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COUNCIL AND OFPFICHRS

OF THE

ZOOLOGICAL

SOCIETY

OF LONDON,

1921.

Patron.

His Masesry Tur Kina,

COUNCIL.

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

ALFRED H. Cocks, Hsq., M.A.

CuHaRLES DrumMonb, Esq.,

Treasurer.

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

THe Rieur Hon.'lue Viscount

Grey, KeG PC!

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

MAC ScD RRS ny aces

President.

Pror. James P. Hitn, D.Sc.,

F.R.S., Vice-President.

WitiiamM Huntsman, Hsq.

Masor THe Lorp Anasrarr

Roserr [nnes-KeEr,

Pror. Ernest W. MacBripe,

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

Cot. Stir Henry McManon,

G.C.M.G., K.C.1.E.

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

Vice-President.

P. Cuatmers MircHeny, Esq.,

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

E.R.S., Secretary.

THE Har or Onstow, O.B.E.

Masor ALBerr Pam, O.B.E.

Tue Lorp QUEENBOROUGH.

His Grace THe DvKeE or

Rurtann, K.G.

Masor Ricwarp 8. Taytor.

A. Trevor-Barryz, Esgq.,

M.A.

Anruony H. WINGFIELD, Esq.,

Vice-President.

A. Smita Woopwarp, Esq.,

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

sident.

PRINCIPAL OFFICERS.

P. Cuaumers Mircuet, C.B.M., M.A., D.Sc., LL.D., F.RB.S.,

Secretary.

R, I. Pococx, F.R.S., F.LS., Curator of Mammals and

Resident Superintendent of the Gardens.

D. Sera-Suiru, Curator of Birds and Inspector of Works.

Epwarb G. BouLencrr, Curator of Reptiles.

Miss L. KE. Cuzxsman, F.E.S., Curator of Insects.

Prof. R. 'T. Lurprr, D.Sc., M.D., Director of Prosectoriw.

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

Dr. N. 8. Lucas, M:B., Ch.B., Pathologist.

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

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

F.W. Bonn, Accountant.

W. H. Coun, Chief Clerk.

LIST OF CONTENTS.

1921, pp. 1--446.

EXHIBITIONS AND NOTICES.

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

Menagerie during the months of November and

December lO One eye. cccni: Las suas atten eakie ee gas onde wer eaters

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

upon, a Black Salamander (Salamandra atra) .........

Iieots do 1, Ili, MRIS, IO /Agse 1 eb-demberanveyay oie cx

remarks upon, a series of lantern-slides of the Foetus

of a Three-toed Sloth (Bradypus tridactylus) .........

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

Menagerie during the month of January, 1921 ......

Prof. G. Exuior Smiru, F.R.S., F.Z.S. Exhibition of, and

remarks upon, photographs of a living example of

er sonishy 4 dalatewll: pare car. kere essnaen. 5 seks aeee te waked.

Dr. R. W. Saurevpt, C.M.Z.S. The last of the Passenger-

Pigeons (Hetopistes migratorius). (Text-figure 1.) ...

Mr. R. i. Pocock, F.R.S., F.Z.S. Exhibition of, and

remarks upon, the skull of a Sumatran Tiger .........

——. Exhibition of, and remarks upon, a Cheetah skin

iromeLancamVvilka, LerritOry ..-....-ses-neseoeeeee2- scree ee

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

Menagerie during the month of February, 1921 ......

Dr. P. Coatmers Mrrcnrny, C.B.E., F.R.S. Exhibition

a sara

of, and remarks upon, lantern-slides of a Baby Chim-

panzee born in the New York Zoological Park ......

Page

183

184

Sir Srpney F. Harmer, K.B.E., F.R.S. Exhibition of, and

remarks upon, a photograph of Elephant Twins ......

Mr. K. G. Bounencer, F.Z.8. Exhibition of Reptiles and

Batrachianrs hee Mei cee me Mee cme Rein Mom epee RE ete dA

Miss L. EK. Corrsman, F.E.S, Exhibition of, and remarks

upon, a nest of Anapha venata (Lepidoptera) .........

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

Menagerie during the month of March, 1921 .........

Mr. R. H. Burne, M.A., F.Z.S8. Exhibition of, and remarks

upon, a series of mounted specimens of young Flat

Fish demonstrating various stages in the transference

of the left eye to the right side of the head ............

Dr. P. CHatmers Mircueny, C.B.E., F.R.S. On a letter

received from Mr. E. H. Bean, Director of the Wash-

ington Park, Milwaukee, Wis., U.S.A., describing

the successful rearing of a Polar Bear Cub ............

Mr. C, Davirs SueErporn, F.Z.8. Exhibition of a coin of

the Saka Dynasty, showing the so-called ‘“ maneless

Wi Ore eee ete seer ceric crete tee tants ei semtaranmtiae is Solaire Bie

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

upon, a series of Jantern-slides of a remarkable nest

offthe Waspi(Vespaigenmanied) si, esecere see rere ae oe

Mr. D. Sera-Smirn. Exhibition of, and remarks upon, a

series of skins of the Australian Budgerigar (Jelo-

psittacus undulatus), showing colour-varieties that

have; been produced imicaptimity: a eeeeeteerer ester scene

Major EK. E. Austex, D.S.0., F.Z.S.. On an apparent

change of habitat on the part of the Common Cricket

(Gryllus domestics) sane ecenne eter eee ecne eee cee ec:

Page

4-43

445

443

444

445

St)

whe

PAPERS.

. The Comparative Anatomy of the Tongues of the

Mammalia.—II. Family 1. Simiide. By CHaruEs

F. Sonnrac, M.D., Ch.B., F.Z.S. (Anatomist to the

Society) im (extateouness N92) eee cca stecceeissdecsserss on

. Note on the Capture (in London) of a rare Parasitic

Fly, Hammomyia (Hylephila) unilineata Zett. By

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

(MexcGSimMoTMMe we Ns gargtsg settee dc ang necks demtiaeeee saree

. The Bases of Classification of the Theriodontia. By

D. M. 8. Watson, F.Z.S. (Text-figures 1-29.) ......

Experiments on Colour-Changes of the Spotted Sala-

mander (Salamandra maculosa), conducted in the

Society's Gardens. By E. G. Boutuncer, F.Z.S.

(Curator of Reptiles). (Plates I. & II.; Text-

DIC ATURE RTE) uch ea Be abi cha tad aba Meio rk ane A AN eae oe

. Contributions to the Morphology, Classification, and Zoo-

geography of Indian Oligocheta. By J. SrepHEnson,

D.Sce., F.Z.S. (Lecturer in Zoology in the University

omebidimbbureh) ay ((ext=tioures lia seeesee cake -.ees saacioe

. On the Structure of the Reptilian Tarsus. By R. Broom,

D.Se., F.R.S., C.M.Z.S. (Text-figures 1-27.) .........

. A Contribution to the Anatomy of the Three-toed

Sloth (Bradypus tridactylus). By CHartus F. Sonnvac,

M.D., Ch.B., F.Z.S8. (Anatomist to the Society).

(Plates I.-IV.; Text-figures 10-15.) ..............2005

. Report on the Deaths which occurred in the Zoological

Gardens during 1920. By N.S. Lucas, M.B., F.Z.S.

GEacholooistite) tine Socleby)) eie.ce: sencecms-cs5c0scsn0ee<=

On the Reproductive Organs of the Ascidian Aiiken-

thalia borealis Gottschaldt. By Dr. AueusTa ARNBACK-

CHrisiie-LinpE. (Text-figures 1-8.) .........0..ecceee

(St)

179

10.

13.

14.

16.

10S);

On the Variation of the Seapula in the Batrachian

Groups Aglossa and Arcifera. By Joan B. Procter,

le Ane iuarel (liven aires tfetinmenye)l-=11(0),)) V4 naaanoeescahondabriaesoon oes

Notes on Marine Wood-boring Animals. —LI. Crustacea.

By Wi A © AL MEAN IDES Conte Zia neces eterinienys ee feet pees

- Colour-production in relation to the Coloured Feathers

of Birds. By A. Mattock, F.R.S., F.Z.S8. (Text-

fiioumies LAAs, 55ers eee meme 9 lo leet

Fossil Bird-remains collected by Dr. Forsyth Major in

Sardinia, Corsica, and Greece. By EH. T. Nrewron,

Le Bi ig ie Ria Noon Gn Aion Sad ob cae AARC ERROR ABABA OD Hic

The Oriental Species of the Genus Callistomimus

(Coleoptera Carabide). By H. i. ANDREWEs.

(SAC rer Ey Neperte c STOEL Ce Cues nalsd an academe on Beare meee:

. On the Molluscan Genus Cochlitoma and its Anatomy,

with Remarks upon the Variation of two closely-allied

Forms. By G. C. Rosson, B.A. (Text-figures 1—9.)

A Revision of the Melolonthine Beetles of the Genus

Lictinohoplia. By Gade Arrow, H-Z.9.) Hanis:

(dca Eth ca Bene et ES Set Mood hae on Suda pHs aa nMmr aN Ne

. The Comparative Anatomy of the Tongues of the

Mammalia.—III. Family 2. Cercopithecide : with

notes on the comparative physiology of the tongues

and stomachs of the Langurs. By Cuarues F.

Sonntac, M.D., F.Z.S. (Anatomist to the Society).

(Text-figures NOSSO) ie arte ere sseare es kites ee ene

. Descriptions of New Moths from South-Kast Brazil.

By E. Duxiyristp Jones, F.E.S., F.Z.8. (Plates

AR Ie) Rane ar RE Li cn Aah dene BAUS Sor G cbt

On some new or little-known Acari, mostly Parasitic in

Habit. By Sranuey Hirst, F.Z.8. (Text-figures

Lie SR ON II RT RUS eee sl oie

. Observations on the Habits of Cochlitoma zebra, var.

fulgurata (Pfeitter), and Cochlitoma zebra, vay. obesa

(Pfeiffer), in Confinement. By Jane Lonesrarr,

Ath Depo Wh © they eon (led tain sis JESSIE) sh Soeesccodaasaeemeodabe

215

379

vil

Page

21. The External Characters and Classification of the

Procyonide (Raccoons,. etc.). By R. I. Pocock,

ERS eee ees a (Mext-tomres Ue Woe )i cere: entineens so 389

22. New or little-known Reptiles and Batrachians from

Southern Annam (Indo-China). By Maucoim A.

Smita, M.R.C.S., L.R.C.P., F.Z.8. (Plates I. & I. ;

Me xt ati Ue Se Ma aM ects. eaciqnecicta sire weidec vrs veeind else A423

Alphabetical Mistroi@ontribwtorse 5. 282.Ge. acca. c.acsnecescess vill

AindexqotlilllistraelOmsi, wencrfe a seieisie seciaect asec cls selac/eos Xl

TRIGGER eT RR SP Xx

ALPHABELVCAL List

OF TIE

COND RB O R's:

With References to the several Articles contributed by each.

(1921, pp. 1-446.)

AnprReEwes, H. EH.

The Oriental Species of the Genus Callistomimus

(Coleoptera, Carabide). (Plate I.) .......................00..

Arrow, G. J., F.Z.S., F.E.S.

A Revision of the Melolonthine Beetles of the Genus

IR gHoolnooliais (OBEN) Is) kackeshon0 sooo ecdnssondseagcoduseas 605

Austen, Major H. H., D.8.0., F.Z.8.

On an apparent change of habitat on the part of the

Common Cricket (Gryllus domesticus) .............0.00.0ce00

Bouuencer, EH. G., F.Z.8. (Curator of Reptiles).

Experiments on Colour-Changes of the Spotted Sala-

mander (Salamandra maculosa) conducted in the Society's

Gardens. (Plates I. & 1J.; Text-figure1.) ...............

Exhibition of, and remarks upon, a Black Salamander

(Salameandre Gtr) ..occecceceenerecneeeenerecnee cee teneeteeteneees

Exhibition of Reptiles and Batrachians ................5-

Page

267

446

Broom, R., D.Sc., F.R.8., C.M.Z.S.

On the Structure of the Reptilian Tarsus. (Text-

PUMICE Se IA he Mcrae ORRER HEA ok sehcaah Maen ASEM and ome uc ses

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

Exhibition of, and remarks upon, a series of mounted

specimens of young Flat Fish, demonstrating various

stages in the transference of the left eye to the right side

Ob ble heads. airy ears eeey ener esc San ener n sa Mn RSS dco ass

Caiman, W. T., D.Sc., F.Z.8.

Notes on Marine Wood-boring Animals.—II. Crus-

LUBY GLSP Nae Bes Baia Oo rR ORE IOUS Bit HN TILE ae Rs a a ol

CurrEsMAN, Miss L. E., F.E.S.

Exhibition of, and remarks upon, a nest of Anapha

WETMEGE UBPUG OP CELA) es toce ceysasctsee eas aetsaeiels aiaises are se Side

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

Note on the Capture (in London) of a rare Parasitic

Fly, Hammomyia (Hylephila) unilineata Zett. (Text-

HisMmene) eee decs Savas sat Seacloshisesia eas uaniee see ee naeietisivets vie sels se

Duncan, F. Martin, F.Z.8, (Librarian to the Society).

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

shdes of a remarkable nest of the Wasp (Vespa ger-

TUCILECE emer Bee IME Rene cte cian teat ce sicke Neate ste pecislascicin ts ¢ odie ss Baines

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

Exhibition of, and remarks upon, a photograph of

In) @olngiais TAINS oo Sok onendubbabdonn sd, JanbbddndesnosonecpBebaTe

Hint, Prof, J. P., F.R,S., F.Z.8.

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

slides of the Fetus of a Three-toed Sloth (Bradypus

PROGLCTAUILS) ee asta Vek elo. s elahesiose eiuish's e's sem erieia'sGucsesvo~ esac seers

Proc. Zoou. Soc.—1921. b

Page

143

444

215

443

445

443

Hirst, Sranuey, F.Z.S.

On some new or little-known Acari, mostly Parasitic

in Jabit.. \Clext-fisures 115.) ieee, serene ceabasetaeeee

Jones, E. DUKINFIELD, F.E.S., F.Z.S.

Descriptions of New Moths from South-East Brazil.

(Plates Peavy as ius, scion epee ae cine. een ceeneeess

Linpr, Dr. Augusta ArnBACK-CHRISTIE-.

On the Reproductive Organs of the Ascidian Awken-

thalia borealis Gottschaldt. (Text-figures 1-8.) .........

LonestaFr, JANE, F.L.S., F.G.S.

Observations on the Habits of Cochlitoma zebra, var.

fulgurata (Pfeiffer), and Cochlitoma zebra, var. obesa

(Pfeiffer), in Confinement. (Plates I.—III.) ...............

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

Report on the Deaths which occurred in the Zoological

(Gree rake) Chbbeniver EPA DE aera gta oe hooncechnesescheannobsénosassanocn

Mattock, A., F.R.S., F.Z.8.

Colour-production in relation to the Coloured Feathers

of Birds, (s(Mext-fisures/ 1 =42)iepes scene pees crereraeerenen:

MircHevit, Dr. P. CHAumers, C.B.E., F.R.8. (Secretary to

the Society).

Report on the Additions to the Society’s Menagerie

during the months of November and December, 1920 ..

Report on the Additions to the Society’s Menagerie

during the month of January, 1921 ................2+--2+0

Report on the Additions to the Society’s Menagerie

during the month of February, 1921 .....................--.

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

Baby Chimpanzee born in the New York Zoological

1 ES He) gape ANTM alah Wa eR nN om aA heel at Ane nary

357

187

379

183

184

Mivrewety, Dr. P. C. (cont.).

Report on the Additions to the Society’s Menagerie

duninesthe miomthvot Marchi, VOQ | igsscuecsrcsssccecsste eyes

On a letter received from Mr. EK. H. Bean, Director of

the Washington Park, Milwaukee, Wis., U.S.A., describing

the successful rearing of a Polar Bear Cub ...............

Newron, E. T., F.R.S., F.Z.8.

Fossil Bird-remains collected by Dr. Forsyth Major in

Mavainre, COrsicamaMOnGVSGCO siyvcanineeiiey, se css sense sisin cis ys

Procrur, Miss Joan B., F.Z.5.

On the Variation of the Scapula in the Batrachian

Groups Aglossa and Arcifera. (Text-figwres 1-10.) ......

Pocock, R. I., F.R.S., F.Z.5. (Curator of Mammals).

The External Characters and Classification of the

Procyonide (Raccoons, ete.). (Text-figures 1-13.) ...... :

Exhibition of, and remarks upon, the skull of a

POLAT PUTO OTE Siatarn nia, sinnaiivicsteleatatealettnateutaetalatirivle(e nila els aa fesse

Exhibition of, and remarks upon, a Cheetah skin from

Mec Ay Uke MOC TNGOUV 9.) eine -siceenundeesise nts PO CREEL

Rosson, G. C., B.A.

On the Mollusean Genus Cochlitoma and its Anatomy,

with Remarks upon the Variation of two closely-allied

Nbsee : Wo wife raya fi iC

Horm (voxtetie ues On) i iiakartgmbeisiiieuvessce sss tauseles

SumrBory, C. Davius, I'.Z.8,

Exhibition of a coin of the Saka Dynasty, showing the

so-called *‘maneless lion” ...........-. SARS O ACO BE On OT aE CER TAG

Suureupr, Dr. R. W., C.M.Z.S.

The last of the Passenger-Pigeons (/clopistes migra-

TORU (MMe RUC ER UMONL:)) wndsisbnraracsinrevenersaerva, ap ruasnn

Page

443

Add

AY)

197,

249

AAD

Smita, D. Seru-, F.Z.8. (Curator of Birds).

Exhibition of, and remarks upon, a series of Skins

of the Australian Budgerigar (Ielopsittacus undulatus),

showing colour-varieties that have been produced in

GEV OMIT cotta SoBde.c coon bbonodaduadsucnecanehabedncs das nop dns

Smitu, Prof. G. E.uiot, F.R.S., F.Z.8.

Exhibition of, and remarks upon, ‘photographs of a

livinssexamplerot Vansvus. «@llate de) «:i.jcceee tanner

Smita, Matcomm A., M.R.C.S., L.R.C.P., F.Z.S.

New or little-known Reptiles and Batrachians from

Southern Annam (Indo-China). ‘(Plates I. & II.;

MMe xi HIG EOS Flim Wie 2g) Wp veces: br miclstster-Tociats o eisicles eio:ciionel de de stan

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

the Society).

The Comparative Anatomy of the Tongues of Mam-

malia.—II. Family 1. Simiide. (Text-figures 1-9.) ...

A Contribution to the Anatomy of the Three-toed

Sloth (Bradypus tridactylus). (Plates I.-IV.; Text-

ritetbtsss) LUO IS)" © Se seeseoa serene do9)3s0n9s3eaa0H5s e008 ncoDe sansa:

The Comparative Anatomy of the Tongues of Mam-

malia.—II. Family 2. Cercopithecide: with notes on

the comparative physiology of the tongues and stomachs

of the Iuangurs. (ext-figures 16-36.) ...................0.

SrePpHEnson, J., D.Sc., F.Z.8.

Contributions to the Morphology, Classification, and

Zoogeography of Indian Oligocheta. (Text-figure 1.) ...

Watson, D., M.S., F.Z.S.

The Bases of Classification of the Theriodontia. (Text-

rigeamnasy (= 9.8)))) pn. Abb gonousunensods oapbeBensordensnconceancapresca:

Page

445

184

277

103

INDEX OF ILLUSTRATIONS,

Acanthodica frigida, Pl. I., p. 323.

Adlurosaurid, Fig. 25, p. 87.

Ailuropoda melanoleuca, Fig. 8, p. 405.

Ailurus fulgens, Figs. 2, 3, 7, 10, 12,

13, pp. 395, 403, 408, 412, 414.

Alaplena castraria, Pl. III., p. 323.

Anisodes antennaria, Pl. IIT., p. 323.

bizaria, Pl. IIL., p. 323.

— carolina, Pl. III., p. 328.

japaria, Pl. III., p. 323.

paranaria, Pl. II11., p. 328.

vigoraria, Pl. III., p. 323.

Anthropopithecus troglodytes, Figs. 1, 2,

pp. 3, 6.

Anteosaurus magnificus, Fig. 28, p. 92.

Apicia geminimacula, Pl. IL., p. 823.

strigularia, Pl. IL., p. 323.

Aplogompha castraria, Pl. I., p. 323.

fumaria, Pl. I., p. 323.

—— setinaria, Pl. I., p. 323.

yaponaria, Pl. I., p. 328.

Arctognathus, Fig. 20, p. 64.

curvimola, Figs. 18, 19, pp. 61,

62.

Arctops willistoni, Figs. 1-3, pp. 37-

39.

Artace regalis, Pl. I., p. 323.

zelina cetana, P\. I1., p. 323.

hanebaria, Pl. I1., p. 825.

Bagodares castra, P|. I., p. d20.

Bassaricyon alleni, Fig. 1, p. 893.

sp., Figs. 3, 5, pp. 397, 401.

Proc. Zoou. Soc.—1921.

Bassariscus astutus, Figs. 1, 3, 6, 13,

pp. 398, 597, 402, 414.

Boarnia nigraria, Pl. IL, p. 328.

Bombinator maximus, Fig. 6, p. 202.

Bradypus tridactylus, Pls. 1.—I[V.,

p. 157; Figs. 10-15, pp. 160, 164,

169, 171, 172, 174.

Broomia, Figs. 17, 18, p. 151.

Broomia perpleaa, Fig. 16, p. 151.

Bryocodia paulina, Pl. I., p. 823.

Bufo hematiticus, Fig. 1, p. 197.

Callistomimus belli, Pl. I., p. 233.

cauliops, Pl. I., p. 233.

chalcocephalus, Pl. I., p. 233.

— coarctatus, Pl. L., p. 233.

@abrewi, Pl. I., p. 288.

—— dux, Pl. L., p. 233.

—— jucundus, Pl. I., p. 283.

littoralis, Pl. I., p. 283.

—— rubeilus, Pl. I., p. 233.

—— sikkimensis, Pl. I., p. 233.

venustus, Pl. I., p. 233.

—— virescens, P1.I., p. 283.

Callurapteryx paularia, Pl. I1., p. 323.

Campatonema marginata, P1.11., p. 323

Carama incolorata, Pl. I., p. 323.

Casea broilii, Fig. 9, p. 148.

Cercocebus ethiopicus, Vig. 32, p. 310.

lunulatus, Fig. 33, p. 311.

Cercopithecus ethiops, Fig. 22, p. 293.

—— albigularis, Fig. 20, p. 288.

| —— burnetti, Fig. 18, p. 286.

X1V

Cercopithecus lalandit, Fig. 20, p. 288.

—— mona, Figs. 19, 20, pp. 287, 288,

—-— nisnas, Fig. 20, p. 288.

patas, Figs. 20, 21, pp. 288, 290.

—— preusst, Fig. 17, p. 284.

sabeus, Fig. 23, p. 294.

schmidti, Fig. 20, p. 288.

tantalus, Fig. 24, p. 297.

Cochlitoma zebra var.

Sulgurata, Figs.

6-9, pp. 259, 260, 252; Pls. I., IIL.,

p. 379.

var. obesa, Figs. 1-5,

251=253, 255, 257 Pls. I -10L.,

p. 379.

Craptoptera fenestraria, Pl. 11., p. 323.

Deiopeus, Fig. 22, p. 72.

—— leptocephalus, Fig. 21, p. 71.

Demodex qgliricolens, Fig. 12, p. 369.

talpe, Fig. 13, p. 370.

Diademodon, Fig. 22, p. 72.

Dibamus montanus, Fig. 2 oe 431.

Dimetrodon, Fig. 22, p. 7

gigas, Big. 21, p. a

Discoylessus pictus, Big, 5, p. 202.

Eariodes flavicilia, Pl. I1., p. 823.

Ectinohoplia affinis, P\. I., p. 267.

flavicauda, Pl. I., p. 267.

inscripta, Pl. I., p. 267.

— latipes, Pl. I., p. 267.

mitidicauda, Pl. 1., p. 267.

—— nitidiventris, Pl. 1., p. 267.

oculicauda, Pl. 1., p. 267.

scutellata, Pl. I., p. 267.

suturalis, Pl. 1., p. 267.

soror, Pl. I., p. 267.

tibialis, Pl. I., p. 267.

Ectopistes migratorius, Fig. 1, p. 441.

Kmydopsis trigoniceps, Fig. 11, p. 149.

Endothiodon ? microps, Fig. 29, p. 95.

Eois lutearia, Pl. IIL, p. 823.

Hosauravus copet, Big. 5, p. 147.

Epiplema ornata, Pl. IL1., p. 528.

rectangularia, Pl. II1., p. 823.

Kuarctos americanus, Fig. 8, p. 405.

Kupithecia aquanivaria, Pl, TIL,

p. 323. ¢

—— cuprearia, Pl, TIL. p. 323.

pp.

INDEX OF ILLUSTRATIONS.

Feather, Diagrammatic drawing of,

Fig. 1, p. 226.

Fimbrios klossi, P\. 1., p. 423.

Galechirus scholizi, Fig. 12, p. 149.

Gorgonops iorvus, Figs. 4-6, pp. 40-42.

Gorilia gorilla, Fig. 3, p. 11.

pe es peguensis, Fig. 1, p. 428.

var. angularis, Fig, 1,

var. irregularts, Fig. 1,

p. 428.

Hemalia magitaria, Pl. TIL, p. 323.

— mantaria, Pl. I1T.. p. 323.

( Hylephila)

Fig. 1, p. 32.

Helioporus pictus, Fig. 10, p. 213.

Howestia browni, Fig. 21, p. 138.

Humming-Bird, Feather from gorget

of, Fig. 2, p. 226.

Hylobates hoolock, Figs 6,9

Fig. 8, p. 26.

—— muelleri, Fig. 7, p, 24.

Hymenomima nivacaria, P|. I1., p. 323.

—— sinuosania, P|, I1., p. 323.

Hypolepis bella, Pl. 1L., p. 828.

Hammomyia unilineata,

1)Oe ell eb

———— hen

Hyria gemmaria, Pl. TIL., p. 328.

Hyriogona montaria, Pl, ITT. p. 328.

—-— santaria, Pl. IIT., p. 328

Tsochromodes elegantaria, P\. IT., p. 328.

Jentinkia sumichrasti,

pp. 402, 414.

Figs. 6, 13,

King Bird-of-Paradise, Feather from

wing of, Hig. 3, p. 226.

Kiikenthalia borealis, Figs. 1-8, pp. 188-

191, 193, 194.

Larentiopsis costtplaga, Pl. IT., p. 323.

Leptotrachelus, Fig. 22, p. 72.

-—— eupachygnathus,

pp. 56, 57.

Limnoscelis paludis, Fig. 4, p. 145.

Listrophorus argentinus, Fig. 10, p, 366.

—— frontalis, Fig. 10, p. 366.

Lobeza irrorata, Pl. 1., p. $23.

Lycosaurus pardalis, Fig. 17, p. 59.

—-— tigrinus, Fig, 20, p. 64.

Figs, 14, 15,

INDEX OF ILLUSTRATIONS,

Macacus fascicularis, Fig. 26, p. 302.

—— nemestrinus, Fig. 29, p. 306.

—— pileatus, Fig. 28, p. 305.

—— rhesus, Fig. 25, p. 301.

—— sinicus, Fig. 27, p. 303.

—— speciosus, Fig. 380, p. 307.

Megalophrys parva, Fig. 8, p. 205.

Megascolecine, Relationships of, Fig.

p. 113.

Merocausta felinaria, Pl. 1., p. 328.

Metasiopsis proutaria, P). I1., p. 325.

Mictochroa pallidula, Pl. 1., p. 823.

—— paulata, Pl. L., p. 828.

Mixosaurus nordenskjoidi, Fig. 14,

p- 150.

Moresa mona, Pl. 1., p. 328.

Mosasaurus lemonnetri, Fig. 26, p. 155.

Narope cingulata, Pl. 1., p. 823.

Nasua nasua, Figs. 2, 3, 4, 11, 12, 13,

pp- 3895, 397, 400, 410, 412, 414.

Nedusia castra, Pl. I11., p. 323.

Nephodia bonitaria, P|. I1., p. 823.

—— paularia, P). IL., p. 323.

Numia striqularia, Pl. U1., p. 328.

3238.

Oospiia altonaria, Pl. IIL., p.

Ophiacodon mirus, Fig. 8, p. 148.

Ophthalmophora TEA,

p. 323.

Oxydia pallidaria, Pl. I1., p. 328.

columbaria, Te

Paleobatrachus sp.?, Fig. 2, p. 198.

Paleohatteria longicaudata, Fig. 18,

p. 150.

Papio anubis, Figs. 34, 36, pp. 314,

319.

—— hamadryas, Figs. 34, 36, pp. 314,

319.

—— mormon, Fig. 36, p. 519.

—— porcarius, Fig. 35, p. 3817.

—— sphinx, Fig. 34, p. 314.

Paracomisies tristaria, Pl. IL., p. 823.

Peacock’s neck, Branches of blue fea-

ther from, Fig. 4, p. 227.

Pelodytes punctatus, Fig. 7, p. 204.

Peloneustes philarchus, Fig. 25, p. 154.

Pelycosaur skulls, Fig. 21, p. 71.

Perophora thermesia, Pl. 1., p. 323.

Phurys fasciata, Pl. J., p, 828.

Phyllomedusa bicolor, Fig. 9, p. 208

| Pipa americana, Fig. 4, p, 201.

Platecarpus abruptus, Fig. 27, p. 155.

Plestosaurus rugosus, Fig. 24, p. lod.

Polla acutaria, P\. 11., p. 323.

Polygrammodes eximia, P\. 1., p. 323.

Potos caudivolvulus, Figs. 1, 3, 5, 9, 12,

14, pp. 3938, 397, 401, 407, 412,

414,

Presbytes cephalopterus, Fig. 16, p. 279.

| Procolophon trigoniceps, Fig. 7, p. 147.

Procyon lotor, Figs. 1, 8, 4, 10, 12,

pp. 393, 397, 400, 408, 412.

Psaliodes aurantaria, P), II1., p. 528.

Ptychopoda campinaria, Pl, III.,

p. 328.

—— delicataria, Pl. I11., p. 323.

—— lelicaria, Pl. ITI., p. 323.

terminaria, P}, II11., p. 323.

Pygmephorus americanus, Fig. 15,

p. 377.

—— tarsalis, Fig. 14, p. 374.

Rana milleti, Pl. I1., p. 428.

—— montivaga, Pl. I., p. 428.

—— sauteri johnsi, Pl. IL., p. 4238.

| Lhinonyssus caledonicus, Fig. 1, p. 358.

—— coniventris, Figs. 6, 7, pp. 362,

363.

——- echinipes, Figs. 3, 4, p. 360.

—— neglectus, Fig. 5, p. 361.

—— waterstoni, Fig. 2, p. 858.

Rhodomena paularia, Pi. I1., p. 325.

—— santaria, Pl. IT., p. 328.

Rhodoneura oxydata, Pl. I., p. 323.

Rhophalodon ¢, Fig. 27, p. 90.

Jischeri, Fig. 26, p. 89.

Rosema pallida, Pl. I., p. 823.

Saccoploca strigaria, Pl. III., p. 323.

Salamandra maculosa, P\s. 1.-I1., p. 99;

Fig. 1, p. 100.

Scincosaurus crassus, Fig. 3, p. 145.

Seymnognathus whaitsi, Figs. 7-

pp. 46-49, 51, 53, 54.

Scymnosaurus watsoni, Figs. 23, 2

pp. 81, 85.

Seymouria baylorensis, Fig. 6, p. 147.

13,

XV1

Simia satyrus, Fig. 4, p. 15.

Sphenacodon ferox, Fig. 21, p. 71.

Sphenodon punctatus, Fig. 22, p. 153.

Sterosternum tumidum, Fig. 15 p. 150.

Symphalangus syndactylus,

p. 18.

Tachychlora flora, P\. I11., p. 328.

Tarache parana, Pl. 1., p. 323.

Tarsius, Photographs of a living speci-

men of, Pl. 1., p. 184.

Testudo sp., Fig. 28, p. 158.

Tetragonodes geminaria, Pl. IL., p. 328.

0 es

Hig. 5,

INDEX OF ILLUSTRATIONS.

Trematops milleri, Fig. 1, p. 144.

Trotogonia castraria, Pl. 11., p. 323.

Uranocentrodon

p. 144.

senekalensis, Fig. 2,

Varanops brevirostris, Fig. 10, p. 149.

Varanosaurus acutirostris, Wig. 21,

p. 71.

Xenopus levis, Fig. 3, p. 199.

Youngina capensis, Figs. 19, 20, p. 152.

INDEX.

1921.—Pages 1-446.

[New names in clarendon type.

Systematic references in italics.

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

Acanthodica frigida, sp. u., 325.

Acanthus cannabina, 231.

obscurus, 230.

—— pratensis, 230.

Acarapis, gen. nov., 378.

Accipiter nisus, 231.

Achatinine, 264.

Agalychnis moreletii, 207,

Aglossa, 197.

Ailuropoda, 417, 420.

melanoleuca, 392.

Ailuropodide, 420.

Ailurus, 420,

Sulgens, 392.

Alauda arborea ?, 229.

arvensis ?, 231.

Alytes obstetricans, 203.

Anapha venata, 443.

Alaplena, gen. nov., 302.

castrafria, sp. n., 302.

. Anas boscas, 231.

Anisodes antennaria, sp. n., 349.

—— bizaria, sp. n., 343.

—— carolina, sp. n., 344,

—— japaria. sp. n., 344.

—— paranaria, sp. n., 344.

—— vigoraria, sp. n., 345.

Anthropopithecus troglodytes, 2.

Proc. Zoou. Soc.—1921,

Anteosaurus magnificus, gen. et

sp. n., 92.

Apicia geminimacula, sp. n., 331.

—— strigularia, sp. n., 330.

Aplogompha castraria, sp. n., 327.

-— fumaria, sp. n., 328.

—— setinaria, sp. n., 328.

—— yaponaria, sp. n., 328.

Aquila sp., 229, 231,

Areifera, 197, 201.

Arctognathus curvimola, 60.

Arctops, 73.

— willistoni, 36.

Artace regalis, sp. n., 553.

Asio accipitrinus ?, 231,

Azelina cetana, sp. n., 336.

— hanebaria, sp. n., 336.

Bagodares castra, sp. n., 332.

Bassaricyon sj)., 892.

Bassaricyonine, 442.

Bassariscus astutus, 392.

Bassariscine, 421.

Batrachopsis melanopyga, 204.

Boarmia nigraria, sp. n., 338,

Bombinator igneus, 202.

maximus, 202.

XVI

Bombinator orientalis, 202.

pachypus, 202.

Boselaphus tragocamelus (4. 8. L.), 442.

Bradypus tridactylus, 157, 188.

Broomia perpleaa, 150.

Bryocodia paulina, sp. n., 324.

Bubo ef. cinerascens, 229.

Bufo galeatus, 438.

granulosus, 209.

melanostictus, 209.

—— vulgaris, 208.

Bufonide, 208.

Caccabis rufa, 231, 232.

Calamaria pavimentata var. uni-

formis, noy., 426.

Callistomimus, 233.

—— quticollis, 237, 242.

——— amabilis, 243.

ee pelili,ispams zor oek

—— cauliops, 236.

—— ceylonicus, 237, 245.

chalcocephalus, 236, 258.

chlorocephalus, 238.

— coarctatus, 237, 246.

—— d’abreui, sp. n., 237, 242.

— dicksont, 256.

—— @u x, sp..n., 237, 244.

eucharis, 237, 247.

—— humeralis, var. nov. (modes-

tus), 237, 243.

——insularis, var. nov. (modes-

tus), 287, 244.

