of D. sollasi was obtained,
but it is quite possible that D. sollasi may have survived to be
contemporaneous with this other type. in any case, though this
type is similar in size to D. sollasi, it is very strikingly different
in nearly every detail.

ewe

FROM THE KARROO BEDS OF SOUTH AFRICA. 655

The skull is short, broad, and deep, and about 4 inches in
length.

The snout is bent down and the caniniform processes are
under the orbits. There is no great thickening of the nasals.
The orbital region is narrow and the parietal region moderately
wide.

The pineal foramen is large and rounded. The preparietal is
small and narrow, and the postfrontals very largely developed.
The parietals are fairly well exposed between the postorbitals,
but are extremely short. The postorbitals in the temporal region
are broad and overhang the temporal fosse.

The nearest ally to this species, which I propose to call
Dicynodon curtus, is Dicynodon dubius Owen, from which it
differs in the unusual shortness of the parietals and in other
proportions.

The following are the principal measurements :—

Greatest antero-posterior length... about 110 mm.
Greatest breadth

Pi Lae aN eine Ae | ao CO
PasaliMlemathe sn waaeunree..cowerccocseueene tee ts LOOT ys
raterogboubaltwidishite tects. ncn. Woon ec ce thcedenis 20). 55
itertemporal width 2 kee-043.4).. cates: PY alae
Between the caniniform processes about 22 .

Text-figure 34.

\
‘

Preparietal and its relations in Dicynodon woodwardi Broom. Nat. size.

DiIcYNODON WoopWARDI, sp.n. (Text-fig. 34.)

The skull which forms the type of this new species was sent to

me from near New Bethesda by Mr. C. Kitching, and I am not

quite sure of its horizon. Pretty certainly it is from some part
of the Cistecephalus zone.

Though the whole of the base of the skull is preserved, much

656 MR. R. BROOM ON ANOMODONT REPTILES

of the upper surface is unfortunately lost, including both post-
orbital arches with the whole of the postorbital bones, practically
the whole of the parietal region, and both squamosals. Still,
what is preserved shows that it is a strikingly new type, and I
have much pleasure in calling it after Dr. Smith Woodward, of
the British Museum.

The skull is moderately small, having measured about 6 inches
in length. The snout is long and narrow, the preorbital portion
of the skull being nearly as ‘long as the postorbital. The nasal
region is imperfect, but there were probably no great bony
thickenings. The maxille have each a round tusk, the head of
which lies under the orbit, and the tusks are relatively farther
back from the front of the beak than in most species.

The frontals are long and moderately narrow. Behind them
lies a most remarkably broad preparietal quite unlike that of
any other known species. By its sides are narrow posterior
processes of the frontals. The postfrontals lie outside the
frontal processes. The postorbitals are lost, but from the con-
dition of the posterior parts of the squamosal it is manifest that
the temporal region must have been comparatively narrow.

The following are the principal measurements :—

xreatest antero-posterior length... about 160 mm.

Greatest breadth .......... . perhaps about 110 ,,
‘Basallplemot ly Me i See eect cee it scnn i 4 145,
dm berorlomucllewiachulia naasce a crear css ae SHO es
Distance betweenstuslss ye sc ee ee, DO nee

DicyNnopon ictinops, sp.n. (Text-fig. 35.)

This new species is formed on the nearly perfect skeleton of a
small animal found by me about 30 miles from Biesjespoort on
the Murraysburg Road. While it is quite possible that the form
is not quite pully grown, it cannot be the young of any known
species.

The head is about 3} inches in length, and the whole animal
from snout to the tip of the tail about 12 21 inches.

The snout is smooth and rounded, without any bony thickening
either on the nasals or in the supraorbital regions, except a very
slight thickening on the nasal just where it overhangs the
nostril. The septomaxillary does not appear to show on the
face. ‘Che lacrimal and prefrontals are small. In the maxille is
a small tusk.

The frontals are large, and a transverse section in the orbital
region would show a convex upper surface, while a section of the
posterior end would have a concave upper surface. The post-
frontals are small, and only a small part appears on the surface.
The preparietal is rather small, its anterior end being on the
plane of the postorbital arch. The pineal foramen is situated far
back. The parietals are small. The postorbitals are very large,

FROM THE KARROO BEDS OF SOUTH AFRICA. 657

and their posterior processes are kept apart farther than in most
Dicynodon species.

On some future occasion the skeleton will probably be described
in detail. For the most part the bones are surrounded by very
hard calcareous masses, and are very difficult to clear.

Text-figure 39.

Top of skull of Dicynodon ictinops Broom.

The following are the principal skull-measurements :—

Greatest antero-posterior length ...... eos ees araucnys
Greaveshmoreagiuht. .. nese thes ie S e 68 ais
hater orgies leawaalitslne yey ea eee ee Tolar
Initextemiporale widths. tunel ee VO S,
Basa lenethia aa aan Anus, atte about SOP tts,
Distance between canines ................5. NNO Nes

DiIcYNODON MACRORHYNCHUS, sp. n. (Text-fig. 36.)

This new species is founded on two small skulls discovered by
me at New Bethesda. For a time I considered they might be
very miniature specimens of Dicynodon platyceps, the common
New Bethesda species, but further careful examination shows
that it is necessary to regatd them as belonging to a distinct
species. The best preserved specimen is taken as the type.

The skull when complete was considerably less than 3 inches

658 MR. R. BROOM ON ANOMODONT REPTILES

in length, and if an adult it represents one of the smallest known
species of Dicynodon. It is chiefly remarkable for the great
length of the orbital and preorbital regions as compared with the
postorbital.

In the nasal region there is a large median thickening re-
sembling that in Dicynodon sollasi. The frontal region is
narrow and the orbits large. The postorbital arch is slender
and the postorbital bones are quite unusual in shape. There is
no trace to be seen, at least on the upper surface, of any post-
frontal, and the postorbital takes its place, having an anterior
process which forms a considerable part of the supraorbital
margin. ‘The posterior process is unusually short, much of the

Text-figure 36.

Top of skull of Dicynodon macrorhynchus Broom.

temporal wall being formed by the parietal. The pineal foramen
is of moderate size, and is situated as shown in the figure given.
The preparietal is long and narrow.

The posterior portion and much of the palate are crushed.

The following are the principal measurements :—

Greatest length ......... probably about 70 mm.
lankerormitallwadith: 2.0. cd0 sae eee CSO. ie
Iinterremporal width... -. eee eeeeeeeee 8 a
From snout to front of pineal foramen... 47 us

Dicynodon macrorhynchus is allied to D. sollasi, D. testudiceps,
and D. ietidops, but being from the middle of the Cistecephalus
zone, must be from a horizon many hundreds of feet higher than
that of any of these others.

seein

FROM THE KARROO BEDS OF SOUTH AFRICA. 659

BArNIA TIGRICEPS (Owen), gen. n.

One of the first species of Dicynodon to be described by Owen
was Dicynodon tigriceps. This is represented in the British
Museum by a very fine skull, said to have come from ‘ Gonzia
River, Kaffraria.” In general appearance it differs very con-
siderably from the typical Dicynodow as represented by D. lacer-
ticeps. The skull is massive and extremely broad and flat, and
the parietal region differs in that the parietals are well developed
and not covered by the postorbitals. The genus Dicynodon is so
large that it would be convenient if we could lop off a few species
and put them into a subgenus, even if the differentiating charac-
ters did not seem of very fundamental importance. But there
is another character that is of great importance. J know of nine
skulls which belong either to Dicynodon tigriceps or closely-allied
species, and all have tusks. It is therefore very highly probable
that the female of Dicynodon tigriceps was tusked, and uot like
the female Dicynodon—tuskless. And if this be so, Dicynedon
tigriceps must be placed in a distinct genus. It is, of course,
impossible to prove that the female was tusked, and even if
50 tusked skulls were found there would still be a doubt. As
it is, I think the probability sufficiently great to make for it a
new genus, bainia, after the father of South African geology,
Andrus Geddes Bain-—one of the most gifted geological geniuses
the world has seen. Even if’a tuskless femaie should turn up,
Bainia must still stand as a subgenus.

BAINIA PEAVOTI, sp.n. (Text-fig. 37.)

This new species is founded on a very fine skull discovered by
me at Biesjespoort. The skull is well preserved and not much
crushed. One zygomatic arch is lost, also part of the base and the
lower jaw. Though very similar in size to Bainia tigriceps, it
differs in many important features; and as it is manifestly a new
species, I have much pleasure in naming it after the late Henry
G. Peavot, who for some years was the Zoological Society’s
Librarian and Clerk of Publications, and whose care and kindly
help all workers so fully appreciated. It is a delight to be able
to add a stone to the cairn of a fellow-worker and friend who laid
down his life on the battlefield.

The skull is of special value in that all the sutures can be made
out with perfect distinctness. In general shape it agrees more
closely with Bb. tigriceps than\with B. laticeps, but differs in the
shape of the zygomatic arch and in many other details which
will be mentioned. ‘The snout and the whole front half of the
skull is much flatter than in any other known Anomodont.

The premaxilla, which is nearly complete, is relatively small
and unusually flat.

The nasals are short but very broad, and have each a well-
developed boss which overhangs the nostril. The suture with
the frontals and prefrontals is nearly transverse.

660 MR. R. BROOM ON ANOMODONT REPTILES

The septomaxilla is not satisfactorily preserved, but is relatively
small.
~ The maxilla is a very remarkable bone, quite unlike that in
any other known Dicynodont. The bone of the left side has lost
the subnasal portion, but is otherwise nearly complete, and it is
so remarkably shallow that one would fancy the palatal half had
been removed; yet if part of the bone has been removed, it must
have been done during the animal’s lifetime. There is no trace
of a tusk, yet there are the remains of the socket of a large tusk.

Text-figure 37.

Sor re
== rs

Skull of Bainia peavoti Broom.

A not improbable explanation of the peculiarity is that the
animal is aged and has lost its tusks, and that in consequence
the palatal portions of the bone have become absorbed, as is seen
in aged human and other mammalian jaws.
The prefrontal is a relatively small trianguar bone which has
a small and not very prominent boss above the front of the orbit.
The lacrimal is small.

FROM THE KARROO BEDS OF SOUTH AFRICA. 661

The jugal is large. It forms most of the lower border of the
orbit, most of the postorbital arch, and a considerable part of
the zygomatic arch.

The frontal is unusually large and wide. Its shape will be best
understood from the figure given. It has apparently a large
articulation with the postorbital, but a thin little strip of the
postfrontal is really wedged in between them. ‘The frontal
passes back to the plane of the posterior end of the pineal
foramen.

The postfrontal scarcely appears on the surface, but though
nearly hidden by the frontal and postorbital, it is really a bone
of appreciable size underneath.

The postorbital is a large and powerful bone. It forms most
of the strong postorbital arch and the whole of the upper border
of the temporal fossa, and pretty certainly articulates with the
squamosal.

The preparietal is unusually large. It passes far forwards
between the frontals, and posteriorly nearly surrounds the pineal
foramen.

The parietals appear to be ankylosed into a fairly large bone
lying mainly behind the pineal foramen. It forms a moderately
large. somewhat concave intertemporal region. Anteriorly it
sends forward on each side a slender process between tie frontal
and postorbital, and posteriorly a long process between the
postorbital and interparietal.

The interparietal is a little broader than the middle part of
the parietal. Laterally it articulates with the large tabular.

The occiput is fairly well preserved, and was probably much
like that of Bainia tigriceps, though some degree of crushing of
the squamosals gives it a rather different appearance. The
occipital condyle is much shorter in the present species.

The squamosals are, as in all Dicynodonts, large and powerful.
The shape of the zygomatic portion can be seen in the figure to
differ appreciably from that in B. tigriceps.

The following are some of the principal measurements :—

Greatest length (oblique) ............... 500 mm.
Greatestioresdtl 2 yo. 0 see. douse ev esewsen 480 ,,
Beisallb lemon iatiiecus cits Sis: ily cy eal ce AGO) ee
Width across nasal bosses ............... DOS,
Interorbutal wadth 27225 eee ene T3307.
Intertemporal width across parietals... 50 ,,
Intertemporal width across post-

OL Italy eee eas AA te I alk 86

BAINIA HAUGHTONI, sp.n. (Text-fig. 38.)

In Haughton’s paper on the Anomodonts in the South
African Museum, recently published, he figures and describes a
skull (8S. A. M. No. 3328) which he believes to be a young

662 MR. R. BROOM ON ANOMODONT REPTILES
I have examined this skull,

specimen of Dicynodon laticeps.
and while it has all the appearance of a young skull, it cannot,

in my opinion, be the skull of a young D. laticeps, as it differs in
having a very large preparietal and very large postfrontals.
Nor can it be the young of either Bainia tigriceps or D. peavoti.
And as it differs from all these three species, but manifestly
belongs to this group, it is necessary to make it the type of a
distinct species which I have much pleasure in naming after
Mr. 8S. H. Haughton, who is doing excellent work in the vast

South African field.
Text-figure 38.

dD
ZAILINNS

EES.

IV Po.

Parietal, preparietal, and frontal regions of Bainia haughtoni Broom.

Mr. Haughton, in addition to describing the specimen, has
given a figure of the preparietal region. While his figure is
essentially correct, a camera lucida drawing I have made of this
region gives, I think, a little more accurately the position of the

sutures.
The following are some of the principal measurements :—

imberorbitale width. .6-cceo see eee 46 mm.
Intertemporal width ..............0...0+- Boas
fda sla oae alae ee Ree ae

Across the nasals

FROM THE KARROO BEDS OF SOUTH AFRICA. 663

EosImMoPs NEWTONI, gen. et sp.n. (Text-fig. 39.)

This genus and species is founded on a skull which I dis-
covered at Victoria West, and which is of special importance as
throwing some light on the age of the rocks which yielded the
unique fossil forms Galechirus scholtzi, Heleosuchus scholtzi,
Arnognathus parvidens, and Heleophilus acutws. When these
fossils were discovered by Mr. Scholtz no other fossil had been
found within 40 or 50 miles, and there was much doubt as to
the age of the deposit. When I described the forms I thought
it very likely that they were some of the land representatives of
the Lystrosaurus zone. Shortly afterwards Mr. A. L. du Toit,
of the Geological Survey, reaching Victoria West from the north

Text-figure 39.

Skull of Hosimops newtoni Broom.

The apparent asymmetry is partly due to crushing, but largely owing to different
degrees of the surface of the bones having been flaked off.

through a very unfossiliferous region, came to the conclusion
that the beds at Victoria West were probably of the Pareia-
saurus zone. For many years I have taken Mr. du Toit’s
opinion, but working from the South I have been compelled to
differ. Biesjespoort, which lies about 20 miles south of Victoria
West, is very rich in fossils, and is certainly at the base of the
Cistecephalus zone. For miles the strata are almost perfectly
horizontal, and it is hard to helieve that the beds at Victoria
West can be much lower than those at Biesjespoort. Unfortu-
nately, the intermediate beds so far as examined are almost
Proc. Zoou. Soc.—1921, No. XLV. A5

664 . Mi. R. BROOM ON ANOMODONT REPTILES

devoid of fossils till we come to the little rich patch at Victoria
West. Soon this locality will be for ever drowned in a oe
dam which is being made above the township.

A small Dicy nodont skull was some years ago discovered in
the deposit, which I referred to Pristerodon aches Huxley,
and took this as confirmatory evidence of the deposit being in
the Pareiasaurus-zone; but we now know that very Se
forms belonging to the genus I am ‘calling Hmydopsis-oceur in the
Oistecephalus zone.

The new Dicynodont skull which I have discovered is nearly
perfect, but being in a hard matrix cannot be very satisfactorily
developed: ‘It is’ about 5 inches long and 4 broad, and is
chiefly remarkable from having the preorbital region very short
and deflected and the parietal region very broad.

The premaxilla is extremely small and short. The nostrils are
large and directed forwards. The nasals,. like the premaxilla,
are also unusually small. The septomaxilla does not show on
the facial surface. The maxilla though fairly well developed is
very short. It has a rather small slightly flattened tusk.

The prefrontal and lacrimal are both small. The frontals are
long and narrow, and form only a small part of the orbital
margin. Between them lies an exceptionally large and long
preparietal. The postfrontals are also very large. The post-
orbitals, on the other hand, have very short posterior processes
which do not meet the squamosals, The parietals are well
developed and broad.

The pineal foramen is unusually small.

The squamosal does not seem to present any unusual features.

The occiput has not been cleaned of matrix,

The species in the folding down of the snout recalls Lystro-
saurus, though the narrow interorbital region gives the upper
surface quite a different appearance. In Dicynodon moschops
Broom we have a very similar folding down of the snout, and at
first sight one might be led to think that this skull was the male
of D. moschops. When, however, we look at the structure we
find very great differences. In D. moschops we find the septo-
maxillary large and forming part of the face. Here it does not
appear on the face. Further, in D, moschops, though the parietals
are also broad the preparietal is small, and there are no post-
frontals showing on the upper surface. The species of Bainia
also show some affinities with the present type, but they all differ
in having the frontal region broader than the parietal. Still we
may conclude that this Victoria West type has aflinities with
D. moschops on the one hand and with the species of Bainia on
the other. As all the known species of Bainia occur in the
Oistecephalus zone, as does also D. moschops, we may, I think,
conclude that the Victoria West animal is either from the
Oistecephalus zone or from the zone immediately below—viz., the
Endothiodon zone. In any case it is pretty certain that the
Victoria West deposit cannot belong to the Pare tasaurus zone.

FROM THE KARROO BEDS OF SOUTH AFRICA. 665

It is certainly very remarkable that, while we know the faunas
of the Hndothiodon and Cistecephalus zones fairly well, nowhere
else have we ever come across any of the forms that occur at
Victoria West.

Though the present Dicynodont is fairly new to the typical
Dicynodons, I think it well to make it the type of a new genus,
which may be called Hosimops, characterized by the very short
deflected snout and the broad .and unusual condition of the
parietal region. The specific name has been given in honour of
Mr. E. T. Newton, F.R.S., whose work I have always greatly
admired.

The following are the principal measurements :—-

Greatest antero-posterior length... about 140 mm.

Greatest) ORE AGU hier i a1 mee Tidy, LOOW fe
Janibesor bitterly, oo) ase ent eevee i BOie ee
linterteniporalawadtiny (amu eek vec SOU

PALEMYDOPS PLATYSOMA, gen. et sp. n. (Tl ext-fig. 40.
»§ i §

This new genus and species is founded on a beautiful skull and
part of the skeleton of a small Dicynodont found by me at

Text-figure 40.

Skull of Palemydops platysoma Broom.

Biesjespoort. ‘The head is barely 3 inches long, and one might

consider the possibility of its being a young Dicynodon. But this

cannot well be the case, as the bones are well ossified and the

tusk apparently fully formed. Further, the structure of the top

of the skull is unlike that in any known species of Dicynodon.
45*

666 MR. R. BROOM ON ANOMODONT REPTILES

In fact, the top of the skull differs so greatly from that of
typical Dicynodons, that I feel justified in making the specimen
the type of a new genus, Palemydops.

The skull is flat and broad. The snout is very short. There
are no indications of any thickenings of the nasal bones. The
maxilla is short and has a well-developed round tusk. The
sutures of the snout elements cannot with certainty be made out.

‘ne frontals are moderately broad, and on the back part of
each is a long deep, probably glandular pit. The postfrontals are
large, and form a considerable part of the orbital margin. The
postorbitals are very large, and form the whole of the inner wall
of the temporal fossa.

The preparietal is relatively small and narrow. The pineal
foramen is small, and lies much behind the plane of the post-
orbital arches. The parietals are large.

The following are the principai measurements of the skull :-—

Greatest antero-posterior length ...... 75 mm.
GREASE eye lie ohisb cite uo Aatctea oat nace Ri aakias Oily was
erntencoxglorivall whit a o2 ee oc ae 5. Sele oye
Intertemporal Wath cock... cder ees AD) ee

EMYDOPSIS TRIGONICEPS (Broom), gen. n. (Text-fig. 41 A.)

In 1904 I described a small Anomodont skull as a species of
Oudenodon and called it Oudenodon trigoniceps. With the much

Text-figure 41.

A. Preparietal region of Hmydopsis trigoniceps Broom.
B. Preparietal region of Hmydopsis sciuroides Broom.

greater knowledge which we now have it became pretty manifest
that it could not be a species of Oudenodon, or rather of
Dicynodon, the parietal region being quite different from the
Dicynodon type» The specimen is now in the Albany Museum,
and Mr. J. Hewitt, the Curator, kindly sent it to me on Joan for
further examination. I discovered on breaking into the maxilla
that there are three or four slender molar teeth. These are long

FROM THE KARROO BEDS OF SOUTH AFRICA, 667

and rounded, and with small serrations on the posterior side of
the upper portions of the crowns. Manifestly the genus is allied

_to Pristerodon and to Hmydops, and also to Dieluwrodon, but it

seems wisest to make a new genus for those forms with three or
four slender posteriorly serrated teeth. Pristerodon has a larger
number of molars—6, 7, or 8—and they are robust teeth.
Emydops has a few teeth which are apparently not serrated.
Diclurodon has a number of molars which have serrations both
in front and behind.

Emudopsis trigoniceps (Broom) may be taken as the type
species of the new genus.

Emypopsis Loneus, sp. n. (Text-fig. 42.)

This species is founded on a collection of eight good and a
number of imperfect skulls found by me at Biesjespoort. ‘They
occur in exactly the same horizon as Diceynodon sollast. As most
of the skulls are about 22 inches in length, and no one is more
than 22 inches, we may safely conclude that they are adult
skulls,

Text-figure 42.

Skulls of Hmydopsis longus Broom.
A. Type skull.
B. Skull of a topotype illustrating the variability of the preparietal.

The snout is very appreciably shorter than the antero-posterior
diameter of the orbit. It is above rounded and moderately
smooth, though the nasal bones are uniformly thickened. The
septomaxillary does not seem to be seen on the face, though it is
impossible to be quite certain.

668 MR. R. BROOM ON ANOMODONT REPTILES

The frontals are long and narrow, and nearly shut out from the
orbital margins by the preefrontals and large postfrontals. They
pass far backwards by the sides of the preparietal. The frontals
are moderately flat.

The preparietal differs in different specimens considerably, but
the appearance of the postfrontals, postorbitals, and parietals is —
very constant. I give figures of the region in two specimens to
illustrate the variation.

The parietals are large and broad and flanked by the post-
orbitals.

In no specimen is there a tusk, but in both upper and lower
jaws are a few slender posteriorly serrated molars. The number
appears to vary with age. In a beautifully preserved lower Jaw
there are three teeth. The first is large and is on the point of
being shed, the replacing tooth being seen below it. The second
tooth is also well developed. he third tooth is very small.

A few years ago Haughton and I described a small skull from
Dunedin, which we provisionally referred to Hmydops and named
Emydops piatyceps. There can be little doubt that this is very
closely allied to the present species, and should be transferred to
the genus Hmydopsis.

The only striking difference between Hmydopsis platyceps and ~
Emydopsis longus is that in the latter the posterior part of the
skull is much longer and broader, and while #. platyceps has
slender tusks, #7. longus has none.

The following are the principal skull measurements (in milli-
metres) of three specimens :—

Type. B. C.
Greatest length ............ 64 68 (2
Greatest width ...... about 48 about 54 about 52
Interorbital width ......... 12 11 12
Intertemporal width ...... 18 18 20
Basaldlencthy sce. ave... 59 about 62 —

EMYDOPSIS SCIUROIDES, sp. n. (T'ext-fig. 41 B.)

The beautiful little skull which forms the type of this new
species was found by me at New Bethesda, the famous locality
which has already yielded so many interesting types. Without
seriously injuring the skull, it will not be possible to say with
certainty if the specimen belongs to Hmydopsis, but the general
resemblance of the skull and lower jaw to those of species of
Hmydopsis is so close as to render it highly probable that the
specimen belongs to a species of this genus.

The next striking characteristics of the skull are the large size
of the orbit and the relative shortness of the parietal region.

The snout is short but relatively broad. A tiny corner of the
septomaxillary appears on the face. The maxilla is deep, and in
this type skull at least, tuskless. The prefrontal and lacrimal
are small.

FROM THE KARROO BEDS OF SOUTH AFRICA. 669

The frontal is very long and narrow, and remarkable in that
the postorbital portion is wider than the orbital. The preparietal
is almost unique among Anomodonts in being very small and
situated entirely in front of the pineal foramen. A similar
condition is only known to occur in Emydopsis arctatus (Owen).
The postfrontal is either entirely absent or does not appear on
the surface. The postorbital is very large,and extending forward
into the region usually occupied by the postfrontal, forms a con-
siderable part of the orbital margin. The parietal is moderately
wide but short, the part behind the pineal foramen being
extremely short. ‘The pineal foramen is very small.

In the lower jaw the dentary resembles that of Hmydopsis
longus, but the post-dentary portion of the jaw is relatively much
shorter.

The following are the principal measurements of the skuil :—

Greatest lemothy 0. 2.2. ..6665tekees so about 60 mm.
Greatest width ......... doubtfuily about 50 ,,
Toterorbitalwidtbr i) a.28).2 ono dae OSes
Wnitertemporal width .0 3.015 ae. get maces Teli 45.
Between caniniform processes ............ Llores 5,
Basa lemau hie hoes ceacca ts one about 48 ,,

Skull of Hmydops parvus Broom.

The skull is slightly crushed, but could not well have been longer than
indicated by the dotted line.

Emypors PARVUS, sp. n. (Text-fig. 43.)

Until the discovery of the present specimen, Hmydops minor
was the smaliest known Anomodont, but this new find gives us a
type which has certainly a shorter skull and possibly smaller in
other respects. This new species is founded on a skull discovered
by me at Bruintjeshoogte in the same stratum as yielded the
type skull of Ictidosuchus longiceps Broom.

The skull is 14 inches long and rather less in breadth, and the
whole animal was probably as small as a newly-born kitten.

670 MR. R. BROOM ON ANOMODONT REPTILES

The snout is very short, but the structure cannot be made out
owing to crushing.

The orbits are small.

The frontal region is relatively narrow, and the temporal
region in front nearly twice as wide. Owing to a finely crackled
condition of the bones, sutures are very difficult to make out, but
the limits of the preparietal can be clearly seen, The pineal
foramen is small and situated well back. The general shape of
the skull will be best indicated by the drawing given.

There is evidence of at least one small unserrated molar,

The following are the principal measurements of the skull :—

Greabestjlemetin, 5... .cceherons see cay about 38 mm.
Greatestuatea dtl, <5. .assh eee ee oy nee
hatererbitalaid tly. oft casas eines SD) i,
Mnibertemnyonall wad tho 5 ees eee Tay ae

EMYDURANUS PLATYOPS, gen. et sp.n. (Text-fig. 44.)

The new genus and species is founded on a small skull dis-
covered by me near Biesjespoort, and is one of the most interesting
Anomodonts known. The specimen was found in the deposit
which has yielded the very numerous skulls of Dicynodon sollasi,
and before being developed it was looked upon as another of the
many duplicates. This was unfortunate, for a beautiful shoulder-
girdle lying against the skull was developed out and a portion of
the skull sacrificed before it was noticed that the skull is really a
very remarkable new type. We have still preserved the greater
part of the skull, minus the arches, but with the contact between
the occiput and the parietal region lost.

The skull is that of a small Anomodont with a broad, flat head
and a wide palate which has on each side three or four teeth.

The premaxilla is broad and shallow. As in typical Anomo-
donts, it forms the greater part of the bony palate. There is a
well-marked median ridge. Its relation to the palatines and
maxillze will be best understood by the figure.

The maxilla is well developed, but is remarkable from the fact
that there is very little of a caniniform process. Some distance
inside of the alveolar margin are three or four teeth in a row.
On the right side the matrix has been left supporting the crowns.
The first is a small sharp-pointed unserrated tooth; the second a
much broader flattened tooth, also without serrations; the third
probably similar to the second; and the fourth a smaller tooth
imperfectly preserved. On the left side the matrix, and with it
much of the teeth, has been removed. The first tooth is seen
to have its root near the suture between the premaxilla and
maxilla, and though it doubtless is fixed in the maxilla, it passes
through the premaxilla. The others are in the posterior part
of the maxilla.

The palatine is well developed and has an unusually large

FROM THE KARROO BEDS OF SOUTH: AFRICA. 671

palatal portion, which is rugose and has probably had a horny
protection. The posterior part of the palate is not preserved.

The nostrils are situated well forward and separated by the
median process of the premaxilla, which passes well back.

The nasals are large. Each has a well-marked boss behind
and above the nostril. The septomaxille cannot be made out.

The frontal is long and narrow, and passes unusuaily far
behind the plane of the postorbital arch. The postfrontal is
small and narrow, but forms a considerable part of the orbital
margin.

Text-figure 44.

A. Top of skull of Emyduranus platyops Broom.
B. Palate of Hmyduranus platyops Broom.
C. First three molar teeth of same, enlarged.

The preparietal is remarkable in being situated quite in front
of the pineal foramen. It islozenge-shaped, the two posterior
sides articulating with the parietals and the anterior with the
frontals.

The postorbitals are large.

The occiput, so far as can be seen, presents no unusual feature.

The following are the principal measurements :—

Greatest length............. eae about 90 mm.
Greatest Widhhtee. "aqme hac Be HDS Nt
Intévorbital width: ee ae Gis once
Intertemporal width ..................... ZO
Between the caniniform processes ...... PS eget

Front of snout to frontof pineal foramen 62

672 _ MR. R. BROOM ON ANOMODONT REPTILES:

This little Anomodont, to which I have given the name
Einyduranus platyops, shows some affinities with Cryptocynodon
simus Seeley, and also with Prodicynodon pearstunensis Broom,
but must be regarded as the representative of a very distinct
genus.

ENDOTHIODON CRASSUS, Sp. n. (Text-fig. 45.)

This new species is founded on a beautiful skull discovered by
me at Dunedin, Beaufort West district. The skull has lost the
lower jaw, and has the parietal region badly weathered and the
squamosal portion of the left temporal arch unperfect, but other-
wise it is almost complete. With the exception of Hndothiodon
whaitsi it is the largest Hndothiodon skull known, and it is little
inferior to that other, though very different-in shape.

Unlike Hndothiodon uniseries Owen and Endothiodon whaitsi
Broom, it is very broad and flat, and the parietal region is only
slightly elevated above the frontal plane. In general shape and
proportions the species which it most resembles is Hndothiodon
paucidens Broom, but that it is very different from this species
will be at once evident on comparing the figures.

The premaxilla when viewed from the front is like a thick
inverted V, the deep notch being for the point of the mandible,
and the two points are the caniniform processes, which here are
formed by the premaxilla and not, as in the tuskless Dicynodonts,
by the maxille. A small knob is situated in front of each
nostril. The superior median process passes between the nasals,
but is short.

The nasals are very broad, and the bones, like all the bones of
the top of the skull, very thick. They are as broad as long, and
the whole preorbital portion of the skull is thus much broader
than long—a condition which differentiates this species of Hndo-
thiodon from ail others.

The prefrontals are of moderate size, and so thickened that
they may be looked upon as forming preorbital bosses.

As in all typical Endothiodons, there are two longitudinal
grooves passing from the front of the frontals down the middle
of the nasals to the upper corners of the nostrils.

The septomaxilla seems to be small and within the nostril,
but the evidence is not satisfactory as to whether it also appears
on the face.

The lacrimal is smaller than the prefrontal, but its limits
cannot be clearly made out.

The maxilla is a large bone. As in all species of Hndothiodon,
it has a long ridge which forms an alveolar border and at a
considerable distance further in a row of teeth. Most of the
teeth appear to be lost, but when complete there was probably a
single row of eight teeth which measured 74 mm. The first
tooth of the series is round in section and has a diameter
of 7 mm.; the last, also round, has a diameter of 85 mm. The
maxilla forms most of the lateral part of the face, and it sends

FROM THE KARROO BEDS OF SOUTH AFRICA. 673

a long and powerful process outwards and slightly backwards
below the jugal to strengthen the jugal arch.

The jugal is, as in other Endothiodonts, more powerful than
in the Dicynodonts, and has a large rounded descending knob.
Tt sends a long process backwards inside the squamosal.

Text-figure 45.

Skull of Hndothiodon crassus Broom. Greatly reduced.

The quadrato-jugal and quadrate (QJ and Qu) of the right side are displaced :
those of the left side lost.

The frontals are large, but the exact limits of the bones cannot
be made out except in front as most of the orbital margins have
been lost. .

The preparietal is large and forms the front of the pineal
foramen wall.

The parietals are narrow but deep.

The squamosals are powerful. The zygomatic portion is more
slender than in Dicynodonts, but the occipital portion is greatly
developed.

674 ON ANOMODONT REPTILES FROM SOUTH AFRICA.

The quadrate and quadrato-jugal are lost from the left side,
but preserved in a slightly displaced condition on the right. As
in Dicynodon, they are apparently ankylosed, and their displace-
ment from the squamosal and paroccipital adds further evidence
to observations I have made in Dicynodon, that while the
quadrate is fixed to the quadrato-jugal, the quadrato-jugal has a
very considerable degree of movement on the squamosal, and
thus relatively to the skull the quadrate is perhaps as freely
movable as in the Lizard, in Anomodonts, and also in
Cynodonts.

The palate is similar to that in other Endothiodonts, and does
not differ greatly from the Dicynodon type. The periotic
processes do not descend to the same degree as in Dicynodon, but
are clasped by the basisphenoids in a similar manner, and have
each a large foramen ovale for the end of the stapes. Hndothiodon
differs from Dicynodon in having a large median basisphenoid boss.

The occipital condyle is flat as in other Hndothiodon species.

The following are the principal measurements of the skull :—

Greatest antero-posterior length...... 502 mm.
Greatest breadth across squamosals.... 400. ,,
Greatest breadth across jugals........ 430 ,,
Breadthacrossmasalay Woe os et 180,
Breadth between nostrils............ 60

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ON THE CICHLID FISHES: OF LAKE NYASSA. 675

36. The Cichlid Fishes of Lake Nyassa. By C. Tarr Recay,
M.A., F.R.S., F.Z.8., Keeper of Zoology, British
Museum (Natural History).

[Received June 7, 1921: Read June 7, 1921. |

(Plates I.-VI.* ; Text-figures 1-30.)

The Fishes of Nyassa have been somewhat neglected in com-
parison with those of Tanganyika and Victoria. hey were first
described by Giinther (P. Z. 8. 1864) from specimens collected
by Sw John Kirk, and nearly thirty years had passed before
the same author again described fishes from the lake sent by
Sir Harry Johnston (P.Z.S. 1893). Later a collection made
by Captain E. L. Rhoades was described by Boulenger (Ann. &
Mag. N. H. (8) 11. 1908). In Boulenger’s ‘ Catalogue of African
Fresh-water Fishes,’ 111. (1915), 38 species of Cichlide are recorded
from Nyassa; one of these, Petrochromis nyasse, may now be
removed from the list. The supposed occurrence of the specialized
Tanganyika genus -Petrochromis in Nyassa was dificult to
explain; re-examination of the type of P. nyassew leads me to
regard it as identical with the more recently described
P. fasciolatus, and I have no doubt that the locality assigned to
it was an error on the part of the collector. The loss of this
species from the Nyassa list is made good by the re-establishment
of Ginther’s Chromis subocularis, placed by Boulenger in the
synonymy of C. johnstoni, so that the number of valid species of
Cichlid hitherto described from Nyassa is 38.

The present revision is based on an examination of the speci-
mens in the collection of the British Museum (Natural History),
including the types of all the species described by Giinther and
by Boulenger, but principally on the study of a very fine collection
made and presented to the Museum by Mr. Rodney C. Wood.
As a result, the number of species is more than doubled, 46 being
described below as new to science. Of the 84 species all but 5
(3 Tilapia, 1 Astatotilapia, 1 Serranochromis) are endemic, and
the proportion of endemic genera is high, 11 out of 15, but more
than half the species belong to the widely distributed genus
Haplochromis. ‘The majority of the Nyassa genera are quite

distinct from any found elsewhere : for example, Lhanyphochromis,
which may be supposed to occupy the same place in Nyassa that
Bathybates does in Tanganyika, is very different from Bathybates.
There are, however, a few remarkable examples of convergent
evolution in Nyassa and Tanganyikat. The fish described below
as Fseudotropheus tropheops bears a great superficial resemblance
to Zropheus, and has the same peculiar dentition ; another new
type, dulonocara, has deep channels with large openings in the
frontal, nasal, orbital, preopercular, and mandibulary bones,
exactly as in Vrematocara.

* For explanation of the Plates see p. 727.
+ For the Tanganyika genera see Regan, Ann. & Mag. N. H. (9) v. 1920, p. 33.

676 ; MR. C, TATE REGAN ON THE

Synopsis of the Genera.

T. Scales cywloid or very finely denticulate.
A. Pharyngeal apophysis formed by parasphenoid on

Teeth in narrow or moderately broad bands, outer bicuspid,

inner tricuspid . ‘sate yee Male tathl lecaboles eatets ED ANee LLC ROTC Cl
Teeth very small, in very “broad bands ...:.... i.e. 2. Corematodus.
Teeth in a few series, slender, with expanded crowns,

the outer obliquely tr uncated . EAE RN ee heat aoe eLbemaunnlapea.

B. Pharyngeal apophysis eet by Deo in middle and prootics at

sides.

Teeth moderate, outer mostly bicuspid, inner tricuspid....., 4. Otophargne.
Meet very Shouts NICU Spee es peck deter ssehie-e seek) see seenenan DEmCe LON OLN aunoa Gl.

II. Scales usually distinctly denticulate. Pharyngeal apophysis formed by para-
sphenoid in middle and hasioccipital at sides.
A. Bones of head with small mucitferous canals with small pores.
1. Premaxillaries without anterior beak-like expansion.
a. ae of membrane of spinous dorsal produced into Jappets.
. Teeth conical, or outer bicuspid and inner tricuspid.

Teeth of iste series in upper jaw first decreasing and then

increasing in size antero-posteriorly ; band of teeth in

lower jaw crescentic ...... 6. Astatotilapia,
Upper jaw with a rather broad ‘band of small ‘cuspidate

teeth anteriorly and a series of enlarged conical teeth

on each side; lower jaw with a band of teeth with

incurved lateral edges, from which a single series runs

back on each side ...... sistestersye, @ Eseudotropheus.
Teeth conical ; lower jaw w ith 4 anterior canines ............ 8. Cynotilapia.
Teeth conical; no canines; 7 to 9 series of scales on cheek. 9. Serranochromis.
Outer teeth decreasing in size posteriorly; no distinct

canines; bands of teeth aa 2 to 6 series of

scales on cheek ...... : 10. Haplochromis:
Teeth very small and slender, forn ming narrow v bands which
are interrupted at the sv mphyses. Para une Ae A eis erh: mops.
g. Outer teeth very broad, compressed, with or without a pair of small
lateral cusps ....... .... 12. Docimodus.
b. Edge of membrane of spinous dorsal 1 running evenly between the tips of
the spines ..... Mauna. \Cyntocanas

. Praemaxillaries with an anterior beak-like expansion.
14, Rhamphochromis.

B. Frontals, nasals, orbitals, preopercular and mandibulary bones deepl
: a v, ply
excavated, with large openings ..........-....0eesee ee 15. Aulonocara.

1. Trnapra A. Smith, 1840
(type 7’. sparmannt A. Smith).

Teeth small or moderate, in several series, the outer bicuspid,
the inner tricuspid (sometimes conical in large fish), Seales
eycloid or feebly denticulate. Pharyngeal apophysis formed by
pavasphenoid only.

Africa; numerous species.

The four species found in Nyassa may be distinguished
follows :—

I. Three anal spines.
A. Caudal scaled only at the base.

Maxillary extending to below eye .............c0ceeccc eee eeseee eee s eee 1. mossambica.
Maxillary not extending to below eye ..............ceceeeeeseesses. 2. melanopleura.
B. Caudal covered with small scales ............ 6.ce i.e. 8. syuamipinnis.

JT. Hourjanallspines: feiss acdc a WANN as era asta ee Ne ERG TEC ROL ce

CICHLID FISHES OF LAKE NYASSA. 677

1, TILAPIA MOSsAMBICA Peters, 1852.

Bouleng. Cat. Afr. Fish, iii. p, 154, fig. 101.
Kast Africa,

2. ‘TILAPIA MELANOPLEURA A Dum., 1859.

Bouleng. Cat. Afr. Fish. iii. p. 190, fig. 123.
West Africa, Congo, and Zambesi.

3. TILAPIA SQUAMIPINNIS Gtnth., 1864.

Bouleng, Cat. Afr. Fish. iii. p. 183, fig. 118.
Nyassa.
4, TILAPIA SHIRANA Bouleng., 1896.

Bouleng. Cat. Afr. Fish, iii. p. 151, fig. 98.
Nyassa and Portuguese H. Africa.

2. Corematopus Bouleng., 1896.

Jaws with very broad bands of very small teeth, the outer
with expanded, compressed, and obliquely truncated crowns, the
inner mostiy pointed, Scales cycloid.,

A single species from Nyassa,

CoREMATODUS SHIRANUS Bouleng., 1896.

Corematodus shiranus Bouleng. Cat. Afr. Fish. iii, p. 494,
fig. 342.

Depth of body 22 in length, length of head 3. Snout with
convex profile, shorter than postorbital part of head. Diameter
of eye equal to preeorbital depth, 5 in length of head: interorbital
width 23. Jaws equal anteriorly; maxillary extending to below
eye. 4 series of scales on cheek. 12 gill-rakers on lower part of
anterior arch. Pharyngeal teeth small. 34 scales in a longi-
tudinal series. Dorsal XVI 11; last spine 2 length of head.
Anal IIIT 9: third spine 4 length of head. Caudal scaly. Caudal
peduncle 13 as long as deep. Body with seven narrow blackish
cross-bars as in Tilapia squamipinnis, the first downwards from
origin of dorsal, the last two on caudal peduncle.

The type, 200 mm. long.

3. Hemirinaria Bouleng., 1902.

Teeth in 3 to 6 series, small, with slender shaft and compressed
and expanded crown, those of outer series obliquely truncated.
Scales feebly denticulate.

A single species from Nyassa.

HEMITILAPIA OXYRHYNCHUS Bouleng., 1902.

Henatilapia ovyrhynchus Bouleng. Cat. Afr. Fish, iii. p. 489,
fig. 339.

678 MR. C. TATE REGAN ON THE

Depth of body 22 to 23 in the length, length of head 3 to 33.
Snout with straight or concave profile, from as long as to
13 diameter of eye, which is 33 to 53 in length of head, in adult
equal to preorbital depth; interorbital width 33 to 4 in head.
Jaws equal; maxillary not extending to below eye. 3 or 4 series
of scales on cheek. 12 or 13 gill-rakers on lower part of anterior
arch. Pharyngeal teeth small. 33 to 35 scales in a longitudinal
series, 5 or 6 from origin of dorsal to lateral line. Dorsal XVI
10-11; last spine from less than #2 to 3 length of head.
Anal ITI 9-10; third spine from less than + to more than # head.
Pectoral as long as or a little shorter than head. Caudal scaly,
truncate or slightly emarginate. Caudal peduncle longer than
deep. A dark spot on operculum, usually two on lateral line
below spinous and soft dorsal respectively and another at base of
caudal. Dorsal and caudal spotted with orange ; anal with long
spots between the rays; males with dorsal, anal, and pelvic fins
darker, pale-edged.

Seven specimens, 90 to 190mm. long. (Voore, khoades, Wood).

4. OvopHaRYNX Regan, 1920
(type Tilapia auromarginata Bouleng.).

As Tilapia, but the prootic forms part of the facet for articu-
lation of the upper pharyngeal on each side.
Nyassa ; two species.

1. OroPpHARYNX AUROMARGINATUS Bouleng., 1908.

Tilapia auromarginata Bouleng. Cat. Afr. Fish. 11. p. 180,
fig. 115. e

Depth of body 23 to 2% in length, length of head 34 to 33.
Snout decurved, as long as or a little shorter than postorbital
part of head. Diameter of eye equal to depth of preorbital,
32 to 4 in length of head; interorbital width 3 to 33. Jaws
equal anteriorly ; maxillary not extending to below eye; teeth in
3 to 5 series, inner tricuspid or conical; 60 to 75 in outer series
of upper jaw, the anterior bicuspid, the posterior conical.
4 series of scales on cheek. 15 to 18 gill-rakers on lower part of
anterior arch. Pharyngeal teeth small. Scales cycloid, 33 or 34
in a longitudinal series, 5 or 6 from origin of dorsal to lateral
line. Dorsal XVII-XVIII 10-11; last spine 2 length of head
or a little more. Anal IJI 9-10; third spine stronger than
dorsals, } head or a little more. Pectoral as long as head, not
reaching anal. Caudal densely scaled, emarginate. Caudal
peduncle 1} as long as deep. Bluish; dorsal and anal dark
blue with yellow edge; dorsal with series of red spots.

Two specimens, 205 and 240 mm. in length, and a skeleton.

CICHLID FISHES OF LAKE NYASSA. 679

2. OTOPHARYNX SELENURUS, sp.n. (Text-fig. 1.)

Depth of body 24 in length, length of head 33. Snout with
straight profile, as long as postorbital part of head. Diameter of
eye nearly equal to depth of preorbital or cheek, 4 to 44 in
length of head, interorbital width 4, Jaws equal anteriorly;
maxillary not extending to below eye; teeth in 3 or 4
series, 50 to 55 bicuspid teeth in outer series of upper jaw.
3 or 4 series of scales on cheek. 12 gill-rakers on lower part of
anterior arch. 35 scales in a longitudinal series, 6 from origin
oH dorsal to iateral line. Dorsal XVI-XVIT 11-12; last spine

length of head, longest soft rays nearly as long as head.
Ian IIL 9; third spine stronger and shorter than last dorsal.
Pectoral as long as head, reaching anal. Caudal sealy, deeply
emarginate. Caudal peduncle 13 as long as deep. Bluish grey,
uniform or with traces of darker cross-bars.

Text-figure 1.

tol

Otopharynx selenurus.

Two specimens, 135 and 155 mm. in total length (Wood).

A smaller example, 90mm. long, is more slender (depth 3 1n
length), but, except for juvenile characters, such as the larger eye
(33 in head), i is very similar to the two deseribed ; ; colour silvery,
ith traces of several cross-bars; an oblong dark spot on lateral
line below middle of spinous dorsal, a band along lower lateral

line.
5. CHILOTILAPIA Bouleng., 1908.

Jaws with an outer series of stout, little compressed, teeth,
and 3 or 4 imner series of smaller subconical teeth, which are
stronger at the sides of the premaxillaries than in front.
Scales cycloid or feebly denticulate.

Nyassa ; a single species,

Proc, Zoou. Soc.—1921, No. XLVI. 46

680 MR. C. TATE REGAN ON THE

CHILOTILAPIA RHOADESII Bouleng., 1908.

Chilotilapia, rhoadesti Bouleng. Cat. Afr. Fish. ii. p. 499,
fig. 366.

Depth of body 21 to 23 1n the length, length of head 33. Snout
short, declivous ; onsen of eye 4 to 42 in length of head,
preorbital depth 4, interorbital width 3. ie aws equal anteriorly ;
mouth wide, with short lateral cleft. 3 or 4 series of scales on
cheek. 11 eill- rakers on lower part of anterior arch. Middle
pharyngeal “teeth moderately strong, subconical. 32 to 34 scales
in a longitudinal series. Dorsal XV—XVI 10; last spine 2 to 4
length of head. Anal III 9-10. Pectoral a little longer. than
head, reaching anal. Caudal densely scaled, emarginate.
Caudal peduncle a little longer than deep. Silvery or bluish;
an opercular spot ; sometimes a broad dark band on each side of
back and another on middle of side; soft dorsal with series
of spots.

The type (220mm.) and two specimens of 180 and 200mm.
(Wood).

6. AsraToTILAPIA Pellegr., 1904
(type Labrus desfontainesti Lacep.).

Near Haplochromis, but posterior teeth of outer series of upper
jaw increasing in size backwards. Teeth in 3 to 5 series,
cuspidate or conical, those of outer series of upper jaw sometimes
bicuspid anteriorly, conical posteriorly ; band of teeth in lower
jaw crescentic. Middle teeth of lower pharyngeal somewhat
‘enlarged. Dorsal XIII-XVIT 8-11. Anal III 7-11. Scales
26 to 36.

Africa.

This genus includes three species placed by Boulerger in 7tlapia,
namely, 7. sywnnertoni, 7’. burton, and 7. calliptera, and four
included by him in Haplochromis, H. strigigena, H. moffati,
H. desfontainesti, and H. moerwensis. These are all closely
related, agreeing in the form and size of the mouth, the rather
short pectorals, comparatively short caudal peduncle (as long as
deep), and the rounded caudal.

ASTATOTILAPIA CALLIPTERA Giinth., 1893.

Tilapia calliptera Bouleng. Cat. Afr. Fish. iii. p. 222, fig. 145.

Depth of body 23 to 24 in length, length of head 23 to 31.
Snout with straight pr ofile, shorter aha postorbital part a head.
Diameter of eye 2 to 43 in length of head, in adult seauge ly
greater than depth of preerbital; interor bital width 33 to 43 in
length of head. Jaws equal anteriorly ; maxillary extending to
below anterior edge of eye; teeth in 3 to 5 series, 40 to a0 fe
outer series of upper jaw, the anterior bicuspid, the last 2 or 3
(young) or 8 or 9 (adult), conical, increasing in size posteriorly.
3 to 5 series of scales on cheek. 8 to 10 gill-rakers on lower part

CICHLID FISHES OF LAKE NYASSA. 68]

of anterior arch. Middle pharyngeal teeth rather strong, conical
in adult. 29 to 33 scales in a longitudinal series, 5 or 6 from
origin of dorsal to lateral line. Dorsal XIV—X VI 8-11; last
spine 2 to 4 length of head. Anal III 7-9; third spine stronger
and usually shorter than last dorsal. Pectoral shorter than head,
not reaching anal. Caudal rounded. Caudal peduncle as long
as deep. An opercular spot and a dark bar from eye to end of
maxillary; body with or without dark cross-bavs and a dark
lateral band; dorsal and caudal sometimes spotted; males with
lower fins blackish, the anal with 2 to 6 rounded orange spots.

Nyassa and Zambezi.

Numerous specimens, 65 to 140 mm. in total length.

7. PsEUDOTROPHEUS, gen. n.
(type Chromis willtamst Giinth.).

Jaws with several series of teeth anteriorly, the outer bicuspid,
the inner small and tricuspid, forming rather broad curved or
transverse bands; upper jaw with a series of conical teeth on
each side posteriorly, more or less sharply differentiated, some
or all larger than the last bicuspid teeth; lower jaw short and
broad, with the lateral margin of the band of teeth incurved,
and with a series of teeth on each side behind the band.
Dorsal X VI-XTX 8-10. Anal III 7-9. Scales denticulate.

Nyassa; five species.

Text-figure 2.

a. Dentition of Pseudotropheus tropheops.
ib: tk Tropheus moorii.

Synopsis of Species.
‘I. Jaws equal anteriorly.

D. XVI-XVII 9. A. III 8. 4 series of scales on cheek........, 1. williamsi.
D. XVIIL9. A.I11 9, 5 or 6 series of scales on cheek.... ... 2. zebra.
Il. Lower jaw shorter than upper; mouth rounded.
D. XVIII 9. A. IL 8. Depth 3im length ........................ 3, novemfasciatus.
DaX Desa As LIL 78" Wepthess maene thems iene | Ln auracus.
III. Mouth subterminal, transverse ; snout very convex.
JD) DRONA RIC TDs en NED GAS ier SRE oo re eo Meee eee uae elas aR 9 éropheops.
46*

682 MR. C, TATE REGAN ON THE

1. PspuporRoPHEUS WILLIAMss Giinth., 1893.

Chromis williamsi Ginth. P. Z.S8. 1893, p. 624, pl. lvi. fig. C.

Tilapia livingstonit Bouleng. P.Z.8. 1899, p. 134, pl. x. fig. 2;
Cat. Afr. Fish. 11. p. 243, fig. 162.

Tilapia williamsi Bouleng. Cat. Afr. Fish. ii. p. 225, fig. 147.

Depth of body equal to length of head, 3 in length of fish.
Upper profile of head convex ; snout as long ag or a little longer
than diameter of eye, which is 34 to 4 in length of head, nearly
equal to or greater than the interorbital width, depth oe cheek,
or pr sorivuanl: Mouth terminal, rounded ; maxillary extending to
below anterior edge of eye; teeth in 5 or 6 series, inner small,
tricuspid; 40 to 52 teeth in outer series of upper jaw, the last
3 or 4 (young) or 6 to 9 (adult) on each side conical, enlarged
and sharply differentiated from the rest, which are bicuspid.
4 series of scales on cheek. 9 or 10 gill-rakers on lower part
of anterior arch. Pharyngeal teeth small, slender. 53 scales
in a longitudinal series, 6 from origin of dorsal to Lewera! line.
Dorsal XVI-XVII 9; last spine a little less than 5 length of
head. suo BEE sey ‘third spine a little shorter than dorsal.
Pectoral ? length of neo not reaching anal. Caudal truncate 2,
densely scaled in basal half. Caudal “peduncle as long as or a
little longer than deep. Body with 6 dark cross-bars, or bars
represented by a series of spots above lateral line and another on
middle of side; an opercular spot and a dark bar or spot on base
of caudal; two round white (? orange) spots on anal fin (pro-
bably absent in @).

Two specimens, types of the species and of 7. livingstoni, 105
and 75 mm. long.

2. PsrUDOTROPHEUS ZEBRA Bouleng., 1899.

Tilapia zebra Bouleng. Cat. Afr. Fish. ii. p. 244, fig. 163 (1915).

Depth of body 24 in length, length of head 34. Snout de-
curved, a little iomec than dnamnetien noe eye, which is 4 in length
of head, a little gr eater than depth of preorbital, equal to depth
of cheek; interorbital width 3 in length of head. Mouth ter-
minal, rounded, rather wide; jaws equal anteriorly : maxillary
extending to vertical from anterior edge of eye; teeth in 4 or 5
series, 56 in outer series of upper jaw, the last 8 or 9 conical.
5 or 6 series of scales on cheek. 12 gill-rakers on lower part of
anterior arch. 31 scales in a longitudinal series, 8 from origin of
dorsal to lateral line. Dorsal X VIII 9; last spine nearly 3 length
of head. Anal III 9; third spine a little shorter than last dorsal.
Pectoral nearly as long as head, not reaching anal. Caudal
probably truncate, densely scaled at base. Candal peduncle a
little deeper than long. A dark bar between eyes, a second from
opercular spot to occiput ; 6 vertical bars on body, the first from
origin of dorsal to base of pectoral, the sixth from soft dorsal to
anal: ; three round pale (? orange) spots on anal in male.

The type, 105 mm. in total length.

CICHLID FISHES OF LAKE NYASSA. 685

3. PSEUDOTROPHEUS NOVEMFASCIATUS, Sp. Nl.

Depth of body 3 in length, length of head 33. Snout with
convex profile, as long as diameter of eye, which is 33 in length
of head, equal to interorbital width, greater than depth of
preorbital or cheek.. Mouth rounded, lower jaw shorter than
upper; maxiilary reaching vertical from anterior edge of eye;
teeth in 5 or 6 series, about 40 in outer series of upper jaw,
anteriorly bicuspid, the last 6 to 8 on each side conical, some
enlarged. 5 series of scales on cheek. 10 gill-rakers on lower
part of anterior arch. Pharyngeal teeth small. 32 scales in
a longitudinal series, 6 from origin of dorsal to lateral line.
Dorsal XVIII 9; last spine 4 leneth of head. Anal ITI 8; third
spine as long as and stronger ‘than last dorsal. Pectoral shorter
than head, not reaching anal. Caudal truncate. Caudal peduncle
as long as deep. Body crossed by 9 dark vertical bars, the first
6 corresponding to the 6 of P. zebra, the last 2 on caudal
peduncle; end of snout and lower jaw blackish; an opercular
spot and a spot on base of caudal; a faint dark band on lateral
line and another on middle of side; spinous dorsal with a dusky
intramarginal band; soft dorsal and caudal with series of spots.

A single specimen, 65 mm. in total length (Wood).

4, PSEUDOTROPHEUS AURATUS Bouleng., 1897.

Tilapia curate Bouleng. Cat. Afr. Fish. ii. p. 246, fig. 164.

Depth of body 3% in length, length of head 33. Snout with
convex profile, a little longer than diameter of eye, which is 4 in
length of head, a little greater than interorbital width, depth of
preorbital, or cheek. Mouth rounded ; lower jaw shorter than
upper; maxillary nearly reaching ver tical from anterior edge of
eye; edn in 5 or 6 series, 45 in outer series of upper jaw, the
last 4 or 5 conical, enlarged. 4 series of scales on cheek. 9 or
10 gill-rakers on lower part of anterior arch. 33 scales in a
longitudinal series. Dorsal X1X 8; last spine 2 length oH head.
Anal III 7; third spine as long as last dorsal. Pectoral 2 2 length
of head, not reaching anal. Tanda densely scaled, eons:
Caudal peduncle a little longer than deep. Lips blackish ; two
black bands between eyes; a black stripe from eye, ending in a
spot on base of caudal; a second near edge of back, a third on
aorsal fin.

The type, 75 mm. in total length.

5. PshUDOLTROPHEUS TROPHEOPS, sp. n. (Text-fig. 3.)

Depth of body 2% to 3 in length, length of head 33. Upper
profile of head very convex; snout as long as diameter of eye.
which is 34 in length of head, ‘slightly exceeds preeorbital depth and
equals depth of cheek ; interorbital region convex, its width 22 to
22 in length of head. Mouth abe minal, Lneteee ; Jaws with 8
series of small cuspidate teeth ; a series of 6 to 8 well-differentiated
conical teeth on each side of see ndleegn 4 series of scales on

684 MR. C. TATE REGAN ON TH

cheek. 9 gill-rakers on lower part of anterior arch. Pharyngeal
teeth small, slender. 33 scales in a longitudinal series, 6 from
fen of dorsal to lateral line. Dorsal XVII 10; last spine
1 length of head. Anal I! 8; third spine stronger and as long
as or a little shorter than last dorsal. Pectoral a little shorter
than head, not reaching anal. Caudal sealy, truncate. Caudal
peduncle as long as or a little longer than deep. Traces of dark
eross-bars on body bearing a series of darker spots above lateral
line and another on andes of side ; an opercular spot and a spot
on base of eaudal ; dorsal wid a blackish intramarginal band.
Two specimens, 116 and 122 mm. in total length (Wood).