—— jucundus, sp. n., 236, 239.

—— lebioides, 237.

.._— littoralis, 237, 249.

—— messii, 238.

——- modestus, 237, 248.

— nuir, 237, 244.

nilgirinus, var. nov. (Nair),

237, 240.

—— quadricolor, 247.

quadriguttatus, 238.

quadristigma vax. chalcocephalus,

236, 238.

- rubellus, 236. :

—— sikkimensis, sp. n., 286, 258.

—— subnotatus, sp. n., 206, 239,

INDEX.

Callistomimus suturalis, 2306.

—— venustus, sp. n., 236, 240.

—— virescens, sp. u., 236, 241.

——— vitalisi, sp. n., 237, 246.

—— westwoodi, 245.

-——— zunnanus, 242.

Callurapteryx paularia, sp. n.,

329.

Calotes microlepis, 428.

mystaceus, 429.

Jalyptocephalus gayt, 210.

Campatonema, gen. nov., 334.

—— marginata, sp. n., 334.

Carama incolorata, sp. n., 354.

Casea broilii, 149.

Ceratophrys boie, 211.

ornata, 211.

Cercocehus ethiopicus, 309.

-—— lunulatus, 311.

Cercopithecus ethicps, 292.

burnetti, 286.

— lalandi, 298.

mond, 289.

patas, 289.

petaurista, 299.

—— preusst, 282.

pygerythrus, 298.

---— pyrrhonotus, 292.

— rufoviridis, 298.

— sabeus, 295.

tantalus, 296.

Cervus canadensis occidentalis (Z. 8. L),

443,

Chelura terebrans, 219.

Chiroleptis australis, 212.

Cinclus aquaticus ?, 230.

Cinnyris gutturalis (z. 8. u.), 444.

Coccothraustes vulgaris, 229.

Cochlitoina, 249.

zebra var. fulgurata, 250, 263,

379.

var. obesa, 250, 263, 379.

Colobus, 282.

Coluber oxycephalus, 426.

Columba ef. livia, 229.

palumbus, 231.

Coracias cf. abyssinica, 229.

Corvus corone, 229,

INDEX.

Corvus monedula, 231.

Coturnix communis, 229, 231.

Cratoptera fenestraria, sp. n.,

Sol.

Cryptotis brevis, 212.

Cypselus apus, 251.

Demodex bovis, 371.

canis var. crinacei, 372.

—— chiropteralis, sp. n., 367.

—— gliricolens, sp. n., 368.

—— melesinus, 372.

—— soricinus, 368.

—— talpe, sp. n., 370.

Dendrelaphis subocularis, 426.

Diademodon, 6d.

Dibamus montanus, sp. n., 431.

Dichogaster, 104, 105, 110.

Didymogaster, 115.

Digaster, 115.

Diplocardia, 104, 105.

Discoglossus pictus, 201.

Dryophis dispar (2. 8. 1.), 444.

Eariodes bimaculata, sp. n., 330.

—— flavicilia, sp. n., 530.

Ectinohoplia, 267.

—— affinis, sp. n., 273.

auriventris, 267.

—— flavicauda, sp. n., 273.

—— formosa, sp. n., 269.

—— inscripta, sp. n., 269.

—— latipes, sp. n., 271.

—- nitidicauda, sp. n., 275.

—— nitidiventris, sp. n., 274.

—— oculicauda, sp. n.,

—— scutellata, sp. n., 27

ss SOLOL Speier nls

suturalis, 267.

—— tibialis, sp. n., 271.

Ectopistes migratorius, 441.

Hlosia bufonia, 210,

Emydopsis, 149.

Endothiodon, 94.

Eois lutearia, sp. n., 546.

Hosauravus copet, 146.

Epiplema ornata, sp. u., 300,

)

X1x

Epiplema rectangularia, sp. n.,

351.

Erithacus rubecula, 230.

Eudichogaster, 103, 105.

Eupemphi«c nattereri, 209.

Eupithecia aquanivaria, sp. n.,

345.

—— cuprearia, sp. n., 342.

—— Mmauvaria, sp. u., 343,

Falco tinnunculus, 231.

Fimbrios klossi, gen. et sp. u., 425.

Fringilla celeos, 229, 251.

Garrulus glandarius, 231,

Gazella bennetti (2. 8. L.), 449.

—— subgutturosa (z.8.L.), 443.

Gorgognathus, 78.

Gorgonops, 74.

toruus, 39.

Gorilla gorilla, 9.

Gryllus domesticus, 446.

Gymnodactylus peguensis var.

angularis, nov., 427.

—— peguensis yar. irregularis,

noy. 428.

Hemalia magitaria, sp. n., 346.

mantaria, sp. n., 345.

Hammomyia (Hylephila) unilineata, 31.

Harrisina mephisto, sp. n., 356.

Heleioporus albopunctatus, 212,

pictus, 213.

Hirundo rustica, 229.

Howesia browni, 158.

Hydrocherus hydrocherus (2. 8.

449,

Hyla lichenata, 207,

maxima, 206.

Hylobates agilis, 19.

hoolock, 20.

lar, 25.

—— muellert, 22.

Hylodes fleischmanni, 211.

—— lineatus, 211.

—— martinicensis, 211,

XxX

Hylodes rantformis, 211.

Hylorhina silvatica, 211.

Hymenochirus boettegeri, 200.

Hymenomina nivacaria, sp. n.,

33.

—— sinuosaria, sp. n., 338.

Hypolepis bella, sp. n., 341.

Hyria gemmaria, sp. n., 348.

Hyriogona montaria, sp. n., 349.

—— santaria, sp. n., 348.

Isochromodes elegantaria, sp. n.,

335.

Jentinka sumichrasti, 392.

Kiikenthalia borealis, 187.

Larentiopsis, gen. nov., 339.

costiplaga, sp. n., 340.

Leptodactylus ocellatus, 211.

pentadactylus, 211.

Leptotrachelus eupachygna-

thus, gen. et sp. n., 55.

Limnodynastes peronti, 211.

Limnoria lignorum, 218.

Limnoscelis paludis, 146.

Liolepsis belliana var. anna-

mensis, nov., 429.

Liponyssus arabicus, sp. n., 365.

—— beriesei, sp. n., 362.

bursa, 365.

serpentium, 365.

Listrophorus argentinus, sp. n

366.

—— frontalis, sp n., 366.

Lobeza irrorata, sp. n., 326.

Lycosaurus pardalis, 58.

Lygosoma corpulentum, sg).

431.

stellatum, 431.

CS)

Macacus fascicularis, 302.

——— inuius, 308,- |

INDEX.

Macacus nemestrinus, 306.

pileatus, 305.

—— rhesus, 299.

sinicus, 303.

speciosus, 307.

Macropus walabates (z. 8. L.), 444.

Megalophrys fee, 206.

hasseltii var. pullus, nov., 440.

—— intermedius, sp. n.. 439.

longtpes, 206.

mijor, 205.

montana, 206.

parva, 205,

pelodytoides, 205.

Megascolecine, 126.

Melopsittacus undulatus, 445.

Merocausta felinaria, sp. n., 327.

Metasiopsis proutaria, sp. n., 349.

Microhyla picta, 437.

Mictochroa pallidula, sp. n., 323.

paulata, sp. n., 323.

Milvus ef. ictinus, 229.

Moniligastride, 129.

Monogaster, 121.

Monopeltis capensis, 443.

Moresa mona, sp. n., 326.

Mosasaurus, 155.

Motacilla lugubris P, 230.

Narope cingulata, sp. n., 354.

Nasua narica, 392.

Nasuine, 422.

Nedusia castra, sp. n., 352.

Nephodia bonitaria, sp. n., 337.

—— paularia, sp. n., 337.

Notiodrilus, 116.

Nototrema marsupiatum, 207.

Numia strigularia, sp. n., 333.

vyctimantis papua, 207.

Octocheting, 128.

Octochetus, 105.

Oligocheta, 112.

Oospila altonaria, sp. n., 349.

Ophiacodon mirus, 147.

Ophthalmophora

sp. n., 329.

columbaria,

INDEX.

Otocorys alpestris ?, 231,

Oxydia pallidaria, sp. n., 334.

Paleobatrachus, 198.

Paleohatteria longicaudata, 150.

Papio anubis, 318.

hamadryas, 314.

mormon, 320.

porcarius, 316.

sphine, 312.

Paracomistes ?, 356.

Pelobates fuscus, 204.

Pelodytes punctatus, 204.

Perissogaster, 115.

Perophora thermesia, sp. n., 353.

Phryniscus nigricans, 443.

Phurys fasciata, sp. n., 320.

Phyllomedusa bicolor, 207.

— burmeisteri, 208.

dacnicolor, 207.

Phylloscopus trochitus, 230,

Pica rustica, 231.

Pina americana, 200.

Pipide, 199.

Platecarpus, 155,

Plesiosaurus, 155.

Plionogaster, 115.

Plutellus, 112.

Polla acutaria, sp. n., 332.

Polygrammodes eximia, sp. n.,

355.

Pontodrilus, 120.

Porzana parva ?, 231.

Potosine, 421.

Potos caudivolvulus, 392.

Pratincola rubetra, 230.

Presbytes, 278.

cephalopterus, 278.

entellus, 282.

Procolophon trigoniceps, 146.

Procyon lotor, 392.

Procyonide, 389, 420, 421.

Procyonine, 422.

Psaliodes aurantaria, sp. n., 342.

Pseudis paradoxa, 210.

Ptychopoda campinaria, sp. n.,

347.

— delicataria, sp. n., 347.

—- lilacaria, sp. n., 347.

xxl

Ptychopoda terminaria, sp. n., 346.

Pternohyla fodiens, 207.

Puffinus ef. anglorum, 229.

— cf. fulginosus, 229.

allied to chlororhynchus, 229.

Pygmephorus americanus, 375.

var, Socotrensis, var. nov.,

376.

— pilosus, 378.

spinosus, 377.

tarsalis, sp. n., 374.

Pyrrhocorax graculus ?, 231.

Pyrrhula ewropea, 229,

Ramiella, gen. noy., 109.

Rana graminea, 437.

milleti, sp. n., 482.

—— montivaga, sp. n., 436.

nigrovittata, 433.

—— sauteri var. johnsi, nov., 434.

Rhinonyssus caledonicus, sp. n.,

357.

coniventris, 361.

echinipes, sp. n. (var. of f.

neglectus?), 359.

——- levenseni, 357.

—— neglectus, sp. n., 359.

— waterstoni, sp. n., 359.

Rhodomena paularia, sp. n., 340.

—— santaria, sp. n., 341.

Rhcdoneura oxydata, sp. n., 354.

Rhopatlodon, 88.

Rosema pallida, sp. n., 326.

Ruticilla phenicurus, 230.

Saccoploca strigaria, sp. n., 3dl.

Salamandra atra, 188.

maculosa, 99.

Saxicola enanthe, 230.

Scaphiopus solitarius, 203.

Scincosaurus crassus, 145.

Scolopax rusticula, 232.

Scymnognathus whaittsi, 44.

Seymnosaurus watsont, 80.

Serinus hortulans ?, 229.

Seymouria baylorensis, 146,

Simia satyrus, 14.

Siphonops annulatus, 443,

Speleorchestes poduroides, 375.

ventriosus, sp. n., 373.

Spenceriella, 115, 120.

Spheroma terebrans, 215.

Sphenodon, 153.

Spreo bicolor (z. s. u.), 442.

Sterna fluviatilis ?, 231.

Strix flammea, 232.

Sturnus?, 231.

Symphalangus syndactylus, 17.

Tachychlora flora, sp. n., 350.

Tarache parana, sp. n., 324.

Tarsius, 184.

Tayassu tajagu (2. 8..), 442.

Telmatobius jelksii, 210.

Testudo sp., 154.

—— migra, 443.

Tetragonodes geminaria, sp. n.,

332.

Tetranycopsis horrida, 372.

Theriodontia, 35.

Totanus calidris ?, 231.

Trematops milleri, 144.

Trichecius brevipes, 366.

Trigaster, 104, 105.

Trigastrine, 129.

INDEX.

Trimeresurus monticola, 427.

Triprion petasatus, 208.

Tropidonotus johannis, 426.

Trotogonia castraria, sp. n., 333.

Turdus merula, 229, 230.

musicus, 229, 230.

—— pilaris, 230,

viscivorus, 230.

Uranocentrodon senehkalensis, 144.

Urside, 420.

Varanops brevirostris, 149.

Varanosaurus, 65.

Vespa germanica, 445,

Vultur cf. monachus, 229.

Xenopus calcaratus, 200.

—— clivii, 200.

—— levis, 199.

Youngina, 151.

Zamenis moi, sp. n., 425.

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1921,

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LIST OF CONTENTS.

1921, Part 2 (pp.-1-186).;

EXHIBITIONS AND NOTICES.

Page

The Szcretary. Report on Additions to the Society's Menagerie during the months of

November and December, 1920 183

Mr. E. G. Boutenenur, F.Z.S. Exhibition of, and remarks upon, a Black Salamander

CSGIGIMOMAE QUA) bis gale o as clea eloie aiun, ech ataysier «ale = el crekele ods cojaee es eee ae 183

Prof. J. P. Hitt, F.R.S., F.Z.8. Exhibition of, and remarks upon, lantern-slides of the

Feetus of a Three-toed Sloth (Bradypus tridactylus) ....ececcececececececeseee 183

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

Diammuatys LODN saints ses mishalnsclete esi oe 184

Prof. G. Exuror Smirn, F.R.S., F.Z.S. Exhibition of, and remarks upon, photographs of

a living example of Zarsivs, (Plate 1.) 1.1... cee ce eee tee cette cn eee ees 184

Contents continued on page 3 of Wrapper.

PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THER

ZOOLOGICAL SOCIETY OF LONDON.

PAPERS.

1. The Comparative Anatomy of the Tongues of the Mam-

malia. II. Family 1. Simiide. By Onarues F.

Sonntag, M.D., Ch.B., F.Z.8., Anatomist to the

Society.

[| Received October 14, 1920: Read February 8, 1921. |

(Text-figures 1-9.)

ConTENTS. Page

Introduction Pads ee 1

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The Orang-Outan SEBO Pier Sie 14

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The Slender Gibbon WS geet hte cape ee 2 NS)

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Toe BOwaeMM (CuO xO sob Sed onavanseauen cuoucbonsJdododeacesn pL

Aiea VVshiteshandeduGubbomereserereaciseeeeemnatrs | 20

Summary and Conclusions 28

Bibliography 29

Introduction.

Many papers dealing with the anatomy of the Anthropoid

Apes contain details of the macroscopic appearances of the

tongues. Most of the authors, however, have limited their

accounts to the description of one or more salient features, such

as the number and arrangement of the vallate papille. More

attention, too, has been paid to the large Anthropoids than to

Proc. Zoou. Soc.—1921, No. I. 1

2 DR. C. F. SONNTAG ON THE ANATOMY

the Gibbons. I have referred to nearly thirty papers, and in no

ease did I find a complete account of any of the Simian tongues.

Even Deniker’s paper on the tongue of the Gorilla, which is

the best, is incomplete, for it deals mainly with the tongue in the

foetus and says little about its condition in the adult.

For the purpose of the present communication I have

examined both fresh and preserved specimens in the Society’s

Prosectorium and in the Museum of the Royal College of

Surgeons, the specimens from the latter being indicated by

the words Mus. R.C.S. Of all the tongues examined only the

measurements of fresh specimens are given, for the dimensions

of preserved ones are worthless. I am indebted to Professor

Arthur Keith for permission to examine some of the tongues

described in this and future papers.

Genus ANTHROPOPITHECUS.

THE CHIMPANZEE (A. troglodytes).

Of the three tongues examined one was fresh, one had been

preserved in the Society’s Prosectorium, and one (No. J. 359.1)

was preserved in the Museum of the Royal College of Surgeons.

These are described respectively as specimens one, two, and

three. The fresh specimen was a male from Landana, 8.W.

Africa.

Several writers have stated that Traill first described the

tongue of the Chimpanzee in 1821, but Traill’s paper is entitled

“The Deseription of an Orang-Outan”’ (206). He certainly

described an arrangement of the vallate papilla which closely

resembles that of the Chimpanzee, but he expressly mentions

the tongue as being that of an Orang *.

My fresh epee? has the following measurements:—Total

length Srl’ Cua, Hong from the apex to the antero-median

vallate papilla 6-1 om. length from that papilla to the epi-

glottis 3 em.; width cote the lingual attachments of the

anterior faucial pillars 4:5 cm.; width of the apex 3 cm.;

thickness in the region of the antero-median vallate papilla

3°5 em.; thickness of the apex °3 cm.

The tongue is long and comparatively narrow, and its width

does not decrease greatly from base to apex. Cunningham (118)

is the only author who points out that its elongated form is due

to the shape of the mouth. He also shows how the tongues of

the Chimpanzee and Gibbons differ more from that of Man in

the disproportion between length and breadth than does the

tongue of the Orang-Outan.

The antero-median vallate papilla stands a short distance

behind the summit of an elevation whence the dorsum slopes

towards base and apex, but the declivity is greater in the latter

direction (text-fig. 1 B, d). In this connection the Climpanzee

* Flower states that Traill really described a Chimpanzee from the Gaboon,

OF THE TONGUES OF THE MAMMALIA. 3

is pre-eminent among the Anthropoids, but the degree of slope

varies in different individuals.

The Apex and Lateral Borders exhibit characters which are

common to many of the Anthropoids. The apex is blunt and

may or may not possess a median notch ; the notch only occurred

-Text-figure 1.

af k.

.The tongue of Anthropopithecus troglodytes.

A, dorsum; B. lateral view; C. inferior surface; D. vertical section of the side

of the tongue. Descriptions in text.

in my fresh specimen. It is roughened by conical papille and

tuberculated by large prominent fungiform papille. Their size

varies in different tongues, but they are always numerous ; some

are smooth, glistening, and hemispherical, others are granular

tee

4 DR. C. F. SONNTAG ON THE ANATOMY

and hemispherical, and others may possess a central boss (text-

fig, 2A, J, m, n). All the conical papille have their points

directed backwards. The lateral borders are full and rounded

(text-fig. 1 D, a); they are beset with both conical and fungi-

form papille, and the lateral organs are situated at their

posterior extremities (text-fig. 1 B, g).

Mesial Sulci.—Mesial sulci may be present on the dorsal and

inferior surfaces. In my fresh specimen (text-fig. 1 A) the

median dorsal sulcus extends from the apical notch to a point

4 mm. in front of the antero-median vallate papilla. It is

irregular and invaded by both conical and fungiform papille ;

posteriorly it is deep and lodges fungiform papillae alone. On

the base of the tongue the median row of vallate papille is

situated on a depressed band of mucosa which is bounded by

large papillae and lymphoid nodules.

In my second specimen there is, instead of a median sulcus, a

row of prominent fungiform papille, and the only representative

of the fissure is a small pit, lodging a fungiform papilla, lying

immediately in front of the antero-median vallate papilla.

In my third specimen (No. J. 359.1, Mus. R.C.S.) there is

neither a median sulcus nor a median row of fungitorm papille,

but there is a pit in front of the antero-median vallate papilla.

The small pit described above inust not be mistaken for the

JSoramen cecum of Morgagni which is absent in the Chimpanzee.

The median ventral sulcus will be described later.

Transverse ridges and sulci are absent.

The Papille.

Papille are present on the entire dorsum, apex, lateral borders,

and a bounding zone on the inferior surface. This general distri-

bution 1S SO common among the Anthropoids that it can be

regarded as the rule. The only exceptions are the Orang-Outan

and the Siamang Gibbon, which have areas on the base which are

devoid of papilla. I am unable to say whether these smooth

areas exist in all Siamangs, but they are not present in all

Orangs.

The Circumvallate Papille (text-fig, 1 B, c & d).

The vallate papille vary in number and arrangement in dif-

ferent individuals, but the relation between the species of

Chimpanzee and the papillary pattern has not been stated by

authors. The following types have been described :—

T-form: Flower (28); Dwight (123); Symington (202);

Huxley and Hunter.

Y-form: Gratiolet and Alix (131); Bischoff (7); Miinch [1].

V-form: Cavanna (109); Ehlers (23).

Cruciate form : Mayer (162).

Linear type: Humphry (142).

The numbers of vallate papille vary from three, as recorded

by Wyman (215), to fifteen mentioned by Humphry (142).

OF THE TONGUES OF THE MAMMALIA, 5

In my three specimens the arrangements are as follows :-—

Specimen No, 1.—There are eight papille arranged in the

Y-form (text-figs. 1A & 2B). Each lateral limb has an

outer compound and an inner simple papilla, and the mesial

antero-posterior row consists of four simple elements.

The papillary bodies of the compound papille are oval on plan ;

the inner papille of the lateral rows are circular; the antero-

median papilla is cireular, and the other three papille are oval

on plan. All the papille are conical on elevation (text-figs.

2 A,r, & 1B, c), the narrow ends of the cones being attached

to the bottom of the fosse. The surfaces are all granulated,

possibly by secondary papille. The vallums are lobulated, and

the fosse are more or less patulous (text-fig. 2, p, q, ¢).

Specimen No. 2.—There are eight papille arranged in the

Y-form, and all of them are simple (text-fig. 2 B), The left

lateral row has three papillee, the right lateral limb has two, and

the mesial antero-posterior limb has three. All the papille of

the lateral limbs are circular and prominent, and the vallums

and fosse are well marked. They are surrounded by papillose

ridges passing inwards from the corresponding lateral organs.

The median row of papille consists of oval elements well recessed

beneath their vallums. The most posterior papilla is difficult to

see unless the pedunculated papille of the base of the tongue are

withdrawn from over it.

In Specimen No. 3 the arrangement of the vallate papilla is

uncommon. There are eight papille in the Y-form (text-fig.

2 B), but the two lateral limbs lie closely side by side, and the

vertical limb has one papilla. The left lateral limb has four

papille and the right one has three.

The Fungiform Papille (text-figs. 1 B, 6, & 2 A, m, n, 0).

Fungiform papille are numerous and cover the dorsum, apex,

lateral borders, and a bounding zone on the inferior surface.

They have the usual arrangement in clusters behind the apex,

transverse rows farther back, and oblique chains in front of the

vallate papille. On the lateral borders they are arranged verti-

cally, and on the inferior surface they are in straight lines

passing from without inwards.

They invade the median dorsal sulcus, and may replace it

altogether. In one of my specimens there is a well-marked

median row of prominent fungiform papille im place of the

median sulcus.

In one of my specimens there are more fungiform papille than

those presented to the naked eye, for the lens “shows how filiform

papille entirely conceal many fungiforms.

At the base of the tongue (text- -fig. | A) there are many long

pedunculated papille which belong to “both fungiform and conical

groups. It is only possible, however, to settle by microscopic

examination to which of them any particular papilla can be

referred, They vary in degree of development in different

6 DR. C. F. SONNTAG ON THE ANATOMY

tongues, and I agree with Humphry (142) that some of them

may be a quarter of an inch long.

These pedunculated papille are smaller and fewer than in the

Gorilla, and larger and more numerous than in the Gibbons.

They are absent altogether in the Orang-Outan. Their surfaces

are smooth or granular. The different forms of papille are

shown in text-fig. 2A, m-o.

Text-figure 2.

The papille and lateral organs of Anthropopithecus troglodytes.

a-l, conical papille ; im-o, fungiform papillex ; p—w, circumvallate papille :

AA’, lateral organ; the three figures on the left side of the bottom row

are vallate patterns, described in the text as 2 B.

The Conical Papille (text-fig. 1 B, a, e, & f).

The conical papille have the same form of arrangement as the

fungiforms, and they are seen to the best advantage when the

tongue is allowed to dry, for they then stand up on the surface.

All their points are directed backwards.

They increase in size from before backwards, and from without

inwards, the largest ones on the anterior part of the dorsum

being in and around the median sulcus. They overlap the fungi-

form papille and conceal some of them entirely. In one of my

specimens they stand on ridges like those on the finger-tips.

They belong to both cylindrical and filiform groups. The

filiform varieties have one or more points. When there is only

one the papilla is tapering, and cireular on section. When

OF THE TONGUES OF 'THE MAMMALIA, 7

there are more than one point the body may be cylindrical, or

flat and thin (text-fig. 2 A, ag). The cylindrical types ave

interspersed among the others. They have a granular surface,

and some of them are very rough. In the latter case, however,

it is difficult to be precise as to their character; they may have

lost. long points as the result of handling (text-fig. 2 A, h, ¢).

The relative proportions of the filiform and cylindrical types

differ in different tongues. Hither type may be in excess, and

they may in some cases be evenly distributed, but it is usual to

find one form predominating.

The different forms are shown highly magnified in text-

fiy. 2 A, nos. al.

Lymphoid Tissue.

The base of the tongue contains much lymphoid tissue which

gives the surface a iopnleted appearance, and there is a great

contrast between it and the rough anterior surface when the

tongue is viewed from the side (text- fig. 1B). The degree of

contrast depends greatly on the areal of filiform papille on

the anterior two-thirds of the dorsum; when these are very

numerous the roughness is greater and the contrast more marked.

The lymphoid nodules vary in size, and the large ones have

small central pits, but it is only possible to tell by microscopic

examination whether any one orifice has ducts of glands con-

nected to it or not. Few of the pitted follicles lie anterior to

the most posterior vallate papilla.

The Lateral Organs (text-figs. 1 B, g, & 2, AA’).

Boulart and Pilliet [2] have stated that the lateral organs of

the Anthropoid Apes are well developed, and mention that the

Chimpanzee has twelve ridges in its organs, but they do not say

whether both organs ave the same number of fissures and

ridges. Dwight (123) showed that each organ is convex in-

ternally. I agree with Dwight, but not with Boulart and

Pilliet. All my specimens have organs convex inwardly, but

there is great varinbility in the number of fissures and ridges

as the following figures show :—

Specimen | (text-fig. 2, AA’). .

Right organ: Length 1:55 cm. Ridges 15. Suter 16.

Left organ: Length 1:6 cm. ftidges 14. Suter 15.

Specimen 2.

Right organ: Length 1:4 cm. Ridges 7. Sulci 8.

Left organ: Length 1:7 cm. Ridges 9. Sulet 10.

Specimen 3.

Right organ : — Ridges 10. Suleti 11.

Left organ : — Ridges 9. Suler 10.

Each organ begins anteriorly as a number of sulci on the

infero-lateral aspect of the tongue, but the pieces of the tongue

8 DR. C. F. SONNTAG ON THE ANATOMY

between them are not raised above the general surface. Behind

these the ridges appear and increase both in length and pro-

minence till a point is reached almost level with the outermost

vallate papilla of the corresponding lateralrow. After that they

diminish in size, but increase in prominence till the posterior

limit of the organ is reached. Many ridges are simple, but a

few are divided by small secondary sulci.

Ridges pass across the dorsum from the upper ends of the

ridges of the lateral organs. These either encircle the vallate

papillee or pass backwards on their outer side.

The Inferior Surface (text-fig. 1 C & 1 B, 2).

The inferior surface presents for examination a mesial sulcus,

two plice fimbriatv, the papillary border, frenum and sublingual

fold, or frenal lamella, so called by Pocock (text-fig. 1 B, ’, &

I OS2)y

The mesial sulcus (text-fig. 1C, a) extends backwards from

the posterior border of the papillary zone to the attachment of

the frenum. It is not occupied by any crest as in the Gorilla.

The plice jfimbriate (text-fig. 1 C, c) are, it is acknowledged,

remnants of the sublingua of the Prosimiew, and in one of my

specimens the two plice and intervening piece of mucous mem-

brane have the appearance of an under tongue.

The plice are two longitudinal folds, almost touching in front,

running backwards and outwards from near the apex to a point

well behind the middle of the tongue, but in one of my specimens

they are nearly parallel to one another. They le on the lateral

aspects of the inferior surface between the mesial sulcus and the

edges of the tongue. Each one has undulating and crenated

edges, and increases in width from before backwards.

When a vertical section of the tongue is made about its centre

(text-fig. 1 D) one sees the mucosa of the floor of the mouth (@)

reflected to form the frenal lamella (c). From the upper surface

of the latter it is reflected on to the plica (6), and thence to the

under surface of the tongue (a). The mucosa between the

plica fimbriate and the sublingual fold is thrown into many

small folds.

The Sublingual Fold (text-fig. 1 C, e).

The sublingual fold, or frenal lamella (Pocock), lodges

Wharton’s Ducts. It is triangular in shape with the base

behind. The apex is bifid and the two ducts open on the points

which are very sharp; the edges are undulating and crenated.

Its mucosa is continuous with that of the floor of the mouth

below and the frenum linguz above.

Some have said that the sublingual fold corresponds to the

sublingua of the Prosimiz, but most anatomists are agreed that

the plicee fimbriate represent it, so the tongue of the Chimpanzee,

which possesses both, is sufficient to disprove the statements of

the former group of observers.

OF THE TONGUES OF THE MAMMALIA. 9

The Papillary Border (text-fig. 1 C, 6).

The narrew papillary border has fungiform and conical

papille, the latter belonging to the filiform and cylindrical

groups. They are arranged in lines passing from without in-

wards. The fungiform papillae are small and most numerous

beneath the apex of the tongue.

The Frenum Lingue (text-fig. 1 C, d).

The frenum is short and thick. It runs from the upper

surface of the sublingual fold to the under surface of the tongue

from the posterior end of the median ventral sulcus backwards.

No lytia is present in the interior of the tongue.

When the apex is examined histologically, no Apical Gland of

Nuhn is seen. In this connection it agrees with the Gorilla and

Gibbons, and differs from Man and the Orang-Outan.

There is only a narrow groove between the base of the tongue

and the epiglottis (text-fig. 1 B, /).

Genus GORILLA.

THE GorILLA (G. gorilla).

The literature containing details of the structure of the

tongue of the Gorilla is not so large as that dealing with the

tongue of the Chimpanzee, but the papers are more complete.

Ehlers (23), Bischoff (7), and Duvernoy (22) have written

accounts of the adult tongue, and Deniker (17) has described

the foetal tongue very fully, but he has said little about the

adult form. ‘Boulart and Pillet [2] have not mentioned the

Gorilla in their study of the lateral organs of the Mammalia.

The specimen which | examined (No. J. 358.1, Mus. R.C:S.)

exhibits features which have not been mentioned by these

authors, or are different from the conditions described by them.

The tongue has not such a great disproportion between its

length and width as that of the Chimpanzee. It is comparatively

broad, and in this connection I agree with Bischoff (7), and -

differ from Duvernoy (22) who said it is narrow. The whole

organ appears rectangular. It slopes gently from the vallate

papillary region to the apex, and more steeply from the vallate

region to the epiglottis; it differs, therefore, from the tongue of

the Chimpanzee.

The apex is square-cut, has no mesial notch, and bears conical

and fungiform papillae which are thickly clustered; the latter are

not so prominent as in the Chimpanzee, but are more marked

than in the Orang-Outan.

The lateral borders have large and medium-sized fungitorm and

conical papille arranged in vertical lines, and they are fissured by

numerous sulci prolonged on to them from the dorsum. At

their posterior extremities one sees the outer ends of the fissures

and Jamine of the lateral organs. In this respect the tongue

agrees with that of the Orang-Outan, and differs from those of

10 DR. C. F, SONNTAG ON THE ANATOMY

the Chimpanzee and Gibbons, in which most of the lateral

organs lie on the lateral borders of the tongue.

Mesial Sulci.—Mesial sulci are present on the dorsal and

inferior surfaces. The dorsal sulcus lies in the centre of the

anterior two-thirds ; it 1s wide and shallow, and contains many

conical and few fungiform papille. Just in front of the antero-

median vallate papilla there is a small pit containing both conical

and fungiform papilla, but this must be regarded as a part of the

mesial sulcus. It must not be mistaken for a foramen cecum,

which does not exist in the Gorilla. There is no median suleus on

the base of the tongue.

The median ventral sulcus (text-fig. 3 B, c) begins in front at

the posterior edge of the papillary zone and runs back, widening

as it goes, till it terminates in a triangular pit into which the

frenum passes. It lodges a median fold of mucous membrane

termed the plica mediana or mesial crest (text-fig. 3 B, d).

Transverse Ridges and Sulci.cSeveral transverse ridges and

sulci are present on the anterior two-thirds of the dorsum. They

are undulating and irregular in direction, and some of them cut

the lateral borders of the tongue and run inwards on the

bounding papillary zone of the inferior surface. The sulci are

narrow and the ridges are covered with papille.

In the interior of the tongue there are neither lytta nor Apical

Gland of Nuhn.

The Papille.

Papille are present on the entire dorsum, apex, lateral borders,

and a bounding zone on the inferior surface. The last is better

developed than in the Chimpanzee and Orang-Outan.

The Circumvallate Papille (text-figs. 3 A & 3 D, a, 6).

In my specimen there are seven vallate papille arranged i

the form of a Y, and the notable feature is that most of them

are compound. Each lateral limb has two papille and the

median one has three.

The two papille of the right lmb are compound (text-

fig. 3 D, a) and lie very close together, for their vallums are only

separated from one another bya narrow fissure. Both fossee and

vallums are prominent and the papillary elements are recessed

below the vallums. The papillary bodies are granulated, possibly

as the result of secondary processes.

The outer papilla of the left lateral limb (text-fig. 3 D, 6) is

compound and the inner one is simple. The fosse are patulous

and the two vallums are in continuity. It seems as if the mner

papilla is an element which has been separated off from the outer

compound papilla. In the case of the compound papilla the

elements are recessed below the vallum, but the simple papilla is

more prominent. The surfaces of the elements are granular.

The outer papille of the lateral limbs are level with the

OF THE TONGUES OF THE MAMMALIA. Wy

Text-figure 3.

' ON Mp

Zs ty ;

Se ae

The tongue of Gorilla gorilla.

A. dorsum; B. inferior surface ; C. lateral view; D. a, 6, d, e, papilla; D. ¢, Jateral

organ showing the lateral vallate papilla above its posterior laminz.

12 DR. C. F. SONNTAG ON THE ANATOMY

posterior extremities of the lateral organs as in the Orang-Outan

(text-figs. 3A & 3D, c). In the Chimpanzee and Gibbons, on

the other hand, they are level with the central rows and sulei of

the lateral organs.

The median row of vallate papille has three elements. The

anterior and posterior ones are simple, but the middle one is

compound.

The following numbers and arrangements have been re-

corded :—

1. Five papille in V-formation—Ehlers (23).

2. Five papille in V-formation in the feetus—Deniker (17).

3. Six papille in V-formation—Duvernoy (22).

4, Seven papille in V-formation-—Bischoff (7).

5. Hight papille in V-formation—Duvernoy (22).

My specimen, therefore, differs in the type of arrangement of

its vallate papillee from that described by other authors.

The Conical Papille.

The conical papille have the same forms and arrangements as

in the Chimpanzee, but they are not so large in the anterior two-

thirds of the dorsum. Behind the vallate region, however, the

large pedunculated papille are larger and much more numerous

than in the Chimpanzee. These latter papille are tapering or

club-shaped, and many of them have small secondary processes

as described and figured by Bischoff (7). Their points are

directed backwards, and some of them overlap the most posterior

vallate papilla (text-fig. 3 D, e).

The Fungiform Papille.

The fungiform papille are not so large or so numerous as in

the Chimpanzee, but they exceed those of the Orang-Outan both

in size and numbers. They are in clusters behind the apex, in

transverse rows farther back, and in oblique chains in front of

the vallate papille. They are most numerous behind the apex

and close to the lateral borders of the tongue, and diminish in

numbers towards the mesial suleus. There are, however, po

fungiform papille overhung by filiform types.

When the tongue is viewed laterally (text-fig. 3 C, FP) one sees

how prominent many of the fungiform papille are, and some of

them appear almost pedunculated. In this connection the Gorilla

differs from all the other Anthropoids.