Text-figure 3. ,

Pseudotropheus tropheops. +.

8. CYNOWILAPIA, gen. n.

Near Pseudotropheus, but teeth conical, in a few series, outer
large, in lower jaw forming distinct canines anteriorly.
Nyassa ; ; a single species.

CYNOTILAPIA AFRA Gtinth., 1893.

Paratilapia ee Bouleng. Cat. Afr. Fish. 111. p. 325, fig. 218.

Depth of body 2? to 3 in length, length of head 3 to 3}. Snout
decurved, as Heine as or a little longer than diameter of eye,
which is 3% to 32 in length of head, greater than preorbital
depth ; intevorbital width 34 in length of head. Jaws equal
anteriorly ; maxillary not quite reaching vertical. from anterior
edge of eye; teeth conical, trisevial, outer large, 28 to 32 in upper
jaw ; lower jaw with four strong anterior canines. 3 or 4 series
of scales in cheek. 14 or 15 gill. rakers on lower part of anterior
arch, 33 scales in a Taneuimdiungl series, 6 from origin of dorsal

CICHLID FISHES OF LAKE NYASSA. 685

to lateral line. Dorsal XVII 9; last spine 2 to 4 length of head.
Anal ITT 8; third spine stronger and as long as or a little shorter
than last ove Pectoral Shor ter than head, not reaching anal.
Caudal? Caudal peduncle as long as or longer than deep.

The types, 85 and 95 mm. in total length.

9. SERRANOCHROMIS Regan, 1920.

As Haplochromis, but third vertebra without inferior apophyses,
fourth with a very small pair. Mouth large; teeth conical ;
cheek deep, with 7 to 9 series of scales.

A single species.

SERRANOCHROMIS THUMBERGI Casteln., 1861.

Paratilapia thumbergui Bouleng. Cat. Afr. Fish. ui. p. 328,
fig. 220.

Nyassa and Zambezi; Katanga and L. Bangwelu; L. Ngami;
Angola.

10. Harptocuromis Hilgend., 1888
(type H. obliquidens Hilgend.).
Céenochromis Pfeff., 1893 (pectoralis).
Champsochromis Bouleng., 1915 (ceruleus).

An outer series of bicuspid or conical teeth, decreasing in size
posteriorly, and one or more inner series of smaller bicuspid), or

Text-figure 4.

oy)

By RA ” ei ie
Hit

Hoe

LH
wea tina sO at
ty

Mis

a. Lower pharyngeal of Haplochromis tetrastigma.
HOS aes, Bs H. placodon.

conical teeth. 2 to 6 series of scales on cheek. Scales usually
distinctly denticulate. Pharyngeal apophysis formed by para-
sphenoid in middle and basioccipital at sides. Third vertebra
with inferior apophyses, which meet below.

52 species of this genus occur in Nyassa, all of them endemic.
In the great number and diversity of species of Haplochromis

686 MR. C. TATE REGAN ON THE

Nyassa resembles Victoria and differs notably from Tanganyika.
In the Nyassa species the caudal fin is truncate or emarginate,
and appears to be always nearly completely covered with small
scales in the adult fish; this feature, the prevalence of a few
distinctive types of acloraion. and the absence of evident
relationship to species found elsewhere lead to the conclusion
that the Nyassa species are a natural group and may perhaps
have evolved in the lake from a single ancestral form. This
conciusion is fortified by the study of such skeletons as are
available.

The differences in the pharyngeal dentition between closely
related species are sometimes very striking; the best examples
of this are H. tetrastigma and A. alaannler, and H. similis and
H. kirkii; these cases make it evident that it is not desirable to
regard the development of large, round, blunt pharyngeal teeth
as warranting generic separation.

Synopsis of the Nyassa Species.

I. Snout not longer than postorbital part of head.
A. Pectoral fin with series of spots on the rays.

Lower jaw projecting .... wecesecseeee * L. Levengstonrs.
Jaws equal; depth of body 2 2 to 3in length . ceseteeeee 2. polystigma.
Jaws equal; depth 22 in length. titdesscettectseenseseenese | Oo NACULIMaNUS.

B. Pectoral fin entnapalane.
1. Preemaxillary pedicels not extending beyond anterior edge of orbits.
a. Body with a series of dark spots on back alternating with a series on or
above middle of side, or uniting with them to form irregular cross-
bars. Jaws equal; outer teeth mostly bicuspid.

Pharyngeal teeth small; spots very large . . 4, simulans.
Middle ‘pharyngeal teeth enlarged, blunt ; veaudal: peduncle

15 to 1? as long as deep .. . 5. subocularis.
Middle pharyngeal | teeth enlarged, plunt ; ae peduncle

a little longer than deep ........ beboonsne, Gh OPDBIGDND

b. Body with 6 dark cross- sai “Thome: jaw projecting; outer teeth
bicuspid.

Pharyngeal teeth small .... ... 7%. johnstoni.
Lower pharyngeal with a eroup of enlar reed plunt teeth ... 8 sexfasciatus.

ce. Body with a dark lateral stripe, slightly curved anteriorly, from head to
caudal tin. Outer teeth bicuspid.
o#. Lower jaw a little projecting; middle pharyngeal teeth stout and

lolunraaeeeeeee shbsachou . el BOR B00
8. Jaws equal; pharyngeal teath eral.
Caudal truncate or slightly emarginate .... ca LOS sunales:
Caudal rather deeply emarginate; 45 to 60 tecth in outer
series ..... Perce alulomeucceps:
Caudal rather ‘deeply emarginate ; “94. to 30 teeth in outer
SELICS ieee othe weeveeeee.. 12. maicrostoma.

d. Body le a iene “inal panel 3 on rea acridine forward as a series of
spots. ‘Teeth conical. Lower jaw projecting. 13. wrotenia.
e. Body with a straight dark band or stripe from head to caudal fin.
Teeth conical. Lower jaw projecting.
2. Mouth little oblique, below level of eye.

Caudal truncate or very slightly emarginate .................. 14. fuscoteniatus.
Caudal distinctly emarginate .......... lveeeedteciea)) bi holotenia.

3. Mouth very oblique, pao sone ¢ ona level with eye.
Depthyot body 2eito,2= invlens th eyes. weresee weed etesseeeet 16. strigatus.

Depth of body Ain dength 20.00... 6.0... ..scecececneerstenesecveeee) Le Gumidraus. ”

CICHLID FISHES OF LAKE NYASSA. 687

Jf. An opercular spot and a blackish spot on or under lateral line below
middle of spinous dorsal ; often a third spot below soft dorsal and a
fourth at base of caudal f.

. 8 to 13 gill-rakers on lower part of anterior arch (? Al. zntermedius).
* Outer teeth bicuspid; lower jaw not projecting.

Maxillary extending to below eye . f 18. awritus.
Maxillary not extending to below eye; . pharyngeal teeth

small . 19. tetrastigma.
Maxillary not “extending ‘to below. eye; pharyngeal teeth

large, rounded, blintey eke tae: secseueeeee 20. placodon.

** Outer teeth oouetel Jerse ‘ee puteetine (? A. intermedius).
Snout a little shorter than postorbital part of head; last

dorsal spine nearly + leneth of head . e 21. intermedius.
Snout as long as postorbital part of head ; last dorsal

spine 3, interorbital width + Jength Os head... Saree 22. modestus.
Snout as long as postorbital part of head ; last dorsal spine

3 to 2, interorbital width 2 or i length Olheadeaaeeeeee 23. woodi.
8. 17 to 28 gill- Pate on lower part of anterior arch.

17 to 21 gill-rakers ; 15 or 16 dorsal spines .................. 24. chrysonotus.
23 to 28 ill- vakers : 17 or 18 dorsal spines ....... wen. 25. quadrimaculatus.

g. A dark band from nape or origin of car cl to base of caudal (sometimes
faint or absent in large specimens).
a, Lower pharyngeal with a group of large, rounded, blunt teeth.

Jaws equal; caudal peduncle 14 to 1} as long as deep ...... 26. spherodon.
Jaws equal; caudal peduncle 13 as long as deep ............ 27. ericotenia.
Lower jaw projecting ........ ....... 28. lateristriga.
8. Lower phar eee daw a ihe 2 maddie series a little enlarged.
Eye 3 to 4 in head (in specimens of 70 to 110 mm.) ......... 29. plagiotenia.
Eye 3 to 33 in head (in specimens of 120 to 170 mm.) ... 30. melanotenia.

y. Pharyngeal teeth small; outer teeth of jaws forming a close-set sevies,
usually bicuspid im young, some or all conical in adult.
* Lower jaw a little shorter than upper... 31. guentheri.
** Jaws equal; 18 or 19 dorsal spines ...... 32. melanonotus.
*k* Jaws equal or lower a little projecting; 15 to 17 dorsal spines.
+ Maxillary extending to vertical from anterior edge of eye.

8 or 9 gill-vakers on lower part of anterior arch ............ 33. brevis.
12 eill-rakers ; 35 or 36 scales; caudal peduncle 15 as long

as deep ... Leese aaa M20LOLENU.
11 to 13 gill- rakers ; 37 to 39 ‘scales ; ‘caudal peduncle 13

WONZpasHlOnptasuel ee phar ass Meeks cae adianadin theese seas onli 35. lepturus.

++ Maxillary not extending to below eye.

Scales 35 to 37; last dorsal spine} to 3 head ............... 36. rhoadesii.
Scales 33 or 34; last dorsal spine quite 2 head ............... 37. atriteniatus.

0. Pharyngeal teeth slender; outer teeth of jaws conical, rather strong,
and spaced.
* Third anal spine as long as last dorsal; pelvic fins shorter than

ea deVeer hats thane ci euebanhea den cs tution wep 38. spilorhynchus.
**< Third anal spine shorter than last dorsal; pelvie fins as long as
head.
Maxillary not quite reaching vertical from anterior aie
of eye . ee. 39. longipes.
Maxillary not ‘nearly. reaching vertical: from: anteric ior edge
need risa : See On ccemulouss

. Coloration Five ery, AG tmes | Ww Hen ree dark ecross-bars.
o#. Maxillary extending to below anterior edge of eye.

Lower pharyngeal with a group of enlarged teeth with

spherical crowns ..... .... 41. macrochir.
Posterior teeth of 2 middle series of lower “pharynge al
somewhat pnlaveed SNA te Re EE eee ones WALA er gUTrOsSOmeds

tI imclude H. modestus herve, but its coloration is unknown; the type, as
preserved, is uniformly brownish.

688 MR. C. TATE REGAN ON THE

8. Maxillary not extending to below eye.
Lower jaw a little the shorter; 12 or 13 gill-rakers on lower

part of anterior arch.. 43. macrophthalmus.
Jaws equal anteriorly ; 11 ‘oill- yakers on “Tower part ‘of

anterior arch .. BbuemAGhe dnetncaBaresdacsabentonaal | CNN IEDKCOSCIRS,
Jaws equal; 15 or 16 gill- yakers. 0... vse. 45. tnornatus.
Lower jaw a little the ‘shorter ; 16 or 17 gill: rakers ......... 46. micrentodon.

2. Premaxillary pedicels extending to between middle of orbits.
47. eucinostomus.
II. Snout longer than postorbital part of head.
A. Lower jaw projecting ; maxillary not extending to below eye.

Teeth cuspidate; caudal emarginate ..................-0.0:.... 48. preorbitalis.

Teeth conical; caudal truncate ..:....0.slo cesses 49: compressiceps.
B. Jaws equal anteriorly ; ee not ae ae to below eye.

15 to 17 gill-rakers on lower part of anterior arch ......... 50. macrorhynchus.

19 or 20 gill-rakers on lower part of anterior arch ......... dl. rostratus.

C. Jaws equal anteriorly ; maxillary extending to below eye.

52. macrostoma.

1. HapPLocHRoMis Livinestonit Giinth., 1893.

Haplochromis livingstonti (part.) Bouleng. Cat. Afr. Fish. 111
p. 286, fig. 194.

Depth of body 22 ‘to 3 in length, length of head 2? to 3.
Snout with straight profile, 14 to 2 diameter of eye, which is 43
to 54.in length of head, interorbital width 33 to 4, depth of pree-
onbrinl 8 3s to 4. Lower jaw projecting; maxillary not extending
to below eye; teeth in 3 or 4 series, outer bicuspid and inner
tricuspid in young, all conical in adult, 50 to 60 in outer series of
upper jaw. 4 or 5 series of scales on cheek. 11 or 12 gill-rakers
on lower part of anterior arch. Pharyngeal teeth small. 33 to
39 scales in a longitudinal series, 6 or 7 Ou eee of dorsal to
lateral line. Dorsal XVI 10- ld last spine 3 to ? length of head.
Anal It 9-10; third spine 3 3 to 2 head. poe al a little shorter
than head, vege himne origin otf anal. Caudal truneate or slightly
emarginate. Caudal peduncle longer than Weep. Silvery (2)
or bluish grey (d ); a dark bar from eye to end of maxillary ;
large opercular spot; body marbled with blackish, the spots hee
bands constantly nearly as in the figure of the type; pectoral
with series of small spots on the rays; dorsal and anal, in male,
blackish with pale edge.

The type, 120 mm. long, and three specimens of 200 to 230 mm.
(Wood).

2. HAPLOCHROMIS POLYSTIGMA, sp.n. (PI. I.)

Haplochromis livingstonii (part.) Bouleng. Cat. Afr. Fish. 111.
p. 286.

Depth of body 2% to 3 in length, lene of head about 3.
Snout with straight progies 13 to 24 diameter of eye, which is 4
to 6 in lene of head, interorbital width 31 to 4, depth of pre-
orbital 37 2 to 43. Jaws equal anteriorly ; maxillary not extend-
ing to below eye; teeth in 3 or 4 series In upper jaw, 2 or 3 in
lower, outer bicuspid and inner tricuspid in young, all conical

CICHLID FISHES OF LAKE NYASSA. 689

in adult, 50 to 65 in outer series of upper jaw. 4 or 5 series of
scales on cheek. 10 to 13 gill-rakers on lower part of anterior
arch. Pharyngeal teeth small. 32 to 34 scales in a longitudinal
series, 6 from origin of dorsal to lateral line. Dorsal XVI 10-
une last spine + ‘to 2 2 length of head. Anal IIT 9-10; third
spine usually a little "shorter than last dorsal. Pectoral as long
as or a little shorter than head, nearly or quite reaching anal.
Caudal truneate or slightly emarginate. Caudal peduncle longer
than deep. Body with large irregular brown spots tending to
run together to form 3 longitudinal bands; head, body, ‘and
vertical fins covered with numerous small dark spots ; pectoral
with series of spots on the rays.

Six specimens (Wood), 120-210 mm. in total length; two
others (Rhoades, Cunningéon) also belong to this species.

3. HAPLOCHROMIS MACULIMANUS, Sp. D.

Paratilapia modesta (part.) Bouleng. Cat. Afr. Fish. 11. p. 326.

Depth of body 22 in length, length of head nearly 3. Snout
with straight profile, shorter than postorbital part of head.
Diameter of eye 5 in length of head, a little less than preorbital
depth, # depth of cheek; interorbital width 4 in head. Jaws
equal anteriorly; maxillary not extending to below eye ; teeth
conical, in 4 series, about 80 in outer series of upper jaw.
5 series of scales on cheek. 11 gill-rakers on lower part of
anterior arch. Pharyngeal teeth small. 33 scales in a longi-
tudinal series, 7 from origin of dorsal to lateral line. Dorsal XVI
Ti Amaliih: 10; seed spine 3 head, a little shorter than last
dorsal. Caudal peduncle a litéle longer than deep. Traces of
dark cross-bars; pectoral with several transverse series of small
dark spots.

A single specimen, 190 mm. long (J/oore).

4, HAPLOCHROMIS SIMULANS, Sp. n. (Text-fig. 5.)

Depth of body 2§ in the length, length of head 5 to 3l. Snout
with straight or eanree profile, 17 to ‘12 diameter of eye, which
is 33 to 42 in length of head, eatlsoer bital width 4, depth of pre-
orbital 4 to 43. Tee equal anteriorly ; : maxillary not extending
to below eye; teeth in 4 to 6 series, outer bicuspid, or posterior ly
conical in adults, 60 to 75 in outer series of upper Jaw. 3 or 4
series of scales on cheek. 10 gill-rakers on lower part of anterior
arch. .Pharyngeal teeth small. 32 to 34 scales in a longitudinal
series, 6 or 7 from origin of dorsal to lateral line. Dorsal XVI
105 duly last spine 3 to more than 2 length of head. Anal III 9;
third spine as long as last dorsal. ” Pectoral as long as or a little
shorter than head, nearly or quite reaching anal. Caudal tiun-
eate or slightly emarginate. Caudal peduncle as long as or a
little longer than deep. Silvery or yellowish ; a dark bar below
eye; an opercular spot: 4 large dark spots on back alternating

690 MR. C. TATE REGAN ON THE

with 4 very large, vertically expanded spots on side, the spots
sometimes confluent to form broad irregular cross-bauds ; dorsal
and anal dusky with yellow margin, or anal pale yellow; caudal
dusky above, yellow below.

Four specimens, 130 to 190 mm. in total length (Wood).

Text-fi gure 5.

Nie

Haplochromis simulans.

This species bears a great resemblance and is probably closely
related to Cyrtocara venusta, but differs in the structure of the
spinous dorsal fin.

5. HAPLOCHROMIS sUBOCULARIS Giinth., 1893.

IE An Sig Os MOPAILG Tolle hie. aie 18>.

Tilapia johnstoni (part.) Bouleng. Cat. Af. Fish. 111. p. 249.

Depth of body 3 to 32 in length, length of head 3} to 32.
Snout with straight or slightly convex profile, a little longer than
diameter of eye, which is 34 to 3? in length of head ; inter-
orbital width 32 to 4 in head, depth of preorbital 44. Jaws
equal anteriorly ; maxillary not extending to below eye; teeth m
4 or 5 series, outer bicuspid, 40 to 50 in outer series of upper
jaw. 3-series of scales on cheek. 10 gill-rakers on lower part of
anterior arch. Middle pharyngeal teeth enlarged, rounded, blunt.
32 or 33 scales in a longitudinal series, 5 or 6 from origin of
dorsal to lateral line. Dorsal XV 11-12; last spine 2 to 4 length
of head. Anal III 8; third spine 3 to 2? head. Pectoral as long
as head, nearly or quite reaching anal. Caudal truncate or slightly
emarginate. Caudal peduncle 13 to 12 as long as deep. Two
dark bars across nape; 4 dark spots at base of dorsal alternating
with a series above middle of side; 2 large spots on caudal
peduncle; dorsal and caudal with series of spots.

Two specimens, 120 and 140 mm. in total length.

CICHLID FISHES OF LAKE NYASSA. 691

6. HAPLOCHROMIS ORNATUS, sp. n. (Text-fig. 6.)

Depth of body equal to length of head, 2? in length of fish.
Snout with slightly convex profile, as long as postorbital part of
head. Diameter of eye 14 depth of preorbital, greater than
depth of cheek, 34 in length of head: interorbital width 4 in
length of head. Jaws equal anteriorly; lips thick; maxillary
not extending to below eye; teeth in 3 series in upper jaw, 4 in
the lower ; 52 bicuspid teeth in outer series of upper jaw. 3 series
of scales on cheek. 12 gill-rakers on lower part of anterior arch.
Middie pharyngeal teeth enlarged, rounded, blunt. 31 scales in
a longitudinal series, 6 from origin of dorsal to Jateral line.
Dorsal XVI 10; spines rather strong, last a little more than 2
length of head. Anal III 9; third spine stronger than dorsals,

'Text-figure 6.

Haplochromis ornatus. 3.
2 head. Pectoral nearly as long as head, reaching anal. Caudal
feebly emarginate. Caudal peduncle a little longer than deep.
Silvery; an opercular spot and a spot above eye; 5 dark spots at
base of dorsal alternating with a series above middle of side;
2 dark bars across upper 3 of caudal peduncle ; dorsal and caudal
with series of spots.

A single specimen, 140 mm. in total length (JVood).

7. HAPLOCHROMIS JOHNSTONI Giinth., 1893.

Tilapia johnstoni (part.) Bouleng. Cat. Afr, Fish. ui. p. 249,
fig. 167. ;

Depth of body 22 in length, length of head 3. Snout with
straight profile, as long as postorbital part of head. Diameter of
eye 4 in length of head, interorbital width 43, depth of pre-
orbital 44. Lower jaw projecting; maxillary not extending to
below eye; teeth in 3 or 4 series, outer bicuspid, 50 in outer
series of upper jaw, 3 series of scales on cheek. 11 gill-rakers

692, MR. C. TATE REGAN ON THE

on lower part of anterior arch. Pharyngeal teeth small, com-
pressed. 31 scales in a longitudinal series, 5 from origin of dorsal
to lateral line. Dorsal XVI 10; last spine a little less than 4
length of head. Anal III 9; third spine stronger than dorsals,
nearly 5 2 head. Pectoral + head, reaching origin of anal. Caudal
truncate ov shghtly emarginate, Caudal peduncle 13 as long as
deep. Silvery; a dark bar from eye to angle of mouth; 6 dark
bars on body, the first downwards from nape, the fifth from end
of dorsal to behind anal; series of spots on dorsal and caudal.
The type, 120 mm. in total length.

8. HaPLOCHROMIS SEXFASCIATUS, sp. n. (Text-fig. 7.)

Depth of body 2% to 24 in the length, length of head 3. Upper
profile of Bend a little concave ; 3 snout 1A to 2 diameter of eye,
which is 4 to 5 in length of Threrudh interorbital width : x to 43,
preorbital depth 4 to ree Lower jaw projecting ; eee not

Text-figure 7.

telo

Haplochromis sexfasciatus.

extending to below eye ; teeth in 3 to 5 series, 40 to 60 bicuspid
teeth in outer series of upper jaw. 3 or 4 series of scales on
cheek. 11 or 12 gill-rakers on lower part of anterior arch.
Lower pharyngeal with a group of moderately enlarged blunt
rounded teeth in the middle posteriorly. 32 scales in a longi-
tudinal series, 6 from origin of dorsal to lateral line. Dorsal XV
11; last spine 2 to nec wly 4 Length of head. Anal III 8— 95
third spine stronger than igh dor al, 4 3 to2 head. Pectoral +to %
length of head, reaching origin of jameld Caudal truncate or
shia ney emarginate. Caudal peduncle 14 as long as deep. Body
with 6 blackish cross-bars; dorsal and camel! with series of spots.
Two specimens, 100 and 170 mm. in total length (Wood).

Lae)

CICHLID FISHES OF LAKE NYASSA. 69

9. HAPLOCHROMIS KIRKIT Giinth., 1893.

Tilapia kirkii (oar ) Bouleng. Cat. Afr. Fish, iii. p. 251, fig. 169.

Depth of body 23 to 22 in length, length of head about 3.
Snout with straight ne 1; to if ‘Uaneie: of eye, which is 32
to 43 in length oF head, in adult equal to preorbital depth; hiner
arbital width 34 to 44 in length of head. Lower jaw a little
projecting ; maxillary extending to between nostril and eye;
teeth in 4 or 5 well- -separated series, inner tricuspid, outer
bicuspid ; 40 to 45 in outer series of upper jaw. 3 or 4 series of
seales on cheek. 11 or 12 gill-rakers on lower part of anterior
arch. Middle pharyngeal teeth stout and blunt. 31 to 33 scales
in a longitudinal series, 5 or 6 from origin oF dorsal to lateral
line. Dorsal XV 10- iM 3 last spine #2 to 3 length of head.
Anal [II 8-10; third spine as long as or a ae shorter than last
dorsal. Pector A as long as or a little shorter than head, reaching
origin of anal. Caudal truncate or shghtly emarginate. Caudal
peduncle 13 to 13 as long as deep. A dark stripe from operculum
to caudal. Usually a stripe or a series of spots above upper
lateral line and another at base of dorsal: ; dorsal and caudal with
series of spots; anal with ocelli in males.

Seven specimens, 100-160 mm. long (Johnston, Rendall, Wood).

Text-figure 8

Haplochromis similis. 2.

10. HaAPLocHRoMIS sIMILIS, sp.n. (Text-fig. 8.)

Depth of body 24 to 2! in length, length of head 3 to 3L.
Snout with straight or slightly convex profile, as long as or
longer than abana of eye, which is 3} to 32 in length of head,
oreater than preorbital depth ; “pte Meal Cait 31 to 3% in

694 MR. C. TATE REGAN ON THE

length of head. Jaws equal anteriorly ; maxillary ending between
nostril and eye; teeth in 4 to 6 series, outer bicuspid, inner
tricuspid, 40 to 52 in outer series of upper jaw. 3 or 4 series of
scales on cheek. 11 to 13 gill-rakers on lower part of anterior
arch. Pharyngeal teeth small. 32 or 33 scales in a longitudinal
series, 5 or 6 from origin of dorsal to lateral line. Dorsal XVI-—
XVII 9-10; last spine 2 to 4 length of head. Anal IIT 9-10;
third spine a little shorter than last dorsal. Pectoral as long as
or a little shorter than head, nearly or quite reaching anal.
Caudal truncate or slightly emarginate. Caudal peduncle 1z to
14 aslong as deep. Coloration as in H. kirku.
Five specimens, 120-150 mm. long (Wood).

11, HaPLoCcHROMIS BREVICEPS, sp.n. (Text-fig. 9.)

Depth of body 3 in the length, length of head 32. Snout a
little shorter or longer than diameter of eye, which is 3 to 33 in
length of head, greater than depth of preeorbitai or cheek; inter-
orbital width-4 in length of head. Jaws equal anteriorly;

maxillary not extending to below eye; teeth in 2 or 3 series;

Text-figure 9.

Haplochromis breviceps. 3.

45 to 60 bicuspid teeth in outer series of upper jaw; 2 series of
scales on cheek. 14 gill-rakers on lower part of anterior arch.
Pharyngeal teeth small. 33 to 35 scales in a longitudinal series,
5 or 6 from origin of dorsal to lateral line. Dorsal XVI-X VIL
11; last spine 2 length of head, Anal II] 9; third spine
stronger than dorsals, 3 or a little more than + head. Pectoral
as long as head, not reaching anal. Caudal emarginate. Caudal
peduncle 13 as long as deep. Silvery; a narrow dusky band
from opercular spot to base of caudal, another along upper lateral
line, a third at base of dorsal. Dorsal and caudal with series of
orange spots; male with ocelli in anal.
Two specimens, 85 and 135 mm. in total length (Wood).

CICHLID FISHES OF LAKE NYASSA. 695

12. HAPLocHROMIS MICROSTOMA, sp.n. (Text-fig. 10.)

Tilapia kirkit (part.) Bouleng. Cat. Afr. Fish. i. p. 251.

Depth of body 3 in length, length of head 34 to 33. Snout
straight, as long as or longer than diameter of eye, which is 33
to 4 in length of. head, a little or considerably greater than pre-
orbital depth ; interorbital width 4 in length of head. Mouth
small; jaws equal anteriorly; maxillary ending a little behind
nostril; teeth in 4 well-separated series, inner tricuspid, outer
bicuspid, 24 to 30 in outer series of upper jaw. 3 or 4 series of
seales on cheek. 11 gill-rakers on lower part of anterior arch.

Text-figure 10.

cS)

. Haplochromis microstoma.

Pharyngeal teeth small. 33 or 34 scales in a longitudinal series,
5 or 6 from origin of dorsal to lateral line. Dorsal XVIT 11;
last spine 2 to 4 length of head. Anal III 9; third spine a little
shorter than last dorsal. Pectoral as long as head, not reaching
anal. Caudal deeply emarginate. Caudal peduncle 13 as long as
deep. Silvery; a dark lateral band from operculum to base of
caudal ; two series of small dark spots, one at base of dorsal, the
other above lateral line; dorsal spotted.

Two specimens, 80 mm. (Wood) and 115 mm. (Rhoades) in
length.

13. HAPLOCHROMIS UROTHNIA, sp.n. (Text-fig. 11.)

Depth of body 23 to 3 in length, length of head 24to 3. Snout
with straight upper profile, in adult as long as postorbital part
of head. Diameter of eye 4 to 42 in length of head, equal to
or a little less than preorbital depth, less than depth of cheek ;
interorbital width 3? to 4in length of head. Lower jaw a little
projecting ; maxillary not quite reaching vertical from anterior
edge of eye; teeth conical in 2 or 3 series, 35 to 45 in outer

Proc. Zoou. Soc.—1921, No. XLVII. 47

696 MR. C. TATE REGAN ON THE

series of upper jaw. 3 or 4 series of scales on cheek. 11 or 12
gill-rakers on lower part of anterior arch. Pharyngeal teeth
slender. 33 scales in a longitudinal series, 5 or 6 from origin of
dorsal to lateral line. Dorsal XVI 9-10; last spine about 1
length of head. Anal IIT 8-9; third spine stronger and shorter

than last dorsal. Pectoral + head, nearly or quite reaching anal.

Text-figure 11.

Haplochromis wrotenia.

Caudal truncate or feebly emarginate. Caudal peduncle 1+ to
12 as long as deep. A series of about 8 dark spots near dorsal
profile, a second above lateral line, a third on middle of side
posteriorly confluent to form a band; dorsal with series of spots.
Three specimens, 170 to 200 mm. in total length (IVood).

14. HAPLOCHROMIS FUSCOTENIATUS, sp. n. (Text-fig. 12.)

Depth of body 22 in length, length of head 23. Head 23 as
long as broad. Snout a little concave in front of eye, thence
straight, nearly twice as long as diameter of eye, which is 5 in
length of head, interorbital width 5, preorbital depth 4. Lower
jaw a little projecting; maxillary not quite reaching vertical
from anterior edge of eye; teeth conical, in 3 series, 60 in
outer series of upper jaw. Cheek: with 4 series of scales, depth

% diameter of eye. 12 gill-rakers on lower part of anterior
arch. Pharyngeal teeth small. 34 scales in a longitudinal
series, 6 from origin of dorsal to lateral line. Dorsal XVI 11;
spines strongly increasing to last, which is 3 length of head.
Anal ITI 10; third spine stronger and a little shorter than last
dorsal. Pectoral + head, nearly reaching anal. Caudal truncate.
Caudal peduncle 1} as long as deep. Sides yellowish; a dusky
band on middle of side, a second backwards from base of pectoral ;
a series of dusky spots at base of dorsal, connected with a second

Col

CICHLID FISHES OF LAKE NYASSA. 697

’

series which form an interrupted band on and above lateral line.
Dorsal with series of spots. Anal with some pale (? red) round
spots.

A single specimen, 175 mm. in total length (IVood).

}
4
*
|

Text-figure 12.

Haplochromis fuscoteniatus.

wie

15. HaPLocCHROMIS HOLOTANTA, sp. n.

Paratilapia dimidiata (part.) Bouleng. Cat. Afr. Fish. iii.
p. 860, fig. 244.

Depth of body 33; in length, length of head 8. Snout de-
eurved, shorter than postorbital part of head. Diameter of eye
43 in length of head, equal to interorbital width or preorbital
| depth, less than depth of cheek. Lower jaw a little projecting ;
maxillary not extending to below eye; teeth mostly conical, in

4 series in upper jaw and 3 in lower, 60 in outer series of upper
jaw. 4 series of scales on cheek. 1 gill-rakers on lower part of
anterior arch. Pharyngeal teeth slender. 36 scales in a longi-
tudinal series, 5 from origin of dorsal to lateral line. Dorsal
ROVE 12); last spine = 2 head. Anal III 10; third Spine stronger
and nearly as long as last dorsal. Pectoral 2 > length of head.
Caudal emarginate. Cundal peduncle 13 as lene as deep. A
dark lateral band from eye to base of caudal fin; soft dorsal
with series of spots.
A single specimen (J/oore), 190 mm. in total length.

16. HAPLOCHROMIS STRIGATUS, Sp. n. (Text-fig. 13.)

Depth of body 22 to 2£in length, length of head 24 to 3. Snout
with straight or slightly « concave p1' ofile, as long as postor bital part
of head. Diameter of eye 4 to 4} in length of head, equal to
depth of preorbital or interorbital width, less than depth of cheek.

aT*

698 MR. C. TATE REGAN ON THE

Mouth oblique; lower jaw projecting; maxillary not extending
to below eye; teeth conical, triserial, 55 to 60 in outer series of
upper jaw. 3 or 4 series of seales on cheek. 12 or 13 gill-
rakers on lower part of anterior arch. Pharyngeal teeth small.
33 or 34 scales in a longitudinal series, 6 from origin of dorsal
to lateral line. Dorsal XV-XVI 11; last spine 2, longest soft
rays 2 to ? length of head. Anal III 10-11; third spine stronger
and shorter than last dorsal. Pectoral nearly as long as head,
reaching anal. Caudal truncate. Caudal peduncle 13 as long
as deep. Silvery; a blackish stripe, half as broad as a scale,
from operculum to base of caudal. Dorsal with series of orange
spots; other fins yellowish (@ ) or pelvics and anal blackish with
red margin (< ),
Three specimens, 170 to 185 mm. in total length (Wood).

Text-figure 13.

Haplochromis strigatus. ¥.

17. HapPLocHROMIS DIMIDIATUS Giinth., 1864.

Depth of body 4 in length, length of head 31. Snout with
straight upper profile, a little shorter than postorbital part of
head. Diameter of eye 5 in length of head, equal to preorbital
depth, slightly exceeding interorbital width. Mouth oblique;
lower jaw projecting; maxillary ending far in front of eye;
teeth conical, in 3 series in upper jaw, 2 in lower; 44 in
outer series of upper jaw; anterior teeth of second series in
upper jaw enlarged. 3 or 4 series of scales on cheek. 11 gill-
rakers on lower part of anterior arch. Pharyngeal teeth small.
35 scales in a longitudinal series, 5 from origin of dorsal to
lateral line. Dorsal XVI 11; last spine 3 length of head.
Anal III J1; third spine as long as and stronger than last

dorsal. Pectoral 2 length of head. Caudal slightly emarginate.

ese ee

CICHLID FISHES OF LAKE NYASSA. 699

Caudal peduncle twice as long as deep. Silvery; an opercular
spot; a blackish lateral stripe, half as broad as a scale, ending in
a spot on base of caudal fin.

A specimen of 165 mm. (Wood), described above, has been
compared with the type, a skin 220 mm. long.

18. Hapnociromis Auritus, sp.n. (Text-fig. 14.)

Depth of body 24 in length, length of head 3. Snout
convex, a little shorter than diameter of eye, which is twice
preorbital depth, 3 in length of head ; interorbital width 43
in head. Jaws equal anteriorly ; maxillary extending to below
eye; teeth in 2 series; 66 bicuspid teeth in outer series of upper
jaw. 3 series of scales on cheek. 13 gill-rakers on lower part of
anterior arch. Pharyngeal teeth small. 32 scales in a longi-

tudinal series, 5 from origin of dorsal to lateral line. Dorsal

Text-figure 14.

SS ee a <

Haplochromis auritus. Nat. size.

XV 10; last spine a little less than 3 length of head. Anal IIT 9;
third spine stronger than and nearly as long as last dorsal.
Pectoral as long as head, reaching anal. Caudal rather deeply
emarginate. Caudal peduncle a little longer than deep. Silvery ;
a blackish opercular spot; a dark spot on lateral line below
spinous dorsal; series of spots on soft dorsal and caudal.

A single specimen, 80 mm. in total length (Wood).

19. HAPLOCHROMIS TETRASTIGMA Ginth., 1893.

Tilapia tetrastigma Bouleng. Cat. Afr. Fish. 111, p, 250, fig. 168.

Depth of body 24 to 24 in length, length of head about 3.
Snout with straight or convex. profile, nearly as long as post-
orbital part of head. Diameter of eye 34 in length of head,
interorbital width 4, preorbital depth 4. Jaws equal anteriorly;
maxillary not extending to below eye; teeth in 3 or 4 series, outer

700 MR. C. TATE REGAN ON THE

bicuspid, about 60 in outer series of upper jaw. 3 or 4 series
of scales on cheek. 10 or 11 gill-rakers on lower part of anterior
arch. Lower pharyngeal a small weak plate bearing small
compressed bicuspid teeth. 30 to 32 scales in a longitudinal
series, 5 from origin of dorsal to lateral line. Dorsal XIV—
XV 10-11; last spine 2 to 4 length of head. Anal III 8-9;
third spine a little shorter than last dorsal. Pectoral as long
as or a little shorter than head, reaching origin of anal or
a little beyond. Caudal slightly emarginate. Caudal peduncle
13 to 1d as long as deep. Silvery; four blackish spots, the first
on operculum, the second on upper lateral line below spinous
dorsal, the third on side below soft dorsal, the fourth. at base of
caudal.
Three of the types, 105 to 110 mm. in total length.

20. HAPLOCHROMIS PLACODON, sp.n. (Text-fig. 15.)

Depth of body 23 to 25 in length, length of head about 3.
Snout with straight or convex profile, shorter than postorbital
part of head. Diameter of eye equal to or greater than depth

Text-figure 15.

lies

Haplochromis placodon.

of preorbital, 83 to 4} in length of head, interorbital width 32
to 33. Jaws equal anteriorly, lower 2 length of head; maxillary
not extending to below eye; teeth in 4 or 5 series, outer
bicuspid, 50 to 55 in outer series of upper jaw. 3 or 4 series of
seales on cheek, 8 or 9 gill-rakers on lower part of anterior
arch. Lower pharyngeal a large and strong plate bearing large
rounded flat teeth. 31 to 33 scales in a longitudinal series,
+ or 5 from origin of dorsal to lateral line. Dorsa! XVI 10-11;

——

CICHLID FISHES OF LAKE NYASSA. 701
last spine 2 to 2 length of head. Anal III 8-9; third spine }
to 2 head. Pectoral as long as head, extending nearly to middle
of anal. Caudal emarginate. Caudal peduncle 12 to 13 as long
as deep. Silvery; a blackish spot on operculum; two large
blackish spots on upper lateral line, expanding upwards towards
spinous and soft dorsal; a blackish spot at base of caudal.
Dorsal and caudal with series of orange spots; males with a
pale edge to dorsal and lower fins dusky, the anal with ocelli.

Five specimens, 130 to 180 mm. in total length (Wood).

21, HaPLocHROMIS INTERMEDIUS Ginth., 1864.

Paratilapia intermedia (part.) Bouleng. Cat. Afr. Fish. iu.
p- 363.

Depth of body 24 in length, length of head 3. Snout with
straight profile, shorter than postorbital part of head. Diameter
of orbit nearly equal to depth of preorbital, which is 43 in
length of head. Jaws equal anteriorly or lower perhaps slightly
projecting; lower jaw # length of head; premaxillary pedicels
not reaching orbits; maxillary not extending to below eye;
teeth small, conical, in 2 or 3 series, about 70 in outer series of
upper jaw. 4 series of scales on cheek. 34 scales in a longi-
tudinal series, 5 from origin of dorsal to lateral line. Dorsal
XVI 11; spines strongly Increasing to last, which is nearly 3
length of head. Anal III 9; last spine 4+ head. Pectoral as
long as head, reaching anal. Caudal rather deeply emarginate.
Caudal peduncle a little longer than deep. Silvery; traces of
two dark spots on upper lateral line; dorsal with series of spots
and a pale edge ; anal with several spots.

The type, a skin, 185 mm. long.
Boulenger’s figure (Cat. Afr. Fish. iii. fig. 247) represents the
body and fins correctly, but the head is inaccurate, the preorbital
being too narrow; also the four black spots depicted are taken

from a specimen of H. quadrimaculatus.

H. intermedius is well distinguished from H. quadrimaculatus
by the deeper preorbital, deeper cheek with 4 series of scales,
shorter premaxillary pedicels, the number of dorsal spines (16
instead of 17 or 18), and the form of the spinous dorsal fin. It
shows much closer agreement with H. placcdon, but has the
teeth in fewer series, smaller, and conical instead of cuspidate ;
also the lower jaw is notably longer.

22. HApLOcHROMIS MoDESTUS Giinth., 1893.

Paratilapia modesta (part.) Bouleng..Cat. Afr. Fish. iii. p. 326,
fig. 219.

Depth of body 3 in length, length of head 23. Snout with
straight profile, as long as postorbital part of head. Diameter
of eye 44 in length of head, a little greater than przorbital
depth, equal to depth of cheek; interorbital width 5 im length:

702 MR. C. TATE REGAN ON THE

of head. Lower jaw projecting; end of maxillary not far short
of vertical from anterior edge of eye; teeth conical, in 3
series, 70 in outer series of upper jaw. 4 series of scales on
cheek. 12 gill-rakers on lower part of anterior arch. Pharyn-
geal teeth slender. 32 scales in a longitudinal series, 5 from
origin of dorsal to laterai line. Dorsal XVI 11; last spine 3
length of head. Anal III 10; third spine stronger and slightly
shorter than last dorsal. Pectoral? Caudal ? (perhaps truncate).
Caudal peduncle as long as deep. Colour ?
A single specimen, 150 mm. in total length.

23. HapPLocHROMIS Woopt, sp.n. (PI. IT.)
Paratilapia modesta (part.) Bouleng. Cat. Afr. Fish. 1i1. p. 326.

Depth of body 23 to 3 in length, length of head 23 to 23.
Snout with straight profile, as long as postorbital part of head.
Diameter of eye 34 to 4 in length of head, equal to or greater
than depth of preorbital or cheek ; interorbital width 6 to 7 in
head. Lower jaw projecting ; maxillary not extending to below
eye; teeth conical, in 3 or 4 series, 55 to 70 in outer series of
upper Jaw. 3 or 4 series of scales on cheek. 11 or 12 gill-
rakers on lower part of anterior arch. Pharyngeal teeth slender.
32 or 33 scales in a longitudinal series, 5 or 6 from origin of
dorsal to lateral line. Dorsal XV-—XVI 9-10; last spine + to
2, longest soft rays 4 to # length of head. Anal III 9-10;
third spine stronger than and about as long as last dorsal.
Pectoral a little shorter than head, reaching anal. Caudal
truncate or slightly emarginate, angles sometimes rounded.
Caudal peduncle as long as or a little longer than deep.
Silvery; about 10 faint dark cross-bars; an opercular spot; a
series of small dark spots near dorsal profile; a large dark spot
on side below spinous dorsal, a second below soft dorsal, a third
at base of caudal. Dorsal and caudal with series of orange spots;
in males dorsal with blackish intramarginal band and orange
edge; pelvics and anal blackish, latter with orange spots.

Six specimens, 130 to 210 mm. long (Wood); two in poor
condition (fendall) also belong to this species.

24, HAPLOCHROMIS CHRYSONOTUS Bouleng., 1908.

Paratilapia chrysonota Bouleng. Cat. Afr. Fish. il. p. 362,
fig. 246.

Depth of body 23 to 3 in length, length of head 3 to 33.
Snout with straight profile, as long as or a little shorter than
diameter of eye, which is about twice depth of przeorbital, 22 to
3} in length of head, interorbital width about 34. Jaws equal
anteriorly ; premaxillary pedicels extending to between anterior
edges of orbits, } length of head; maxillary not extending to
below eye; teeth small, conical, in 3 or 4 series. 2 or 3 series
of scales on cheek. 18 to 21 gill-rakers on lower part of anterior

CICHLID FISHES OF LAKE NYASSA. 793

arch, Pharyngeal teeth small. 31 to 34 scales in a longitudinal
series, 5 or 6 from origin of dorsal to lateral line. Dorsal X V—
XVI 9-11; last spine 2 to 4 length of head. Anal III 9-11;
third spine about 2 head. Pectoral as long as head, reaching
anal. Caudal scaly, slightly emarginate. Caudal peduncle as
long as or a little longer than deep. Silvery with four blackish
spots. one on operculum, the second on and under lateral line
below spinous dorsal, the third below soft dorsal, the fourth at
base of caudal; males usually darker, often with yellow kack.
Several specimens, 100 to 130 mm. long.

2). HAPLOCHROMIS QUADRIMACULATUS, Sp. n.

Paratilapia intermedia (part.) Bouleng. Cat. Afr, Fish. 111.
p- 363.

Depth of body 21 to 2% in length, length of head 3 to 33.
Snout with straight profile, shorter ham postorbital part of head.
Diameter of eye 4 to 42% in length of head, depth of preeorbital
5 to 54, interorbital width 33. Jaws equal or lower slightly
projecting ; Peery pedicels extending to between anterior
edges of orbits, 2 2 length of head; maxillary not extending
to below eye; teeth sei conical, in 2 or 3 series. 2 or 3
series of scales on cheek. 24 to 27 gill-rakers on lower part
of anterior arch. Pharyngeal teeth small. 34 to 36 scales ina
longitudinal series, 5 or 6 from origin of dorsal to lateral line.
Dorsal XVII-X VITE 11-12; spines canes or slightly increasing
from seventh or gil, last 2 to 4 length of head. Anal IIT
10-12; third spine 1 to 2 head. Pectoral as long as or shorter
than head, not or barely reaching anal. Caudal scaly, emarginate.
Caudal peduncle a little longer than deep. A blackish spot on
operculum, a second under lateral line below spinous dorsal,
usually a third between lateral lines and a fourth at base of
caudal.

Seven specimens (Rhoades, Whyte), 170 to 200 mm. long.

26. HapLOCcHROMIS SPH@RODON, sp.n. (Text-fig. 16.)

953.

Tilapia later “istriga (part.) Bouleng. Cat. Afr. Fish. iti. p.
SFA!

Depth of body 25 to 3 in length, length of head 38 to 33.
Snout with straight profile, as long as or a little shorter than
diameter of eye, which is 3 to 34 in length of head, interorbital
width 4, preorbital depth 5. Jaws equal anteriorly; maxillary
not extending to below eye; teeth in 4 to 6 series, 40 to 60
bicuspid teeth in outer series of upper jaw. 2 to 4 series of
scales on cheek. 9 or 10 gill-rakers on lower part of anterior
arch. Middle pharyngeal teeth large, with spherical crowns.
31 or 32 scales in a longitudinal series, 5 from origin of dorsal
to lateral line. Dorsal XV-XVI 10-12; last spine about 2
length of head. Anal III 8-9; third spine usually a little
shorter than last dorsal. Pectoral about as long as head, reaching

_—

704 MR. C. TATE REGAN ON THE

anal. Caudal slightly emarginate. Caudal peduncle 1+ to 13
as long as deep. An opercular spot; a blackish band from
nape to caudal.

Five specimens, 80 to 125 mm. long (Wood, Rhoades), and a
skeleton. ‘

Text-figure 16.

Haplochromis spherodon.

ol
°

27. HAPLOCHROMIS ERICOTHENIA, sp.n. (Text-fig. 17.)

Depth of body 31 to 34 in length, length of head 3 to 33.
Snout with straight profile, nearly as long as or shorter than
diameter of eye, which is 2? to 3 in length of head, interorbital

Text-figure 17.

Haplochromis ericotenia. Nat. size.

width 41, preorbital depth 54 to 6. Jaws equal anteriorly ;
maxillary not extending to below eye; teeth in 4 series, 35
bicuspid teeth in outer series of upper jaw. 3 series of scales on
cheek. 10 gill-rakers on lower part of anterior arch. Middle
posterior pharyngeal teeth enlarged and obtuse, probably spherical

—_ ——

CICHLID FISHES OF LAKE NYASSA. 705

in adult. 31 scales in a longitudinal series, 7 from origin of
dorsal to lateral line. Dorsal XV-XVI11; last spine 2 or a
little more than 2 length of head. Anal IIT 9-10; third spine
as long as last dorsal. Pectoral + head, not reaching anal.
Caudal slightly emarginate. Caudal peduncle 13 as long as deep.
Silvery ; an opercular spot; 8 faint dark ecross-bars on body; a
blackish band, more or less broken up into a series of spots on
the bars, from nape to middle of base of caudal.
Two specimens, 65 and 72 mm. in total length (Wood).

28. HapLOCHROMIS LATERISTRIGA Giinth., 1864, (Text-fig. 18.)

Chromis lateristriga (part.) Giinth. P. Z.8. 1864, p. 312.
Tilapia lateristriga (part.) Bouleng. Cat. Afr. Fish. iii. p. 253.
Tilapia lethrinus (part.) Bouleng. t. c. p. 254.

Depth of body 2? to 3 in length, length of head 23. Snout
with straight upper profile, as long as postorbital part of head.
Diameter of eye 4 to 5 in length of head, interorbital width 34
to 43, depth of preorbital 33 to 4. Lower jaw a little projecting ;

Text-figure 18.

LUN WR)
ZO NINN:

Haplochromis lateristriga. ¥-

maxillary exposed distally, ending below nostril; teeth in 3 or 4
series, inner small; 40 to 55 bicuspid teeth in outer series of
upper jaw. Cheek with 3 to 5 series of scales. 153 gill-rakers
on lower part of anterior arch. <A group of large blunt teeth on
posterior part of lower pharyngeal. 34 scales in a longitudinal
series, 6 from origin of dorsal to lateral line. Dorsal XV-XVI
10-11; last spine 3 to 2 length of head. Anal III 9; third
spine stronger and a little shorter than Jast dorsal. Pectoral ?
or + head, nearly or quite reaching anal. Caudal densely scaled,
slightly emarginate. Caudal peduncle 14 as long as deep, A
blackish band from in front of dorsal fin to upper part of base of
caudal; an opercular spot; dorsal and caudal spotted.

_ Three specimens: the type, a skin 160 mm. long, a specimen
of 185 mm. (Wood), and another of 100 min. (Rhoades).

706 MR. C. TATE REGAN ON THE

29. HAPLOCHROMIS PLAGIOTENIA, sp.n. (Text-fig. 19.)

Tilapia lateristriga (part.) Bouleng. Cat. Afr. Fish. i. p. 253.

Depth of body 22 to 33 in length, length of head 3 to 3}.
Snout with straight or shghtly convex profile, as long as or a
little longer than diameter of eye, which is 3 to 4 in length of
head, greater than preorbital depth, equal to or greater than
depth of cheek ; interorbital width 3} to 32in head. Jaws equal
anteriorly ; maxillary not extending to below eye; teeth in 2 to
4 series; 35 to 50 bicuspid teeth im outer series of upper jaw.
3 or 4 series of scales on cheek. 8 to 11 gill-rakers on lower part
of anterior arch. Pharyngeal teeth small, compressed, those of
the series on each side of middle line a little enlarged. 31 to 33
scales in a longitudinal series, 5 or 6 from origin of dorsal to

lateral line. Dorsal XV-XVI 10- 11; Jast spine about 3 length

Text-figure 19.

Haplochromis plagiotenia. %.

of head. Anal III 8-9; third spine 2 to 4 head. Soft dorsal;
anal, and pelvic fins produced i in males. Pectoral as long as or a
little shorter than head, nearly or quite reaching anal. Caudal
slightly emarginate. Caudal peduncle longer than deep. A dark
band from nape to middle of base of caudal ; dorsal and caudal
usually with series of spots; males darker in colour, with dark
fins, the dorsal with a pale edge.

17 specimens, 70 to 110 mm. in total length (2hoades, Wood),
all but one with 15 dorsal spines.

30. HAPLOCHROMIS MELANOTENIA, sp.n. (Text-fig. 20.)

Depth of body 2§ to 2? in length, length of head 23 to 3.
Snout straight, declivous, weeny as long as or a little longer than
diameter of eye, which is 3 to 32 in length of head, equal to or
greater than interorbital width, greater than preorbital depth,

CICHLID FISHES OF LAKE NYASSA. 707

not less than depth of cheek. Jaws equal anteriorly 5 maxillary
extending to below anterior edge of eye; lips thick; teeth in
4 or 5 well-separated series, conical, or outer bicuspid and inner
tricuspid in young, 40 to 50 in outer series of upper Jaw. 3 or 4
series of scales on cheek. 9 or 10 gill-rakers on lower part of
anterior arch. Lower pharyngeal stout; middle teeth conical or
slightly compressed, moderately strong, set well apart, only 8 in
the series on each side of middle line. 33 scales in a longitudinal
series, 5 from origin of dorsal to lateral line. Dorsal XV-XVIT
10-11; last spine about 2 length of head. Anal Ill )9)s third

2)
9

spine stronger than dorsals, + to 2 head. Pectoral as long as

Text-figure 20.

Haplochromis melanotenia. +.

head, reaching anal. Caudal truncate or slightly emarginate.
Caudal peduncle 14 to 12 as long as deep An opercular spot; a
blackish band from nape to caudal; series of orange spots on
dorsal.

Three specimens, 120 to 170 mm. in total length (Wood).

31. HAPLOCHROMIS GUENTHERI, sp. n.

Chromis lateristriga (part.) Giinth. P. Z.8. 1864, p. 312.

Lilupia lateristriga (part.) Bouleng. Cat. Afr. Fish. iii. p. 253,
fie LO,

Paratilapia dimidiata (part.) Bouleng. t. ¢. p. 360.

Depth of body 23 to 3 in length, length of head 3} to 32.
Snout with straight or slightly convex protile, shorter than post-
orbital part of head. Diameter of eye 34 to 5 in length of head,
greater (young) or less (adult) than depth of preorbital, less than
depth of cheek ; interorbital width about 4 in length of head.
Lower jaw a little shorter than upper; maxillary not extending
to below eye; teeth in 3 or 4 series, outer bicuspid in young,
conical in adult, 40 to 70 in outer series of upper jaw; anterior
outer teeth of lower jaw directed outwards. 3 or 4 series of

708 MR. C. TATE REGAN ON THE

scales on cheek. 13 or 14 gill-rakers on lower part of anterior
arch. Pharyngeal teeth small, none enlarged. 33 to 35 scales
in a longitudinal series, 5 from origin of dorsal to lateral line.
Dorsal XVI-XVII 10-12; Jast spine 2 to 4 length of head.
Anal III 8-10; third spine stronger and shorter than last dorsal.
Pectoral as long as head, usually not reaching anal. Caudal
slightly emarginate. Caudal peduncle longer than deep. Silvery ;
an opercular spot; a blackish band from nape to caudal; usually
a series of dark spots at base of dorsal; series of orange spots on
dorsal fin.

Seven specimens of 130 to 190 mm. (Wood), the one figured by
Boulenger (Rhoades), and one of the types of C. lateristriga a
skin 245 mm. long.

32. HAPLOCHROMIS MELANONOTUS, sp. n. (Text-fig. 21.)

Depth of body 22 to 3 in length, length of head 3}. Profile
of head evenly decurved. Snout % to 13 diameter of eye, which
is 3 to 32 in length of head, greater than depth of preorbital

Text-figure 21.

Haplochromis melanonotus. 2

or cheek; interorbital width 31 to 4 in head. Jaws equal
anteriorly ; maxillary not extending to below eye; teeth in 4 or
5 series; 40 to 60 biscuspid teeth in outer series of upper jaw.
4 series of scales on cheek. 12 gill-rakers on lower part of ante-
rior arch. Pharyngeal teeth small, none enlarged.~ 34-35 scales
in a longitudinal series, 6 from origin of dorsal to lateral line.
Dorsal X VIII-XIX 10; last spine 2 to 3 length of head, longest
soft rays } head. Anal III 10; third spine from less than 3 to
more than 2 head. Pectoral as long as or a little shorter than
head, nearly or quite reaching anal. Caudal sh ehtly emarginate.
Caudal peduncle 1} as long as deep. Silvery; a blackish band
from nape to upper half of base of caudal.

Two specimens, 85 and 200 mm. in total length (Wood).

CICHLID FISHES OF LAKE NYASSA. 709

33. HapLocHRoMis BREVIS Bouleng., 1908.
Tilapia brevis Bera Cat. Afr. Fish. 111. p. mn ciel Wee
Depth of body 24 in length, length of head 2§. Snout convex,
pier ‘ter than hee of eye, which is twice preorbital depth,
21 to 23 in length of head; interorbital width 4 in head. Jaws
equal anteriorly; maxillary extending to below anterior edge of
eye; teeth in 2 or 3 series; 65 to 80 bicuspid teeth in outer series
of upper jaw. 3 series of scales on cheek. §& or 9 gill-rakers on
lower part of anterior arch. Pharyngeal teeth small. 31 scales
in a longitudinal series, 6 from origin of dorsal to lateral line.
Dorsal XV 12; last spine a little less than 3 length of head.
Anal III 9; third spine stronger than dorsals, 2 2 head. Pectoral
as long as head, reaching anal. Caudal emarginate. Caudal
peduncle as long as deep. Silvery; an opercular spot; a dark
band from nape to base of caudal.
Two specimens, 70 and 75 mm. in total length.

34. HapLocHRomis NoToT#n1A Bouleng., 1902.
Paratilapia nototenia Bouleng. Cat. Afr. Fish. iii. p. 359.

Depth of body 25 to 3 in length, iength of head 22 to 3.
Snout decurved, ohoabes than postor bital part of head. Lyunees
of eye 32 to 42 in length of head, interorbital width 33 to 33,
preorbital Seah 4 to 44. Jaws equal anteriorly; maxillary
extending to below anterior margin of eye; teeth in 3 to 5
series, outer biscuspid in young, conical in adult, 75 to 90 in
outer series of upper jaw. 3 series of scales on gheek, 12 gill-
rakers on lower part of anterior arch. Pharyngeal teeth small.
35 or 36 scales in a longitudinal series, 6 from origin of dorsal
to lateral line. ‘Dorsal XVI-X VIi 10-12; last spine } length of
head. Anal ITI 9-10; third spine j te 2 head. Pectorala little
shorter than head, reaching origin of anal. Caudal emarginate.
Caudal peduncle 135 as long as deep. An opercular spot : a
blackish band on each side of “back from origin of dorsal to caudal ;
dorsal and caudal with bars or series of spots; lower fins tinged
-with orange.

The type and three specimens of 180 to 250 mm. (Wood).

35. HAPLOCHROMIS LEPTURUS, sp.n. (Text-fig. 22.)

Paratilapia rheies (part.) Bouleng. Cat. Afr. Fish. iii. p. 361.

Depth of body 3£ to 33 in length, length of head 3 to 3.
Snout decurved, a ittle shorter than postorbital part of head.
Diameter of eye 44 to 6 in length of head, interorbital width
33 to 4, depth of preorbital 4 ie Ai. Jaws equal anteriorly ;
maxillary extending to vertical from anterior edge of eye;
teeth conical, in 3 to 6 series, 70 to 90 in outer series of
upper jaw. 4 or 5 series of scales on cheek. 11 to 13 gill-
rakers on lower part of anterior arch. Pharyngeal teeth small,
37 to 39 scales in a longitudinal series, 6 or 7 from origin of

*
710 MR. C. TATE REGAN ON THE

dorsal to lateral line. Dorsal XVII 11-13; last spine from a
little less than 1 to nearly 2 length of head. Anal IIT 10-11;
third spine stronger anda little shorter than last dorsal. Pectoral
2 to Z length of head, not reaching anal. Caudal emarginate.

4
Caudal peduncle 14 to 2 as long as deep. Silvery ; back darker ;

Text-figure 22.

Haplochromis lepturus. 3.

an opercular spot; a dark band from origin of dorsal to caudal
(absent in the largest specimen, which is darker than the others);
series of spots on dorsal.

Five specimens of 170 to 240 mm. (Wood) and one of 360 mm.
(Lthoades).

36. HapLoCHROMIS RHOADESII Bouleng., 1908.

Paratilapia rhoadesii (part.) Bouleng. Cat. Afr. Fish. iii. p. 361,
fig. 245.

Depth of body 22 to 3 (adult} or 32 (young) in the length,
leneth of head 24 to 3. Snout decurved, as long as postorbital
part of head. Diameter of eye 4 to 53 in length of head, inter-
orbital width 4 to 435, depth of preorbital 34 to 44. Lower jaw
usually a little projecting ; maxillary not extending to below eye ;
teeth cuspidate in young, conical in adult, in 3 to 5 series, 60 to
80 in outer series of upper jaw. 4 or 5 series of scales on cheek.
11 or 12 gill-rakers on lower part of anterior arch, Pharyngeal
teeth small. 35 to 37 scales in a longitudinal series, 5 to 7 from
origin of dorsal to lateral line. Dorsal XV—X VII 11-13; spines
equal or slightly increasing from sixth or seventh, last 7 to +
length of head. Anal IIT 10; third spine stronger than dorsals,
4 to 3 head. Pectoral 3 to % head, reaching vent or origin of
anal, Caudal emarginate. Caudal peduncle 14 to 12 as long as
deep. An opercular spot; body with or without faint dark cross-
bars; a blackish band from nape to caudal; dorsal with series of
spots.

The type (250 mm.) and a series of specimens 120 to 250 mm,
long (Wood).

_
CICHLID FISHES OF LAKE NYASSA, 711

37, HAPLOCHROMIS ATRITENIATUS, sp. n. (Text-fig, 23.)

Depth of body 2% in length, length of head 3. Snout with
straight profile, nearly as long as postorbital part of head.
Diameter of eye 4 to 4} in length of head, interorbital width 4,
depth of preorbital 4 to 44. Lower jaw slightly projecting ;
maxillary not quite reaching vertical from anterior edge of eye;
teeth in 3 series, outer bicuspid or some conical, 70 in outer series
of upper jaw. 4 series of scales on cheek. 11 gill-rakers on lower
part of anterior arch. Pharyngeal teeth small. 33 or 34 scales
in a longitudinal series, 6 or 7 from origin of dorsal to lateral]
line. Dorsal XVI 10-11; spines equal or slightly increasing

from the sixth, last 2 or a little more than 2 length of head.

‘Text-figure 23.

Ne
PNY
A es
We
WS

Haplochromis atriteniatus. x.

Anal III 9; third spine + head. Pectoral as long as head,
extending a little beyond origin of anal. Caudal emarginate.
Caudal peduncle 13 to 12 as long as deep. An opercular spot ;
a blackish band from nape to caudal; dorsal and caudal with
series of spots. :

Two specimens, 150 and 170 mim. long (Wood).

38, HAPLOCHROMIS SPILORHYNCHUS, Sp. n. (Pl. VI. fig. 2.)

Paratilapia dimidiata (part.) Bouleng. Cat. Afr. Fish, iii. p. 360.
Depth of body 314 to 33 in length, length of head about 3.
Snout decurved, as long as postorbital part of head. Diameter of »
eye 43 to 54 in length of head, less than depth of preorbital or
cheek ; interorbital width 43 to 53 inlength of head. Lower jay
a little projecting; maxillary not extending to below eye; teeth
conical, in 3 or 4 series, outer strong and spaced, 25 to 40 in,
upper jaw. 5 or 6 series of scales on cheek. 10 or 11 gill-

Proc. Zoou. Soc.—1921, No, XLVIITI, 48

712 MR, ©, TATE REGAN ON THE

rakers on lower part of anterior arch, Pharyngeal teeth slender.
36 scales in a longitudinal series, 6 or 7 from origin of dorsal
to lateral line: Dorsal XVI 12; last spine 3 lensth of head.
Anal III 9-10; third spine as long as and ee than Jast
dorsal. Pectoral ? to + head, not reaching anal. Caudal
emarginate, Caudal peduncle 12 13 as long as deep. A large black
spot on preorbital; an Soanonlen spot; a blackish band on each
side of back from origin of dorsal to caudal.

Seven specimens (Wood, Rhoades, Rendall) 150-240 mm. in
total length,

39, HAPLOCHROMIS LONGIPES, sp.n. (Pl. V. fig. 2.)

Depth of body 3 in length, length of head 3. Snout decurved,
twice as Jong as diameter of eye, which is 54 in length of head,
less than depth of preorbital or cheek ; interorbital width 45 in
length of head. Lower jaw a little projecting ; maxillary not quite
reaching vertical from anterior edge of eye. ‘Teeth conical, in
3 series, outer rather strong, 45 in upper jaw. 9 series of scales
on cheek; 13 gill-rakers on lower part of anteriorarch. Pharyn-
geal teeth slender. 57 scales in a longitudinal series, 6 from
origin of dorsal to lateral line. Dorsal X VI 12 ; spines strongly
increasing to last, which is 2 length of head. Anal IIE 10; third
spine ? length of head. Soft dorsal and anal produced, pointed.
Pectoral 2 length of head; pelvics as long as head. Caudal
emarginate. Caudal peduncle 13 as long as deep. An opercular
spot; a blackish band on each side of back from nape to base of
caudal.

A single specimen, 260 mm. in total length.

40, HAPLOCHROMIS CHRULEUS Bouleng., 1908.

Champsochromis ceruleus Bouleng. Cat. Afr. Fish. 111. p. 433,
fig. 295.

Closely related to the preceding, differing as follows :—Depth
of body 4 in length, head 32. Snout 2 diameter of eye, which
is 6 in length of “head. icror bia width 4in head, Maxillary
not aeanlty reaching vertical from anterior edge of eye. 11 gill-
rakers on lower part of anterior arch. Caudal peduncle twice as
long as deep.

Total length 255 mm.

41, HAPLOCHROMIS MACROCHIR, sp. n. (‘Text-fig. 24.)

Depth of body 22 in length, length of head nearly 3.
Snout with straight or slightly convex profile, as long as
diameter of eye, which is 3{ in length of head, 13 depth of
preorbital or cheek; interorbital width 4 to 43 in head. Jaws
equal anteriorly ; maxillary extending to vertical from anterior
edge ef eye; teeth very small, in 2 or 3 series, outer bicuspid,
about 80 in outer series of upper jaw; 4 series of scales on

- —"

CICHLID FISHES OF LAKE, NYASSA, 713

cheek. 13 or 14 gill-rakers on lower part of anterior arch. A
group of enlarged teeth with spherical crowns on posterior half of
lower pharyngeal. 33 scales ina longitudinal series, 6 from origin
of dorsal to lateral line.. Dorsal XVI 10; last spine 2 to 3,
longest soft rays # to # length of head. Anal III 9; third spine
3 to 2 head. Pectoral longer than head, reaching middle of anal.
Caudal emarginate. Caudal peduncle longer than deep. Silvery ;

Text-figure 24.

oS)

Haplochromis macrochir.

an opercular spot; about 10 faint dark cross-bars; dorsal and
caudal with series of spots; anal, in male, with several ocelli.
Two specimens, 115 and 130 mm. long (Wood).

42. HAPLOCHROMIS ARGYROSOMA, sp. n. (Text-fig. 25.)

Tilapia macrophthalma (part.) Bouleng. Cat. Afr. Fish. ii.
p. 261.

Depth of body equal to length of head, 3+ in length of fish.
Snout decurved, shorter than diameter of eye, which is 3 in length
of head, twice depth of preorbital or cheek; interorbital width 5
in length of head. Jaws equal anteriorly: maxillary extending
to below anterior edge of eye; teeth in 2 or 3 series, 50 in outer
series of upper jaw, mostly bicuspid, but the last few on each side
conical ; 2 series of scales on cheek. 11 gill-rakers on lower part
of anterior arch. Last few teeth of two middle series of lower
pharyngeal enlarged. 33 scales in a longitudinal series, 5 from
origin of dorsal to lateral line. Dorsal XVII 10; last spine
nearly 2 length of head. Anal JII 9; third spine less than 2 head.
Pectoral as long as head, reaching anal. Caudal emarginate.
Caudal peduncie 14 as long as deep, Silvery; dorsal and caudal
with series of spots.

48*

714 . MR. GC. TATE REGAN ON THE

A single specimen, one of the types of 7. macrophthalma,
75 mm. in total length.

Text-figure 25.

Haplochromis argyrosoma. Nat. size.

43, HAPLOCHROMIS MACROPHTHALMUS Bouleng., 1908.

Tilapia macrophthalma (part.) Bouleng. Cat. Afr. Fish. iil.
pe 26d, ie ioe

Depth of body 23 to 3 in length, length of head 3. Snout with
slightly convex profile, shorter than diameter of eye, which is
25 in length of head, twice depth of preorbital or cheek, or inter-
orbital width. Lower jaw a little shorter than upper; maxillary
nov extending to below eye; teeth im 2 series, 50 or 60 bicuspid
teeth in outer series of upper jaw, anterior outer teeth of lower
jaw divected outwards. 2 series of scales on cheek. 12 or 13
gill-rakers on lower part of anterior arch. Last few teeth of 2
middle series of lower pharyngeal a little enlarged. 32 or 33
scales in a longitudinal series, 5 from origin of dorsal to lateral
line. Dorsal XVI 10; last spine nearly $ length of head. Anal
Til 8; third spine as long as last dorsal. Pectoral a little longer
than head, extending to above anal. Caudal emarginate.
Caudal peduncle 13 as long as deep, Silvery.

Four specimens, 70 to 80 mm. in total length (/hoades).

44, HAPLOCHROMIS LEUCISCUS, sp.n. (Text-fig. 26.)

Depth of body 31 to 33 in length, length of head 3} to 33.
Snout with straight or slightly convex profile, a little shorter
than diameter of eye, which is 3 in length of head, twice pre-
orbital depth; interorbital width 44 in head. Jaws equal
anteriorly ; maxillary not extending to below eye; teeth in 2 or
3 series; 45 bicuspid teeth in outer series of upper Jaw. 2 or 3
series of scales on cheek. 11 gill-rakers on lower part of anterior
arch. Pharyngeal teeth small. 31 to 33 seales in a longitudinal
series, 5 or 6 from origin of dorsal to lateral line. Dorsal X VI—
XVII 11; last spine 2 length of head. Anal III 9; third spine

CICHLID FISHES OF LAKE NYASSA. 715

Stronger and as long as or:a little shorter than last dorsal.
Pectoral. ¢ to % head, not reaching anal. Caudal emarginate.
Caudal peduncle 14 as long as deep. Silvery ; dorsal and caudal
with series of spots.

Two specimens, 75 and 80 mm. long (hoades, Wood).

Text-figure 26.

Haplochromis leuciscus. Nat. size.

45, HAPLOCHROMIS INORNATUS Bouleng., 1908.

Tilapia inornata Bouleng. Cat. Afr. Fish. ui. p. 263, fig. 178.

Depth of body 3} to 33 in length, length of head 37. Snout
shorter than diameter of eye, which is 2? to 3 in length of head
and twice depth of preeorbital or cheek; interorbital width 4} in
length of head. Jaws equal anteriorly ; maxillary not extending
to below eye; teeth small, in 3 series; 50 bicuspid teeth in outer
series of upper jaw. 2 or 3 series of scales on cheek. 15 or 16
gill-rakers on lower part of anterior arch. Pharyngeal teeth
small. 34 to 36 svales in a longitudinal series, 5 from origin of
dorsal to lateral line. Dorsal XVI-X VIII 11-12; last spine 2
length of head. Anal Il] 9; third spine nearly as Jong as last
dorsal. Pectoral as long as head, not reaching anal. Caudal
emarginate. Caudal peduncle 13 as long as deep. Silvery.

Two specimens, 85 and 95 mm. in total length (2hoades).

46. HAPLOCHROMIS MICRENTODON, sp. n. (Text-fig. 27.)

Depth of body 23 to 23 in length, length of head 3} to 34.
Snout with slightly convex profile, a little shorter than diameter
of eye, which is 3 in length of head, 13 preorbital depth, 14
depth of cheek ; interorbital width 4 in length of head. Lower
jaw a little shorter than upper; maxillary not extending to
below eye; teeth in 2 or 3 series, outer bicuspid, about 70 in
upper jaw; inner teeth of lower jaw minute, almost invisible,
outer anterior teeth directed outwards. 2 or 3 series of scales

716 ( MR. C. TATE REGAN ON THE

on cheek. 16 or 17 gill-rakers on lower part of anterior arch.
Pharyngeal teeth very small. 33 scales in a longitudinal series,
5 from ae of dorsal to lateral line. Dorsal XVI 10-11; last
spine 2 to 3, longest soft rays 3 to 2 length of head. ‘Adal
IIT 8-9; third spine a little shorter than last dorsal. Pectoral
longer than head, extending beyond origin of anal. Caudal

Text-figure 27.

Haplochromis micrentodon. 4.

emarginate. Caudal peduncle longer than deep. Silvery, with
isiees of several dark ecross-bars ; Gora and caudal with series
of spots.

Two specimens, 110 and 115 mm. long (Wood).

47. HAPLOCHROMIS EUCINOSTOMUS, sp. n. (PI. IV. fig. 1.)

Depth of body 31 to 34 in length, length of head 3}. Snout
with straight profile, as long as "ie ater of eye, sytnichit is 33 in
length of “head, rauch greater than preorbital depth; inter-
orbital width 37 to 33 in. length of head. Mouth small, oblique ;
jaws equal anteriorly ; premaxillary pedicels extending to be-
tween middle of orbits, 2 length of head; maxillary not
extending to below eye; teeth small, in 2 or 38 series, outer
bicuspid. 3 series of scales on cheek. 16 or 17 gill- rakers on
lower part of anterior arch. Pharyngeal teeth smail, slender.
35 scales in a longitudinal series, 5 from origin of dorsal to
lateral line. Dorsal XVI-XVII 11-12; last spine 2 length of
head. Anal IIT 10; third spine stronger than dorsals, 3 head.
Pectoral a little shorter than head, not reaching anal. ante
emarginate, densely scaled in basal half. Caudal peduncle 13 as
long as deep. Silvery, with traces of dark cross-bars.

Two specimens, 85 and 100 mm. long (Wood), the smaller (? d )
much darker in colour than the larger.

————

—e

CICHLID FISHES OF LAKE NYASSA. furs

48. Haprocuromis PRHORBITALIS, sp. n. (PI. IIT.)

Depth of body 23 to 22 in length, length of head 22-3.
Snout longer than postorbital part of head ; upper profile straight,
obliquely descending. Diameter of eye 5 in length of head,
interorbital width 43, depth of prezorbital 3. Lower jaw pro-
jecting ; maxillary ending not far behind nostril; teeth in 3
series in upper jaw, 4 in lower, outer bicuspid, about 70 in upper
jaw. 4 series of scales on cheek. 9 or 10 gill-rakers on lower
part of anterior arch. Pharyngeal teeth small, slender. 35 or
36 scales in a longitudinal series, 6 from origin of dorsal to
lateral line. Dorsal XV—XVI 12-13; last spine 2 or d length
of head; longest soft rays 2 or 2 head. Anal III 10- il third
spine stronger than dorsals, # or 4 head. Pectoral a little shorter
than head, reaching anal. Caudal emarginate. Caudal peduncle
14 as lone as deep. Greyish ; an opercular spot; dorsal with
oblique str ipes or series of spots.

Two specimens, 210 and 240 mm. in total length (Wood), The
larger has the dorsal fin higher than the other and the lower fins
dusky; it is probably a male.

49, HAPLOCHROMIS COMPRESSICEPS Bouleng., 1908.

Paratilapia compressiceps Bouleng. Cat. Afr. Fish. ii. p. 331,
fig. .222.

Depth of body 3 in length, length of head 23 to 2%. Head
4 times as long as broad; snout a little concave in front of eye,
thence straight, longer than postorbital part of head. Diameter
of eye 54 to 6 in length of head, equal to interorbital width, from
a little more than $ to nearly 3 depth of preorbital, Lower jaw
projecting ; chin deep; mavxillary ending a little behind nostril ;
teeth conical, in 3 series. 3 or 4 series of scales on cheek. 11
to 13 gill-rakers on lower part of anterior arch. Pharyngeal
teeth small. 33 to 35 scales in a longitudinal series, 5 or 6 from
origin 2 dorsal to lateral line. Dorsal XV-XVI 11-13; last
spine 4+ length of head. Anal III 10-11; third spine as long as
last dienes Caudal scaly, truncate. Caudal peduncle 14 to 12
as long as deep. Silvery; three yellow-green bands,,one along
upper outline of head and body, the second above upper lateral
line, the third along middle of side.

The type, 160 mm. tong, and 3 specimens of 180-200 mm.
(Wood).

50. HaPLOCHROMIS MACRORHYNCHUS, sp. n. (Text-fig. 28.)

Tilapia rostrata (part.) Bouleng. Cat. Afr. Fish. ii. p. 255.

Depth of body 22 to 22 in length, length of head 23 to 22.
Snout with straight or convex vrofile, longer than postorbital
part of head. Diameter of eye 5 to 53 in lensth of head, inter-
orbital width 3} to 33, depth of preorbital 3? to 4. Jaws equal
anteriorly ; maxillary not extending to below eye; teeth in 4 or

718 _ MR. C. TATE REGAN ON THE

5 series, outer bicuspid or posteriorly unicuspid, about 70 in
outer series of upper jaw. 3 or 4 series of scales on cheek. Gill-
rakers rather long, 15 to 17 on lower part of anterior arch.
Pharyngeal teeth small. 34 or 35 scales in a longitudinal series,
6 or 7 from origin of dorsal to lateral lime. Dorsal XV-XVI 11;
last spine 3 length of head. Anal ITT 9; third spine # length of
head. Pectoral # length of head, nearly or quite reaching anal.
Caudal emarginate. Caudal peduncle 13 to 14 as long as deep.

Text-figure 28.

Haplochromis macrorhynchus. +.

Upper half of body with 6 dark cross-bars broken up into 3
series of spots, the first of the lowest series on operculum, the
sixth at base of caudal; dorsal with series of spots.

Three specimens, 190 to 210 mm. long (d, Md, Moore).

51. HAPLOCHROMIS ROSTRATUS Bouleng., 1899.

Tilapia rostrata (part.) Bouleng. Cat. Afr. Fish. i. p. 255,
fig. 172.

Depth of body 34 in length, length of head 25. Snout with
convex profile, longer than postorbital part of head. Diameter
of eye 4 in length of head, interorbital width 43, depth of pre-
orbital 43. Jaws equal anteriorly; maxillary not extending to
below eye; teeth in 3 well-separated series, outer bicuspid, 54
in outer series of upper jaw. 3 series of scales on cheek.
Gill-rakers short, 19 or 20 on lower part of anterior arch.
Pharyngeal teeth small. 34 scales in a longitudinal series,
6 or 7 from origin of dorsal to lateral line. Dorsal XVI 12;
last spine nearly 2 length of head. Anal JII 10; third spine

3 length of head. Pectoral? Caudal emarginate. Caudal .

peduncle i2 as long as deep. Colour nearly as in H. macro-
rhynchus, but the posterior bars more oblique and the fins
unspotted.

The type, 105 mm. long.

et Ee ee

CICHLID FISHES OF LAKE NYASSA. “719

52. HAPLOCHROMIS MACROSTOMA, sp.n. (PI. IV. fig. 2.)

Depth of body 3 in length, length of head 23. Snout with
profile a little concave in front of eye, thence straight, declivous,
22 as long as diameter of eye, which is 55 in length of head, less
than preorbital depth, 3 depth of cheek ; interorbital w idth 4 3
in length of head. Jaws equal anteriorly ; maxillary consider-
ably exposed distally, extending to below anterior 4 of eye;
teeth small, conical, in 6 series. 6-series of scales on cheek.
12 gill-rakers on lower part of anterior arch. Pharyngeal teeth
slender.. 35 scales in a longitudinal series, 6 from origin of
dorsal to lateral ae Dorsal XVI 10; last spine 4, longest soft
rays less than 4 length of head. Anal II 10; third spine
stronger than dorsals, + head. Pectoral 3 hee not quite
reaching anal. Caudal very slightly emarginate Caudal
peduncle lias long as deep. A series of about 8 dark spots on
each side of back; a second series below lateral line, posteriorly
confluent to form a band.

A single specimen, 260 mm. in total length (Weod).

1]. LeruHrinops, gen. n.
(type Chromis lethrinus Giinth.),

Differs from Haplochromis in the dentition. Teeth very small
and slender, in a few series, forming narrow bands which are
interrupted ‘at the symphyses : ; outer teeth mostly bicuspid, often
unicuspid posteriorly, inner uni- or tri-cuspid.

ae four pheeies.

Synopsis of the Species.

I. Pharyngeal teeth all small.
Snout shorter than postorbital part of head; maxillary not far

short of vertical from anterior edge of eye ............ 0000005. 1. albus.
Snout longer than postorbital part of head; maxillary ending
MCh ka EMM CNMOSHRI ees ae ety ec ecctes ee tatstesauterens amacronhynenus.
IJ. Last few teeth of two middle series of lower pharyngeal somewhat enlarged.
10 gill-rakers on lower part of anterior arch ........................ 3. lethrinus.
12-13 gill-rakers on lower part of anterior arch .................... 4. leptodon.

1. LETHRINOPS ALBUS, sp. n.

Tilapia macrophthalma (part.) Bouleng. Cat. Afr. Fish. iii.
p. 261.

Depth of body equal to length of head, 3 in length of fish.
uae with straight profile, as long as diameter of eye, which is

z in length of head, interorbital width 5, depth of przorbital
Al. dase equal ; maxillary not far short of vertical from
anterior edge of eye; teeth in 2 series. 3 series of scales on
cheek. 10 gill-rakers on lower part of anterior arch. Pharyn-
geal teeth small. 32 scales in a longitudinal series, 5 from

720 _ MR. C. TATE REGAN ON THE

origin of dorsal to lateral line. Dorsal XVI 10. Anal IIL 9;
third spine 2 length of head, a little shorter than last domsal,
Pectoral as long as head, extending beyond origin of anal.
Caudal scaly, emarginate. Caudal peduncle as lone as deep.
Silvery ; an opercular spot.

A single specimen, 100 mm. in total length (Whyte).

2. LETHRINOPS MACRORHYNCHUS, Sp. 0.

Tilapia lethrinus (part.) Bouleng. Cat. Afr. Fish. i. p. 254.

Depth of body 24 in length, length of head 22, Snout with
straight profile, longer than postorbital part of head. Inter-
orbital region flat. Diameter of eye 44 in length of head,

interorbital width 43, depth of preorbital 3. Lower jaw pro--

jecting ; maxillary ending not far behind nostril; teeth in 3
series. 3 series of scales on cheek. 10 gill-rakers on lower part
of anterior arch. Pharyngeal teeth small. 33 scales in a longi-
tudinal series, 5 from origin of dorsal to lateral iine. Dorsal
XVI 11; last spine 3 length of head. Anal III 9; third spine
less than } head. Pectoral as long as head, extending nearly
to middle of anal. Caudal scaly, emarginate. Caudal peduncle
12 as long as deep. A blackish band from nape along upper
lateral line to base of caudal.
A single specimen, 180 mm. long (Whyte).

3. LErHRINoPs LETHRINUS Ginth., 1893.

ees lethrinus (part.) Bouleng. Cat. Afr, Fish. 11. p. 254,

seulivulk

Se of body 23 to 22 in length, length of head 22 to 3.
Snout with straight profile, as long as postorbital part of head.
Interorbital region flat. Diameter of eye 3? to 47 in length of
head, 3 nterorbital width 4, preorbital depth 3 to 33. Lower jaw
slightly projecting ; maxillary extending to between nostril and
eye; teeth in 3 to 5 series. 3 or 4 series of scales on cheek.
10 gill-rakers on lower part of anterior arch. Lower pharyngeal
with the last few teeth of the two middle series a little enlarged.
32 to 34 scales in a longitudinal series, 5 or 6 from origin of
dorsal to lateral line. Dorsal XV-XVI 11; last spine from
less than 4 to 3 length of head. Anal III 9; third spine 3 to 2
head. Pectoral a little shorter than head, reaching origin of
anal. Caudal densely scaled, emarginate. Caudal peduncle 14
to 14 as long as deep. A blackish lateral stripe running back-
wards above lower lateral line; sometimes a second above Jateral
line and a third near edge of back, or these may be represented
by series of spots; dor cal spotted.

Three specimens, 140 to 200 mm. long (Johnston, Moore).

CICHLID FISHES OF LAKE NYASSA, een

4, LETHRINOPS LEPTODON, sp.n. (Text-fig. 29.)

Depth of body 22 to 3 in length, length of head 25 to 33.
Snout with steep, straight or slightly convex profile, as long as or
alittle shorter than postorbital part of head. Interorbital region
convex. Diameter of eye 32 to 4 in length of head, interorbital
width 34 to 5, depth of preorbital 3 to 33. Lower jaw slightly
projecting; maxillary not extending to below eye; teeth in 3 or
4 series. 3 or 4 series of scales on cheek. 12 or 13 gill-rakers
on lower part of anterior arch. Lower pharyngeal with the last
few teeth of the two middle series a little enlarged. 31 to 34
scales in a longitudinal series, 5 or 6 from origin of dorsal to
lateral line. Dorsal XV-XVI 10-13; last spine 2 to } length of
head. Anal III 9-10; third spine + to 2 head. Pectoral nearly

Text-figure 29.

Lethrinops leptodon. 3.

as long as head, reaching anal. Caudal scaly, emarginate.
Caudal peduncle 1} to 14 as long as deep. Silvery or bluish,
with faint dark cross-bars; an opercular spot; an oblong dark
blotch on lateral line below spinous dorsal, from just below
which a dark band runs backwards to the caudal; dorsal and
caudal with series of orange spots; dorsal sometimes with a
yellow edge; anal with several ocelli in males.
Eight specimens, 140 to 180 mm. in total length (Wood).

12. Docimopus Bouleng., 1896.

Differs from Haplochromis in the dentition. Teeth compressed,
uni- or tri-cuspid, in 4 or 5 well-separated series; outer teeth
large, forming a close-set series of about 20 in each jaw,

Nyassa; a single species.

722 MR, C. TATE REGAN ON THE

Docimopts joHNnstont Bouleng., 1896.

Docimodus johnstonii Bouleng. Cat. Afr. Fish. iti. p. 282,
fig. 192.

Depth of body 2? to 3 in length, length of head 3 to 33.
Snout convex, 17 to 13 diameter of eye, “which is equal to or
greater than depth of preorbital, 4 to 43 in length of head,
interorbital width 3. to 33. Jaws strong ; lower projecting ;
maxillary not extending to below eye. 3 or 4 series of scales on
cheek. 10 or 1} gill- fakers on lower part of anterior arch.
Pharyngeal teeth small, COM DUE essed, bicuspid. 34 to 36 scales in
a longitudinal series, 5 from origin of dorsal to lateral line.
Dorsal XVI-XVII 9-11; last spine 4 to 2 length of head.
Anal IIT 9-10; third spine as long as or a little “shorter than
dorsal. Pectoral as long as head, reaching anal. Candal scaly,
truncate, with slight median notch. Caudal peduncle 14 to 132
as long as deep. A dark lateral band from shoulder to base of
caudal; a series of dark spots at base of dorsal; an opercular
spot; dorsal with series of spots, in male with a dark intra-
marginal stripe and pale edge; anal and cafidal, in the male,
dusky with pale edge.

Four specimens, 160 to 250 mm. long (Johnston, Wood).

13. Cyrrocara Boulenger, 1902
(type C. moorit Bouleng.).

Differs from Haplochromis in the structure of the spinous
dorsal fin, which has the edge of the membrane straight between
the tips of the spines. Teeth in several series, the outer enlarged,
conical, or some bicuspid.

Nyassa.
Synopsis of the Species.
Jaws equal; caudal truncate or slightly emarginate .............. ..... 1. venusta.
Jaws equal; caudal crescentically emarginate........................2..... 2. anmectens.
IDo\iiGNO TPN RON XENUNVES! BAmeUE UA ao 0 SAG eh guduewcdabogiaasdusuer tdvoeaioaden sey! Sy BACON:

1. CyRTOCARA VENUSTA.

Haplochromis venustus Bouleng. Cat. Afr. Fish. ii. p. 287,
fio. 195,

“Depth of body 24 to 22 in length, length of head 3 to 34.
Snout with straight profile, as tone as postorbital part of head.
Diameter of eye “Ag to 5 in lengéh’ of head, equal to or a little
less than preorbital depth : interorbital width 34 to 4 in length
of head. Jaws equal anteriorly; maxillary not pcndine” to
below eye; teeth conical, or some of the outer bicuspid, in 4 or 5
series, outer larger. 3 or 4 series of scales on cheek. 11 or 12
gill rakers on lower part of anterior arch, 32 to 34 scales in
a longitudinal series, 6 or 7 from origin ef dorsal to lateral
line. Dorsal XV-XVI 10-11; last spine 2 length of head.

Anal IIT 10; third spine 4 head. Pectoral as long as head,

a ae
ae ee eee

- CICHLID FISHES OF LAKE NYASSA, TOS

about reaching anal. Caudal Sy truneate or slightly emar-
ginate. Caudal peduncle 11 to 13 as long as deep. Bluish, with
large vertically expanded blue-black spots tending to form
inresular eross-bars ; dorsal and anal with broad orange margin.

Four specimens, 170 to 200 mm. long (Lhoades).

Boulenger’s figure shows a slight daunentees of lappets, but
where the dorsal membrane is not torn its margin runs evenly
between the tips of the spines.

2. CYRTOCARA ANNECTENS, sp. n. (Text-fig. 30.)

Depth of body 24 to 22 in length, length of head 32 to 34,
Occiput convex, out straight or slightly concave, a little shorter
than postorbital part of head. Diameter of eye 4to 44 in length
of head, equal to preorbital depth; interorbital width 32 in
length os head. Jaws equal anteriorly ; maxillary not quite
reaching to below eye; teeth conical, in 3 or 4 series, outer

Text-figure 30.

Cyrtocara annectens. +.

larger. 2 or 3 series of scales on cheek, 12 or 13 gill-rakers on
lower part of anterior arch. 34 to 36 ous in a longitudinal
series, 0 or 6. from origin of dorsal to lateral line. Dorsal
XVI_XVII 11-12; ia spine 2 head. Anal III 8-9; third
spine stronger and shorter than last dorsal. Pectoral as long
as or a, little longer than head, reaching anal. Caudal densely .
scaled, crescentically emarginate. Caudal peduncle 13 to 14 as
long as deep. Uniformly eich black.

inrae specimens, 175 to 195 mm. in total length.

This species connects C. venusta with C. moorii, having the
mouth formed as in the former, but in most srihenaaeeeneace more
nearly agreeing with the latter.

124 ._ MR. C, TATE REGAN ON THE

3. CYRTOCARA MOORIL.

Very near the preceding species, but lower jaw projecting,
some of the outer teeth pointing outwards, and maxillary
extending to below eye. Occiput very convex. 3 or 4 series of
scales on cheek. 11] gill-rakers on lower part of anterior arch.
Dorsal XV-XVI 1]. Anal IIJ-IV 8-9.

14. RHAMPHOCHROMIS, gen. n.*
(type Hemichromis longiceps Giinth.).

Form elongate. Snout produced, nearly or quite 3 length of
head. Mouth with lateral cleft, ending far in front of eye;
premaxillaries with an anterior beak-like expansion ; lower Jaw
deep ; teeth conical, acute, biserial, or sometimes a third series
anteriorly in upper jaw; teeth of outer series strong or mode-
rately strong, set well apart; anterior teeth of second series in
upper jaw enlarged. 16 to 18 gill-rakers on lower part of anterior
arch. Scales 36 to 44. Dorsal XVI-XX 11-13; spines slender,
rather short; soft fin rounded. Anal III 9-11; third spine
about as long as and stronger than last dersal. Pectoral rather
short. Caudal scaly, emarginate.

Nyassa.

Synopsis of the Species.
I. Scales 36 to 40. Depth 4 to 42 in length. Depth of caudal peduncle not less

than half its Jength. ;
A. Interorbital width about 4 in head, diameter of eye 53 to 6 (in specimens

of 200 to 240 mm.); 3 or 4 series of scales on cheek.
Teeth moderate, 20 to 25 in outer series on each side............ 1. longiceps.
Teeth strong, 10 to 15 in outer series on each side ...,........... 2. ferox.

B. Interorbital width about 5 in head ; teeth strong.

Diameter of eve 5 in head (in specimens of 200 to 230mm.) ;
Stor Avsenies/ oh scales onicheekec.s-nass-+ ser eer aee eee enah os
Diameter of eye 6 to 8 in head (in specimens of 160 to

3. macrophthalmus.

330mm.); 5 or 6 series of scales on cheek....,................ 4. qwoodi.
If. Scales 43 or 44. Depth 4% to 54 in length.
Caudal peduncle 24 as long as deep; snout convex ............... 5. esow.
Caudal peduncle 2% as long as deep; snout straight ............ 6. leptosoma,

1. RHAMPHOCHROMIS LONGICEPS.

Hemichromis longiceps (part.) Giinth. P. Z. 8. 1864, p. 313.

Champsochromis longiceps (part.) Bouleng. Cat. Afr. Fish. ii,
p. 434 (1915).

Depth of body 4 in length, lengtn of head 24. Diameter of
eye 22 in length of snout, 53 in length of head ; interorbital
width 4 in length of head. Jaws meeting anteriorly; chin
prominent; teeth moderate, 20 to 25 on each side in upper jaw.
4 series of scales on cheek. 36 to 38 scales in a longitudinal

* In the ‘Zoological Record’ for 1916, I named C. ceruleus as the type of
Champsochromis; consequently a new name has to be found for the other species
included in Champsochromis by Boulenger.

CICHLID FISHES OF LAKE NYASSA. 125

series, 5 or 6 from origin of dorsal to lateral line. Dorsal X VII-—
XVIII 11-12; last spine 7 length of head. Anal III 9.
Pectoral 4 length of head. Caudal peduncle 15 as long as deep.
Silvery; back darker ; an opercular spot.

Description mainly based on a specimen 240 mm. long
(Moore), which has been compared with one of the types, a skin
of 220mm. A skeleton has 36 (18+18) vertebre.

2, RHAMPHOCHROMIS FERQX, Sp. n.

? Hemichromis longiceps (part.) Giinth. P. Z. 8. 1864, p. 313.
Champsochromis longiceps (part.) Bouleng. Cat. Afr. Fish. iii.
434 (1915).

Depth of body 4 in length, length of head 3. Snout a
little less than 4 length of head. Diameter of eye 6 in length
of head, interorbitai width 43. Jaws meeting anteriorly; chin
prominent ; teeth strong, 10 to 15 on each side of upper jaw.
3 or 4 series of scales on cheek. 38 or 39 scales in a longi-
tudinal series, 5 or 6 from origin of dorsal to lateral line,
Dorsal XVII 12; last spine 7 length of head. Anal IIT 10.
Pectoral 4 length of head. Caudal peduncle twice as long as
deep. Silvery; back darker; an opercular spot.

Two spirit-specimens (Moore, Rendall), 200 and 240 mm. in total
length. Gunther’s second type-specimen of H. longiceps probably
belongs to this species.

3. RHAMPHOCHROMIS MACROPHTHALMUS, sp. n. (PI. VI. fig. 2.)

Depth of body 4 to 44 in length, length of head 22, Diameter
of eye 2 in length of snout, 5 in length of head, equal to
interorbital width. 3 or 4 series of scales on cheek. 38 to 40
scales in a longitudinal series, 5 to 7 from first dorsal spine to
lateral line. Dorsal XVIIL-XIX 11- 12; last spine 1 length
of head. Anal III 10. Pectoral 2 length of head. Caudal
peduncle twice as long as deep. Silvery ; back dar ker ; a blackish
opercular spot; dorsal and caudal greyish, pelvies and anal
orange.

Three examples, 200 to 230 mm, in total length (Wood).

4, RHAMPHOCHROMIS WOODT, sp, n.

Champsochromis longiceps (part.) Bouleng. Cat. Afr. Fish. iii.
p. 434, fig. 296 (1915).

Depth fice body 4 to 4% in length, length of head 22 to 23.
Snout 3 length of head. Diameter of eye 6 to 8 in lepgth a
head, jae eoriiel width 42 te 5. Lower jaw projecting. 5 or 6
sevies of scales on cheek. “38 to 40 scales in a lonvitudinal series,
6 or 7 from origin of aaa to lateral line. Dorsal XVII-XIX
11-12; last spine + to + length of head. Anal III 10-11.
Pectoral 2 or a little ee than 2 length of head. Caudal peduncle

726 MR, GC. TATE REGAN ON THE

13 to 2 aslong as deep, Silvery, back darker; an opercular spot ;
dorsal and caudal spotted.

Four specimens, the one figured by Boulenger, 330 mm. long
(Rhoades), and three of 160 to 215mm. (Wood). A skeleton has
38 (18 +20) vertebre.

5. RHAMPHOCHROMIS ESOX.

Paratilapia esox (part.) Boulenger, Ann, & Mag. N, H. (8) 11.
1908, p. 240.

Champsochromis longiceps (part.) Boulenger, Cat. Afr, Fish. i.
p. 434 (1915).

Champsochromis esox (part.) Boulenger, t. c. p. 435, fig. 297.

Depth of body 42 in length, length of head 3. Snout with
_ convex upper profile, 3 length of head.) Diameter of eye 72 to 8i
in length of head, interorbital width 32. Lower jaw projecting;
only the inner edge of Sfaginecallleny sheathed by preeorbital.
5 or 6 series of scales on cheek. 43 or 44 scales in a longitudinal
series, 7 or 8 from origin of dersal to lateral hne. Dorsal X VITI-—
XX 12; last spine + length of head. Anal IIT 10. Pectoral
2 length of head. Caudal peduncle 27 as long as deep. Silvery;
back darker; an opercular spot and a lateral band.

Two spirit-specimens, one of 370mm. the type figured by
Boulenger, the other 330mm. long (Joore), and a skeleton with
39 (19 + 20) vertebree.

6. RHAMPHOCHROMIS LEPTOSOMA, Sp, 0.

Paratilapia esox (part.) Bouleng. Ann. & Mag. N. H. (8) ii.
1908, p. 240.

Champsochromis esox (part.) Bouleng. Cat. Afr. Fish. ii, p. 435
(1915).

Depth of body 53 in length, length of head 3. Snout
2 length of head, with straight upper profile ; diameter of eye 74
in length of head, interorbital width 4. Distal half of premaxil-
lary sheathed by preorbital for nearly its whole width; jaws
meeting anteriorly, but chin prominent. 6 series of scales on
cheek. 43 scales in a longitudinal series, 7 from origin of dorsal
to lateral line. Dorsal XIX 13; last spine 4 length of head.
Caudal peduncle 2? as long as deep. Silvery; back darker ;
a dark opercular spot and a lateral band.

A single specimen, one of the types of P. esox, 380 mm. in
total length.

15, AULONOCARA, gen. n.

Fronfal, nasal, orbital, preopercular, and mandibular bones
with large channels with wide openings as Trematocara, from
which 4ulonocara may be distinguished externally by the bicuspid
outer teeth, the more numerous ‘dor sal spines, and the presence of
two lateral lines.

CICHLID FISHES OF LAKE NYASSA. 127

AULONOCARA NYASS&, sp.n. (PI. V. fig. 1.)

Depth of body 24 to 3 in length, length of head 22 te 3.
Snout with ee or slightly convex profile, as long as post-
orbital part of head. Diameter of eye 33 to 33 in length of head,
equal to or greater than preorbital depth, greater Ales ree
orbital width. Jaws equal anteriorly; maxillary eanieealen, not
extending to below eye; teeth small and slender, in 4 or 5 series,
outer bicuspid, scarcely larger than inner. 9 or ‘10 gill-rakers on
lower part of anterior arch. Pharyngeal teeth small, slender,
bicuspid. 31 or 32 scales in a longitudinal series, 5 or 6 from
origin of dorsal to lateral line. Dorsal XV-XVI 10-11; last
spine 2.to 4 length of head. Anal III 9; third spine stronger
and as long as or a little shorter than last dorsal. Pectoral as
long as head, reaching anal. Caudal scaly, emarginate. Caudal
peduncle longer than deep. Silvery; an opercular spot; about
10 faint dark cross-bars; dorsal with series of orange spots ;
lower fins orange (9) or blackish, the anal with orange
spots (¢).

Three specimens, 100-125 mm. long (Wood).

EXPLANATION OF THE PLATES.

Prats I. Haplochromis polystigma.
Puate II. 5 woodi.
Puate IIT. . preorbitalis.
Pruate LV. fig. 1. s eucinostomus.
fig. 2. 0 macrostoma.
Puate V. fig. 1. Aulonocara nyasse.
fig. 2. Haplochromis longipes.
Prate VI. fig. 1. Rhamphochromis macrophthalmus.
fig. 2. Haplochromis spilorhynchus.

Proc. Zoot, Soc.—1921, No. XLIX, AQ

ON THE SYSTEMATIC POSITION OF SAGARTIA LUCIA. 729

37. Note on the Systematic Position and Distribution of the
Actinian Sagartia luaw. By J. Puayrarr McMovrricn,
OMEZ:S:

Received September 19, 1921: Read October 18, 1921.]
(Text-figures 1-4.)

During the last twenty years the small Actinian known as
Sagartia lucie has frequently been an object of study, and much
is known as to its variations, methods of reproduction, powers of
regeneration, reaction to light, locomotion, and general ecology.
Ne yet, lew eee, it has not been subjected fe an anatomical
investigation sufficiently detailed to reveal the features by which
its systematic position may be determined.

It was first described in 1898 by Verrill, who gave a fairly
full description of its external form and coloration, without,
however, considering its anatomical structure. Later, Mrs. G. C.
Davenport (1903) in a study of the variations in the number and
arrangement of the vertical orange stripes, which are usually
such a characteristic feature in the coloration of the column-
wall, showed that reproduction by longitudinal division or by
basal fragmentation was of frequent occurrence, variations in the
number of the orange stripes resulting therefrom, as well as
probably variations in the arrangement of the mesenteries, which,
in the individuals with twelve stripes: were stated to be ‘usually
arranged in twenty-four pairs. But still there was lacking a
sufficiently detailed account of the anatomy of the species to
determine definitely its systematic affiliations, and other papers
that have since dealt with it have not altered the situation in
this respect. It may be well, therefore, to put on record some
observations that were made several years ago, especially as they
led to interesting conclusions not only as to the systematic
position of the fori, but also as to its distribution.

Concerning the disk and tentacles little need be said, further
than to note the weakness of their musculature, the ectodermal
fibres of the tentacles forming a simple layer on the smooth, or
but slightly folded, surface of the mesoglea; on the disk they
may even be absent. Between the bases of the outermost ten-
tacles and the apparent margin of the column there is a deep
fosse, which is very characteristic of the species (text-fig. 1). It
is evidently equivalent to the thin portion of the column-wall that
intervenes in Metridium senilis between the outermost tentacles
and the so-called collar, this latter being the upper edge of the
strong mesogleeal sphincter. But whereas in Wetridiwm this
thin portion of the column is fully exposed in expansion, in
Sagartia lucie it remains introverted, thus producing the
characteristic fosse. ‘The mesoglea forming its walls is very thin

and smooth on both surfaces, aid both the endoderm and ectoderm
4.9%

730 MR. J. P. MCMURRICH ON THE SYSTEMATIC

are also thin, the latter containing a few scattered and almost
spherical mucous glands, without distinct indication of muscle
fibres. The outer wall of the fosse joins the column proper at the
apparent margin, and at once a change of structure is observable.
The mesoglea becomes decidedly thicker and its outer surface
irregular; the ectoderm thickens and becomes richly supplied
with gland cells, which, with the greater thickness of the layer,
assume an oval or pyriform shape; and a weak circular endo-
dermal musculature becomes evident. But there is no sign of a

5
sphincter, either endodermal or mesogleal. One would expect

Text-figure 1.

Sections through the margin and outermost tentacles of an individual from
Woods Hole. ¢=tentacle ; m=margin.

to find a mesogleal sphincter just below the apparent margin,
but in the several individuals that were sectioned I looked in
vain for even traces of it.

In two particulars, then, Sagartia lucie differs notably from
the typical members of the genus to which it has been assigned,
namely, in the presence of a- deep fosse and in the absence of a
mesoglwal sphincter. In the one feature it shows some resem-
blanee to Metridiwm, and in the absence of a sphincter it is

POSITION AND DISTRIBUTION OF SAGARTIA LUCLA, 731

paralleled by some members of the genus Aiptasia; neither of
these: genera, however, belonging to the same subfamily as
Sagartia.

When one comes to consider the arrangement of the mesen-
teries, difficulties are at once encountered, because of the tendency
of the species to reproduce by longitudinal fission. Some halt-
dozen individuals of which serial sections were prepared all
showed irregularities in the arrangement of the mesenteries that
may be reasonably attributed to this process, but nevertheless,
they permit of inferences as to what the arrangement may have
been before fission occurred. Three individuals may be taken as
examples (text-fig. 2). In each of these in sections through the
middle of the column there was a deep siphonoglyph at one end
of the long axis of the stomodeum, and to this a pair of directive
mesenteries with well-developed muscle pennons was attached.
Opposite it was a second siphonoglyph which was relatively quite
shallow, and to this a second pair of directives was attached, the

Text-figure 2.

Diagrams showing the arrangement of the mesenteries in three individuals

from Woods Hole, Mass.

muscle pennons of these being, however, very imperfectly de-
veloped. On either side of the fully-developed directives there
were representatives of two cycles of imperfect mesenteries, and
then came on either side a pair of perfect ones. So far the
arrangement is quite regular except for the imperfect develop-
ment of one siphonoglyph and its directives, but in two of the
individuals, one member of each of the lateral pairs (II. and VI.)
was decidedly less developed than the other. In these same two
individuals (B and C) the pair II. is suceeeded by another perfect
pair (III.) poorly developed, and next this is the second pair of
directives (IV.), no imperfect pairs occurring in the interspaces
between IJ. and III. and III. and IV. Similarly, another pair
of perfect mesenteries (V.) with fully-developed muscle pennons
occurs between IV. and VI., and while no imperfect pairs occur
in the interval between these and IV., in the interval on the other
side there is in one individual a single small pair and in the

ayy MR. J. P. MCMURRICH ON THE SYSTEMATIC

other two such pairs. These two indivicuals have, then, six
pairs of perfect mesenteries and a number of imperfect. ones
disposed unsymmetrically. It is to be noted that this arrange-
ment is that found in sections passing through the aboral half
of the stomodezeum ; higher up, close to the bases of the tentacles,
«number of additional rudimentary mesenteries may be found,
but these need not be considered at present.

In the third individual (A) there were but five pairs of perfect
mesenteries in sections passing through the aboral half of the
stomodeum. Higher up, however, six occurred, with repre-
sentatives of two cycles of imperfect ones in the intervals on
either side of the fully-developed directives and in that between
Vi. and V., but only a single feeble pair in each of the other
intervals. Following the series of sections aborally it is found
that the change from the conditions showing six pairs of perfect
mesenteries to that showing only five is due to the disappearance
of a member of each of the pairs IT. and IIT. together with the
intervening imperfect mesenteries, the remaining members ot
the perfect, pairs thus being brought into apposition and seeming
to form a single pair.

The explanation which seems to fit these cases is that each is
an example of an individual that has undergone longitudinal
fission. The half of the body with the deep siphonoglyph and the
representatives of three cycles of mesenteries shows the normal
symmetrical arrangement characteristic of the species, and the
irregularities and feeble development in the other half are
indications of an attempt to regenerate the disturbed symmetry.
It may be concluded that in the original individuals developed
from ova there were three cycles of mesenteries arranged
hexamerously, only those of the first cycle being perfect. The
reproductive elements were not sufficiently developed in any of
the individuals collected at Woods Hole to allow of a determina-
tion of their distribution, but in specimens collected at Plymouth
it was found that they were limited to the pairs of the second
and third cycles, 7.e., to the imperfect mesenteries. It may be
added that both oral and marginal stomata occurred in the
perfect mesenteries. Acontia are present, but ave not abundant.

The structural features, then, that may be supposed to be
characteristic of individuals that have not undergone fission are
(1) the occurrence of a well-defined fosse; (2) the absence of a
mesogleal sphincter; (3) the ocenrrence of but six pairs of
perfect mesenteries, which are also sterile, the reproductive
elements being limited to the imperfect cycles. But before
considering the significance of these features it will be well to
consider the geographical distribution of the species.

Verrill’s original description of Sagartia lucie (1898) was
based on individuals collected in Long Island Sound. He states
that his attention was first directed to it in 1892 and eXPIresses
his conviction that it did not occur in any quantity in that
region throughout the period 1865-1890. Furthermore it was

yl St

POSITION AND DISTRIBUTION OF SAGARTIA LUCIA. faa

stated to occur at Woods Hole in 1898, and Verrill collected there
extensively during the period 1871-1887 without finding it.
Parker (1902) added to these data observations on its occurrence
at Newport in 1895 and at Nahant in 1899, it having been
unknown in either of these localities before the dates mentioned,
and he also records its occurrence at Salem, Mass., in 1901, that
being apparently the northern limit of its distribution at that
time. The records brought together by Parker seem to show
clearly that the species has rapidly increased its distribution, at
first eastward and then northward, until in ten years after it
was first observed in Long Island Sound it had reached Salem.
I have nothing to add to these records, except to state that in
the summers of 1889 and 1890 I made special efforts to collect
representatives of all the Actinian species occurring in the Woods
Hole region and feel certain that, if Sagartia lucie had occurred
in that locality at that time in any considerable numbers,
I could not have failed, as I did, to find it.

_ But whence did it come to Long Island Sound? Verrill
suggests that it might have been brought from farther south on
the shells of oysters that.were annually brought north in large
quantities and deposited in the waters of the Sound, and this
possibility seems to find support in the following observations.
In 1887 I described as Sagartia pustulata a form found on dead
shells in shallow water in the vicinity of Beaufort, N.C. Its
specific name was suggested by the appearance given to the
column by the longitudinal and circular furrows that beset it, an
appearance that may frequently be observed in living examples
of S. lucie, which form it also resembled in its coloration,
except that the orange stripes of the column were lacking. This
may seem an important difference, but undoubted individuals of
S. lucie without the characteristic stripes have been observed
(Davenport, 1903; Walton, 1908). Unfortunately my prepara-
tions of S. pustulata are unsatisfactory and the preserved material
has long since disappeared, so that [ cannot determine the extent
of its similarity to S. lucie. This much, however, is certain, it
has no sphincter; but there were only slight indications of the
fosse, and it was not possible to determine the arrangement of
the mesenteries. The points of resemblance shown by the two
forms suggest their identity, but further observations are
necessary to establish this and with it the extension of the
distribution of the species so far to the south.

But the story of the distribution is far from being completed
with the consideration of the west coast of the North Atlantic.
Walton (1908) has placed on record the discovery at Plymouth
in 1896 of a form which Mrs. Davenport later identified with
S. lucie. Previous to the date mentioned it had not been found
in that locality, notwithstanding the extensive collections that
had been made there, and from the Millbay Docks, where it was
first observed, it has extended throughout the entire harbour and
is now one of the common forms inhabiting the district. The

734 MR. J. P. MCMURRICH ON THE SYSTEMATIC

evidence points to its being an intreduced form, possibly from
the other side of the Atlantic, but Walton suggests that neither
New England nor South Devon was its original home, it having
been introduced into both localities at approximately the same
time. 2

There is another possibility, or rather a strong probability.
In 1846 Mr. W. P. Cocks captured at St. Ives, on the north
coast of Cornwall, a form which he later (1851) described as
Actinia chrysosplenium. He sent drawings and a description of
it to Johnston, who included it in his ‘ History of the British
Zoophytes’ (1847), and he conferred the same favour on P. H.
Gosse, the drawing being this time the coloured one which is to
be found reproduced in plate vi. of the ‘Actinologia Britannica’
(1860). Gosse referred it, with some doubts, to the genus
Sagartia and added three additional localities at which it was
found by Mr. Cocks, all on the Cornish coast, but neither he nor
Johnston had personal knowledge of the form, their descriptions
being based on notes furnished by Cocks.

Cocks’s original figures do not suggest much resemblance to
S. lucie; they show relatively large circular spots scattered over
the column, looking like verruce, but representing really, as may
be gathered from the description, spots of bright yellow, the
ground-colour varying “ from a bright pea-green to the dark holly-
leaf tint.” But it is also stated that the yellow may be arranged
in stripes instead of spots, and in the figure reproduced by ‘osse
it 1s so represented, the similarity to the coloration of S. lucie
being thus very great. In Gosse’s figure, however, the tentacles
are shown as if very few in number and plump; Cocks’s original
figure, on the other hand, represents them as much more numerous
and tapering, quite as they are in S. lucie. Combining the two
figures, then, one would have a fairly accurate representation of
a S. lucie, both as regards its external form and its coloration.
It is true that no acontia were observed ; but S. lucie is somewhat
sluggish in emitting them, and the comparison that Cocks makes
in a letter to Gosse of the appearance of the column-wall to that
of a piece of “‘india-rubber when pierced with a pin” may well
have been suggested by the somewhat pustulous appearance
frequently presented by S. lucie. Indeed; making allowances
for the imperfections of descriptions of Actiniz written before
1860, I do not hesitate to express the opinion that when
examples of Cocks’s species, taken in his localities, are studied,
they will be found to be identical with 9. ducie.

If this opinion be correct then the species has been located on
the southern coast of England at least since 1846, and its
appearance at Plymouth in 1896 may have been merely an
easterly extension of its area of distribution, similar to what
occurred on the New England coast between 1891 and 1901.
This would seem a simpler explanation of the facts than the
supposition that it had been introduced from some distant
locality.

a a ee ee

» 6 = gee 4m

POSITION AND DISTRIBUTION OF SAGARTIA LUCIE. 735

I have sectioned a number of examples collected at Plymouth
and find that they agree in all essential points with those from
the New England coast. The irregularity of the mesenteries
was somewhat more pronounced, one individual (text-fig. 3), for
example, showing in sections through the aboral half of the
stomodeum only two pairs of perfect mesenteries, situated
opposite one another, one pair being directives attached to a
well-developed siphonoglyph, while in the other pair the muscle
pennons were on adjacent surfaces and there was no siphono-
glyph. The members of this second pair, however, were attached
to the stomodzeum opposite the middle of its longer axis, and,
although there were no mesenteries of younger cycles intervening,
it seems probable that they represent members of two different
pairs, IT. and VI., there having been a failure of regeneration,
after longitudinal fission, of the typical mesenteries. In the
intervals between the directives and each of the other perfect
mesenteries there were representatives of two other imperfect
cycles, symmetrically arranged, those representing the second
cycle bearing reproductive elements.

Text-figure 3.

VI ff

Diagram of the arrangement of the mesenteries in an individual from Plymouth.

Nor is the story of the distribution yet complete. In the
summers of 1909 and 1911 I collected at the Canadian Biological
Station at Departure Bay,on the east coast of Vancouver Island,
an Actinian whose superficial resemblance to S. lucie of New
England was most striking, the only noticeable difference being
a somewhat greater average size. They were of the same green
colour, the column was adorned with the same characteristic
longitudinal stripes of yellow or orange, and the tentacles showed
the same form and coloration. They were found on Jesse Island,
not far from the station, among the barnacles that closely
covered the face of a large rock, in such a situation that for hours
each day they were exposed by the tide. This was the only
location in which they were observed in these years; but ona
third visit to the station in 1912, I found them quite abundant
in slight depressions and crevices in the sloping rocks forming

736 MR. J. P. MCMURRICH ON THE SYSTEMATIC

the shore at the station, again well above low-water mark. so
that they remained exposed to the air for several hours at each
tide. Iam confident that they did not occur on these rocks in
the earlier years; had they then secured foothold upon them
they could hardly have escaped observation. The rock on Jesse
Island is not far from the station, but their extension to the
latter locality is of interest in connection with the migrations of
S. lucie on the New England coast and at Plymouth. Many
years ago Mr. Alexander Agassiz gave me a number of drawines
of Actinians that he had collected in the Gulf of Georgia in 1859
and among these there were two that were unmistakably
representations of the form under consideration, The drawings
were made from specimens collected on Galliano Island, and
furnish certain proof that the form has been in existence on the
Canadian Pacific coast for over sixty years at least.