On the sides of the tongue the fungiform papille are in

vertical chains, and they are in rows passing from without

inwards on the inferior surface (text-fig. 3 C, /'P).

The Lateral Organs (text-fig. 3C, LO, & 3 D, c).

The lateral organs of the Gorilla have been omitted from

Boulart and Pilhet’s paper. They are situated on the edges of

the dorsum of the tongue, beginning posteriorly on a level with

OF THE TONGUES OF THE MAMMALIA. he

the external vallate papille of the lateral rows, and extending

forwards to a point level with the posterior extremity of the

mesial suleus. The numbers of ridges and elevations are as

follows :—

Right organ: Length 1-3 em. Ridges 11. Sulci 12.

Left organ : Length 1°35 em. Ridges 12. Suler 13.

In his study of the tongue of the fcetal Gorilla, Deniker (17)

has shown that the lateral organs are better developed than in

the adult.

The ridges and sulci just cut the lateral borders of the tongue.

They are long, narrow, and tapering ; some are twisted and some

have secondary fissures.

The Inferior Surface (text-fig. 3 B).

The inferior surface presents for examination a papillary

border, a frenum, a sublingual fold, two plice fimbriatee, a mesial

crest, and a mesial suleus which has already been described. It

has, therefore, the same structures as the tongue of the Chim-

anzee, with the mesial crest in addition.

The papillary border maintains the same width across the

under surface of the apex, but it widens out from before

backwards along the lateral borders. It bears conical and

fungiform papille whose method of disposition is of the usual

type (text-fig. 3 B, a).

The most notable feature is a row of closety-set club-shaped

conical papille bounding the zone internally (text-fig. 3 B, /).

They increase in size from before backwards, and they he flat

against the surface of the tongue. Some are entire and others

are subdivided into lobules as shown in text-fig. 3 D,d. In no

other Anthropoid tongue is there a uniform row of these papille.

The mesial crest, ov plica mediana, is a fold of mucous mem-

brane occupying the ventral mesial sulcus (text-fig. 3B, d). It

runs along the anterior border of the frenum, and thins out and

disappears on the dorsal surface of the sublingual fold. Deniker

(17) considers that it is a remnant of the sublingua of the

Prosimie.

The plice fimbriate (text-fig. 3B, 6) are united anteriorly at

the posterior border of the papillary zone on the inferior surface.

When they are traced backwards they diverge and become wider

and more prominent. They have undulating free margins, and

ave pale in colour, The mucosa between them is pale and the

vemainder is pink in preserved specimens, so the plicx and inter-

vening part together look like a sublingua. A vertical section of

the tongue of the Gorilla resembles that of the Chimpanzee

(text-fig. 1 D).

The frenal lamella, or sublingual fold, is triangular in shape

and has a rounded entire apex (text-fig. 3B, gy). Wharton’s

Duets open on its dorsal surface, and the bristles shown in

text-fig. 3 B, e, pass into them,

14 DR. C. F. SONNTAG ON THE ANATOMY

Bischoff (7) denied the existence of a frenum, but Ehlers (23)

and Deniker (17) saw one. In my specimen it runs from the

dorsal surface of the sublingual fold to a triangular depression on

the inferior surface of the tongue (text-fig. 3 B).

Genus SIMIA.

THe Orane-Ouran (S. satyrus).

The tongue of the Orang-Outan resembles that of Man in its

relative proportions of length and width and certain of its struc-

tural peculiarities. J examined three preserved specimens in the

Museum of the Royal College of Surgeons, and the following

description refers to No. J.421.3. It is designated as specimen

No. 1 here.

The apex is rounded, has no mesial notch, and possesses conical

and fungiform papille, but both forms are small.

The lateral borders are massive, and have both conical and

fungiform papille distributed in the usual manner. Only the

outer ends of the lamine and sulci of the lateral organs cut the

lateral borders of the tongue.

The dorsum has no median or transverse sulci.

The Papille.

Papille cover most of the dorsum, apex, lateral borders, and a

bounding zone on the inferior surface.

The Circumvallate Papille.

The following numbers and arrangements of the vallate panillee

have been recorded :—

1. Ten papille in V-formation—Flower (28), Boulart and

Pilliet [2].

2. Hight papillae in V-formation—Miinch [1].

3. Seven papille in V-formation—Sandifort (271).

4, Three papille on each limb of a V—Fick (255).

5. Hight papille in the T-form—Traill (206) *.

I observed the following numbers and arrangements of the

papillee :—

Specimen No. 1. (text-fig. 4 A).—The papille are arranged in

the V-form, but the angle embraced by the two limbs is more

acute than in Man. ‘There is an apical papilla and three papille

on each limh, the whole series standing on a raised smooth band

of tongue. The papille of the left limb are all simple, and the

most external one has a small umbilicus (text-fig. 4A). The

most external papilla of the right limb is simple, and the middle

and inner papille are compound. The apical papilla is com-

pound. All the papillary bodies are round or oval on plan and

conical on elevation, with the free broad end of the cone over-

lapping the vallum. The fossz are well marked.

* See footnote on page 2.

OF THE TONGUES OF THE MAMMALIA. 1)

Specimen No. 2.—The papille which are twelve in number,

are arranged in the form of a V, but they do not stand on a

raised band of tongue as in specimen number one. Several of

them are compound.

Text-figure 4.

The tongue of Simia satyrus.

A. dorsum; B. ventral surface. The mucosa has been dissected back to show

Wharton’s Ducts; the arrows point to the actual positions of the sublingual

glands, and the dotted lines indicate the positions of the plicee fimbriate.

The Fungiform Papille.

The fungiform papille are very small. They have the same

arrangement in clusters and rows as occurs in the Chimpanzee

and Gorilla. In the centre of the dorsum they are concealed by

long filiform papille. On the inferior surface of the tongue they

are larger than on the dorsum, and they are disposed in lines

passing from without inwards. A double row forms a prominent

arch round the anterior extremity of the frenum (text-fig. 4 B).

The Conical Papille.

The conical papille are marked features on the anterior two-

thirds of the dorsum (text-fig. 4), but they are more pronounced

in some individuals thanin others. They have the same arrange-

ment in clusters, transverse rows, and oblique chains as in the

other Anthropoids, and they increase in size from before back-

wards and without inwards. They are very long on an area in

16 DR. C. F. SONNTAG ON THE ANATOMY

the centre of the anterior two-thirds of the dorsum, but, as their

points face in all directions, they appear tangled and without any

definite method of arrangement. In that central area, again,

fungiform papille are concealed by the conical forms. There is,

therefore, a similarity to the condition already described in the

Chimpanzee, and shown in text-fig. 1 A, but the papille are more

diffused in the latter.

The papille belong to the filiform and cylindrical types, but

the former predominate, thereby giving the dorsum its shaggy

appearance.

The base of the tongue is devoid of the long conical papille

which are present in the other Anthropoids, and it resembles

that of Man in this respect.

On the sides of the tongue they are in vertical chains, and on

the inferior surface they are in lines passing from without

inwards.

In all situations, except on the central area of the dorsum

described above, the points are directed backwards.

The Lateral Organs.

Boulart and Pilliet [2] state that the lateral organs have each

twelve lamine, but I found that the numbers of fissures and

ridges differ on the two sides as follows :—

Right organ: Length 16cm. Ridges 14. Sulei 15.

Left organ : Length \-6em. Ridges 12. Sule 13.

The ridges run obliquely from below upwards, and they

diminish in size both forwards and backwards, but the organs

are continued by simple folds of the mucosa.

The lateral organs end posteriorly on a level with the most

external vallate papilla. The greater part of each lies on the

dorsum as in the Gorilla, but some of the anterior lamine and

sulci project considerably on to the lateral borders of the tongue,

Lymphoid and Glandular Tissue.

The Orang-Outan agrees with Man and differs from all other

Anthropoids in the possession of the Apical Gland of Nuhn.

Mucous and serous glands occur as usual on the base of the

tongue, but there are no large follicles with central pits as in

Man and the Chimpanzee.

On the inferior surface there are several small pits surrounded

by a raised zone (text-fig. 4 B), but histological examination alone

will reveal their true nature. J have been unable to study these

as my specimens have been preserved too iong in formalin for

satisfactory histological work.

The frenum lingue is well-marked, as in Man, the plice

Jimbriate are of no greater development, and the frenal lamella

is as in the human tongue; it is not a triangular process as

OF THE TONGUES OF THE MAMMALIA. 17

in the other Anthropoids, but simply a fold over Wharton’s

Ducts.

The tongue of the Orang-Outan, therefore, resembles that of

Man in the following particulars :—

. Its general proportions.

. Its rounded apex.

. The V-type of vallate papille.

. The absence of long conical papille on the base.

. The frenum lingue.

. The small proportions of the fimbriate plice.

. The nature of the sublingual fold.

. It has an Apical Gland of Nubhn or Blandin.

CO NI Oo Ol H Oo DO

In all these particulars it differs from the Chimpanzee and

Gorilla, so it must be placed next to that of Man in a classifica-

tion of tongues.

SYMPHATANGUS.

THE SIAMANG (S. syndactylus).

(Specimen No. J. 357.3, Mus. R.C.S.)

The tongue is long and narrow, and the distance between the

antero-median valiate papilla and the epiglottis is long (text-

fig. 5).

The apex is rounded and devoid of a notch. It bears conical

and fungiform papille which have the usual disposition.

The lateral borders are full and rounded. They bear conical

papille with backwardly-directed points, prominent fungiform

papille, and the chief parts of the lateral organs. This condition

is present in all Gibbons, but the fungiform papille vary in size

and prominence in different species.

Mesial and transverse sulci are absent on the anterior two-

thirds of the dorsum. Both are present on the base of the

tongue, but these may have been induced by the preserving fluid.

The mesial ventral sulcus begins at the posterior border of the

papillary zone, and runs backwards to open into a triangular

depression into which the frenum passes.

The Papille.

Papillz are present on the apex, lateral borders, the entire

anterior two-thirds of the dorsum, the sides of the posterior

third of the dorsum, and the hounding zone on the inferior

surface.

On the posterior third of the dorsum there is a large central

area devoid of papille (text-fig. 5). It stretches forwards

anterior to the median row of vallate papille, and sends out

limbs on which the lateral papille stand. It extends right back

to the epiglottis, and is ridged and furrowed at its posterior part.

Anteriorly it is bounded by small conical papille, and large

Proc. Zoou. Soc.— 1921, No. I]. 2

18 DR. C. F. SONNTAG ON THE ANATOMY

conical papillae oceupy the spaces between its lateral borders and

the edges of the tongue.

The Orang-Outan is the only other Anthropoid which has a

bare area (text. fig. 4 A), but itis V-shaped and smaller in size.

The Circumvallate Papille.

Five vallate papillz are arranged in the form of a Y ; of these,

three form a mesial row, and there is a right lateral and ai left

lateral papilla. All the papille are circular on plan and conical

on elevation, and the fossa are well-marked. But the vallums

are not demarcated off from the smooth area on which the

papille stand. All have smooth bodies, and the right and left

lateral ones are umbilicated.

Text-figure 5.

The tongue of Symphatangus syndactylus, showing the dorsum, inferior

surface, and right lateral organ with lateral vallate papilla.

The Fungiform Papille.

The fungiform papille are present over the anterior two-thirds

of the dorsum except for a zone along the mid-line. They have

the usual distribution in clusters and rows, and are never very

large nor concealed by conical ,papille. On the lateral borders

and inferior surface they are arranged in lines running vertically

or from without inwards.

The Conical Papille.

They have the usual form of distribution in clusters and ridges,

and their characters are visible to the naked eye right forward

to the apex. Most of them are filiform with the points eine cid

backwards, or backwards and inwards.

OF THE TONGUES OF THE MAMMALIA. 19

The conical papillee on the base increase in size from before

backwards, but they never attain the degree of development

which is exhibited on the tongues of the Chimpanzee or Gorilla.

They belong to the cylindrical type, and many of the large ones

at the base have long hair-like processes. These basal papille

are arranged in two groups—one on each side of the non-

papillary area.

No foramen cecum, lytta, or plice fimbriate are present.

The Lateral Organs (text-fig. 5).

The two lateral organs, which are convex inwardly, have the

following measurements, etc. :—

Right organ: Length1-3em. Ridges 11. Sulei 12.

Left organ : Length 13cm. Ridges 12. — Sulet 13.

The organs are situated chiefly on the sides of the tongue, but

the inner extremities of their laminze and sulci extend on to

the dorsum. Anteriorly and posteriorly they are continued by

simple folds of mucous membrane. These characters are common

to all the Gibbons and the Chimpanzee.

The lateral vallate papillze are level with the lamine and ridges

of the posterior halves of the organs, but they are not so far back

as in the Gorilla or Orang-Outan, and not so far forwards as in

the Chimpanzee and many other Gibbons.

There are no lymphoid nodules with central pits on the base of

the tongue.

The frenwm is short,and runs from the floor of the mouth to a

triangular depression on the inferior surface of the tongue.

The Siamang is the only Gibbon which has no bifid triangular

sublingual fold through which Wharton’s Ducts pass. The latter

open on caruncule sublinguales, as in the Orang-Outan, but it was

not at all clear if these were overlapped by plice in the specimen

which I examined.

In the presence of the bare area on the base of the dorsal

surface and in the characters of the openings of Wharton’s

Ducts, the tongue of the Siamang agrees with that of the Orang-

Outan, and differs from the tongues of all the other Simiide.

Genus Hy LoBATEs.

THE SLENDER GrBBon (ZZ, agilis).

I have not had the opportunity of examining the tongue of

H. agilis, but Flower (28) points out that it has the following

characters :—

1. The tongue narrows slightly from base to apex.

2. The apex is obtusely rounded.

3. The vallate papille are small, irregularly placed, and adopt

the V-formation.

4, Fungiform papillz are large and evenly distributed

20 DR, C. F. SONNTAG ON THE ANATOMY

5. Conical papille short and thick on the anterior part of the

dorsum.

6. Conical papille at the base of the tongue large, soft, and

pointed. ;

7. Lateral organs distinct.

8. Sublingual fold bifid.

Tue Hoorock Gipson (f. hoolock).

(Specimen No. J. 357.2, Mus. R.C.S.)

The tongue is short, compact, and rough, and tapers from base

to apex (text-fig. 6).

The apex is rounded and has no mesial notch. It bears conical

and fungiform papill, but the latter are small and inconspicuous.

The Jateral borders are the same as in S. syndactylus, but are

shorter.

The Papille.

The papille have the usual general distribution, but there is

no smooth non-papillary area at the base as in S. syndactylus.

The Circunwallate Papille.

There are four vallate papille arranged in the Y-form. Two

are mesial, and there is a right and a left lateral papilla

(text-fig. 6).

The right and left lateral papille are level with the central

lamine and sulci of the lateral organs. They are circular on

plan and conical on elevation, the narrow ends being attached to

the bottoms of the fosse. The fosse are plain, and the vallums

are lobulated and surrounded by conical papillee.

The two mesial papille (text-figs. 6 & 9, 19) are close together.

The anterior one is circular and the posterior one is oval on plan,

and both are conical on elevation with the broad ends free.

Both Jie within a depression surrounded by a prominent lobu-

lated ridge of mucosa, and the space between them and the latter

is crowded with small cylindrical conical papille. The fosse are

not very prominent. All around the common rampart there are

conical papillee.

The Fungiform Papille.

The fungiform papille are disposed over the whole of the

anterior two-thirds of the dorsum. As many of them are con-

cealed by the conical papilla, it is necessary to employ a lens to

detect them all. The largest ones on the dorsum are immediately

in front of the vallate papille, but all the others are small.

There are few on the lateral-borders of the tongue. On the

papillary zone of the iferior surface they are larger than any-

where else. They occupy most of the zone beneath the apex, but

farther back they are arranged in an irregular double row, on

which some of the papille are hemispherical and others are

OF THE TONGUES OF THE MAMMALIA. 21

pedunculated. Some run into the lateral organs (text-fig. 6

FP).

The Conical Papille.

On the anterior two-thirds of the dorswm the conical papille are

strong and coarse, and give the tonguea rough appearance. Their

disposition in clusters and rows is not very clearly marked, but is

maintained. Several groups are arranged in an undulating

manner, and the points of the papillae point backwards, back-

wards and outwards, or backwards and inwards (text-fig. 6).

They conceal several of the fungiform papille.

Text-figure 6.

OS VO,

The tongue of Hylobates hoolock, showing the dorsum, inferior surface,

and right lateral organ.

Behind the vallate papille the tongue is covered with cylindrical

conical forms. On the area lying between the lateral vallate

papille and the level of the posterior pole of the rampart common

to the two mesial vallate papille they are small, but behind that

level they are large, and many have strong single processes (text-

fig. 9,5). There are some small cylindrical forms anterior to

the vallate papillee.

The characters and disposition of the conical papillae on the

sides and inferior surface are the same as in all other Anthropoids.

The Lateral Organs (text-fig. 6).

Right organ: Length 1:2 cm. Ridges 12. Sulci 13.

Left organ: Length 1:25em. Ridges 10. Suler 11.

22 DR. C. E. SONNTAG ON THE ANATOMY

Both organs are convex inwardly and lie on the lateral borders,

with the inner extremities of their ridges and sulci on the dorsum.

In the specimen (No. J. 357.2, Mus. R.C.S.) which I examined

more of the left organ is on the dorsum, but that may be

produced by mechanical distortion. Most of the ridges are sub-

divided, and fungiform papille invade the anterior ones on both

organs, Simple folds of the mucosa lie anterior to each organ,

and the lateral vallate papillz are level with the central laminz

and sulci. han

The following structures are absent :—

1. Large lymphoid nodules with cential pits.

2. Lytta.

3. Plice fimbriate.

4. Apical gland of Nuhn.

5. Foramen cecum.

The frenum is of moderate length.

The ventral mesial sulcus is narrow, deep, and lodges an antero-

posterior crest. It runs back from the posterior border of the

papillary zone to the triangular depression into which the frenum

passes.

The bifid sublingual fold is not present in the specimen in the

Museum of the Royal College of Surgeons, and the frenum passes

straight from the triangular pit on the inferior surface of the

tongue to the mucous membrane of the floor of the mouth.

THE Bornean Gipson (/7. muellerd).

The specimen described helow was obtained from a female

Gibbon from Borneo which died in the Society’s Menagerie.

Measurements.—Total length 6°5 em. ; length from the apex to

the central vallate papilla 5°25 cm.; length from the central

vallate papilla to the epiglottis 1:25 cm.; width between the

attachments of the anterior faucial pillars to the edges of the

tongue 2°8 cm.; width of the apex 1:4 cm.; thickness at the

central vallate papilla 1:2 em. ; thickness of the apex °6 cm,

The tongue is, therefore, long and narrow, and it is pig-

mented.

Pigmentation.—The dorsum in front of the vallate papille is

bluish black in eolour, and the glistening bluish-black fungiform

papille appear prominently on it, as their colour is darker than

the rest of the dorsum. The dorsum behind the vallate papillee

is not pigmented, is white, and the lateral organs lie at its antero-

Jateral aspects.

The inferior surface has a central unpigmented area shaped

like the head of a spear, whose apex reaches the posterior border

of the papillary bounding zone. From the posterior extremity

of the hastate central area a narrow clear band runs on each side

to the lateral organ, and thus brings the clear areas of dorsum

OF THE TONGUES OF THE MAMMALIA. 25

and inferior surfaces into continuity. The rest of the inferior

surface is bluish black in colour.

A pex.—The obtuse apex has a fine central notch. It bears fine

conical papille on the dorsum and lateral borders, and both

conical and fungiform papille on the inferior surface. The

conical papille are of the cylindrical and filiform types, and the

latter have their points directed backwards. The numbers of

points differ greatly.

Median sulci.—Median longitudinal sulci are present on both

dorsum and inferior surfaces. The median dorsal suleus runs

back from the apex for a distance of 1:4 em. The median

inferior sulcus begins at a point *5 cm. posterior to the apex for

1-5 cm., becoming wider as it goes, till it opens into a triangular

area to which the frenum is attached. It is shut off from the

apex by the papillary border.

Lateral borders.—The lateral borders have fungiform and

conical papille. The latter are both cylindrical and filiform

with their points directed backwards.

The inferior surface.—The inferior surface is surrounded by a

papillary border which is rough to the touch; it varies in width

from *2 cm. at the apex to °5 cm. at the lateral organs, so it

becomes wider when it is traced backwards. It bears beth fungi-

form and conical papillae. The former are arranged in two rows,

of which the inner one consists of closely-set elements, but the

outer papille are more discrete. There is, however, no disposi-

tion according to size, for both rows have different sizes of

members. ‘The conical papille are cylindrical and filiform with

their points directed backwards.

Internal to the papillary border the mucous membrane is

smooth and, with the exception of the mesial sulcus, furrowless.

Plice fimbriate, lytta, foramen cecum, and the Apical gland of

Nuhn are absent.

The Papille.

Papille cover the entire dorsum, the apex, lateral borders, and

a bounding zone of the inferior surface.

The Circumvallate Papille.

There are three vallate papille arranged in the form of a

triangle whose apex is in the mid-line posteriorly, and the vertical

angle included is obtuse. The lateral papillee, which are simple,

are *7 cm. distant from the compound apical papilla and 1:3 em.

from one another. «

The apical papilla is very prominent, is oval in shape with the

long axis antero-posterior, and measures *6 em. by “3 cm. Its

fossa is narrow and irregular, and lodges two elements (text-

figs. 7 & 9,17). These have smooth glistening bodies, and the

anterior one has a small central depression. The entire series of

elements is dumb-bell-shaped and the vallum sends small lobules

24 DR, C. F. SONNTAG ON THE ANATOMY

into its hollows. The vallum is finely lobulated. Perhaps this:

papilla has been formed by fusion of elements similar to those in

the mesial limb of H. hooloch (text-fig. 9, 19).

The two simple lateral papille are ovoid, and the papillary

bodies and vallums are granular (text-fig. 7).

All the papillz are conical on elevation, the broad ends of the.

cones projecting beyond the vallums (text-fig. 9, 17).

The Conical Papille (text-fig. 9, 1-13).

The conical papille have the usual arrangement in clusters and

rows of varying degrees of obliquity, and belong to the cylindrical

and filiform series. The latter have one or more points, all of

which are directed backwards. Those shown in text-fig. 9,

nos. 1-5, have shafts circular on section, but the papille (text-

fig. 9, 6-8) are flat.

Text-figure 7.

The tongue of Hylobates muelleri.

a, central fungiform papilla; 6, d, conical papille; ¢, epiglottis; 7, papillary bounding

zone of inferior surface; m, part of frenum; 2, frenal lamella; 0, p, mesial

ventral sulcus: g, pigmented area ; 7, unpigmented area.

At the base of the tongue there are large conical papillae whose

points are directed backwards and inwards. Their surfaces are

granular, and they may have prominent secondary papille. They

are not so well developed as in the Gorilla and Chimpanzee, but

their secondary processes are larger than in either of these

animals. ‘They are shown in text-fig. 9, 9-13. Nei

The Fungiform Papille (text-fig. 9, 14-16).

There is little to note about the fungiform papilla. They have

the same type of arrangement as in the other Anthropoids, and

OF THE TONGUES OF THE MAMMALIA, 25

they are evenly distributed. They are most numerous behind the

apex on the dorsum and inferior surface.

The Lateral Organs.

The lateral organs, as in the Chimpanzee, lie on the sides of

the tongue, with the upper ends of their fissures and ridges pro-

jecting on to the dorsum (text-fig. 7). Hach organ is convex

inwards, and has both plain and subdivided ridges. The two

organs are compared as follows :—

Right organ: Length*9cm. Ridges 12. Sulci 13.

Left organ : Length 9 cm. Ridges1l. Sulei 12.

The middle ridges and sulci of the lateral organs are level with

the lateral vallate papille.

Lymphoid Tissue.

Small and large lymphoid nodules are present on the base of

the tongue, and some of the large ones have central pits which

are, however, larger than in the Chimpanzee.

The Frenum.

The frenum is short and thick. It runs from the upper surface

of the sublingual fold to be inserted into a triangular depression

on the under surface of the tongue and from its under surface to

the floor of the mouth.

The Sublingual Fold.

The sublingual fold is triangular, and has a bifid apex and

crenated edges.

THE WHITE-HANDED GrBBon (JH. lar).

T examined two preserved specimens (Nos. J.357.1 & J. 357.2)

in the Museum of the Royal College of Surgeons. The former is

oval in shape and the latter is conical, but the differences in form

are possibly due to varying degrees of muscular contracture.

They both differ also in the character of their papille and lateral

organs. Unless differences are specifically stated here, the

characters described are common to both.

The apex is similar to that of S. syndactylus.

The lateral borders are full and rounded. They possess both

conical and fungiform papille, and the latter are very prominent ;

some are hemispherical and others are pedunculated. At their

posterior extremities the lateral organs are situated.

No mesial sulci are present on any part of the dorsum.

The Papille.

The papille have the same distribution as in H. muellert and

H. hooloek, and there is no smooth area on the base of the tongue

as in S. syndactylus,

26 DR. C. F. SONNTAG ON THE ANATOMY

The Circumvallate Papille.

In specimen No. J.357.1 there are four papille arranged in the

Y-formation. These are so disposed that there is a right lateral

papilla, a left lateral papilla, and two mesial papille—anterior

and posterior. All are small in size.

Text-figure 8.

Ht)

HEA

ie i

NVI

The tongue of Hylobates lar.

Two forms are shown, with dorsum (upper row) and inferior surface (middle row).

The figures R and L are lateral organs of the figure on the right hand, and

. -the unmarked lateral organ belongs to the left one.

The lateral papille are oval and subdivided into two by fissures,

and the vallums and fosse are not very obvious.

OF THE TONGUES OF THE MAMMALIA, 27.

The mesial papille are small,the vallum of the anterior papilla

consists of two ecrescentic ridges, and the vallum of the posterior

papilla is large and fusiform.

The Fungiform Papille.

The fungiform papille have the usual disposition in rows and

clusters. They tuberculate the apex. There are few on the

lateral borders, but they are numerous and prominent on the

anterior part of the papillary zone of the inferior surface. They

, vary in prominence in different tongues, and the degree in which

they are concealed by conical papille also varies. There is a

prominent cluster immediately in front of the vallate papille, and

some invade the vallate Y and are hard to distinguish from

them.

The Conical Papille.

Most of the conical papille are of the filiform type, and have

the usual type of arrangement on the dorsum, lateral borders, and

inferior surface. Their points are directed backwards, or back-

wards and inwards, and they increase in size from before

backwards and without inwards. . They: conceal the anterior

fissures and ridges of the lateral organs. | '

At the sides of, and posterior to, the vallate papillary Y they

become large and oval or round, with the apices directed back-

wards and inwards, and the round ones have each a long central

spine.

The Lateral Organs.

Specimen 1 (text-fig. 8, R. & L.).: |

Right organ: Length °8 em. Ridges 5. Sulci 6.

Left organ: Length -8 em, Ridges 4. Sulci 5. -

Specimen 2 (text-fig. 8).

Right organ: Length 1:2 cm. Ridges 10, Svdler 11.

Left organ: Length 1:25em. Ridges 10. Suler 11.

The lateral organs differ in character in the two specimens.

In specimen 1 there are six or five sulci appearing as slits

running obliquely forwards and upwards, and the intervening

lamine are coarse, short, and not greatly raised above the level of

the surface of the tongue. In the second specimen the organs

have the appearance usual in all the other Gibbons. .

The following structures are absent :—

1. Lymphoid nodules with central pits.

2. Lytta.

3. Foramen cecum.

4, Plice fimbriate.

The sublingual fold is triangular in shape with a bifid apex.

From its upper surface the short Jrenum passes to the inferior

28 DR. C.F. SONNTAGOON: THE ANATOMY

surface of the tongne and Hoe ‘its: ander: surface to the floor of

the mouth. > —

The ventral mesial ends Tagine, anteriorly at the posterior

border of the papillary zone. It is narrow and deep, and lodges

a small median crest. ee 0

Text-figure 9,

Papilla of the-Gibbons,

1-13, conical papilla; 14-16, fungiform papille ; 19, vallate papillae of H. hoolock.

SUMMARY AND. CONCLUSIONS.

1. Apical notches and mesial dorsal sulci are more frequently

absent than present, and are of no particular value for compa

tive purposes.

2. The lateral borders have the same characters as regards

papille in allforms. In the Chimpanzee and Gibbons they lolge

the greater part of the lateral organs, but in the Gorilla and

Orang they only lodge a small -part.

3. The vallate papille are usually. fewer than in Man (7-12).

In the Orang they are in V-formation, but they usually assume

the Y-type in others. Compound papillee are common.

4, Filiform papillae are the predominating type of the conical

group on the anterior two-thirds of the dorsum.

5. The long conical papillee on the base are largest in the

Gorilla, smaller in the Chimpanzee, smallest in the Gibbons, and

absent in the Orang. :

6. Only the Orang and Siamang have smooth non-papillary.

areas on the base.

7. Plice fimbriatz are abceut fo the Gibbons.

8. Median ventral crests occur in the Gorilla and. some

Gibbons.

9. The Orang and Siamang have no triangular sublingual fold,

but all the other Simiz. possess one. In the Gorilla the apex is

entire, but in all the rest it is bifid. |

OF THE TONGUES OF THE MAMMALIA. 29

10. Only Man and the Orang have the Apical Gland of Nuhn,

11. The tongue of the Orang resembles that of Man more

closely than any other one does, and the tongue of the Bornean

Gibbon resembles, in many ways, those of the Cercopitheques,

which will be described in my next paper.

BIBLIOGRAPHY.

The numbers included in round brackets correspond with those

in Professor Arthur Keith’s “‘ Introduction to the Study of the

Anthropoid Apes” (Natural Science, vol. ix. 1896, Rep. 1897).

‘hose included in square brackets refer to the following papers

not mentioned by Professor Keith :—

1. Miincou.—Morphol. Arbeiten, Bd. 6, pp. 605-690. 1896.

2. Boutart & Pinurer.—Journ. de ]’Anat. et de la Physiologie,

Paris, 1885, p. 337.

[I was unable to see the characters of the part of the frenum

running between the inferior surface of the frenal lamella and

the floor of the mouth in the specimens in which I have not

specifically described this part. |

ON A RARE PARASITIC FLY. 31

2. Note on the Capture (in London) of a rare Parasitic Fly,

Hammomyia (Lylephila) unilineata Zett. By Lt.-Col.

S. Moncx'ron Copeman, F.R.S., M.D., F.R.C.P., F.Z.S.

| Received October 14, 1920: Read February 8, 1921. ]

(Text-figure 1.)

The exceptional rarity of this fly, of which I have had the

pleasure of presenting to the British Museum (Natural History)

the only specimens in our national collection, affords reason for a

record of its capture in considerable numbers in London (Prim-

rose Hill) during the past two years, for some discussion of its

nomenclature, and for a brief account of its seasonal prevalence

and habits so far as they are at present known.

In a previous communication I have set out some facts as to a

flourishing colony of a burrowing bee (Andrena fulva) on a

localized area on the south-western slopes of Primrose Hill,

which has been kept under observation over a period of six years,

during the latter half of which period the number of individual

burrows has, for reasons which were explained in my previous

paper, increased to a considerable extent. It was while watching

the operations of the bees on an outlying portion of this colony,

on May 16,1919, that, for the first time, I observed and obtained

specimens (two in number) of a fly which, from its actions,

which I studied carefully for some time, is apparently a parasite

of the burrowing bee.

On the morning (about 10 a.m.) of this day, which was bright,

without obvious breeze, I was lying motionless on the grass

watching the bees leaving and returning toa collection of burrows

on a small patch of bare earth, when my attention was attracted

by a homeward-bound bee, which, on approaching its burrow,

dived stright down into it, instead of, as usual, hovering around

for several seconds before doing so. As it approached, I noticed

that it was followed by a couple of flies, of a species unknown to

me, which appeared for the moment to be as astonished as I was

at its precipitate retreat into its burrow. After what looked like a

fight with one another for a few moments over the surface of the

ground, one of them followed the bee down into the hole, re-

appearing almost immediately, and then again descending into

the burrow, this time backwards. Meanwhile, the second fly

remained on guard outside. When the first fly at length emerged

head foremost, both of the flies were eventually trapped in a

match-box, a net not being available at the time. But unfor-

tunately one of them managed to escape before they could be

taken home and killed. The remaining specimen was despatched

at once to Major E. E. Austen, D.S.O., at the British Museum

(Natural History), with a request for its identification, as the

species was unknown to me. Under the circumstances, it may

32 LT.-COL. 8. MONCKTON COPEMAN ON

be of interest to quote from Major Austen’s reply, of May 17th,

1919, as follows :—

“The fly enclosed with your letter is a most interesting thing,

and I am more than sorry that I have been unable to name it

FOr YOU 0. 0% The species of the genus Wiltogramma (fam. 'Tachi-

nide) and its nearest allies behave exactly as you described, and on

more than one occasion I have watched one of these flies stalking

Text-figure 1.

Hammomyia (Hylephila) unilineata Zett., 2.

(The mark in the centre of the thorax is due to the fly having been

pinned before the photograph was taken.)

Magnified x10.

a solitary bee as a stoat does a rabbit—moving when the bee

moved, stopping when the bee stopped, and so on. Judge of my

surprise, therefore, when, on opening the box, I found an insect

such as, to the best of my belief, in nearly thirty years’ experience

of Diptera, I have never seen before! ‘The species undoubtedly

belongs to the Anthomyide—not to the Tachinide,—but is aber-

rant in more than one respect. It is not represented either

in our own collection or in the series presented by the late

A RARE PARASITIC FLY. 33

Mr. Verrall (the result of some fifty years’ collecting). My col-

league, who works at Diptera, dees not know it, and [ have spent

some hours.... trying to work it out with books ete., but without

success. Itis a most surprising thing, especially in view of the

locality. I hope that you will be able to secure more specimens,

and of both sexes.”

Mr. Austen, at first, was of the opinion that the sex, not only

of this particular fly, but also of each of a number of similar

specimens that I was subsequently able to send him, was male,

but when, later on, at the suggestion of Mr. Collin, who had

learnt of my find through Professor Poulton (to whorh I had given

some specimens for the Hope Collection at Oxford), one of the flies

was dissected, leading to the discovery of a perfect ege, it became

obvious that the flies that had been captured must be females.

And itis a curious and interesting fact that among about fifty

specimens of this fly which have been caught (all in the same

locality) up to the end of their seasonal prevalence in the early

part of June of the present year (1920) not a single individual

of the male sex has been secured. I learned, however, from Mr.

Collin that he now possesses three specimens of the male in-

sect—two taken by the Rev. A. H. Haton at Seaton (Devon) and

the third taken by himself at Long Sutton in Hampshire, all in

May 1919, curiously enough the same year and month in which

the first specimens (females) were found by myself on Primrose

Hill. My. Collin further informed me that he possesses female

specimens from the following localities :—‘* Newmarket, Shoebury-

ness, and Dunsford (all June captures); Cuckmere (Sussex)

taken in May; and a spetimen taken by Col. Yerbury at

Charlton in April.

The difficulty as to the sex of my first captures arose from the

masculine character of the approximation of the eyes, which is the

cause of the very narrow “ frons,” which, however, appears to be

peculiar to, and distinctive of, the female in this group of An-

thomyids—a feature which is well shown in the photograph, and

which, so far as | am aware, has not previously been illustrated.

NoMENCLATURE.—As stated above, Major Austen and _ his col-

league in the Diptera department of the British Museum (Natural

History) were unable, at first, to identify this fly, but on calling

there on May 28th, 1919, with somefurther captures, Major Austen

informed me that Dr. P. H. Grimshaw of the Royal Scottish

Museum, who had recently paid him a visit, had stated, when the

specimens were shown to him, that he recognised the fly, iden-

tifying it as Chortophila buccata Kallén, although the species was

not (as Major Austen understood him) included in the Scottish

collection.