Text-figure 4.

Diagrams showing the arrangement of the mesenteries in three individuals
from Departure Bay, British Columbia.

As has been stated, these Pacific forms resemble S. lucie in
their external form and coloration ; their anatomical structure is
also the same. They show a similar fosse, complete absence of a
sphincter, and a similar arrangement of the mesenteries. The
irregularities of the mesenteries were not so marked as in the
Plymouth examples, but were nevertheless quite evident, as may be
seen from the diagrams (text-fig. 4), which represent the arrange-
ment observed in three individuals. In each case the individual
was monoglyphie, with but a single pair of directives, and in one
case there were only four pairs of perfect mesenteries, in the
other two five pairs. Two cycles of imperfect and fertile
mesenteries were present, In one case arranged symmetrically
with regard to the four perfect pairs, but in the others showing
some irregularity, and in the example with four pairs of perfeet
-mesenteries representatives of a third imperfect cycle, without
mesenterial filaments or reproductive elements, occurred in two
of the interspaces. Acontia were present, but not abundant.

There can be no doubt as to the specific identity of the

POSITION AND DISTRIBUTION OF SAGARTIA LUCIA, Gan

Vancouver Island forms with those from New England and
Plymouth, so that we now know of three areas inhabited by the
species, separated from one another by wide intervals, two of
them by the North Atlantic Ocean and the other from these by
the whole width of the North American Continent. Further-
more, there is the certainty that the species has been ‘in
residence”’ on thé Canadian Pacific coast for over sixty years,
and if the suggestion be correct that it has been secondarily
introduced into the other two areas, the Pacific may have been
its original home. But Mr. Cocks’s discovery of his 4. chryso-
splentwm on the Cornish coast dates back seventy-five years, and
this lessens the probability of the introduction hypothesis. There
is another possibility, however, namely, that originally the species
had a cireumpolar distribution, like Meéridiwm senilis, Urticina
felina, and a number of other Colenterate forms, and that its
present areas of distribution are but separated remains of a much
larger area. If further observations should reveal its presence
on the Asiatic side of the Pacific the probability of this
suggestion would be greatly increased*; in the meantime it is
merely offered as an alternative to the introduction hypothesis.

But no matter what the original home of the species may have
been, the evidence is clear that in each of its known localities
it has in recent years more or less markedly extended its dis-—
tribution. The cause of this is also obscure. It scores largely
in favour of the introduction hypothesis, but is not necessarily a
proof of it. It may be a phenomenon in some respects and on
a lesser scal2 comparable to the migration of the Colorado
Potato Beetle many years ago, but as to the influences that
determine it in three widely separated areas I have no suggestion
to make.

To return now to a consideration of the systematic affinities of
the species. The possession of acontia marks it as a member of
the family Sagartiade, a family in which several subdivisions are
now recognized. From the subfamily Phelline it is excluded
by the thinness of the column-wall and by the fact that acontia
may be extruded through the wall, and it finds no place among
the Sagartiine, since it has not more than six pairs of perfect
mesenteries, these being also sterile. This leaves only the
Metridiinze and the Aiptasine for its reception, and it is very
doubtful if the separation of these two groups can be maintained.
For the Aiptasiine differ from the Metridiinze only negatively,
in the lack of a mesogloal sphincter, and since it may be
supposed that they are descendants of forms possessing that
structure, its absence in them is due tu a process of reduction
and is of less importance than the arrangement of the mesenteries.
Furthermore, I have found in diptasia (Heteractis) lucida faint
indications of a mesogleal sphincter, and, I may add, since this

* Since possibilities are being discussed I may suggest that perhaps the form

collected by Stimpson in Hong Kong Harbour and described by Verrill (1869) as
Sogartia lineata may be the Asiatic representative of S. lucie.

738 MR. J. P. MCMURRICH ON THE SYSTEMATIC

observation has not been confirmed by others (Duerden, Pax)
who have studied that species, that a quite distinct though feeble
mesogleeal sphincter occurs in A. pallida. In the arrangement
of the mesenteries the Aiptasias agree with the Metridinz, so
that their separation from that subfamily seems to be artificial
and unnecessary.

S. lucie agrees with the majority of the Aiptasias in lacking
a mesogloeal sphincter and in the possession of not more than six
pairs of perfect mesenteries, and, with the Aiptasias, should be
assigned to the subfamily Metridiine. That means that it is
not entitled to the generic term Sagartia. When Gosse (1859)
established the genus Sagartia he included in it all the forms
known to him to possess acontia, with the exception of Adamsia
palliata, which had already been assigned to a special genus by
Forbes. In the list of the forms belonging to the new genus,
Gosse placed viduata (effeta) first, and it might therefore be
taken as the type species, but later, in the ‘Actinologia Britannica’
(1860), Gosse proposed the subdivision of Sagartia into a number
of subgenera, retaining the original name for a group of forms
of which miniata is the type, while viduata is referred with
troglodytes (wndata) and parasitica (polypus) to a subgenus
Cylista. This complicates matters; for miniata and the other
forms that Gosse associated with it are, apparently, referable to
the older genus Cereus, and if this be the case Sagartia becomes
merely a synonym, unless vidwata (effeta) be accepted as its
type species. This seems the proper thing to do, for otherwise
the confusion that now exists in Actinian nomenclature would
become still worse confounded.

Sagartia viduata, or, to give it its more correct name, S.efeta L.,
has no fosse and a well-developed mesoglceal sphincter ; my pre-
parations from specimens collected at Plymouth do not allow of
certainty as to the arrangement of the mesenteries, though the
indications were that more than six pairs were perfect, but
Carlgren (1893) has shown that this is the case. Sagartia, then,
as is seemly, belongs to the subfamily Sagartiine, and our
S. lucie cannot be referred to it. What, then, is the proper
generic term for this species? It has some resemblance to
Aiptasia, but, lacking the characteristic double row of permanent
cinclides of that genus, it cannot well be included in it. It has
already been pointed out that there is a strong probability,
indeed, I believe it is more than a probability, that it is identical
with the A. chrysospleniwm of Cocks, and that form Gosse
recognized as a Sagartia, referring it to a special subgenus
Chrysoela. If my belief as to its identity is well founded,
S. lucie should: be known as Chrysoela chrysosplenium (Cocks)
Gosse. At all events it is not a Sagartia, nor can it be assigned
to any of the genera now recognized.

1847.
1851.

1855.

1860.
1869.

1887.

1893.
1898.

1900.

1902.

1903.

1905.

1907.

1908.

LOL.
1 OU.

T319:

POSITION AND DISTRIBUTION OF SAGARTIA LUCIA. 739

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vil.

Parker, G. H.—Pedal Locomotion in Actinians. Journ.
Exp. Zool. xxii.

D. W. Davis.—Asexual multiplication and regeneration in
Sagartia lucie Verrill. Journ. Exp. Zool. xxviii.

Hausman, L. A.—The Orange-striped Anemone (Sagartia
lucie Vervill). An Kcological Study. Biol. Bulletin,

XXXVI.

THE ANATOMY OF THE TONGUES OF THE MAMMALIA. 741

38. The Comparative Anatomy of the Tongues of the Mam-
_malia.—V. Lemuroidea and Tarsioidea. By CHARLES
F. Sonntag, M.D., F.Z.S., Anatomist to the Society.

[Received August 8, 1921: Read November 8, 1921.]

(Text-figures 66-69.)

ConTENTS.

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INTRODUCTION.

The literature of the tongues of the Lemuroidea and Tarsioidea
is extensive, but it deals mainly with the organs of taste and the
characters and significance of the sublingua. A few tongues have
been described in detail in monographs on certain animals *,
but little attention has been paid to some features which are of
value for purposes of classification. The present paper is based
on the examination of specimens in the Society’s Prosectorium
and the Museum of the Royal College of Surgeons, the latter
being described as Mus. R.C.S.

In the Lemuroidea and Tarsioidea the foramen cecum and
apical gland of Nuhn are absent. Some have no lateral organs,
and the lytta appears for the first time in the Primates. The
frenal lameilze are sometimes very complex.

The Sublingua.

The first description was contained in Tiedemann’s paper on
Stenops (18), but John Hunter had observed it previously and
likened it to the tongue of a bird. It.is horny, devoid of taste
buds, and fixed to the mandible by a frenum. Small muscle
bundles derived from the hyoglossus and genioglossus run into it.

It is lyrate (Lemuride and Indrisidee) or tongue-shaped
(Loriside and Galagide), and has a variable number of apical
denticles which differ in character in different animals; these are
absent in Zarsius and Chiromys, but the latter has a strong
projection on the centre of its anterior border.

* See papers 2, 3, 6, 7, 13, 14, and 16 in Bibliography.

742 DR. C. F. SONNTAG ON THE COMPARATIVE ANATOMY

Crests are present on the ventral surface in all species. They
are three in number in the Indriside and Lemuride, except
Microcebus, but only one is present in the Loriside, Galagide,
Microcebus, Chiromys, and Tarsius. The dorsal surface bears a
crest of variable size in the Loriside and Galagide, the most
pronounced one occurring in Perodieticus potto.

The consistence varies greatly. Im ali the Lemuroidea the
sublingua is horny, but in Tarsiws it is soft. In the Lemuride
and Indriside the entire sublingua is uniformly thiek, but in the
Loriside, Galagide, and Chiromys the central parts are thicker
and harder than the lateral parts. The horny covering is pro-
duced by an excessive thickening of the stratum corneum.

In Lemur the sublingua can be separated to a considerable
extent from the under surface of the tongue, but in Zarsius they
cannot be separated, for the sublingua is demarcated from the
tongue by a slight groove only. In the Loriside aud Galagide
the degree of mobility is intermediate, and in Chiromys it is
greater than that described in the writings of Gegenbaur (5) and
Pocock (15).

I have already summarised the differences between the sub-
lingue of the Prosimiz and Marsupialia (20).

Flower (4) and Pocock (15) believe that the function of the
sublingua is to clean the posterior surfaces of the incisor teeth ;
but others think it has no function, and regard it purely from
the point of view of phylogeny. I believe that the sublingua of
the Marsupialia is entirely a vestigial structure.

The Lyltta.

The tongues of the Lemuroidea and Tarsioidea have lytte
which are of two kinds:—1. The lytta of the tongue. 2. The
lytta of the sublingua.

The lytta of the sublingua is absent in Zemwr, double in
Tarsius, and single in Perodicticus and Loris; in all these forms
it is internal, but in Chiromys it forms a strong, external,
nodulated ridge ending anteriorly in a hook.

The lytta of the tongue is present in Perodicticus, Loris, and
Chiromys, but Gegenbaur and Owen missed 1t in the latter. It
is connected to the lytta of the sublingua by connective tissue.

I have nothing to add to existing accounts of the histology of
the sublingua and lytta.

Family LEMURIDA.

Genus Lemur.

In all species the apex is rounded and covered with small
conical and fungiform papille, but has no notch. The lateral
borders increase in thickness from before backwards and have
lateral organs at their posterior extremities. The oral part of the
dorsum possesses several wide, shallow transverse ridges and

sulci, and there is a smooth, non-papillary area in front of the

OF THE TONGUES OF THE MAMMALIA. 743

epiglottis bisected by the median glosso-epiglottic fold. Small
ridges may be present under the apex and send forward pointed
processes (text-fig. 66 C).

Text-figure 66.

The tongues of Lemur catta (A) and Nycticebus tardigradus (B); Cand D.
the under surfaces of two examples of the tongue of Lemur rufifrons.

The circumvallate papille are arranged in Y-formation in most
species. They are round or oval, and are surrounded by an
annular or lobulated vallum. They may be equal in size, or
the most posterior papilla may be the largest. The following
arrangements were seen by me or recorded by others :-—

1. Lemur catta: Six papille in Y-formation (text-fig. 66 A).
2. L. fulvus: Nine papille in Y-formation (text-fig. 67 A).
3. L. fulvus rufifrons: Nine papille in Y-formation (text-
fig. 67 E).
4. IL. macaco: Kight papille in Y-formation (text-fig. 67 F).
5. L. varius: Five papille in Y-formation.
6. L. mongoz: Nine papille in Y-formation (text-fig. 67 I).
7. ZL. mongoz: Four papille forming a double pair.
8. L. melunocephala: Nine papille in Y-formation (Minch).
9. L. rubriventer: A paix of papille (Munch).
Proc. Zoou. Soc.—1921, No. L. 50

744. DR. GC. F. SONNTAG ON THE COMPARATIVE ANATOMY

The papille may be concealed by the long conical papille on
the base of the tongue (text-fig. 67 C), or there may be a clear
zone between them (text-fig. 67 D).

The fungiform papille, which are small and numerous, stretch
right across the dorsum, but have the usual arrangement in
clusters and rows. The transverse ridges extend far forwards at
the expense of the apical cluster. On the ventral papillary zone
they are scanty and only form a single row,

Text-figure 67.

The vallate papille of the Lemuroidea: A-D. Lemur fulvus; EH. L. rufifrons ;
H-H. L. macaco; 1. L. mongoz; Ja-Jce. plan and elevation of papillee of
L. rubriventer ; Jd. fungiform papille of Lemar.

The conical papille (text-fig. 68 A) have the usual arrange-
ments in clusters and rows, and their points run in the usual
directions. Those on the oral part of the dorsum are small, but
those on the pharyngeal part are big and cylindrical] or scale-like.
These two groups are separated by a line of demarcation concave
forwards, lying a little in front of the vallate papillary area. The
points of the basal papiile are straight or recurved. This
arrangement is also present in Hapalemur, according to Beddard
(1), and Chiromys, but is absent in Microcebus, the Loriside,
Galagide, and Z'arsius.

The lateral organs (text-fig. 68 B-E) consist of lamineze and
sulci on the dorsum alone (L. macaco) or on the lateral borders
and inferior surface. The central lamine are larger than the
lateral ones, and the inner border of each organ is convex towards

_ OF THE TONGUES OF THE MAMMALIA. 745

the lateral vallate papille as in the Simiide. The relative
position of these papille to the organs varies in different species,
lying level with the central or posterior lamine. In Microcebus
the lateral organs are absent, and this condition, together with
the number and arrangement of the vallate papillee and state
of the conical papille, links the tongue to those of the Loriside
and Galagide, and distinguishes it from those of Zemw. The
numbers of laminz and sulci frequently differ in the two organs
of the same tongue, and the secondary sulci may be well marked

or indistinct.
Text-figure 68.

/s ( A
@
NN

@
AT tana alin,
Seo Se?

A. Conical papille of Lemur; B-E. lateral organs of ZL. eatta (B), L. fulvus (C),
I. macaco (D), L. mongoz (E); F. cross sections of the tongue of Chiromys
in the anterior (a) and middle (b) thirds: (s. sublingua, 1.1. lytta uf tongue,

ls. lytta of sublingua).

The ventral papillary zone is narrow and has many conical but
few fungiform papille, and the ventral mesial sulcus may contain
a mesial] crest; it is absent altogether in some examples of each
species, but present in others.

The sublingua (text-fig. 69) is triangular or lyrate, and the
apex is divided into a small group of slender denticles. The
edges are serrated in L, catia, L. macaco, and L. varius, and entire
in L. fulvus and L. fulvus rufifrons. The median crest is well
developed, but the lateral ventral crests vary in size. Between
the base of the sublingua and the frenal lamelle is a deep sulcus
which is sometimes V-shaped.

The frenum consists of two lax portions separated by the sub-
lingua. The first extends from the floor of the mouth to the
under surface of the sublingua, and the second runs from the

upper surface of the sublingua to the mesial ventral sulcus of
5()*
50

146 DR. C. F. SONNTAG ON THE COMPARATIVE ANATOMY

the tongue. The frenal lamelle, which are sometimes complex,
have the following characters :-—

1. LZ. catta: lamelle are two blunt rounded processes.

2. L. coronatus: \amelle form a parallel-sided flap.
3. L. macaco: triangular with rounded apex and lobulated
sides.

A. L, varius: two broad flaps with rounded anterior borders
and sharply cut mesial borders.

5. L. mongoz: lamelle form a parallel-sided flap.

The tongues of the species of Lemur resemble those of the
Chimpanzee and Gibbons in the following respects :—

1. Several vallate papillae form a Y.

2, The conical papille on the pharyngeal part of the tongue
are much larger than those on the oral part.

3. The lateral organs are situated at the posterior parts of
the lateral borders and are convex towards the lateral vailate
papille.

The tongues differ from those of the Orang-Outan and Man,
which have no large conical papille on the pharyngeal part of
the tongtie, and the vallate papille form aV. ‘These also have
the Apical Gland of Nuhn.

They differ from those of the Gorilla and Orang-Outan, whose
lateral organs form ladder-like patterns on the dorsum.

Genus Microcrsus.
Smirn’s Dwarr Lemur (JZ. murinus).

Three vallate papille form a triangle with the base anterior ;
the basal papille are small and circular, but the apical papilla is
large, round, and granular.

The conical papille imerease in size from the apex of the
tongue to the epiglottis, and those on the base are not dispro-
portionately large as in Lemur and Hapalemur.

Lateral organs are absent.

The sublingua is a flat horny plate with a row of denticles on
its anterior border.

One can see, therefore, that there is no resemblance in any of
the essential points between the tongues of Lemur and Micro-
cebus, but the latter resemble those of some of the Lorises,
Galagos, and Pottos, described below,

Family INDRISID&.

Genus [yprRis.

Flower (4) described the tongue of J. brevicaudatus as
follows:—“The tongue is long and narrow, with an obtuse
point at the apex, being less truncated than in Lemur. It is

had,

OF THE TONGUES OF THE MAMMALIA. TAG

24 inches long, and 3 inch broad at its widest point. The dorsal
surface is covered with a very close-set velvety pile of small
papille, apparently all of one kind. There are two small circum-
vallate papille, 2; inch apart, just in front of the attachments of
the palato-glossal folds.”

The sublingue resemble those of the Lemuride.

Text-figure 69.

I. ue K. L,

The sublingua of Lemur catta (A.B), L. fulvus (C), L. rufifrons (D), L. macaco
(HE); Nycticebus tardigradus (E.G); Loris gracilis (H); Perodicticus potto
(1); Hemigalago demidoffi (J); Galago crassicaudata (K); Tarsius spec-
trum (L).

Family Lorisip &.

yenus NYCTICEBUS.
THE Stow Loris (WV. tardigradus).

Three preserved specimens were examined, and their lateral
borders were seen to have no lateral organs. The apex, sulci,
and bare area in front of the epiglottis were similar to those of
Lemur.

Three vallate papille (text-fig. 66 B) form an equilateral tri-
angle with the base behind. All are circular, prominent, and

748 DR. C. F. SONNTAG ON THE COMPARATIVE ANATOMY

finely granular; the fosse are well marked and the vallums
appear as clear zones. They are conical on elevation, the bases
of the cones projecting beyond the fosse. In two specimens the
papillae are of equal size, but in the third (Mus. R.C.S8.) the apical
papilla is smaller than the basal ones.

The fungiform papille are not numerous, and form a dorsal
bounding zone on which they have the usual arrangement in rows
aud clusters, but the apical cluster is small; and they are few in
number on the ventral papillary zone.

The conical papille have the usval arrangement in clusters and
rows ; they increase in size from apex to epiglottis, and there is
no sharp transition between small conical papille on the oral and
large papille on the pharyngeal part of the dorsum as in Lemur
and Hapalemur.

Lateral organs are absent.

The sublingua differed in two of my specimens. In the first it
is tongue-shaped with a broad base whose angles are rounded,
and its area is half that of the inferior surface of the tongue.
The denticles on the centre of the anterior border are large, but
the more lateral ones are smaller. The central part of the sub-
lingua is thick and strong, but the lateral parts are thin. On
the ventral surface there is only one strong median ridge, and
the upper surface has a strong ridge which is received into the
median ventral suleus of the tongue. The frenum is long and
lax, so the sublingua is freely movable. In my second specimen
the sublingua has no rounded basal angles, the apical denticles
are more closely packed, there is only a faint ridge on the upper
surface, and the whole organ is more fixed to the under surface
of the tongue,

The frenal lamelle of the first specimen are broad, with bluntly
rounded apices, but they are narrow and pointed in the second
example. Supplementary lamellz were present in Pocock’s spe-
cimen (15).

The median ventral sulcus has no crest, but receives the ridge
on the upper surface of the sublingua.

Genus Loris.
Tur SuenDER Loris (Loris gracilis).

The conical tongue narrows rapidly from base to apex, and its
proportions are small.

Three vallate papille form an isosceles triangle with the apex
behind. All are circular, furrowed, and granular, and are sur-
rounded by prominent vallums. None are overlapped by conical
papille.

The fungiform papille differ from those of Vycticebus tardi-
gradus in that they extend right across the dorsum. They have
the usual arrangement in rows and clusters, but the transverse

OF THE TONGUES OF THE MAMMALIA. 749

rows extend far forwards at the expense of the apical cluster.
Those at the posterior part of the oral division of the dorsum are
large, hemispherical, polished, and close together. Few are pre-
sent on the inferior surface of the tongue.

The conical papille resemble those of Vycticebus tardigradus
in their mode of enlargement from before backwards, and differ
from those of Lema and Hapalemur. Most of the papille are
cylindrical, and are surrounded by zones of interpapillary
dorsum.

Lateral organs are absent.

The following account of the sublingua (text-fig. 69) 1s supple-
mentary to those of Tiedemann (18), Otto, Vrolik (16), and
Gegenbaur (5). It is leaf-shaped and w rapped round the convex
inferior surface of the tongue, so 1t 1s concavo-convex on cross
section. The edges are finely crenated and the apex is sharp.
The denticles are smaller, less numerous, and more closely packed
than those of WVycticebus tardigradus, but the characters of the
central and lateral parts are similar in both cases. The mesial
dorsal crest is not so pronounced as that of some examples of
WVycticebus, but is equal to that of others. Both species have
equally mobile sublingue, but the freedom is less than that of
Lemur.

The frenal lamelle are two small, rounded processes whose
edges extend postero-laterally for a considerable distance. No
accessory lamelle are present.

The median ventral sulcus resembles that of Vycticebus tardi-
gradus.

Genus PERODICTICUS.
BosmAn’s Porro (Perodicticus potto).

The tengue, which narrows slightly from base to apex, has no
apical notch and no mesial dorsal sulcus, but possesses a few
small, narrow transverse sulci in the posterior part of the oral
division of the dorsum.

The circumvallate papille.—Van der Hoeven (6) described and
figured three large papille in a triangle with the apex behind.
In the specimen in the Museum of the Royal College of Surgeons
there are three large excavated papille forming a triangle. In
my specimen there are three in a triangle, but the apical papilla
is divided into two large elongated parts enclosed in the one
fossa.

The fungiform and conical papilla do not differ materially
from those of Vycticebus tardigradus.

Lateral organs are absent.

The sublingua is tongue-shaped and has nine apical denticles
as in P, ibeanus. Its strong median ventral crest is bifurcated
posteriorly, and the dorsal ridge is larger and sharper than that
of any other species which I examined. The edges are devoid of

750 DR. C. F. SONNTAG ON THE COMPARATIVE ANATOMY

serrations and the degree of mobility is equal to that which
exists in Vycticebus and Loris (text-fig. 69),

Nussbaum (11) pointed out that the sublingua has a rod-like
core, which he termed the ‘“lyssa of the sublingua.”” It has a
connective-tissue sheath and consists of fat, cartilage, and
connective tissue. It is connected above to the lyssa of the
tongue which contains fat, connective tissue, and muscle fibres.

The frenal lamelle are two conical processes whose crenated
edges extend postero-laterally almost as far as the palato-glossal
folds.

The median ventral suleus has no crest, but receives the
prominent median dorsal ridge on the sublingua.

Family GALAGID&.

Genus GALAGO.
THe BusHy-TAILED GaALaco (G. crassicaudata). Mus. R.C.S.

The tongue was preserved, so no measurements were made.

Three vallate papille are arranged in the form of an isosceles
triangle with long sides. They are large, oval, and excavated,
the fosse are patulous, and the vallums appear as clear zones.
The basal pair are opposite the attachments of the palato-glossal
folds. Flower (4) also recorded three papille in the form of a
triangle. ;

The fungiform papille are small, rounded, and arranged in the
usual manner.

The conical papille increase in size from before-backwards,
and there is no sharp line of contrast between those on the oral
and those on the pharyngeal parts of the tongue as in Lemur and
Hapalemur. Flower states that they are thick on the base of
the tongue. They have denticulated points directed backwards.

The lateral organs are absent, the mesial ventral sulcus has no
crest, and the ventral papillary zone has no fungiform papille.

The sublingua (text-fig. 69), which is flat, horny, and tongue-
shaped, has a rounded anterior border bearing sixteen squat
sharp-pointed denticles. It covers rather more than half of the
width of the under surface of the tongue and half of the length
of the free part between the apex and frenum. ‘The sides are
parallel and. entire. Flower’s specimen had upturned edges and
a downwardly-directed apex which adapted its form to that of
the under surface of the tongue.

Genus HEmIcaLaco.
Demiporr’s Hremicataco (H. demadoffi).

The small conical tongues of both my specimens have three
vallate papille arranged in the form of a triangle with the apex

OF THE TONGUES OF THE MAMMALIA. (oil

directed backwards ; the basai papille are small and circular, and
the apical papilla is large, cireular, and granular, with a patulous
fossa.

The fungiform papille are not numerous, but stand up
prominently. They have the usual arrangement, but the apical
cluster is small. They form a well-marked row of closely-set
elements on the posterior third of each lateral border of the
tongue. None are present on the inferior surface of the tongue.

The conical papille give the surface of the tongue a finely-
granular appearance. ‘They increase in size trom before back-
wards, and there is no sharp contrast between the papille on the
oral and pharyngeal parts of the tongue as in Lemur and
Hapalemur. When the tongue is dried they stand up promi-
nently and reach a higher level than the summits of the
fungiform papille. They form a narrow ventral papillary zone.

Lateral organs are absent.

The sublingua (text-fig. 69), which is flat, horny, and tongue-
shaped, has a rounded, denticulated, comb-like anterior border,
and nearly parallel, entire edges. It has a well-marked median
crest, but no lateral ridges, and it covers a larger area of the
under surface of the tongue than that of the Galago.

The frenal lamelle are small and pointed, and the mesial
ventral suleus does not contain a crest.

Pocock (15) described nine apical denticles on the sublingua of
Galago monteire.

Family CurroMYID&.

Genus CHIROMYs.
Tur Ayr-Ave (C. madagascariensis).

The following description is supplementary to the existing
accounts by Owen (18) and Peters (14) :—

Measurements.— Total length 4:6 cm.; oral part 3:1 cm.;
pharyngeal part 5 em.; width between the lingual attachments
of the palato-glossal folds 1:6 em.

The apex is thick and rounded, but has no notch, and the
massive lateral borders are devoid of lateral organs. Mesial
dorsal and mesial ventral sulci ave absent.

Three circumvallate papille are arranged in the form of a
triangle with the apex behind. All are circular and granular,
and surrounded by nodulated vallums. Owen described two
papille, about two lines apart, lying about an inch and a half
behind the apex of the tongue. Peters recorded three papille
arranged in a triangle, and figured nodulated vallums.

The fungiform papille are not numerous. They extend right
across the dorsum, but are scanty in the middle line. They have
the usual arrangement, but the apical dorsal cluster is small.
One large papilla bisects the base of the vallate papillary triangle.

152 DR. GC. F. SONNTAG ON THE COMPARATIVE ANATOMY

None are present on the inferior surface of the tongue. They
are smooth or granular, and are hemispherical or pedunculated.

The conical papille have the usual arrangement. They are
closely packed and exhibit considerable mutual compression.
Some are low and cylindrical, but others stand up prominently.

The papille on the oral part of the tongue are smaller than
those on the base, and a line of demarcation, concave forwards,
separates the two groups. The tongue resembles those of Lemur
and Hapalemur in this respect, and differs from those of the
Lorises, Pottos, and Galages. There is also an area devoid of
papille in Paierh of the epidlottis ; this is bisected by the median
glosso-epiglottic fold.

The sublingua has been described by Owen (13), Gegenbaur (5),
and Pocock (15), but several details are omitted, or not sufficiently
emphasised, in their accounts.

It is a flat horny plate with entire lateral margins and
rounded anterior border with a central projection, whose point
marks the place where the strong, denticulated median ventral
ridge bends downwards in a hook. This crest increases in depth
from behind-forwards, and its hook hes 9 mm. posterior to the
apex of the tongue.

It was shown in the descriptions of the sublingue of the
Lorises that there is a thick central strip and two thin lateral
parts. The same parts are present in Chiromys, but differ from
those of the Lorises in that the central part is adherent to the
under surface of the tongue, but the lateral parts are free; in
the Lorises, on the other hand, both parts are free. Gegenbaur
and Pocock both describe a free lateral margin, but do not give
any idea of the extent of the free part. Jn my specimen the
total width of the sublingua is 1-2 em.; the central firm adherent
part is *6 em. wide, and each free lateral part is °3 em. wide. A
probe can be passed for a considerable distance between the
under surface of the tongue and the sublingua. No denticles
project from the anterior border.

Pocock (15) deseribed the two small glandular (?) pockets on
each side of the frenum, and pointed out that the frenal lamelle
are narrow.

Gegenbaur (5) described the horny nature of the sublingua,
but did not show how the thickness varies in different parts.
Although one thinks that the central parts are thicker than the
lateral parts, when the tongue is entire, one sees that the reverse
is the case when sections ave made. The apparent thickness of
the central part is due to a downward projection of the lingual
muscles to which the central part of the sublingua is closely
applied (text-fig. 68 F).

The Lytta —The tongue possesses a strong median ridge on
the under surface of the sublingua, which Owen termed the
“lytta.” He did not describe sections of the tongue, for a well-
developed lytta is present close to the sublingua. Gegenbaur
made the same omission, but Nussbaum (11) recorded its

OF THE TONGUES OF THE MAMMALIA. 753

presence. It is united to the upper surface of the sublingua in
the median part of the latter. ‘lhe condition resembles that of
Perodicticus, in which there are lingual and sublingual lytte.
In the latter species both lytte are internal. The position of the

lytta in different parts of the tongue is seen in text-fig. 68 F.

Family TaRrsiip 4.

Genus TARSIUS.

Tue Tarsier (7’. spectrum).

The classical work of Burmeister (2) and the subsequent
papers by Gegenbaur (5) and Pocock (15) have dealt with most
points, so the. work of any observer is now comparatively limited.

In the specimen in the Museum of the Royal College of
Surgeons the shape of the tongue and the characters of the
vallate, fungiform, and conical papille are similar to those
described by Burmeister. That author does not mention that
the lateral organs are absent. There is no great contrast between
the conical papille on the oral part of the tongue and those on
the base.

The sublingua is soft and defined from the under surface of
the tongue by a groove only ; it is the most adherent sublingua
among the Prosimie. The median ventral rod does not termi-
nate in a knob as in Pocock’s specimen, but its free anterior
extremity is rounded (text-fig. 69), and is slightly turned
downwards as in Chiromys; as the specimen was preserved, I
am unable to say whether this curving of the ridge was produced
by the alcohol.

The frenal lamella are two conical tapering processes, and
differ from the conditions recorded by Burmeister and Pocock.

Summary and Conclusions.

1. The tongues of the species of Lemur resemble those of the
Chimpanzee and Gibbons in the Y-formation of their vallate
papille, their convex lateral organs, and the contrast between
the large conical papille on the pharyngeal part of the tongue
and call ones on the oral part. They differ from the tongues
of Man and the Orang-Outan in that the latter have no large
conical papille on the base of the tongue, and their vallate
papille form aV. They differ from the tongues of the Gorilla
and Orang-Outan, whose lateral organs form ladder. like patterns
on ale dorsum of the tongue.

The tongue of JLicr Daehue differs in the essential features
a those of the species of Lemur, and resembles those of the
Lorises, Galagos, and Pottos.

3. The tongues of the Loriside and Galagide differ from those
of Lemur in so many features that they are important for
“purposes of classification. Adding them to other characters

754 DR. C. F. SONNTAG ON THE COMPARATIVE ANATOMY

deseribed by Pocock, Forsyth Major, and others, we obtain the
following comparison :—

Lemur. Loriside and Galagide.

1. Vallate papille form a Y. . Vallate papille form a tri-

angle.

2. Lateral organs convex. 2. Lateral organs absent.
Conical papille small on the 3. Conical papillae increase
oral part of the tongue gradually from the apex
and large on the base. of the tongue to the epi-
glottis.

4, Sublingua triangular or 4. Sublingua tongue-shaped,
lyrate, with apical den- with. denticles along its
ticles. anterior border.

5. Urethra opens above tip of 5. Clitoris traversed by urethra.
clitoris.

6. Ectotympanic annular or 6. Ectotympanic external to
horseshoe-shaped and in- bulla, of which it forms
side bulla. the outer wall.

I have not had the opportunity of examining the tongues of
the Indriside, but published accounts and illustrations allly them
to those of Lemar.

4. The tongue of Chiromys has its own characteristic sub-
lingua, which has a larger area free from the tongue than
existing accounts lead one to expect. It has no lateral organs,
but its basal conical papille are large. It has two or three
vallate papille. Its characters resemble those of Zemzz on the
one hand and those of the Loriside and Galagide on the other.

5. The tongue of Zarsius spectrum has the most adherent
sublingua. It stands by itself in the consistence and size of the
sublingua and the characters of its vallate papille.

6. The frenal lamelle are frequently very complex, and
supplementary lameile are present in Vycticebus tardigradus.

7. The foramen cecum and Apical Gland of Nuhn are absent.

8. The lytta is present in several forms, and is frequently
accompanied by a central axis, or lytta, of the sublingua. In
the other Primates this is absent.

Bibliography.

1. Bepparp, F. E.—‘‘ On some Points in the Structure of
Hapalemur griseus.” P.Z.S. 1884, p. 391.

Burmeister, H.—‘ Beitriige zur niheren Kenntnis der
Gattung Zarsius.” Berlin, 1846.

Campren, F. A. W. van.—-Verhand. der Konig. Akad. van
Wetenschappen, Amsterdam, 1859.

Firowrer, W. H.—‘‘ Lectures on the Anatomy of the Organs
of Digestion of the Mammalia.” Med. Times and Gazette,
1872

O00) Se lo aon

10.
abt:
12.
13.

14.
15.
16.

17.
18.
19.
20.

lo |

OF THE TONGUES OF THE MAMMALIA. (3:

. GucENBAur, U.—‘* Uber die Unterzunge des Menschen und

der Siiugethiere.” Morph. Jahrb. 1884, p. 428.

. Horven, J. van der.— Bijdrage tot de Kenntnis van den

Potto van Bosman.” Amsterdam, 1851.
Houxtey, T. H.—‘“On the Angwantibo (Arctocebus cala-
barensis).” Proe. Zool. Soc. 1864, p. 314.

. Kinema, P. H.—‘ Eenige Vergelijkend-Ontleedkundige

Aanteekeningen over den Otolicnus Peli.” Leyden, 1855.

Mecxet, J. I'.—System der vergleichenden Anatomie.
Teil 4. Halle, 1829.

Mitncun, F.—‘ Die Topographie der Papillen der Zunge des
Menschen und der Siiugethiere.’” Morphol. Arb. 1896.

Nusspaum, J.—Anat. Anz. pp. 551-561, 1896; and pp. 345-
358, 1897.

OppreL, A.—Lehrb. der vergl. mikr. Anat. vol. iii, pp. 251—
270 and 391-393. .

Owen, R.—‘* Monograph on the Aye-Aye.” Trans. Zool. Soc.
London, vol. v. p. 33, 1863.

Perers, W.—‘ Uber die Saugethiergattung Chiromys.”
Abh. der Konig]. Akad. der Wiss. zu Berlin, 1865.

Pocock, R. I.—*‘ On the External Characters of the Lemurs
and of Tarsius.” P.Z.S. 1918, pp. 19-53.

Scureper, J. L. C., vAN DER Koxnk, and W. VroiimK.—
Bijdragen tot de Dierkunde, uitgegeven door het Konin-
klijk Zoologisch Genootschap “ Natura Artis Magistra.”’
Kerste Deel. Amsterdam, 1848-54.

Sonntac, C. F.—Papers on the Tongues of the Primates.
P.Z.S. 1921, pp. 1-29, 277-322, 497-524.

TrepEMANN, F’.—Arch. f. Anat. u. Physiol. vol. v. pp. 348-356,
STO:

Tuckerman, F.—Journal of Morphology, vol. vii. pp. 69-94,
1891.

Sonnrae, C. F.—P. Z.8. 1921, pp. 547-577.

ROUT AN
ie ee

at
BN:
ny)

sel ;

HE ANATOMY OF THE TONGUES OF THE MAMMALIA. 757

39. The Comparative Anatomy of the Tongues of the Mam-
malia.—VI. Summary and Classification of the Tongues
of the Primates. By Cuaries F. Sonntag, -M.D.,
H.Z.S., Anatomist to the Seciety.

[Received August 8, 1921: Read November 8, 1921.]

INTRODUCTION.

In the five preceding papers of this series I described the
structure of the tongues of the Primates and gave a short
account of the phy siolog gy of the lingual glands. I also drew
attention to a number of points bearing on “the phylogeny of the
tongue. In the present paper I have summarised these observa-
tions and pointed out how the lingual characters support systems
of classification which have been based on external and skeletal
characters.

SUMMARY.

Pigmentation.—Most of the pigmented forms are included in
the Cebide. The colour, which is yellow, green, brown or bluish-
black, especially the latter, is uniformly distributed all over the
dorsum, or the pigmented an colourless parts may form definite
patterns ; in Ateles grisescens, for example, there is a white cross
on a brown background, The vailate and fungiform papille may
be pigmented (e. g. in Hylobates lar) or colourless, but the
lateral organs and central parts of the inferior surface of the
tongue are always white. If several examples of each pigmented
species are examined it will be seen that the colour varies
considerably, so is of no value for purposes of classification.
Most specimens of Cercopithecus patas, for example, have reddish-
yellow tongues, but some tongues are colourless, and the fungi-
form papille of Cercopithecus tantalus ave yellow or colourless.

The bluish-black colour persists longest in preserving fluids.

Form and Proportions.—Most tongues are conical, but a few
are oval, spatulate or rectangular; and shape is of no value for
comparative purposes.

In most of the Primates the tongue is long and comparatively
narrow, but in Anthropopithecus troglodytes and Gorilla gorilla
the tongue of the young animal is relatively wider than that of
the adult. I did not, however, see such variations in Cerco-
pithecus patas or Macacus sinicus, of which I examined very
young and adult examples.

Cunningham showed that the tongue of Simia satyrus resembles
that of Homo most closely in the relative proportions of length
and width.

The Apex is round, truncated or pointed, and may or may not
have a notch, but the latter is usually absent from the fresh
tongue, It is comparatively smooth, or roughened and tubercu-

758 DR. CG. F. SONNTAG ON TILE COMPARATIVE ANATOMY

lated by sinall conical and fungiform papille. The relative
quantities of fungiform papille on and immediately behind the
apex vary; in the Simiide and Cercopithecide they are
numerous and thickly clustered, but in the other families they
are few and discrete.

In some species of Zemur the apex exhibits a number of sharp-
pointed processes prolonged forwards from ridges on the inferior
surface of the tongue, but these vary even in different examples
of each species.

Sulci and Ridges.—Few fresh tongues have median dorsal
sulci on the oral part of the dorsum, but many preserved
specimens do; and I observed a well-marked median sulcus on
the pharyngeal part in Anthropopithecus troglodytes alone. The
most pronounced mesial dorsal sulcus which I observed occurred
in Mystax ursulus. Median dorsal ridges are present in some
Lemuroidea and Hapalidee.

Wide, shallow transverse sulci separating low wide transverse
ridges are present on the fresh tongues of Lemur catia and
Hapale jacchus; and fine narrow sulci are seen in Gorilla gorilla
and Perodicticus potto. Some of the fine sulci and ridges remind
one of the patterns of fissures and ridges on the finger-tips.

The median ventral sulcus is present in most tongues, and is
never an artefact induced by preserving fluids as is the dorsal
one in bottled specimens. It is narrow and deep, or wide and
shallow, and it frequently opens posteriorly into a triangular
fossa which recieves the upper end of the frenum. In some of
the Lemuroidea it receives the median dorsal crest of the
sublingua, and it recesses the crest on the dorsal surface of the
frenal lamella of Cebus fatuellus.

In Gorilla gorilla, some species of Hylobates and some of the
Lemuroidea it contains a fixed crest; and it has been stated that
the crest is a remnant of the sublingua, but the presence of both
these structures in some Lemurs would seem to disprove that
theory.’ .

Lateral Borders.—The edges of the tongue are sharp, or full
and rounded, and increase in thickness from before backwards in
most species. Those of Chiromys madagascariensis are more
massive in proportion to the size of the tongue than those of any
other species of the Primates. Of the Pithecoids Simia satyrus
and Anthropopithecus have the most massive lateral borders.

In Tarsius spectrum, Microcebus, Chiromys, and all Loriside
and Galagide the lateral borders are devoid of lateral organs
and, as I hope to show in a future paper, this has an important
bearing on phylogeny. In Gorilla gorilla and Simia satyrus
only small parts of the lateral organs are found on the lateral
borders, but in all other Primates the greater part is found
there.

‘he conical and fungiform papille on the lateral borders are
arranged in vertical rows and the points of the former are
directed backwards.

OF THE TONGUES OF THE MAMMALIA. 159

Distribution of the Papille.—In all Primates except some
specimens of Simia satyrus and Symphatangus syndactylus, some
Lemuroidea and Homo, papille cover the entire dorsum, apex,
lateral borders, and a bounding zone of the inferior surface. In
all these species there are smooth non-papillary areas on the base
of the tongue, and in the Lemurs the area is bisected by the
median glosso-epiglottic fold.

The ventral papillary zone varies greatly in width, and its
characters ave of limited value for purposes of classification. It
is wide in the Simiide, Cercopithecidee, and in Cebus, Ateles, and
Lagothrix, but is narrow or absent in all other Primates. In
species with a wide zone the conical and fungiform papille are
numerous, but in those with a narrow zone there are few or no
fungiforms. In Goriila gorilla many of the fine transverse sulci
on the dorsum cut the lateral borders and run inwards on the
ventral papillary zone.

The Circumvallate Papille.—In my paper on the tongues of
the Cercopithecide I showed that all the papillary patterns
present in the family will be seen in most species if sufficient
examples of each ave examined. The whole series may quickly
appear or it may be necessary to examine many. I am not
prepared, however, to apply this rule to any other family except
the Hapalidz, as insufficient specimens have come to hand.

In the following list of papillary patterns, P means two
. papille forming a pair, and D.P. indicates four papille arranged
in a double pair. The Y type means that there are several
papillae present and does not include forms in which the four
papille of a double pair are arranged in a Y.

Family. Patterns of vallate papille.
PELL A ee ye elke se hia DP Leia Ay A
Cercopithecide ......... A Vi op ln DP
Celiedee reales weeds Ne ene ey yh do abe
Hlanpabiclesa My s503 3: Xe
Wgenaumrcbestasi ale. Goh an NG [SS ee
LORUsiGeey Henkes. sae Ne ;
Gala gileagrey ise. Hecs see iNew ee
@hivouiyides) 45) sgh 3) 42: TANCE OLE
Damen lee tigated saeco aetsicr Three papille arranged in a line.

One can see, therefore, that the Simiide and Lemuride are
the only families whose tongues possess more than four papillee
arranged in a Y, and it will be shown later that they differ from
all Scher Primates i in other respects.

The papille are round or oval on plan and conical on elevation,
with the bases of the cones projecting beyond the vallums.

The Fungiform Papille stretch right across the dorsum, or are
absent from the centre thereby forming a dorsal bounding zone.
They form a cluster behind the apex, but are arranged in rows

Proc. Zoou. Soc.,— 1921, No. LI. 51

760 DR. C. F. SONNTAG ON THE COMPARATIVE ANATOMY

of varying degrees of obliquity behind that. The apical cluster
is large in the Simiide and Cercopithecide, but in all other
families the transverse rows extend far forwards at its expense.
In the majority of tongues with large apical clusters there are
many fungiform papille on the ventral papillary zone,

It has been shown by Tuckerman that the fungiform papille
of the apical cluster have many taste-buds.

In some specimens of Anthropopithecus troglodytes there is a
row of prominent fungiform papillae occupying the mid-dorsal
line of the tongue and replacing the median dorsal sulcus.

It is sometimes impossible to tell whether a papilla at the
posterior part of the oral division of the dorsum. is a large
fungiform or small vallate form, for fossa and vallum may ihe
indistinguishable even through a strong lens. Histological
examination is the only proof, The funeiior m papille may have
no taste-buds or these, if present, lie on ine free upper surface of
the papilla; in the vallate papille, on the other hand, the taste-
buds never le on the free upper surface of the papilla, but are
deep down on one or both sides of the fossa.

In some tongues there are more fungiform papille than are
visible to the naked eye, for some are entirely concealed by
overhanging conical papille (e.g. Anthropopithecus troglodytes).

The fungiform papille on the ventral zone may be thickly
clustered at the apex of the tongue and scanty further back, or
vice versa, and the examples, and the rows in which they are
arranged are close together or discrete.

From the point of view of classification the most important
features are the size of the apical dorsal cluster and the presence
ov absence of the fungiform papille on the ventral papillary zone.
‘Although their presence or absence in the centre of the oral
part of the dorsum varies greatly, it is not a character of
sufficient distinctness to be of “value for purposes of classification.

The Conical Papille vary im size and arrangement in the
different families, and there are three types of the formere=

1. The papille on the pharyngeal part of the tongue are
small :—--Homo, Sema satyrus.

2. The papille on the oral part of the tongue are compara-
tively small, but those on the pharyngeal part are large and
prominent :— Gor ‘lla aorilla, Anthropopithecus troglod Wee all
species of Hylobates, all species of Lemur, Chiromys madagas-
cariensis, and Tarsius spectrum. ‘his type also occurs in some
lower Mammalia. ‘

3. The papille gradually increase in size from the apex of the
tongue back to the epiglottis :—Cercopithecidee, Cebide, Hapalide,
Loriside, and Galagide. This arrangement is also ‘present im
Microcebus in which the vallate papille form a triangle, so the
papille are of value for distinguishing it from Lemur.

The arrangement of the papille distinguishes most of the
Cebide from all other families. In the latter they form a cluster
behind the apex and rows of varying degrees of obliquity behind

OF THE TONGUES OF THE MAMMALIA. 761

that, but in the Cebide they are dotted irregularly all over the
dorsum.

The tongues with smooth non-papillary areas on the pharyngeal
part of the dorsum have been enumerated above.

In most Primates the points on the oral part of the dorsum
look backwards or backwards and inwards, but in some specimens
of Simia satyrus and Cercopithecus cethiops those on the centre of
the orai part run in all directions.

In the Cebidee, Hapalide, Lemuroidea, and Tarsioidea the
conical papille are mostly pointed, and cylindrical and globular
forms are uncommon. In the other families there is a good
admixture of all types. ‘

The Lateral Organs present numerous forms and are of value
for purposes of classification :—

1. Organs absent :—AMicrocebus, Chironvys, Tarsius, the Lovi-
side and Galagidee.

The lamine and sulci form ladder-like patterns on the
dorsum of the tongue, and only their outer ends cut the lateral
borders of the tongue :—Gorila gorilla and Simia satyrus.

3. The organs are convex towards the lateral vallate papille:
Anthropopithecus troglodytes, Hylobates (all species), Lemur (all
species ).

4. There are rows of short lamine and sulci on the lateral
borders of the tongue :—Cercopithecide,

. The inner borders of the organs are concave towards the
lateral vallate papile :—Cebide.

6. A few faint irregular laminz and sulci are present on the
lateral borders of the tongue :—Hapalide.

In very few specimens did I find an equal number of laminze
and sulci in the two organs of the same tongue. But one must
be careful net to mistake simple folds of the mucosa at either
end of the organs for lamine. Histological examination is the
only true test in doubtful cases, for it reveals the presence of
taste-buds in the true lamine.

In a few species of Cercopithecus one may find fungiform
papillee situated on the lamine of the lateral organs.

The degree of protrusion of the lamin and depth of the sulci
vary not only in different animals but in several examples of
each.

The Lingual Glands are divided into apical and basal parts, but
the former is most variable.

The Apical Gland of Nuhn is present only in Homo and Simia
satyrus. Oppel believes that it is a piece which has become cut
off from a forward prolongation of the basal glandular mass.
The basal mass in the Marsupialia sends forwards two prolonga-
tious of variable stoutness, and it is possible that the Apical
Gland of Nubn has been cut off from one of these. If that were
so it would support Gegenbamr’s view that the tongues of the
Primates have evolved from those of the Marsupialia.

The serous and mucous glands on the pharyngeal part of the

762 DR. C. F. SONNTAG ON THE COMPARATIVE ANATOMY

tongue are developed to an equal degree in the Primates, but
the degree of development of the entire basal mass varies.

The pharyngeal part of the tongue possesses a variable degree
of development of lymphoid nodules, and a variable number of
orifices of pits and the ducts of glands; and the latter are
of value for distinguishing the tongues of the various genera of
the Cercopithecide from one another. Orifices are visible in all
Cercopithecide, but are absent from most Cebide, so are of
classificatory importance when taken in conjunction with the
characters of the lateral organs and mode of arrangement of
the conical papille. The following are the Cheever of the
orifices in the Cercopithecidee :—

Genus Presbytes :—Orifices larger and more patulous than in
any other genus, and lie in the centres of large round glandular
areas. The salivary glands are enormous.

Genus Cercopithecus :—Orifices well-marked, discrete and not
patulous, and no white areas present around them.

Genus MZacacus :—Orifices like pin points.

Genus Cercocebus:—Small duct orifices present at the sides of
the base of the tongue.

Genus Papio ._Targe duct or Unees present at the sides of the
base of the tongue. :

In many of the Cebide the sour on the base form zones,

with concave anterior borders, in front of the epiglottis, but 1 did _

not observe a similar condition in any other family.

The Prenal Lamella varies greatly in the Primates, and appears
to belong to the floor of the mouth rather than to the tongue.
It varies even in several examples of the same species, so is of
limited value for purposes of classification.

In Homo, Simia satyrus, and some specimens of Symphatangus
syndactylus it appears as two simple folds over Wharton’s Ducts ;
but Mr. Pocock informs me that he observed two well- marked
processes in a young Simia satyrus. IJ cannot believe, however,
in conformity with niy observations on the tongues of other
animals, that reduction in the lamella is a change due to
advancing years. In other species the lamella appears as a
tviansulat or tongue-shaped process with an entire or divided
apex ana with both edges entire, serrated, or bearing small sharp
points. The sharpest and most prominent points on the edges
occur in Cercopithecus preussi and some specimens of Cebus

fatuellus.

The Orifices of Wharton’s Ducts vary in their position on the
lamella. as follows :—

1. On the upper surface— Gorilla gorilla.

On the apex— A nthropopithecus troglodytes.

3. On the under surface— Pithecia satanas.

The apical position is the commonest form, however.

Tuckerman descvibed a rich nerve plexus with peripheral nerve
endings in the lamella of Ateles ater, and Gegenhaur considered
that the endings were tactile in function. As this condition has

Ne a oe See en ee

OF 'THE TONGUES OF THE MAMMALIA. 763

not been found in any other tongue there is insufficient material
in which to work out its phylogenetic significance.

The Subtingua.—The various Lemuroidea are differentiated
from one another by the shape, mobility, and characters of the
crests and denticles as follows :—

1. Sublingua triangular or lyrate, has three ventral crests,
and-is very mobile. Phe apex 1s aivatted into a brushwork of
denticles :— Lemur, Hapalemam, Indriside.

Sublingua a large flat plate adherent to the under surface
of the tongue by its central parts; no denticles present, but a
strong keel-like ridge on its under surface projects forwards in
the middle line :—Chiromys.

3. Sublingua large, tongue-shaped, but not quite so free as that
of Lemur. ‘Vhere is only one median ventral crest, bat’ the
dorsal surface has a cress of variable prominence. This crest, the
characters of the denticles, and the variations in the consistence
of various parts of the sublingua ure important :—-

In Microcebus the sublingua is uniformly thick, the median
dorsal evest is slight, and the denticles are moderately long.

In WVyeticebus and Loris the central parts of the sublingua ave
thicker than the lateral parts, the median dorsal crest is small
and the denticles are of moderate length ; they are discrete in
the former and close in the latter, but there is no essential
difference between the sublinguz of these genera.

In Perodicticus the median ventral crest is bifurcated pos-
teriorly, the median dorsal crest is very prominent, and the
denticles are long and slender.

In Mecrocebus, Galago, and Hemigalago the anterior border of
the sublingua is broad, but in Loris and Vycticebus it is more or
less pointed.

The Plice Pimbriate of the Simiide are derived from the sub-
lingua by a process of phylogenetic reduction, and I showed that
the plicee of Anthropopithecus troglodytes with the intervening
piece of mucosa form a soft triangular field resembling a sub-
lingua; this is even more marked in the tongue of the new-born
child, as deseribed and figured by Gegenbaur. The plice of
Phascolurctos cinereus, however, do not beund such an svea.

The tongues of the Gibbons, Cercopithecide, Cebide, and
Hapalide illustrate the ultimate stage of reduction, for no traces
of the sublingua or plica are present as a rule in the extra-uterine
stage. In the fetal Gibbon, as shown by Deniker, there is a
well-marked sublingua; and T observed two minute plicee in a
young Cercopithecus patas, so it is probable that the feetuses of all
Primates have sublingue.

If one examines a series of human tongues at different ages,
one finds that the new-born child has well- -leveloped plicee or an
actual sublingua provided with taste-buds. As age advances the
buds disappear and the plicee diminish in size. These taste-buds
_ probably account for the more acute sense of taste in the child.
Experimental methods also demonstrate that the sense of taste

764 DR. C. F. SONNTAG ON THE COMPARATIVE ANATOMY

elicited by applications of solutions to the centre of the oral part
of the dorsum diminishes as age advances.

This atrophy of structure following loss of function may have
played an important part in the reduction in and ultimate loss of
the primitive Mammalian tongue.

The Lytta—Vwo forms are to be recognised :—The lytta of
the tongue and the lvtta of the sublingua, And Gegenbaur
showed that the latter, when present, appears in one of two
forms. In Stenops it forms a str ong central supporting rod, but
in Tarsius ib is double. In Lemur itis absent altogether. Owen
described the keel on the ventral surface of the sublingua as the
lytta, but sections through the tongue show a well-marked lytta
inside ; it is connected in the middle to the sublingua.

CLASSIFICATION.

The structures which are of value for purposes of classification
are :—

1. The number and arrangement of We vallate papille.

2. The arrangement of the conical papille on the oral part of
the dorsum.

3. The mode in which the conical papiile increase in size from
before backwards.

4, The size of the cluster of fungiform papille on the dorsum
behind the apex of the tongue.

5. The characters of the lateral organs.

6. The width of the ventral papillary zone, with the number,
characters, and arrangement of its papille.

7. The presence or absence and characters of the lytte of
tongue and sublingua.

8. The presence or absence of the apical gland of Nuhn, and
the characters of the orifices of ducts and pits on the pharyngeal
part of the dorsum

9. The sublingua and plice fimbriate.

These features are of specific value in the case of all the
Simiide, except Hylobates, and many of the Lemuroidea, but
they are of generic value ouly in all otber Primates. They lend
additional weight to some schemes of classification Lased on
external and skeletal characters; in some cases they help us to
settle points which are not supported by a large body of very
conclusive evidence.

The tongue of Simia satyrus resembles that of Homo most
closely. Both have rounded apices, and their form and general
proportions are similar. They have no large conical papille on
the base of the tongue, and their vallate papille forma V. The
frenal lamelle are two small folds over Wharton’s Ducts, and
their plice fimbriate are equally developed, but smaller than
those of other Primates; and they are the only Primates possess-
ing an Apical Gland of Nuhn. In Simia satyrus the lateral

OF THE TONGUES OF THE MAMMALIA. 765

organs are better developed, and the conical papiile on the oral
part of the dorsum include more pointed forms, but Homo alone
of all the Primates possesses a foramen cecum.

The ladder-like patterns formed by the lateral organs distin-
guish Simia satyrus and Gorilla gorilla trom all other Simiide
which have organs convex towards the lateral vallate papille, but
the tongue of Gorilla gorilla has enormous conical papille on the
pharyngeal part of the tongue, large plice fimbriate, a median
ventral crest, and a large triangular frenal lamella; it has no
Apical Gland of Nuhn, and its vallate papille form a V or Y.

The only character which really differentiates Anthropopithecus
troglodytes on the one hand from Hylobates and Symphatangus on
the other is the possession of plice fimbriate, for there is a
similarity between them in all other features ; and Hylobates has
a well-marked, triangular, bifid lamella, whereas Symphatangus
syndactylus has two small folds over Wharton’ s Ducts similar to
those of Homo and Simia satyrus. Their vallate papillae form
a Y.

In the Cercopithecide, Cehide, and Hapalide the vallate
papillae never form a Y, the conical papille on the base of the
tongue are never disproportionately large, and the lateral organs
are neither ladder-like nor convex towards the lateral vallate
papille. The chief differences between them lie in the characters
of the lateral organs, the size of the apical cluster of fungiform
papille, and the mode of arrangement of the conical papille on
the oral part of the dorsum.

In the Cercopithecide the lateral organs form rows of lamin
and sulci on the lateral borders, the apical cluster of fungiform
papillae is large, and the conical papillz on the oral part of the
dorsum are regularly arranged. ‘I'he genera are distinguished by
the characters of the orifices of ducts and pits on the pharyngeal
part of the dorsum (page 762).

In the Cebide the inner borders of the lateral organs are con-
cave towards the lateral vallate papille, the conical papille on
the oral part of the dorsum are crowded together without any
definite arrangement, and the apical cluster of fungiform papille
is small.