Shortly afterwards Professor Poulton arranged with Major

Austen that Mr. Collin should be afforded opportunity of

examining the specimens I had presented to the Museum,

for the reason that, as Mr. Collin informed me, his uncle, the

late Mr. Verrall, had suggested that a fly, apparently identical

Proc. Zoou. Soc.—1920, No. III, 3

34 ON A RARE PARASITIC FLY.

with mine, should be known as Hammomyia unilineata Zett.

rather than WZ. (Chortophila) buccata Fallén.

Subsequently Mr. Collin wrote to me that he possessed several

continental specimens of H. buccata Fln., including a pair so

named by Zetterstedt himself, and that he was of opinion that

these were abundantly distinct from the British species, which is

undoubtedly Hammomyia (Hylephila) unilineata Zett. Further-

more, he kindly directed attention to the fact that in describing

unilineata Zetterstedt wrote ‘ Feminee ad nidis Andraenarum

invente, in quarum larvis larvee hujus speciei Ariciz verisimiliter

parasite vivant observante D. Professore Wahlberg.” Appa-

rently, therefore, buccata Fallén requires confirmation so far

as our British fauna is concerned, as Mr. Collin is convinced that

he has never seen a British specimen of the true buccata.

SEASONAL PrEVALENCE.—AS stated above, my first specimens of

this fly were captured on Primrose Hill on 15th May, 1919. On

the same cireumscribed area of ground I caught ten more speci-

mens on May 19th, ten on May 20th, and seven on May 21st.

Then, after an absence from town of several days, a further seven

flies were caught on May 27th and three on May 29th—at

which date I had again to leave home for about a fortnight.

Just before my return the fine weather gave place to rain,

accompanied by a considerable fall in temperature, with the

result that, subsequently, neither bees nor flies were to be found,

notwithstanding careful search on several successive days.

It may be mentioned that, in 1919, Andrena bees were first

seen on May 6th, a warm and sunny day, when a number of

males, which always emerge before the females, were flying

about; while females were not found, with the exception of one

or two solitary specimens, until several days later. So that, as

might be expected in view of what is known as to the parasitism

of the fly in question, it is obvious that the seasonal prevalence

of the female of both fly and bee tallies closely.

In consequence, doubtless, of the abnormal meteorological con-

ditions during the present year (1920), both bees and flies appeared

at an earlier date than in 1919, the first flies, three in number,

having been obtained on May Ilth. Careful search was made

for male specimens of the fly, but again without success. By

the end of May both bees and flies had disappeared.

It will be noted that, with some exceptions, the specimens of

the fly in Mr. Collin’s collection were captured in the month of

May—the month during the latter half of which, as my obser-

vations on Primrose Hill, extending over a period of several

years, have shown, the female of the bee Andrena fulva 1s

more particularly prevalent.

ON THE CLASSIFICATION OF THE ‘THERIODONTIA, =)

3. The Bases of Classification of the Theriodontia.

By D. M. 8. Watson, F.Z.S., University College, London.

[Received October 19, 1920: Read February 8, 1921.]

(Text-figures 1-29.)

Among the first series of reptiles from the Karroo system

of South Africa sent home by Andrew Geddes Bain were a few

poor fragments of animals with a heterodont carnivorous denti-

tion. Later collections from the same rocks included more

satisfactory remains of these animals, which were described by

Owen, who recognised their mammalian appearance and despite

his ante- evolutionsur y views even suggested that they were mam-

malian ancestors. Prof. Seeley’s visit to South Africa marked

a turning-point in our knowledge of these reptiles, because he

showed that their remains were found in rocks of widely different

ages, and that the latest assemblage—Diademodon, Cynognathus,

and 7irachodon—were more mammal-like in their dentition than

were their earlier forerunners. He showed also that they pos-

sessed a mammal-like secondary palate, but failed to arrive at a

satisfactory interpretation of that region in the less complete

remains of the earlier forms known to him. Neither Owen,

Seeley, nor Lydekker was able to draw up any useful classification

of these reptiles on account of the paucity of material, and the

first definite step in so doing was made by Broom, when in 1904

he showed that Scylacosaurus sclateri, a form from the lowest

zone of the Beaufort beds, differed from the ‘‘Cynodonts” of

the highest zone of that formation in lacking any trace of a

secondary palate.

Subsequent work by Broom added many new generic types to

those included with Scylacosawrus in that primitive division of

the carnivorous Therapsids whose members lacked a secondary

palate and had uncusped molar teeth. This division Broom

made into an order and called Therocephalia.

No further important additions were made to our knowledge

of the skull of any of these reptiles till, in 1911, the writer gave

a very detailed account of the skull of Diademodon and Broom

a more general description of the skull in all the Cynodonts.

The first important addition to our knowledge of the earlier

Theriodonts was the description by the present author of the

posterior half of a skull from the Cistecephalus-zone, which

agreed with Gorgonops in having a broad parietal region, the

parietal bone being excluded from the margin of the temporal

fossa. In the same paper some of the more salient features of

the palate of Gorgonops were described, and it was indicated

that the form showed the beginnings of the Cynodont secondary

palate, the skull known as Arctognathus curvimola showing an

intermediate condition. Whilst I was writing this paper in

36 MR. D. M. S. WATSON ON THE

London, Broom in Seuth Africa was describing two very com-

plete Gorgonopsid skulls, one associated with the anterior part

of a skeleton. Of these skulls Broom gave a good description,

bringing out the whole structure of the face and parietal region,

but not giving us so satisfactory an account of the palate and

occiput. Broom, sometimes in conjunction with Haughton, sub-

sequently added many new and often strange forms to the Gor-

gonopsidee—on the whole, emphasizing rather their resemblance

to the Deinocephalia and Dicynodontia and even Pelycosauria

than those which they show to the “Cynodontia.” In 1914 I

was able to show that known Gorgonopsids could be arranged

as a morphological series giving a gradual passage in the

structures of the occiput, and of the basicranial and otic regions

between Dimetrodon a Pelycosaur and Diademodon a “*Cynodont.”

In the same paper I described the palate of the ‘ Cynodont”

Bauria, showing that it differed much from the Cynognathids

and resembled the non-Gorgonopsid Theriodonts with a primi-

tive palate more than the Gorgonopsids. In consequence, purely

as a temporary measure, | revived the order Theriodontia and

divided it into four sub-orders—the Therocephalia, the Gor-

gonopsia, the Bauride, and the Cynodontia. Since that paper

was written, Haughton has published descriptions of certain new

forms and made important new additions to our knowledge of

the brain-case of the earlier Theriodonts. In his most recent

paper he uses provisionally my 1914 classification, emphasizing

its insufficiency.

In revising a paper on the relative ages of the Paleozoic and

Triassic reptile- bearing rocks, which has occupied me at intervals

for some years, 1 was forced to deal with the problems presented

by the fauna of the copper-bearing Permian sandstones of the

Orenburg district of the Urals. One of the most noteworthy

forms from this locality is Rhopalodon, an animal whose skull,

which alone is certainly known, presents many resemblances to

the Gorgonopsids. The necessity of discussing the systematic

position of this form led me to an examination of all the

Theriodont material available, with the results which are set out

below.

It is convenient to begin with a description of the material at

my disposal, then to discuss the morphological results which

arise from it, and, finally, consider the evolution of the group

and the relationship of Deuterosaurus to it.

ARCTOPS WILLISYONI Watson, Proc. Zool. Soc. 1914, p. 1026.

Type and only known material: a skull lacking the anterior

part of the snout, the quadrates and quadrate rami of the

pterygoids, otherwise complete and practically undistorted.

From Howse Post, near Fort Beaufort, 8. Africa, not improbably

E,NDOTHIODON zone.

I deseribed and figured the oceiput and basicranial region in

the oviginal description.

CLASSIFICATION OF THE THERIODONTIA. 37

Arctops has a depressed and very massive skull, the snout,

when broken off about 5 em. in advance of the orbit, being

rectilinear in section, bounded by a straight dorsal surface which

passes somewhat abruptly into straight, nearly vertical, lateral

surfaces. The orbit is small, placed high in the skull, and nearly

laterally directed; its upper margin is continued forward by a

ridge separating the dorsal surface from the lachrymal region,

which is excavated into a shallow depression. ‘I'he interorbital

region is very wide, forming a flat surface bounded laterally by

shallow bays over the orbits.

Text-figure 1.

5a. Tae, |Par. BOc. ExOc. Au.Ga.

Arctops willistoni Watson. ‘Type-skull.

Dorsal aspect. 2.

B.Oc., basioccipital; I.Par., interparietal ; Sq., squamosal ; Tas., tabular.

The parietal region, also flat, is even wider than the interorbital

surface ; ib separates the very small temporal fossxe which face

more largely laterally than dorsally. The occipital surface is

very wide and is separated from the parietal by a sharp corner.

The squamosal is small, consisting mainly of a vertically standing

plate passing directly outward from the end of the massive

paroccipital process. The posterior surface of the bone at this

articulation is produced backwards into a ridge which forms the

38 MR. D. M. S. WATSON ON THE

inner wall of the auditory groove. Sutures over the outer

surface are not clearly recognisable.

I have already described the basieranial region, but would

again call attention to the flat, laterally directed, plate-like basi-

pterygoid processes. The narrow ridged girder formed by the

parasphenoid and pterygoids extends forward to the front of the

orbit, where it suddenly passes into the wide posterior end of the

palate. The pterygoids pass outward to form thick downwardly

directed flanges, not very deep when compared with later forms,

but of great antero-posterior extent. The middle region of the

posterior part of the palate forms a slightly raised area separating

two concavities, whose surface lies mainly on the ectopterygoids,

large square bones which only take a small part in the great

flanges. The middle part of the palate forms a shallow groove

Text-figure 2.

Arctops willistoni Watson. Type-skull.

Right lateral aspect. x2.

B.Sp., basisphenoid; Pr., pterygoid.

beginning at the extreme posterior end and running forwards to

the posterior nares. At about the level of the anterior end of

the ectopterygoid, this groove is overhung by a pair of small

processes rising from the ptervgoids. Further forward the floor

of this groove is cut into by the narrow slits which represent the

posterior ends of the posterior nares, These are separated by

a narrow bar of considerable vertical depth. The structure of

this bar is not quite certainly determinable, but on the curved

fracture which forms its present front termination it is certain

that its upper surface consists of a pair of ridges separated

by a parallel-sided cleft not more than a millimetre wide and

nearly a centimetre deep. From this slit a suture seems to be

continued on to the palate. Further back two lateral ridges are

CLASSIFICATION OF THE THERIODONTYA, 39

added to the original pair, the gaps between them being also

apparently continued as sutures on to the palate. ‘Thus the

posterior part of the bar separating the internal nares seems to

be built up of two pairs of bones, of which the outer terminates

not far in front of the posterior ends of the nares. It is probable

that the inner pair are prevomers and the outer the anterior ends

of the pterygoids, which hence form a part of the border of the

posterior nares.

Text-figure 3.

Arctops willistoni Watson. 'Type-skull.

Palatal aspect. X $.

Ee.Pr., ectopterygoid; Par.Oc., paroccipital ; Pr., anterior end of pterygoid;

P.V. ?, prevomer, posterior end of internarial bar. Parts in broken lines

restored without evidence.

GorGoNnops ToRVUS Owen, Cat. 8. Afr. Rept. 1876.

Type: a skull with the zygomatic arches broken away, the

basis cranii only represented by a fractured surface passing

horizontally through the basisphenoid. The paroccipital pro-

cesses only represented by the impression on the matrix of the

anterior face of that of the right side, the posterior part of the

palate represented only by the impression of its dorsal surface.

40 MR. D. M. 8. WATSON ON THE

Otherwise the skull is complete and, on the whole, extremely well

preserved. It is represented by beautiful and most accurate

lithographic drawings in Owen’s Catalogue. From Mildenhals,

Fort Beaufort. Another more complete, but less well-preserved,

skull from the Hndothiodon-zone of Beaufort West is in the

American Museum of Natural History.

Text-figure 4.

Ban Peis lee rae neds

Gorgonops torvus Owen. ‘Type-skull.

Dorsal aspect. Xz. Parts in broken line restored without evidence.

Fr., frontal; I.Par., interparietal; Pr.FR., prefrontal; P.O., postorbital ;

Pr.FR., postfrontal; S.Mx., septomanxilla; S.Oc., supraoccipital.

The skull of Gorgonops has a flat dorsal surface, which passes

through a chamfered corner into the nearly vertical sides of the

snout. The orbit is large, directed almost entirely laterally and

of considerable depth. Immediately in front of it the snout is

CLASSIFICATION OF THE THERIODONTIA. 4]

nearly square in section, the lachrymal region being excavated

into shallow hollows prenhane by a thickening of ine prefrontal,

which forms the lateral border of the flat dorsal surface. Fur ther

forward the snout becomes deeper, the nearly flat lateral surface

passing by a rounded corner into the dorsal surface. There is a

long broad swelling on the maxilla over the root of the canine.

The interorbital and parietal regions are both very broad, the

latter passing smoothly into the broad occipital surface. The

part of the occiput preserved consists mainly of the very broad

interparietal, whose sutures with the tabulars are shown. ‘The

supraoceipital has only a very shallow exposure below the inter-

parietal. A peculiar feature of this skull is the irregular shape

of the pineal foramen and the fact that that opening is raised

on a little column standing up above the general level of the

parietal region.

Text-figure 5.

al) Fe Fr. Na. SMx.

x SSS

pore —— —

if it Kj / | ]

pe ah oe oy

er Wrens ali: ; PMx.

Gorgonops torvus Owen. Type-specimen.

Right lateral aspect. 3%. Parts in broken lines restored without evidence.

Ju., jugal; Lac., lachrymal; Mx., manxilla; P.Mx., premavxilla.

The general structure of the dorsal and lateral surfaces will be

best understood from text-figs. 4&5, The cruciform shape of

the pair of frontals is noticeable.

The structure of the external nostril is very well shown in the

specimen. The dentigerous part of the premaxilla is deep, and

articulates directly with the anterior end of the maxilla, which

overlaps on to it. The dorsal surface of the two bones is the

lower margin of the nostril and forms the emplacement of the

septomaxilla. Behind the nostril the maxilla rises to a long

suture with the facial part of the septomaxilla; behind this bone

it reaches the nasal. The nasals forma slightly coved roof to the

olfactory chamber and reach forward almost to the end of the

nose, where they terminate in a nearly straight margin, from the

middle of which arises the narrow process which articulates with

42, MR. D. M. S. WATSON ON THE

the facial processes of the premaxilla. The facial part of the septo-

maxilla articulates with the lower margin of the nasal, but the

two bones separate before the nasal terminates, so as to leave

that bone overhanging the nostril like the eaves of a roof. After

its separation from the nasal the septomaxilla passes downwards

and gives off a process from its anterior border, which passes

inwards towards the middle line, following the curve of the

Text-figure 6.

Gorgonops torvus Owen. ‘Type-specimen.

Palatal aspect. 32. Unshaded areas surrounded by continuous lines present

but mutilated, broken lines parts restored without evidence.

P.V.?, internarial bar, ? prevomers ; Pat., palatine; Vo., ‘ vomer.”

anterior border of the nasal. The lower part of the septo-

maxilla is a rounded column, swelling out to a base which rests

on the premaxilla.

T have already (1912) given an account of the general features

of the palate of Gorgonops, but, as further study of the specimen

in the light of other material has enabled me to make out some

CLASSIFICATION OF THE THERIODONTIA 43

interesting features not previously recorded, I give here a more

detailed description. The premaxilla has a narrow dentigerous

surface, with five sockets for the roots of the incisors; above the

border the bone thickens, forming a deep wall, from whose ad-

median half the palatine process arises; this is at first a flat

expansion, but soon becomes a rounded, backwardly directed

tubercle, separated from its fellow and resting in a groove on the

lower surface of the “‘ prevomer.” The internarial bar is a narrow

rod which anteriorly is comparatively wide. Anteriorly its

palatal surface bears a median ridge which separates two channels

bounded by other lower ridges, which form the lateral borders of

the bone. Further back the lower surface of the bar becomes

flat, and the whole of the posterier part is only represented by a

broken surface, which probably originally supported a deep

median ridge.

The internal nares are very large openings bounded by the

premaxilla in front, where they are very wide, and contracted

posteriorly by the thickening of the maxille necessitated by

the large sockets for the canines. The maxille form their

outer borders for some distance and are then excluded by the

palatines. Finally, the posterior border is formed by the semi-

circular margin of a bone whose nature has to be discussed.

Between the internal nares and the pterygo-parasphenoidal bar

the palate forms a large area of complicated shape. The height

above the lower margin of the premaxilla at which the palatal

processes start, and the deep step in the lower border of the

maxilla just in front of the canine make the ventral surface of

the internarial bar lie much dorsal to the lower edges of the

maxille in the cheek-region. Thus at the back of the nares the

palate is very much vaulted. Behind the canine the palatal

exposure of the maxilla, which bears no trace of cheek-teeth, is

very broad and its admesial surface forms a deep vertical plate.

This surface when followed caudally passes into a similar face

carried by the palatine, which stands almost vertically, tightly

attached to the maxilla by an obvious and deeply interdigitated

suture, and with its lower edge forming with that bone a broad flat

face in the area where cheek-teeth would naturally be expected.

These teeth must have been functionally replaced by a hard gum,

possibly cornified so as to form a crushing plate.

The wide groove formed by the palate at the posterior end of

the internal neres is rapidly divided into three, each groove of the

lateral pair is deep and narrow and cylindrical; it shallows

rapidly when traced backwards, finally becoming flat when it

reaches the ectopterygoid. The bottom of the lateral groove

has a suture running the whole of its length, which is completely

exposed on the right side, but concealed by matrix except for its

anterior end on the left side of the type-skull. This suture,

which seems to be truly a suture and nota crack, unites the pala-

tine with the pterygoid, which bone hence forms the posterior

margin of the posterior nares,

44. MR. D. M. 8. WATSON ON THE

The middle groove of this part of the palate becomes narrower

as it is traced backward, but remains deep. Its hinder end is

separated from the lateral grooves by triangular raised areas,

which are roughened, but seem not to bear teeth.

The internarial bar is continued back into this part of the

palate as a narrow slip separated from the pterygoids by a pair of

open and very obvious sutures. These sutures rapidly approach

and fuse, being continued backwards by an obvious median suture

for about 5mm. This open suture, with a visible strip of matrix

in it, then suddenly ends and is with certainty not continued

backward in the middle line. It is, however, apparently replaced

by a pair of much less obvious sutures, between an overlapping

median bone and the pterygoids, which pass outward to the

margins of the median groove and seem then to be continued

backward by still less obvious sutures running along these

borders. Further back the wide, essentially flat palate gives

origin to the descending flanges. The ectopterygoids are separated

by obvious sutures and ave comparatively small bones not taking

any large part in the flange.

The palate of Gorgonops thus seems to show large pterygoids

reaching forward to the posterior nares and widely separating the

palatines, which are small bones simply continuing the ectoptery-

goids forward. In that part of the palate which lies in front of

the transverse flanges the pterygoids do meet each other for a

very small distance in the middle of their length, but posteriorly

are separated by a median vomer and anter iorly by the posterior

end of the internarial bar which is clasped between their distal

ends. There is no evidence to show whether or not these two

median bones are really separated, but as the anterior passes

dorsal to the pterygoids, whilst the other overlaps their ventral

surface, there is great probability that they do not represent parts

of the same element?

SCYMNOGNATHUS WHAITSI Broom, Proc. Zool. Soc. 1912, p. 861.

Type: a figured skull, nearly complete, but considerably

erushed and showing little of the structure. Other imperfect

skulls and other bones.

The individual of which, under the name of Scymnognathus

whaitsi, 1 described the lower jaw (1912) and the brain-case and

occiput (1914), does not belong to this species, and is deseribed in

this paper as a new genus and species. There are in the British

Museum three specimens of JS. whaitst:—R. 4053 collected by the

Rev. J. H. Whaits, as a very large number of small fragments

which, fitted together, form a skull from the front of the orbits

backwards with the pro-atlas and atlas in position, the anterior

end of the snout and a mass of separate fragments representing

the major part of the face; of these a small bit of the posterior

part of the palate is of great morphological interest. The

back of the skull built up from these remains is quite un-

distorted and has been very completely developed, now showing

CLASSIFICATION OF THE THERIODONTIA. 45

the whole lateral surface of the brain-case, the structure of the

zygomatic arches and roof of the skull and occiput with great

perfection. It is in many ways the best Gorgonopsid skull

known.

R, 4052. A skull retaining a well-preserved and only slightly

sheared snout, with a much crushed posterior part, permittin;

the definite identification of R. 4053.

49369. A snout, somewhat distorted and not very well pre-

served which has been cut into slabs. It agrees well with the

corresponding part of R. 4052.

All the material of Seymnognathus whaitsi comes from the

Endothiodon-zone of Beaufort West. The skull, as a whole, is

remarkable for the marked distinction between the relatively

narrow snout and palate and the wide postorbital region. Owing

to this shape, the orbits look as much forward as outward. The

temporal fossee are very large and face more upwards than out-

wards. ‘The parietal region is, in consequence, narrow and the

occiput deeply cupped, owing to the backward swing of the

squamosals from their union with the postorbitals.

The snout is much more rounded than in Gorgonops or Arctops,

although towards the orbits it is still somewhat “ square-cut.”

The external nares closely resemble those of Gorgonops, and there

is the same step between the lower edges of the premaxilla and

maxulla.

The structure of the dorsal and lateral surfaces of the skull

are obvious from text-figs. 7 & 8, but it is necessary to give some

account of the brain-case and palate.

The basioccipital is fused with the exoccipital and paroccipital,

and its suture with the basisphenoid has been destroyed by

a fracture. It isa long narrow bone, terminating behind in a

single condyle, which is probably partly exoccipital. This condvle,

as seen in section, 1s much wider than it is high, the dorsal sur-

face being excavated by the lower part of the foramen magnum.

The posterior part of the basioccipital is thus thin.

The exoccipitals are of the ordinary Gorgonopsid or Pelycosaur

pattern, but thei upper surfaces are concealed by the overlapping

elements of the proatlas. Further forward the lower surface of

the basioccipital and of the paroccipital fused with it project

down as a short, powerful, obscurely bilobed process, whose outer

part supports the fenestra ovalis; with this process the powerful

tuber basisphenoidalis articulates dorsally, though ventrally the

two projections are separated by a gap.

The paroccipital and pro-otic are fused, not only with each

other, but also with the basioccipital; the suture between the

pro-otic and the basisphenoid remains open.

The paroccipital process is extremely massive, passing out

from the side of the basioccipital on the lower surface of the skull

to its broad abutment on the squamosal. ‘he anterior and lower

faces of this process are excavated by a groove which leads inward

to the large irregular opening, which is the fenestra ovalis. The

co)

46 MR. D. M. S. WATSON ON THE

Text-figure 7.

Scymnognathus whaitsi Broom. Back from R. 4053. Snout from R. 4052.

B.M.N.H. Dorsal aspect. x4.

At., atlantal neural arch; Ju., Jugal; P.Par., preparietal ; Pr.At., pro-atlas.

CLASSIFICATION OF THE THERIODONTIA. A7

paroccipital process is bounded above by the small oval post-

temporal fossa, which lies at the level of the floor of the foramen

magnum.

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In the pro-otic on its front face, above, and in front of the

fenestra lies the outer opening of the foramen for the facial

nerve. This opens downwards and has below it a little hollow

for the geniculate ganglion. iis

Immediately above and a little in front of the facial is another

much larger foramen opening directly forward ; its outer margin

48 MR. D. M. S. WATSON ON THE

is carried by a spout-like projection from the pro-otic, its inner

border 1s basisphenoid. That bone immediately in advance of

the foramen has a deep depressed groove. There can be no

doubt that this foramen is for the fifth nerve, the cavity before

it having housed the Gasserian ganglion. Above and in front of

this foramen the pro-otic is continued forward, having a suture

with the basisphenoid, until its anterior margin or that of the in-

distinguishably fused supraoccipital is cut into by a notch, which

is very nearly converted into a foramen by the basisphenoid.

This foramen must be venous; it is in part the homologue of one

which is almost constantly represented in Therapsids.

The supraoccipital is as always spread out into a wide plate,

but from the anterior part of this expansion a special thickening

Text- figure 9.

Far. | Par Tap.

Oil. Rarer Bee

Scymnognathus whaitsi Broom. KR, 4053. B.M.N.H.

Occipital aspect of skull, with the anterior ends of the proatlas attached. x 5.

Par., parietal ; Qu., quadrate; T.B.Sp., tuber basi-sphenoidalis.

is carried forward, forming the roof and part of the side-wall of

the brain-case. It is this thickening whose margin forms the

dorsal border of the venous notch. With the sides of the upper

part of the supraoccipital in the region of this thickening the

interparietal articulates, stretching far forward in contact with

the parietal above and the supraoecipital below, and widely

exposed in the outside view of the brain-case.

The basisphenoid is a remarkable bone, which in the speci-

men is broken off in front. As far as it is preserved, how-

ever, it consists of « body which is articulated with the front of

the basioccipital largely through the intermediary of the two

SIFICATION OF THE THERIODONTIA.

CLA

inassive downward projections, which are its tubera.

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articulation the bone becomes narrower where it is attached to the

Its lateral face here bears the groove for the Gasserian

ganglion, above which the bone again widens to the continuation

pro-otie.

Proc, Zoou. Soc.—1921, No, IV.

50 MR. D. M. 8S. WATSON ON THE

of its suture with the pro-otic. Above and in front of the

termination of this suture the basisphenoid is still continued

upward as a slender process, whose upper margin is the lower

border of the great venous notch and whose lower margin meets

its fellow in an open median suture below the brain, immediately

in advance of the hypophysis. The rostral part of the basi-

sphenoid is a vertically placed plate arising from the body of the

bone and separated from the upper parts, just described, by a

deep narrow notch, the open side of the pituitary fossa, From

its sides arises the thick flat expansions, which are the basi-

pterygoid processes. These incline downwards at the back at an.

angle of about 45°, and whilst their dorsal surface is sharply

separated from the vertical face of the medial lamina, which lies

above them, their ventral faces pass smoothly down to form a

blunt ridge on the lower surface of this part of the basisphenoid.

The parietal is composed of a plate of bone lying on the roof of

the skull with an almost plane dorsal surface. Its postere-lateral

corner is drawn out into a long process, which passes backwards

to touch the extreme tip of the squamosal. The posterior edge

of the whole bone is in contact with the interparietal towards the

middle line and with the tabular laterally. By far the greater

part of the outer margin of the parietal is in contact with the

postorbital, which completely excludes it from participation in

the margin of the temporal fossa. From the lower surface of the

parietal a powerful ridge is developed, which marks the side-wall

of the brain-case. Posteriorly this ridge just touches the anterior

end of the supraoccipital. Immediately in front of this bone it

has a suture with the epipterygoid ; further forward its lower

edge is free, but gradually declines, until at or about the front

end of the parietal it vanishes. The lower surfaces of the pre-

parietal and frontals form the roof of the brain-case in this

region, and the lower surface of the anterior part of the brain

is supported by an ethmoid ossification. This is a thin hem1-

cylindrical shell of bone with a rib along its ventral surface in the

middle, which indicates that it rested on a deep median septum

now broken away and lost.

The posterior end of the ethmoidal cavity is widely open. The

opening of the anterior end is much contracted and les clese up

to the skull-roof.

The floor of the cavity close to its anterior end is perforated

by a pair of large oval foramina, which face downward. These

are separated only by a narrow septum and must be for the optic

nerves, which hence had a remarkably long intracranial course.

The epipterygoid is only represented by its upper end, which,

though narrow antero-posteriorly, is thin. It has a suture with

the parietal and with the front end of the supraoccipital, the

latter connection being of considerable morphogenetic importance.

There is a medium-sized foramen for the Xth nerve, opening

downwards and backwards below the exoccipital well above the

CLASSIFICATION OF THE THERIODONTIA, 51

Text-figure 11.

alll

: Face

Scymnognathus whaitsi Broom.

Posterior part from R. 4053. Snout reconstructed from a series of transverse sections

of 49369 completed from R. 4052. Ectopterygoid region + detached area

including the vomer R. 4053. #4. B.Pr., basipterygoid process.

52. MR. D. M. S. WATSON ON THE

bottom of the skull. There isa single hypoglossal foramen in

the usual position.

Taken as a whole, it is obvious that the brain-cavity was very

small in relation to the skull, and especially that the cerebral

hemispheres were still of much less bulk than the cerebellum.

At the same time the very great proportional length of that part

of the brain which lies in front of the fifth nerve foreshadows

the great cerebral development which occurred in later allied

forms.

The palate of Seymnognathus whaitsi is still not known as a

whole, but the anterior part is very well shown in the sections

of 49369, from a reconstruction made from which text-fig. 10 is

mainly drawn, and in the solidin R, 4052. The pterygoidal flange

and one transverse bone are preserved in position in R. 40538, and

that individual retains a small fragment from the middle of the

palate just in front of the anterior end of the long pterygo-

parasphenoid bar. This fragment shows a pair of much raised

ridges, which lie on the pterygoids and diverge outwards as they

are traced forward. These are covered with a shagreen of small

teeth. Between these the palate is deeply grooved ; lateral to

them it is depressed into deep hollows. The dorsal surface

of the fragment bears a deep median keel. This fragment has

been cut across by a tranverse cut, so that it now shows three

sections. That at the back shows that the keel is formed by a

single bone whose lower edge is received into a groove on the

upper surface of the fused pterygoids, which meet below it.

In the middle section this median bone has a deeply grooved

lower edge, the two thin ridges which form the side-walls of this

groove being received in slits in the pterygoids. These latter

bones meet in a median suture on the palate and here bear the

massive tooth-bearing ridges. On the front section the median

bone is exposed on the palate, forming the roof of the median

groove and separating the pterygoids. ‘The median bone thus

corresponds exactly in position and relations with the posterior

median bone in Gorgonops and the back of the vomer in Diade-

modon. The anterior part of the palate resembles that of Gor-

gonops in the relation of the internarial bar to the palatine

process of the premaxillse and in its shape.

Near its anterior end the internarial bar is a single bone

with a convex dorsal surface from which a ridge rises. This ridge,

which is detached, apparently by fracture, extends upwards and

backwards, obviously representing an ossification in the nasal

septum. The lower surface has a low median ridge separating

two well-defined grooves. As this bone is traced backward it

gradually becomes narrower from side to side until in the region

of the first molar tooth, where it is seen in section (text-fig. 12). it

has become converted into a plate 35 mm. in depth and only two

millimetres thick at the lower edge, where it is widest. The

dorsal centimetre of this narrow septum is clasped between two

CLASSIFICATION OF THE THERIODONTTIA. 53

thin films of bone, whose outer surfaces ave in contact with

another pair of similar slender processes. Hven in this region

the lower edge of the median bone still les considerably dorsal of

the tooth-bearing margin of the maxilla.

In the next section, about 1 cm. further back, the median

plate is shallower, its dorsal margin being curved downwards.

The two pairs of plates which support its upper edge are thicker,

but still retain their same relations.

The next slab has fortunately been split longitudinally and

somewhat developed, so that it gives conclusive evidence that

the lateral pair of processes described above are part of the

Yext-figure 12.

Scymnognathus whaitsi Broom.

Series of transverse sections at about 1 cm. interval, across the snout of No. 49369.

B.M.N.H. 1, anterior section; I.N.B., internarial bar; in 2-5 only the

internarial bar is represented; in 6 the maxillz and palatines are shown ;

in 7 only the anterior ends of the palatines and pterygoids. 5.

palatines. The inner pair pass down to the ventral surface and

there form a little boss on the palate, which separates the two

deep grooves on the palatines. These grooves are so overhung by

the more ventral parts of the palatines that their floor can

scarcely be seen in a direct ventral projection No sutures can

54 MR. D. M. S. WATSON ON THE

be seen in the little median boss, and it is probable that the

median element of the internarial bar has terminated in it.

The vertically standing part of the palate, which forms the

side-wall of the groove just described, descends to the level of the

lower border of the maxilla, where it passes into a flat, thick,

horizontally lying plate, which extends outward to the lower edge

of the maxilla with which it has a suture.

This horizontal part of the palatine forms the greater part of

that bone, and extends backward and forward in contact with the

maxilla, until by narrowing and increasing in depth it becomes

converted into a mainly vertically disposed plate, which forms

part of the side-wall of the posterior part of the very large

internal nares. It then terminates.

Text-figure 13.

HOUERUE Ve aie Wy

WLU DSRS

tae TTT \ Th tee r

AY AINA Mh | i

FINVE) amulet dee,

Scymnognathus whaitsi Broom.

Reconstruction of internal aspect of the left side of the snout cut through in the

middle line. Internarial bar and ossification in the nasal septum unshaded

and surrounded by a thick continuous line; anterior end of the pterygoid

represented by a line of small crosses. Parts of palatine seen through other

bones in broken line. Reconstructed from the sections of 49369, checked

by R. 4052. & 2.

Thus the anterior part of the palate is essentially a flat plate

of bone, whose middle part is cut out by a narrow but gradually

widening groove which plunges steeply downward to the deeply

sunk posterior margin of the posterior nares. This groove is

divided into two by a narrow vertical septum, which descends

nearly to the level of the general plane of the palate.

CLASSIFICATION OF THE THERIODONYTTIA. 55

There can be little doubt, especially when the conditions in

Arctops and Gorgonops are considered, that the inner pair of pro-

cesses which support the imternarial bar are the anterior ends

of the pterygoids. It remains to be shown by other materiai

whether the median internarial bone and the median vomer in

the back of the palate are parts of the same bone or are, as is

more probable, separated.

The strange way in which the median internarial bar rises as

a thin but very deep septum from the much sunk posterior nares,

nearly to the general level of the palate, seems to be only

explicable if its ventral border supported the middle of a small

soft secondary palate stretched between the maxille and the

palatines.

I have already described the mode of articulation of the

squamosal with the brain-case and with the fused quadrate and

quadrato-jugal in Proc. Zool. Soc. 1914, p. 1034, fig. 6.

The squamosal above the level of the post-temporal fossa bows

out backward, so as largely to increase the size of the dorsal

opening of the temporal fossa. It thus makes the occiput very

deeply cupped, the interparietal region being narrowed and the

outer part of the tabular running nearly antero-posteriorly.

At the extreme postero-lateral corner of the skull, the

squamosal turns sharply into a process passing forward and

Inward in the zygomatic arch. This process is clasped by other

bones both admesially and externally. One of these bones is

the jugal. The other conceivably also jugal, but much more

probably postorbital. A gap about 2 cm. long in both sides of

the specimen prevents a definite decision on this point.

The squamosal at the corner is made of a very peculiar,

extremely dense, though finely cancellous bone. This structure

is found in this region in all Theriodonts I have examined,

LEPTOTRACHELUS EUPACHYGNATHUS, gen. et sp. nov.

Type: a skull and lower jaw, described in error as Seymno-

gnathus whaitsi by the writer (Ann. & Mag. Nat. Hist. ser. 8,

vol. x. p. 578, fig. 3, and Proc. Zool. Soc. 1914, pp. 1027, 1032,

figs. 3, 4, & 5).

The material is a largely disarticulated skull varying in

preservation, with one complete and one pavtially disarticulated

ramus of the lower jaw. ‘The skull is represented by the brain-

case, interorbital region, left nasal, lachrymal, prefrontal, jugal,

and squamosal in natural articulation, the right jugal, lachrymal,

and prefrontal in natural articulation, but separated from the

skull, an isolated maxilla, and quadrate and quadrato-jugal.

The mode of articulation of the quadrate with the squamosal is

clear, and with the perfect lower jaw gives the length of the skull

and the position of the maxilla. The large articulated part of

the skull gives practically all the dorsal and the posterior

part of the lateral surface directly, The occiput is essentially

56 MR. D. M. 8S. WATSON ON THE

completely preserved. All the sutures except those of the

pavietals with each other and the preparietal are well shown.

In text-fig. 14 it is probable that the anterior part of the snout

is made a little too narrow.

I have already described and figured the basicranial and otic

regions, the occiput, and the interior of the brain-case.

Text-figure 14,

Leptotrachelus eupachygnathus, gen. et sp.nov. Type-skull.

Dorsal aspect. X §.

The outside of the brain-case is illustrated in text-fig. 16. The

foramen for the VIIth nerve lies just above and in front of the

fenestra ovalis, opening downwards through the pro-otic. The

CLASSIFICATION OF THE THERTODONTIA.

trigeminal ‘ foramen ”

dorsal in position.

lies considerably forward and is move

It lies at the end of a Jong slit and is pre-

sumably really only an incision and not a foramen.

The length

Text-figure 15,

aie

Leptotrachelus eupachygnathus.

Type-skull.

Right lateral aspect. X F.

Text-figure 16.

Hy)

i Abe

LYE i, & | zy

For Vi. PBR VE Fen.Ov. B.Oc.

TLeptotrachelus eupachygnathus, Type-skull.

Left lateral aspect of brain-case, the parts of the skull lateral to the post-temporal

fossa being removed as in text-fig. 10.

of the slit is rendered uncertain by the fracture of the anterior

end of this part of the brain-case.

Tn the part of the brain-case

preserved there is no evidence of the large venous foramen

58 MR. D. M. S. WATSON ON THE

described above in Scymnognathus whaitsi. There is no trace

of the great anterior projections of the basisphenoid which in

Seymnognathus meet in median suture in advance of the pituitary.

There is an ethmoid, which, so far as its very incomplete

exposure allows it to be seen, does not difter from that of

Scymnognathus.

The squamosal of Leptotrachelus is remarkable for the great

length of its lateral projection and the extreme suddenness of

the postero-lateral corner. As in Seymnognathus its distal end

is received between two bones, here almost certainly the jugal

and postorbital.

The cup-shaped depression in the widened lower edge of the

squamosal into which the head of the quadvrate fits is very narrow,

not half the width of the projection of the squamosal.

The quadrate is a relatively large bone about 30 mm. high by

15 mm. wide; it forms a nearly parallel straight-sided figure, the

lower edge being a little marked off by a groove and forming the

articular: surface. The upper end is rounded and fits snugly into

the hollow in the squamosal. The quadrato-jugal is fused with

the articular margin of the quadrate; it then separates from

that bone, leaving a small quadiate foramen, whilst farther

dorsally it spreads out into a flat sheet of bone which covers the

outer margin of the quadrate and laps over its posterior surface.

When articulated with the squamosal the quadrate and

quadrato-jugal are largely visible from behind.

The maxilla of Leptotrachelus shows a single canine in use,

with traces of the crown of a successional canine high up in the

alveolus, and four cheek-teeth ; it is possible that there was really

a fifth cheek-tooth.

LycosaurRvUs PARDALIS Owen, Cat. Foss. Rept. 1876, p. 15,

pl. 14.

The type-skull of Lycosawrus pardalis was re-examined and

discussed by Broom, Proc. Zool. Soc. 1911-12, p. 1079, who gave

it a dental formula: 1. 5, c. 2, m. 4.

The type-skull (R. 1717, B.M.N.H.) from the Cistecephalus-

zone (%) of the Sneeuberg 1s considerably crushed laterally, but

has the anterior end of ae snout complete and well-preserved ;

behind the canine on the left side the outer surface of the skull

is complete to the orbit, the whole orbital margin is present and

a bit of the edge of the parietal region. The other side is a

weathered face which cuts further and further imto the skull

until it so far crosses the middle line as to expose the admedian

surface of the left epipterygoid and completely to remove the

brain-case. The squamosals are completely destroyed. The

right lower jaw is, however, nearly perfect, having suffered only

the loss of the posterior part of the angular so as to expose

the articular—the position of the quadrate is thus fixed. The

parts of the skull remaining are quite well-preserved and show

many sutures,

CLASSIFICATION OF THE THERIODONTIA. 59

It is obvious that the snout is short, high, and narrow, with no

trace of the square section which occurs in all the Gorgonopsids

deseribed above.

The tooth-bearing edge of the maxilla is curved and passes

gently into that of the premaxilla without the step of Gorgonops

; : s

or Scymnognathus. ‘here is, however, a diastema between the

closely-set incisors and the canine.

There are clearly 5 incisors, 1 canine, and 4 or possibly 5 molar

teeth. Thesmall canine recorded by Broom immediately in front

of the large one does not exist on the well-preserved left side,

and his views seem to have been founded on a small strip of

tooth in this position on the right side, which is really an exposed

portion of the lower canine. The maxilla is short and deep.

Text-figure 17.

Fe. Fr, Niet Guha

Lycosaurus pardalis Owen. Type-skull.

Reconstruction of the right lateral aspect, X 2. The parts represented in

broken lines hypothetically restored.

The external nostril of Zycosaurus differs considerably from

that of Gorgonops. It faces more laterally and is much larger ;

it is no longer overhung by so large a corner of the nasal,

although there is still a trace of the older structure.

The facial part of the septomaxilla is much smaller, and the

foramen between that bone and the maxilla is not only smaller

but opens more directly outward. The septomaxilla in front of it

seems to be rounded and grooved. Finally, the internarial

process of the premaxilla is longer and stands more vertically, so

that the end of the snout is deeper and less rounded in side-view.

The interorbital region is narrow, the postfrontal being a

narrow pointed strip, as in the skull of Arctognathus curvi-

60 MR. D. M. 8S. WATSON ON THE

mola subsequently described (text-fig. 18). The parietal region

is obviously of the Gorgonopsid type and cannot be very wide,

although its width cannot be determined with any pretence to

accuracy.

ARCTOGNATHUS CURVIMOLA (Owen), Cat. Foss. Rept. 1876, p. 71,

pl. 68.

The skull (No. 47339 B.M.N.H.) deseribed by Owen as Lyco-

saurus curvimola was found with Dicynodon tigriceps i the

Cistecephalus beds of the Kagaberg, near Bedford, 8. Africa.

Its palate was developed by Mr. Hall and described by Prof.

Seeley, Phil. Trans. B. 185. The skull was then examined by

Dr. Broom, who noted that the parietal region seemed to be

broad, and made for it the genus Arctognathus.

The actual preservation of this skull is good; but before it

was buried the left maxilla and ectopterygoid, together with the

bit of lower jaw in their vicinity, were separated from the rest

of the head by a nearly plane split, moved outwards for about

15 mm. and there fixed in the sediment. How this very peculiar

result was brought about is very difficult to understand, although

tentative suggestions might be made.

Whilst lying at the surface the nodule containing the skull

was exposed to weathering, which has cut down into it so as

completely to remove the right squamosal, the parietal region

beyond the middle line, and the postorbital bar.

Fracture has removed the occipital condyle and part of the

paroccipital process, but has left the stapes and quadrate with

the lower jaw in articulation on the right side. Enough of the

occiput is left to make the structure clear. ‘The palate is well

exposed and very well preserved, the right ramus of the mandible

is perfect and weli-exposed.

On the dorsal surface of the parietal region the suture between

the parietals and the pineal foramen are very well shown on a

weathered face, which lies a little below the original dorsal

surface; the right side of this region retains its natural surface

and shows the structure clearly.

The skull is short, broad, and deep. The snout is rounded in

section and terminates in front in the internarial premaxillary

processes, which form the extreme front end of the skull over-

hanging the oral margin.

The very large nostril faces largely outward and is not over-

hung by an outstanding corner of the nasal. The septomaxilla

is small, and the foramen between it and the maxilla very small.

The interorbital width is considerable, but the orbits look

upward and forward as largely as outward. The frontal does

enter into the orbital margin, but only throygh a short distance

The postfrontal is a narrow strip of bone wedged in between the

frontal and the postorbital.

No trace of a preparietal is to be seen on the parts preserved,

CLASSIFICATION OF THE THERIODONTIA. 61

the median suture is clearly shown from the front of the pineal

foramen to a point between the frontals, and the well-marked

suture between the frontal and parietal passes very little in

advance of the pineal opening. These sutures are, however,

exposed at a plane below the orginal dorsal surface, and there is

a remote possibility which cannot, although very improbable, be

entirely excluded, that the preparietal was represented by a

Text-figure 18.

Arctognathus curvimola Owen. ‘Type-skull.

Restoration of the dorsal aspect, the parts in broken lines being hypothetically

restored.

minute scale of bone lying on the dorsal surface. ‘The parietal

region 1s about as wide as the interorbital.

The maxilla is short and deep, its co aA margin 1s

much curved and passes smoothly with no trace of a step into

that of the premayilla, The canines appear not to be completely

erupted, and the four small cheek-teeth are also not very firmly

62 MR. D. M. 8S. WATSON ON THE

planted. The four incisors, though powerful, are not so dispro-

portionately large as they are in earlier Gorgonopsids.

The palate of Arctognathus is very well-preserved, but its

anterior end and two strips along the maxille are concealed by

unremoved matrix and by the lower jaw.

The basioccipital is broken off through the vagal foramen,

where it is thin and not very wide. The posterior part of the

Text-figure 19.

Arctognathus curvimola (Owen). Type-skull.

The palate, X 3. Stippled area covered by matrix and the lower jaw.

Parts in dotted lines hypothetical.

Ae A, direction of the section of ‘‘ Lycosaurus tigrinus,” text-fig. 20.

St., stapes.

basisphenoid forms a triangular area with raised lateral margins,

representing the tubera of earlier forms. Above the edges the

sides of the bone are flat and vertical, posteriorly they terminate

in the region of the fenestiz ovales, these openings beimg con-

cealed by the foot of the stapes. Anteriorly these vertical sides

of the basisphenoid approach together until they are only

CLASSIFICATION OF THE THERIODON'TIA, 63

separated by a narrow ridge which runs forward to the palate

proper. From the vertical sides of the basisphenoid horizontal

processes arise; these basipterygoid processes support the ptery-

goids. From their articulation with the basisphenoids the

pterygoids pass backward towards the quadrates, but do not

articulate with those bones, as they appear to terminate in free

points before reaching them. The pterygoids pass forward,

forming with the median ridge which continues the basisphenoid

a bar whose ventral surface is almost cylindrical, broken only by

the median fillet. At the hinder end of the palate the pterygoids

suddenly widen, forming very deep powerful flanges. This part of

the bone has a transverse suture with the ectopterygoid. Medially

the two pterygoids meet in a visible suture which lies at the bottom

of a small depression. ‘his suture soon terminates at the brim

of a much deeper and more sharply-marked hollow, which, as it

passes forward, widens and is converted into a deep open groove

forming anteriorly the whole roof of the much vaulted palate.

Throughout its extent this groove has well-marked, indeed often

vertical, sides. Anteriorly this groove is divided into two by a

ridge which rises from its surface. At about the level of the

last maxillary tooth this groove is bounded by roughened areas

of bone, which appear to have borne teeth. These areas are

undoubtedly on the pterygoids and are separated by visible

sutures from the palatines, which le laterally to the pterygoids

in front of the ectopterygoids. Further forward these sutures,

which form the inner border of the palatines, approach one

another and descend into the groove, so that its side-walls are in

front formed by the palatine. The ectopterygoids are separated

from the palatines by visible sutures.

There is no trace of a suture down the mid-line of the groove,

and its roof seems to be formed by a median bone, which

terminates at the sudden end of the groove and must be bounded

by sutures with the pterygoids along its edges ; of these presumed

sutures nothing can be seen in this specimen.

The type-specimen of Lycosaurus tigrinus Owen seems to throw

light on the structure of the palate of Arctognathus curvimola.

It consists of a fragment of a snout, broken off through the

premaxille in front and by an oblique fracture on the left side,

but showing much of the right maxilla. It has been so developed

as to show a small strip of the surface of the right palatine and

shows a section of the palate on the hinder end. ‘This species is

referred by Broom to a new genus Arectosuchus, and said to have

a dental formula, i. 5, ¢c. 1, m. 4 ov 5, representing a much more

primitive type of Theriodont than Arctognathus. The type-

specimen only shows two incisors, a canine, and a few cheek-

teeth, and it seems certain that Broom examined and used for

his description a snout of Seymnognathus whaitsi which Lydekker

had referred to Z. tigrinus.

Except in the larger size and somewhat different direction of

its canine, the type-specimen of ZL. tigrinws seems to agree

64 MR. D. M. S. WATSON ON THE

exactly in size and every point which can be compared with

A. curvimola. The section of the palate shown on its posterior

fracture is represented in text-fig. 20.

There is a median element bearing a high dorsal ridge, now

detached, with a very deep groove on its mid-ventral surface and

earried out laterally in a long wing, which is overlapped by the

pterygoid. This bone bears a powerful irregular roughened

projection carrying small teeth: laterally its surface is smooth,

and is continued to the hinder end of the maxilla by that of the

ectopterygoid. he two bones scarcely meet, but are joined

together by a thin film of bone, undoubtedly the palatine, which

covers their dorsal surfaces.

Text-figure 20.

Aretognathus. Type-specimen of Lycosaurus tigrinus Owen. X 5.

A) } iy 3

Obliquely transverse section as a plane corresponding to A...... A in text-fig. 19.

If this section be compared with that which the palate of

Arctognathus would present if cut along the line A~A, there can

be no doubt of the close aftinity—indeed, specific identity—of

the two forms, for even the possible measurements are in very

close agreement.

Thus we have confirmation for the view that the roof of the

median groove in the palate of Arctognathus is formed by a

median vomer.

The epipterygoid of the type-specimen of A. curvimola 1s

shown to meet the parietal in a long suture, exactly as does that

of Diademodon.

The preceding series of description is based on the more

complete and satisfactory remains of Gorgonopsids in the British

Museum, largely Hndothiodon-zone forms. Of recent years

Broom and Haughton, either independently or together, have

described many complete Gorgonopsid skulls, chiefly from the

Cistecephalus-zone. ‘They have, however, never given so complete

an account of any form as that of Seymnognathus included in

this paper, and it is seldom that they have given more than one

CLASSIFICATION OF THE THERIODONTIA. 65

or, at most, two drawings of any one skull. Thus it is difficult

to carry out any detailed discussion of the mutual relations of the

known Gorgonopsids,

In order to discuss with any satisfaction the classification of a

group, it is necessary to know the main outlines of its history, to

understand the direction of the advances which make the struc-

ture of all late members of it differ from their ancestors, and

to work out any adaptive modifications which characterize its

different branches.

It is now generally agreed that the Gorgonopsids include the

ancestors of the Cynognathide, and that the Pelycosaurs are a

group which includes self-contained lateral branches springing

from the very base of the stem of the mammal-like Reptiles.

Thus by comparing the two extreme terms, Varanosaurus and

Diademodon, we can gain at once a knowledge of the advances in

structure which have occurred in the Anomodonts, and on the

assumption that these changes have proceeded regularly we can

determine the trend of advance during the evolution of the group.

Discussion of intermediate forms will then enable us to decide

whether this trend really expresses a true view of the mode of

evolution, or whether the actual observed differences between

the extremes represent the result of a series of fortuitous changes

of indeterminate direction.

The work of Broili, of Case, of Williston, and the present writer

has led to the view that Varanosaurus is the most primitive known

Pelycosaur, forming a morphological ancestor to Dimetrodon,

through a Deiopews-like form. The view that Diademodon or

Trirachodon is the most advanced of known Anomodonts results

from the work of Seeley and Broom.

The differences between the skull of Diademodon and Varano-

SaUrUs are :—

In General Shape.

In Varanosaurus the snout is very long, square-cut in section,

and roomy, compared with the rest of skull, with lateral nostrils

and a long straight tooth-row. The large orbits are entirely

laterally directed. The smail temporal fossa lies entirely on the

side of the skull and is almost hidden from above by the very

broad parietal region. The occiput slopes forward, but is not

deeply cupped. The sides of the skull are nearly straight. The

skull is higher than wide.

In Diademodon the snout is short, small in volume, rounded in

section, with nostrils looking more forward than laterally. The

tooth-row is short and curved The orbit is comparatively small

and looks very largely forward. ‘The temporal fossa is very

large, lies entirely on the top of the skull, and is not visible from

the side, the parietal region being drawn up into a narrow crest.

The occiput slopes «a little forward and is deeply cupped. The

sides of the skull gradually approach one another to the orbits

Proc. Zoou. Soc.—1921, No. V. 5

66 MR. D. M. 8S. WATSON ON THE

but their direction is then changed as they form the slender

snout. The skull is much wider than high.

In the Brain-case.

[The structure of the anterior face of the pro-otic and supra-

occipital is not known in Varanosaurus and other regions are not

very well shown. The following account of the Pelycosaurs is

based on Deiopeus and Dimetrodon. |

In Pelycosaurs the basioccipital is thick, ending in a large

rounded condyle. The large fenestrz ovales are placed on the

bottom of the skull, far out at the side of the deep well-developed

tubera. ‘The paroccipital process is short and slender, supporting

the squamosal and touching the tabular; it les well above the

lower surface of the skull. The pro-otic is small, its anterior face

in no way overhanging the notch for the fifth nerve. The supra-

occipital is entirely plate-like, not forming a roof over the brain

in advance of the Vth nerve. The basisphenoid is massive,

forming a sloping floor to the posterior part of the brain-case.

It bears definite Sphenodou-like basipterygoid processes, anteriorly

it is in Varanosaurus continued forward by a long channel-shaped

parasphenoid.

The parietal does not form any part of the side-walls of the

biain-case. ‘The epipterygoid is a slender rod of circular section.

The whole brain-cavity is very small in comparison with the

size of the skull.

In Diademodon, on the other hand, the basioccipital is small

and plays at most a subsidiary part im the pair of occipital con-

dyles. The small fenestree ovales are placed on the bottom of the

skull, not very far separated. Basisphenoidal tubera are repre-

sented merely by the edges of the triangular lower face of the

basisphenoid. The paroccipital is a long powerful process sup-

porting the squamosal and touched by the tabular; it lies on the

lower surface of the skull.

The pro-otic is large, being carried forward by a great process

which completely overhangs the trigeminal foramen.

The supraoccipital is produced forwards by two wings, which

cover and form side-walls to a great deal of the brain-cavity in

advance of the Vth nerve.

The basisphenoid is a small bone forming a nearly horizontal

floor to the brain-case. It has small lateral basipterygoid pro-

cesses with the pterygoids attached to thei flat lower surfaces ;

anteriorly it is carried forward by a slender process which reaches

the palate and there spreads out into a broad vomer in the roof

of the posterior part of the nasc-pharyngeal ducts.

The parietal forms a good deal of the side-wall of the brain-

case. ‘The epipterygoid isa flat plate forming the side-wall of

the brain-case for some distance and articulating with the anterior

edges of the pro-otic. The brain-cavity is relatively very large.

The ear of a Pelyeosaur, so far as can be inferred from the

bone which housed it, lies low down on the side of the brain-ease,

CLASSIFICATION OF THE THERIODONTTA. 67

has small simple semicircular canals, and has a cochlea which

leaves no evidence of its existence on the bones. ‘lhe fenestra

rotunda is represented by a notch on the ridge, which in the

bony skull separates the vestibular cavity from the vagal foramen,

and thus opens inside the brain-eavity. The stapes is always

very large and is perforated, the fenestra ovalis being a large

irregular hole. There is no groove for the external auditory

meatus.

In Diademodon the inner ear retains its original position low

down on the side of the brain-case and still shows only simple

semicircular canals. It has, however, a well-defined cochlea

housed in a erypt passing forward and inward, and curved

forward through about a quadrant of a cirele. The fenestra

rotunda is a complete foramen, which opens indeed into the vagal

foramen, but does so on the outer surface of the neural cranium,

exactly as if does in the young Ornithorhynchus.

The stapes, although still of good size, is much smaller than in

Pelycosaurs, and the fenestra ovalis is a neat round hole of

small size. The external auditory meatus is housed by a deep .

groove.

The nose of Diademodon occupies a smaller space than that of

Dimetrodon, but the area of its sensory epithelium seems to have

been increased by a great development of turbinal cartilage, now

only represented by a series of ridges, on the inner surface of

the nasals and prefrontals which once supported them. Nothing

of the kind occurs in Pelycosaurs,

Many other features in the nose of the Anomodonts can only

be discussed in connection with the septomaxilla, palate, etc.,

and then only im a detailed discussion of individual forms.

The Roof of the Skull.

In Varanosaurus the parietals ave short, very broad, and with

the pineal foramen very far back. Their edges are separated

from the temporal fossee by a union of the postorbital and

squamosal. There is a large postfrontal lying on the roof of the

skull. The frontal is a large bone always entering into the

orbital margin. The prefrontals are large bones, almost equally

divided between the dorsal and lateral surfaces of the skull,

each bearing a depression on the outer face just in front of the

orbit. The nasals are narrow slips of bones. There is a small

supratempoval,

In Diademodon the pavietals are long, very narrow, and with

the pineal foramen between their anterior ends. ‘They form the

inner margins of the temporal fosse for a very long way, the

squamosal and postorbital being widely separated. ‘There is no

postfrontal. The frontal is a small bone, not entering the

orbital margin. ‘The prefrontals are small bones on the rounded

snout, with no depression in front of the orbit. The nasals are

wide, especially posteriorly. There is no sjpuniioutypant ls

68 MR. D. M. S. WATSON ON 'THE

In side-view the skull of Varanosaurus shows a long shallow

maxilla, completely separated from the nasal by the long lachrymal

extending from the orbit to the septomaxilla. There is no very

marked specialization of the dentition, all the teeth from the

premaxilla backward being similar in form and not very dis-

similar in size. The quadrato-jugal is exposed on the side-wall of

the skull and the jugal stops considerably before the quadrate.

Diademodon has a short deep maxilla reaching the nasal in a

long suture. The lachrymal is a small short bone. The dentition

1S fully divided into incisor, canine, ‘‘ premolar,” and “ molar

teeth.”

The quadrato-jugal no longer appears on the surface of the

skull and the jugal extends back to its extreme hinder end.

The suspensorium of Varanosaurus consists of a large quadrate

with a definite pterygoid wing, whose posterior surface J 1s covered

by the pterygoid... The outer edge of the quadrate is attached to

the quadrato-jugal, there being no quadrate foramen. The upper

part of the posterior surface of the quadrate is covered by

the squamosal, that bone passing so far inwards as to touch the

pterygoid.

In Diademodon the quadrate is a very small bone, either with

or without a pterygoid wing, but in no case articulating with the

pterygoid. ‘The outer edge of the quadrate is fused with the

quadrato-jugal, from which it is separated only by a small foramen,

the articular surface being formed about equally by either bone.

The whole posterior surface of the joint bone is covered by the

very large squamosal, which extends down to the condylar edge.

The primitive Pelycosaur palate has the following characters :—

The pterygoid is a triradiate bone, articulating “by a movable

facet with the basipterygoid process, from which point the

quadrate ramus rises and runs backward as a vertically placed

sheet of bone, passing behind the quadrate. The lateral wing of

the pterygoid passes directly outward from the region of the

basipterygoid and terminates in the usual flange.

The anterior part of the pterygoid forms a large part of the

essentially flat palatal surface and articulates with ‘the prevomer.

It meets its fellow in median suture in Varanosaurus. he

dorsal surface of the pterygoid is raised into a ridge near the

middle line of the skull. In later forms (Dimetr afl, e. g.), the

ridge is much exaggerated and its median surface passes

smoothly into the ventral surface. The palatines are small flat

bones. The prevomers are distinct. The anterior end of the

palate is not known in any primitive Pelycosaur, but from

the conditions in later forms there is no doubt that the posterior

nares were small and lay in the general plane of the rest of the

palate, which was essentially Ane In such later Pelycosaurs as

Dimetrodon, owing to the step in the lower edge of the maxilla,

the palate is considerably vaulted, and the posterior nares lie

above the level of the cheek-teeth.

In Diademodon the pterygoid articulates by a rigid suture with

CLASSIFICATION OF THE THERIODONTTA. 69

the lower surface of the basipterygoid process, and there is no

quadrate ramus, the bone terminating at its attachment. The

transverse ramus does not arise from the basipterygoid region,

but very much further forward, the posterior part of the bone

being a slender strip, which is united with its fellow and the

parasphenoid to form a slender bar.

The transverse ramus arises very suddenly from this bar, the

bone being drawn downward into deep and very powerful flanges.

There is no anterior vamus, the bone ending in a transverse

suture with the palatine at the region where it joins the jugal.

The transverse bone is very small and is not included in the

flange at the foot of which it lies.

The palatine is a very large bone, forming a great deal of the

posterior part of the palate, then turning downward to form the

lateral wall of the posterior part of the nasopharyngeal duct,

and, finally, developing a secondary plate, which forms a floor to

that passage.

The two palatines are separated by a median vomer, which

forms the roof of the nasopharyngeal passage, apparently ter-

minating behind in a pointed slip separating the anterior ends of

the pterygoids, but really passing backward to the basisphenoid.

The prevomers seem to have vanished entirely.

The maxillz send inward secondary plates, which continue those

of the palatine forward. The premaxille have palatal processes,

which pass dorsally to the secondary plates of the maxille and

touch the anterior end of an ossification in the nasal septum,

viz., a mesethmoid.

In Varanosaurus the epipterygoid is a slender rod rising

vertically from the dorsal surface of the pterygoid just in advance

of the basipterygoid articulation.

In Diademodon the epipterygoid is a large flat sheet of bone

forming a side-wall to the brain-case and articulating by a very

long suture with the parietal and frontal above, having a suture

with the pro-otic behind, articulating with the basipterygoid

process below, and ending in a suture with the pterygoid.

Behind the basipterygoid and below the point of exit of the

maxillary, mandibular, and motor portions of the fifth nerve, the

epipterygoid is continued backward by a process occupying the

position of a quadrate ramus of the pterygoid. The ramus in

certain species articulates with the front face of the quadrate.

The preceding pages record the more important differences

between the most primitive and the most advanced known

Anomodont; they bring out the direction of the evolutionary

advances and show how enormous is the structural gap between

them, a gap represented in time by the relatively small interval

between the bottom of the Permian and the middle of the

Triassic system.

It remains to show how completely this morphological gap can

be bridged by the material available. Although, as Case and

Williston have repeatedly and emphatically pointed out, the Pely-

cosaurs are a self-contained group dying out with Dimetrodon and

70 MR. D. M. S. WATSON ON THE

Edaphosaurus, they show evolutionary changes which are in the

main in the same direction as these which lead from Varanosaurus

to Diademodon, coupled, of course, with many individual speciali-

zations. Although it is not essential for the purpose of this

paper, it is, I think, useful to point out certain of the more

striking of these advances, using Varanosaurus, Deiopeus, and

Dimetrodon as the series of forms. These animals lived side by

side, and this series is only a morphological one.

In general shape Dimetrodon shows an advance on Varano-

saurus in that the snout is much deepened, is square in section

only immediately in front of the orbit, and is, in general, wedge-

shaped with a rounded dorsal edge. The par ‘ietal region 1s

narrower, and leaves the temporal fossee visible from above. The

oceiput is more vertical. In the brain-case Dimetrodon shows an

advance over Deiopeus in that the basioccipital is thinner, the

basisphenoidal tubera smaller, and the paroccipital process larger.

The anterior margin of the pro-otic lies furtherin advance of the

internal auditory meatus. The brain-cavity is considerably

deeper and wider posteriorly. The fenestra ovalis of Dimetrodon

is smaller than that of Deiopews and the stapes lighter. The roof

of the skull of Dimetrodon differs from Varanosaurus in the

following ways:—the parietals are Jess wide and the pineal

foramen further forward. The postorbital is visible from above.

The pair of frontals have acquired a cruciform shape owing to a

widening of the interorbital surface. Deiopeus provides an exact

intermediate, the increased width of the interorbital surface

having arisen by an increase in size of the pre- and post-frontals,

so as to leave a gap which is filled up by a special process of the

frontal.

In side-view the skull of Dimetrodon shows a shortened and

deepened maxilla touching the nasal in a short suture. The

dentition is sharply divided into incisors and cheek-teeth by a

diastema. The third maxillary tooth is much larger than the

first two, and the lower border of its socket lies well below the

dentigerous border of the premaxilla.

The lachrymal does not reach the septomaxilla. The orbit is

placed high up in the skull.

It seems probable that the deepening of the maxilla and the

“step”? depend on the necessity of finding room for the roots of,

and the development of, replacing teeth for the greatly lengthened

maxillary teeth.

The condition of the canine may depend on the following con-

siderations:—A. large canine in the upper jaw presupposes a

similar tooth in the lower jaw; such teeth, which are designed

for killing animals, are most useful in the front of the mouth. The

lower jaw, as a whole, bites inside the upper jaw. The first tooth

of the lower jaw cannot be much enlarged, because of the difficulty

of making a pit for its reception near the middle Ime in the

palatal process of the premaxilla; hence the lower canine cannot

be quite at the end of the jaw. A large lower canine almost

involves a diastema in the upper jaw for it to bite into, because

CLASSIFICATION OF THE TILERIODONTIA, 71

Text-figure 21.

Series of dvawimgs of the right lateral aspect of Pelycosaur skulls,

reduced to the same length.

A. Varanosaurus acutirostris Broili, restoved from the type-skull in Munich, partly

after Broili. B. Deiopeus leptocephalus Cope, restored from the type-skull

in the American Museum. C. Sphenacodon ferow Marsh, after Williston.

D. Dimetrodon gigas Case, from a photograph published by Case.

72 MR. D. M. S. WATSON ON THE

being laterally placed, the formation of a pit in the palate

fer its reception so narrows the dentigerous part of the maxilla

outside it as to leave no room for a tooth. The same factors will

render difficult any great enlargement of the next tooth, and the

upper canine thus comes to lie behind the diastema and separated

from it by at least one small tooth. Considering the mechanics

of the whole arrangement, it appears useful to make the dentition

more or less symmetrical about the diastema by enlarging a pre-

maxillary tooth to correspond with the canine, so that the single

lower canine forces the prey into the gap between two large upper

teeth. The dentition so designed is realized in Dimetrodon.

Text-figure 22.

Re es mn ee

A. Superimposed outlines of sagittal sections of the brain-cases of Deiopeus in

broken line and Dimetrodon in continuous line reduced to the same length,

to show the thinning of the basis cranii, the enlargement of the brain-cavity,

and the forward growth of the pro-otic.

B. Superimposed outlines of sagittal sections of the brain-cases of Leptotra-

chelus in broken line and Diademodon in continuous line reduced to the

same length, to show: the thinning of the basis cranii, the forward growth

of the pro-otic, the enormous increase of the cerebellar cavity, and the

relatively slight growth of the cerebral region.

The great deepening of the maxilla demanded by the canine

automatically squeezes out of existence the anterior part of the

lachrymal.

Deiopeus and Ophiacodon provide an exact intermediate between

CLASSIFICATION OF THE THERIODONTIA. . 73

Varanosaurus and Dimetrodon in dentition, as Case has already

shown.

In the palate Dimetrodon differs from Varanosaurus in that the

transverse flanges of the pterygoids have moved much forward

from the basipterygoid region. The condition in Deiopeus is not

known. Thus many of the structural changes which separate the

advanced from the primitive Pelycosaurs are in the same direction

as the general advances of the Anomodonts from first to last.

The earliest known Gorgonopsid is Galesuchus gracilis Haugh-

ton, from the Zapinocephalus-zone.

This skull I have not seen, but judge that it is so much

weathered that the original shape of the squamosals cannot be

seen. Only the dorsal aspect has been figured. The skull is very

remarkable, differing much from all later Gorgonopsids and even

more from Pelycosaurs. Its only primitive features seem to be

the extremely sloping occiput (the apparent slope being possibly

exaggerated), the high position of the foramen magnum and

foramen jugulare, the large deep paroccipital processes, the

great size of the frontal, and the lateral direction of the orbits.

It is, however, so incompletely known that a full discussion of

its affinities is impossible.

The most primitive known Gorgonopsid is Arctops. This re-

tains as primitive features, which it shares with the Pelycosaurs :—~

the square section of the snout with a depression on the preorbital

surface overhung by the prefrontal, the lateral direction of the

orbits, the extreme parietal width, and the shortness of the

parietals. The resemblances in the basicranial and otic regions

have been discussed (P. Z. S. 1914, p. 1027).

The palate retains a primitive structure in the non-fusion of

the prevomers.

The skull shows advances over Varanosaurus in the following

ways :—Owing to the need of increasing the size of the temporal

muscles the squamosal is bayed outward, a modification which

makes the temporal fossa visible from above; this change not

only allows of greater thickening of the temporal muscles during

their contraction, a function which is believed by Gregory and

Adams to be the factor which determines the origin of fenes-

tration, but also enables the outer part of the temporal muscle to

acquire a new origin on the upper edge and inner surface of the

zygomatic arch, thereby establishing an independent masseter

muscle. At the same time this widespread zygomatic arch passes

much laterally to the quadrate and leaves that bone, with

the quadrato-jugal attached to its outer margin, lying entirely

within the back of the enlarged temporal fossa; these bones,

having thus lost the support that they originally received from

the junction of the quadrato-jugal with the jugal and squamosal

by sutures, can only be adequately supported by a more powerful

abutment on the paroccipital process and by an extension of the

squamosai down their posterior surface.

At the same time the lateral extension of the squamosal renders

14 MR. D. M. S. WATSON ON THE

the back of the skull much wider than the neck, and makes the

tympanic membrane lie much nearer the middle line than the side

of the skull. As it is necessary to keep this membrane exposed

to the outer air, any swelling of the neck will automatically lead

to the formation of an external auditory meatus; the beginnings

of this passage are seen in Arctops in a groove on the posterior

surface of the squamosal, just outside the end of the paroccipital

process.

On the vential surface Arctops has advanced over Varano-

sawrus in the shift forward of the great pterygoidal flanges to

a position far in advanee of the basipterygoid processes ; this

change results in a further great enlargement of the cavity for the

temporal muscles, and allows the development of great pterygoidal

muscles with an insertion on the dorsal surface of the palate.

This shift forward of the flanges has occurred in Dumetrodon,

but the conditions in Arctops differ from those in the earlier

form in the fusion of the posterior parts of the pterygoids and

parasphenoid into a massive ridged girder. This change adds

greatiy to the strength and rigidity of the skull, but its details

cannot readily be explained by mechanical considerations; the

most important of these is the replacement of normal basiptery-

goid processes by the laterally directed flat lappets which occur

in all Theriodonts.

In the palate itself the more important changes are the

development of a median groove, a necessary preliminary to the

establishment of a secondary palate, which is brought about by

that change in the dentition involving the development of a

‘step ” between the maxillary and incisor teeth which has been

discussed under Dimetrodon, and the extreme posterior position

of the hinder ends of the posterior nares; this latter change

is itself probably to be associated with the incipient secondary

palate, leading as it does to a longer air-passage whose posterior

end is not so easily closed by the presence of food in the anterior

part of the mouth.

Another advance in Arctops is the more vertical position of the

occiput.

Glorgonops is in some ways as primitive as Arctops, with which

it shares very broad parietal and interorbital regions and a

square sectioned snout. The orbit faces laterally in both forms,

and each has a remarkably broad interparietal, a feature in which

they resemble Deinocephalians.

Gorgonops has a pair of large frontals, which are cruciform in

plan exactly as in the later Pelycosaur.

The prefrontal is a large bone which overhangs a well-marked

depression on the preorbital surface.