The vallate papille are two in number, and there is a well-
marked median suleus on the pharyngeal part of the dorsum in
Aotus, but in all other genera there are more than two vallate
papille, but no basal dorsal suleus.

The ventral papillary zone is wide, and has few fungiform
papille in Lagothria, but it has many papille in Ateles and Cebus.
In Afeles the ventral fungiform papille are most numerous
round the apex, but in Cebus they are most numerous farther
back.

The ventral papillary zone is narrow in Pithecia, Alouwatia,
Callicebus, and Cacajao. It has many fungiform papille in
Pithecia but few or none in the others; and Alowatta is the only

766 DR. C. F. SONNTAG ON THE COMPARATIVE ANATOMY ‘

genus possessing large glandular orifices on the base of the tongue.
Callicebus has many nodules and few orifices on the base, but
Cacajao has neither.

The Hapalide always have a vallate triangle and a few
irregular lamine and sulci in their lateral organs. The pha-
ryngeal part of the tongue occupies a relatively small part of the
whole, and the ventral papillary zone is small or absent. The
conical papille are regularly arranged and the apical cluster of
fungiform papille is small. The lingual characters are not
sufficiently distinctive to classify the genera.

The Lemuroidea and Tarsioidea differ from the Simize in the
possession of a sublingua whose size, consistence, mobility,
denticles, and minute structure vary in different families.

In the Tarsioidea the sublingua is soft, small, devoid of
denticles, and only delimited at the sides from the under surface
of the tongue by a narrow groove. There are no lateral organs,
no large conical papille, and few apical fungiform papillee.

In the Lemuroidea the sublingua is large, horny, denticulated,
and is move or less movable.

In Lemur and Hapalemur the sublingua is triangular or lyrate,
very free, and has three ventral crests. No lytta is present,
The vallate papillee form a Y, and the conical papille on the base
ave disproportionately large. The lateral organs are convex
towards the lateral vallate papille.

In Chiromys the sublingua is tongue-shaped, adherent by its
central parts, has one ventral crest which is keel-like and
nodulated. It has a lytta inside the tongue. Vallate papille
form a pair or triangle, but never a Y. Conical papillae on base
of tongue large. No lateral organs.

In Microcebus, the Loriside and Galagidz the sublingua is a
flat plate with one median ventral crest and a variable dorsal
crest. It varies in mobility, consistence, and denticles, and the
value of these characters has already been described (page 763).
The vallate papille form a triangle, but never a Y, the conical
papilla on the base of the tongue are not disproportionately large,
and lateral organs are absent. :

Tbe special points arising from these notes on classification
are :—

1. The tongue of Sima satyrus resembles that of Homo.

2. Simia satyrus and Gorilla gorilla differ from all other
Simiidz in the characters of the lateral organs, but there the
resemblance between these two species stops.

3. The tongue of Hylobates differs from that of Anthropopi-
thecus troglodytes in having no plice.

4. The separation of Symphatangus from Hylobates.

5. The great value of the lateral organs for purposes of
classification.

6. The value of the lingual glands for distinguishing between
the genera of the Cercopithecide.

—

ee a a a ae

OF THE TONGUES OF THE MAMMALIA. 767

7. The genera of the Hapalide cannot be distinguished from
ene another by the characters of the tongue.

8. The tongues of Lemur and Hapalemur resemble those of
the Simiide most closely.

9. It is impossible to ally the tongue of Wicrocebus with those
of Lemur and Hapalemur, but it resembles those of the Loriside
and Galagide.

10. The characters of the tongue at once distinguish the
Lorises, Galagos, and Pottos from Lemur.

11. The tongue of WZarsius resembles neither those of the
Simiide nor of Lemur, and is hard to distinguish from those of
the Loriside and Galagide. It has three vallate papille in a
line, but these may be elements of a triangle which has been
flattened out.

12. The keel on the under surface of the sublingua is not the
true lytta as described by Owen.

ON SOME NEW PARASITIC MITES. 169

40. On some new Parasitic Mites.
By Stanuey Hirst, F.Z.S8.

(Offered for publication by permission ae the Trustees of the British Museum.)

[Received September 18, 1921; Read October 18, 1921.]
(Text-figures 16-43.)

The present paper deals with various parasitic Acari, mostly
belonging to. the family Gamaside, including eight new species
of Liponyssus and a new species of Rhinonyssus, also two new
genera closely allied to Rhinonyssus. A new Tarsonemid mite
belonging to the genus Podapolipus is also described, and a new
‘Red Spider” (Paratetranychus trinitatis) found on grapes at
Trinidad.

Text-figure 16.

Rhinonyssus nove-guinee, sp. u., %. Ventral view.

Genus Rutyonyssus Trouessart.
RHINONYSSUS NOVE-GUINES, sp.n. (Text-fig. 16.)

©. Abdomen not elongated. The spinules or hairs usually
present on the venter in this genus have practically disappeared,
being mostly replaced by exceedingly minute and inconspicuous
circular pores. Anal plate distinct, being elongate pyriform ; it
is furnished with a pair of hairs. Second and third coxe each

770 MR. STANLEY HIRST ON SOME

with a pair of very stout cones or tubercles; the other segments
of the legs are provided ventrally with similar stout cones and
also a few spinules.

Length (including capitulum) *93 mm.

Host: A bird (Craspedophora magnifica), New Guinea. Pro-
fessor Trouessart’s collection. ;

RHINONYSSOIDES, gen. nov.

Closely alhed to Rhinonyssus Trouessart, but the chelicera is — f
very differently shaped, consisting of a short stout basal portion
and a long thin distal style, the latter has a very minute lobe
near the distal end, which is very slightly curved, being in the
form of a minute claw-like denticle (text-fig. 17 6).

Text-figure 17.

TERZIJ

Rhinonyssoides trouessarti, sp.u., 2. Ventral view.
a. Palp from below. b. Chelicera.

RHINONYSSOIDES TROUESSARTI, Sp. n. (Text-fig. 17.)

2. Body shaped as in Rhinonyssus, the abdomen not elongated
however. Anterior hairs on venter short but quite fine. A
number of hairs are situated posteriorly on the venter; they are
longer than the anterior pairs of hairs and are fairly slender, the

‘

NEW PARASITIC MITES. (erRil

basal half being only slightly thickened, whilst the apical half is
fine. Anal plate present, being pyriform i in shape and furnished
with three hairs as in Liponyssus ete. First free segment of
palp apparently with a process. There is a median row of
minute denticles on the ventral surface of the base of the
eapitulum. Anteriorly the second coxa is produced to form a
stout process or spur rather similar to that present in the same
position in Liponyssus but thicker and blunter. The spinules or
hairs present on the proximal segments of the legs in Rhinonyssus
are replaced by curious globular structures in this species; a pair
being situated on each of the first three coxe and a single globule
on the fourth coxa. Trochanter of first leg with two chitinous
globules ventrally, second and third trochanters each with three
globules and a short pointed spinule, fourth trochanter with
four globules and a short pointed spinule. Distal segments of
legs furnished with short pointed spinules. Claws of first legs
only slightly curved (the claws of the other legs are missing in
the unique specimen).

Length (with capitulum) 92 mm.

Host: A bird (Sphecotheres mawillaris), Australia. Professor
Trouessart’s collection.

NEONYSSUS, gen. nov.

Allied to ZLiponyssus but with the palpi shorter, the sternal
plate longer thau wide, and the hairs on the body and legs very
few in number. Differing from Lhinonyssus in the presence of
two dorsal shields (an anterior and a posterior one), instead
of only a single anterior shield.

NEONYSSUS INTERMEDIUS, sp. n. (Text-figs. 18, 19.)

2. Two dorsal shields both fairly large, their surface reticulated
and with numerous punctations, posterior shield with the hinder
end rather bluntly rounded off. Sternal shield longer than wide,
apparently there are no hairs on it, but there are three minute
dots or circles on each side. Grenito-ventral plate unusually short.
Anal plate similar to that of Liponyssus, being pear-shaped.
Palpus short, being very similar to that of Rhinonyssus.
Apparently there is no median groove on the ventral surface of
the capitulum. With the exception of the pair on the anal plate
there seem to be no spines or hairs on the ventral surface.
Peritreme opening dorsally asin Rhinonyssus. Legs rather short,
apparently only the distal segments are furnished with hairs, a
dorsal hair on the tarsus of the third leg being longer than the
others, but still it is not very long ; there is also a pair of distal
spines on the ventral surface of this segment. Except for the
dorsal spur on the anterior surface of the second coxa which is
present but quite short (obsolete), the coxe are without hairs or
spines.

It is possible that some of the hairs on the body and limbs of

772

Text-figure 19.

Text-figure 18.

MR. STANLEY HIRST ON SOME

Neonyssus intermedius, sp. n., 9. Dorsal view.

a. ‘Tarsus of first leg from above.

Neonyssus intermedius, sp. n., 9. Ventral view.

NEW PARASITIC MITES. 773

the single specimen have been destroyed by the caustic potash
used in preparing it. It seems probable, however, that this
absence is natural for in general appearance the mite is very
like a Rhinonyssus, and the spines may be reduced or almost
suppressed in the latter genus. There is a fully-developed larva
inside the specimen and it resembles the parent in having only a
few hairs on the legs. The claws of this larva are well developed
and strongly curved.

Length (with capitulum) *53 mm.

Host: The above description is based on a single specimen
labelled ‘“*Sur un Oiseau de Madagascar.” Professor E. L.
Trouessart’s collection. .

Genus Liponyssus Kolenati.

The species of this genus are found both on birds and mammals.
Only the species that I have been able to examine personally
are included in the key given below. A number of species are
not in dur collection, and these have been left out. Two dorsal
shields are present in the nymphal stages of Liponyssus, whereas
there is usually a single undivided dorsal shield in the adults.
In the adult females of a few species, however, there are two
dorsal shields as in the nymphs: for instance in LZ. viator, sp. n.,
where they are shaped exactly as in the nymphs of certain other
species. The adult females can always be distinguished by the
presence of separate sternal and genito-ventral plates on
the venter, and by the delicate fan-shaped or semi-circular
genital operculum situated at the front end of the latter plate.

Key to the females of the species of the genus Liponyssus
present in the British Museum Collection :—

ie with two shields.............
Ale

2
Worsomauwithtarsmpdetshield sy. et tek aes cee: wcicete 8
aie of the two dorsal shields very small ......... 3.

2.
4

Posterior of the two dorsal shields fairly large .........

( Anterior dorsal shield small and separated from the

| minute posterior shield by a considerable space... L. serpentiwm Hirst.
| Anterior dorsal shield rather long and only separated

i. from the posterior one by a short space ................ I. natricis Gerv.

Sternal plate quite short, the two posterior hairs on it
n long. Anterior spur on second coxa short............. LZ. viator, sp. n.
Sternal plate longer

ae margin of sternal plate strongly chitinised .

OD on

Posterior margin of sternal plate weakly chitinised...
{ Posterior dorsal shield with only three pairs of fairly
J Jone hairs S. pss eee ee ee aa slate) cen, eheropteralis,
6. nom, nov.
| Posterior dorsal shield with a number of fairly long
feeeap IMDS: ss vetna re mete tte are Saeed vcr eactiness¢ Lan WYASEC, SPs Ds

774 MR. STANLEY HIRST ON SOME

‘Sternal plate with strong reticulate markings. A few
of the hairs at the posterior end of the body longer
7 2 than the other hairs

| Sternal plate smooth. Hairs at posterior end of body
not much longer than the others ..

( Genito- ventral plate truncate pedoonly nel fae
nished with three pairs of hairs g

ioe)

Genito-ventral plate rounded off or pointed pos-
Re HERO My desc06 a0 doe acd ob aoa soa voGnba as vee aaa ToOSLU eee cod ehE coo
Sternal plate with a pair of well-defined circular

| stigma-like markings. Genito-ventral plate with
9.4 THI PUIEE) JOHN TRS) OIE WEMDES Us cag ngscoaddoecoateasoendoce penboandio:

| Sternal plate without any circular markings. Genito-
{ventral plate with only a single pair of hairs.........
( Sternal plate with the posterior margm thickened......

10.
( Sternal plate not thickened posteriorly ..............

Coxe of legs furnished with distinct dentiform spurs.

11-% Coxe of legs without any spurs, except an anterior
dorsal one on the second CoxaA ............cee cee cee eens
Coxa of second leg with a very large curved outer

12. spur 4

Coxa of second leg without any very large outer spur.
Coxa of second leg with two posterior spurs ............
13

Coxa of second leg with only one posterior spur ......

(Second coxa with a large curved inner (anterior)
| spur ventrally. Second tarsus usually ending in
14.4 two spurs, one of w nchrisiveny stout 1o 0s. sass

{ Second coxa without any anterior inner spur ventrally.

(Only the anterior dorsal spur of the second coxa and
1s J a little outer posterior spur on the third coxa present
oO.

(Several other spurs present on the coxee as well ......
(Sestenon end of dorsal scutum rounded off or blunt..
16. ‘

| Posterior end of scutum pointed ................

Second coxa with only a very short enmiedl : spur
posteriorly rate
lips

Second coxa with a large pointed spur posteriorly
( Posterior end of eenito-ventral plate rather wide and
| blunt or rounded off .
18.4 :
| Posterior end of genito-ventral plate narrower and
{usually pointed Case HEY Re EE
Anal plate rather angular in Genes
119),
a plate pear-shaped, the sides being curved.........
( Size larger. Dorsai shield elongated and rather narrow.
Sternal plate comparatively long and without any
| distinct markings. Peritreme rather short .........
4
|
L

20.

shorter and with afew deep curved littleimpressions
on each side in the anterior angles. Peritreme long.

Size smaller. Dorsal shield wider. Sternal plate .

L. biscutatus, sp. n.

L. javensis Oudimns.

LL. madagascariensis,
sp. n.

L. sternalis, sp. n.

10.
LL. ethiopicus, sp. n.

nae
12.

18.

L. creightoni Hirst.

13.
LL. sciurinus, sp. n.

14.

LL. liberiensis Hirst.
oe
LL. blanchardi

Trouessart.
16.

L. arcuatus C. L. Koch

Wie
LL. confucianus Hirst.
LL. macedonicus, sp. n.
19.

PA
LL. flavus Kolenati.

20;

L, ellipticus Kolenati.

LL. granulosus
Kolenati.

NEW PARASITIC MITES. Tigo
oles plate with three pairs of hairs..................... 22.
21.
Sternal plate with two pairs of hairs ..............00... 23.
Dorsal shield narrow, especially posteriorly (parasitic ;
a5 OUMTAUS)) Seay oekaki. necealas . cat muh rma tes eee ea CoO may ELL STiR
Dorsal shield wider (parasitic on hen)... ................. D. bursa Berlese.
( Hairs on dorsum only slightly eurved.........0........... DL. silviarum
23. Can. & Fanz.
Hairs on dorsum very distinctly curved.................. 24),

{ Dorsal shiela rather short. Anal plate rather wide
Ieee nigh Ov boweeen ce iaet creat «elk, Rae Ri Ra

* 24.<
| Dorsal shield more elongated. Anal plate narrower
Raye Gere oe eee bse ctins cudiwe bos cehecduelcoobedaannte dl Suarounaen on OOK hbaniist

L. lacertinus Berlese.

LIPONYSSUS NATRICIS Gerv.

Liponyssus arabicus mihi described from a female specimen
found on a lizard (Agama adramitana) is apparently identical
with ZL. natricis Gerv. I have examined examples of the latter
found on a snake at Florence kindly sent me by Prof. Berlese.
A very minute posterior shield is present on the dorsum in the
adult female, and Berlese seems to have overlooked it. It is
very similar in shape to that present in Z. serpentiwm mihi, but
is longer and only separated from the anterior dorsal shield by a
short space, whereas in the L. serpentiwm the two dorsal shields
are widely separated from one another.

LIPONYSSUS VIATOR, sp. n. (Text-fig. 20.)

Q. There are two dorsal shields, shaped as shown in figure,
the anterior one being slightly produced in the middle posteriorly,
whilst the posterior one has the anterior margin rather strongly
concave ; posterior dorsal shield shaped rather like a tongue, the
lateral margins being at first convex and then narrowing to form
the bluntly pointed posterior end. Hairs on dorsal shields short,
some of the hairs on the margins of the anterior dorsal shield
are a little longer, and the hairs of the terminal pair on the
posterior shield are longer than the others on its surface.
Lateral hairs on body fairly long and the posterior hairs on
uncovered part of dorsum very much longer than those on the
shields. Sternal plate very short, being in the form of a narrow
transverse strip; it bears three pairs of hairs, those of the
anterior pair are much shorter than the posterior ones, the latter
being much longer than usual. Genito-ventral plate narrow, the
posterior eud rather blunt. Anal plate pear-shaped. Peritreme
reaching as far forwards as the middle of the second coxa or
somewhat further forwards. A minute and inconspicuous spine
or spur is present on the anterior surface of the second coxa, but
it is much shorter than in most species of Liponyssus; the other
cox apparently are without spurs. It is not easy to make out
the denticle on the first free seginent of the palp in the specimens

Proc. Zoor. Soc.—i921, No. LIT. 52

776 MR. STANLEY HIRST ON SOME

from Cypselus affinis, but it is quite distinct in the example from
Gecinus vaillanti, being sharply pointed.
Length of body (not including capitulum), 2 °65--87 mm.
Host: Several specimens from nests of Cypselus affinis,
Calcutta, collected by C. Paiva (28. vi. 1909) and forwarded to
me by Dr. Nelson Annandale. Also an example from Gecinus
vaillanti, Tangier (May 1892), presented to the Museum by
. Dr. James Waterston.

Text-figure 20.

Liponyssus viator, sp. u., 9. Dorsal view.

a. Sternal plate.

Nore.—The specimen from Gecimus vaillantt agrees closely
with those from Cypselus affiis in the structural details given
above, but the posterior dorsal shield is rather more narrow, and
the denticle on the first free segment of the palp distinct.

LIPONYSSUS CHIROPLERALIS, Nom. nov.

2. Two dorsal shields are present, their surface being finely
reticulated. The shape of these shields varies somewhat in
specimens from different localities; the posterior one is tongue-
shaped, its hinder end being narrowed and blunt. Hairs on the

NEW PARASITIC MITES. ue

shields of moderate length ; there are three pairs of these fairly
long hairs on the posterior shield and also three or four pairs of
very minute and inconspicuous hairs at its posterior end. Hairs
on softer parts of body very numerous, of moderate length and
rather thick. Sternal plate trapezoidal, its posterior margin is
very much thickened, beg strongly chitinised as in LZ, ethio-
picus and L, nyasse ; anterior pair of hairs on it quite short, the
other two pairs usually longer. Genito-ventral plate with the
posterior end sharply pointed. Anal plute pear-shaped. Peri-
treme only reaching as far forwards as the middle of the second
coxa or even shorter. First free segment of palp with a well-
developed process. Second cowa with the anterior spur well-
developed ; apparently there are no other spurs or spines on the
coxe. Bristles on dorsal surface of legs very numerous and
rather stout.

Hosts: Bats. On Pipistrellus, Ambleside, Windermere
(11. ii. 1912) (J. FL. Cormack); ex Dr. J. Waterston’s coll.
On Myotis myotis, Oristano, Sardinia (Dr. A. HW. Krausse); ex
Hon. N. C. Rothschild’s coll. On bats, at Ain Sefra, S.W.
Algeria (14. v. 1913); ex Hon. N. CU. Rothschild’s coll. Off
Ctenodactylus gundi, Gorge of Tilaton, Kl Kantara; specimens
collected by the Hon. L. W. Rothschild and Dr. E. Hartert.
Also a specimen from a bat, Tabgka, Tiberias (P. £. Schmitz) ;
ex the Hon. N. C. Rothschild’s coll.

Norn.-—Canestrini, relying upon the colour of living specimens,
considered this species of Liponyssus with two dorsal shields to be
L. arcuatus C. L. Koch (see Prospett. Acar. Ital. i. pp. 120-121,
1885), and Berlese is also of this opinion in his Acari ete. Reprt.
in Italia (Ordo Mesostigmata, p. 22, 1892). Oudemans gives the
name L. arcuatus to quite a different species with an undivided
scutum (see Arch. Naturg. Abt. A. Heft ix. pp. 68-75, 1913).
According to this author the name of this species with two dorsal
shields should be Z. masculinus C. Li. Koch—a mite found on the
house-mouse (see Tijdschr. Ned. Dierk. Ver. (2) vii. p. 293, 1902).
It is impossible to determine the two species said by Kolenati to
have two dorsal shields, viz., Steatonyssus periblepharus and
S. brachypeltis (see Sitzb. K, Akad. Wiss. Wien, xxxv. pp. 186 &
187, 1859). As it seems to be impossible to ascertain the real
name of this mite, I propose the new name JL. chiropteralis
for it.

LIPONYSSUS NYASSA, sp. DD.

2. Two dorsal shields are present in this species, the anterior
one being very wide; the posterior one is much narrower and
shaped like that cf Z. cethiopicus, sp. n., and L. chiropteralis,
being tongue-shaped and with the anterior margin distinctly
concave. It differs from the posterior dorsal shield of Z. chiro-
pteralis in having a number of moderately long hairs on its
surface instead of only three pairs as in that species. Hairs on
uncovered (softer) part of dorsum numerous, stiff, of moderate

52*

778 MR. STANLEY HIRST ON SOME

length, and sharply pointed much as in L. chiropteralis. Sternal
plate rather weakly, chitinised except the posterior margin, which
is very much thickened and slightly concave as in L. chiropteralis
and “. wthiopicus; there are numerous fine transverse linear
markings on its surface and the three pairs of hairs are long and
fine. Genito-ventral plate with the posterior end rather bluntly

Text-figure 21.

Vitae ice

THRZI—

Liponyssus biscutatus, sp.n., 2. Dorsal view.

pointed, and it has only one pair of hairs on it. Anal plate pear-
shaped. Hairs on vente numerous, fine and of moderate length.
Peritreme extending a little further forwards than the middle of
the second coxa. First free segment of palp with a ventral
process. With the exception of the well-developed anterior spur
on the second coxa, the coxe are without spurs or spines, Legs
furnished with numerous fine hairs or sete.
Host: Elephant shrew, Chiromo, Nyasaland; ex Coll. of
Imperial Bureau of Entomology.

NI
“I
io)

NEW PARASITIC MITES.

LIPONYSSUS BISCUTATUS, Sp. n. (Text-figs. 21, 22.)

2. Dorsal shields two in number, both are fairly wide, the
outline of the posterior one is shaped rather like a thimble or
cone, the blunt apex of which is the posterior end. Hairs on
scutum not numerous and arranged as shown in figure, those in
the middle of its surface are short, but the ones on the margins
of the anterior plate are longer. Hairs on softer part of dorsum
much longer than those on the dorsal shields, a few at the
posterior end of the body being considerably longer than the
others. Hairs on venter fine and of moderate length. Sternal
plate large and its surface is strongly reticulated ; its posterior

Text-figure 22.

Lhd Aasy \ 11
Ss as a’
LA ls im

TERS

Liponyssus biscutatus, sp.n., §. Ventral view.

margin is concave; three pairs of hairs are present on it, the
first pair being short and weak, the posterior pairs very long.
Genito-ventral plate narrowed posteriorly to form a sharp point.
Anal plate pyviform and rather longer than usual. Peritreme
reaching as far forwards as the middle of the second coxa or
slightly further. First free segment of the palp furnished with
a strong pointed tooth-like process. Anteriorly the second coxa
has the usual spur, otherwise the cox are devoid of spurs.
Hairs on legs quite fine, none of them being spiniform.

Length of body (not including capitulum) °65 mm.

Host-: Dendropicus cardinalis, South Africa,

780 : MR. STANLEY HIRST ON SOME

LIPONYSSUS MADAGASCARIBNSIS, Sp..n. (Text-figs. 23, 24.)

@. Dorsal shield shaped rather hke that of LZ. bursa (see text-
fig. 23), the hairs on its surface short, the terminal pair at the
posterior end somewhat longer than the others. Hairs on the
softer part of the dorswm numerous, they are short but mostly
slightly longer than those on the shield. Sternal plate large,
being longer than usual; it is trapezoidal in shape, the posterior
margin being straight except in the middle where it is slightly
coneave; there are three pairs of hairs on it and they are rather
short. Genito-ventral plate unusual in shape for a species of

Text-figure 23.

SOY J. :
N My f I Ny TERZIW
Liponyssus madagascariensis, sp. n., 9. Dorsal view.

Liponyssus, for instead of being pointed or rounded off posteriorly
it is truncated ; also there are three pairs of hairs on it instead
of only one pair, one pair being situated opposite the fourth coxe,
the two others at the posterior end of the plate, one of these
pairs being placed on its posterior margin. Anal plate pear-
shaped and more elongated than usual. Peritreme extending a
little beyond the first coxa. Besides the usual dorsal spur, there
is a ventral (inner) spine anteriorly on the second coxa, it is stout
but curved. There is a similar anterior spine on the third coxa.
Both the second and third coxe are without any posterior spurs.

NEW PARASITIC MITES. 781

There is, however, a short but blunt posterior spur on the fourth
coxa. Legs furnished with hairs and bristles, there is a short
spine ventrally at the distal end of the tarsi of the posterior
legs.

Text-figure 24.

Liponyssus madagascariensis, sp. n., 9. Ventral view.

Length of body (not including capitulum), 2 -79--96 mm.
Host: Lemur mongoz var. albifrons, Diego Suarez, Madagascar
(G. Melow); ex the Hon. N. C. Rothschild’s coll.

LIPONYSSUS STERNALIS, sp. n. (Text-figs. 25, 26.)

2. Dorsal shield large and undivided, it is much narrowed
anteriorly, in the middle the sides are straight and parallel,
posteriorly it is rather abruptly narrowed to form a short tail-
like portion which is truncated posteriorly. Anterior hairs on

782 MR. STANLEY HIRST ON SOME

scutum of moderate Jength and so are also two pairs at the
posterior end, one pair being terminal, the rest of the hairs on
its surface are very short. Hairs on soft ‘uncovered part of
dorsum of moderate length. * Sternal plate trapezoidal in shape,
its posterior margin slightly concave; on each side in the
anterior angle there is a very well-defined circular mark consisting
of an outer rim and a pitted imterior; three pairs of hairs are
present on the sternal plate and they are long and fine. Genito-
ventral plate rounded off posteriovly and with three pairs of hairs
on its surface, viz., the usual pair situated opposite the fourth

Text-figure 25.

Liponyssus sternalis, sp. n., 9. Dorsal view. |

coxze and also two pairs at the posterior end of the plate, one of
them being terminal. Anal plate large and pear-shaped. Hairs
on venter very numerous and of moderate length. Peritreme
extending forwards about as far as the middle of the first coxa.
Palp ventrally with a lobe-like process on the first free segment
as in L. granulosus etc. The usual anterior dorsal spur is
present on the second coxa; there are also chitinous raised lines on
the coxe but no distinct spurs. There are numerous bristles
on the legs, especially on the dorsal surface.
Length of body (not including capitulum) *80 mm.

§
"4
\
4

NEW PARASITIC MIYES. 783

Host: A bat found in a cave near Lake Beshik, Salonika
(27. v. 1918), Dr. James Waterston.

Text-figure 26.

5
cs

x,
ede An SN
ay ITT . YQ
Ppp :
14

Liponyssus sternalis, sp.n., 2. Ventral view.

a. Anterior angle of sternal plate greatly enlarged.

LIPONYssUS ATHIOPICUS, sp.n. (Text-fig. 27.)

2. Dorsal shield shaped as shown in text-fig. 27, being elongated
and comparatively narrow; it is widest opposite the interval
between the second and third legs, being narrowed anteriorly,
whilst posteriorly it is still narrower gradually decreasing in
width, and being rather drawn out and tapering, the end bluntly
pointed. Hairs on its surface numerous; they are not very
long and are usually plain, but some of them have a few incon-
spicuous hairlets. Hairs on uncovered part of dorsum also
numerous; they are not much longer than those on the scutum,
some of them are slightly bifid at the end and furnished with

784 MR. STANLEY HIRST ON SOME

inconspicuous accessory hairlets, especially those at the hinder
end of the body, which are also slightly stouter and blunter than
the others. None of the posterior hairs are especially long.
Sternal plate trapezoidal, not short, its surface finely reticulated ;
its posterior margin is much thickened and slightly concave.
Genito-ventral plate narrowed and tapering to a point posteriorly.
Anal plate pear-shaped. Hairs on venter rather numerous; they
are finer and rather longer than those on the uncovered part of

Text-figure 27.

Liponyssus ethiopicus, sp.n., 2. Dorsal view.

the dorsum. Peritreme reaching about as far forwards as the
middle of the first coxa. Anteriorly the second coxa has a strong
forwardly directed spine; otherwise the coxe are devoid of
spines or spurs. First leg long, especially the tarsus, the fourth
legs also long; second and third legs much shorter, the former
being rather stout. First free (proximal) segment of palp
furnished with a process which seems rather flattened and not so
spiniform as in some species,

NEW PARASITIC MITES. 785

3. Ventral plate with reticulate markings and rather narrow,
being scarcely widened posteriorly; there are eighteen hairs on
it arranged in nine pairs and also an unpaired posterior hair.
The usual strong anterior spur is present on the second coxa.
There is also a rather strong curved spine on the anterior margin
of the third coxa. Bristles on legs mostly fine and none of them
are very stout. Peritreme reaching the first coxa.

Norre.—The male is described from specimens from the Sudan
only.

Length of body (not including capitulum), 9 °61 mm. (distended
female specimens up to '98 mm.); ¢ °43—--45 mm.

Hosts: Elephant shrew, Chiromo, Ruo River, Nyasaland,
1916 (R. C. Wood); ex Coll. of the Imperial Bureau of Ento-
mology. Also off bats at Singa, Sennar Province, Sudan, 1913
(Capt. W. #. Marshall); ex the Hon. N. C. Rothschild’s coll.

Text-figure 28.

a. Liponyssus sciurinus, sp.n., §. Dorsal view.
b. Liponyssus blanchardi Trouessart, 2. Dorsal view.

LIPONYSSUS SCIURINUS, sp. n. (Text-figs. 28 a, 29.)

2. Body oval, being widest some distance before the posterior
end, sometimes it is slightly bilobed posteriorly. Dorsal shield
large and undivided, being shaped as shown in text-fig. 28 a,
posteriorly it is evenly narrowed, the end being bluntly pointed.
Hairs in middle of this shield exceedingly minute and fine, even

786 MR. STANLEY HIRST ON SOME

the marginal hairs are quite short, the terminal posterior pair
being, however, slightly longer than the others. Hairs on
uncovered part of dorsum short and fine. Sternal plate longer
than is the case in L. carnifex and L. arcuatus, the posterior
margin being only slightly concave; the three pairs of hairs on
its surface are fairly long. Genito-ventral plate of moderate width,
its posterior end rounded off. Anal plate pear-shaped, Hairs on
venter fairly numerous. Anterior (dorsal) spur of second coxa
well-developed; there is also a little curved hook-like spur (or

Text-figure 29.

TERZE—~
til H
i
L

Liponyssus sciurinus, sp. n., 9. Ventral view.

rather spine for it is articulated) on the inner surface of this
coxa anteriorly, and two large spurs or teeth on the posterior
margin, the outer one being almost as large as the inner. Third
coxa also with two spurs on its posterior margin, but the outer
one is much weaker than the inner. Peritreme extending forwards
as far as the middle of the first coxa or a little further
forwards, There are a number of subspiniform bristles on the
distal segments of the legs; the paired stout little spurs or
spinules present at the distal end of the second tarsus in

NEW PARASITIC MITES. 787

L. carnifex and L. arcuatus ave absent in this species from
squirrels.

3. Genito-ventral plate vather slender and it has nine pairs of
hairs on it, besides the unpaired posterior hair, it is not divided
into two portions by a transverse line as is the casein L. flavus ete.
Peritreme veaching as far forwards as the middle of the first coxa.
Spurs and spines on the coxze as in the female sex. “At the distal
end of the second tarsus there is a pair of very. strong short
spinules ventrally. Fourth leg not enlarged, the spiniform
bristles on this limb are not very thick.

Text-figure 30.

Liponyssus blanchardi Trouessart, 2. Ventral surface showing coxe, sternal
and genito-ventral plates.

Length, 2 (including capitulum and palpi) 83-90 mm.; of
body only -73-—81 mm.; greatest width ‘40-42 mm.
3. Length of body 53 mm.; width °31 mm.

Host: Squirrels at Briangon, France (26. ix. 1887). Collection
of the Paris Faculty of Medicine.

LiPonyssvus BLANCHARD! Trouessart. (Text-figs. 28 6, 30, 31.)

Leiognathus blanchardi Trouessart, 1904, Arch. Parasit. viii.
pp- 558-561, 2 text-figs.

2. Dorsal shield large and undivided, narrowing posteriorly,

788 MR. STANLEY HIRST ON SOME

the tip being blunt (almost pointed). Hairs on dorsal shield all
short, but the marginal ones are longer than the others. Hairs
on uncovered part of body about as long as the longest of the
marginal hairs on the shield or somewhat longer, a few at
the posterior end of the body are longer than the others, but
still none of them are very long. Sternal plate trapezoidal, not
very short, and furnished with the usual three hairs which are of
moderate length; the posterior margin is straight. Genito-

Text-figure 31.

Liponyssus blanchardi Trouessart, &.

a. Ventral surface showing coxze and ventral plate.
b. Fourth leg greatly enlarged. é
ventral plate of moderate width, the posterior end rounded off.
Anal plate pear-shaped, Hairs on venter short and not very
numerous. Leritreme reaching about as far forwards as the
middle of the first coxa. With the exception of a distinct (outer)
pointed tooth or spur on the posterior edge of the third coxa, and
the usual anterior spur of the second coxa, the coawe are devoid
of spurs or teeth. None of the spines on the legs are very stout.

S$. Ventral shield becoming rather wide behind the last cox,
its sides here being convex but afterwards narrowing to form the

ee ee ee

NEW PARASITIC MITES. 789

usuil neck-like portion connecting with the anal part of the plate;
the transverse line present on this plate in Z. flavus and
L. granulosus is absent in this species. here are nine pairs of
hairs on the ventral plate and also the usual unpaired posterior
hair. Besides these hairs on the plate itself there are a few
lateral hairs posteriorly on the venter and they are fairly long.
The curved anterior seta on the inner side of the third coxa is
rather stout, being almost spiniform ; this coxa has also a very
slender outer posterior spur. Fourth coxa also usually with a
very slender posterior spur. First leg more slender than the
others, the second and third being short and stout; the fourth
leg is enlarged as in ZL. carnifew C. L. Koch, some of the spines
on this limb being rather stout, especially in the specimens from
the Prairie Marmot. Second tarsus with a pair of distal spinules
ventrally which are very stout and quite short.

Length. 2 (without capitulum) ‘73-78 mm.; ¢ °58—63 mm.

Host: Alpine Marmot. ‘The above description is based on a
number of female specimens mounted on a slide (one of Professor
Blanchard’s preparations in the collection of the Paris Faculty
of Medicine). Also examples of both sexes taken on marmots at
Zermatt (M. Biner); ex N.C. Rothschild’s coll. Also examples
of both sexes from specimens of the Prairie Marmot (Cynomys
ludovicianus) bred by Sir KH. Loder at Leonardslee, Horsham,

England.

LIPONYSSUS CONFUCIANUS, nom. nov.

Lipynyssus berlesei Hirst, Proc. Zool. Soc. 1921, p. 362.

Canestrini has already described a species of Leiognathus=
Liponyssus from a bat under the name ZL. berlesei (Atti Soc. Ven.
Trent, xi. pp. 142-143, 1888), so it is necessary to find a new
name for my species from MJyospalax scansus. 1 propose the
name L,. confucianus for this species.

LIPONYSSUS MACEDONICUS, Sp. n. (Text-figs. 32, 33.)

2. Differing from L. sciwrinus ete. principally in the shape
of the dorsal shield, which is rather abruptly narrowed at the
posterior end so as to practically form a short tail. Hairs on
dorsal shield exceedingly minute, a few near the anterior end on
the margins being a little longer and the terminal pair at the
posterior end of the shield also a little longer than the rest, but
still quite short. Hairs on uncovered part of body also short,
but two pairs at the posterior end (one of them ventral) are
somewhat longer than the others. Sternal plate trapezoidal and
not short, the posterior margin straight ; the three pairs of hairs
on this plate ave rather short. Genito-ventral plate short and
fairly wide, its posterior end rounded off. Hairs on venter a
little longer than those on the dorsum. <dAxal plate rather
elongate pyriform. Peritreme reaching almost as far forwards
as the anterior margin of the first coxa. Second coxa with the

790 MR. SLANLEY HIRST ON SOME

anterior (dorsal) spur large, ventrally this coxa has a large
posterior spur (larger than in Z.arewatus). Third coxa with two
posterior spurs, the inner fairly well developed, the outer weaker.
Fourth coxa apparently without any spur. Femur of first leg
dorsally with two rather noticeable but not very long spiniform
sete near its distal end and there are also some quite short ones
on this limb, Ventrally the tarsus of the second leg has two
stout but very short spurs or spinules. arsi of third and fourth
legs with a spine near the distal end. There are also some
weaker spines and setze on the legs.

Text-figure 32.

Liponyssus macedonicus, sp. n., 9. Dorsal view.

dé. Ventral plate without any transverse line, it is expanded
just behind the last coxe and afterwards narrows again to form
the neck-like portion connecting it with the anal part; the anal
part of the plate has a very minute denticle or cone on each of
its margins just before the striated tip; there are nineteen hairs
on the ventral plate including the posterior unpaired one. Coxe
furnished with spurs as in the female, but there is also a slender
(outer) spur on the last coxa. Second leg stout but rather short,
the fourth leg longer than the others and considerably swollen,
one of the spines on the patella and another on the tibia are

4
.
}
}
i
d

NEW PARASITIC MITES. 791

stouter than the rest. A pair of spinules are present ai the
distal end of the tarsi of the three posterior pairs of legs.

Length of body (not including capitulum), 2 °65—67 mm.:
36 57mm.

Host: Spalax monticola thermaicus. Doiran, Macedonia.
Several specimens collected by Dr. James Waterston (24. x

1918).

Text-figure 33.

Liponyssus macedonicus, sp.u., 2. Ventral view.

Lirponyssus FLAVUS Kolenati. (Text-figs. 34, 35.)

Dermanissus flavus Kolenati, Parasit. Chiropt. 1857, p. 19.

Liponyssus lobatus Oudemans, Tijdschr. Ned. Dierk. Ver.
1904, (2), pp. 18-22, pl. i. figs. 3-17.

@. A little before the posterior end the dorsal shield becomes
narrowed, but the tip is not pointed but widely rounded off
(almost truncated) as in L. carnifex. Hairs on its surface very
short and fine; they are not numerous and are distributed as
shown in the figure. Hairs on uncovered part of dorsum very

Proc, Zoou, Soc,—1921, No. LIII. 53

792 MR. STANLEY HIRST ON SOME

numerous, being very short and rather stiff. Sternal plate
trapezoidal, the posterior margin being strongly concave; the
three pairs of hairs on this plate shorter than its length. Grenito-
ventral plate curiously sculptured, especially the anterior part of
it (genital operculum) which has scale-like markings ; posterior
end of this plate rounded off. Anal plate more triangular than
pear-shaped, the sides being straighter than usual. Venter with
numerous very short stiff hairs. Peritreme reaching as far as
middle of the first coxa or a little further forwards. First
free segment of palp with a large flattened process. Second

Text-figure 34.

TERZI—

Liponyssus flavus Kolenati, 2. Ventral surface showing cox, sternal and
genito-vyentral plates.

coxa with the usual strong anterior spur, but without any
distinct spurs or tooth-like processes posteriorly ; on the outer
half of its posterior border, however, it is furnished with a slight
bet distinet curved raised edge.’ Third coxa also with a slight
curved raised edge posteriorly, the outer half of this coxa is
semewhat produced and bears a hair. Fourth coxa with a
chitinous line near the posterior margin; sometimes there is
a tiny inconspicuous spine connected with an outer series of very
minute denticles on the extreme outer side of this coxa, but it is

NEW PARASITIC MITES. 793

easily overlooked. There are some rather thick spiniform sete
on the dorsal surface of the legs, those on the femora being
especially noticeable.

3. Posterior end of dorsal shield narrowed and angular, being
almost pointed, but blunt instead of rounded off as in LZ. granu-
losus. Ventral plate divided into two regions by a transverse
line as in LZ. granulosus. There are five pairs of hairs on the
anterior part of this plate and eighteen hairs on the posterior

Text-figure 35.

Liponyssus flavus Kolenati, g. Ventral view.

a. Sternal plate of Liponyssus granulosus Kolenati, 2.

part of it. Hairs on venter numerous and quite short. Peritreme
apparently somewhat longer than in the male of ZL. granulosus,
almost reaching the middle of the first coxa. Fourth leg not
specially enlarged.

Length of body (not including capitulum), 9 ‘65mm.; ¢
54mm.

Host: Bats. The above description is based on Kolenati’s
types in our collection labelled Dermanissus flavus There are
also specimens from a Noctule bat, Aylesford, Kent (27. x. 1919),
collected by the author,

53*

794 MR. STANLEY HIRST ON SOME

LIPONYSSUS GRANULOSUS Kolenati.

Dermanissus granulosus Kolenati, Parasit. Chiropt. 1857,
p. 20.

Dermanissus glutinosus Kolenati, ib. p. 20.

@. Dorsal plate rather like that of Z. flavus in shape. The
hairs on its surface very minute, those on the anterior margin
longer, however. Sternal plate trapezoidal in shape, the posterior
margin strongly concave; in the anterior angle of this plate
there are a few rather deep impressions or markings, the rest of
its surface being practically smooth and without sculpturing ;
the three pairs of hairs on this plate are longer than in L. flavus,
being longer than the plate itself. Genito-ventral plate with the
markings on its surface very similar to those present in
L. flavus, the posterior end is rounded off. Anal plate pear-shapeds
the sides being more convex than in L. flavus. Venter with
numerous short hairs, which are slightly longer than in L. flavus.
Peritreme reaching as far forwards as the middle of the first coxa
or slightly further. First free segment of paip with a large
flattened process as in Z. flavus. Coxze of legs very like those of
L. flavus, the second and third having slight raised curved edges
posteriorly, which perhaps represent obsolete spurs.

3. Posterior end of dorsal shield wider and rounded off
posteriorly (instead of almost angular and bluntly pointed as in Z.
flavus). Two of the hairs at the posterior end of this shield are
much longer than the others. Hairs on uncovered part of dorsum
very sparse. Ventral plate divided into two regions by a strong”
transverse line (situate almost opposite the posterior end of the
last coxe). Hairs on the posterior part of this plate Jess
numerous than is the case in LZ. flavus and slightly longer than in
that species. There are five pairs of hairs on the anterior part of
it as in LZ. flavus, but only 9-10 on the part of it situated behind
the transverse line and they are arranged regularly in pairs
(whereas there are eighteen on this part of the ventral plate in
L. flavus). airs on venter few in number and slightly longer
than those of Z. flavus. Peritreme shorter than in the female,
apparently only reaching a little further than the middle of the
second coxa. Last pair of legs not enlarged and the bristles and
spines are not very stout.

Chelicera furnished with a rather long pointed process besides
the two short fingers.

Length of body (not including capitulum), 9 *63—"68 mm. ;
3 55mm.

{Norn.—The male of this species is described from a single
example of this sex from’ the tube labelled Z. glutinosus by
Kolenati, for there were no males in the tube labelled L. granu-
losus. |

Host: Bats. Two tubes (each containing a few specimens)
labelled’ Dermanissus granulosus and Dermanissus qlutinosus.
They bear the British Museum Register no. 56.163 (December,

NEW PARASITIC MITES, 795

1856) and are described in the register as types presented by
Prof. Kolenati. We have also speciinens of this species taken on
Vespertilio myotis at Begamo, ex N.C. Rothschild’s collection.

Text-figure 36.

Liponyssus ellipticus Kolenati, 2. Ventral view.
a. Dorsal shield of same.

LIPONYSSUS ELLIPTIOUS Kolenati. (Text-fig. 36.)

Caris elliptica Kolenati, Parasit. Chiropt. p. 16 (1857).

@. Size large for a Liponyssus. Dorsal shield more elongated
and narrower as compared with its width than that of L. granu-
losus Kolenati, posteriorly it is narrowed, the end being practically
truncate; the hairs on the posterior part of the scutum, especially
the marginal ones, are much longer than inthat species. Hairs on
softer uncovered part of dorsum fairly numerous and of moderate
length. Sternal plate rather large, being longer than that of
L. granulosus, its posterior margin is concave and not thickened ;

796 MR. STANLEY HIRST ON SOME

the three pairs of hairs on it fairly long. Genito-ventral plate
with the posterior end rounded off, and it only bears a single
pair of hairs. Anal plate pear-shaped, the sides of the anterior
part of it being well curved, whilst the pointed posterior end is
elongated. Peritreme rather short. First frée segment of palp
with a process ventrally. Anterior surface of second coxa with a
long slender spur; except for this one, the coxe are entirely
devoid of spurs or spines. Numerous fine sete are present on
the legs.

Text-figure 37.

Liponyssus longimanus Kolenati, . Ventral view.
a. Chelicera of same. i

Length (owing to the damaged condition of the unique specimen
it is impossible to give the exact length).

Host: The name of the host is not given on the label, but
Kolenati gives it as M/yotis murinus in the work quoted above.

LIPoNYssUS LONGIMANUS Kolenati.. (Text-fig. 37.)
Caris longimana Kolenati, Parasit. Chiropt. p. 16 (1857). —
3. Dorsal shield with faint reticulate or scale-like markings;

NEW PARASITIC MITES. 197

its posterior end is narrowed and practically truncate, the hairs
on its surface are not very long, one or two of the marginal ones
posteriorly are a little longer, however, and the pair at the end
considerably longer than the others. Ventral plate with about
twenty-five hairs on its surface, including the posterior unpaired
one; it becomes greatly enlarged in width behind the last coxe
and is not divided by any transverse line. Hairs on venter very
few in number and of moderate length (not very long). Movable
finger of chelicera only about half the length of immovable one.
Peritreme short, only reaching the second coxa. With the excep-
tion of a long slender spur on the anterior surface of the second
coxa, the coxe are without spurs or spines.

Length of body (without capitulum) *59 min.

Host: The name of the host is not mentioned on the label,
Kolenati says the species is found on Miniopterus schreibersti and
Rhinolophus clivosus.

Nore.—It seems probable to me that this mite is the male of
Liponyssus ellipticus Kolenati, but it is better to keep the names
separate until proof of this is forthcoming.

Family TARSONEMID4&.
Genus Tarsonemus Can. & Fanz.

TARSONEMUS TRANSLUCENS Green. (Text-figs. 38, 39, 40.)

3. Body of the male rather long and narrow, being more than
twice as long as its greatest width; it is widest at the point at
which the posterior pairs of legs are inserted, this part of the
abdomen being slightly wider than the cephalothorax. Posterior
end of abdomen narrowed and rather tail-like, each of the last
segments being considerably narrower than its predecessor. The
curious anal structure at the extreme end of the body is only 20
wide, the segment preceding it 27-30 » wide, and the third from
the end 40-50 » wide. When viewed laterally the body is seen to
be very high at the level of the posterior legs, but anteriorly it is
much lower and the abdomen is narrowed posteriorly as noted
above. Hairs on body all rather short. Legs. Second leg a
little longer than the first and the third is the longest. Fourth
leg really the shortest, but its position at the highest and widest
part of the body makes it appear longer than the first and second
legs: this leg is stowter than the others and its penultimate
segment has an angular internal spur or process distally, there is
also a long stout bristle situated near it; fourth tarsus curved or
bent and a little beyond the middle of its length it is furnished
with a very long fine hair, after which the segment becomes
slightly reduced in width. Tarsus of the three first legs with a
pulvillus, but the terminal segment of the fourth has only a short
knob-like structure at the end, it is possible that this is the
rudimentary tarsus. There is a stout striated sensory seta on
the anterior tarsi, that on the second tarsus being larger than
the one on the first. There are also four slender but stiff sensory
setz on the first tarsus.

798 MR. STANLEY HIRST ON SOME

2. Body oval, being considerably longer than wide. Hairs on
body few in number and quite short and inconspicuous. Pseudo-
stigmata almost circular in shape. First leg the shortest, being
slightly shorter than the second. Claw of first tarsus rather large
and only slightly curved. The largest of the striated olfactory sete
on this segment is long but slender and situated near the middle,
there are also two short striated sete of the same nature near the
proximal end, one of them being club-shaped. In addition to the
fairly stout sete just mentioned there ave two pairs of long

Text-figure 38. ~

Tarsonemus translucens Green, g. Ventral view.

slender tactile sete. Second tarsus with a rather long striated
seta near its proximal end, but it is a little shorter than the
longest of the stout ones on the first tarsus.

Measurements. 6. Body (including capitulum) 157-167 » in
length; width of cephalothorax 72-75; greatest width of
abdomen about 80. Length of first leg 75 p, of second 96 p, of
third 122 mu, of fourth (not including the proximal segment) 72 p.

2. Body 190-207 » in length ; in width 107-115 p.

_ Hab. On Capsicum, Trinidad (fF. W. Urich). On tea, Ceylon
(Hutsoi). At)

circa Si eee

799

NEW PARASITIC MITES.

Text-figure 39.

Tarsonemus translucens Green, &. Lateral view.

Text-figure 40.

Tarsonemus translucens Green, 2. Ventral view.

800 MR. STANLEY HIRST ON SOME

Genus PopAroxuipus Grassi & Rovelli.

PODAPOLIPUS AHARONII, Sp. n. (Text-fig. 41.)

2. Body elongated, being from over three to more than three,
and a half times as long as wide. There are usually a, large
number of embryos or eggs in the body. Cephalothorax minute.
and bearing the appendages. Anterior pair of legs minute but
segmented, the segments apparently being three in number;
a seta is present dorsally at the distal end of the first segment of ©

Text-figure 41.

THR Al J

Podapolipus aharonii, sp.n., 2.

} a. Dorsal view of capitulum and anterior end of body showing the legs, etc.
b. The same from below.
e & d. Outline of body of the entire mite showing variation in length.

this leg ; the terminal segment has a weill-curved hook-like claw’
at the end. Second pair of legs reduced to smooth little
unsegmented sacs. Just above the first leg there is a tiny cone
or tubercle at the apex of which is the stigmal orifice. Movable
finger of chelicera curved and styliform, being quite slender.
This species is much more elongated than LP. reconditus Grassi
& Rovelli and P. grassit Berl., resembling P. bacillus Berl.

NEW PARASITIC MITES. 801

(from Java) in this respect, but Opsomala cylindrica, the host
of the new species described above, is confined to the Mediter-
ranean Region.

Length of body 122-171 p, its width 40-49 p.

Host: A number of specimens from the wings of Opsomala
cylindrica Marsch., Ekron, Palestine, 15. ii. 1921 (J. Aharon).

The specimens were red in colour when alive.

Fam. TETRANYCHIDG.

Genus PARATETRANYCHUsS Zacher.

PARATETRANYCHUS TRINITATIS, sp.n. (Text-figs. 42, 43.)

3g. Hairs on body quite short, they are distributed as shown in
the figure, some of them seem plain, but others are furnished

Text-figure 42.

Paratetranychus trinitatis, sp.n., 6. Dorsal view.

a. Traces of palp. b. Penis from the side, greatly magnified.

with inconspicuous accessory hairlets. Anterior eye distinct
although small, the posterior is inconspicuous and easily over-
looked, being more weakly chitinised. Penis curved at the end

- 802 ON SOME NEW PARASITIC MITES.

and shaped as figured. Dorsal spine on palp very short. Dorsal
sensory finger stouter than and a little shorter than the two
neighbonring rod-like sete, the latter are of unequal length, one
being slightly longer than the other. ‘Terminal sensory finger of
palp apparently absent. .

Text-figure 43.

|
}
t

Leaf of vine showing discoloration caused by Paratetranychus trinitatis, sp.n.

Length of male (including capitulum) 342-375 p, of hard
external part of penis about 11-15 y, of rod-like sete of tarsus of
palp 43-6 yw, of dorsal sensory finger of palp 33-4 p.

Host plant: Grape vine, Trinidad (f. W, Urich).

ON THE EXTERNAL CHARACTERS OF THE MUSTELIDA, 803

41, On the External Characters and Classification of the
Mustelide. By R. I. Pocock, F.R.S8., F.Z.S8.

[Received June 6, 1921; Read November 8, 1921.1]
(Text-figures 27-39.)

CoNnTENTS. Page
Entroducttonerte cree reo cero e eer cae one eS
Dieskynimariumis eu eek eee esc eee eee eee BOS
Phe MacialliVilbrisssa lan kN Sa eet en Le
DPhrethxternalar: cAMet ha wi wen oe coe sake OL
ST G gH Seti za Shad seh Bese e ele es SRR cca Me ta Ric cate COU:
The Anus and External Genitalia........................ 827
The Classification of the Mustelidee..................... 829

Introduction.

In 1869 Gray (Cat. Carnivorous etc. Mammals Brit. Mus.
pp. 79-142) relegated the genera which by all modern zoologists
are referred to the family Mustelide to two families, Mustelide
and Melinide. The Mustelidz were divided into the following
tribes :—(1) Mustelina for JMJartes, Mustela, Putorius, Gulo,
Galera, Grison; (2) Lutrina for Lutra and other genera of
typical otters ; (3) Enhydrina for Enhydra. The Melinide were
similarly divided into the tribes:—(1) Melina for Arctonyz,
Meles, Pawidea, Mydaus ; (2) Mellivorina for Mellivora ; (3) Me-
phitina for Mephitis, Conepatus, Spilogale; (4) Zorillina for
Zorilla {| =Ictonyx|; (5) Helictidina for Helictis.

From my present standpoint the interest of Gray’s classification
lies in its influence upon Gill, who in 1872 (Smiths. Mise. Coll.
xi. pp. 64-65) followed Flower (P. Z. 8. 1869, pp. 11-15) in fusing
the Melinide and Mustelide into one family, for which the latter
name was retained, and elevated all Gray’s tribes to the rank
of subfamilies—Musteline, Melinz, Zorilline, Mephitine, ete.
As will appear in the sequel, this classification in my opinion
better expresses the affinities of the genera dealt with than any
that followed it *.

In 1883 Flower (Encycl. Brit., Mammalia, pp. 439-440) divided
the Mustelide into the following subfamilies :—

(1) Melinze for Mephitis, Arctonyx, Meles, Mydaus, Taxidea,
Mellivora, Helictis, Ictonyx.
- (2) Musteline for Galictis| Tayra + Grison], Mustela[ = artes},
Putorius |= Mustela], Gulo.
(3) Lutrine for Lutra, Aonyx, Enhydra [= Lata].

This arrangement, based mainly on the structure of the feet,

* T have intentionally ignored such family names as Mellivoride, Zorillide, etc.,
proposed by Rochebrune and Mephitids proposed by Rhoads, because no attempt
was made to define the families thus casually designated and no reasons were given
for assigning that rank to the groups.

804 MR. R. I. POCOCK ON THE

was adopted by Mivart (P. Z. 8. 1885, pp, 393-395); and with
the addition of Pecilogale and Lyncodon to the Musteline was
repeated by Flower and Lydekker in 1891 (‘ Mammalia,’ pp. 567-
591). It was also accepted by Sedgwick (‘ Students’ Text-Book
of Zoology,’ i. p. 624, 1905).

In 1902 de Winton (‘Zoology of Egypt, Mammalia,’ p. 245),
trusting apparently to the structure of the teeth, modified
Flowev’s system by transferring Mephitis, Conepatus, Mellivora,
and Jctonyx from the Meline to the Musteline. Max Weber in

1904 (Die Siiug. pp. 536-538) followed de Winton with respect
to the allocation of Mellivora, but otherwise agreed with Flower.
And in the same year Trouessart (Cat. Mamm. ., Suppl. pp. 187-
215) also followed Flower, but put. Pecilogale into the Meline.
He further subdivided. Musteline into two tribes :—Gulonee for
Gulo, Galictis [= Tayra+Grison]|, and Lyncodon, and Mustelez
for Mustela [= Martes|and Putorius [= Mustela}, with subgenera
Lutreola and Arctogale {= Mustela].

In 1912 Miller (Cat. Mamm. Western Europe, pp. 340-341)
divided the Mustelide into four subfamilies :—(1) Meline con-
taining “ about a dozen genera,” of which only JJeles is quoted and
dealt with ; (2) Lutrine containing “ four genera,” Lutra alone
being discussed; (3) Gulonine containing G'ulo and probably
Mellivora; (4) Musteline restricted to the genera Martes, Mustela
(with its subgenera Putorius and Lutreola and Mustela), and
Vormela *

Although Miller, like de Winton, seems to have relied mainly
on the teeth, it seems clear that a large number of the genera
placed by de Winton in the Musielinz were left by Miller in the
Meline, where Flower placed them. Another point to be noticed
in connection with Miller’s classification is his severance of Gulo
from the Musteline as the type of a special subfamily t, with the
guess that Mellivora is a related genus.

Setting aside for the moment the classifications of Gray and
Gill, two conclusions come out from this brief review :—(1) There
is complete agreement amongst recent writers with regard to the
status of the Lutrinz; (2) Although Flower’s subdivisions have
held the field until the present day, there is a great variety of
opinion as to the constitution of the Melinz and Musteline. It
is admitted by all that Meles, Arctonyx, Tawidea, Mydaus, and
Helictis are Meline; and that Martes, Mustela (with related
generic or subgeneric forms, such as Vormela, Putorius, Lutreola),
as well:as Zayra, Grison, and Lyncodon are Musteline. But
Mephitis, Spilogale, Conepatus, Mellivora, Ictonyx, and Pecilogale

* Tn 1911 Satunin (Mitt. Kauk-Museums, v. p. 267) defined the genera Vormela
and Putorius, admitting the following subgenera of the latter, Putorius (s.s.),
Lutreola, Ictis, and Kolonokus. The genus Putorius is equivalent to Mustela as
understood by Miller, Ictis (=subgenus Mustela) being employed for the stoats
and weasels. Miller was perhaps not acquainted with this paper when he wrote his
‘Catalogue of the Mammals of Western Europe.’