Gorgonops shows advances over Varanosawrus which, so far as

the parts are known, include all those which occur in drctops,

with the following additions :—Lhe external nostril in Gorgonops

is much complicated by the great development of the septomaxilla

and the foramen behind it. This foramen, first recognised hy

CLASSIFICATION OF 'THE THERIODONTTA. 75

Case in Dimetrodon, occurs in all the more primitive Anomodonts

and is still of uncertain function. In all Pelycosaurs the septo-

maxilla is a small bone resting on the maxilla and premaxilla at

the back of the nostril, being touched by the nasal and forming

with the other bones the aie amen, which I propose to call the

septomaxillary foramen.

In Dimetrodon the lower edge of the septomaxilla, where it

rests on the maxilla, is turned inwards and forms a partial floor to

the nasal passage. The anterior border of the septomaxilla is

provided with a process which partially divides the external

nostril into a lower and an upper part.

It is probable that the process on the anterior edge of the

septomaxilla is associated with the original turbinal, simply

forming the anterior end of that ridge. iia the living lizards and

snakes, “Whe septomaxilla lies inside the nostril as it does in the

Captorhinids, the collateral ancestors of the Pelycosaurs; it has

in them a characteristic and uniform situation, in that it is

ossified in the membrane dividing the main nasal cavity from

Jacobson’s organ, running nearly horizontally from the maxilla

to the cartilaginous nasal septum.

In evomotionts I have only heard one possible suggestion for

the function of the septomaxillary foramen, that it ser rved as an

outflow for the ductus naso-lachrymalis, the liquid poured out

from it serving to keep the muzzle wet asin Artiodactyls. This

view is in harmony with the known position of the duct in early

amphibia and reptiles. It, however, does not afford any satis-

factory explanation of the eveat size of the foramen in Gor gonops.

The very peculiar conditions in the Deinocephalian Jormo-

saurus (Proc. Zool. Soc. 1914, p. 757, figs. 1 & 2) suggest another

explanation. In this animal the ordinary external nostril, which

in early reptiles always lies between the premaxilla, the septo-

maxilla, and nasal, appears to be represented by a minute foramen

between the septomaxilla and the nasal. The large opening

which is the functional nostril seems to be really a septomaxillary

foramen, as it lies between the premaxilla, the maxilla, and

septomaxilla, a situation which is never occupied by the ordinary

nostril in early reptiles, but agrees with that of the septo-

maxillary foramen in the earlier Vheriodonts. Thus this foramen

must be of the nature of a nostril. The characteristic position

of the septomaxilla in Squamates suggests that the foramen leads

into Jacobson’s organ, and it will follow that that organ was

the functional olfactory organ of Mormosaurus; the shallowness

and small size of the upper part of the nasal cavity, which

distinguishes Mormosaurus from such Deinocephalia as J/oschops

with a normal septomaxillary foramen and nostril, can thus be

accounted for.

The advances over Varanosaurus which are shown in the nose

of Gorgonops ave:—(1) the direction of the nostril forwards

instead of laterally, a change which renders the appreciation of

odours coming from the direction in which the animal is pro-

76 MR. D. M. 8. WATSON ON 'THE

ceeding more delicate, because owing to a Pitot effect more air

will be driven into the nasal cavity; (2) a great increase in the

size of the septomaxillary foramen, possibly associated with a

further elaboration of Jacobson’s organ ; (3) a great increase in

the size of the facial part of the septomaxilla—this may be due

to purely mechanical reasons.

Another advance very clearly shown in Gorgonops is a great

deepening of the maxilla and a concurrent reduction of the

lachrymal, the prefrontal showing little reduction.

The palate of Gorgonops is advanced in the great size of the

internal nares, in the width, depth, and backward extension of

the median groove, in the internarial bar being single, and in

the occurrence of a median vomer in the back of the palate.

The very large size of the pterygoids and the forward position

of the roughened and possibly tooth-bearing areas on these bones

are primitive features.

The basicranial region shows no structural detail.

Scymnognathus whaitsi shows many resemblances to Gorgonops

1m its snout.

It shows advances over Avctops in the thinner basioccipital,

smaller basisphenoidal tubera, and less massive paroccipital pro-

CeSses.

The brain-case is advanced over that of the Pelycosaur

Dimetrodon in the great forward extension of the pro-otice and

supraoccipital and the junction of the latter bone with the

epipterygoid ; a remarkable feature is the forward process of

the basisphenoid, which forms a floor to the brain-case in

advance of the pituitary fossa.

The most striking advances over Arctops are the great redue-

tion in width of the parietal region, the lengthening of the

pavietals, and especially the enormous increase in spread of

the squamosals. The turning backward of the upper part of the

squamosal at the posterior margin of the temporal fossa makes

that opening even larger, and increases the length of certain

fibres of the temporal muscles.

So far as known, similar differences separate Scymnognathus

from Gorgonops, the latter genus having much larger frontals

than the former.

By the great expansion of the width of the back of the skull,

the orbits of Scymnognathus ave made to look partly forward.

Another small but important advance in Scymnognathus is

that certain fibres of the temporal muscle have secured an origin

from the dorsal surface of the parietal region.

Seymnognathus is probably less advanced than Arctops and

Gorgonops in its less vertical occiput. In the palate Seymno-

gnathus is probably less advanced than Gorgonops in the small

size of the median groove, and its restriction to the anterior

part of the palate and to a narrow space round the hinder end

of the posterior nares.

The incompletely known Leptotrachelus shows an important

CLASSIFICATION OF THE THERIODONTIA, re

stage in the advance in structure of the basieranial region,

already discussed in P.Z.S8. 1914, p. 1027, fies. 3 & 4; it is in

this region more advanced than any other known Hndothiodon-

zone Gorgonopsid. It retains as primitive features the very large

quadrate, large postfrontal and frontal, and a sloping occiput.

Thus the “Yndothiodon-zone Gorgonopsids show definite ad-

vances over the Pelycosaurs in the direction of Diademodon.

Kach form is advanced in éertain features whilst retaining a

more primitive structure in others, so that an imaginary animal,

built up by throwing together the most advanced features found

in all the actual animals, would be far more advanced than any

one is on the average; although in no point would it be more

advanced than a known form. tn fact, the evidence existing

here, small though it is, suggests that tihee is a limit to the

total amount of advance possible to the members of a group in a

given time, and that these changes may be distributed either

over the whole animal or concentrated on a definite region, which

will then present a structure of much more advanced type than

is found in allied contemporaneous forms. A somewhat similar

conclusion seems to have been reached by W. D. Matthew from

the study of the more abundant material of fossil mammals.

Discussion of the Gorgonopsids of the Cistecephalus-zone is

rendered difficult by two factors—the incomplete descriptions

and insufficient figures of many of the perfect skulls in S. Africa

and New York, and the fact that the Cistecephalus-zone is a

long one and that we do not know the relative ages of the

Gorgonopsids from it. It will appear from the evidence to be

brought forward in this paper that the forms from Dunedin and

Nieuweveld localities are early, those from New Bethesda and

the Kagaberg which are associated with Dicynodon tigriceps

considerably later in time. There is, however, no stratigraphical

evidence that this is so.

From the Cistecephalus-zone Broom and Haughton have

described several forms as species of Scymnognathus—s. tigriceps

B. & H., S. parvus Br., S. minor Bv., S. angusticeps Br., S. serra-

tidens Tau., are all from the Nieuweveld. These forms may

very possibly be congeneric; they agree with Scymnognathus in

having 1. 5, c. 1, m. 4-5, but quite certainly do not belong to

that genus. They differ from Scymnognathus whaitsi in the

following characters :—

The snout is very much deeper, its anterior end instead of

being rounded is vertical (¢f. Broom, P. Z.8. 1913, p. 225, pl. 36),

the external nostril is much larger, the septomaxillary foramen

smaller. The anterior end of the nasal does not fully overhang

the nostril. ‘The top of the snout may be ridged, and the square

section with a preorbital depression overhung by the prefrontal

is entirely lost (cf. S. serratidens, Ann. South Afr, Mus. vol. xii.

p. 89, pl. xii.). The snout is much shorter and the prefrontal

in consequence smaller,

78 MR. D. M. 8S. WATSON ON THE

There is no step in the upper jaw, the curved tooth-bearing

edge of the premaxilla passing smoothly into that of the maxilla.

In_ 8. tigriceps thé squamosals are not nearly so much spread, the

skull being much deeper in proportion to its width. If we may

trust the existing description of a not very satisfactory prepara-

tion, the palate of S. tigriceps differs from that of S. whaitst in

the loss of the anteor projection of the pterygoid and the great

extension of the palatines. On the other hand, judging from

the description by Haughton of its endocranial surface, the

brain-case of S. tigriceps may have greatly resembled that of

S. whaits.

Thus the Cistecephalus-zone animals referred to Seymno-

gnathus do not belong to that genus, but differ from it by a

series of advances which will be seen to be all in the direction

leading to Diademodon. Certain of these species appear to

resemble Lycosaurus pardalis considerably, agreeing with that

animal in dentition, the short high snout with a rounded dorsal

surface, the large nostril, the vertical imternarial bar, the small

exposure of the septomaxilla, the absence of a step in the upper

jaw, the short and deep maxilla, and the small prefrontal. Ail

of them, however, seem to retain a large postfrontal bone.

Two other remarkable forms, apparently from the lower part

of the Cistecephalus-zone, Scylacops capensis and Corgognathus

longifrons, ave of interest because they strongly recall Hndothiodon-

zone forms.

Gorgognathus with its Spaaneprenneel yp Jong low snout somewhat

resembles Scymnognathus whaitsi, and its ver y broad interorbita]

and intertemporal surfaces agree soratlh Glorgonops. It is, however,

advanced in the following characters :— The loss of the step in

the jaw, the rounded snout, and especially the vertical occiput.

Haughton has pointed out another advanced feature in the

structure of the basicvanial region. Seylacops is a small unusual

form with a low broad snout: it is advanced in the exclusion of

the frontal from the orbital margin, in the rather vertical occi-

put, in the loss of the step in the upper Jaw, and especially in the

loss of the anterior ramus of the pterygoid: it appears to retain

a rather primitive Gorgonops-like nose and has only small

temporal fossee. Broom’s figure of the occiput suggests that it

is advanced in the shallowness of the paroccipital processes. [It

is probable that the fragment of a Gorgonopsid skull which I

described (Ann. & Mag. Nat. Hist. 1913, vol. xi. p. 65, figs. 1-4)

belongs, if not to Seylucops, at any rate to a clos ely ailucdl ton |

Arctognathus curvimola is a far more advanced form than any

so far discussed in this paper ; 1t pr comma ly comes from a higher

horizon in the Cistecephalus-zone than Gorgognathus, oe,” Thi

shows advances in the following features :—The snout is short,

narrower than the orbital region, rounded over the mid-line.

The nostrils are very Jarge and the septomaxillary foramen

smal]. The nasals do not overhang in front. The interorbital and

intertemporal surfaces are narrow, the orbits facing outwards,

CLASSIFICATION OF THE THERIODONITTA. 79

upwards, and a little forwards. The postfrontal is very

small, the preparietal is absent. ‘The maxilla is very short and

deep, there is no step in the upper jaw. ‘The prefrontal is short

and probably small. I have already shown that the basicranial

and otic regions are very advanced (Proc. Zool. Soc. 1914,

p- 1028). The epipterygoids are widened and flat, very much

as in Diademodon. The posterior end of the quadrate ramus of

the pterygoid no longer reaches the quadrate. The pterygo-

pavasphenoidal girder, instead of having a flat ventral surface

from which a deep median crest rises, is rounded, with the crest

represented by a low fillet.

The palate is most conveniently compared with that of

Gorgonops, which represents an earlier stage leading to it.

The great median groove is deepened and its roof is entirely

formed by a median vomer, which presumably represents a

forward growth of the posterior median bone of the Gorgonops

palate. Anteriorly the ridge rising from the vomer in <Arcto-

gnathus suggests that there was a soft secondary palate into

which a secondary bony plate may have grown out in the

concealed part of the palate.

The tooth-bearing roughened area of the anterior ramus of

the pterygoid hes much further back than in Gor gonops, and the

pterygoid no longer reaches the posterior nares.

The quadrate of dArctognathus is much smaller than that of

Scymnognathus.

The conversion of Aretognathus into a Cynodont Tike 0 yno-

gnathus demands only the following changes :—Still further thin-

ning of the basis cranil, further reduction of the quadrate wing of

the pterygoid, the development of a connection between the

quadrate wing of the epipterygoid and the paroccipital ; further

retraction of ane unterior ramus of the pterygoid, so as to reduce

the roughened areas to a pair of small knobs on each side of the

posterior end of the median groove; a little reduction of the

ectopterygoid; the development of secondary plates from the

maxille and palatines in the existing soft secondary palate ;

the conversion of the narrow intertemporal area into a sagittal

crest, to increase the length of the temporal muscles; the loss

of the postfrontal, and a further reduction of the frontal and

prefrontal, leading to an increase in size of the posterior part of

the nasals. ‘These changes are all in the same direction as those

which convert a Pelycosaur like Varanosaurus into a Theriodont

like Gorgonops, and an animal like Gorgonops into a form like

Arctognathus, and are, on the whole, smaller than those which ave

necessary to carry out the earlier improvements ; in fact, Arcto-

gnathus, which is technically a Gorgonopsid, is structurally closer

to Cynognathus than it is to Gorgonops.

Amongst other advanced forms allied to the Gorgonopsids and

coming from the Cistecephalus-zone are Cynosuchus and Whaitsia,

which have both been excellently described, though not com-

pletely figured, by Haughton.

80 MR. D. M. S. WATSON ON THE

Whaitsia is a remarkable form with a somewhat primitive

basicranial region, and a palate which in general agrees with

Gorgonops, but differs in the development of a special process

passing out on each side of the internarial bar so as to divide

each posterior nostril into two. {The meaning, morphology, and

function of this arrangement are quite uncertain, the anterior

vacuities are not exactly homologous with the anterior palatine

incisions of mammals and Cynodonts, because the posterior

border of these incisions is always formed by the anterior edge

of the secondary plate of the maxilla.| Whactsia is very advanced

in the reduction of the wide parietal region to a narrow sagittal

crest, which characterizes it, and in the extreme reduction of the

dentition.

The preceding discussion shows that the Gorgonopsids include

a series of forms which exhibit in their skulls a gradual series of

changes by which so primitive an animal as Arctops passes in-

sensibly into a Cynognathid. It establishes clearly the existence

of a series of evolutionary trends, which persist without change

from the beginning of the Anomodonts in Varanosaurus to their

end in Diademodon, and indeed to lead on to mammals. It

remains to discuss the other primitive Theriodonts included in

Broom’s order Therocephalia and the Deinocephalia, to see how

far these evolutionary trends apply also to them, and to consider

the relation of these forms to the Gorgonopsids, which are

plainly the central group of the ‘Theriodonts.

No Therocephalian is at all well known, despite the very large

number of forms which have been described. We know the

dorsal and lateral surfaces of the skull in a good many forms

(Seylacosaurus, Lycosaurus, Scaloposaurus, etc.), the palate 1s

known more or less completely in others (Scylacosaurus, Seyn-

nosaurus, Scaloposaurus, ete.). The basicranial region is known

in no Therocephalian, neither has any occiput been figured.

Haughton has described the brain-case of Alopecognathus, but

his figure is not in all points (e.g., the character of the supra-

occipital and the relations of the interparietal and parietal) very

convincing.

The most important materials of Therocephala in the British

Museum are the more or less complete skulls of Scaloposaurus

from the Cistecephalus-zone and Scylocosaurus and Seymno-

saurus watsoni from the T'apinocephalus-zone.

ScyMNOSAURUS WAtTSONT Broom, Proc. Zool. Soc. 1915, p. 169,

fig. 6.

Lycosuchus? Watson, Proc. Zool. Soc. 1914, p. 10386, fig. 7.

Type: a skull with seven cervical vertebre in natural articu-

lation, other vertebre and fragmentary limbs doubtfully

associated. Tapinocephalus-zone, Uitkyk, Dist. Prince Albert,

Cape Province.

The skull of the type is curiously preserved: it is embedded in

a caleaveous nodule, which breaks with a conchoidal fracture and

CLASSIFICATION OF THE THERIODONTIA, 81

is so loaded with a very fine siliceous mud as to be glass-hard,

completely blunting a carefully hard-tempered chisel after a

single blow.

The skull is broken through along the plane of the palate, part

of that structure adhering to each block. ‘he skull is then

broken through by a split which passes through the brain-cavity

and removes the whole left posterior corner of the skull behind

the orbit. Finally, another spht traverses the occiput, part of

that region adhering to each surface.

Where weathering has softened the matrix, very good pre-

parations are easily made, and in these regions, especially where

Text-figure 25.

Qu. inlooe Oe. 1 B.de Civek

Scymnosaurus watsoni Broom. Type-skull.

Occipital aspect. 4.

the bone has been cleaned by weathering, the preservation 1s

extraordinarily good. Development of the unweathered regions

is a very ditticult and extremely slow and tedious process. Never-

theless, I have been able to make a satisfactory preparation of the

inner surface of the cranial cavity.

The occiput now shows nearly every detail of its structure,

although a direct view of its posterior surface cannot be seen.

Its outline is well shown and the posterior surfaces of the quad-

rates and squamosals ave clean,

Dr. Broom’s figures give a good idea of the general shape, the

structure of the dorsal and lateral aspects not being shown.

Proc. Zoo. Soc.—1921, No. VI. 6

82 MR. D. M. 8, WATSON ON THR

My former figure gives a good idea of the palate, whose struc-

ture is well shown. It, however, does not clearly illustrate the

exact mode of articulation of the quadrate to the squamosal and

the structure of the extreme postero-lateral corner of the skull

and the auditory groove. This inaccuracy, which is not of a very

serious nature, was due to a misunderstanding by about 30° of

the orientation of the detached left corner, which was at that

time the better exposed.

The basioccipital condyle is largely concealed by the attached

atlas, but is partly exposed from below and cut by fractures which

give sections through it. It is rounded and nearly twice as wide

as itis deep. It is shortand immediately in front of it, at a plane

a little in front of the general occipital surface, lie the very

broad and massive basisphenoidal tubera. These have a flat

posterior surface overhung by the projecting exoccipitals, which

are separated from them by the small vagal foramen and per-

forated by small foramina for the XIIth nerves. The structure

of the anterior part of the basisphenoid has already been described.

The paroccipital process is very massive; not only is it thick

from back to front, but the small post-temporal fossa is placed

high up so that the process is deep. Although the fenestra ovalis

is not visible there can be no doubt, from consideration of the

general structure, that it lies far out.

Very little of the supraoccipital is visible from behind, the inter-

parietal terminating only a short distance above the foramen

magnum.

The joint supraoceipital and interparietal form a very thick

mass whose posterior surface stands nearly vertical.

The occiput, as a whole, forms an equilateral triangle with an

angle at the top. ‘l'o the lower parts of the lateral sides of this

triangle two others are added, standing out as fins; these are

composed of those parts of the squamosals which articulated with

the jugals.

The Jower and median parts of the main triangle are flat and

stand vertically, the lateral borders are turned back, so that

viewed as a whole the occiput is deeply cupped, the back- turned

margins gradually approach one another and, finally, fuse to form

the very deep sagittal crest.

The extreme upper part of the occiput is formed by the

parietals, the interparietal terminating far below the summit.

Laterally the parietals are covered by the tabulars, which form

the margin of the occiput for some distance, strengthened by

production of the parietals along their anterior faces and more

laterally by a similar covering of squamosals, which, indeed, over-

lap onto the parietals.

The squamosals articulate, as just described, with the parietals

and tabulars, and then extend outward into powerful processes,

their upper parts being turned backward so as greatly to increase

the size of the dorsal opening of the temporal fossze.

The ventral halves of their posterior surfaces are vertical,

CLASSIFICATION OF THE THERIODONTIA. 83

continuing the plane of the paroccipital processes, with whose ends

they are rigidly articulated. The posterior surface of the lower

part of the squamosal is separated from that of the paroccipital

by the usual ridge, which borders a groove for the external

auditory meatus.

This ridge continues upwards until, just below the level of the

post-temporal fossa, its hinder margin is turned inwards as

a seroll with a thickened edge in a very unusual manner, not

understood by me when I published my figure of the palate.

Text-figure 24.

Scymnosaurus watsoni Broom. Type-skull.

: 5 ; ; 2 ge ern Meter ety

Brain-case in sagittal section seen from the left side. Hpipterygoid of left

; i P .

side seen in outer view. 3.

J.Av.M., internal auditory meatus.

This scroll overhangs the occiput, but very rapidly subsides

into the general surface of the squamosal, just at the point where

that bone begins to bend backward. _

The lower edge of the squamosal is thin and is splht by two

notches associated with the attachment of the quadrate.

The quadrate, or in all probability the fused quadrate and quad-

vato-jugal, is relatively small, nothing of its hinder cae being

)

4 MR. D. M. S. WATSON ON ‘THE

visible except the extreme articular edge and the two processes

which interlock with the squamosal. There is evidence that it

is no higher than the paroccipital process, but its upper edge and

front face are not seen. The bone extends only very slightly

laterally of the outer process, forming the extreme end of the

lower margin of the occiput. Above its end the border of the

squamosal runs upward and slightly outward, its front face being

supported by the jugal.

The plan of this very remarkable occiput is repeated in a less

exaggerated form in Scylacosaurus sclateri, so far as the still very

incomplete preparation of the British Museum skull allows of a

comparison.

The brain-cavity of Scymnosaurus behind the epipterygoid is

now fairly well exposed. Its general characters will be best under-

stood from text-fig. 24. The foramen magnum is extremely

small, but the brain- cavity is somewhat latger than would be

anticipated ; ; although not high it is fairly broad, especially in

advance of the exit ok the vagus. It is of normal Anomodont

type with an opening to the inner ear placed very low down and

with this opening confluent with the foramen for the Xth nerve.

The sunken edge which separates these openings is continued

upwards by a ridge on the wall of the brain-cavity, which

separates the narrow medullary from the wider cerebellar region.

The pituitary fossa, although not cleared of matrix, is undoubtedly

shallow. There is a powerful process below the notch for the

Vth nerve, and the supraoccipital, with possibly a strip of the

pro-otie, extends forward as side-walls as far as the epipterygoid,

passing median of that bone.

The limb-bones doubtfully associated with the type-skull are

small and very slender, so that if they really belong to it the

proportions of the animal must have been like Hycenodon. It

they do belong the agility which they imply may be the ex-

planation of the unexpectedly large cerebellar cavity, which,

however, shows no trace of floecular fossee.

It is interesting to compare Scymnosaurus with Scymncgnathus,

which is of about the same size, though later in time. The two

animals are carnivorous and have very similar dentition, especially

in the feebbe molay series.

The Therocephahan is the more advanced in the following

characters :—

1. The reduction of all parts lying below the base of the

brain, the basioccipital, basisphenoid, and especially

the quadrates.

The reduction of the intertemporal region te a narrow

sagittal crest.

3. The shortening of the snout.

4. The lengthening of the temporal fossee.

Seymnosaurus rather vecalls Seymnognathus in its square-cut

snout, Scylacoswurus is much more advanced in the rounding of

CLASSIFICATION OF THE THERIODONTIA. 85

the dorsal surface of the nose, in both forms the extreme lowness

of the face in early Gorgonopsids is lost, the maxilla being deep

and the anterior part of the skull in general high.

The incisor-teeth in Scylacosaurus are small and the premaxilla

shallow below the nostril.

There is no step between the canine and the incisors, the teeth

forming a curved row like that of the latest Gorgonopsids and the

Cynognathids.

A preparietal appears to be lacking in Therocephalia, other-

wise the interorbital region does not difter greatly from that of

Gorgonopsids.

In the palate the Gorgonopsids are all more primitive than the

two Therocephalians in not possessing a suborbital fossa. They

are, however, all far more advanced in their possession of the

vaulted palate, which leads so directly to the development of

a secondary palate, and in the complete suppression of an inter-

pterygoid fossa.

The median part of the palate of Seymnosaurus even projects

slightly above the general level, that of Seylacosaurus is essen-

tially flat.

Scymnosaurus shows an advance on Scylacosaurus in the median

vomer which appears on the palate.

Both Therocephalians agree with one another in certain special

features, such as the extent to which the ectopterygoid con-

tributes to the pterygoid flange (in which they differ from the

Gorgonopsids); and in their general appearance, in the structure

of the occiput, ete., they in no way recall the Cynognathids, as do

all the Gorgonopsids dealt with in the preceding parts of this

paper.

Scymnosaurus and Seylacosaurus are Tapinocephalus-zone

forms, and there is no evidence of any animals with similar

structure in the succeeding Hndothiodon- and Cistecephalus-

zones.

No certain Therocephalian is known in the Hndothiodon-zone

(as I understand it), unless Broom’s /etidognathus is of that age.

In the Cistecephalus-zone Scaloposaurus, represented only “by

the type-skull from Stylkrantz, is the only satisfactorily preserved

form. It has been well described and figured by Owen and Broom,

whose accounts should be referred to. The little skull differs

exeeedingly from Scymnosaurus, the temporal fosse are short,

there are no pronounced sagittal and lambdoid crests, the squamo-

sals are not expanded, and the postorbital apparently does not

‘reach the jugal behind the orbit. {This may be only on account

of weathering of the surface. |

On the palate there is evident a wide interpterygoid vacuity,

agreeing with tle much narrower opening in Scymnosawrus; there

are large suborbital vacuities as in that form. Nothing is shown

of the anterior part of the palate, nor are the details of the basis

cranii well displayed.

Thus the extant material of Therocephalia sheds no light on

86 MR. D. M. S. WATSON ON THE

the evolution of that group, and the known forms are so few that

no classification is possible.

Tt is, I think, reasonable to assume that the Therocephalia

have sprung from the Gorgonopsid stock, and that they represent

a series of distinct branches which display a much more rapid

advance in structure than the conservative main stem. These

advances are, on the whole, along the trends of Gorgonopsid evo-

lution, quite early Therocephalians thus agreeing in certain

features with Cynognathids. It is, as I have already shown,

probable that Bauria and its allies are descendants of Thero-

cephalia, representing the product of a parallel series of changes

to that which resulted in Cynognathwus imposed on a different

ancestor.

Im the preceding part of this paper, I have dealt only with a

selected series of Gorgonopsids which present resemblances to the

Cynognathids, and have tacitly assumed that these forms are

the main stock. There are, however, many other Gorgonopsids

which appear to represent side-branches, displaying either

accelerated evolution of certain features or else individual

specialisations.

Of these forms the earliest and one of the best known is #luro-

saurus felinus Owen. This form was first described by Owen,

Q.J.G.S. vol. xxxvil. p. 261, pl. ix., Seeley subsequently figuring

an incompletely prepared palate. Broom later corrected certain

features of Seeley’s description of the side of the skull.

The British Museum includes, in addition to the type, two

snouts which were regarded by Lydekker as Z. felinus. Broom

left manuscript-labels concurring in the identification, and

a detailed examination which I made of them showed that the

external surface and dentition are in complete agreement. One

of the specimens had no lower jaw attached and the palate has

been developed, with the remarkable result that it is shown to

differ very considerably from the type, being probably generically

distinct. The whole circumstance is of importance, because it

shows that a specimen showing only the outer surface of the

snout and dentition of a Gorgonopsid may be an inadequate type.

In text-fig. 25 I give three slightly reconstructed views of the

snout of B.M.N.H. R.855 from the Hndothiodon-zone? of Beau-

fort West. This type is very advanced in the depth and rounded

section of the snout, in the supression of a step in the maxilla

before the canine, and i in the relatively slight overhang of the

anterior border of the nasal. It retains a very large septo-

maxillary foramen and a large facial exposure of the septomaxilla.

The palate unfortunately shows no sutures, but gives a good view

of the general form. In general form this palit differs S very

considerably from that of Arctops, Gorgonops, Scymnognathus, ete.

The median region is excavated into a very narrow groove

bounded laterally by massive processes, whose palatal surfaces

bear small teeth in sockets. Lateral to the process is a small

fenestra or possibly a very deep pit with a well-defined margin ;

CLASSIFICATION OF THE THERIODONTIA, 87

further laterally the palatine forms a deep groove, flattening

out as itis traced forward until it becomes the nearly vertical

surface of that anterior part of the bone which bounds the outer

side of the posterior nares. The pterygoid flanges are powerful,

but differ from the ordinary Gorgonopsid type in that the ecto-

pterygoids extend down to their summits. There is a single inter-

narial bar whose ridged lower surface lies far above the lower edge

of the maxilla.

Text-figure 25.

DS Poy.

—

7 NN

" = eID LNB.

meu

Mx. io

Sai

Ailurosaurid, ? gen. et sp.

Dorsal, right lateral, and palatal views of the anterior part of a skull.

R. 855, B.M.N.H. X 3.

The palate is of the same type as that of Hlurosaurus felinus,

but differs in the much smaller development of tooth-bearing

areas and in the much more caudal position of the hinder ends

of the posterior nares.

An analysis of the structure presented by a series of animals

belonging to the Theriodontia thus suggests that that group is a

88 ON THE CLASSIFICATION. OF THE THERIODONTIA.

natural one, the conservative main stem which leads on to the

Cynognathids being represented by a series of Gorgonopsids of

which Gorgonops itself is one of the most primitive members.

From this stem side-branches arise, which retain the broad

parietal region and other primitive features, but present either

an accelerated development of certain regions or are indi-

vidually specialised. From still earlier members of the main

stem arose the groups of animals, resembling one another in

the precocious conversion of the broad intertemporal region into

a Sagittal crest and in the acquirement of suborbital vacuities,

which are usually included in the Therocephalia and belong to

many independent stirps, each in all probability being dependent

on the main Gorgonopsid stock. It is shown that there is a

series of evolutionary trends which persist throughout the whole

group of Anomodonts from Varanosaurus to Diademodon, and

that the special rapid advances which separate the Therocephalia

from the Gorgonopsids, in the main, merely follow out the pre-

determined evolutionary track proper to the group.

Thus any classification of the Theriodontia is necessarily

complicated, as involved and difficult of construction as that of

the Theria themselves. Existing material is so incomplete that

any attempt at detailed classification, even if only into families,

is dangerous, in that it will load the literature with undefined

groups, whose characteristic forms may only be known from the

front end of the skull or the dentition.

The detailed descriptions of skull-structures in this paper show

how unreliable, even for generic distinction, are the characters

presented by the teeth of Theriodonts.

Thus, for the present, | am inclined to retain my former

division of Theriodontia into Gorgonopside, Therocephalide,

Cynognathide, and Bauromorpha, fully recognising that these

groups—or, at any rate, the first two—cover a multitude of forms

not directly of common origin and only held together by two

or three striking characters.

It remains to discuss the connections of the Theriodontia

with the other groups of South African Anomodonts—the

Deinocephalia, Dromosauria, and Dicynodonts.

In the copper-bearing sandstones and associated limestones of

the Ural Mountains, which immediately succeed the Artinsk

beds and are shown by a comparison of reptile and amphibian

faunas to be slightly older than the Tapinocephalus-zone, are

found three types of Anomodonts, each represented by skulls or

jaws: of these Deuterosaurus is clearly a Deinocephalian of the

Tapinocephaloid group recalling many South African forms.

Deinosaurus (= Cliorhizodon) is represented by jaws, in one

case associated with a palate whose dorsal surface is well exposed.

Rhopalodon is a name covering not only several jaw-fragments

but also a complete skull, which was described by Prof. Seeley.

Of this skull, remarkably beautiful lithographic drawings of the

‘.

‘

‘ , rls es

foe h aa aN POM AON

i ~, =\\

t \ rare iak

H EAEN H SS \ U ~\

lL We 1 SS ‘ \

| A) AS Hi Gn

H ie NE | ‘ 1

Loy tf wa -

te ay is > + 0)

Soon eg

po ceeweses Sean e apse Se = ee Se ee eee ee el

oeesesfO

WSS

‘ TT

; |

ES 5

Rhopalodon fischeri Kutorga.

Restoration of skull from the figures published by Seeley, Phil. Trans. B. 185, 1894.

and the brain-case figured by von Meyer, ‘ Paleontographica.’

Parts in broken lines hypothetical, sutures in dotted line suggested by the

original figures.

90 MR. D. M. 8. WATSON ON THE

dorsal surface and right side were published by Seeley, and that

author also gave lee intelligible figures of the palate and of

the much damaged occiput.

This skull The the lower jaw in position, and except for the loss

of the end of the snout appears to be (as are the other bones

from the same rocks) very well preserved and undistorted.

The skull itself is in Russia and quite unreachable, but in the

light of our present knowledge of Anomodont structure it is

possible by a careful study of the drawings and of Prof. Seeley’s

description to gain a clear idea of its more important features.

In text-fig. 26 I have drawn four reconstructions of this skull

on the indications available.

For the occiput I have used that figured by von Meyer as

Deuterosaurus, which cannot belong to that genus because it is

Text-figure 27.

Brain-case of Rhopalodon ?

Right lateral aspect. X 4.

From a cast in the British Museum of the specimen figured by von Meyer

as Deuterosaurus.

only half the size of that in the skull figured by Seeley, and does

not appear to agree at all in structure. It is, on the other hand

of very nearly the same size as the occiput of Seeley’s Rhopalodon’

skull, shows no features incompatible with the wreck of that

region in this skull, and must presumably belong either to

Rhopalodon or the very similar Deinosaurus.

Rhopalodon at once recalls the Pelycosaurs in appearance and

in certain structural features. It has a high compressed snout

passing backward into a square-cut lachrymal region, with a

depression overhung by a projecting ridge on the pr efrontal, just

as in Dimetrodon. The jugal in its shape at once recalls that of

the earlier genus. It differs from Dimetrodon in the much

CLASSIFICATION OF THE THERIODON'TTA, on

larger temporal fosse, visible from above, in the outward bowing

of the zygoma, in the vertical occiput, and in the smaller

lachrymal—all changes which follow the ordinary trends of

Theriodont development.

The neural cranium, as shown in von Meyer’s figures and in

text-fig. 27, is Pelycosaur-like in general build, and especially in

the complete absence of that forward growth of the supraoccipital

and pro-otic above the notch for the Vth nerve which occurs in

Theriodonts, and in the occurrence of a special notch for a vem

above the incisura prooticis.

This occiput is, however, specialised in the development of a

mass of bone below the basioccipital condyle, which is presumably

associated with a very vertically placed fenestra ovalis. The

development of this plate is the explanation of the extreme

depth of the pituitary fossa.

The palate of Rhopalodon recalls that of Dimetrodon in its

massive flanges on the pterygoids and in the row of teeth which

decks them. The very large internal nares also recall certain

advanced Pelycosaurs.

In certain ways the skull of Rhopalodon resembles that of the

more primitive Gorgonopsids ; the dorsal surface of the skull, for

example, is very like that of Gorgognathus. ‘The side view

differs, however, in the great depth of the snout and in the

relatively powerful molar dentition.

Although Seeley’s figures are not very readily interpreted in

that region, it seems that the quadrate of Rhopalodon is large

and well exposed from behind, and that its outer edge lies on the

outer surface of the skull exactly as in the South African

Deinocephalia, and not at all as in the Gorgonopsids, although it

represents a state from which that in the latter group could

readily be derived. The brain-case of Rhopalodon differs from

that of any Gorgonopsid in the vertical plate below the condyle

and in the non-extension of the supraoccipital, etc., forward.

It is unfortunate that no part of the palate behind the flange

is preserved, but judging from the front of the fragment of basi-

sphenoid preserved in the occiput, and the general structure, there

can have been no narrow bar separating the subtemporal fosse

as in Gorgonopsids, but the conditions must have been more as in

the South African Deinocephalian Mormosaurus. There is, in

fact, no doubt that Rhopalodon is not a primitive Gorgonopsid,

but is a primitive Deinocephalian, with the members of which

group it agrees in all the characters in which it differs from the

Theriodonts. Its general resemblance to primitive ‘Theriodonts

suggests, however, that we are very near the point of separation

of these two orders.

The present seems a suitable opportunity for adding to the

description which I gave (Proc. Zool. Soc. 1914, p. 770, ete.) of a

skull referred to Vitanosuchus. I recently found a block fitting

on to the fragment of maxilla of that specimen which contains

the anterior end of the maxilla, parts of the premaxille, septo-

MR. D. M. 8. WATSON ON THE

maxilla, ete. This addition shows that there are only three

premaxillary teeth, that the end of the snout is longer than in

. . a 2 . . [a al =

the original figure, and that it is rounded. Thus the skull

belongs to a new genus and may be called Anteosaurus magnificus,

gen. et sp. nov., holotype R. 3595, B.M.N.H.

Text-figure 28.

1 4b

=| \

= Th

=f Yy Mx.

B ‘V4 ZG) y

S \2

S 7% Fa.

Se 74

Right lateral and palatal views of the anterior part of the skull of the holotype

of Anteosaurus magnificus, gen. et sp. n.

Parts in broken lines restored, areas surrounded by thin irregular lines

present in the specimen. XX 4.

The general structure will be best understood from text-fig. 28.

The septomaxilla is a small bone lying within the nostril, to

which it forms a floor, passing inward nearly to the middle line.

CLASSIFICATION OF THE THERIODONTIA. 93

It articulates with the nasal posteriorly, with the maxilla in the

middle, there being apparently no septomaxillary foramen, and

with the premaxilla in front.

The palatine extends forward to the middle of the canine

tooth, forming the greater part of the outer wall of the long

narrow posterior nares; at the hinder end of the opening it has a

suture with the prevomer, which appears to form the whole

inner border of the nostril.

The prevomers are unfused, each provided with a high, thin

dorsal ridge, similar flanges from the pterygoids passing between

and separating those of the prevomers.

The whole structure is like that of the ‘apinocephaloid

Mormosaurus, and especially like that of the detached nose which

seems to belong to Laniasaurus.

The structure of these palates raises doubts as to the forma-

tion of the internarial bar in Gorgonopsids by the fusion of a

pair of prevomers, because in the Deinocephalia the pterygoids

separate the posterior ends of the prevomers, whilst in Gorgo-

nopsids they clasp the outer sides of the posterior ends of the

internarial bar. The difficulty is not, however, an insuperable

one.

The relation of the Dicynodonts to other Anomodonts is a

subject ou which there has been much difference of opinion, but

which can be more satisfactorily discussed now that many details

of Gorgonopsid structure are known.

The characteristic features of all Dicynodonts are :—

1. The occipital condyle is triple, the exoccipitals forming

its upper parts.

2. The supraoccipital is only slightly drawn forward to

form side-walls to the brain-case.

3. The fenestra ovalis lies at the end of a long tube com-

municating with the vestibule.

4, The temporal fosse are very large.

5. The face is short.

6. The premaxille are edentulous and the maxilla is carried

out laterally to the molar teeth, if any be present, and

its margin is a sharp ridge covered by a horny sheath.

7. There is a rudimentary secondary palate.

8. The prevomers are fused, forming a roof to the depressed

median part of the palate.

9. There are no definite pterygoid flanges.

10. There is an interpterygoid vacuity reaching back to the

basipterygoid process and forward to the prevomer.

11. Both quadrate and quadrato-jugal form the articular

condyle for the lower jaw.

12. The squamosal is of characteristic shape with a wide flat

zygomatic part rising from the upper part of a flat body,

the lower part of whose front face is covered by the

quadrate and quadrato-jugal.

13. The dentaries are fused and extremely massive.

94 MR. D. M. S. WATSON ON THE

When the structure of the skull of a Gorgonopsid was dis-

covered, Broom and I independently pointed out the many

resemblances which it presented to Dicynodonts in the inter-

temporal, basicranial, and other regions.

These resemblances are real, but, with the exception (?) of the

occurrence of a preparietal, lie entirely in the common possession

of primitive Anomodont characters, such as the broad parietal

surface and the main features of the basis cranii. When con-

sidered in more detail, the structure of such a Dicynodont as

Endothiodon seems to show no such resemblance to that of a

Gorgonopsid as to imply any closer connection between the

two groups than either of them bears to the Deinocephalia

or Dromosauria. The secondary palate of Hndothiodon is dif-

ferent in type from that of Diademodon and all the forms of

Gorgonopsids ieading up to it. In them, as in mammals, the

original vaulting of the palate is brought about by a down-

growth of the tooth-bearing edge of the maxilla below that of

the premaxilla ; the internarial bar remains attached to the

ends of the palatal processes of the premaxille, and when the

secondary plates ‘of the maxille grow out they lie ventral to

the palatal parts of the premaxille. In fact, the original surface

of the palate lies on the roof of the naso-pharyngneal duct, the

sides of the palate growing down below it.

In Endothiodon (text-fig. 29), on the other hand, the palatal

surface of the premaxille lie in the same plane as that of the

maxille and transverse bones, the posterior nares open into a

deep groove, excavated in the original palate, which is roofed

by a great forward growth of the prevomers over the region

formerly occupied by the large posterior nares.

Thus it appears that the palate of Dicynodonts does not

present a real resemblance to that of Gorgonopsids, but repre-

sents a different mode of development of a secondary palate,

identical with that of a Chelonian. One of the most striking

features of the Dicynodont palate is the loss of the pterygoid

flanges, which are only represented by slight eminences on the

edge of the narrow posterior part of the palate over the pterygo-

transverse suture. ‘[his loss seems to be due to the very great

expansion of the temporal muscles squeezing them out of exist-

ence, their presence not being necessary to insure accurate closure

of the mouth in an animal without a closely-fitting dentition—in

some cases their function being taken over by the long canines,

which often present wear-faults on their inner sides in large

Dicynodonts.

When allowance is made for the changes resulting from the

development of the horny covering of the edges of the jaws and

the extension of the crushing palate, it is readily seen that the

main plan of the anterior part of the Andothiodon palate is

reducible to that found in Therocephalia, except for the absence

of suborbital fenestree.

Posteriorly, however, there is in Dicynodonts no trace of the

CLASSIFICATION OF THE THERIODONTIA, 95

long narrow girder which in the Theriodonts extends from the

basipterygoid processes to the flanges of the pterygoids, and it is

therefore probable that the Dicynodonts separated from the

Theriodonts before the establishment of this feature—that ?s, at

about the same time as the Deinocephalia. At the same time, in

Text-figure 29.

Palate of Endothiodon ? microps Broom.

Drawn from R, 4044, B.M.N.H.; the quadrates, quadrato-jugals and jugals,

and quadrate rami of the pterygoids, from other material.

the Dicynodonts as in the Deinocephalia and all other 8. African

Anomodontia in which the facts are known, the descending

flanges of the pterygoids lie well in advance of the basipterygoid

processes, a position which is markedly different from that found

96 MR. D. M. S. WATSON ON THE

in all non-mammal-like reptiles, but which has arisen within

the group, as it does not occur in Varanosaurus.

The relation of the Theriodontia to the Dromosauria 1s obscure,

because of our very slight knowledge of the detailed skull-strue-

ture of the members of that group. In general build the

Dromosaur skull differs from that of the Theriodonts and also

of the Pelycosaurs in the extreme shortness of the face. In this

feature, and also in the depth of the squamosal below the root

of the zygoma, it recalls the Dicynodonts, differing from them,

however, in retaining a very short temporal fossa. The Dromo-

saur skull recalls that of Bolosaurus, and also still more strongly

that of Palewobathria, it my interpretation of Crednev’s figures of

that animal be correct. There are in the structure of the

post-cranial skeleton no characters which show significant resem-

blances to any other South African form, and it seems not

improbable that the group represents the end members of an

Artinskian group which survived into Upper Permian times.

Thus, in my opinion, the three orders Deinocephalia, Dicyno-

dontia, and Theriodontia may have arisen from a common stock

whose direct conservative descendants are the Gorgonopsids,

and the Dromosauria may represent a more widely separated

stock of the Anomodontia.

One other line of argument which I have not before considered

is concerned with the dentition,

In a discussion of the Dimetrodon dentition on p. 70, 1 pointed

out that it is desirable in a carnivorous animal with an enlarged

canine towards, but not actually on, the front end of the lower

jaw to have enlarged premaxillary teeth before the diastema to

balance the large canine behind that gap. In such earlier

Gorgonopsids as Gorgonops and Scymnognathus, the incisors are

actually of great relative size, much bigger than the cheek-teeth,

though, of course, not rivalling the great canines.

In Deinocephalia, both Tapinocephaloids (e. g., Deuterosaurus)

and Titanosuchids (e. ¢., Anteoswurus), the incisors may be of very

great size, and in the former group very curiously converted into

effective crushing-teeth. Both these animals have an enlarged-

canine, followed in the one case by a single molar, in the other

by a row of eight very small teeth ; in Deinosaurus (Clior hizodon)

there is a single enlarged canine follow ed by ten smaller cheek-

teeth, all larger than the single incisor preserved.

Thus in this feature the Deinocephalia present a definite

resemblance to the Gorgonopsids, one which is shared also by

Dimetrodon.

In later Gorgonopsids, as I have shown above, the incisors

become relatively smaller, and the ‘“ molars,’ although not

increased in number above the original five, become relatively

larger.

There is some evidence of an irregular replacement of all the

teeth of a Gorgonopsid.

CLASSIFICATION OF THE THERIODONTIA. 97

When the Cynodontia are reached we find a still further

decrease in the relative importance of the incisors, together with

an increase in the number of cheek-teeth in Cynosuchus to 8, all

still much smaller than the incisors, in WVythosaurus to 8,

all larger than the incisors, Galesaurus 210, Cynognathus 9,

Diademodon ¢12, Trirachodon 9. This sudden inerease in the

number of cheek-teeth seems to be associated with the develop-

ment of a bony secondary palate, which allows of prolonged

mastication being carried on without obstruction of the naso-

pharyngeal passages.

It is important to note that the anterior five ‘* molars” of

Cynognathus crateronotus are sharply distinguished from those

which sueceed them, and that in Diademodon, when the four

anterior cheek-teeth are similarly distinguished by structure,

these teeth and these alone amongst the cheek-teeth give indica-

tions of replacement.

In fact, the evidence brought forward by Broom (Bull. Amer,

Mus. Nat. Hist. vol. xxxii. p. 465), although it is not quite so

conclusive as one could wish, does tend to show that these

Cynodonts had a thoroughly mammalian dentition with premolars

replacing milk-predecessors and molars never replaced.

Comparison with Gorgonopsids suggests that the premolars are

the original cheek-teeth, the molars representing a new backward

growth of the dental lamina, in which the teeth all belong to a

single generation, and from their origin have never had either

predecessors or successors.

I hope to return to a consideration of the whole problem of

dental succession In early reptiles shortly.

The Gorgonopsids being characterized throughout their history

by the possession of a very short series of molar teeth, we have to

consider the problem presented by the fact that whilst some

Therocephalia, e.g., Scymnosaurus and Hycwnosuchus, resemble

them in this feature, others, Alopecodon ete. amongst large forms

and Jcticephalus and Scaloposaurus amongst the small, retain

a large series of molars, without, so far as known, having a

secondary palate, and possess in addition small incisors.

It seems reasonable to regard these latter animals as unmodified,

the dentition being derived directly from their ancestors, whilst

Hycenosuchus represents a parallel reaction to that of the Gor-

gonopsids to similar feeding habits. All detailed discussion of

the dentitions of Theriodontia are rendered nugatory by our

complete absence of knowledge of the postcranial skeleton, for

only by a study of the whole structure is it possible seriously to

consider the habits and adaptations of an animal, and teeth react

perhaps more quickly than any other structures to external

impressions.

In any case the occurrence of these Therocephalians with a

long tooth-row and small incisors, and of a similar structure im

Olicrhizodon, shows that the heavy incisors of Gorgonopsids

Proc. Zoou. Sec.—1921, No. VII. 7

98 ON THE CLASSIFICATION OF THE THERIODONTIA.

and Deinocephalia have been independently acquired by these

two groups.

The study of Theriodont structure and evolution, which is the

eontent of this paper, thus leaves us still without any satisfactory

classification of that group, but in its establishment of a series of

evolutionary trends, which persist throughout the history of the

Anomodontia, has I hope laid a solid foundation on which a

natural arrangement may in future be built up when increased

knowledge allows an examination of the “ adaptive radiation ” of

the order to be entered on. Meanwhile, the forms whose skulls

are well known fill in with considerable completeness the great

morphological gap which exists between the Lower Permian

Pelycosauria and the Lower and Middle Triassic Cynodonts, and

in this way enable us to understand the material steps in the

evolution of almost all the structures of a Cynodont skull from

those in so primitive a reptile as Seymouria or, indeed, in the

still more primitive Embolomerous amphibia. Dealing as it does

with many diverse faces of the subject with which it is concerned,

this paper does not lend itself to summarisation, but the point of

widest interest brought out in it is undoubtedly the demonstration

of the occurrence of the same evolutionary trends in so many

allied branches in the Pelycosaurs, Gorgonopsids, Therucephalia,

and Deinocephalia; and the fact that the changes brought about

in accordance with these trends often serve an adaptive purpose

and appear to depend on mere mechanical necessities.

My thanks are due to the Percy Sladen Trustees for defraying

part of the expenses of my visits to South Africa and Texas.

To Dr. A. Smith Woodward and Dr. ©. W. Andrews I am

indebted for many facilities and much kindness in the British

Museum, and to the Department of Industrial and Scientific

Research I owe the stipend which has enabled me to carry out

this work.

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Bale & Danielsson, Ltd.

COLOUR- CHANGES OF THE SPOTTED SALAMANDER (SALAMANDRA MACULOSA).

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ON COLOUR-CHANGES OF THE SPOTTED SALAMANDER, 99

4, Hxperiments on Colour-changes of the Spotted Sala-

mander (Salamandra maculosa), conducted in the

Society’s Gardens. By E. G. Bounenerr, F.Z.S.

(Curator of Reptiles).

‘Received October 29, 1920; Read March 8, 1921. |

(Plates I. & II.: Text- figure 1.)

The remarkable experiments conducted in recent years at the

Biologische Versuchs-anstalt, Vienna, by Kammerer on the

action of modified environment on certain Batrachians have

attracted much attention, and a good deal of criticism. Prof. E.

W. MacBride, F.R.S., to whom I am indebted tor much help

and advice, suggested to me last year that an attempt to repeat

the experiments conducted by Kammerer with the Spotted

Salamander (Salamandra maculosa), on the correlation between

the coloration and the conditions under which Salamanders may

be subjected to, would be of special interest. According to

Kammerer, an increase of the bright colour takes place on a

yellow soil, especially when the animals are kept in terraria

coloured yellow in such a manner that yellow rays of light fall

on the Salamanders, whilst the reverse takes place when they

are kept in the dark on black humus. Further, he claimed to

have succeeded in the extremely difficult task of inducing his

Salamanders, which he kept under the above conditions, to breed

in captivity, and states that the offspring inherited the characters

of their parents, being yellower or blacker than the normal,

according to the surroundings in which the first generation had

been placed. During the past year I have been keeping under

conditions similar to those referred to by Kammerer a large

number of Salamanders received from the French Jura, thanks to

the kindness of Mme. Phisalix of the Pasteur Institute, but it

is as yet too early to expect any striking changes, and the

present paper deals with the results obtained by repeating

certain experiments conducted by Kammerer’s pupil, Seceroy,

who kept Salamander larve in aquaria under yellow or black

conditions *. Secerov unfortunately started his experiments with

only four larve, the offspring of a Salamander of the variety

teniata. Two of these larve were placed in an aquarium

coloured yellow and two in an aquarium coloured black. . The

results he obtained were as follows:

That Salamanders kept in yellow surroundings when in

the larval stage were on transformation yellower than their

mother: that the spots were larger and had a tendency to

fuse together.

That the reverse takes place when Salamander lnrvee are

kept in black surroundings, the transformed animal being

darker than the mother: that the spots are smaller and

more numerous.

# * Biologisches Centralblatt,’ 1914.

100 MR. E. G. BOULENGER ON COLOUR-CHANGES

In January of this year (1920), I was given a pregnant Sala-

mander of the variety twniata, which I was informed was

received from Western Germany. In this specimen (see text-

fig. 1), in which neither the black nor the yellow can be said

to predominate, the dorsal spots are confluent into broad bands

along each side.

On February 2nd this Salamander gave birth to 32 young,

and half the larvee were immediately placed in aquaria 16 inches

square, entirely painted orange-yellow, whilst the other half

were placed in similar aquaria painted black all over. In the

Text-figure 1.

course of their larval life 12 of these young Salamanders died,

7 of those placed in the yellow aquaria, and 5 of those in the

black aquaria. Of these, however, two in each set died only just

prior to transformation, having developed the yellow and biack

pigments of the perfect animal. It will be observed from the

figures on Plates I. & 11. that in these experiments the results

obtained by Secerov were fully confirmed, the majority of the Sala-

manders which had been kept in vellow surroundings when in

their larval condition being on transformation yellower than the

mother, and the spots larger and fused together : the Salamanders

OF THE SPOTTED SALAMANDER. 101

kept in black surroundings being on transformation darker than

the mother, and the spots smaller and more numerous.

To appreciate fully the markings on these Salamanders it is,

however, necessary to bear in mind the fact that it is possible

to define two forms of S. muculosa. As I pointed out in a paper

brought before this Society some years ago*, authors in dividing

this Salamander into a number of varieties have dwelt on

supposed structural characters, whilst ignoring the disposition

of the markings. I showed that two principal forms existed,

namely, the typical form and the variety teniata: the former, in

which the yellow appears as markings of various shapes disposed

anyhow over the body, being more or less an eastern form ; the

latter, in which the dorsal spots are regularly disposed in two

parallel series continuous with the markings on the parotoids,

being a more western form. The markings in the last referred

to variety may form regular bands or be broken up into

numerous small spots, which, however, always retain their

duplex disposition, not encroaching over the black vertebral!

area, or, if they do so, as is rarely the case, they will be

connected by a cross-bar in an H-shaped form. The habitat of

the forma typica appears to be bounded to the west by the Evz

Mountains, the Danube, the Alps, and the Rhone, Salamanders

from east and south of that line belonging to it. All over

France, west and north of the Rhone, Belgium, $. Holland,

Western Germany, Spain, and Portugal the variety teniata

alone occurs, with very rare exceptions. Both are found together

only on the line of demarcation, such as in the French Jura,

where the variety teniata predominates, but is not completely

fixed.

Even amongst Salamanders from the neighbourhood of the

line of demarcation it is rare to experience any difficulty in

classifying them into either of the two forms, and I was

therefore much puzzled by Secerov’s results, as neither of his

Salamanders kept during their larval life under yellow sur-

roundings were quite normal ¢eniata. One of these Salamanders

was specially abnormal, as two spots on the anterior part of the

body fused together over the vertebral area; and I should have

felt some doubt in referring it to the variety teniata, had I not

known the mother to be a very typical specimen of this form.

The results I have obtained help to solve the problem, as they

show that in the majority of cases, if the offspring of a Salamander

of the western variety teniata is kept when in the larval

conditions in yellow surroundings, the markings will differ in

their disposition from those of the parent, and be similar in

their arrangement to what we find in the typical eastern form,

in which the spots are disposed in an irregular fashion over the

vertebral area.

* “A Contribution to the Study of the Variation of the Spotted Salamander,”

P.Z.S. 1911,

102 ON COLOUR-CHANGES OF THE SPOTTED SALAMANDER.

Plate II. shows the nine Salamanders kept during aquatic life

in yellow surroundings on or just prior to their transformation,

and it will be observed that six are referable to the typical form,

two are somewhat doubtful, and one only could possibly be

regarded as belonging to the variety teniata. Plate I. shows

those kept in black aquaria, and it will be noticed that the spots

are disposed in parallel series, as would be expected from the

offspring of Salamanders of the striped variety.

On transformation the young of specimens of the variety

teniata from the French Jura, which is, as referred to above,

on the line of demarcation where both forms occur, are often

referable to the typical form; the spots, however, in the course

of time change their disposition, breaking up, and disappearing

on the vertebral area, so that within a year the Salamanders are

similar to their parents in the arrangement of their markings.

T have no doubt that we shall find this to be the case with the

Salamanders which in their larval form have been brought up im

yellow surroundings, and that the spots will in time assume

the same duplex disposition as presented by their parents,

and brethren brought up when in the larval form in black

surroundings.

ON INDIAN OLIGOCH ATLA. 103

9. Contributions to the Morphology, Classification, and

Zsogeography of Indian Oligocheta. By J. SrEPHEN-

son, D.Sc., F.Z.8., Lecturer in Zoology in the

University of Edinburgh.

[Received November 6, 1920: Read February 22, 1921.]

(Text-figure 1.)

CoNTENTS. Page

I. The ‘Affinities and Systematic Position of the Genus 103

Hudichogaster Mchisn., and some related questions ...

II. On Polyphyly in the Oligocheta... poe peli

III. Some general Considerations on the Geonmaplical Divine

butionkofalndiantOhpochsetanis..as.sssse eee een LAs

1.—The AFFINITIES AND Sysrematric Posrrion or THE GENUS

HUDICHOGASTER MCHLSN., AND SOME RELATED QUESTIONS.

The genus Hudichogaster was established in 1902 by Michael-

sen (2) for LH. ashworthi, then first made known, and for several

other worms which had been originally described as species of

Dichogastey (or Benhamia), but which were placed by Michaelsen

in the Tierreich volume of 1900 in the genus Zvrigaster. The

separation of these three genera is based on the presence or

absence and on the position of the calciferous glands; Vrigaster

has none, Hudichogaster has them in xi, and xil. ~ (with, ‘it may be,

3G, Oi iimle 1m addition), Dichogaster has them in xiv., xv. and Xvi.,

or in xv., xvi. and xvi. The diagnosis of the genus Eudichogaster

is as follows :—

Sete four pairs per segment. Prostatic pores two pairs on

Xvi, and Xix., or one pair on xvil. (? or xviii.). Spermathecal

pores two pair's ON Viil. and 1x., or one pair on viil., or in groove 7/8.

Two gizzards in front of the testis segments. Calciferous glands

two or three pairs, in xi. and Xii., or x., xi. and Xil., or Xi., xil.

and xiii. (in one species no proper calciferous glands recognizable).

Micronephridial. One or, more usually, two pairs of testes.

Prostates tubular.

In the paper in which the genus was established, Michaelsen

placed it in the Trigastrinz. In 1903, however (3), he leans to

the view that it is to be derived from Octochetus, and there-

fore to be included in the Octocheetinze, though he does not carry

out this implication in the tables. In 1909 (4) he definitely

adopts this view. In 1910 (5) he abandons it, and, deriving

Ewdichogaster from Trigaster, again places it in the Ty igastrine.

The object of the present communication 1s, by bringing - forward

additional evidence, to decide the question in favour of its

inclusion in the Octochetine, by showing that it is descended

from Octochetus, not from 7Z'rigaster.

104 DR. J. SLEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,

The classification of the Megascolecide, to which both sub-

families, the Octochetine and 'Trigastrine, belong, proceeds on

phylogenetic lines; and a very considerable degree of success has

been reached in the filiation, and consequently in the definition

and arrangement of genera. The origin of the family, as is now

generally recognized, is to be sought in the “ original Acantho-

driline,” a form which is represented at the present day by the

genus ‘Wotiodrilus as defined by Michaelsen in the Tierreich

volume (1). Its essential characters (for our present purpose) are

as follows: .A pair of male pores on xvill., two pairs of prostates

opening separately on xvii. and xix., a single cesophageal gizzard,

four pairs of sete per segment, one pair of meganephridia per

segment, no calciferous glands.

The important characters of the genera which enter into the

following discussion may be stated thus :——

Diplocardia, one remove from the original Acanthodriline, and

the ancestor of the Trigastrine, differs from the original Acan-

thodriline only in having two gizzards. It is found in North

and Central America.

Prigaster, descended from Diplocardia, differs from the latter

genus in being micronephridial ; the posterior male organs are

either of the scanthodriline type or they may have undergone

the ‘* microscolecine reduction ” (disappearance of the posterior

pair of prostates, and union of the male pores with the anterior

prostatic openings on xvil.); there are two or three gizzards, but

no calciferous glands. Zrigaster is found in Mexico and the

West Indies.

Dichogaster is similar to Trigaster, except that there are

calciferous glands in two or three of the segments xiv.-xvu. A

point to which attention has been called is that while in Vrigaster

the setal interval cd is greater than ab, in Dichogaster these

intervals are approximately equal. Hudichogaster was supposed

to resemble Z7igaster in this respect, but in a few species ab 1s

equal to ed. Dichogaster is endemic in tropical Africa, and

probably in Central America and the West Indies, but has spread

widely in the tropics and warmer temperate ZONES.

The first development, therefore, in the above line of descent

(Diplocardia-Trigaster- -Dichogaster), aod the primary charac-

teristic of the Trigastrine (I include Diplocardia in the

Trigastrine, v. post.), is the reduplication of the gizzard.

The genus Octochetus belongs to a different line. It differs

from the original Acanthodriline in being micronephridial, and

in having a pair of calciferous glands in segment KV 10) samen!

few species, to which more particular reference is made subse-

quently, are without calciferous glands); it has the single gizzard

and other characters of the ancestral form. It occurs in India

and New Zealand.

The other genera of the subfamily need not be mentioned here.

The first development in the Octochztine, and therefore their

AND ZGOGEOGRAPHY OF INDIAN OLIGOCHATA. 105

primary characteristic, is the splitting up of the nephridial

system.

Original Acanthodriline

(single gizzard ; meganephridia ;

no calciferous glands).

Octochetus Diplocardia

(single gizzard; micronephridia ; (double gizzard; meganephridia ;

calc. glands in xv. or xvi. 1m most no cale. glands).

species).

Trigaster ’

(double or triple gizzard ;

micronephridia; no calc. glands).

Dichogaster

(double gizzard; micronephridia;

calc. glands in region xiv.-xvil.).

Eudichogaster, whose position is now under discussion, has a

double gizzard, is micronephridial, and has calciferous glands

(except in one species) in two or more of segments X.-xill.; In

some species the posterior male organs have undergone the

microscolecine reduction. It is purely Indian in distribution.

The view, now held by Michaelsen, that Hudichogaster is derived

from J'rigaster, and therefore to be included in the Trigastrine,

is based primarily on the close anatomical similarity between

the two genera; the only essential difference is that calciferous

glands are absent in Vrigaster and present in Hudichogaster.

Moreover, there is one species of Hudichogaster in which calciferous

glands can scarcely be said to be present at all. Michaelsen,

in describing H. bengalensis (5) says:—‘a pair of lateral

ealeiferous glands in each of segments x.—xiii., not externally

demareated.” In giving additional notes on the same species (9)

T have stated that “the bulgings of the esophagus in segments

x.-xiii. are thin-walled and are not at all set off from the lateral

walls of the cesophagus; they are not calcareous glands any more

than the similar part of the tube in, for example, Pheretima

posthuma is a series of calcareous glands”; and in support there

follows a description of the appearances seen on opening the

tube. Here, then, is a form which according to strict definition

isa Urigaster; though I imagine no one will quarrel with either

Michaelsen or me for including it, on geographical grounds, in

Hudichogaster.

Michaelsen also regards the similarity of the nephridial

condition as being decisively in favour of the derivation of

ELudichogaster from Trigaster vather than from Octochetus. The

species investigated for the purpose of this comparison were

Eudichogaster ashworthi, Trigaster lankesteri subsp. calwoodi,

and Octochetus thurstoni. In Hudichogaster ashworthi theve

occurs in each segment a number of small loose micronephridial

106 bx. J. STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,

tufts, each apparently with a funnel; in addition, in the hinder

segments there is on each side, near the ventral nerve cord, a

larger nephridium in the form ‘of a fairly large rosette, with a

funnel in the preceding segment. Trigaster lankesteri has the

same arrangement, while Octochwtus thurstoni has numerous micro-

nephridia throughout the body, without any trace of the larger

organs. The only morphological change, therefore, which is

necessary in order to evolve Hudichogaster from Trigaster is the

development of calciferous glands; and indeed these are scarcely

present in one species of Hudichogaster.

Michaelsen admits that the geographical facts do not at first

sight appear favourable to this view. As has been noted,

Eudichogaster is purely Indian, while T’rigaster belongs to

Mexico and the West Indies. But the widespread occurrence of

Dichogaster, a descendant of Trigaster, in tropical Africa is

evidence, according to Michaelsen, of a former land-bridge across

the Atlantic; and Michaelsen supposes that either (1) Eudicho-

gaster originated from Trigaster in America, spread eastwards

ACL‘OSS ihe! land-bridge to Africa, colonized anes or parts of it,

and then spread eastwards again across another land-bridge to

India (it is not found in Africa at the present day because it

has been extirpated there by the dominant Eudrilines and the

later evolved Dichogaster); ov (2) that Trigaster itself spread

eastwards by the same bridges, and gave rise to Hudichogaster at

the eastern extremity of its range, 7.e. in India, itself later being

extirpated in the middle portion of its range, 7. e. in Africa, by

the Kudrilines and its own descendant Dichogaster, as before.

In putting forward the view that the ancestor of Hudichogaster

is Octochetus and not Trigaster, it may be admitted that, as

Michaelsen says, Hudichogaster aud Trigaster ave very much alike

morphologically. At the same time, I think we now possess

evidence of a much closer similarity between Hudichogaster and

Octochetus (ov at least some species hitherto reckoned as Octo-

chetus) than Michaelsen was aware of. The gap between the

two genera is bridged almost, if not quite, as completely as that

between Hudichogaster and Trigaster. The points to be discussed

are the gizzards. the calciferous glands, and the uephridia.

I have deseribed (11) in Octochetus pallidus a commencing

doubling of the gizzard:—‘“‘The gizzard is barrel-shaped, in

segment vi.; the cesophagus is distinctly strengthened in seg-

ment v. also, where shining longitudinal muscular bands are seen.

This seems to be the beginning of a double gizzard, such as seen

in Hudichogaster, Dichogaster and Trigaster: 1 do not, however,

suggest at present that any of these genera are derived from this

species, or indeed from the genus Octochetus at all.”

I may here call attention to the relation of the septa to the

condition of duplicate gizzard. Octochetus pallidus is one of the

rather few species of the genus which retain all the septa in the

anterior part of the body (behind the level where they first

AND ZOOGEOGRAPHY OF INDIAN OLIGOCH ATA. 107

definitely begin). The presence of septa between the successive

gizzards seems to be a necessity for their development as separate

structures. and two gizzards probably could not develop in the

majority of species of Octochetus, where one, two, or three septa

are absent in the gizzard region; an extension of the muscularity

of the esophagus would simply result in an increase in the size

of the existing gizzard. In Trigaster, with two or three gizzards,

the septa are all present. Iam not acquainted with the facts in

all the numerous species of Dichogaster, but the septa are certainly

often present; in D. malayana, where there is no septum 5/6,

separate gizzards are, according to my observation (9), scarcely

discernible in segments v. and vi.—they seem to have “run

together,” as it were. In the single species of the genus J/ono-

gaster—essentially a Dichogaster in which there is only one

gizzard—the septa in the gizzard region are wanting (6) and the

two gizzards of the Dichogaster ancestor have doubtless “run

together.” In those species of Dichogaster where septa are absent

in the region of the gizzards (e.g. D. crawi) we may perhaps

predict that the gizzards will not remain long separate, and that

the condition of Monogaster will be arrived at. We may conclude

that the duplication of the gizzard, while impossible in the

majority of species of Octochetus, would be possible in the primi-

tive group consisting of O. bishambari, pachpaharensis, and pallidus,

and seems to be in process of accomplishment in O. pallidus.

Next with regard to the calciferous glands: in Hudichogaster

these organs are in segments x.-xii. or thereabouts, in Octochatus,

usually, in segments xv. or xvi. It would, I think, be difficult to

derive Hudichogaster from Octochetus if these were constant

characters of the two genera (though Michaelsen, deriving Dicho-

gaster from Hudichogaster (4), sees no difficulty, apparently, in

assuming a (dislocation of the glands backwards; in deriving

Hudichogaster from Octochetus—trom the usual type of Octochetus

that is—the dislocation would have to be forwards). But the

more primitive species of the genus Octochetus (O. bishambari,

pachpaharensis, and pallidus) have no calciferous glands; in

Eudichogaster bengalensis, as has been seen, they are at a very

low level of development ; the morphological similarity, in this

respect, between these species of Octochetus and Hudichogaster is

just as close as that between Trigaster and Hudichogaster, and the

derivation of the one from the other just as easy.

It will be remembered that one reason for deriving Vudicho-

gaster from Trigaster rather than from Octochetus was the

similarity of the nephridial condition in H#. ashworthi and

T'. lankesteri, and the dissimilarity between H. ashworthi and

O. thurstont. But a wider survey of the nephridia of the latter

two genera shows that not all Eudichogasters are in the same

condition as H. ashworthi, and that not all species of Octochetus

are like O. thurstoni. Of the six species of Hudichogaster where

the descriptions are sufficiently detailed to be of use, in only one

other (4. prashadi) is the nephridial system capable of being

108 DR. J. STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,

described in the same way asin /. ashworthi; it follows there-

fore that if EZ. ashworthi is similar to Trigaster, the majority of

species of Mudichogaster, so far as known, are not. Speaking

very broadly, there is indeed some similarity between all these

six species of Hudichogaster and Trigaster lankestert ; in all, a

certain number of the micronephridia are of large size, much

larger than in Pheretima, for example, or Hutypheus, to take two

well-known micronephridial genera. But this feature occurs also

in some species of Octochwtus—-in exactly those three species

previously referred to; in these there are seven, three, or even

apparently only one nephridium on each side in each segment,

which make up in size what they lack in number. The majority

of species of Hudichogaster, in fact, approach in their nephridial

condition somewhat more closely to such forms as Octochctus

pallidus and O. pachpaharensis than to Trigaster lankesteri ™.

T am, however, not inclined to attach very great weight to

any argument from the nephridia, The possession of a certain

number of micronephridia of fairly large size does not necessarily

show genetic relationship: it occurs, for example, in species of

Megascolides and Megascolea, which belong to a different sub-

family, the Megascolecinew. Indeed there are very diverse

conditions within these two genera themselves.

T think the above considerations show that the passage from

Octochetus to Eudichogaster is just as easy morphologically as

that from Vrigaster, and that there is no difficulty in deriving

Eudichogaster from an Octochetus ancestor which had the

characters of the group pallidus, pachpaharensis, and bishambare.

But if the morphological evidence is equally balanced, the

geographical evidence is strongly on the side of the descent of

Eudichogaster from Octochetus. Octochetus is a characteristic

Indian genus, found throughout the land; Hudichogaster is

exclusively Indian, and is found in a broad belt across the middle

* In Budichogaster ashworthi, towards the hinder end of the body the innermost

of the transverse series of micronephridia enlarges so as to resemble a mega-

uephridium ; the number of micronephridia in each segment appears to be small,—

in var. kinneari it is about six on each side. In KH. prashadi much the same

occurs,—there are about five on each side, regularly arranged behind each other m

succeeding segments till towards the hinder end, where fhe innermost becomes

larger and the others smaller, less regular, and more numerous. In H. barodensis

the three most dorsally situated micronephridia on each side of each segment are

larger than the rest, while at the hinder end the innermost (most ventral) also

enlarges. In EH. bengalensis there are two pairs of large nephridia per segment in

addition to a number of small micronephridia; towards the hinder end the inner of

the two larger nephridia becomes more conspicuous than the other. In EF. chitta-

gongensis there are three or four nephridia on each side in each segment, arranged

behind each other in succeeding segments, the outermost in each transverse row

being the longest; near the hinder end the imnermost increases in size and becomes

more conspicuous. In ZF. trichochetus there are four longitudinal rows on each

side of the body, but here the innermost series is the smallest. In H. parvus,

though the nephridia are “ diffuse,” they are of considerable size.