+ This was originally done by Gray in 18265 (Ann. Phil. xxvi. p. 339); but in his
latest classification he abandoned the group named (xuloninee.

tie a a al

EXTERNAL CHARACTERS OF THE MUSTELID®. 805

have been placed according to the fancy of authors in either
subfamily. It is quite clear therefore that there is something
wrong with the classification, and that the characters used for
defining the two subfamilies are insufficient for the purpose.

In 1920 (Proce. Zool. Soc. pp. 179-187 and pp. 424-436)
I attempted to show that Miller's guess as to the kinship between
Gulo and Mellivora was untenable; and that there is practically
no evidence of sufficiently close relationship between MMeles and
Taxidea to justify their retention in the same subfamily. Grulo
was affiliated with Martes; Gill’s subfam:ly Mellivorine was
resuscitated for Wellivora; Taxidiine was proposed as a new
subfamily group for Yawidea; and Meles and Arctonyx were
provisionally left as the sole representatives of the Meline.

Subsequently (Proc. Zool. Soc. 1921, pp. 473-486 and pp, 535--
546) a study of the auditory bulle in the Mustelide supplied
additional characters for the definition of the subfamilies ; and
reasons for the admission of several genera of Lutrinz were
brought forward.

In the following pages the external characters of all available
genera, apart from G'ulo, Mellivora, Meles, Taxidea, and the
genera of Lutrinz, which I dealt with in the papers just mentioned,
are described in detail; and a summary of the results setting
forth my views on the classification of the Mustelide is given at
the end.

Most of the material forming the subject-matter of the paper
consisted of specimens which had lived in the Zoological Gardens
and were examined after death at the Society’s Prosectorium.
Mr. Oldfield Thomas, however, has given me all the facilities at
his command for studying the splendid collection under his
charge at the Natural History Museum. Iam much indebted to
him and to Mr. Martin Hinton for the help received in this way.
Tam also particularly grateful to Mr. Cyril Hopwood, a member
of the Council of the Rangoon Zoological Society, for sending me
through Mr. Max Friedlander, the Superintendent of the Gardens,
a spirit-specimen of Melogale personata; and to Mr. KE. Jacobson,
for procuring for me at Mr. Boden Kloss’s request, a similarly
preserved example of Plesiogale nudipes * from Fort de Kock on
the west coast of Sumatra.

The Rhinariwm.

The most primitive type of rhinarium is found in MJartes and
Gulo. It is naked above, the lateral slits of the nostrils extend
nearly to its posterior margin, the infranarial portions are shallow
or moderately deep, the inferior edge is convex or angled, but it
is not, strictly speaking, contintied over the upper lip—that is to
say, the gutter which cleaves the lip consists of smooth skin
differentiated’ by its texture from the rhinarium above. This

* Gray referred this species, with others, to his genus Gymnopus (see infra, p. 818).
That name, however, is preoccupied. Hence I propose Plesiogale as a substitute.

806 MR. R. I. POCOCK ON THE

Text-figure 27,

Aty

TH \
Lp MUA yeni ENS
as LM vi RS
z

A. Side view of head of Putorius putorius,
B. Front view of rhinarium of the same.

C. The same of Gale nivalis.

D. Side view of rhinarium of the same.

KE. Front view of rhinarium of Plesiogale gymnopus.
KF, Side view of rhinarium of the same,

G. Side view of head of Charronia flavigula.
H. Front view of rhinarium of the same.

I. Upper view of rhinarium of the same.

K. Front view of rhinarium of Martes foina.
L. Upper view of rhinarium of the same.

EXTERNAL CHARACTERS OF THE MUSTELIDA. S07

gutter is continued as a groove up the middle line of the rhinarium
between the nostrils. (Text-fig. 27, K, L.)

In Charronia the gutter is obliterated, except close to the
convex lower margin of the rhinarium, which is grooved above,
and the upper lip is scantily hairy in the middle line. (Text-
fig. 27, GI.)

In Gale, Mustela,'and Putorius it is also obliterated or nearly
so, and the infranarial portions of the rhinarium are shallower.
In Gale nivalis the hair of the muzzle encroaches on the upper
surface. (Text-fig. 27, A-D.)

In Plesiogale nudipes the rhinarium is large, with a median
groove, deep infranarial portions, and naked above back to the
posterior ends of the nostril shts. (Text-fig. 27, E, F.)

In Jctonyx, as in Putorius, the rhinarium is lightly biconvex
above and mesially grooved in front, with the inferior edge
horizontal in the middle, convex at the sides where it curves
backwards ; but the infranarial portions are deeper than in Gale,
Mustela, or Putorius, and this area of the rhinarium is wider than
the upper portion. There is no trace of gutter and the con-
tinuously hairy upper lip is about as deep as the height of the
rhinarium. (Text-fig. 29, A-B.)

In an example of Grison furax, or an allied species, from
Brazil, the upper surface of the rhinarium is evenly rounded from
side to side, the groove is on the lower half of the front surface,
and the nostrils are wide apart. The infranarial portion is deep
in the middle, but quite narrow laterally, where it passes beneath

the slit of the nostril; and this narrewed piece is defined from
the median piece by an oblique groove. There is no trace of
gutter and the depth of the lipi is less than that of the rhinarium.
Two or three specimens of Grisonell« (sp. ¢) from Cordova in the
Argentine show individual variation in the rhinarium. In one
the upper surface is lightly biconvex, the median groove is
distinct, and theve is no gutter, in the other the median groove is
obsolete, the upper surface is uniformly convex as in Grison, and
there is a very shallow gutter. In both the imfranarial portion
is tolerably deep and mesially angled below, but the lateral
portions, which in Grison pass backwards beneath the nostril slits,
are practically obsolete. (Text-fig. 28, A-C.)

In Zayra barbara the upper side of the rhinarium is grooved
and the anterior surface deeply and widely grooved. The nostrils
are larger and closer together than in Grison and the infranarial
portions are deep both mesially and laterally, and are marked in
front, aS In Gruson, by a fine oblique groove on each side which
meets its fellow of the opposite side in the middle line above the
inferior angle of the rhinarium in front. The rhinarium is about
twice as deep as the upper lip, which has no gutter. (Text-
fig. 28, D-F.)

The rhinarium of Tawidea recalls that of Gale, Mustela, and
Putorius in the shallowness of the infranarial portions and the
presence of the median groove. It is, however, relatively wider,

Proc. Zoou. Soc.—1921, No. LIV. 54

808 MR. R. I. POCOCK ON THE

has the nostrils a little closer together, and the upper surface
overgrown with hair nearly to the ‘anterior margin. ‘The lip has
no cutter and is not so deep as the rhinarium.

In Mellivora the vhinarium is wide as in Taxidea, and the

Text-figure 28.

wn
{o3

of, Ait: aN
ein HAR SS

ph aly!

aK Sey Nike SS

A ¢ ats MAWES

fe (=

Side view of head of Giison furax or a closely allied species.
B. Front view of rhinarium of the same.
C. Upper view of the same.
. Side view of head of Layra barbara.
E. Front view of rhinarium of the same.
F. Upper view of the same.

EXTERNAL CHARACTERS OF THE MUSYELID, 809

upper lip is shallow and without gutter; but the infranarial
portions are somewhat deeper than in that genus; there is no
median groove and the lateral slits of the nostrils are more
abbreviated, and the upper surface is quite naked.

In Jeles the rhinarium is very large and naked, the infranarial
portions being exceedingly deep and often convexly produced in
the middle below. ‘he nostrils are tolerably close together, but
the lateral slits, although narrow, are long. There is no median
groove, and the upper hp, shallower than the rhinarium, has no
gutter.

In the Skunks the range in variation of the rhinarium is very
instructive. In Sprlogale it is very like that of Martes, having
shallow infranavia! portions defined by faint grooves from the
median portion, a median groove in front in the lower part, an
angled inferior border, and a complete gutter dividing the upper
lip, which is not quite so deep as the rhinarium. (Text-
fig. 29, C.)

In Mephitis the infranarial portions are always deep, wide,
and well-defined laterally beneath the posterior sht of the nostrils
as in Meles, and when the groove between the nostrils and the
two defining the Renremeell portions and the gutter on the lip
ave absent, “the resemblance to that genus is particularly close.
But in a specimen, said to have come from Manitoba, the
rhinarium is lower and somewhat wider than in other examples,
the lip is cleft by a shallow gutter continued on to the lower half
of the rhinarium, and the lateral portions are defined by shallow
grooves running from the nostrils to the middle line of the
Fitenen edge. (Lext- fig. 29, D, E.)

In Conepatus proteus the chimera is twice as wide as high,
about equal in depth to the upper lip which has no gutter, its
upper edge is evenly convex, its lower edge horizontal, and,
although there is a short groove between the nostrils, the nostrils
themselves are wholly visible from the anterior view, the lateral
slits being short and not continued to the eee oe edge, thus
aeseqlin the rhinarium of Jellivora. (Text-fig. 29, F, G.)

In Melogale personata the rhinarium is See for the
extreme depth of the mesially grooved infranarial portions which
are produced inferiorly, so that their median depth is equal to
the depth of the area above it and of the upper lip. They are
also very deep laterally beneath the slits of the nostrils, which
are continued to the posterior border of the rhinarium. The
upper lip is ungrooved and is only sparsely hairy mesially and
laterally as far back as the angle of the mouth. ‘(Text- ee. 29,
By, 1; 30, G.)

The nee ium of Mydaus is even more aberrant and recalls
that of the Suide in the sense that it consists of a some-
what rounded dise with two subcircular nostrils on its anterior
surface, the slits of the nostrils being very narrow and short.
The infranarial portions are much deeper than the upper portion

of the rhinarium and are everywhere of equal depth. They are
54*

810 MR. R. I. POCOCK ON THE

defined by a vertical groove running downwards from the nostril,
and each of these grooves is continuous with the outer edge of a
long, narrow, parallel-sided philtrum, of the same texture as the
rhinarium, which extends to the edge of the upper lip and is
marked by a groove passing upwards on to the rhinarium. In
front of the rhinarium above them is a naked strip of skin on
the muzzle. (Text-fig. 25, K, L.)

Text-figure 29.

yh iy
Hp AA
ifattuyer

WSS
NUNS
oe

A. Side view of head of Ictonyx capensis.
3. Front view of rhinarinm of the same.
C. The same of Spilogale putorius.
D, The same of Mephitis mephitica.
EK. Side view of the same.
K. Front view of rhinarium of Conepatus proteus.
G. Side view of the same.
H. Front view of rhinarium of Melogale personata.
J. Side view of the same.
K. Front view of rhinarium of Wydaus meliceps (from dried skin).
L. Side view of the same.
x ac

EXTERNAL CHARACTERS OF THE MUSTELID 4. 811

The genera may be grouped as follows by the rhinarium :—

a. No complete and true philtrum resembling the rest of the
rhinarium in texture dividing the upper lip, which is at most
grooved.
6. Upper lip divided by a complete groove of smooth naked
skin, giving mobility to the right and lett halves ............ Gulo, Martes.

6’. Groove on upper lip obliterated or nearly so, the two halves
of the lip not separately movable, or only slightly so.

ec. Infranarial portion of rhinarium much shallower beneath
lateral slits of nostrils than above them.

d. Intranarial portion laterally very narrow...... Mustela, Gale, Putorius,
Charronia, Grison, Taxidea.
d’, Infranarial portion laterally moderately deep... Tayra, Ictonyx, Spilo-
gale, Mephitis, Conepatus.

ce’. Infranarial portion very deep beneath lateral slits,
approximately as deep as portion above them ............... Meles, Melogale.

a. A long narrow philtrum, resembling the rest of the rhimarium,
dividing the upper MPs infranarial portions everywhere very
ALG e py Bice ras dae cepnctnosiatelas Dee eenleeee wacca bie mentee ee suamoca tance eaten My LAUESE

The Facial Vibrisse.

The number of tufts of facial vibrisse characteristic of the
Carnivora is typically present in the Mustelidee; but the vibrisse
themselves are longer and more numerous in predatory forms
(like Maries, Mustela, and J/ctonya) than in fossorial forms (like
Mellivora, Meles, and Mephitis). They are especially well
developed in the matter of rigidity and number in the truly
semi-aquatic genera like Lutra and Amblonyx; but in the
tropical West- “African Otter, Paraonyx, Hinton records the com-
plete absence of the superciliar y and upper genal tufts. In other
genera, like JMJeles, the upper genal tuft is sometimes apparently
absent; but a noticeable point about this tuft in all genera,
when it is present, is its high position on the cheek, only ¢ a little
below and behind the posterior corner of the eye. In the one
fresh example of Z'ayra barbara examined, I could find no trace
of the interramal tuft. (Text-figs. 27-29.)

In Melogale personata the vibrisse are all long and numerous as
in predatory forms, the submentals being unusually well developed
on the area of nearly naked skin which runs along the lower lip
as far as the angle of the mouth, corresponding to that of the
upper lip. (Text-fig. 30, G.)

The Heternal Ear.

The external ear shows every grade between high specialisation,
the primitive condition, and almost souiplete degeneration. In
its highest types, as exemplified by Martes* and “Char ronid, it 1s
of tolerably large size and is pr ovided with a well- -developed bursa,
the posterior flap of which is large, has a strongly convex margin,
and is attached above behind the straight-edged portion of the

* See my paper on the Pine and Beech-Martens (P. Z. S. 1914, pp. 1062-1068).

S12 MR. R. I. POCOCK ON THE

posterior border of the ear which constitutes the anterior ‘flap.
The supratragus has a simple subglobular thickening. The tragus
and supratragus are well defined and separated by a distinct.
notch, and the antero-internal and postero-internal thickenings
descend behind or within the tragal and antitragal prominences
respectively. (Text-fig. 27, G.)

Text-figure 30.

we ae aor

Har of Gale nivalis.
» Lecilogale albinucha (from dried skin).

» Mephitis mephitica.

- 4 Conepatus proteus.

BF. ,, Spilogale putorius.
G. Side view of head of Melogale personata.

ID EE Gae <i.

A.
B.
C. ,, Lyncodon patagonicus ( s 4 )e
Dv.
E

In Mustela, Gale, Plesiogale, and Putorius the ear is relatively
smaller and rounder than in MWartes, and further differs in that

EXTERNAL CHARACTERS OF THE MUSTELID. 813

the thickening of the supratragus is larger and valvular, and the
edge of the cavity has a small soft membranous flap just above
the supratragus, which is especially well developed in Plesiogale.
‘Moreover, the antero-internal thickening curves sharply beneath
the supratragus posteriorly. In Gale, Mustela, and Plesiogale
the bursa is almost or quite as well developed as in Martes, but in
Putorius it is reduced to a marginal slit, owing to the reduction
in size of the posterior flap, the. edge of which is near ly straight.
(Text-figs. 27, A; 30, A.)

Gulo has the pinna like that of JMartes, but relatively smaller
and provided with a smaller bursa; and the pinna of Melogale
is very like that of Mustela and Gale, the bursa, tragus, and
antitragus being well developed, but the supratr agus is less
valvular, the antero-internal ridge descends more vertically, and
there is a little pouch above the hollow. (Text-fig. 30, G.)

Tayra and Grison also resemble Jfustela and Gale in the pre-
sence of the upper membranous flap, the valvular supratragus,
the posterior curvature ot the antero-internal ridge beneath the
supratragus ; but differ in the reduction of the bursa to an even
erveater extent than is seen in Putorius. The posterior flap is
continuous above and below with the margin of the pinna, and
the anterior flap is represented by a very shallow ridge of in-
tegument a little in front of it, so that the poueh is reduced to a
minimum. In Grison and Grisonella the valvular supratragus
and the flap above it are larger than in Tayra; and in Grisonella
the anterior ridge of the bursa is shorter than i in Grison. (Text-
He 28, A, Ly)

in eve yx, and also in Pecilictis and Pecilogale, judging from
dried skins, the pinna differs essentially from that of Jusiela,
Gale, and Putoriws in the total absence of all trace of the bursa
and of the membranous flap above the supratragus. Otherwise
it is similar. (Text-figs. 29, A; 30, B.)

In Meles and Tamidea the pinna is smaller than in /ctonyz, and
the supratragus has a simple subglobular thickening as in J/artes.
Otherwise they are tolerably similar; the pinna is less salient in
Tasxidea than in Meles.

In Lutra the pinna, although much reduced, is of the same
simple type as in JMeles. In Hydrictis it is simplified by the
almost complete suppression of the tragus and antitragus and of
the supratragus apart from the subglobular thickening.

In Spilogale, Mephitis, and Conepatus a noticeable peculiarity
in the pinna is the entire absence of the tragal thickening, so
that the external orifice of the ear is exposed and not hidden in
a cup-like hollow; and the cavity of the pinna has no definite
anterior rim. The low antero-internal ridge curves posteriorly
beneath the valvular supratragus, which appears to be set higher
in the ear than in other genera. The pinna is of comparatively
large size in Spilogale; but in Conepatus—at all events, in
C. protews—it is much reduced and scarcely stands away from the

head. There is no trace of the bursa. (Text-fig. 30, D-F.)

814 MR. R. I. POCOCK ON THE

In Zyncodon also the pinna is as much reduced as in Conepatus,
standing away from the head as a shallow short ridge, the lower
end of which does not reach nearly to the aditus inferior. here
is no bursa, but the structure of the ridges could not be deter-
mined on dried skins. (Text-fig. 30, C.)

The pinna of Wellivora surpasses even that of Conepatus and
Lyneodon in reduction in size. It is represented merely by a
thickening of the integument, without any free edge, round the
small cavity; and, judging from the descriptions, the pinna of
Mydaus is very similar to that of Mellivora, but I am unable to
say whether the orifice is exposed as in Mephitis, Spilogale, and
Conepatus or concealed by a tragal thickening.

The following table shows briefly the principal modifications in
the external ear :—

a. Bursa retained.
b. Bursa marginal, upper end of its posterior flap hidden by
HITED LOVE HIFEPNOW IY) suo cdoonn cod ecu, Gaannheisees one cen Mustela, Gale, Plesiogale,
Putorius, Martes, Charronia, Gulo, Melogale.
b’, Bursa remote from margin, shallow, its posterior flap in
no way concealed by anterior flap................. Tayra, Grison, Grisonella.
a’. Bursa absent.
c. Auditory orifice concealed, usually by tragal thickening.
d. Pinna laminate, standing away from head.

e. Supratragus a well-developed valvular flap ............ LIectonyx, Pecilictis,

Pecilogate.
e’. Supratragus at most a ridge with rounded thickening.

Ff. Pinna moderately large ..............:-00000cseeeeessss. Meles, Paxidea.
jf o JSEWOA, TOEIN TACHA 6... -o3 ace codsay coand6voN oes LEe oor ~ Lutra, Amblonyzx, ete.

d’. Pinna a thickened rim of integument .. Boapoenunodoreccn. JURA GGG

e’. Auditory orifice CES not concealed Ihe ‘hace thicken-
SUC ROS ar Scie each tn hota kea ea tea eS eey ison ‘Spilogale, Mephitis, Conepatus.

The Feet.

In describing the feet of the Mustelide I start with two
assumptions. First, that the primitive feet were adapted for
running or climbing, or both combined, and not for digging or
swimming, which hamper, if they do not prevent, the first-
mentioned activities. Second, that the feet were naked below,
provided with short curved claws, interdigital webbing up to the
proximal end of the pads, large 4-lobed plantar pads, double
carpal and metatarsal pads, and a hairy heel; and that the digits
were unequal in length, the first being the shortest, the second
and fifth next and approximately equal in length, and the third
and fourth the longest and also approximately equal, so that the
digital pads, when not spread, formed a strongly curved line
round the anterior margin of the plantar pad. And possibly in
the hind foot the third and fourth digits were more closely tied
together than to the second and fifth respectively. .

The genera whose feet most nearly approach these hypothetical
feet are Zayra and Grison. It is therefore needless to describe
them in detail. From the material examined I have not been

EXTERNAL CHARACTERS OF THE MUSTELIDS. 815

able to satisfy myself that there is any fundamental difference
between the feet of the two, unless it be that in Gvison the
plantar pads are defined from the carpal and metatarsal pads by
a deeper and wider groove. In Yayra these pads are confluent
throughout their width, there is only a very slight trace of
division between the two moieties of the carpal pad and none in

Text-figure 31.

A. Right hind foot of Tayra barbara.

B. Right fore foot of the same.

C. Right hind foot of Charronia flavigula.
D. Right fore foot of the same.

816 MR. R. I. POCOCK ON THE

the case of the metatarsal pad*. In an example of Grison furax
or an allied form, from Brazil, there is a comparatively large area
between the plantar and carpal pads, the latter are well defined
but mesially in contact, and there is some naked skin above the
large external pad. Similarly, in the hind foot there is some
naked skin above the somewhat rounded metatarsal pad, which
except internally is distinctly separated from the plantar pad.
These differences from the feet of Zayra are, however, not so
well defined in examples of Grisonedla from Cordova in the
Argentine. In both genera the pads are granular and tessel-
lated, and the third and fourth digits of the hind foot are a little
nearer together than they are respectively to the second and
fifth. Nevertheless, there is quite a marked extent of web
between their pads. (Text-figs. 31, A, B; 32, C, D.)

Of these two genera Vayra is an active tree-climber like
Martes, and Grison an agile ground-hunter like JZustela; and in
Martes and Mustela and genera related to them the highest type
of feet subservient to those modes of life are found. The claws
are shorter and more curved, the digits are more widely separable,
and the third and fourth of the hind foot are not closer together
than to the second and fifth respectively. The underside of the
webs is more or less hairy, there is at least a patch of hair
between the carpal and plantar pads, the carpal pads are quite
senonied, the metatarsal pad is absent, the hair of the caleaneum
extending down to the plantar pad, and all the pads are
narrower and transver sely or concentricaily ridged and grooved.

Of the genera related to J/artes, Charron has feet which
depart least from the type seen in Grison and Tayra, although
still very distinct. The area between the digital and plantar
pads has four large patches of hair on the interdigital webs ; but
these do not reach the edge of the webs distally or the plantar
pads proximally, and the lower side of the digits is also naked ;
and there is a similar patch of hair surrounded by naked skin on
the area between the carpal and plantar pads, which are well
developed, the pollical and hallucal elements of the latter being
widely confluent with the plantar, while the internal and ex-
ternal moieties of the carpal pads are large and have a small area
of naked skin above them. (Text-fig. 31, C, D.)

In Martes martes and M. foina the area between the digital
and plantar pads is covered with hair, except for narrow naked
strips radiating along the digits; the area between the plantar
and carpal pads and round the latter is aiso continuously hairy ;

* Lonnberg (Arkiv fdr Zool. vill. no. 16, p. 10, 1913) figured the hind feet of
two forms of Tayra from different altitudes and localities in Ecuador, to show that
the combined plantar and metatarsal pad—there being no line of demarcation
between them—is longer in the race from the lowlands. Defects in the drawings of
these feet may be attributed to their being taken from skins softened in water after
being dried.

r ‘In 1918 I resuscitated this genus of Gray’s, based upon Martes flavigula, when
I tonne that the baculum differs greatly from that of Martes martes and M. foina
(Ann. & Mag. Nat. Hist. (9) i. pp. 308-310).

EXTERNAL CHARACTERS OF THE MUSTELIDA. 817

the plantar pads are smaller, more subdivided, their pollical and
hallueal elements are connected by a narrow strip with the main
pad, and the carpal pads are much reduced and separated by a
longer space from the plantar pad. J. foina, however, has
much larger digital, plantar, and carpal pads than JZ. martes*, and
thus more nearly approaches Charronia. In Jf. martes the
digital pads are narrowly ovate, the plantar pads are broken up
into four areas connected by narrow naked strips, and the carpals
are quite small.

Text-figure 32.

. Right hind foot of Pecilogale albinucha (from dried skin).
. Right fore foot of the same.

Right fore foot of Grisonella sp. from Cordova, Argentine.
. Right hind foot of the same.

Sein >

In Mustela erminea and hibernica, Gale 7 nivalis and africana,
Putorius putorius, furo, and nigripes the feet, except that they
are relatively shorter and stouter, generally resemble those otf

* For figures illustrating these differences, subsequently verified on other
specimens, see my paper on these species of Martes (Proc. Zcol. Soc. 1914, pp. 1¢62—

1068).
+ Distinguishable from Mustela by the shape of the baculum.

818 MR. R. I. POCOUK ON THE

Martes in hairiness and pad development. But in G. africana
alone have I found the plantar pad broken up into four elements
to the same extent as in MW. martes; and only in @. africana
could I find the inner moiety of the carpal pad; but, judging
from its minute size in this species, it may have been overlooked
in the others. (Text-fig. 33, C—F.)

Writing of the Muinks (Lute cola), Flower and Lydekker
G imme Living and Extinct,’ p. 586), said that they differ from
the Polecats, Stoats, and Weasels “ by the toes being partially
webbed and by the absence of hair in the intervals between the
naked pads of the soles of the feet.” From this statement it
seeins quite clear tnat neither of these authors was aware that
the toes of Polecats, Stoats, and Weasels are fully webbed up
to the digital pads. As for the hairiness of the soles of the feet, I
cannot discover from examining the long series of dried skins of
Minks in the British Museum that there is any essential
difference between them and Polecats, Weasels, and Stoats in
that or any other particular. The area between the plantar and
digital pads is covered with hair; and it appears that Baird was
perfectly correct when he said that the feet are well furred
between the pads, though the hair is scantier in summer than in
winter.

In some members of this group, however, the feet are quite
naked beneath. Gray, for example, based the genus Gymnopus *
upon certain Oriental species related to Mustela and Gale, namely
nudipes, strigidorsa, and kathiah; and to these africuna was
added. According to the diagnosis the soles of the hind feet in
these species have three oblong pads, an arched and bald space
behind them, and the heel hairy. But in the Egyptian example of
Gale africana above veferred to the feet are approximately as
hairy as in G. nivalis; and in kathiah, which I provisionally retain
in Mustela, pending the discovery of the structure of the baculum
or other characters to settle its affinities, there is a patch of hairs
on the lower side of the webs and also between the carpal and
plantar pads, and the hair extends down to the plantar pad on
the hind foot. Thus both africana and kathiah fall outside the
genus according to the diagnosis.

In Plesiogale nudipes and strigidorsa the feet are quite naked
beneath, and the plantar and carpal pads are relatively con-
siderably larger than in Mustela, Gale, and Putorius, and
apparently less coarsely striated. The plantar pads are very
distinctly four-lobed, the three main lobes being widely in
contact, while the pollical and hallucal lobes are attached to the
posterior end of the inner of the three main lobes. The two
carpal pads are well developed and almost in contact. Just
above them there is a narrow naked space, and there is a
corresponding space, which, however, seems to vary individually

* Quoted in this paper as Plesiogale (see above p. 805), of which nudipes is the
type-species. I premise that the group is at least as much worth nominal
recognition as Lutreola or Kolonokus.

EXTERNAL CHARACTERS OF THE MUSTELIDA. 819

in P. nudipes, and lies behind the plantar pad of thefhind foot.
(Text-fig. 33, A, B.)

The feet of these two species are of great interest. It might
be claimed that their nakedness is a secondarily acquired
character, correlated with life in warmer latitudes than those

Text-figure 33.

A. Right hind foot of Plesiogale nudipes.
B. Right fore foot of the same.

C. Right hind foot of Gale nivalis.

D. Right fore foot of the same.

E. Right hind foot of Putorius putorius.
¥F, Right fore foot of the same.

820 MR. R. I. POCOCK ON THE

frequented by typical Stoats, Weasels, and Polecats. If nudipes,
which ranges from the Malay Peninsula to Borneo, were alone
concerned, that view would be more defensible than, in my
opinion, it is. But strigidorsa occurs much farther to the
north, in Sikhim. Again, it must be remembered that the
tropical American species brasiliensis, which may be taken as the
type of Gray’s genus WVeogale, if it be resuscitated, has hairy feet.
I incline, therefore, to the opinion that in Plesiogale we have the
most primitive type of foot met with in the subfamily Musteline
as here restricted—a foot which differs mainly from that of
Grison and Grisonella in the complete suppression of the meta-
tarsal pads.

The feet of Gulo* resemble in all essential respects those of
Martes, except that they are relatively broader.

The feet of Pesilogale ave very like those of Plesiogale, but the
claws of the fore feet are a little larger and the third and fourth
digits of the hind foot less widely separated. In both these
particulars Pacilogale, so far as the feet are concerned, connects
Plesiogale with Ictonyx, which in the length of the claws belongs
to the fossorial group of Mustelidee—Pwerlogale being one of
those genera in which the distinction between fossorial and
cursorial Mustelide breaks down. (Text-fig. 32, A, B.)

In the remaining genera the feet are fossorial and characterised
by long and blunt claws, which are especially long on the fore
feet. The only other point they have in common 1s close union,
sometimes amounting to basal fusion, between the digital pads of
the third and fourth digits of the hind feet—a phenomenon
foreshadowed in the feet of Grison and Tayra and Pecilogale.

Fossorial feet which come nearest to the feet of the Weasels,
Polecats, and Martens, although differmg in the characters
mentioned in the last paragraph, are found in Jctonyv and
Pecilictis. The feet, closely resembling those of Pacilogale, are
tolerably narrow, and the digits, apart from the third and fourth
of the hind foot whieh are close together, are widely separable ;
the metatarsal area is without trace of pads and covered with
hair down to or nearly down to the plantar pad; the digital pads
ave oval, weli defined, and coarsely striate—at least apically ; the
plantar pad is narrow, deeply emarginate behind, and very
distinctly four-lobed, and the lobes are coarsely striate and are
in contact to about the same extent as in Martes foina; there
ave two striate carpal pads, a smaller inner and a larger outer,
separated from each other and from the plantar pad by about
the same distances as in Martes foina, although the sizes of these
pads and the distances above mentioned vary according to the
species. The carpal pads are largest and the distances in question
smallest in the species in which the area round them and the
avea between the plantar and digital pads are naked. In other

%* See my paper on the external characters of this genus (Proc. Zool. Soc. 1920,
pp. 179-187).

EXTERNAL CHARACTERS OF THE MUSTELIDA. 82]

species the hair of the carpus extends to the carpal pads, overlaps
the inner and spreads into the posterior emargination of the
plantar pad; and there is a scanty patch of hair upon the lower
side of the interdigital integument*. (Text-fig. 34, A-D.)

Text-figure 34.

A. Right fore foot of Ietonyx capensis, from S. Africa, with hair on
interdigital integument.

B. Right fore foot of the same.

C. Right hind foot of Ictonyx sp., from the Sudan, with naked soles.

D. Right fore foot of the same.

* In Pacilictis the feet are more like those of typical Mustela, in
the sense that the soles are much more thickly hairy and the
plantar pad is smaller and more subdivided, no doubt in con-
formity with the sandy habitat of the animal.

The feet of Zawidea resemble those of /ctonya in the absence

* Thus Thomas’s statement that Ictonyx may be distinguished from Pecilictis
by the nakedness of the soles of the feet does not hold good.

822 MR. R. I. POCOCK ON THE

of the metatarsal pad, but in most respects they are very different.
The digits are shorter, closer together, with the pads larger, less
well defined, and the interdigital webs (although narrower) extend
further up the pads, especially on the fore foot. The plantar

ad of the broad fore foot, although wider than long, is narrower
than the foot itself, indistinctly lobate, and much less deeply
emarginate behind; there is a single small carpal pad, representing
the outer * of the normal two, set a little distance behind the
plantar, and the area behind this and the plantar pad is partially
overgrown and overlapped by hairs on the inner or pollical side
of the foot as in some examples of /etonyx. In the hind foot the
plantar pad is longer than wide, narrower than the foot, irre-
gularly heart-shaped, and very indistinctly subdivided. The pads
are granularly roughened.

The feet of Lyncodon, judging from dried skins, are peculiar in
the almost complete suppression of the interdigital webs, which
extend only a short distance beyond the plantar pads. The
underside of the digits and the area between the digital and
plantar pads is naked, but the sides of the digits are fringed with
longish hairs. In the fore foot the claws are long and fossorial,
the plantar pad is strongly arcuate and short, the hairs of the
carpus encroaching upon it in the middle line behind. The
carpal pad is represented only by the outer moiety, which is
continuous distally with the outer lobe of the plantar pad. The
carpal vibrissx are retained, although the claws are fossorial.

The hind foot closely resembies the fore foot, but the claws are
short; and there is no trace of metatarsal pads, the metatarsus
being covered with hair, which extends over the middle of the
plantar pad. (Text-fig. 35, A, B.)

In all the other genera of fossorial footed Mustelide the meta-
tarsal pad is retained and is of large or comparatively large size.

In Melogale personata tT the fore foot is a little wider than the
hind foot, the digits are webbed to the proximal ends of the well-
defined digital pads, the soles are entirely naked as far back as
the proximal ends of the carpal and metatarsal pads, and all the ~
pads are tolerably coarsely striated, although less coarsely than
in Mustela, Maries, Gulo, and Ictonyw. The fore foot otherwise
tolerably closely resembles that of Meles, but the digits are rather
more separable, the digital pads are better defined, the plantar pad
is relatively narrower, more distinctly four-lobed, with its anterior
and posterior margins more curved, the two elements of the
earpal pad are in contact or very nearly so in the middle line, and
the carpal vibrisse are well developed. In the hind foot the
digits are markedly more widely separable than in Meles, there

* In my paper on Tawxidea and Meles (Proc. Zool. Soc. 1920, p. 428) I inad-
vertently described the carpal pad as representing the inner or radial carpal of
Meles.

+ Hodgson figured the hind foot of this species, but wrongly labelled it Urva
eancrivora (Journ. As. Soc. Bengal, xvi. pl.ii.). A figure of the fore foot is in his
unpublished drawings.

EXTERNAL CHARACTERS OF THE MUSTELIDA. 823

being quite a distinct space between the third and fourth,
although the space is narrower than that between the second and
third or fourth and fifth. As in the fore foot the digital pads
are better defined, the plantar pad is narrower, much more
strongly curved, and very distinctly four-lobed, the hallucal lobe,
like the pollical lobe of the fore foot, being as large as the others
individually. There are two well-defined elongated metatarsal
pads, confluent in the middle line proximally, and the hairy
caleaneal area is a little shorter relatively than in Meles meles.

(Text-fig. 35, C, D.)

Text-figure 35.

A. Right hind toot of Lyncodon patagonicus (from dried skin).
B. Right fore foot of the same.

C. Right hind toot of Melogale personata.

D. Right fore foot of the same.

C, D, x i.

The feet of Meles meles 1 have already described and figured
(Proc. Zool. Soc. 1920, pp. 426-428, fig. 20, A, B). In addition
to the differences mentioned above, it may be added that they
are of a more decidedly fossorial type than in dMelogale, the hind

Proc Zoo. Soc.—1921, No. LV. 55

824 MR, R. I. POCOCK ON THE

foot being narrower as compared with the fore foot, with the pads
of the third and fourth digits confluent. .In both the fore and
hind foot the plantar pads are very wide, the carpal pads of the
fore foot are separated, and there is a patch of hair on the area
behind the plantar pad. In the hind foot the metatarsal pads
are shorter and more confluent, and closer to the plantar pad
than in Melogale, and all the pads are granular or papillate, not
striated. ;

Text-figure 56.

A. Right hind foot of Spilogale puterius.
B. Right fore foot of the same.
C. Right hind foot of Mephitis mephitica.
D. Right fore foot of the same.

xs.

Judging from dried skins and from Hodgson’s unpublished
drawings referred to in my paper quoted above, the feet of
Arctonyx seem to resemble those of Meles in all essential par-
ticulars. The soles of the feet are quite naked. The large naked
area above the plantar pad of the hind foot has two elongated

EXTERNAL CHARACTERS OF THE MUSTELID®. + 825

thickenings representing metatarsal pads; the plantar pad of
the hind foot is about as wide as long and markedly three-lobed
(Hodgson probably accidentally omitted the hallucal element)
‘and narrower than the foow; the digital pads of both feet are oval
and well defined. .In the fore foot the plantar pad is markedly
four-lobed and wider than long, and separated from it by a con-
siderable space are the two rather large carpal pads, which are
together about as wide as the plantar pad. The pads are coarsely
granular and tessellated.

The fore foot of Mellivora resembles that of V'axidea rather
than that of Meles in the large size of the digital pads and the
extension of the webbing along their edges, but the plantar pad
is as wide as in Meles, and the pollex is shorter than in either
genus. The carpal pads also are much larger than in either, the
outer moiety being a large subspherical mass at least as long and
half as wide as the plantar pad, while the much smaller, smooth
pollical portion is also larger than in Meles, and the two are
separated from the plantar pad by a narrower strip of integument,
which has no hairs upon it. The hind foot is like that of Meles,
except that the digital pads are larger and more fully webbed,
the plantar and metatarsal pads are larger, and the rest of the
metatarsal area covered with naked wrinkled skin.

In Spilogale, Mephitis, and Conepatus the pads are granular
and the lower surface of the feet is naked, at least as far back as
the posterior border of the carpal and metatarsal pads, which
ave well developed. Moreover in all the examples of Jephitis
and of Conepatus examined the heel from the metatarsal pad
upwards is covered with naked wrinkled skin. Spilogale seems
to have the feet less specialised for digging than the other genera,
in the sense that the plantar pads are more distinctly broken up
into four lobes and the carpal and metatarsal pads more reduced,
the latter being, at all events in some cases, marked by a distinct
longitudinal groove defining the two elements; and the third and
fourth digits of the hind foot, although closer to each other than
to the second and fifth respectively, have the digital pads unfused.
(Text-fig. 36, A, B.)

In the several unlocalised examples of Mephitis examined there
is naturally some variation m the size and shape of the pads,
the two carpals being sometimes large and in contact, sometimes
smaller and separated ; but in all cases the pads are relatively
larger than in Spilogale and less subdivided, and the digits are
shorter and more fully webbed, the pads of the third and
fourth of the hind feot being fused in the middle lire. (Text-
fig. 36, C, D.)

A further stage of specialisation is seen in Conepatus proteus,
where the plantar, carpal, and metatarsal pads are as well deve-
loped as in any example of Mephitis; but the digital pads are
larger and are united by webbing almost up to their proximal
ends. (Text-fig. 37, A, B.)

The most highly specialised type of fossorial foot is perhaps

5d*

826 MR. R. 1. POCOCK ON THE

found in Mydaus. The pads of the second, third, fourth, and
fifth digits are fused together, except quite at them distal ends,
as completely as the pads of the third and fourth digits of the
hind foot are sometimes fused in JMeles. ‘The pads of the pollex
and hallux, however, are independently movable, although
attached by shallow webbing to the pad of the second digit.
The soles are quite naked, The plantar pads are as wide as the

Text-figure 37.

A. Right hind foot of Conepatius proteus.
B. Right fore foot of the same.

C. Right hind foot of Mydaus meliceps.
D. Right fore foot of the same.

feet, much wider than long, lightly arcuate, and four-lobed, the
pollical and hallucal elements being large. ‘There are two well-
developed, slightly separated carpal pads, the inner being much
the smaller of the two. Conjointly they are almost as wide as

EXTERNAL CHARACTERS OF THE MUSTELIDA, 827

the plantar pad, from which they are separated by a short space,
and there is an area of naked skin above them. In the hind foot
the metatarsal area is naked to the heel and is provided with
two long pads as wide as the foot, the inner being the wider of
the two. (Text-fig. 37, C, D.)

The Carpal Vibrisse.—The occurrence of these tactile hairs in
many different orders of Mammals suggests that they are a
primitive mammalian character. J have not mentioned them
under the separate generic headings because they are present in
most of the forms discussed. They are especially well developed
in the active predaceous forms like Mustela, Martes, Tayra,
Grison, and the Lutrine. In the fossorial forms they appear to
be of less importance, judging from the frequent difficulty there is
in distinguishing them amongst the long hairs of the leg. They
are present, however, in Lyncodon and in Lctonyx. In the latter
they are not always to be de ected with certainty, and the tuft
may be reduced to one bristle. But the sense of touch locates
their position by the perception of a small wart-like swelling on
the skin, where, as pointed out by Beddard, the nerve supplying
the vibrisse ends. ‘This sweiling is present in such highly
specialised diggers as Mellivora and Tawxidea, although I failed to
differentiate the vibrisse themselves. In Meles, Mephitis, and
Conepatus I could find neither swelling nor vibrisse.

The Anus and Heternal Genitalia.

So far as is known, the normal pair of anal glands is present in
all genera of the family, but they vary greatly in development.
In many cases, 7. ¢. in Martes, Gulo, Tayra, Lutra, Taaxidea*,
Meles, the glands are not enlarged and the secretion is not par-
ticularly offensive in scent to Mah, and is not apparently used by
the animal itself for defensive purposes. But in other genera,
like Mellivora, Grison, Ictonyx, Mephitis, Conepatus, the glands
are enlarged, the secretion is copious, and is ejected at an enemy
in case of attack or when two individuals of the same species fight
together. And these genera are unusually coloured, being either
grey above and black below or striped with white along the
dorsal area tT. Putorius also is accredited with the same habit,
although the scent of the secretion is not so pungent and lasting
as in the other genera mentioned. Nor is the coloration so
striking; but the underfur is creamy white when the coat is
lifted and this pallid hue often pervades nearly the whole body ¢.

* At least in the single female example I examined.

+ See my paper, “ The Warning Characters of Musteline Carnivora” (Proc. Zool.
Soc. 1908, p. 955). In this paper I threw out the suggestion that the similarity in
colour between Tayra and the Canine genus Speothos might be an instance of
mimicry, on the supposition that the secretion of the anal glands in Tayra might
turn out to be offensive as it is in Grvison. Since subsequent ohservation has shown
this not the case, the suggestion must be withdrawn. It does not seem likely that
Speothos is the offensive model mimicked by Tayra.

{ See my paper on the warning coloration of P. nigripes (Proc. Zool. Soc. 1911,
pp. 559-560).

828 MR. R. I. POCOCK ON THE

The anus itself is typically exposed as a button-shaped convex
disc, everywhere surrounded by hair as in Martes, Mustela, Gulo,
Lutra, Ictonyx, and others; but in some eases, as in Mellivora,
it is sunk in a circular thickening of corrugated integument,
whereas in Meles there is a more delinite thinner-walled pouch
round it. In Vayra, again, there is above the anus a shallow
depressed area, bounded on each side and posteriorly towards the

Text-figure 38.

\\ \ Ny,
in

az
Wee

BEEN
— 5 y
are SSA \\
\ \ \

Re ¥
Wimes
yf 0

Pye

¥
y Y

GOs")

YE ZEB

C= ZZ

at
eZ:
4

j i sy)

te
=a

A. Anal region of Tayra barbara g, showing the naked skin around the
anus and the depression above it.

B. The same of Melogaie personata Z, showing the scrotum close beneath
the anus.

C. The same of Grisonella sp. from Oordova, Argentine.

root of the tail by a low ridge of skin. Possibly some modi-
fication similar to this was the origin of the deep subcaudal
glandular pouch present in WMeles and, according to accounts, in
Arctonyx. (Text-fig. 38, A-C.)

The perineal region between the anus and vulva or scrotum is
thickly overgrown with hair in Gulo, Martes, Mustela, Ictonyx,

EXTERNAL CHARACTERS OF THE MUSTELIDA. 829

and Lutra. It is similarly hairy in a male Grisonella, but in a
female of the same species it is naked or scantily hairy. It is
also scantily hairy in Mellivora, Tayra, Taxidea, Meles, Mephitis,
and Conepatus.

In all cases examined the vulva is a fusiform excrescence with
a vertical slit-like orifice, bounded on each side by two labia
enclosing the clitoris and the urinary and genital orifices. (Text-

fig. 39, A-C:)

Text-figure 39.

A. Anus and vulva of Coxepatus proteus.
B. The same of Grisonella sp. from Cordova, Argentine.
C. The same of Plesiogale nudipes.

The penis is very variable in shape, in accordance with the
shape of the baculum; but the prepuce is always situated well
in advance of the scrotum, as in the Canide, Procyonide, and
Ursidee.

The Classification of the Mustelide.

In the classification which follows I have adopted the method
employed by Gray of combining analytical headings, which give
prominence to particular characters, with the definitions ordinarily
found in the text-books, where the characters are set forth as if
of equal value. For the interpretation of the table it is necessary
to remember that contrasted headings are marked by the same
letter—e. g., a—a’, b-b', and so on. I have adopted all the sub-
families originally proposed and defined by Gray and Gill, and
consistency in treatment has compelled the addition of several
new groups of that rank. It is not claimed that they are all of
equal value. The Guloninz, for instance, have been admitted
rather as a concession to the opinion of a distinguished contem-
porary than as a mark of my conviction that the characters
deserve the rank assigned to them. Systematists of the old

830 MR, R. I. POCOCK ON THE

school will probably criticise the classification on the grounds
that most of the subfamilies are monotypical. That objection
is, however, 12 my opinion, illogically hollow. Difterentiatmg
characters and degrees of affinity are in danger of being com-
pletely lost sight of, if structural variations are disregarded
because of their restriction to an isolated genus represented by
two or three, or even a single species. And I can see no logical
halting-place between some such amplification of Gill’s classifica-
tion as that here proposed and the classification of Turner (Proc.
Zool. Soc. 1848, p. 86), who admitted no names for subordinate
sections in the group. Hither method of dealing with the —
numerous genera 1s preferable to the superficial and dissatisfying
system proposed by Flower, to which recent authors have
adhered.

a. Hind feet larger than fore feet *; head, fur, tail, and limbs modified in
the normal mammalian fashion for aquatic life; kidneys lobulate.

Subfamily Lurrinm Gray & Gill.

Skull shaped very much as in the typical Musteline; cavity
of bulla, where known, as in Martin, but with roof depressed
upon auditory annulus and not extending to paroccipital process ;
palatine foramina maxillary. Basicranial and basifacial axes
approximately in one plane. Teeth powerful, upper carnassial
with talon large, nearly as long as blade; molar wider than long,
but about as large as carnassial; lower carnassial with long wide
heel and large metaconid. Hind feet with normally proportioned
digits, the second and fifth much shorter than third and fourth.
Tail long.

Genera Lutra, Hydrictis, Lutrogale, Amblonyx, Aonyx, Paraonyx,
and possibly others.

In my paper on ‘‘ The External Characters of some Species of
Otters” (Proc. Zool. Soc. 1921, pp. 535-546), reasons were given
for regarding the Otters as modified descendants of the Musteline-
Martine stock of the Mustelide rather than of the Meline stock
as suggested by Miller.

Subfamily Laraxin#, nom. nov.
(=Enhydrine Gray & Gill.)

Differing from the Lutrine in having the hind feet very large
and paddle-like, with digits subequal, but the fifth slightly longer
than the third or fourth; the tail short, the basicranial axis

depressed at an angle upon basifacial axis; the angular of the

* Flower’s frequently copied statement that the feet in the Lutrinse are “ short
and rounded” is unintelligible. The feet of the Meclinz, on the contrary, he described
as *‘ elongated,”

CLASSIFICATION OF THE MUSTELIDA. 831

mandible remote from the condyle; the teeth with rounded cusps,
and the median lower incisor suppressed.

Genus Latax.

No one can doubt that Latav is a Lutrine profoundly modified
in adaptation to marine life. The change in its method of feed-
ing, from the capture of swift-swimming fish to the picking up
and crushing of sessile molluscs or Bigae moving crustaceans, has
modified the teeth and skull in the way indicated ; aaa the
increase in size and alteration in structure of the heat feet,
- accompanied by loss of length in the tail, suggests the need for
skiiful turning amongst euinmney ged rocks, rather than for swift

progression through the water and comparative activity upon
the land.

a'. Hind feet usually smaller, never larger than fore feet ; no structural
modifications subservient to aquatic life; kidneys not lobulate.

b. Cavity of bulla completely divided into two subequal chambers by oblique
partition passing from the stylomastoid foramen forwards and inwards
to the carotid foramen.

Haxictipina Gray & Gill.

Bulla not communicating with periotic hollow, its roof close to
auditory annulus. Upper carnassial with large bicuspid talon ;
molar wider than long, smaller than carnassial ; lower carnassial
with moderate heel and large metaconid. Bhinarium very deep
below nostrils, lateral shit of nostrils long and complete ; no phil-
trum ox groove on upper lip. Har with well- -developed bursa.
Feet fossorial, naked below, but narrow, with webbing only
extending to proximal end of digital pads, which are widely
separable ; all the pads coarsely striate ; two long metatarsal pads.
No anal pouches. ;

Genera Helictis and Melogale.

Although Helictis has been allowed by almost universal consent
to retain a place in the Melinz, where it was placed by Flower,
recent examination has justified its separation from that sub-
family, as I suggested last year, and its assignment to a special
subfamily as proposed by Gray and Gill. The auditory bulla is
quite peculiar, and the teeth are neither Meline nor Musteline ;
the rhinarium may be described as Meline and the ear as Muste-
line; the feet, although fossorial in the matter of the claws, are
uot so specialised for digging as those of Meles ; the tail, moreover,
is long and there is no trace of the subcaudal pouch found in that
genus. Apart from the auditory bulla Helictis is a less specialised
form than Meles, and is a type intermediate between the “ fos-
gorial” and ‘‘non-fossorial” members of the Mustelide. Its
trivial name Ferret-Badger is not inappropriate.

832 MR. R. I. POCOCK ON THE

b’. Cavity of bulla, when divided, separated into an anterior larger and a
posterior smalier chamber by a transverse partition running from the
stylomastoid to the foramen Jacerum posticum.

ec. Walls of bulla everywhere thick and permeated with air-cells continuous
with those ot the mastoid, the cells of which may communicate with
the cavity of the bulla.

Subfamily MusTE.in@, s. s.

Skull with long cranial and short facial portions. Teeth
sectorial, talon of upper carnassial small, anterior ; molar much
wider than long; lower carnassial with cuspidate heel, meta-
conid retained or absent; pm. 1 lost above and below. Tympanic
annulus in contact with rceof of bulla. Palatine foramina
maxillary. Rhinarium small. Bursa of ear marginal, large or
moderately so. Feet cursorial, usually hairy below, no metatarsal
pads; other pads moderate or small, coarsely striate, claws short
and curved ; digits webbed to proximal end of pads, the 2nd, 3rd,
and 4th widely separable. Baculum with a narrow, long, deep
groove in its distal third beneath, ending in a median rounded
apex.

Genera Mustela, Gale, Plesiogale, Putorius (with such subgenera
of one or the other as Zutreola and Kolonokus). Vormela.

The restriction of the term Musteline to the Stoats, Weasels,
and Polecats is justified by the number of characters by which
they differ from other members of the Mustelide. The group
contains a larger number of species than any other subfamily, |
and, apart from the Lutrine, is more widely distributed. When
the tropical forms are better known, other genera may be added;
but perhaps characters negativing some statements in. the
description of the subfamily may come to light.

ce’. Walls of bulla thin, its cavity either closed behind or opening into
a spacious periotic hollow.

d, Cavity of bulla closed behind.

e. Teeth sectorial, approximately as in Musteline, the upper carnassial
being longer than wide, with the talon anterior and narrow-necked,
and the molar wider than long, etc., and the line of the upper incisors
nearly straight. Feet scansorial and cursorial, with short sharp claws.

Subfamily MARTIN«, s. s.

Two carpal pads separated by hairy space from narrow plantar
pad, digital webs hairy below; no metatarsal pads. A well-
developed marginal bursa on ear. No trace of pouch above anus.
Pm. 1 retained above and below. Baculum with two or four
‘branches.

Genera Martes and Charronia.
Although usually associated with the Stoats, Weasels, and

CLASSIFICATION OF THE MUSTELID. 833

-Polecats, which they generally resemble in the structure of the
feet and teeth, the Martens differ from them principally in
the structure of the auditory bull, but also in their longer jaws,
less sectorial teeth, the retention of pm. 1, and the position of
the posterior palatine foramina on the suture.

Subfamily Guitoninaz Gray & Miller.

Chiefly distinguishable from the Martine by having the pre-
orbital foramen in front of the orbit, the bulle small, the auditory
tube long, the occipital region short, the mastoid large, the upper
anterior premolars separated from the lower, and the metaconid
of the lower carnassial suppressed.

Genus Galo.

Subfamily Tayrin#, nov.

Distinguishable from the Martine and Gulonine by having
the two carpal pads fused into a large mass as large and as wide
as the large plantar pad and in contact with it, by the presence
of a large metatarsal pad and naked interdigital webs and a very
shallow bursa remote from the posterior edge of the ear, by the
loss of pm. 1 above and below, the presence of a shallow subeaudal
pouch, and an apically unbranched baculum.

Genus Tayra (=Calera).

The Tayra was for quite unintelligible reasons formerly
regarded as congeneric with the Grison, the two being quoted as
ralictis, They are at least as different as the Polecat is from
the Marten.

e’. Teeth not of the sectorial type, the upper carnassial about as wide as

long, with the talon median and arising by a long base from the blade ;
the molar about three times the size of the carnassial, as long as wide;
lower carnassial with a long wide heel, and the metaconid as large as
the other cusps; the upper incisive line curved, Feet fossorial, with
long powerful claws.

Subfamily Muti, s. s.

Two carpal pads separated by a naked or hairy space from the
wide plantar pad ; interdigital webs naked below, narrow between
the digital pads; third and fourth digits of hind foot in contact ;
a well-developed, partially divided metatarsal pad. No trace
of bursa on ear. Rhinarium very deep beneath lateral slits of
nostrils. A deep subcaudal glandular pouch.

Genera Meles and Arctonyx.

I have seen no fresh specimen of Arctonyw, but the external
form is exceedingly Badger-like. The teeth, too, are very like
those of Meles. The cranial differences Letween them are well
known. In the extension of the mastoid below the auditory

834 MR. R. I. POCOCK ON THE

ovifice the two genera appear to be peculiay in the family,
although this character is foreshadowed in Galo.

d’, Cavity of bulla opening into a large or very large hollow space in the
mastoid portion of the periotic.

f. Prlate produced posteriorly, so that the mesopterygoid fossa is far
behind the molaz teeth ; baculum, where known, a stout bone.

g. Rhinarium a subcireular flattish disc, continued to the edge of the upper
lip by a narrow long philtrum resembling it in texture; nostrils
anterior, above middle of rhinarium ; the lateral slits almost obliterated.
Feet fossorial, with pads of 2nd, 3rd, 4th, and 5th digits united nearly
to their apices.

Subfamily Mypatn= Gray.

Skull with muzzle and dentition much as in the Meline, but
with frontal postorbital processes absent, zygomata and mandible
slender, and mastoid not produced below auditory orifice. Cavity
of bulla ceasing just behind petrous portion of periotic, and
opening laterally into hollow of mastoid portion of periotie by a
single orifice as in the Mephitine. Pinna of ear reduced to
a simple rim. No subeaudal pouch.

Genus Mydaus.

I have seen no fresh specimen of this genus, and the structure
of the baculum, if developed, is apparently unknown. The
Teledu has well-marked peculiarities, especially in the structure
of the rhinarium and feet. The rhinaritm is unique; and the
only genus which shows an approach to the fusion of the digital
pads is Conepatus. The latter also has the pinna of the ear
greatly reduced with the orifice exposed, and the bulla of Mydaus
seems to resemble tolerably closely that of the Mephitine ; but
in other respects, particularly in the long jaws, the curved upper
incisive line, the long posterior palate, the backward position of
palatine foramina, the genus more nearly approaches the Meline.
If Ferret-Badger is a good name for Helictis, perhaps Skunk-
Badger would be a suitable appellation for Mydaus.

g’. Rhinarium normal, nostrils much nearer its lower than its upper edge,
the lateral slits long; no trace of philtrum and no groove on upper
lip. Pads of 2nd, 8rd, 4th, and 5th digits free.

h. Tympanic annulus projecting as a free ridge into bulla, not confluent
with its roof, the posterior rafter arched high above petrous, leavmg a
large space between the chambers, of which the posterior is not multi-
loculate; bullse encroaching on glenoid in front.

i. Teeth sectorial, ina general way like those of Martine ; occipital crest
inclined backwards and inwards from mastoid, not semicircular from

back view.
Subfamily Mptiivorin® Gray & Gill.

Periotic hollow, comparatively small, not extended to tip of
mastoid, which, like the paroccipital, stands prominently away

CLASSIFICATION OF THE MUSTELIDA. 835

from bulla, the latter rounded in front and remote from hamular ;
palatine foramina on suture. Lower carnassial without meta-
conid; last lower molar lost; upper molar with wide cingulum.
Feet very broad, digits closely tied, with large carpal and meta-
tarsal pads in contact with wide plantar pads. Pinna of ear
reduced to an integumental thickening. A glandular pouch
round anus.

Genus JMellivora.

Subfamily Lcronycuin 2”.
(=Zorilline Gray & Gill.)

Periotic hollow larger, extending to tip of mastoid, which, like
the paroccipital, is close to the bulla; the latter pointed in front
and fused with hamular; palatine foramina on maxilla. Lower
carnassial with high acute metaconid ; last lower molar retained ;
upper molar without wide cingulum. Feet narrow, but claws
long, the digits widely separable; pads small, carpals separated
from plantar pad; metatarsal absent. Ears with large pinna.
Circumanal integument unmodified.

Genera [ctonyx (Zorilla) and Pecilictis. (? P Kecilogale.)

i’. Teeth not sectorial ; upper carnassial enormous, with very large bicuspid
talon; molar a little smaller than carnassial, triangular, its inner and
anterior borders subequal as in Meline; lower carnassial with large
cuspidate heel and large metaconid; occipital crest rising nearly
vertically from mastoid, and nearly semicircular from posterior aspect.

Subfamily Taxiprn2 Pocock.

Feet broad and fossorial with long claws as in Mellivorine,
but plantar pads narrower, a single small carpal pad remote from
plantar pad, and no metatarsal pad. No glandular pouch
associated with anus. Ear with pinna well developed.

Genus 7J'axidea.

h’. Tympanic annulus confluent at its summit with roof of bulla; posterior
aatter-like partition of the bulla close down upon the petrous, leaving
a narrow passage between the two chambers, the posterior of which is
markedly multilocular; bullzes remote from glenoid in front.

Subfamily Grisonin#, nov.

Teeth as in Musteline, but talon of upper carnassial much
larger, occupying nearly half the inner surface of the tooth ;
palatine foramina maxillary. Feet not fossorial, almost as in
Tayrine, but with carpal pads farther from plantar pad. Pinna
of ear well developed with bursa present, but small and in front
of posterior margin. No subecaudal pouch.

Genera Grison, Grisonella.

836 MR. R. I. POCOCK ON THE

Subfamily Lyncoponrin®, nov.

Differing from the Grisonine in cranial and more particularly
in external characters. Bulle inflated, not flattened externally
towards auditory orifice, with anterior edge obliquely transverse,
not emarginate ; mesopterygoid fossa longer ; pnv. 2 lost above and
below. Feet with long fossorial claws, but unique in the family ;
digits only webbed for a short distance beyond plantar pads ;
hairy laterally, mesially naked below, like the reduced webs.
Carpal pads reduced to the external moiety which is m contact
with the plantar pad, the rest of the carpal area covered with
hair, which encroaches over the middle of the plantar pad; no
trace of metatursal pads; plantar pad of hind foot reduced like
that of fore foot. Upper lip completely grooved. Har greatly
reduced, no trace of bursa or of valvular supratragus.

Genus Lyncodon.

I have seen no fresh examples of this small Patagonian genus,
my observations having been made upon dried skins and skulls
in the Natural History Museum. The coloration is very like
that of Grison, with which Lijncodon was affiliated by Matschie,
and I suggested that the likeness might be a case of Miillerian
‘or genuine mimicry (Proc. Zool. Soc. 1908, p. 953). It cannot
be claimed that the genus is closely related to Grison. Even the
evidence that it 1s a very aberrant form of the Grison-group is
not convincing, despite the resemblance between the two in the
internal structure of the bulla suggested: by my unavoidably
imperfect examination of this portion of the skull in Lyncodon.
The feet differ from those of Grison at least as profoundly as the
feet of Mustela or Maries differ from those of JMJeles or Mellivora.

jf’. Palate not produced posteriorly, the mesopterygoid fossa reaching
almost up to the line of the upper molar teeth. Baculum unossified or
ossified as an exceedingly slender undifferentiated rod.

Subfamily Mepnmm#« Gray & Gill.

Cavity of bulla simple, ceasing close behind petrous, communi-
cating with large periotic hollow by means of a small circumscribed
orifice set forwards in a line with the outer end of the petrous
and in front of the stylomastoid foramen, which is set far in and
narrowly separated from the foramen lacerum posticum. Bulla
not nearly extending to paroccipital, but encroaching on glenoid
infront. Muzzle massive, but zygomata and postor bital processes
weak. Teeth resembling those of the Melinz and Mydaine, but
the upper Incisive row nearly straight, the upper molar not twice
the size of the carnassial and wider than long; pm. 1 lost above
and below; palatine foramina maxillary. Feet fossorial, with

CLASSIFICATION OF THE MUSTELIDA. 837

soles naked, two carpal pads, and metatarsal pad; pads of 3rd
and 4th digits of hind foot basally united. Har with orifice
not concealed behind tragal ridge. No pouches associated with
anus.

Genera Spilogale, Mephitis, Conepatus.

Although the Skunks constitute a well-developed subfamily,
they seem to be linked in a measure with the Meline through
Mydaus. The three genera show considerable range in structure,
Spilogale being the least, and Conepatus the most, specialised
form, the former standing nearest to the more typical members
of the Mustelide, such as /ctonyx, and Conepatus, in its feet, ears,
and rhinarium, foreshadowing the peculiarities of Mydaus.

eas

ON FOSSIL HYRACOIDS FROM THE FAYUM. 839:

42, Megulohyrax Andrews and Titanohyrax, g. n.—A Revision
of the Genera of ae from the Haye tim, Hgypt.
By H. Matsumoto *

Received October 18, 1921: Read November 22, 1921. |
(Text-figures 1-6.)

In the course of my re-study of the fossil Hyracoidea of the
Fayum, Egypt, based upon the material belonging to the American
Museum and the British Museum, I found that Schlosser’s
identification of Andrews’s genus Megalohyrac is incorrect, ‘the
type-specimen of the genotype Meg. eocenus Andrews belonging
not to Schlosser’s “ Megalohyraw,” but to his ‘* Mixohyrax.”
Consequently, his “iW twohyres” should be called Megalohyrax,
while his *‘ Megalohyrax” wants a new generic name. I propose
to call it Titanohyrax, g. n.

I have the pleasure to express here my hearty thanks to
Dr. Arthur Smith Woodward and Dr. Charles W. Andrews,
by whom I was permitted to examine the material belonging to
the British Museum.

Genus MreGgaLtouyrax Andrews.

Megalohyrax Andrews, Geol. Mag., Decade iv. vol. x. 1903,
p. 841; Andrews, Brit. Mus. Cat. Tert. Vert. Fayum, Egypt,
1906, p. 92.

Mixohyrax Schlosser, Zool. Anz., Bd. xxxv. 1910, p. 502;
Schlosser, Beitr. z. Pal. u. Geol. Osterreich-Ungarns u. d. Orients,
Bd. xxiv. 1911, pp. 98.& 115.

This genus is to be re-diagnosed as follows :—

Long-skulled and long-snouted. Premaxillze long, but not
especially elongated supero - posteriorly. Nasals long, with
acutely-pointed anterior ends, which le nearly as anteriorly as
the anterior ends of premaxiile and a certain, but not very great,
distance in front of the anterior ends of naso-premaxillary sutures.
External nares not retired at all. Upper surface of skull very
rough, with irregular pits and grooves with intervening ridges.
Mandibular ramus shallow, with a large ee like opening on
the inner surface. Dental formula : oe ee {i and I, enlarged
and tusk-like. Cheek-teeth bunoselenodont, ee yea
Upper Pay and _ usually also P’, three- -cusped ; P*_M” , and

* Communicated by Dr. C. W. AnpReEws, F.R.S., F.Z.S.
Proc. Zoot. Soc.—1921, No. LVI. D6

840 MR. H. MATSUMOTO ON THE GENERA OF

very weak, and especially so in the more postcrics premolars and
in the oe Mesostylar fold absent in Pon Parastylar fold
of P™ * very feeble; that of P rather Le both parastylar

and mesostylar folds of M~ ° especially strong, rounded, and
stout. Ridges of lower P,—M, W-shaped. Metaconid simple,

without differentiated fae Pistol: Posterior talon of M, very
large.

Genotype: Megalohyrax eocenus Andrews, 1903; non Schlosser,
1900

This genus includes Megalohyrax minor Andrews, 1904;
Mixohyrax niloticus Schlosser, 1910; Mix. swillus Schlosser,
1910; and Megalohyrax pygmeus, sp. n., besides the genotype.

Synopsis of Species of Megalohyrax.

(1) Extremely donee species ; length of upper Ban

and of M * vespectively 75 mm. and 86 mm.
CAndrewsisrtty ye: MesSo02 et Ev. iak ene eoceenus.

(2) Large species; length of upper P!4 and of uM
respectively 63 (Andrews’s type: C. 8818)—64
(Am. Mus. 13332) mm. and 74 (Andrews’s
type)—78 (Am. Mus. 13332) mm.; that of

lower P,, and of M,,, ca. €9 (Am. Mus.

13345)—70 (Schlosser) mm. and ca. 78 (Am.
Mus. 13338)—85 Boe Am. Mus. 13345)

MINA, TESPCCLUVELY <8 esse silo teal sitcene chee [ss weneee minor.
(3) Rather ie ge species; length of upper P** and

of M’” respectively ca. 54 mm. and ca. 62 mm.
(Schlosser’s fig.) ; that of lower P, | and of M_ ,,

55 (Schlosser)—57 (Schlosser; Am. Mus.

13334) mm. and 68 (Schlosser)—75 (Am.

Mis; 13334)/mm. respectively = 20 iy eas . niloticus.
(4) Small species; length of lower Spe) 46 mm.

(Schlosser) ; that of lower M, ais 32 mim.

(Schlosser), whereas the same measurement
in the immediately preceding and the next
species is 42 mm. and 24-25 mm. respectively . swillus.

(5) Very nel species; length of upper P'* and

of M™ * respectively 35°5-36 mm. and 37 mm.
(type-specimen: Am. Mus. 14454); that of

lower ae and of ey 3° 31°5 (M. 9480)--33

(type-specimen) mm. and ca. 38 (Am. Mus.
14463) —-40 (type-specimen) mm. respectively . pygmcus.

FOSSIL HYRACOIDS FROM THE FAYUM. 84)

(1) Megatonyrax rocanus Andrews,

M. eocenus Andrews, Geol. Mag,, Decade iv. vol. x. 1903,
p. 340, text-fig. 1; Andrews, Brit. Mus. Cat. Tert. Vert. Fayim,
Egypt, 1906, p. 92, pl. vi. figs. 1 & 2 (non text-fig. 39).

M. 8502; type-specimen ; fragment of right premaxilla and
maxilla, bearing C—M® in situ.

Schlosser has failed to understand this type-specimen, hence
also to identify this species and this genus. He had the opinion
that the teeth of this specimen, as figured by Andrews, were
erroneously restored. According to my actual observation of
this specimen, Andrews’s illustration proves to be fundamentally
correct, though the measurements of the teeth of this specimen
were confusingly misprinted (width as “length,” and length as

“width” in P°— M). The more important dental characters
of this specimen are:—(1) The premolars have no proper
mesostyle (a distinctive character from Zitanohyraa), though
the external basal cingulum of ieee (unknown in P*) has a
raised tubercle just outside the metacone. (2) The parastylar
and mesostylar folds of the molars are very strong, rounded, and
stout (a distinctive character from Zitanohyrax). (3) The para-
conal and metaconal folds of the molars are exceedingly feeble
and insignificant (a diecinetive character from Geniohyus and

Bunohyrax). (4) The P’” have no hypocone; P’, a very rudi-
mentary one; and Pp’, a more or less well-developed one (in
Gentohyus and Bunohyraaw, all P’ have no hypocone ; in certain
other species of Megalohyraa, Pry hake no hypocone and P* has
one; and in Trtanohyrax, all P’* have a well- developed PEGs
cone). (5) The posterior surface of the protocone of PM
is not rounded, but is provided with a well-rounded ridge; the

anterior inner surface of the hypocone of M’” is not rounded,
but is provided with a well-developed metaloph, which is as
strong as or even stronger than the ridge on the posterior surface
of the same cone (distinctive characters from itanohyrazx).
(6) The surface of enamel of the teeth is not smooth, but very
rough (a distinctive character from Titanohyr an). These
characters indicate that Megalohyrax typified by this specimen
does not correspond to Schlosser’s “‘ Megalohyrax” (= Titarohyrax)
at all, but to his “ Mixohyrax” (= Megalohyrax).

As Andrews’s measurements of the teeth of this specimen were
confusingly misprinted, I here give the measurements taken by

me (in mm.) :—

56%

842, MR. H. MATSUMOTO ON THE GENERA OF

Length. "idth.

Oe ta sc 2 ee 18 Ios
EPH 2d Ss, AUPE repo 17 18
D2 ee aa ames 17 AAlliess,
Pe ac tae ee tea S25 21°5
1S ePU NRE iin aaa 1) a=
ITU Meer oe ea 24 —
MG Wake fae 28-5 a
Miah! yaneatia 34 37

Length of | eee eat Ge Sey 75

Leneth of Ms tek ia SO

(2) MecaLoHyrax minor Andrews,

M. minor Andrews, Geol. Mag., Decade v. vol. 1. 1904, p. 213 ;
Andrews, Brit. Mus. Cat. Tert. Vert. Fayim, Egypt, 1906, p. 97
(vars), pl. vil. fig. 1 (non figs. 2 & 3).

Minohyrax andrewsi Schlosser, Zool. Anz., Bd. xxxv. 1910

Ly ’ ; D
p- 503; Schlosser, Beitr. z. Pal. u. Geol. Osterreich-Ungarns u. d.
Orients, Bd. xxiv. 1911, p. 115, pl. x. (i1.) figs. 9-11.
C. 8818; type-specimen; fragment of left maxilla, bearing
1 Mare :
P—M in situ.

Schlosser kept the specific name “ minor” for a certain speci-
men of Andrews’s and adopted a new name, “ andrevwsi,” for the
present species, which includes this type-specimen of J. minor.
Such a statement as Schlosser’s is, of course, against the law
of priority: the name “ minor” must be kept for the present

; : Saale) 13 ; :
species. The length of P and of M of this type-specimen is
63 mm. and 74 mm. respectively.

M. 9478; fragment of left mandibular ramus, bearing Poe
in situ, which are strongly worn.

The mandibular ramus measures 52 mm. and 64 mm. in
depth at the anterior side of P and of M, respectively. The

teeth measure as follows (in mm.) :—

Length. Width.
Dense ade ts -s 18 12°5
MOS. t ae 22°5 14:5
Wy cugeu sdadade oss 25°5 fh
We Je se paliadoacinoan 38 19
dfengthvor Ws nese as 85

M. 9419; small fragment of left mandibular ramus, bearing
M, in site. The molar measures 26 mm. in length and 17 mm.

in width.

FOSSIL HYRACOIDS FROM THE FAYUM. 843

(3) MrcaLonyrax NiLoricus (Schlosser).

Mixohyrax niloticus Schlosser, Zool. Anz., Bd. xxxv. 1910,
p- 503; Schlosser, Beitr. z. Pal. u. Geol. Osterreich-Ungarns u.
d. Orients, Bd. xxiv. 1911, p. 116, pl. xi. (iii.) fig. 9, pl. xii. (iv.)
figs. 3 & 6, pl. xv. (vil.) figs. 1, 4, & 8.

Mixohyrax suillus Schlosser, loc. cit. 1911, p. 118 (pars), pl. x.
(ii.) fig. 6.

(4) MrGaLoHyRax surLLus (Schlosser).

Mixohyrax suillus Schlosser, Zool. Anz., Bd. xxxv. 1910,
p. 503; Schlosser, Beitr. z. Pal. u. Geol. Osterreich-Ungarns u.
d. Orients, Bd. xxiv. 1911, p. 118 (pars; non pl. x. (ii.) fig. 6).

(5) MEGALOHYRAX PYGMUS, Sp. n.

Saghatheriwm magnum Andrews, Geol. Mag., Decade v. vol. iv,
1907, p. 99 (row Andrews, 1904), text-fig. 2.

For the full description of this new species, see another
paper of mine, now in preparation. The type-specimen and
paratypes of this species belong to the American Museum. I
will cite here only one specimen belonging to the British
Museum, which is to be referred to this species.

Text-figure 1.

Megalohyrax pygmeus, sp. n., Matsumoto. Crown view of left lower C—M3;
natural size; M. 9480; referred specimen, not the type.

M. 9480; left mandibular ramus bearing C—M., in situ (text-

fig.1). he mandibular ramus is very shallow (though crushed just
a little, it can by no means be restored to a ramus so deep as in
Saghatherium) and has a large fenestra-like opening on the inner
surface (distinctive characters from Saghatherium).

A fall description of this new species is given elsewhere.

The measurements of the teeth were as follows (in mm.) :—

Length. Widtt.
Co ees eieeh le Ab 2-7
Ba eae dal 7 4-4
| ra aut oar 77 5
Bafa cr tention 8 6
Papadive: peers 4 8-7 7-2
M, ub te adiee HORNE ie 10:7 (is)
M, eno Sereye ae 11:8 8°D
M, PE AR Nt arate 17:5 9°6

Length of PD igcreteesnecnee sense 31°d

Length of M_ ..........-.- seen ee

844 MR. H. MATSUMOLO ON THE GENERA OF

Genus TITANOHYRAX, g. n.

Megalohyrax Schlosser (now Andrews, 1903), Zool. Anz.,
Bd. xxxy. 1910, p,. 9025 Schlosser, Beitr.) zal. jusitneal:

Osterreich -Ungarns u. d. Orients, Bd. xxiv. 1911, pp. 97 & 104.

This genus is diagnosed as follows :—

Skull very imperfectly known (laying aside the problematical
specimen CU, 8801, which may possibly be referred to this genus),
but judging from the shape of premaxile and mandible
and from the dental series of both jaws, this genus might be
more or less short-skulled and short-snouted. Premaxille
exceedingly elongated supero-posteriorly ; their anterior ends
lying a ‘eveat distance in front of the anterior ends of naso-
premaxillary sutures. Judging from the shape of the premaxille,
the external nares may be retired backwards instead of being
terminal. Mandibular ramus shallow, without fenestra-like

: ae ees .
opening. Dental formula: on; IT enlarged and tusk-like ;
none of lower incisors tusk-like. Cheek-teeth selenodont,
: 1 3

brachyodont, though rather high. All upper P —-M° four-cusped.
Paraconal and metaconal folds somewhat indistinct except
eres 1 3

in P. Mesostylar fold present in all P—M’. Parastylar and
mesostylar folds trenchant instead of being rounded. Ridges of
lower Ds W-shaped. Metastylid well differentiated from
metaconid. Posterior talon of M 2 rather small.

Genotype: Iegalohyrax paleotherioides Schlosser, 1910.
This genus includes 7’. ultimus, sp. n-; 7’. schlossert, nom. n. ;
and 7’. andrewsi, sp. n., besides the genotype.

Synopsis of Species of Titanohyrax.

(1) Extremely gigantic species, being the largest of
all the hyracoids hitherto known, upper and
lower M 2 measuring about 40 mm. in length,
whereas those of the next species measure
abode 3 Oma. sli ce ets Bent ales, asa, Aken ultimus.

(2) Cae species; length of upper P'* and of
M” respectively *0 mm. and 84% mm.
(Schlosser); that of lower P, , and of M_ .,
ca. 70mm. and 94 mm, respectively (Seblorsel), schlosseri.

(3) Large species; length of upper P*“ and of M*”
respectively ca. 73 mm, and 75 mm. (Schlosser) ;
length of lower M, 22 mm. (Schlosser), whereas

the same of the immediately preceding and the
next species measures 24°5 mm. and 20 mm.
respectively ; lower cheek-teeth of long and
narrow type; snout rather long, the distance

FOSSIL HYRACOIDS FROM THE FAYUM. 845

from the tip of mandibular symphysis to the
posterior side of lower Dm, measuring ca.

114 mm. in a young individual with functional
milk-molars (Schlosser’s fig.) ............+-. paleotherioides.

(4) Rather small species; length of lower M, ,

76 mm. (type-specimen: C. 8822-3); lower
cheek-teeth of short and wide type; snout
very short, the distance from the tip of
mandibular symphysis to the posterior side
of lower le measuring only 86 mm. in an old

individual with much worn premolars and
molars (type-specimen) ...........2-....0 PLN andrewsi.

(1) TiranoHyRAX ULTIMUS, sp. n.

M. 12057 ; type-specimen ; isolated, right upper M
M. 12058; paratype; isolated, left lower M,.

M. 12059; paratype; isolated, left upper Be
M. 9479; paratype; isolated, left upper C.

Text-figure 2,

Titanohyrax ultimus, g. & sp. n., Matsumoto. Crown view of right upper M?;
natural size; M. 12057 ; type-specimen.

The M’, the type-specimen, is strongly worn (text-fig. 2). The
acute mesostylar fold is a generic character ; the parastylar folds
are broken away in this specimen. The outer surface of both the
paracone and metacone are distinctly concave, without distinct
paraconal and metaconal folds, also a generic character. The
inner to posterior surface of the protocone and the anterior to
inner surface of the hypocone are smoothly rounded, without
any ridge, also generic characters. The anterior and inner basal
cingula are continuous to each other, extending from the

846 MR. H, MATSUMOLO ON THE GENERA OF

anterior side of the protocone to the anterior half of the inner
surface of the hypocone,

Text-figure 3.

Titanohyrax ultimus. Crown view of left lower Mz; natural size; M. 12058;
paratype.

The M, is moderately worn (text-fig. 3). The worn area, as well

as the lophids as a whole, is W-shaped. The differentiation of
the metastylid from the metaconid (a generic character) is rather
feeble, being feebler than that im the other smaller species of this
venus, The entostylid is differentiated from the entoconid, being
as well differentiated as the metastylid is from the metaconid ;
this differentiation is obviously more intense than that in the
other smaller species. The anterior and posterior basal cingula
are well developed; the latter has a distinct tubercle, just behind
the posterior transverse ridge and outside the entostylid. The
outer basal cingulum is feeble, being represented by a few rather
insignificant crenule between the bases of the protoconid and

hypoconid.

Text-figure 4.

Titanohyrax ultimus. Crown view of left upper P4?; natural size; M. 12059;
paratype.

The P’ is well worn (text-fig. 4). The parastylar and meso-
stylar folds are very acute. The outer surface of the paracone is
distinctly concave, while that of the metacone is rather flat. The
worn area of the protocone is oval, with an acute prolongation

FOSSIL HYRACOIDS FROM THE FAYUM, 847

exter-anteriorly. The inner to posterior surface of the protocone
is smoothly rounded, while the outer-anterior side of the hypo-
cone has a feeble ridge. The anterior and inner basal cingula
are well developed, but not continuous to each other. .

Text-figure 5.

Titanohyrax ultimus. Crown view of left upper C ; natural size; M. 9479;
paratype.

The upper C is moderately worn (text-fig. 5). It is ovoid in
palatal view. The worn area has an acute prolongation, which
stretches backwards. The inner cingulnm is well developed, ex-
tending from the anterior end to the posterior end of the tooth.

The teeth above described measure as follows (in mm.) :—

Length. Width.
Uren Oca 16-5 12-5
Pe a 39 34-5
Magee ene 4] 46
NIN wh 39 28

(2) TrrANOHYRAX SCHLOSSERI, nom. n.

Megalohyrax eocenus Schlosser, Beitr. z. Pal. u. Geol. Osterreich-
Ungarns u. d. Orients, Bd. xxiv. 1911, p. 105 (non Andrews,
1903), pl. xi. (ai.) fig. 7.

Judging from Schlosser’s description and figure, there is no
doubt that his specimens of his “ Megalohyrax eocceivus” belong
really to his ‘‘ Megalohyraw,” viz. Titanohyrawv, though Andrews’s
Megalohyrax eocenus does not. So that Schlosser’s specimens
want a new hame.

(3) TrrANOHYRAX PALHOTHERIOIDES (Schlosser).

Megalohyrax paleotherioides Schlosser, Zool. Anz., Bd. xxxv.
1910, p. 502; Schlosser, Beitr. z. Pal. u. Geol. Osterreich- Ungarns
udsOrients, Bdexxiv. 1911p. 106, pl. xi. (ii1.) fig. 1, pl. xi.
vanes 1.

(4) TrrANOHYRAX ANDREWSI, sp. n.

Megalohyrax minor Andrews, Brit. Mus. Cat. Tert. Vert.
Faytim, Egypt, 1906, p.97 (pars; non Andrews, 1904), pl. vil.
figs. 2 & 3 (non fig. 1); Schlosser, Beitr. z. Pal. u. Geol. Osterreich-
Ungarns u. d. Orients, Bd. xxiv. 1911, p. 105.

MR. H. MATSUMOTO ON THE GENERA OF

848

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aN Si} ul Sy, puv Fq) &yj—eq pu Ty 141M ‘snurer avpnqipwew gfe, Fo MotA Todd -ojounszey] “u “ds wp 3 Vsmarpuy vvwhyoungzyy,

9 OIG y-4XoT,

FOSSIL HYRACOIDS FROM THE FAYUM. 849

C. 8822-3; type-specimen (text-fig. 6); large fragment of right
mandibular ramus, bearing P,—M, i sitw, and large fragment of
left mandibular ramus, including a greater part of symphysis,
bearing ub His and P,—M, in situ; it is almost certain that these
two rami belong to one and the same mandible.

The symphysis of this mandible is quite different from that of
the other genera of the Fayfim hyracoids: it is rather short and
very deep, with the anterior lower side very steep; it rather
resembles the mandibular symphysis of a tapir. None of the lower

incisors is tusk like. Both I, are alike spatulate, with flat,

wide, and thin crowns; their roots are clavate, with constricted
proximal parts. Unfortunately, the part of the dental arch cor-
responding to I, and C is broken away in this specimen ; I, and
C must be very reduced, if they were present at all, as can be
judged from the very short distance from I, to premolars. The

mandible measures:as follows (in mm.) :—

Menethy of symphysis «yi... ost aan <5 Weal 50
De prot tnersanMiey ery csoh 2. ss 82. yeens sees 37
Length from tip of symphysis to posterior

end Of Pw... e erect eee eeee eee eee eeee ees 86
Depth of ramus at anterior side of P Beare AQ
Ditto at anterior side of M, Pe ear Meee 1 Do

All the cheek-teeth are moderately to strongly worn. The
entostylid is not well differentiated from the entoconid. All the

cheek-teeth are short and wide. The M,, , have a tubercle, as a

part of the posterior basal cingulum, just behind the posterior
transverse ridge.
The teeth of this specimen measure as follows (in mm.) :—

hight. Left.
Length Width
Length. Width. (transverse (antero-posterior
diameter in I). diameter in I).
LT ceeeseees — — 14:3 TD
Ty ceseeeeee — — aC ae
Dieteeceece Lud 11-4 (14) (roots). —
eee 17 13-5 16-5 135
LE ease 18 16°5-a. 18 15
Meese 19 16°5 20 16
ih rsa 935 19 - 93+5 18-5
Ni ae 325 19 an se

Length of Me fo OSM 76

850 ON FOSSIL HYRACOIDS FROM THE FAYUM.

M. 9220; paratype; fragment of left mandibular ramus, bear-
ing P,—M, in situ; P, M,, and M, being imperfectly repre-
sented. The mandibular ramus, as well as the teeth, is smaller
than that of the foregoing specimen. It may be possible that
this specimen belongs to a female, and the foregoing one to a
male.

The ramus measures as follows (in mm.) :—

Depth of ramus at anterior end of P) ............... 34
Ditto abjantertoniend of) MMPs er awe sy saree. Wau su 37
Ditto at posterior end of Mi int. .nttcrsseet-Weraee, 45
Height of ascending bar at condyle ..........,....... 120+
Minimum height of the same at the concavity

between condyle and coronoid process ............ 116
Minimum antero-posterior width of the same just

below comdiyile. 6,5. pach alc ts eine aan 5D

The teeth measure as follows (in mim, ) :—

Length. Width.
een ee A 16-5 12°5
Ps), Wiese = 14
2) ene et = 155+
EN bebe 238 21 18
I eee 30+ —

C. 8801; fragment of posterior part of skull, including occiput
(¢this genus and species).

It is almost certain that this skull, Ulustrated by Andrews in
his text-fig. 39, Cat. Tert. Vert. Fayiam, Egypt, 1906, does not
belong to either Geniohyus or Megalohyrax, since it differs very
much from the known skulls of these genera. Naturally, there
may be a possibility that it belongs to Titanohyraxv, and even to
the present species, though of course it is not certain at present.

ON THE VISCERAL ANATOMY OF THE MARSUPIALIA. 851

43. Contributions to the Visceral Anatomy and Myology
of the Marsupialia. By CHartes F. Sonnrac, M.D.,
F.Z.S., Anatomist to the Society.

[Received October 5, 1921: Read November 22, 1921. |}
(Text-figures 70-79.)

ConrTeENTs.

Page | Page
Wentnallesentciyeecee ster corte Coll) Urinary, Orcans) 9). 0- soe ene as Gil
NG MOSeHMissWeN terres eens Sole |) DuctlesssGlands),.. 15s.) ee SES
Cervical Lymphatic Glands ......... 851 | Respiratory. Organs .................. 870
Muscular System’. <6. cises ores 2 802 | Vagus and Sympathetic ............ 873
Viguilauee eee een pte etisaee §, SHANA MOIVSLEMALI CMs forests cleek. es aT,
Salivary Glands ........................ 857 | Summary and Conclusions ......... 880
PAimeniannys Ganley ey sae Ola stb IOs Lapiiys cv ses. dese sete ace) SOL
Organs of Circulation ............... 865

The present paper 1s based on the examination of several
Polyprotodont and Diprotodont Marsupials * which died in the
Society's Gardens; and the material comprised both fresh and
preserved specimens.

I have used my paper on the Koala and Vulpine Phalanger for
purposes of comparison (15), and I have drawn attention to the
differences between the anatomy of Perameles obesula and that of
Conolestes obscurus, recently described by Osgood (9).

When the abdomen of Dendrolagus ursinus was opened an
avascular peritoneal fold, representing the remains of the ventral
mesentery, was seen running from the pelvic floor posteriorly to
beyond the urinary bladder anteriorly. Beddard (1) observed a
similar condition in D. bennetti and Ornithorhynchus.

Adipose Tisswe:—Owen (10) states that the omentum of the
Opossums has no fat when there are accumulations elsewhere.
In Petaurus sciureus the omentum is heavily laden with fat, and
there are deposits in other parts. The fat in the anterior part of
the thorax contains the thoracic lobes of the thymus gland.

Cervical Lymphatic Glands :—The relative positions of the two
groups of glands in the Marsupialia have already been described
in my paper on the Koala (15), but the number of glands in each
varies. The anterior group has only one in Perameles obesula
and Cenolestes obscurus, but several are present in LDendrolaqus

* Didelphyide: Didelphys azare, D. marsupialis, D. albiventris, Metachirus
opossum, Philander laniger, Marmosa elegans; Dasyuride: Dasyurus viverrinus,
Sarcophilus harrisi; Peramelide: Perameles obesula; Macropodide : Macropus
bennetti. M. giganteus, M. rufus, M. melanops, Dendrolagus wrsinus, Onychogale
lunata, Bettongia penicillata; Phalangeridee: Petawrus sciureus, P. breviceps,
P. breviceps papuanus, Pseudochirus peregrinus, Phalanger orientalis, Petaurus
australis. Duplicates of some adult forms, and a number of pouch specimens of
different ages were examined. :

852 DR. C. F. SONNTAG ON THE VISCERAL ANATOMY

ursinus. Phascotarctos cinereus has the greatest number in the
posterior group, and the parathyroids may be included among
them.

The only Marsupial with exoccipital processes touching the
deep aspect of the platysma is Phascolarctos cinereus. ;

The Muscular System.

As considerable attention has already been paid to the muscles
of the Marsupialia, the present section is limited to some points
in the myology of the head and neck.

The Platysma :—The origin, insertion, and relations are similar
in all Marsupials, but the characters differ considerably. In
Dendrolagus ursinus and Macropus rufus it is muscular in the
face, but almost aponeurotic in the neck. In Perameles obesula,
on the other hand, it is muscular and thick in the neck, but more
aponeurotic and firmer in the face; and it arises posteriorly from
an equally strong and thick panniculus carnosus. In some parts
these muscles are nearly two millimetres thick, and it is difficult
to tell where the one ends and the other begins. The panniculus
is very thick over the pectoral region. These conditions are
reversed in Ceenolestes obscurus, 11 which Osgood described a thin

anniculus and platysma. The platysma of Surcophilus harrist
(text-fig. 70 A) is muscular throughout, and its fibres cross at the
anterior extremity of the interramal space. Beneath it there lies
a stratum of more or less transverse fibres which fuse with it at
the sides of the neck. Although the latter cannot be traced to an
attachment in the face, they may correspond to the sterno-facialis
of some of the higher Mammalia, e. ¢. Octodon and the Carnivora.
The condition present here may indicate that the sterno-facialis
arises from the platysma by splitting.

The Sterno-mastoid and Cleido-mastoid :—Carlsson (2) showed
that they are fused throughout the greater part of the neck in
Dendrolagus dorianus and Petrogale penicillata, but not in Tricho-
surus vulpecula; and they are similarly fused in Dendrolagus
ursinus. In a pouch specimen ‘of Macropus gigantews they fuse
much farther forwards; but the muscles of Petawrus sciurews run
parallel to one another, and are only fused at their cranial
insertions. Osgood (9) records a similar condition in Uewnolestes
obscurus, so it differs greatly from that in Perameles obesula, in
which the cleido-mastoid arises from the cephalo-humeral and fuses
with the cranial end of the sterno-mastoid. In Philander laniger,
Metachirus opossum, and Didelphys the sterno-mastoid, cleido-
mastoid, and cleido-occipital form a group of three almost parallel
muscles traversing the neck obliquely from behind forwards. In
ali species examined except D. azare they had separate origins
from the clavicle; but in that species the cleido-occipital was
implanted By tendon into the cleido-mastoid at its posterior
extremity.

The posterior triangle of the neck is obliterated in Dendrolagus
ursinus and Petrogale xanthopus, in which the trapezius is in

—— oe

AND MYOLOGY OF THE MARSUPIALIA, 853

contact with the sterno-mastoid. In all other Marsupials the
posterior triangle is evident, and its most prominent structure
is the external jugular vein. The descending cutaneous nerves
emerge from between the muscles in Dendrolagus (text-fig. 74).

The Omo-hyoid is present in all Marsupials, and is strap-like
or fusiform. A central tendon is present in MJacropus rufus (19)
and a slight one exists in Petrogale wanthopus (13). In a pouch
specimen of Macropus giganteus (text-fig. 71a) it crosses the
lateral thyroid lobe which projects beyond the pretracheal muscles,
and Osgood (9) figures it crossing the anterior lymphatic gland in
Cenolestes obscurus.

The Sterno-hyoids form a uniform strip from origin to insertion
in all Marsupials except a male pouch specimen of Macropus
giganteus (text-fig. 71c),in which they diverge at their hyoid
attachments, and the larynx protrudes between them. They are
easily separated, as a rule, from the subjacent sterno-thyroids,
but that cannot be done in Perameles obesula. This would show
that these muscles arise from a single sheet by splitting.

In my paper on the Koaia and Vulpine Phalanger (15)
I showed that the mylo-hyoid, digastric, and sterno-hyoid muscles
of the former form a thin sheet playing over, but in no way con-
nected to the hyoid bone; but I did not observe this condition in
any other Marsupial; it may also be an abnormality in the
animal examined by me. And in no other were there so many
small muscle bundles running between the digastrics and sur-
rounding muscles. The fusion of the digastric and mylo-hyoid
supports Gegenbaur’s view that these muscles were differentiated
by splitting.

The Diaastric has a small central tendon in Macropus, but
none is present in Dendrolagus, Perameles, and Metachirus. In
Petrogale xanthopus (18) there are small fibrous areas, but no
true central tendon. The mandibular insertion is usually about
half the length of the bone between the angle and the sym-
physis, and is usually muscular, but Osgood (9) gives a more
extensive attachment from angle to symphysis in Cwnolestes
obscurus ; and the anterior part is more aponeurotic.

The Hyoglossus was absent in my specimen of Phascolarctos
cinereus, and is not described in Macalister’s paper on the same
animal. It is present in all other Marsupials. Its absence in
the Koala is probably due to the long interval between the tongue
and the hyoid bone, which has few attached muscles.

Carlssen (2) described the myology of Dendrolagus dorianus in
detail, and grouped the muscles in three sets :—

1. Those resembling the muscles of the terrestrial Macro-
podide and differing from those of the Phalangeride.

2. Muscles resembling those of the Phalangeride and differing
from those of the terrestrial Macropodide.

3. Conditions which differ from those of the terrestrial Macro-
podide and Phalangeride, and are secondary adaptations for an
arboreal life.

894 DR. C. F. SONNTAG ON THE VISCERAL ANATOMY

My own observations on Dendrolagus ursinus confirm those of
Carlsson.

The stylo-hyoid ligaments are most prominent in Phascolarctos
cimereus.

I was unable to detect any condition foreshadowing the sterno-
maxillares of the Edentata in the insectivorous Peramelide.

The Mouth.

The lips are fleshy in all Marsupials, and the upper one is cleft
to a variable depth in the Macropodide, but entire in all Poly-
protodonts. An incipient cleft is present in Canolestes obscurus.
They are relatively thicker and always entire in the new-born
animal. They bear numerous hard, rounded, or pointed tubercles
in many species; in the Didelphyide they are absent in
Metachirus opossum, Philander laniger, Marmosa elegans, and
LIndelphys marsupialis, but they are situated close to the angles
of the mouth in Didelphys azare. In Dasyurus viverrinus they
are situated at the level of the canine teeth. They are close to
the angles of the mouth in Dendrolagus, Pseudochirus, and
Trichosurus. The largest tubercles observed by me were in
Dendrolagus ursinus. They are never so numerous as the
papille in the Felide. In some forms the lips have callous pads.

Both labial frenums are present in all genera except
Dendrolagus and Pseudochirus, which have only the upper one.
In the young animal it passes between the incisor teeth to be
attached to the anterior end of the incisive palatal pad, but it is
fixed to the gum in theadult. Ina young Epyprymnus rufescens
Tues continuous with the median erect running back on the
anterior end of the palate. ‘The lower lip is united to the sides
of the gums by comb-like crests in Dendrolagus and Pseudochirus,
arid these form shaggy pads in the vestibule (text-fig. 70 H, a).

Labial labrets are present in Phascolomys and Cenolestes.
Owen (10) pointed out that the labial glands are well developed
in Dasyurus, but I did not observe orifices in D. viverrinus.

The mueosa lining the vestibule is smooth in all Didelphyide,
Sarcophilus harrisi, Trichosurus vulpecula, and Petaurus sciureus.
It is covered with innumerable closely-set, rounded papille of
moderate size in Dasyurus viverrinus and Phascolarctos cinereus,
but these are not arranged in rows like the papille of the Ungu-
lata. In many Marsupials it is traversed by ridges of various
kinds, which, in some cases, if notin all, are modified and fused
papille. They are thick and entire in Perameles obesula and
Petaurus breviceps papuanus, and the spaces between them
probably serve as traps for insects in the former. Their free
borders are nodulated in the latter. They are thin and comb-
like in Macropus, “pyprymnus Pseudochirus, and Dendrolagus
(text-fig. 70 D, > & KH, 6); and their function may be to keep the
spaces between the teeth clear of food particles, for they fit into.
them.

AND MYOLOGY OF THE MARSUPIALIA. 855

Cheek-pouches have been described in several species.
They are large in Chironectes (10) and Cenolestes (9), inter-
mediate in size in Phascolarctos, and small in Peragale lagotis
(10); but I do not think that the small fosse of Perameles obesula
ean be dignified by the name of cheek-pouches. Forbes mentions
rudimentary ones in Phascolomys. No Marsupial has comb-like
structures on the tongue like the Ungulata.

Text-figure 70.

A. The platysma of Sarcophilus harrisi showing superficial (a) and deep (4) fibre
B-F. The palates of Sarcophilus harrisi (B), Onychogale lunata (C), Den-
drolagus wrsinus (D), and Macropus bennetti (F). E. Vestibule of Dendro-
lagus ursinus ; c. papille. Other letters in text.

The Hard Palate:—In many Marsupials the anterior end has
an eminence, or ‘incisive pad,” whose long axis usually coincides
with that of the palate; it is at right angles to it, however, in
Sarcophilus harrisi (text-fig. 70 B,p). It is smooth, papillose, or
tuberculated, and varies greatly in prominence. In Phascolarctos
cinereus and some of the Phalangeride it is replaced by a cluster
of incisive tubercles, and its place is taken by a short, incisive
ridge in “pyprymnus rufescens.

The complete palatal ridges are most numerous in the Perame-
Proc, Zoou, Soc,—192!, No, LVII, 57

856 DR. C. F. SONNTAG ON THE VISCERAL ANATOMY

lide and Oenolestes obscurus, fewest in Trichosurus vulpecula,
and largest in Phascolarctos cinereus, but the commonest number
is eight. They are most variable in the Polyprotodonts, and the
numbers of complete and incomplete ridges are of considerable
value for purposes of classification, as shown in the table of
formule given below.

In all Marsupials the ridges on the anterior part of the hard
palate are larger than those on the posterior part, and the pos-
terior margin of the palate frequently appears as a well-marked
ridge. The spaces between the ridges are occupied by papille,
tubercles, or incomplete ridges, and the latter are present in
Didelphys, Perameles, Dasyurus, Sarcophilus, and Cenolestes. The
incomplete ridges cross the mid line, and differ from those of
many of the higher Mammalia, in which they run from a median
raphé to the sides of the palate.

No Marsupial has a median raphé running the whole length
of the palate.

In a very young pouch specimen of Onychogale lunata the lips
are entire, the tongue is excavated to form a bed for the nipple,
and the incisive pad and anterior palatal ridges are outlined and
hardly elevated; but the posterior palatal ridges are well-marked.
The conditions are such that there is a firm area of contact
between the nipple and mouth. In an older pouch specimen of
Macropus bennetti the upper lip is cleft, the incisive pad is more
prominent, and the anterior palatal ridges are more developed
than those of Onychogale; and in pouch specimens of Macropus
giganteus and Phalanger orientalis of still more advanced ages
the conditions resemble those of the adult—namely, large incisive
pads, strong anterior ridges, and weak posterior ones.

Tn animals requiring a firm contact between the mouth and
nipples large anterior ridges would be disadvantageous, and they
are unnecessary while the diet consists entirely of milk.

The vestibule and cavum oris communicate posteriorly behind
the last molar teeth, and in several Marsupials the diastemata
form anterior communications.

The Soft Palate is frequently covered with papille, which are
small, but visible to the naked eye. It has no uvula, and thins
out posteriorly where it forms the anterior and lateral boundaries
of the posterior extremity of the nasal tube; the orifice of the
tube is round and small, or long and pyriform or slit-like. The
larynx may be gripped by the tube, or the opening of the tube
may overlap its superior aperture. Inthe mammary feetus the
larynx passes into the nasal tube so that the entrance of air is
not impeded in suckling *.

In all Marsupials the orifices of the Eustachian ~tubes are
within the nose. And these have been figured in my paper on

* In a manuscript note Garrod states that the soft palate shuts off the glottis
from the mouth in Tamandua tetradactyla, and suggests that this arrangement
comes into use when the animal is collecting masses of termites before swallowing.
No insects can possibly enter the larynx in consequence, ;

AND MYOLOGY OF THE MARSUPIALIA, 857

the Koala and Vulpine Phalanger (15). They are not surrounded
by a cushion.

The characters of the palate can be conveniently expressed by
means of formule. In the following list C denotes complete
ridges, | incomplete ridges, P incisive pad, T incisive tubercles,
Ra median incisive ridge or raphé, and U uvula. For the area
surrounded by the incisor and canine teeth the term “ incisive
area” is suggested.

POLYPROTODONTS. DIPROTODONTs,
Metachirus opossum C,Ip5P U- Phascolarctos cinereus C,I,TU—
Philander laniger ... C,I,P+U— Trichosurus vulpecula C,I6P—U—
Didelphys azare ... C,1,P+U-— Pseudochirus pere-

i marsupralis C,I,P + U— GUMS So eos C,1,.P —U—
Marmosa elegans ... CjIb6kP—U— Petaurus sciureus... C,1,TU —
Perameles obesula ... C,,1,P+U— » breviceps pa-

[DUWORRUS SoraaccesoocSes C,1,P + U—
Cenolestes obscurus ... C,1,P +U—
Apyprymnus rufescens C-I5RU —
Dendrolagus (all

SPECIES iin ec datic ais aye C,1oP + U—

Macropus (all species) C,I,P + U—
Onychogale lunata... CzI5P+U —
Phalanger orientalis... C,Ib5P-+U—

Dasyurus viverrinus C,1,P+U—
Sarcophilus harrist... C-I,P+U —

The only Marsupial in the above list in which the palate
narrows from before backwards is Lpyprymnus rufescens. In
the Peramelids the palate is long and narrow.

I have nothing to add to existing descriptions of the tonsils
and fauces. And in no Marsupial could I detect Waldeyer’s
lymphatic ring in the pharynx.

The Salivary Glands (text-figs. 71-74).

The Parotid Glands (m) are small in all Polyprodonts, but vary
considerably in thickness and superficial extent in the Dipro-
todonts. ‘hey are thick, and extend from the auricles to the
clavicles in Dendrolagus and Dorcopsis, and Garrod has figured
their surface markings in the latter (4). Their relations in the
former are shown in text-fig. 74; and the ducts, which are
capacious and thin-walled within the glands, become thicker and
contracted where they cross the masseter muscles. They are
equally thick, but only reach the middle of the neck in Phasco-
larctos cinereus. And in none of these genera do they pass
mesially to the inner borders of the sterno-mastoid muscles. In
other genera they are thin, and cover a variable extent of the
neck. They reach the clavicles in Petaurus, but do not extend
so far in Macropus, Hpyprymnus, and Onychogale.

The most extensive parotids are present in Pseudochirus pere-

grinus, in which they extend from the auricles to the clavicles,
57*

858 DR. C. F. SONNTAG ON THE VISCERAL ANATOMY

and pass inwards to cover the posterior fourth of the ventral
surface of the neck. The glands are composed of numerous small
portions surrounded by connective tissue, but it is not always
possible to discern the ductules. They were prevented from
reaching the mid line of the neck by the cervical thymic lobes in
my specimen of Vrichosurus vulpecula, but Symington (17) saw
them cover the thymus in his. Although their extent is great
they are thin,

Text-figure 71

The neck of Macropus giganteus. a. omo-hyoid; b. thyroid gland; ce. pretracheal
muscles; d. fascia; e. submaxillary gland; f. mandible; g. small gland;
h. digastric; 7. mylo-hyoid ; 7. larynx; &. cervical thymus; /. sterno-mastoid ;
m. parotid gland in outline.

The Submaaillary Gilands (e) are, generally speaking, large in
Polyprotodonts and small in Diprotodonts, but there are excep-
tions in the latter. They are large, single, and oval in the Didel-
phyide and Dasyuride, and in some of the former they reach from
the angles of the mandible to the clavicles. They are immense and
lobulated in Perameles obesula (text-fig. 72), in which a copious

AND MYOLOGY OF THE MARSUPIALIA. 859

viscid secretion is required to surround and entangle the insects
trapped in the cheek-pouches (?) and spaces between the palatal
ridges and oral vestibule; and large glands are also present in
the insectivorous Cwnolestes obscurus (9)*. In Sarcophilus
harrisi the total mass of salivary glands is small, but the sub-
maxillaries form the chief part,

In the Diprotodonts the submaxillary glands are single or lobu-
lated, and the largest ones are present in the various species of

Text-figure 72,

The neck of Perameles obesula. e’. concealed lobe of submaxillary gland ;

E.T.V. external jugular vein. Other letters as in text-fig. 71.

Dendrolagus. Small isolated glands are present between the
main masses in Macropus giganteus (text-fig. 71, g), but the small
glands are united to the main masses in Dendrolagus wrsinus
(text-fig. 74,9). And in these genera I traced small sympathetic

* Tn insectivorous Marsupials the glands are relatively smaller than those in the
insectivorous Edentata.

860 DR. C. F. SONNTAG ON THE VISCERAL ANATOMY

nerves into the small glands (text-fig. 78, sgn). The ducts do not
open on frenal lamell as in the Primates.

One must be careful not to confuse the sulbrtemallare) glands
with the cervical lobes of the thymus gland, and _ histological
examination must be carried out in doubtful cases.

The Sublingual Glands of many Marsupials have already been
described, and Oppel (8) has collected the various observations.

In Dendrolagus ursinus they are long, thin, and narrow, and
resemble those of D. dorianus already described by Carlsson (2).

“Text-figure 73.

DS
HH

The neck of Petawrus sciureus. Letters as in text-fig. 71.

Osgood (9) described large round glands in Cenolestes obscurus.
Those of Peramebes obesula are oval and do not extend so far
forwards. Posteriorly they are in contact with the submaxillaries.
The submaxillaries are large and the sublinguals small in
P. obesula, but the conditions are reversed relatively in
C. obscurus.

When the total bulk of the salivary glands of each Marsupial
are compared, it is seen that Dendrolagus has the largest mass.
And they conceal large areas of the muscles in the vicinity of the
hyoid bone.

AND MYOLOGY OF THE MARSUPIALIA. 861

I have nothing new to add to our knowledge of the arterial
supply and venous drainage of the salivary glands. And I could

not detect any connection between them and the anterior cervical
lymphatic glands.

Text-figure 74.

The neck of Dendrolagus wrsinus (A) and the cervical thymus B. 2, 0, cutaneous
branches of the cervical plexus; p. trapezius; g. common carotid artery.
Other letters as in text-fig. 71.

The Alimentary Canal.

The Esophagus :—The mucosa of the abdominal part exhibits
transverse ruge in Didelphys, but in no other genus. |

The Stomach :—The form depends on the degree of distension,
but the latter is not stated in all existing accounts. Owen
deseribed a globular stomach in several species, but I found that
the empty stomach is more or less triangular in a number of
Polyprotodonts. . Rane

In Perameles obesula (text-fig. 75 B) the empty stomach 1s tri-

862: DR. C. F. SONNTAG ON THE VISCERAL ANATOMY

angular with the apex formed by the cesophageal orifice. The
left rounded basal angle represents the fundus, and the duo-
denum joins it at the right one. The pylorus does not project
into the duodenum, and the sphincter is weak.. The fundus is
rugose. No gastric gland is present as in Cenolestes (9), but
there is a small lymphatic gland on the lesser curvature.

Text-figure 75.

The stomach in the Marsupialia. A. Dendrolagus wrsinus open to show—a. central
tract; b. sacculi; ¢. gland strips; d. esophagus; e. pyloric region. B. Pera-
meles obesula. C. Metachirus opossum. D. Petaurus breviceps. E. Petaurus
Sciureus.

The empty stomach of Metachirus oposswm (text-fig. 75 C)
differs from the above in the thickness of the pyloric region, the
existence of well-marked ruge on both walls, and the great
strength of the pyloric sphincter. There is a small lymphatic
gland on the lesser curvature as in Perameles obesula.

AND MYOLOGY OF THE MARSUPIALIA, 863

In Petaurus sciureus (text-fig. 75 E) the globular fundus is
directed forwards and to the left. ‘The duodenum, whose com-
mencement is notched, leaves its dorsal aspect. In P. breviceps
(text-fig. 75 D) the duodenum has no notches, but a groove
marks the pyloro-duodenal junction. ‘The greater curvature is
sacculated. ,

In Dendrolagus ursinus (text-fig. 75 A)* the greater part is
concealed by the liver, the only parts visible being the cardiac
cul-de-sac and greater curvature. The form and general cha-
racters resemble those of other species already described by
Beddard:(1), Owen (12), and Carlsson (2). These authors com-
pared the stomachs with those of other Macropodide, and
described the extent of the glandular and non-glandular parts of
the mucosa.

Owen and Beddard described groups of follicular glands lying
along the sides of the central tract (a4) in D. imustus and
D. bennetti. In D. ursinus they are replaced by long glandular
strips (c), into which branches of the abdominal parts of the
vagus nerves can be traced, and the vagus branches probably
contain sympathetic fibres. From the examination of the
stomachs of animals belonging to most of the mammalian orders,
I have come to the conclusion that the branches of the vagi are
greatly increased in numbers when there is a special gastric
glandular apparatus. These branches pass directly to the glands
as in D. ursinus, or run through a plexus with ganglia as in
Phascolarctos cinereus (15). The pylorus does not project into
the duodenum in Dendrolagus.

Osgood described a special gastric gland in Canolestes obscurus,
but did not mention its nerve supply.

The histology of the stomachs of the Marsupialia has already
received considerable attention, and Oppel (8) has collected and
analysed the various papers. I have nothing new to add to
these accounts. Nematode worms are present in many stomachs.

The Intestinal Tract :—The observations recorded here are
supplementary to those of Chalmers Mitchell (6, 7), Cunningham
(3), Owen (10), and Flower (20).

In Metachirus opossum the well-marked duodenal loop is wide,
thin-walled, and extends posteriorly for a considerable distance.
The flexure between its second and third parts is connected
by the usual peritoneal ligament to the abdominal parietes and
meso-colon, but is not continuous with the splenic ligament.
The cecum is two and a half inches long. It is capacious, and
its interior is devoid of folds and septa. Meckel’s Tract and the
large intestine differ in no respect from that of Didelphys
virginiana (6). The cecal mesentery has a concave free edge
and the angle between the eewcum and Meckel’s Tract is bisected
by an artery (text-fig. 76 A).

The cecum of Perameles obesula (text-fig. 76 B) is one and

* The specimen was specially hardened by injections of formalin to show the
topography of the stomach.

864 DR. C. F. SONNTAG ON THE VISCERAL ANATOMY

three-quarter inches long. It is longer and thinner than that of
Peragale lagotis figured by Chalmers Mitchell (6), and it has a
mesentery whereas that of P. lagotis has not. In other respects
the intestinal tracts of these Peramelide are similar. Their
ceca differ from the minute conical appendage of Canolesies
obscurus as figured by Osgood (9), and the latter is devoid of a
mesentery.

Text-figure 76.

A. Czxecum of Metachirus opossum; B. cecum of Perameles obesula; C-¥. aortic
arches with innominate (IA), right subclavian (a), right common carotid (6),
left common carotid (ce) and left subclavian (d) arteries; G—J. the spleens of
the Marsupials (names in text).

The ligament running backwards from the duodenum in
Perameles obesula crosses the front of the rectum and fuses with
the splenic ligament. They enclose a pocket containing the
rectum, and a probe passed into it runs backwards into the
pelvis.

The intestinal tract of my nearly adult female Dendrolagus
ursinus differs in several respects from that of a young specimen
described by Chalmers Mitchell (7). The duodenal loop is sharply
differentiated, but runs tmsensibly into Meckel’s Tract. ‘The coils
of Meckel’s Tract do not form a double spiral, and there is only

AND MYOLOGY OF THE MARSUPIALIA. 865

one cecum. The latter is fusiform, but is devoid of longitudinal
bands, sacculi, and a vermiform appendage, and the superior
mesenteric artery does not occupy the free edge of its mesentery.
At the point where the ileum enters the colon there is a trans-
verse vein. At a point two inches proximal to the ileo-cecal
junction there is a single Peyer’s Patch measuring one and a
quarter inches in length. .

No trace of a second cecum was found in my pouch specimens
of Macropus bennetti, Macropus giganteus, Onychogale lunata, and
Betiongia penreillata.

I have refrained from giving a detailed list of the measure-
ments of the different parts of the intestinal tract in my speci-
mens as they had been preserved for varying periods. The data,
if given, would be of little value in consequence.

The Pancreas.

In all Marsupials examined except Phascolarctos cinereus, I
found the pancreas much subdivided and dendritic, but the
arrangements of the subdivisions varied considerably. In Meta-
chirus opossum they are aggregated to form a compact mass in the
posterior part of the duodenal loop, and a long, narrow part
stretches across the abdomen to the spleen. Small branches are
given off from the body. The duct opens into the duodenum
along with the common bile-duct. Perameles obesula also pos-
sesses a long, narrow body with lateral processes, and the duodenal
loop contains an almost complete ring of pancreatic tissue such as
is found insome Rodents and Carnivora.

In Petaurus sciureus, P. breviceps, and P. breviceps papuanus the
pancreas consists of long strips and small isolated pieces arranged
along the course of the mesenteric vessels. It is difficult to detect
the ducts of the small lobules, and it is impossible to do so when
there is much fat in the omentnm and mesentery. The same
dendritic arrangement is seen in Dendrolagus ursinus, in which
the pancreatic and common bile-ducts open separately into the
duodenum. In JAlacropus, on the other hand, the ducts are
confluent (Owen).

Osgood describes a dendritic pancreas in Cenolestes obscurus (9),
so its condition is no more highly evolved than that of Perameles
obesula.

It is evident, therefore, that the pancreas of Phascolarctos
cinereus is of a higher type than the dendritic, more or less
diffuse, organ of other forms. ’

The Organs of Circulation.

The Pericardiwm adheres to the diaphragm in all Marsupials
except Trichosurus vulpecula, and it adheres to a variable extent
to the sternum and ribs. In Dendrolagus ursinus and pouch
specimens of Macropus giganteus and JL. bennetti the lungs were
seen to overlap the base of the heart, and the precordia, or
uncovered part, was adherent to the sternum and ribs.

866 DR. C. F. SONNTAG ON THE VISCERAL ANATOMY

The Heart:—In all Marsupials the apex is formed by the left
ventricle, and the right ventricle falls short of it by a variable
interval. Cunningham (3) stutes that the right ventricle in
Lhylacinus stops short of the apex by 13 inches, but I never
observed such a long interval in any Marsupial. In very young
pouch specimens she right ventricle is relatively longer than in
adults.

Owen (11) describes bifurcation of the right auricular appendix
as one of the chief characteristics of the Marsupial heart, but
some species have no trace of a division, and in others the division
has no reference to the ascending aorta.

The appendix is conical and bifid in Petaurus sciureus and
pouch specimens of Macropus bennetti and Macropus giganteus,
but the two divisions are small. In Dendrolagus ursinus and
Perameles obesula it is large and round; and the former has a
well-marked process lying on the dorsal aspect of the aorta, but
no trace is present in the latter. Cunningham pointed out that
there is no bifurcation in Thylacinus and Dasywrus, but there: is
a strong division in Cuscus and Trichosurus. A slight division is
present in Phascogale, but it has no reference to the ascending
aorta (3). In Phascolarctos cinereus the secondary process is large,
and the right precaval vein passes into it.

The ahietrantens of the interior of the auricles and ventricles
of many Polyprotodents and Diprotodonts have already been
described by Cunningham.

The Aortic Arch:—In all Marsupials it describes a full curve
and ends at the level of the fourth dorsal vertebra, and the
relations are similar in all forms to those described in the Koala
(15). The branches differ considerably, for six types are present,
as shown in text-fig. 76 C-F.

1. Both carotids and subclavians arise separately, so no inno-
minate artery is present— Phascolarctos cinereus T.

. The two subclavian and the innominate arteries are given
off, and the latter divides into the two common carotid arteries—
Dasyurus macrurus (C).

3. The branches arise as in Man--Phascolarctos cinereus*,
Petaurus breviceps*, and Phascolomys mitchelli (D).

4, The innominate artery gives off the left common carotid
artery, and divides into the right subclavian and right common
carotid arteries— T'richosurus vulpecula t.

5. The innominate artery gives off the right subclavian artery,

and is continued as the bicarotid trunk which divides into the
two common carotid arteries—Thylacinus harrisi, Didelphys can-
crivora, Dasyurus viverrinus, und Petaurus australis * (EK).
6, The innominate artery gives off the right subclavian and
both common carotid arteries at the same level-~Cuscus macu-
latus, Dendrolagus ursinus, Phascolarctos cinereus *, and Macropus
giganteus (EF),

* These observations are recorded in hitherto unpublished notes by Garrod.

+ These types have been figured in my paper on the Koala.

AND MYOLOGY OF THE MARSUPIALIA. 867

The left subclavian artery is the only constant branch, and the
left superior intercostal artery sometimes arises from the arch a
little distal to it.

I agree with Cunningham that there is no trace of the
obliterated ductus arteriosus. And I have nothing to add to
existing descriptions of the remainder of the arteries.

The Vena TZ'ransversa is usually horizontal and drains the
mouth, tongue, salivary glands, and some of the cervical muscles.
In a few cases it receives laryngeal veins. In some specimens of
Trichosurus vulpecula it is connected to the internal jugular veins
by vessels corresponding to the anterior jugular veins of the
‘Ocelot and Man. I have already figured these in my paper on
the Koala and Vulpine Phalanger (15).

The Haternal Jugular Veins in all Marsupials are similar to
those described in my paper on the Koala.

In all Marsupials examined by me there are two precaval veins,
but Garrod mentions in a hitherto unpublished note that the two
innominate veins unite about half an inch in front of the heart
to form a single precaval vein in Petaurus breviceps.

The Postcaval Vein covers the posterior half of the abdominal
aorta in all Marsupials examined by me except Phascolarctos
cinereus, and Hochstetter found it covering the aorta in all
except Petaurus taguanoides. Beddard found it covering the
aorta in all his specimens.

The Urinary Organs.

The kidneys are conglobate in all Marsupials, and their relative
positions vary as follows :—

1. Right kidney entirely anterior to the left one—Phasco-
larctos cinereus.

2. Posterior pole of right kidney levei with the hilum of
the left one—WMetachirus opossum, Marmosa elegans, Perameles
obesula, Petaurus sciwreus, and Trichosurus vulpecula.

3. Both kidneys situated at the same level— Macropus

giganteus, Macropus bennetti, Dendrolagus ursinus, and Cenolestes
obscurus.
_ On section only one papilla is present, and it may be sharp and
conical, or broad and flat. Small nodule-like elevations may be
present on broad, flat papille. No fat is present in the kidney
pelves in any of my specimens.

The course of the ureters is similar to that already described
in my paper on the Koala and Vulpine Phalanger (15).

Tha ventral surface of the bladder is connected to the ventral
abdominal wall by a ligament, which varies in prominence: in
different species. It is thin in the Polyprotodonts and thick
in Dendrolagus ursinus. In that species, as in D. bennetti (1), its
attachment to the abdominal wall is very extensive. In all
species it is completely anangious. It represents the remains of
the ventral mesentery. Lateral vesical ligaments are absent.

868 DR. C. F. SONNTAG ON THE VISCERAL ANATOMY

The dorsal surface of the bladder of Dendrolagus ursinus is
connected to the dorsal abdominal wall at the sides of the
vertebral column by two strong ligaments, which divide the pelvic
cavity into a median and two lateral compartments. The central
one contains the rectum and uterus, and the oviduets curl over
the free edges of the ligaments.

In Perameles obesula the uterus is separated from the rectum
by the wall of the peritoneal pocket formed by the fusion of the
splenic and duodenal ligaments. |

I have nothing to add to the existing descriptions of the
generative organs of either sex, nor to Osgood’s réswmé of our
knowledge of the marsupial bones (9). In the latter paper the
nipples are also enumerated, and it is shown that they are
asymmetrically arranged in the Didelphyide alone.

The Ductless Glands.

The suprarenal capsules are situated ‘on the mesial aspects of
the anterior poles of the kidneys in all Marsupials examined by
me except Phascolarctos cinereus, and that species has a very
prominent ligament uniting the left kidney and its capsule.

The Thymus Gland.

The Polyprotodonts differ from the Diprotodonts in the charac-
ters of the thymus gland. In the former it consists of two
elongated or oval bodies in the anterior part of the thorax, but
in the latter there are two cervical lobes in addition; and one
must be careful to distinguish them from salivary glands.
Osgood (9) found four thoracic glands in Cenolestes.

Topography :—The cervical lobes lie immediately under cover
of the platysma. They are oval or pyriform, with the narrow
ends turned forwards. They lie at the root of the neck, as in
Dendrolagus ursinus (text-fig. 74 B), JMJacropus giganteus
(text-fig. 71, &), and Macropus rufus, or they may extend far
forwards and even conceal one of the submaxillary glands, as in
Petaurus sciureus (text-fig. 73, k). They may touch the parotid
glands anteriorly, or be in contact with them throughout the
entire length of the neck as in Zrichosurus vulpecula. In Pseudo-
chirus peregrinus they are concealed by the parotid glands. ‘They
may be in contact with the mid line, or an interval in which the
pretracheal muscles are seen may separate them. The subjacent
structures are the sterno-mastoids, pretracheal muscles, and part
of the sternum and clavicles.

The surfaces may be finely or coarsely lobulated, and in no case
were they invested by a connective-tissue capsule derived from
the cervical fascia.

Symington (17) described and figured the topography of the
thoracic lobes, and described the gland in a number of Marsupials.
His observations on the Polyprotodonts are limited to Didelphys
virginiana and Dasyurus.cancrivora. He could not detect cervical

AND MYOLOGY OF THE MARSUPIALIA. 869

lobes in either of these species, and I was unable to detect them
in the Didelphyide, Dasyuride, and Peramelide. He believes
that the primitive condition is present in the Dasyuride, and the
cervical lobes of the Diprotodonts are a specialization in asso-
ciation with the vegetable diet and the call for a larger amount
of thymic tissue. If, as Swale Vincent suggests, the cervical and
thoracic lobes perform different functions, it is difficult to ascribe
any use other than dietetic to the former.

As age advances, the thymus-—both cervical and thymic—
diminishes in size and degenerates.

Phe Thyroid Gland.

In Perameles obesula the left lobe is long and narrow, ‘but the
right one is short and oval, and partly under cover of the pec-
toralis major. No isthmus is present; and this arrangement is
the reverse of that which I have already described in Phasco-
larctos cinereus (15). >

In all other Marsupials, I found the gland to be composed of
two small, oval lobes lying on the sides of the larynx alone
(Petaurus sciureus), or the posterior part of the larynx and a
‘variable number of tracheal rings; and the only animal possessing
an isthmus is 7richosurus vulpecula.

Each lobe receives a single artery from the common carotid,
and the veins join the internal jugular veins.

The lobes were concealed by the pretracheal muscles in all
animals examined except a male pouch specimen of Macropus
giganteus.

The Spleen (text-fig. 76 G—J).

The observations recorded here are supplementary to those of
Colin Mackenzie (5), Owen (11), and Retiterer (21).

In most genera there is little variation in form in the various
species, but the spleen has different shapes in the species of
Dendrolagus. In D. ursinus it is long and Y-shaped, with
rounded extremities (text-fig. 761). In D. inustus (12) it is
T-shaped, and it is spoon-shaped in D. bennetti (1). Carlsson (2)
states that it 1s pointed in D. dorianus.

In Petaurus sciureus (text-fig. 76 J), P. breviceps, and P. breviceps
papuanus the spleen is triangular, with sharp angles not prolonged
into processes, and the base of the triangle is applied to the
greater curvature of the stomach.

The spleen of Phalanger orientalis resembles those of the
Petaurists, the only difference being that its apical angle is 90°,
whereas those of the latter are larger.

Colin Mackenzie described and figured the spleen of Macropus
and my specimens differ in no way from his description; and a
pouch specimen of Onychogale lunata has a Y-shaped spleen
similar to that of the Kangaroos. The form does not alter from
birth to adult life,

870 DR. Cc. F. SONNTAG ON THE VISCERAL ANATOMY

In Perameles obesula (text-fig. 76G) the spleen has a massive
triangular body with thick sides, and the angles are prolonged by
cylindrical or flattened processes of splenic tissue. If the pro-
cesses are removed, the resulting organ resembles that of
Phascolomys described and figured by Colin Mackenzie. Garrod
states in a hitherto unpublished note that the spleen of the
Wombat forms an equilateral triangle. In a specimen of
Phascolomys mitchelli it had a lateral piece. In Perameles nasuta
it is thin and triangular, without prolongations.

The Bandicoots are the only Marsupials in which I observed
a continuity between the splenic and duodenal peritoneal liga-
ments. And the looseness of connection between the spleen and
stomach differs considerably ; these viscera were very closely
united to one another in Dendrolagus ursinus.

Osgood (9) showed that the spleen of Cenolestes obscurus has
an elongated body and a lateral piece, so it differs from that of
Perameles obesula.

In Metachirus opossum it is T-shaped, with one of the hori-
zontal limbs short (text-fig. 76 H1).

The external appearances of the spleens differ considerably
even in different species. Histological examination of the spleens
of many Marsupials shows that there is a very strong trabecular
network.

The Respiratory Organs.

The epiglottis is large, and its apex, which is entire or notched,
is frequently emarginate. It lies vertically or inclined forwards,
and it is sometimes gripped by the posterior extremity of the
nasal tube. And it is closely related to the base of the tongue in
all Marsupials except Phascolarctos cinereus. As the form of the
epiglottis and prominence of the aryteno-epiglottic folds vary,
the shape and size of the superior aperture of the larynx differ
considerably (text-fig. 77). .

In Didelphys azare the epiglottis is large, conical, connected
to the arytenoids by prominent ligaments, and strongly emar-
ginate, without any trace of a notch. The superior aperture of
the larynx is long and narrow, and cuts into the posterior surface
of the epiglottis. In D. marsupialis, on the other hand, the
entire epiglottis is only slightly emarginate, the arytenoid carti-
lages are plainly visible, and the superior laryngeal aperture is
wide and round. i{n Philander laniger and Marmosa elegans the
epiglottis is broad, and its apical margin is slightly concave but
not emarginate; and the superior aperture of the larynx is
triangular. Their condition is intermediate between those of
Didelphys and Metachirus opossum, in which the non-emarginate
apex of the epiglottis is notched. The epiglottis, therefore,
-is of considerable value for differentiating the genera of the
Didelphyide from one another.

The epiglottis is large, prominent, and entire in the Dasyuride.
In Dasyurus viverrinus it. is uniformly thick and the apex is

AND MYOLOGY OF THE MARSUPIALIA, 871

slightly emarginate. Its connections to the arytenoid cartilages
are also thick, and the superior aperture of the larynx is large
and open. The rima glottidis has very thick boundaries. In
Sarcophilus harrisi, on the other hand, the epiglottis is large
and thin, and its posterior parts are folded inwards and lean
against the anterior parts, so the superior aperture of the larynx
is a transverse slit. And the arytenoid cartilages and rima
glottidis are entirely concealed.

Text-figure 77.

Didelphys Didelphy. Philender Metachirus Marmosz

ezereae. marsupials. laniger: opossum. elegans:

» l
A
) Ml

beasyurus Sarcophilus _ Ferametes Dendrolagus Macropus
viverrinus. Aarrtst. o6esu/2. ‘ursinus. bennett.

| Ny ml ) M _ Pbalenger

Aepyprymnus Petaurus Petaurus Petaurus brevi- ag an talys
rufescens. scuireus, 6reviceps. ceps papuanus. ;

The larynx of the Marsupialia. The convex dotted lines crossing the epiglottis
are the levels of emargination.

The epiglottis of Perameles obesula has a broad, emarginated,
notched apex; and there is a small fossa with a crescentic
orifice between the epiglottis and the tongue. The characters of
the fossa in P. lagotis have already been described by Owen (10).

In the Macropodide there is considerable variation in the
characters of the epiglottis. In Dendrolagus ursinus it is entire
and the sides are folded inwards, so the superior laryngea

Proc. Zoou. Soc.—1921, No. LVITI. 58

872 DR. Cc. F. SONNTAG ON THE VISCERAL ANATOMY

aperture has the form of the figure 8, and the rima glottidis iS
invisible. In MZpyprymnus rufescens, Macropus bennetti, and
Macropus giganteus the apex of the prominent epiglottis has a
small sharply-cut notch resembling that in Trichosurus vulpecula,
and much emargination is present. The notches in the Polyproto-
dont, on the other hand, are wider and shallower.

Owen (10) states that the epiglottis of the Phalangers is broad
and bifid, but I found it entire and slightly concave in a mam-
mary foetus of Phalanger orientalis. It is entire in Pseudochirus
peregrinus, Petaurus sciureus, Petawrus breviceps, and Petawrus
breviceps var. papuanus. In Petawrus sciureus the degree of
emargination is considerable.

The large entire epiglottides in Phasolarctos cinereus (15) and
Conolestes obscurus (9) have already been described. é

The characters of the other cartilages, and the cords, ventricles,
and muscles are described by Owen.

The trachea is usually long and narrow, but it is short and wide
in Phascolarctes cinereus. The cartilages frequently form com-
plete rings, but in many species they are defective behind and a
membrane fills the gap. Some of the rings are expanded and
thickened on one side, and the numbers observed were :— Meta-
chirus opossum 19; Perameles obesula 34; Petaurus sciureus 35;
Dendrolagus ursinus 35. The numbers in other Marsupials have
been recorded by Cunningham (3), Osgood (9), and Owen (10).
In Marsupials in which the vagus and sympathetic are separate,
the tracheal and pulmonary plexuses are large.

The Lungs :—My observations differ from those of Owen (10)
in a number of cases. In the Didelphyide the right lung is
trilobate and has a well-marked azygos lobe. The left lung is
entire in Wetachirus opossum, but is bilobate in Marmosa elegans.
In Perameles obesula the right lung has three lobes and an azygos
appendage, and the left one is entire ; Owen also states that the
left lung of Perameles is undivided. The Phalangeride have
bilobate left lungs, and the right ones are trilobate with the
azygos lobe in addition. In the Macropodide, of which Dendro-
lagus ursinus, Macropus bennetti, and Macropus giganteus were
examined, both lungs have deep median sulci dividing them into
anterior and posterior parts, but these are not entirely separated
from one another, and the azygos lobe is large. ‘The conditions
differ from those of other Kangaroos described by Owen (10).
The mesial borders of the lungs are thin in the Macropodide.

The division of the lungs appears to differ considerably even in
the same species. Osgood (9) observed variations in Oanolestes
obscurus, and Owen, Forbes, and I have recorded three different
arrangements in Phascolarctos cinereus. I am unaware of any
account of the lobes of the lungs of the uterine feetus, and it is
important to ascertain whether there is fusion of lobes of the left

lung during development similar to that which takes place in
Bradypus tridactylus, in which the feetal lungs are divided into
lobes ; but those of the adult are entire.

AND MYOLOGY OF THE MARSUPIALIA. 873

In a Dendrolagus ursinus hardened by formalin injections it is

seen how the lungs conceal the large vessels and part of the right
auvicular_appendix.

Text-figure 78.

(
|

oy — ey —

a=)

The cervical parts of the vagi and sympathetics in Dendrolagus ursinus.
Letters in text.

The Vagus and Sympathetic Nerves*.

No account of the viscera of any animal is complete without a
description of these nerves, which control their activities.
The cervical parts are arranged in two ways in the Marsupialia.
* The lettering on text-figs. 78 and’79 corresponds, so far as possible, to that of
text-figs. 62 and 63 im my paper on the Koala (15).
58*

874 DR. C. F. SONNTAG ON THE VISCERAL ANATOMY

In the first type, which is present in both Polyprotodonts and
Diprotodonts, they are fused to form vago-sympathetic cords.
In the second form, which is found only in Diprotodonts, they
are connected by a variable number of communicating branches.
In no Marsupial did I find both types present in the neck as I
observed them in Tamandua tetradactyla and Lutra maculicollis.
These forms were present as follows in my specimens :—

Type 1:—WMetachirus opossum, Didelphys azare, Didelphys
marsupialis, Philander laniger, Dasyurus viverrinus, Sarcophilus
harrisit, Perameles obesula, Phascolarctos cinereus, Trichosurus
vulpecula, and Petawrus sciureus.

Type 2:—Dendrolagus ursinus, Macropus giganteus, Macropus
bennetti, Pseudochirus peregrinus, and Aipyprymnus rufescens.

The course of the vagi from the base of the skull to the point
where they reach the cesophagus in the posterior part of the
thorax has already been described in my paper on the Koala (15),
but the branches vary considerably.

The vagus (V) and hypoglossal (XII) nerves are fused in
Phascolarctos cinereus, but branches of communication run
between them in other Marsupials.

In Dendrolagus ursinus the pharyngeal (a) and superior laryn-
geal (6) nerves are given off from the sympathetic on the mght
side, but from the vagus on the left. The cord giving off the
right nerves supplies the submaxillary glands (s.g.n), and
possesses a ganglion (g'); it probably contains vagus fibres.

The vagus and sympathetic communicate. In the first type
branches run between the vagus and superior cervical ganglion
(s.c.g). In the second type there are several branches of com-
munication (c’) throughout the neck as well.

At the root of the neck the right vagus gives off its recurrent
branch (d), and the vago-sympathetic cord is resolved again into
its component vagus (v) and sympathetic (s) elements. I observed
no communications between the right and left recurrent laryn-
geal nerves, but the right nerve and sympathetic cord com-
municated.

Cardiac Nerves:—Iin type 1 the vagus transmits sympathetic
cardiac nerves, but in type 2 (text-fig. 79) the cardiac branches of
the vagus (f) and sympathetic are separate throughout. And the
cardixe branches of the sympathetic (c.b.s) come from the middle
ganglion or main cord, but never from the superior ganglion.
The cardiac and pulmonary plexuses have well-marked ganglia in
Dendrolagus ursinus, and that species has large tracheal and
cesophageal plexuses.

The Posterior Thoracic Parts of the Vagi:—In the Koala, as in
some reptiles, birds, and Man (Wirsung’s case), they are fused to
forma combined trunk; but the trunk of the Koala is not
resolved again into the two vagi. And many gastric nerves are
given off in the thorax.

In Metachirus opossum the left vagus runs along the ventral
surface of the cesophagus. It gives off a branch which accom-.

AND MYOLOGY OF THE MARSUPIALIA. 87)

panies the right vagus on the dorzal aspect on the cesophagus.
And no branches of communication run between the vagi*. In
Perameles obesula the cords have a similar disposition, but com-
municating branches run between them.

The conditions are more complex in Dendrolagus wrsinus
(text-fig. 79 A). The vagi are linked together by numerous
branches of communication (c¢.b.v). The left one has a large oval
expansion (EV) beyond which it bifureates, and the right half fuses
with the right vagus. The branches of supply to the cesophagus
are very numerous.

Text-figure 79.

v. [oA chs cbs. uv

G@SOPHAGUS

STOMACA.

The vagus and sympathetic in Dendrolaqus ursinus. A. In anterior two-thirds of
thorax. B. Posterior thoracic and abdominal parts. op. cesophageal plexus ;
tp. tracheal plexus; f. right cardiac branches of vagus; g. pulmonary nerves?
C.P.G. ganglia of cardiac and-pulmonary plexuses. Other letters in text.

In no case did I find any communication between the vagus
and phrenic nerves.

Abdominal Parts of the Vagi:—The vagi terminate in three
ways in the Marsupialia. The combined trunk ends in the solar
plexus, and gives branches to the stomach in the Koala as in some

* In Nandinia binotata, the most primitive Carnivore, the conditions resemble
those in Metachirus.

876 DR. C. F. SONNTAG ON THE VISCERAL ANATOMY

lower Vertebrata. In Metachirus opossum and Perameles obesula
the left vagus ends in the ventral wall of the stomach, and the
right one gives branches to the stomach and ends in the solar
plexus. In the third type, which is exemplified by Dendrolagus
ursinus (text-fig. 79 B), the main masses of the vagi end in the
stomach, and branches of communication run to the solar plexus.
Numerous branches (9.p.1) run to the special gastric gland.

I never observed any other mode of termination in the
Mammalia, but the branches of communication between the vagi
and the abdominal sympathetic plexus are more numerous in
some of the Kutheria.

The Cervical Sympathetic :—The arrangement in animals with
the first type is figured in my paper on the Koala (15), and that
in the second type is shown in text-fig. 78.

The superior cervical ganglion (S.C.G) is always present ; the
inferior one is frequently present, but the middle one (M.C.G) is
very variable. It is well-marked in Dendrolagus ursinus, but
absent in <d¥pyprymnus rufescens. In a pouch specimen of
Macropus giganteus it appears as a slight increase in the diameter
of the nerve. When both middle ganglia are present one may
be minute and give off no branches. ‘The branches of the cervical
sympathetic are :—

1. Communicating to the cervical plexuses, vagi, and right
recurrent laryngeal nerve.

2. Branches accompanying the common carotid (4.6), sub-
clavian, and vertebral arteries.

3. Cardiac nerves (c.b.s.) which arise from the middle or inferior
ganglia or main cords themselves.

I was unable to trace the filaments accompanying the vertebral
arteries to the brain. In some higher Mammals they can be
traced nearly to the interpeduncular space.

The sympathetic cords terminate in one of three ways. They
may end in the solar plexus and give off no branches to the aortic
plexus; and in this and the next form no splanchnic nerves are
present. In the second type the main mass of the sympathetic
ends in the solar plexus, but sends a small communication branch
to the aortic plexus. In the third type, which is the com-
monest, the cords run back to the sacral region, and splanchnic
nerves are present.

It has been shown by Volkmann, Bidder, and others that the
sympathetic becomes included more and more in the vagus as we
descend the vertebrate scale, and I believe that the form in
which they are fused in the neck is more primitive than that in
which they are separate. The nerves are fused in the neck in
Ornithorhynchus (Knox).

In Dendrolagus wrsinus the phrenic nerves bifurcate beyond
the roots of the lungs, and the halves pass to the ventral and
dorsal parts of the diaphragm. The nerve to the omo-hyoid
comes from the cervical plexus, and has no connection with the
hypoglossal nerve as in Petrogale xanthopus.

AND MYOLOGY OF THE MARSUPIALIA, 877

Systematic.

When the structures described above are arranged in sys-
tematic order, the following results are obtained :—

Sub-order POLYPROTODONTIA.

Characters common to all Genera:—Lips entire. Both frenums
present. Three cireumvallate papille, a well-developed sub-
lingua, and no lateral organs. Tonsils compact. Stomach simple
and has no special gland. Parotids small. Submaxillaries
large. Intestinal tract short. No cervical thymus. No thyroid
isthmus. Post-caval vein covers aorta, Vagus and sympathetic
fused.

Family Dipetpuyip2.

Characters common to all Genera :—Ovral vestibule smooth.
No cheek-pouches. Duodenal loop differentiated. Caecum well-
developed. Thyroid gland anterior. Mamme asymmetrical.
Patella not ossified.

Genus Didelphys :—LUabial tubercles variable. Palatal formula
C,I,.P+U—. Epiglottis conical, entire, emarginate. Lower
cesophagus transverely rugose. Mamme 5-13.

Genus Metachirus:—No labial tubercles. Palatal formula
CI,P—U—. Epiglottis notched. Lower esophagus not trans-
versely rugose. Mamme 7-9.

Genus Philander:—No labial tubercles. Palatal formula
C,1,P+U—. Epiglottis has concave anterior border and
rounded angles. Lower cesophagus not transversely rugose.
Mamme 7.

Genus Marmosa:—No labial tubercles. Palatal formula
CI,P—U—. Epiglottis with concave anterior border and sharp
angles, Lower cesophagus not transversely rugose. Mamme
9-15.

Family DasyuRriIpD&.

Characters common to all Genera :—No cheek-pouches. Duo-
denai loop not differentiated. Czcum absent. Thyroid gland
anterior. No transverse cesophageal ruge. Mamme sym-
metrical. Patella not ossified.

Genus Dasywrus:—Labial tubercles opposite canine teeth.
Oral vestibule papillose. Palatal formula C,I,P+U—. Hpi-
glottis thick, rounded, entire. Mamme 6-8.

Genus Sarcophilus:—No labial tubercles. Oral vestibule
smooth. Palatal formula C,I,P+U-—. Epiglottis thin and
parts crushed together. Mamme 4.

Family PERAMELIDA.

Genus Perameles :—No labial tubercles. Oral vestibule with
hard ridges. Cheek-pouches (?) small fosse. Palatal formula
C,,J,P+U—. Epiglottis notched. No transverse esophageal

878 DR. C. F. SONNTAG ON THE VISCERAL ANATOMY

ruge. Duodenal loop differentiated. Czecum well-developed.
Spleen massive and triangular. Thyroid gland posterior.
Mamme symmetrical. Patella ossified. Mamme 6-8.

Sub-order DIPROTODONTIA,

The Diprotodontia have few characters common to all their
genera in contradistinction to the Polyprotodontia. And these
characters are:—Parotid glands large. Tongue has reduction in
the sublingua and shows lateral organs. Thymus gland has
two cervical and two thoracic lobes. Meckel’s Tract long.
Cecum large.

Family Macropopip&.

Characters common to all Genera:—Lips cleft. No cheek-
pouches. Oral vestibule ridged. CHsophagus has no transverse
ruge. Stomach sacculated. Lungs have small degree of
division. Vagus and sympathetic nerves separate. Thyroid
gland anterior and has no isthmus. Mamme symmetrical.
Patella not ossified.

Genus Macropus:—Both labial frenums present. Palatal
formula C.I,P+U—. Epiglottis notched and wide. Spleen
with lateral process. Stomach has patches of follicular glands
and the area of digestive epithelium is small. CHisophagus enters
stomach near cardiac end. Mamme 4.
~ Genus Dendrolagus :—-Large labial tubercles. No lower labial
frenum. Palatal formula C,I,P+U—. Epiglottis entire and
folded. Stomach with large area of digestive epithelium and
follicular glands in patches or strips. Spleen Y-, T-, or
spoon-shaped. Cisophagus enters stomach near cardiac end.
Mamme 4,

Genus .Lpyprymnus :—No labial tubercles. Both labial
frenums present Palatal formula C_.I,RU—. Epiglottis wide
and notched. Cisophagus enters stomach near pyloric end.

Hamily PHALANGERID#.

Characters common to all Genera:—Lips entire. No cheek-
pouches. CHsophagus has no transverse ruge. Stomach simple
and has no special gland. Duodenum not a very specialized
loop. Meckel’s Tract short. Czcum well-marked. Mamme
symmetrical. Patella not ossified.

Genus Petaurus:—No labial tubercles. Both labial frenums
present. Oral vestibule smooth. Palatal formula varies. Epi-
glottis entire and rounded. Spleen an isosceles triangle. Vagus
and sympathetic fused. Mamme 4.

Genus Pseudochirus:—Lower labial frenum absent. Large
labial tubercles. Oral vestibule ridged. Palatal formula
C,I,P—U—. Hpiglottis entire. Vagus and sympathetic sep-
arate. Mamme 4.

tubercles.
Kpiglottis with shallow notch.
Vagus and sympathetic fused.
Genus Zrichoswrus:—Both labial frenums present.
Oral vestibule smooth,
Epiglottis with sharp notch.
Vagus and sympathetic fused.

triangle.
tubercles present.
1
C,
piece.

tubercles.

AND MYOLOGY OF THE MARSUPIALIA.

Genus Phalanger :—Both labial frenums present.
Oral vestibule smooth. Palatal formula C,I,P+U—.

me ye

879

No labial

Spleen a rectangular scalene

Mammee 4.

Labial
Palatal formula

Spleen with lateral

Family PHASCOLARCTID.

Genus Phascolarctos :—Both labial frenums present.
Oral vestibule papillose.

No labial
Palatal formula C,I,TU—.

Kpiglottis large, entire, emarginate. Spleen an isosceles triangle.

Stomach has a special gastric gland.

fused. Mamme 2 and are symmetrical.

Vagus and sympathetic
Patella absent.

Osgood (9) described the resemblances between Canclestes
obscurus and modern Peramelids, and the following table shows
the differences between it and Perameles obesula:—

Perameles obesula.

. Four large follicles in the sub-

mental triangle.

. Platysma and panniculus strong.
. Digastric muscular and less exten-

sive.

. Palatal formula Cy,1;P + U—.

. Absent.

. Epiglottis notched.

. No special gastric gland.

. No precardial lymphatic gland.
. Parotid glands small.

- Sublingual glands small.

. Cheek-ponehes minute.

. Two thoracic thymic lobes.

. Spleen triangular.
. Thyroid gland in posterior part of

neck,

. Well-marked czecal mesentery.
. Lower pole of right kidney level

with hilum of left one.

. Mamme 8.
. Large marsupium in adult.

. Arterial canal of seventh cervical

vertebra incomplete.

Coenolestes obscurus.

. Absent.

. Weak.

. Digastric has aponeurotic attach-
ment between symphysis menti and
mandibular angle.

. Formula CoIl;P + U—.

. Labial labrets present.

. Entire.

. Present.

. Large.

. Four thoracic thymic lobes.

. Spleen with lateral piece.

. In anterior part.

. Absent.

. Kidneys at same level.

. Mamme 4.

. Marsupium absent in adult* and
rudimentary in the young.

. Complete.

In my paper on the Koala and Vulpine Phalanger I tabulated
the differences between them, and stated that the Phalanger
has characters possessed in common by many Marsupials. The

880 DR. C. F, SONNTAG ON THE VISCERAL ANATOMY

following list enumerates the points in which all forms differ from
Phascolarctos cinereus :—

Hyoglossus muscle present.

Vena transverse horizontal.

Anterior palatal ridges convex forwards.

Tongue close to epiglottis.

Pylorus does not project into duodenum.

Tonsils in fauces.

Pharynx has no dorsal ridges.

Innominate artery present.

Postcava covers abdominal aorta.

Gall-biadder does not extend as far as the middle of the
duodenum.

11. Pancreas dendritic.

12. Patella present.

13. Right lung has an azygos lobe.

SOOIH OF wr

I have nothing new to add to the well-known external and
skeletal characters of the Marsupialia.

Summary and Conclusions.

1. The mylo-hyoid and sterno-hyoid muscles are attached to
the hyoid bone, and the hyoglossus muscles are present in all
Marsupials except Phascolarctos cinereus.

2. The sterno-facialis muscle may arise from the platysma by
splitting.

3. The characters of the mouth and epiglottis are very variable
in the Marsupialia, and I would urge that they be added to those
at present in use for purposes of classification,

4. The right auricular appendix is not always bifid, as stated
by Owen, and sometimes the division has no relation to the
ascending aorta. :

5. When a special gastric gland is present, the abdominal
branches of the vagi are very numerous.

6. The Polyprotodonts have small parotids and large sub-
maxillaries, but the reverse is the case in Diprotodonts. The.
largest glands are present in Dendrolagus.

7. Lhe thyroid isthmus and anterior jugular veins are only
found in T'richosurus vulpecuia.

8. The postcaval vein covers the aorta in all Marsupials except
Phascolarctos cinereus and Petaurus taguanoides.

9. The vagus and sympathetic nerves are fused in the neck in
all Polyprotodonts and some Diprotodonts, but they are separate
in a few Diprotodonts. From the examination of these nerves
in all classes of vertebrates I have come to the conclusion that
the former is the more primitive type.

AND MYOLOGY OF THE MARSUPIALIA, 881

10. The Polyprotodonts have more characters common to all

‘genera than the Diprotodonts. And they retain primitive cha-
racters in their tongues, intestinal tracts, thymus glands, and
nervous systems.

co 00

10.
ln
12.

13.
14.
15.

16.
17.

BIBLIOGRAPHY,

. Bepparp, F. E.—“The Visceral Anatomy and Brain of
Dendrolagus bennett.” Proc. Zool. Soc. Lond. 1895,
pp. 131-137. |

. Carusson, A.—‘* Uber Dendrolagus dorianus.” Zool. Jahrb.
Jena, Systematik, xxxvi. pp. 547-617. 1913-14.

. Cunninenam, D. J.— Challenger’ Reports. Zool., V., pt. xvi.

pp. 1-192. 1882.

. Garrop, A. H.—‘“On the Kangaroo called Malmaturus
luctwosus by d’Albertis, and its Affinities.” Proc. Zool.
Soc. Lond. 1875, pp. 48-59.

. Macxenziz, Corry.—Journal of Anatomy and Physiology,

1916, pp. 1-18.

MircHeti, P. Cuatmers.—‘On the Intestinal Tract of
Mammals.” Trans. Zool. Soc. Lond. xvii. pt. v. pp. 437—
He oelOOD:

. Mrrcuent, P. Coatmers.—‘‘ Further Observations on the

Intestinal Tract of Mammals.” Proc. Zool. Soc. Lond.
1916, pp. 190-199.

OppEeL, A.—Lehrbuch der verg. mikr. Anat. vols. i.—ili.

Oscoop, W.—‘‘ A Monographie Study of the American
Marsupial Conolestes.” Field Museum of Nat. Hist.,
Zool., Chicago, May 1921.

Owrn, R.—The Comparative Anatomy of Vertebrates,
vol. ill.

Owen, R.—Art. ‘‘ Marsupialia” in Todd’s ‘ Cyclopedia of
Anatomy and Physiology,’ vol. i111.

Owen, R.—‘‘ Notes on the Anatomy of the Tree-Kangaroo
(Dendrolagus inustus Gould).” Proc. Zool. Soc. Lond.
1852, pp. 103-108.

Parsons, F. G.-—“‘On the Anatomy of Petrogale vanthopus,
ete.” Proc. Zool. Soc. Lond. 1896, pp. 683-714.

Povutron, E. B.—“< On the Tongues of the Marsupialia.”
Proce. Zool. Soc. Lond. 1883, pp. 599-628.

Sonnrac, C. F.—“ The Comparative Anatomy of the Koala
and Vulpine Phalanger.” Proc. Zool. Soc. Lond. 1921,
pp. 547-577.

Swan, J.— The Comparative Anatomy of the Nervous
System of Vertebrates. 1864.

Symineron, J.—“The Thymus Gland in the Marsupialia.”
Jour. Anat. Phys. xxxii. pp. 278-291. 1898.

. Swate, Vincenr.—Internal Secretion and ‘the Ductless

Glands. London, 1912, pp. 353-366.

882 THE VISCERAL ANATOMY OF THE MARSUPIALIA.

19. Winpte, B., & Parsons, F.—‘‘ On the Anatomy of Macropus
rufus.” Jour, Anat. Phys. xxxu., N.S. xii. pp. 119-
134. 1898.

20. Frowrer, W. H.—‘“ On the Organs of Digestion in Mam-
malia.” Medical Times and Gazette, 1872, pp. 645-647.

21. Rerrerer.—C. R. Soc. Biol. vol. 78, p. 535. 1915.

The papers by Carlsson, Osgood, and myself contain extensive
bibliographies. Owen’s article on the Marsupialia in Todd’s
‘Cyclopedia of Anatomy and Physiology’ enumerates the older
papers. The external characters of the Diprotodonts have been
described by Pocock, P. Z.S. 1921, pp. 591-608.

THE SECRETARY ON ADDITIONS TO THE MENAGERIE, 883

EXHIBITIONS AND NOTICES.
October 18th, 1921.

Sir 8S. F. Harmer, K.B.E., Se.D., F.R.S., Vice-President,
‘in the Chair.

The Secrerary read the following Report on the Additions to
the Society’s Menagerie during the months of May, June, J uly,
August, and September, 1921 :—

May.

The registered additions to the Society’s Menagerie during the
month of May were 307 in number. Of these 60 were acquired
by presentation, 206 were deposited, 18 were purchased, 11 were
received in exchange, and 12 were born in the Menagerie.

The following may be specially mentioned :—

2 Cheetahs (Cyncelurus jubatus), 2 , from Hargeisa, Somaliland,
presented by Major E. H. Ward, R.M.A., and Lieut. A. M.
Hutchinson on May 30th.

2 Tayras (Galera barbara) and 1 Kinkajou (Potos caudivolvulus),
from Ambalema, Colombia, presented by Henry C. Vaughan on
May 17th.

4 Crab-eating Dogs (Canis thous) and 2 White Amphisbenas
(Amphisbeena alba), from Morro Velho, Brazil, presented by
George Chalmers, C.M.Z.S., on May 23rd.

2 Small Egyptian Jerboas (Jaculus jaculus) from Giza, 2 Large
Egyptian Jerboas (Jaculus orientalis), from Maryit, and 2 Four-
toed Jerboas (Scarturus tetradactylus), new to the Collection.
from Maryut, presented by the Giza Zoological Gardens on
May 21st.

JUNE,

The registered additions to the Society’s Menagerie during the
month of June were 399 in number. Of these 167 were acquired
by presentation, 125 were deposited, 11 were received in ex-
change, 52 were purchased, and 46 were born in the Menagerie.

The following may be specially mentioned :—

2 young Lionesses (Felis leo) and one Bull Buftalo (Bos caffer),
from Naorobi, presented by Maj.-Gen, Sir E. Northey, K.C.M.G.,
on June /th.

A collection of Mammals from Burma, including a pair of
Thamin (Cervus eldv), a pair of Hog-Deer (Amis porcinus), a pair
of Muntjac (Muntiacus muntjak), a Tigress (Felis tigris), and two
Burmese Pig-tailed Macaques (Macacus leoninus), received in
exchange fron the Rangoon Zoological Gardens on June 25th.

884 THE SECRETARY ON ADDITIONS TO THE MENAGERIE.

A female Black Leopard (felis pardus) and a Leopard-Cat
(Ff. bengalensis), from Thayetmyo, Burma, presented by Mr. J. A.
P. Stuart.

3 Persian Gazelles (Gazella subguttwrosa) and a Gmelin’s Sheep
(Ovis orientalis), from Persia, presented by Sir Percy Cox,
K.C.M.G., Capt. R. E. Cheesman, C.M.Z.S., and Major Daly on
June 6th.

3 American Bluebirds (Stalia sialis) and 2 Sacred Ibises (bis
cethiopica), bred in the Menagerie.

4 Kastern Paradise Whydah-birds (Steganura paradisea ver-
reauxt) and 4 Hast-African Colies (Colius affinis), both new to
the Collection, presented by Mr. W. G. Keeping.

1 Greenland Falcon (Hierofalco candicans), captured at Boncath,
Pembrokeshire, and presented by Col. Cecil Spence Colby, C.M.G.,
D.8.0.

A large collection of North-American reptiles and batrachians,
including 10 Horrid Rattlesnakes (Crotalus horridus), 2 Copper-
heads (Ancistrodon contortriv), and 3 Mocassins (Ancistrodon
piscivorus), presented by the New York Zoological Society.

A Striped Tree-Frog (Megalixalus fornasinit), new to the
Collection, from Zanzibar, presented by Capt. W. H. Ingrams,
E.Z.S.

JULY.

The registered additions to the Society’s Menagerie during the
month of July were 180 in number. Of these 112 were acquired
by presentation, 20 were deposited, 1 was received in exchange,
21 were purchased, and 26 were born in the Menagerie.

The following may be specially mentioned :—

A collection of Mammals and Birds including 1 Serval (Felis
serval), 2 Spotted Hyznas (Hyena crocuta), 2 Harnessed Ante-
lopes (L’ragelaphus scriptus); 3 Collared Guinea-fowls (Vumida
meleagris), 2 Marabou Storks (Leptoptilus crameniferus), 1 Ostrich
(Struthio camelus), 1 Black-necked Crowned Crane (Balearica
pavonina), from Gambia, presented by H.E. Capt. C. fel Armitage
on July 19th.

il Malayan Tapir (Zapirus indicus), born on July 4th.

2 Nylg ghaie (Boselaphus tragocamelus), born on July 5th.

2 Cape Hyrax (Hyrax capensis), born on July 31st.

1 Great Kangaroo (Macropus giganteus), 1 Red Kangaroo
(Macropus rufus), purchased on July 6th.

1 Pygmy Flying Phalanger (Acrobates pygmea), new to the
Collection, deposited on July 6th ; 1 Noisy Rat (Rattus vociferans),
new to the Collection, from Kuala Lumpur, presented by C. Boden
Kloss, Esq.

3 One-Streaked Hawks (Asturinula monogrammica) from the
Northern Province of Nigeria, presented by J. R. C. Stephens on
July 25th.

4 Ostriches (Struthio camelus) from an Ostrich Farm in Tunis,
purchased on July 27th.

.THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 885

AUGUST.

The registered additions to the Society’s Menagerie during the
month of August were 231 innumber. Of these 61 were acquired
by presentation, 42 were depesited, 5 were received in exchange,
111 were purchased, and 12 were born in the Menagerie.

The following may be specially mentioned :—

A collection, principally from New Guinea and Aru, obtained
by Mr. Frost, and comprising a few mammals and reptiles and
about 100 birds, including four species of Birds-of-Paradise, some
rare Doves and Parrots. The following species are new to our
Collection :—3 Orange-fronted Fruit-Pigeons (Péilopus awrantii-
frons), 4 Stephani’s Green-winged Pigeons (Chalcophaps stephani),
1 Lesser White Goshawk (Astur lewcosomus), 3 Red-bellied Rails
(Hulabeornis castaneiventris), 2 Brown-billed Brush-Turkeys
(Talegallus fuscirostris), 1 Amboina King-Parrakeet (Aprosmictus
amboimensis), 8 New Guinea Terrapins (Lmydura nove-guinee),
1 Striped Gecko (Gecko vittatus): purchased on August 27th.

A collection of Mammals and Birds from Para, including 2 Red
Brockets (Mazama rufa), 2 Crab-eating Raccoons (Procyon can-
ertwvorus), 2 Maguari Storks (Huxenura maguart), 2 Cocoi Herons
(Ardea cocoi), 2 Scarlet Ibises (Hudocimus ruber), and others:
purchased on August 9th.

1 Lechee Antelope (Cobus leche), and 1 Wart-Hog (Phacocherus
ethiopicus), from 8. Africa, deposited on August 31st.

1 Finch-billed Bulbul (Spizivus canifrons) and 1 White-winged
Mynah (Poliopsar nemoricola), new to the Collection, from
Yunnan, presented by J. D. La Touche, C.M.Z.8., on August
10th.

1 Nigerian Hare (Lepus camnopus), new to the Collection, from
Kano, N. Nigeria, deposited on August 11th.

SEPTEMBER.

The registered additions to the Society’s Menagerie during the
month of September were 131 in number. Of these 56 were
acquired by presentation, 20 were deposited, 14 were received in
exchange, 37 were purchased, and 4 were born in the Menagerie.

The following may be specially mentioned :—

1 Abyssinian Oribi (Ourebia montana), from Kenya Colony,
K. Africa, deposited on September 24th.

1 Broad-billed Roller (Hurystomus crassirostris), from the Aru
Islands, new to the Collection, deposited on September 13th.

Mr. M. A. C. Hinton exhibited, and made remarks upon, an

embryo African Elephant.

886 THE SECRETARY ON ADDITIONS TO THE MENAGERIE.

Mr. E. G. Boutencer, F.Z.8., exhibited, and made remarks
upon, photographs of a Common Grass Snake swallowing a

Viper.

Mr. F. Martin Duncan, F.Z.8., exhibited a series of cinema-
tograph films of Marine Animals, and asked the Society to accept
the series for its Zoological Film Library.

Mr, D. Seru-Suirn, F.Z.S., exhibited a series of photographs
of the Sacred Ibis nesting in the Society’s Gardens, and said that
the last occasion on which the birds had bred in the Gardens was
in 1877, as recorded by the late Dr. P. L. Sclater in ‘ The Ibis’
(1878, ser. 4, vol. 11. p. 449).

‘Prof, G. Extior Suiru, F.R.S., F.Z.8., gave an account of the
habits of Varsius, and illustrated his remarks by a series of
photographs of living specimens which had been taken by
Mr. W. E. Le Gros Clark, F.Z.S., in Borneo.

November 8th, 1921.
Dr. A. Surra Woopwarp, F.R.S., Vice-President,

in the Chair.

Mr. Evcar P. Cuancz, M.A., M.B.O.U., gave an account of
his investigation of the laying-habits of the Cuckoo (Cuculus
canorus) and the life of the young Cuckoo, and illustrated his
remarks with s striking series of cinematograph films and photo-
graphs,

November 22nd, 1921.

Prof. E. W. MacBring, F.R.S., Vice-President,
in the Chair.

The Sncrerary read the following Report on the Additions
to the Society’s Menagerie during the month of October, 1921 :—

The registered additions to the Society’s Menagerie during the
month of October were 147 in number, Of these 48 were acquired
by presentation, 36 were deposited, 10 were received in exchange,
34 were purchased, and 19 were born in the Menagerie.

A HUMAN SKULL AND OTHER REMAINS FROM N. RHODESIA, 887

The following may be specially mentioned :—

1 Bosman’s Potto (Perodicticus poito), from Ilorin, Northern
Provinces of Nigeria, presented by J. R. C. Stevens, Hsq., on
October 16th.

2 Maholi Galagos (Galago maholi), from Pretoria, received in
exchange.

2 Snowy Owls (Nyctea scandiaca), from Northern Baftin’s Land,
presented by Henry Toke Munn, F.R.G.S., on October 8th.

1 Salvin’s Razor-billed Curassow (Mitwa salvini), from Eeuador,
new to the Collection, deposited on October 11th.

2 Regal Horned Lizards (Phrynosoma regale), from Arizona,
presented by E. Newitt on October 17th.

Mr. A. 8S. Le Sovér, C.M.Z.8., exhibited, and made remarks
upon, a series of lantern-slides illustrating the early life-history
of Ornithorhynchus.

Dr. A. SmirnH Woopwarp, F.R.S., V.P.Z.8., exhibited, and
made remarks upon, a Human Skull and other remains from
Broken Hill, North Rhodesia, upon which he had founded the
species Homo rhodesiensis. In comparing the Rhodesian skull
with a Neanderthal skull from La Chapelle, Dr. Smith Woodward
stated that the former may prove to be the next grade after
Neanderthal in the ascending series.

Proc. Zoou. Soc.—1921, No. LIX. a9

No. 220.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*
October 18th, 1921.

Sir 8, F. Harmer, K.B.E., Sc.D., F.R.S., Vice-President,
in the Chair.

The Secretary read a Report upon the Additions to the
Society’s Menagerie during the months of June, July, August,
and September, 1921.

Mr. M. A. C. Hinton exhibited, and made remarks upon, an
embryo African Elephant.

Mr. EK. G. Boutenctr, F.Z.S., exhibited, and made remarks
upon, photographs of a Common Grass-Snake swallowing a
Viper.

Mr. F. Martin Duncan, F.Z.S., exhibited a series of cinemato-
graph films of Marine Animals, and asked the Society to accept
the series for its Zoological Film Library.

Mr. D. Seru-Suirs, F.Z.S., exhibited a series of photographs
of the Sacred Ibis nesting in the Society’s Gardens, and said that
the last occasion on which the birds had bred in the Gardens was
in 1877, as recorded by the late Dr. P. L. Sclater in ‘ The Ibis’
(1878, ser. 4, vol. i. p. 449).

* 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.

Prof. G. Exxior Surry, F.R.S., F.Z.8., gave an account of the
habits of Yarsius, and illustrated his remarks by a series of
photographs of living specimens which had been taken by
Mr. W. E. Le Gros Clark, F.Z.8., in Borneo.

In the absence of Mr. Sranuey Hirst, F.Z.S8., a réswmé of his
paper ‘‘On some new Parasitic Mites,” was given by Prof. 8.
Mautik, F.Z.8.

In the absence of the Author, Prof. J. Puavrarr McMurricn’s
paper ‘‘ Note on the Systematic Position and Distribution of
the Actinian, Sagartia lucie,” was taken as read.

The next Meeting of the Society for Scientific Business will be
held on Tuesday, November 8th, at 5.30 P.m., when the following
communications will be made :—

Epear P. Cuance, M. ».M -B. O.

The Laying- habits of the aes (Cuculus canorus) and the
Life of the young Cuckoo.
W. Rast Suerzirrs, M.A., D.Sc.

Evolution within .the Genus.—Part I. Dendronephthya
(Spongodes), with Descriptions of a Number of Species.
Part II. Description of Species (Alcyonaria) taken by the
‘Siboga’ Expedition.

Dr. Cuas. F. Sonntag, F.Z.8.

The Comparative Anatomy of the Tongues of the Mam-
malia.—V. Lemuroidea and Tarsioidea. VI. Summary and
Classification of the Tongues of the Primates.

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

The External Characters and Classification of the Mustelidz.

3
The following Papers have been meebived —
Cuas. F. Sonntac, M.D., F.Z.S.
Contributions to the Visceral Anatomy and Myology of the
Marsupialia.
C. W. Hosuey, C.M.G., C.M.Z.8.
The Fauna of Hast Africa and its Future.
H. Matsumoto.

Megalohyrax Andrews and Titanohyrax, g.n.—A Revision,
of the Genera of Hyracoids from the Fayim, Egypt.

The Publication Committee desire to remind Fellows
that it is only by courtesy that they are allowed to
bring guests to a Scientific Meeting. For the present
no Fellow can be permitted to bring more than one
guest to a Meeting.

Communications intended for the Scientific Meetings should
be addressed to

P. CHALMERS MITCHELL,
Secretary.
ZOOLOGICAL Society oF LonpoN,

ReGent’s Park, Lonpon, N.W. 8.
October 25th, 1921.

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No. 221.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

November 8th, 1921.

Dr. A. Sutra Woopwarp, F.R.S., Vice-President,
in the Chair.

Mr. R. I. Pocock, F.R.8., F.Z.8., gave a résumé of his paper on
“The External Characters and Classification of the Mustelide.”

In the absence of the Author, Dr. W. Rar SHeErrirr’s paper
‘¢ Evolution within the Genus.—Part I. Dendronephthya (Spon-
godes), with Descriptions of a Number of Species. Part II. Des-
cription of Species (Alcyonaria) taken by the ‘ Siboga’ Expedi-
tion,’ was taken as read.

Dr. Cuas. F. Sonnrac, F.Z.S., gave a réswmé of his paper
“The Comparative Anatomy of the Tongues of the Mam-
malia.—V. Lemuroidea and Tarsoidea. VI. Summary and
Classification of the Tongues of the Primates.”

Mr. Epear P. Cuance, M.A., M.B.O.U., gave an account of
his investigation of the Laying-habits of the Cuckoo (Cucwlus

* 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 subseribe to the Publications; but it may be obtained on the
day of publication at the price of Stapence, or. if desired. sent post-free for
the sum of Six Shillings per annum, payable in advance.

canorus) and the Life of the young Cuckoo, and illustrated his
remarks with a striking series of cinematograph films and photo-

graphs.

The next Meeting of the Society for Scientific Business will
be held on Tuesday, November 22nd, at 5.30 p.m., when the
following communications will be made :—

Dr. A. Suri WoobwarD, je. ZS., E.RSS.

Hehe Be a Receil Haugan Skull from Broken Hill,
North Rhodesia.

Cuas. F. Sonntag, M.D., F.Z.S8.

Contributions to the Viscer al Anatomy and Myology of the
Marsupialia.

C. W. Hosiery, O.M.G., C.M.ZS.

The fee Ef Bast inex and its Future.

H. Matsumoto.

Megalohyrax Andrews and Titanohyrax, g.n.—A Revision
of the Genera of Hyracoids from the Fayim, Egypt.

The following Paper has been received :—
J. SrepHENSON, D.Sce., F.Z.8.
Contributions to the eee Clselien ca, and Zoo-
geography of Indian Oligocheeta :—
ITV. On the diffuse Production of Sexual Cells in a Species
of Chetogaster (Fam. Naidide).

V. On Drawida japonica (Michlsn.), a Contribution to the
Anatomy of the Moniligastride.

VI. On the Relationships of the Genera of Moniligastride ;
with some Considerations on the Origin of Terrestrial Oligo-
cheeta.

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 or Lonpon,
Recent’s Park, Lonpon, N.W. 8.
November 15th, 1921.

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No. 222.

ABSTRACT OF THE PROCEEDINGS

LOOLOGICAL SOCIETY OF LONDON,*
November 22nd, 1921.

Prof. E. W. MacBripz, F.R.S., Vice-President,
in the Chair.

Mr. A. S. Le Soviir, C.M.Z.S., exhibited, and made remarks
upon, a series of lantern-slides illustrating the early life-history
of Ornithorhynchus.

Dr. A. Smirn Woopwarp, F.R.S., V.P.Z.S., exhibited, and
made remarks upon, a Human Skull and other remains from
Broken Hill, North Rhodesia, upon which he had founded the
species Homo rhodesiensis. In comparing the Rhodesian skull
with a Neanderthal skull from La Chapelle, Dr. Smith Woodward
stated that the former may prove to be the next grade after
Neanderthal in the ascending series.

With the consent of the Author, the communication of
Mr. C. W. Hosiey’s paper on “The Fauna of East Africa and
its Future,’ was postponed.

* 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. Jt 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 Siwpence, or, if desired, sent post-free for
the sum of Six Shillings per annum, payable in advance.

Dr. Cuas. F. Sonnrae’s paper “ Contributions to the Visceral

Anatomy and Myology of the Marsupialia,” and Prof. H. Mar-
suMoro’s paper on ‘ Megalohyraw Andrews and Titanohyraa,

n.—A Revision of the Genera of Hyracoids from the Faytim,

Egypt,” were taken as read.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, February 7th, 1922, at 5.30 p.m.

A notice stating the Agenda for that Meeting will be circulated

in J anuary.

The following Papers have been received :—

. STEPHENSON, D.Sc., F'.Z.8.

Contribution to the Morphology, Classification, and Zoo-
geography of Indian Oligocheeta :

IV. On the diffuse Production of Sexual Cells in a Species
of Chetogaster (Fam. Naidide).

V. On Drawida japonica (Michlsn.), a Contribution to the
Anatomy of the Moniligastride.

VI. On the Relationships of the Genera of Moniligastride ;
with some Considerations on the Origin of Terrestrial Oligo-
cheeta.

R. Broom, D.Sc., F.R.S., C.M.Z.8.

On the Temporal Arches of the Reptilia.

The Publication Committee desire to remind Fellows

that it is only by courtesy that they are allowed to
bring guests to a Scientific Meeting. For the present
no Fellow can be permitted to bring more than one
guest to a Meeting.

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 Sociury or Lonpon,
Recent’s Park, Lonpon, N.W. 8.
November 29th, 1921.

36.

37.

40,

ele

PAPERS.

Page

. On some new Genera and Species of Anomodont Reptiles from the Karroo Beds of

South Africa. By R. Broom, F.R.S., C.M.Z.S. (Text-figures 28-45.) .........+0 647

The Cichlid Fishes of Lake Nyassa. By C. Tats Rucay, M.A., F.R.S., F.Z.S., Keeper
of Zoology, British Museum (Natural History). (Plates I-VI.; Text-figures 1-80.) 675

Note on the Systematic Position and Distribution of the Actinian Sagartia lucie.
By J. Puayrairs McMorricn, C.M.Z.S. (Text-figures 1-4.) ..........-.22 2c 00s 729

. The Comparative Anatomy of the Tongues of the Mammalia.—V. Lemuroidea and

Tarsioidea. By Craruus F. Sonntac, M.D., F.Z.8., Anatomist to the Society.
(Gliese etme sh GG = OOF) isc herrea e atte dine is race apetthayh caste plviemitme Aaj c Gimaioueelencne ina

. The Comparative Anatomy of the Tongues of the Mammalia.—VI. Summary and

Classification of the Tongues of the Primates. By Cuarnus F. Sonnrac; M.D.,
ZL Sees An ALOIS EaLORUeR SOULE iy rates «02 she cpae) cesaciaceys eves ery suave ten eecbel wince) nataeerienees ee OW

On some new Parasitic Mites. By Stanney Hirst, F.Z.8. (Text-figures 16-43.) .... 769

On the External Characters and Classification of the Mustelide. By R. I. Pocock,
IRIE Shy BL Alse Casa snl Paolo) 66 Geo upiad aeiuo Dod con deOo cd uecueooUodolbe 803

2. Megalohyruz Andrews and Titanohyrax, g. n.—A Revision of the Genera of Hyra-

coids from the Fayiim, Egypt. By H. Marsumoro. (Text-figures 1-6.).......... 839

Contributions to the Visceral Anatomy and Myology of the Marsupialia. By
Cnarces F, Sonntac, M.D., F.Z.S., Anatomist to the Society. (Text-figures 70-79.) 851

PME ues lel shy Ole OOM EOUNONS eye oft) ages s Feys Glamis gs esravcln a o)< cee)srois th le vig aw ste eleva, VILL

Henclexcten allt iivertit coarse went eines tate co a eos naa Na checae ee araga eu ncts aia otiaic ral vase han .co.e se ceatenarn ates xii

LS OE Alas:

1921, Parr IV. (pp. 647-887).

Page
C.Tarm Regan: Pls. L-VI. Cichlid Fishes of Lake Nyassa ...... 675

NOTICE.

The ‘ Proceedings’ for the year are issued in fowr parts, paged consecutively,
so that the complete reference is now P. Z. 8.1921, p.... The Distribution
is usually as follows :—

Part I. issued in March,
2 rretalalie % June.
sell bd Bs af September,
aro MEV ieee December.

‘ Proceedings,’ 1921, Part IIT. (pp. 447-646), was published on September 9th,
1921. :

The Abstracts of the ‘Proceedings,’ Nos. 220-222, are
contained in this Part.

‘The dates of Publication of ‘Proceedings’ 1830-1858 will be found in the ~
« Proceedings’ for 1893, pave 436.

The dates of Publication of ‘ fTransactions’ 1833-1869 will be found in the
‘Proceedings’ for 1915, page 814.

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