In Octochetus pallidus the micronephridia in the post-clitellar segments are

about seven on each side in each segment, and they increase in size from the

ventralmost to the fifth, the two most dorsal being smaller again ; this difference in

size disappears towards the hinder end. In O. pachpaharensis there are three on

each side per segment behind the genital region, and in front even fewer.

AND ZOOGEOGRAPHY OF INDIAN OLIGOCHANTA. 109

of the country. Z'rigaster is not known outside Mexico and the

West Indies. Deriving Hudichogaster from Octochetus, we need

no such hypothesis as that advanced by Michaelsen—-the origin

of Hudichogaster from Trigaster in America, and its spread by

means of land-bridges across the Atlantic and Indian Oceans (or

alternatively the spread of Vrigaster itself by the same means) as

far as India. Hudichogaster would have arisen where we find

it-—in India, where its ancestor also lives.

I conclude, therefore, that Hudichogaster arose from Octochetus

in India. It must therefore go into the Octochetine, not the

Trigastrine.

RAMIELLA, gen. noy.

T propose now to consider the more primitive species of the

genus Octochetus to which reference has been made in the pre-

ceding paragraphs.

In 1914 (7) L described a worm which I placed in the genus

Octochetus under the naine O. bishambari, although it differed

from all species of Octochetus then known in having no calciferous

glands, and in having only one nephridium on each side in each

segment. Measured by its size, indeed, this nephridium would

be a meganephridium, and the worm would not be an

Octochetus at all, but an Acanthodiiline—a ‘“ Notiodrilus,”—and

would correspond to the original Acanthodriline, the origin of

the Megascolecide. This, however, seemed impossible; there are

no representatives of the Acanthodriline in India (except one

introduced species of Microscolev) ; and the single nephridium

does not, according to the evidence of sections, come into relation

with the septum in the normal way, and 1s therefore to be locked

on as a hypertrophied micronephridium, the only one left of a

former larger series.

In 1920 (11) two more forms closely related to the preceding

came to light. While both, Octochetus pachpaharensis and O.

pullidus, ave without calciferous glands, the first has only three

(or anteriorly perhaps fewer) micronephridia on each side per

segment, and the second only about seven.

This reduction in the number of micronephridia is probably

—certainly in the case of O. bishambari—to be looked on as

secondary, while the absence of calciferous glands is probably

primitive. Other primitive features are the presence of all the

septa in the anterior part of the body (behind the level at which

they first definitely begin), and the absence of spines or teeth

on the penial sete.

It is apparently from this group that Hudichogaster has arisen,

as I have argued above. Since the group is a well-defined one,

is differentiated from the remaining species of Octochetus by

morphological characters of importance-—absence of calciferous

glands, reduction in the number of micronephridia—and has

different relationships from those other species, I propose to

erect for them a new genus, Ramiella, which J associate with

110. DR. J. STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,

the name of my former colleague, Prof. Shiv Ram Kashyap of

Lahore.

Diagnosis :—Setie eight per segment. Male pores on xviil.;

two pairs of prostatic pores, on xvii. and xix. Spermathecal

pores two pairs, in 7/8 and 8/9, or on vill. and ix. Gizzard in

vi. All septa present after their commencement. No calciferous

glands. Micronephridia; micronephridia relatively large, few

in number. Testes and funnels free in x. and xi.

Distribution :—India (Mahableshwar, $8. Rajputana, Saharan-

ur).

It will be noted that the species extend in a line from the

Western Ghats to the Western Himalayas, the most primitive

(at least the one in which the reduction in the number of

nephridia has made least progress) being at the southern end,

the most modified at the northern.

The relationships of the genera of Octochetine may be set

forth in the accompanying tree :—

“Original Acanthodriline.”

(Howascolex ?).

ne ns

Octochetus. Hoplochetella. Ramiella.

eae SS

— =a i

Dinodrilus. Butypheus. Eudichogaster.

Brythreodrilus.

(On the question of the inclusion of Howascolew in the ancestral

line of the Octochetine see (8), and the references there given.

On the inclusion of Hoplochetella and Erythreodrilus in the

Octochetine see (10).)

Probably no genealogical tree expresses relationships with

exactitude; every genus is strictly speaking at the end of a

short side line. Thus Ramiella probably comes off the main

stem shortly above the position of Howascolex; the original

meganephridium seems to have broken up in a different way in

these two genera—in Howascolex to have become one still fairly

large and a number of minute nephridia, in Ramiella to have

dissolved into a few moderate-sized organs.

The Derivation of the genus Dichogaster.

There remains the question of the origin of the genus

Dichogaster. In 1903 and 1909 Michaelsen regarded it as

derived from Hudichogaster; while in 1910 he states that

morphologically it is best derived from Eudichogaster, though

geographically it would appear easier to derive it from T’rigaster ;

the geographical argument is, however, not by any means

absolutely cogent—there is nothing in the facts essentially

opposed to the derivation from Hudichogaster. ;

To this view of the origin of Dichogaster I cannot agree. In

AND ZOOGEOGRAPHY OF INDIAN OLIGOCH RTA. Tabi

Eudichogaster the calciferous glands are in segments xX.—xiii., or

some of them; in Dichogaster as a rule in xv.—xvii. It is not so

easy for me as it is for Michaelsen, apparently, to imagine a

“dislocation backwards” of the glands; it is easier for me to

conceive Dichogaster arising from Tvrigaster which has no such

glands, than from Hudichogaster which has them, but in a

different place. Apart from that, the geographical argument

seems to me decisive: Trigaster belongs to Mexico and the West

Indies, and these regions are probably part of the endemic home

of Dichogaster ; Hudichogaster is altogether Indian, and it is very

doubtful whether there is any endemic species of Dichogaster

in India at all——certainly there is none anywhere near the

Kudichogaster vegion. The place of origin of Dichogaster was

pretty certainly not India. I derive Dichogaster therefore from

Trigaster.

Diplocardiine and Trigastrine.

There is a line of descent, the Megascolecinss, which leads

from the “original Acanthodriline,’ and in which the initial

change is the disappearance of the anterior prostates and the

union of the posterior prostatic pores with those of the vasa

deferentia on segment xviii. ‘There is another line, the

Octochxtine, in which the initial change is the breaking up of

the meganephridia into mieronephridia. Similarly, there is a

third line, the initial change here being the reduplication of the

gizzard.

This third line (there are still several others) has commonly

been divided up into the two subfamilies of the Diplocardiinz

and Trigastrine. These subfamilies, indeed the two combined,

are smaller than the Megascolecine or the Octochetine ;

the Diplocardime comprise only Diplocardia and Zapoteeia

(Diplocardia having two and Zapotecia three gizzards, a

distinction which is not held to be of generic importance in the

ease of Trigaster, which contains species with both) ; while the

Trigastrine, after the removal of Hudichogaster, comprise

Trigaster, Dichogaster, Monogaster, and Hutrigaster (with three

gizzards, and three pairs of calciferous glands in segments xv.,

xvi. and xvii.). I believe there would bea gain in uniformity

and an increase in convenience in uniting the subfamilies under

the one head of Trigastrine.

* Original Acanthodriline.”

Diplocardia.

eee

Ris

Zapotecia. Trigaster.

Dichogaster.

rane

| ibis

Monogaster. Eutrigaster,

112 DR. J. STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,

References to Literature.

1. Micuartsen, W.—Oligocheta, in: Das Tierreich. Berlin, 1900.

2, m Neue Oligocheten und neue Fundorte alt-bekannter. Mitth.

Naturhist. Mus. Hamburg, vol, xix. 1902.

3. i Die geographische Verbreitung der Oligochiten. Berlin,

1903.

4. fe The Oligochzeta of India, Nepal, Ceylon, Burma, and the

Andaman Islands. Mem. Ind, Mus., vol. 1. 1909.

5. nt Die Oligochiaten-fauna der vorderindisch - ceylonischen

Region. Abh. Naturw. Verein Hamburg, vol. xix. 1910.

6. * Oligochaten, in: Ergebnisse der zweiten deutschen Zentral-

Afrika-Expedition 1910-1911. Leipzig, 1915.

7. SrEPHENSON, J.—On a collection of Oligochzeta mainiy from Northern India.

Ree. Ind. Mus., vol. x. 1914.

8. za On some Indian Oligocheta, mainly from Southern India

and Ceylon. Mem. Ind. Mus., vol. vi. 1915.

9. Hs Ona collection of Oligocheta belongmg to the Indian

Museum, Rec. Ind. Mus., vol. xii. 1916.

10. Ps On a collection of Oligochzeta from various parts of India

and Further India. Rec. Ind. Mus., vol. xi. 1917.

11. it On a collection of Oligocheta from the lesser known parts of

India and from Eastern Persia. Mem. Ind. Mus., vol. vii.

1920.

T1.—On PouyPpHyLY IN THE OLIGOCHATA.

The material which I wish to use in this discussion is derived

from the Megascolecide, and largely from the subfamily Mega-

seolecine. It will be necessary first to show how the various

genera of this subfamily are related (v. text-fig. 1).

The whole of the family Megascolecidee is to be derived from

an original form which has essentially the characters of the genus

Notiodrilus as defined by Michaelsen in the Tierreich volume of

1900. These are: Testes and funnels two pairs, free in seg-

ments x. and xi.; vasa deferentia of each side uniting in their

backward course so that there is only one pair of male apertures,

on xviii. ; two pairs of tubular prostates, with unbranched central

canal, opening on xvii. and xix.; sete four pairs per segment ;

one pair of meganephridia per segment; a single gizzard far

forward, in segment v. or Vi.

The Megascolecine, however, take their origin from a form

which is one remove from this— Diplotrema, in which the anterior

pair of prostates have disappeared and the posterior pair of

prostatic pores have moved forwards to open on xviii. near the

apertures of the vasa deferentia.

In Plutellus, the first genus of the Megascolecine, the prostatic

pores have fused with the openings of the vasa deferentia on

xvill. so that there is but one pair of pores; and this remains

throughout the subfamily as its distinguishing character. In the

remaining genera the changes are of three chief kinds—the sete

may take on the perichetine arrangement, the four pairs multiply

in number and become spread out to form a more or less com-

plete ring round each segment ; the nephridia may be broken up,

113

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114. DR. J. STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,

with the substitution of a number, sometimes a very large number,

of small micronephridia for the single pair of meganephridia in

each segment ; and the single central canal of the prostate may

branch, with the consequence that the organ is no longer tubular

and cylindrical in form, but racemose. In a small group of

genera there is a development of two or more gizzards, instead of

the single gizzard of Plutellus.

Plutellus, then, has meganephridia, eight sete per segment

arranged in four pairs (the lumbricine arrangement), and a pair

of tubular prostates opening on segment xvill. in common with

the vasa deferentia. From Plutellus is derived Megascolides, in

which the nephridia are breaking up or have broken up; this

apparently does not always take place in the same way: in one

group of forms there are three or four nephridia on each side of

each segment, all about the same size, while in other cases there

is one large one anda number of quite small ones; however, all

stages of the process are united in this genus, so long as the

prostates and sete retain their original condition. The next

stage is WVotoscolew: the prostates now become branched ; in a

number of cases the branches of the central canal are so insig-

nificant that they have no effect on the form of the gland, and

can only be demonstrated in sections—the genus, however, is

defined as including all forms in which there is any branching at

all. Following this we come to Megascolex, where the setze take

on the perichetine arrangement; here again there are a number

of intermediate stages; in a number of species the anterior

segments retain the Tumbricine arrangement, and the increase in

the number of sete takes place gradually as we move backwards;

in others the anterior segments show an increase too, but he

paired arrangement still holds—there are six pairs, or eight pairs,

instead of four; and soon. The last genus along this line is

Pheretima; the essential characters are those of J/egascolea, but

the gizzard is further back, the testes and male funnels are

enclosed in testis sacs instead of being free in the segments (this

oecurs occasionally in JJegascolex), and on the whole the ring of

sete is more closed up—has smaller gaps in the dorsal and

ventral lines than is usual in MJegascolex.

But there are other lines starting from Plutellus. In the line |

just considered the first change was the breaking up of the

nephridia; in another line the multiplication of the sete comes

first. This change, occurring in the basal genus Plutellus, gives

Diporocheta, the generic characters of which are therefore

tubular prostates, meganephridia, and perichetine sete. It is,

of course, impossible to derive this form from any of the first

line, since those all have micronephridia; the meganephridial

condition is the primitive one, and a meganephridial cannot be

derived from a micronephridial form. From Diporocheta is

derived Perionyx, in which the prostates have branched; this

genus therefore possesses meganephridia, perichetine seter, and

racemose prostates. Asin the case of Megascolides and Notoscolex,

AND ZOOGEOGRAPHY OF INDTAN OLIGOCH ATA. 115

the transition between these two genera is gradual, and in some

eases the branching of the central canal of the prostate or its

absence can only be determined by microscopic examination.

ie n the third line which starts from Plutellus the initial change

the modification of the prostates; Woodwardia, having thus

racemose prostates, lumbricine sete, and meganephridia, cannot

be placed on either of the other lines, since in them either

the sete or the nephridia are modified from the start. From

Woodwardia is probably to be derived Comarodrilus, in which

the nephridia in front of the clitellum, but only these, ave broken

up, the gizzard has become vestigial, ‘and the originally paired

sper mathecal pores have fused in the middle line.

The genus Spenceriella has the primitive form of prostate, but

is mieronephridial, and has the perichetine arrangement of

sete; it is probably to be derived from Megascolides by mul-

tiplication of the sete. It could however equally come from

Diporocheta by the breaking up of the nephridia.

A group of small genera are characterized by the reduplication

of the gizzard. Digaster and Didymogaster have two gizzards,

and are distinguished from each other by the number and

position of the spermathece ; Perissogaster has three gizzards

situated anteriorly, as in the two former species. The condition

of the other systems indicates that these are all to be derived from

Notoscolex. Plionogaster, in which there are several gizzards

more posteriorly situated, at the beginning of the intestine, is to

be considered as or iginating from Megascolex.

Finally Pontodrilus is to be mentioned. The majority of

species are littoral in habitat; one is terrestrial, and one is limnie.

It is derived directly from Plutellus; the eizzard has become

vestigial, and nephridia are absent from the first twelve or

fourteen segments.

Attention may here be drawn to two points. The first is

that the genera of this subfamily can be arranged in phylo-

genetic order, We know which characters are primary, which

secondary—and therefore we know which forms must have come

first In evolution; in addition, we have in several cases inter-

mediate forms (between Diplotrema and Plutellus, which are

united by Michaelsen (14a); between MJegascolides and Noto-

scolex ; between NVotoscolex and Megascolex, which Michaelsen also

merges (16); and between Diporocheta and Peri ‘tonya, which again

are united by Michaelsen (16)). The whole tree is still before

us, and all stages in the evolution of the subfamily are there for

detailed examination. While there is room for doubt in some

details, the main outline will probably stand firm.

The second point is that evolution has proceeded along a few

definite lines; the essential changes are confined to a few

systems, and follow a definite direction in each case. We have

the change in the arrangement of the sete, from the lumbricine

to the perichetine; the change in the nephridial system,

116 DR. J. SPEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,

from the meganephridial to the micronephridial; and that in

the prostates, from the tubular to the racemose form; in some

cases we have changes in the gizzard, in the direction of reduction

or reduplication. Especially, in this subfamily, the first three

systems are the important ones; in the majority of genera the

gizzard remains the same, and the genera are defined by the

condition of the sete, nephridia, and prostates. Since classifi-

cation represents relationships, or ought to, as closely as

possible, and since these three systems are those which have been

affected in the course of evolution, it is these which form the

basis of our generic definitions.*

Without going into detail, I may refer, in amplification of the

foregoing, ‘o init other Silanes of the same large family.

The Octochatine, like all the subfamilies of the Megascolecide,

are ultimately derived from the original Votiodrilus form. The

first change is the breaking up of the nevhridia; the whole

subfamily is therefore micr onephridial. In two of the branches

of the subfamily the increase in the number of the sete takes

place. In one genus we have a doubling of the gizzard

(Hudichogaster, which I place in the Octochetine, cf. No. 1 of

the present series). When changes in the male organs occur

they take a different line from those which characterize the

Megascolecine ; the “ microscolecine reduction” which is found

in certain genera, consists in the disappearance of the posterior

pair of pr ostates and the translation forwards of the openings of

the vasa deferentia to join the anterior prostatic pores on

segment Xvil.

The Trigastrine, like the Octochztine, area much smaller sub-

family than the Megascolecine. The first aha ngein the Votiodrilus

ancestor along this line is the doubling of ‘the gizzard ; subse-

quently we may have the breaking up of the nephridia and the

microscolecine reduction of the anal apparatus. The only other

development is the appearance in certain genera of calciferous

glands; the genera are therefore defined in ferns of the gizzards,

nephridia, “le apparatus, and calciferous glands.

In these subfamilies we thus see the same thing—evolution

proceeds along a few definite lines, in a definite direction in each

system. Sometimes one, sometimes another system is the first

to start evolving, sometimes one, sometimes another follows—

and so we get a number of combinations, which characterize the

different genera.

* T do not mean to say that the diagnoses of the genera of Megascolecide, as we

usually have them, are confined to the systems mentioned, though there is a

tendency so to limit them ; compare, for example, the diagnoses of the Megascolecinz

given by Michaelsen in the Tierreich (6) with those by the same author in 1907 (9).

A diagnosis usually contains an element of description, statements which hold good

for all hitherto known individuals of a species, or species of a genus, but which are

not necessary parts of our conception of the species or genus. Thus we often see

recorded the description of species which necessitate the widening of the current

diagnosis of a genus.

AND ZOOGEOGRAPHY OF INDIAN OLIGOCHAETA. TA

T now wish to argue that a number of genera of Megascole-

cid are probably polyphyletic. I take as a text a sentence of

Benham’s, in his paper on the Oligochetes of the Subantarctic

Islands of New Zealand (4) :—‘ According to Michaelsen, species

of Microscolex may arise at different times, in different parts of the

world, from different species of Votiodrilus. This thesis involves

so profound a modification in the accepted ideas of evolution that

space will not permit me to discuss the problem here.” I think

we may take it, then, that the general view with regard to the

multiple origin of species, genera, and larger groups is one of

scepticism; the orthodox view is that each group has arisen

once and once only.

But we have to remember that the essential variations in the

geneva of this family are not innumerable, but limited. It is

not the case that the modes of variation are so numerous, and

the possible combinations therefore so greatly more numerous

still, that there is no reasonable chance of the same combination

of characters ever being repeated—this seems to be the foundation

of the orthodox view. The combinations of characters that

distinguish the various genera can be obtained in move than one

way, and the characters and their combinations are few enough

to render it possible that this has happened; I would even say

probable that this has happened, and perhaps often.

Let us remember that the perichetine arrangement of sete is

secondary to the lumbricine, the micronephridial condition

secondary to the meganephridial, and the racemose prostate to

the tubular, and that these changes have demonstrably taken

place more than once; and let us take sucha form as Megascolea,

with pericheetine sete, micronephridia, and racemose prostates.

It may have arisen from a form with lumbricine sete, micro-

nephridia, and racemose prostates (i.e. Notoscolex) by the multipli-

cation of the sete; or it may have arisen from a form with

perichetine sete, racemose prostates, and meganephridia

(i.e. Pertonyx) by breaking up of the nephridia. Both these

modes of origin have, in fact, been suggested; the point seems to

be decided—tfor some species of Megascolex, at any rate—by

finding a number of intermediate forms between WVotoscolex and

Megascolex. There is a third possibility, from a form with

pevicheetine sete, micronephridia, and tubular prostates, by

the change of the tubular into the racemose prostate—this

would mean that Spenceriella was the ancestor.

Or take Perionyax, with perichetine sete, racemose prostates,

and meganephridia. It might be derived from a form with

lumbricine sete, meganephridia, and racemose prostates (i. e.

Woodwardia) by multiplication of sete; or from one with peri-

chetine sete, tubular prostates, and meganephridia (i.e.

Diporocheta) by the branching of the prostatic lumen. Here the

existence of intermediate forms has decided in favour of the

latter.

Spenceriella is a very small genus, with one species in India

118 Dk. J. SYEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,

and two in Victoria in Australia. It can be derived from

Megascolides by multiplication of setz, or from Diporocheta by

the breaking up of the nephridia. Both these genera, Megascolides

and Diporocheta, occur both in India and Victoria. It is at

least not improbable that Spenceriella has been evolved separately

in India and Australia, from one or other of these genera,

perhaps from the same, perhaps from a different one, in the two

parts of its range.

It is to be noted also, that confining ourselves to that con-

siderable group of genera of the Megascolecine with a single

gizzard, which are distinguished by the characters of these three

systems, the logical end of any line of evolution is J/egascolew.

The order in which the changes in the three systems have

occurred varies in the different lines; the nephridia may be the

first to undergo their characteristic evolution (J/egascolides line),

or the sete (Diporocheta line), or the prostates (Woodwardia line).

But as we follow out the lines other changes are added ; and if,

in any line, all three systems pass from the primitive to the

derived condition, we arrive at JMJegascolex, with perichwtine

sete, micronephridia, and racemose prostates.

There seems therefore to be an @ priort probability that groups

of worms possessing certain combinations of characters, that is

certain genera, may have arisen more than once, and that the

genera as we have them to-day are polyphyletic.

But we are not dependent altogether on a priori reasoning.

The case to which Benham refers is the multiple origin of

Microdrilus from Notiodrilus. The two genera are distinguished

by the condition of the posterior male organs—in Jicroscolex the

‘‘microscolecine reduction” has taken place (this indeed is the

origin of the term). On Possession Island (one of the Crozet

group, some distance south-east of Cape Colony) Michaelsen found

two species of worms, obviously very closely related, one with the

original condition of the male organs (i.e. a Wotiodrilus), the

other with the microscolecine condition (i.e. a Microscolec) ;

the deduction is that the latter species has been evolved on the

island from the former. The two species agree in the peculiar

pigmentation, in the arrangement of sete, glandular modification

of the integument in the neighbourhood of the genital pores, in

the form of the penial sete, form of the prostates, and size and

shape of the spermathecal diverticula. The MJieroscolex was

represented by numerous examples, and so was not an individual

chance variation; and Michaelsen is doubtless right in holding

(8, 9) that it has been evolved in this remote region from the

Notiodrilus which occurs there. Of course, other species of

Microscolex have evolved elsewhere, from other species (con-

ceivably from one species) of Notiodrilus; and thus MJicroscolex

has arisen at least twice, from different ancestors and at different

times.

AND ZOOGEOGRAPHY OF INDIAN OLIGOCH ALA. 119

Benham, however, is wrong if, in the sentence I have quoted

above, he means to imply that Michaelsen holds heretical views

on evolution. Michaelsen is strictly orthodox; he will not have

it that different species of a genus may arise at different places

from different species of a parent genus; he merges the two

genera concerned, and calls all the species Microscolex (8).

Again, in the Abor country, in a remote spot in the

Hastern Himalayas, a worm is found named by me Pertonya

annulatus (18); like other examples of the genus it has racemose

prostates and perichtine sete ; but while the rest of the genus

has only meganephridia this worm has, in addition to mega-

nephridia, micronephridia also in all the postgenital segments.

But the presence of micronephridia is just what distinguishes

Megascolex from Perionyx, and by definition the worm should go

in Megascolea. A large number of species of Pertonya, however,

have a rather characteristic appearance—the dorsal surface is

deeply pigmented, of a dark purple colour; the sete are

exceptionally numerous, and the breaks in the middorsal and

midyventral lines are very small; the male poresand spermathecal

pores are close together neax the midventral line and, internally,

the gizzard is considerably reduced. These characters are not set

down in the generic diagnosis; some of them are scarcely definite

enough, and they are not features of all the species, though, in

varying degree, they are of many; they are, however, all possessed

by Perionyx annulatus. Lastly, Perionyx annulatus occurs in the

heart of the Perionya region, and more than a thousand miles

from the Indian Megascolea region. There is only one possible

conclusion—that this worm, by definition a J/egascolex, has

evolved where we find it from a Perionyx, and that it has nothing

to do in its origin with any other Indian or Australian Megascoles.

Very similar is Megascoler dubiws, which also seems to have

avisen, far away from the JJegascolea region, from a Perionyx.

I have already said, however, that Jegascolex has originated

from iVotoscolea (umbricine sete, micronephridia, and racemose

prostates) by increase in the number of the sete; ; and indeed we

get so many intermediate stages in this increase that this is no

doubt true for at any rate a large number of species ; Meygascole«

is therefore diphyletic.

But this does not end the complexity. Michaelsen (14) has

pointed out the close relation of certain Ceylon species of

Megascolex to certain Ceylon species of Wotoscolew—the group

of Megascolex travancorensis to that of Notoscolex ponmudianus.

The argument is the same as in the case of the Motiodrilus and

Microscolew of Possession Island ; ; the species of Megascolex have in

all probability arisen from the local representatives of otoscolex.

There is also a similar correspondence between species of

Notoscolex and species of JMegascolev in another restricted area,

the N. Island of New Zealand; here, too, the inference is that

the second have arisen from the former.

120 DR. J. STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,

Once more, Spenceriella possesses the perichietine arrangement

of sete, micronephridia, and tubular prostates, differing only

in the latter respect from J/egascolex, which has the more ad-

vanced racemose form of the glands. But Michaelsen has lately

(16) transferred two species of Spenceriella to Megascolex ; though

the branching of the central canal was not to be inferred from

anything in the external form of the glands, it was found to exist

in a slight degree on microscopical examination of sections. It

is scarcely rash to look on these apparently transitional species

as descended from species of Spenceriella, which they so much

resemble. They can hardly be descended from either Votoscolex or

Perionyx; the transitional species in these cases are characterized

by the incomplete setal rings, or by the incompletely broken up

nephridia, and have, apparently, the fully developed racemose

prostates, as is usual in Votoscolex and Perionyx.

In other words, species which anatomically belong to the

same genus, Megascolex, have arisen from two less specialised

genera, Votoscolex and Perionyx, and at least at three separate

times; quite possibly Spenceriella is the origin of certain other

species. Probably, of course, this much understates the truth ;

it is only a few small groups of species of M/egascolew that we can

thus trace back at present; the great bulk of species have

probably originated at still other times and in still other places.

Michaelsen, having before his eyes the separate origin of

Megascolex trom Notoscolew i in New Zealand and Ceylon, sets over

the polyphyletic difiiculty by merging the two genera into one (16).

But this is too short a way with the difficulty ; if, wherever

we find a polyphyletic origin, we merge the genera concerned,

then of course no genus will be polyphyletic, and orthodoxy will

triumph. And it may be noted that even this device of fusion

is not effective where a genus has a double origin from two

other genera. Assuming that some species of Megaseclex have

arisen from Wotoscolex, others from Perionye, the fusion of

Notoscolew, Megascolea, and Perionyx into a single genus leaves us

where we were, since the genus now has a double ovigin from

Megascolides and Diporocheeta (cf. text-fig. 1).

Take now the case of Pontodrilus. From its ancestor Plutellus

it differs in two primary respects—the gizzard has become

vestigial, and there are no nephridia at all in the first twelve

segments ; it is littoral in habit, and is very widely distributed

in the warmer regions of the globe. Benham in 1903 (3

discovered in a lake in New Zealand a worm with the above

anatomical characters, which he called Plutellus lacustris,

on the ground that the features wherein this worm

agreed with Pontodrilus and differed from Plutellus appeared to

be adaptive and related to an aquatic habitat ; he implies, though

he does not expressly state, that this worm had an origin from

Plutellus independent of that of the bulk of the species of

Pontodrilus, and cannot therefore be united with them in the

AND ZOOGEOGKAPHY OF INDIAN OLIGOCH ATA, 121

same genus. Michaelsen transferred the worm to Pontodrilus

(9), adding later (12) that it might be a Plutellus, an example of

convergence—though besides the primary features there were

others also which characterized both the new worm and the

previously known species of Pontudrilus. Benham (4) appears

to take the same view. Lastly, Michaelsen (11, p. 22) appears to

have definitely adopted the yiew that it is a Pontodrilus,

since he speaks of the apparent absence of Plutellus from New

Zealand.

Some time ago I found an entirely terrestrial Pontodrilus in

material from the centre of Ceylon (19); this may, possibly, be

the ancestor of all the littoral forms (their littoral habit is of

course secondary) ; on the other hand, it may equally well be a

descendant of some one of the species of Plutellus which are

indigenous in Ceylon, while the bulk of the species of Pontodrilus

originated elsewhere. In any case, we seem to have a distinct

possibility—I will not say more—that worms which must, ana-

tomically, be placed in the genus Pontodrilus have arisen at

various times and in various places.

An extremely curious case is afforded by a genus newly

deseribed by Michaelsen as Monogaster (15). It is esseutially

a Dichogaster (subfam. Trigastrine) in which the two gizzards

have, as it were, run together again, probably in consequence of

the disappearance of the septum between them. In the evolution

of Dichogaster the steps from the original Notiodrilus ancestor

have been as follows :—First the doubling of the gizzard, then the

development of the miecronephridial condition, and then the

development of calciferous glands in certain postgenital segments.

In Monagaster, therefore, the gizzards having secondarily united,

the essential characters are the micronephridial condition, the

ealciferous glands, and a single gizzard. But these are exactly

the characters of Octochetus, which belongs to an altogether

different subfamily, the Octochetine. In this line the initial

change was the breaking up of the nephridia, and this has been

followed in Octochetus by the development of calciferous glands,

here too, as in Dichogaster and Monogaster, in the segments behind

the ovaries; the gizzard has never been double. There is nothing

in the arrangement of the male organs to distinguish JMJ/onogaster

from Octochetus ; the calciferous glands in Monogaster ave three

pairs, in segments xv., xv1. and xvil., while in Octochetus they

are one or two pairs, In Xv., Xvi., or both ; but this could not be

a ground for generic distinction. There is a difference in type

between the micronephridia in the two genera,—numerous and

tubular in Octochetus, fewer and saclike in Monogaster; but

beyond this the only distinction is in the distribution—J/ono-

gaster comes from Africa, from the /ichogaster region, while

Octochetus has never been found farther west than the Malabar

coast of India. The line of descent of each is perfectly plain;

still the case illustrates my contention, that the same eud may

122 DR. J. SCEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,

be reached by different paths; in other words, certain groups

with the same anatomical characters may have a polyphyletic

origin. ™

Must it then be an article of faith that each genus has arisen

once and once only? Variations may be innumerable—no doubt

every organ and part may vary and does vary independently ;

but the variations that mean anything, that come to anything

from the point of view of evolution, are not innumerable—they

are limited both in their seat and in the direction they take.

And it would seem that similar steps are being taken in many

parts of the range of a group; with the consequence that the end

condition is similar also. In other words, we have a polyphyletic

origin of certain groups.

Objections may be raised to the above line of argument. It

may be said, for example, that what I have been discussing are

cases of convergence, which nobody has ever denied. Or it may

be said that if, as I have claimed, Megascolex or any other genus

has a multiple origin, then it is not a true genus, and that the

group we know as J/egascolex really consists of several genera

with different lines of descent.

The term ‘“ convergence ” is applicable to the case of M/onogaster

and Octochetus, just discussed; it is applicable to that of the

group of species of Megascolew descended from Perionyx and the

group descended from Wotoscolex, as well as that descended from

Spenceriella, if this origin should be confirmed. But it is not

applicable to the different groups of species of Megascolea

descended from different Motoscolex forms, nor, generally, to the

multiple origin of one genus from another single genus:

there is no convergence here—the developments are parallel.

Further, along with the use of the term convergence there

seems to go an idea that a careful morphological examination, or

a consideration of distribution, will ultimately suftice to distinguish

groups of different origin, and that a different descent will always

betray itself to sufficiently careful and minute investigation. My

point is that it may not do so. Naturally, in giving specific

instances to support the @ prior? argument, I have had to give

cases where some features of certain species of a genus seem to

countenance a separate origin for these species ; otherwise if there

had been no anatomical features at all to support the hypothesis

of a double origin, the argument could have been deductive only.

I have tried to show that in the genus degascolea we can with

some probability separate off small groups herve and there which

have originated at different times, from different ancestors—

these ancestors, too, belonging to more than one genus. But,

even if these groups were separated off, are we prepared to say

that the large number of species which remain (the great

majority of the genus) own a single origin ?—that we have heen

able to discriminate all such independent groups? Such a claim

would be, to my thinking, extremely rash.

AND ZOOGEOGRAPHY OF INDIAN OLIGOCHA&TA. 123

As to the further point, that Jegascolex is not a true genus

but a group of genera, the matter is largely one of words. If

anyone wishes to paraphrase my conclusion, and say that

‘different lines of descent may give rise to forms that it 1s

impossible to separate generically by anatomical characters,” I

have no quarrel with him. I believe, however, that the sentence

“genera may be polyphyletic” expresses this conclusion with

equal correctness. For what is a genus? It is, of course, an

assemblage of species having certain characters in common: are

we justified in going further, and saying ‘A genus is an

assemblage of species having certain characters in common, and

owning a common descent ” ?

I do not think so. In the first place, the term genus was used

in the first sense long before the doctrine of descent had won

acceptance. Another and more important reason is that, as must

be evident from what has preceded, we do not know what the

lines of descent certainly are, even in so well known a group

(and one so favourable for our purpose) as the Megascolecidee. If

we choose the second conception, we shall find it tnpossible, in

the present state of knowledge, to divide up the polyphyletie

group known as the genus J/egascolex into assemblages of species

having certain characters in common and owning a common

descent.

Yet genera, definite assemblages, we must have ; we must have

groups above species, and these groups must necessarily have

limits of some kind. Since we cannot, in the present state of

knowledge, define these groups by their descent, we must define

them by their anatomical characters, and perhaps by their

distribution. In some cases we can say that in all probability a

group so defined is a genetic unity; ju many cases we do not

know whether this is so or not; in some we shall suspect or feel

eonvineed that it is not. But till we are able definitely to mark

out new groups on genetic lines, we cannot relinquish the old

anatomical groups.

A classification is one thing, a phylogenetic tree another. No

one believes more firmly than I do that phylogeny ought to be

the basis of classification ; but candour must admit that as yet

it is incapable, in many cases, of constituting such a basis. Our

classification must necessarily, for practical reasons, present itself

as a complete scheme; our phylogenetic trees are and will long

remain woefully incomplete. As our ideas of phylogeny become

more and more settled, our classification must be revised to

correspond with it. But so long as we see anatomical groups

which we suspect, or can demonstrate, to be of diverse origin,

without being able definitely to separate them up according to

their descent, so long we shall have to put up with polyphyletic

genera.

For bibliography see end of next section.

124 Dk. J. STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,

ILL. Some GENERAL CONSIDERATIONS ON THE GEOGRAPHICAL

DISTRIBUTION OF INDIAN OLIGOCH ATA,

The ae distribution of Indian Oligocheta has been

treated at length by Beddard and Michaelsen (Beddard 1, 2:

Michaelsen 7, 10, 11). The earlier writings of both nacho are

valuable for the discussions of the means by which the migrations

of Oligocheta are in general effected ; but the large collections of

Indian worms investigated by Michaelsen in 1909 and 1910 (10,

11) increased our knowledge of the actual facts of distribution to

such an extent that the Special conclusions in the later works

supersede those arrived at in the earlier.

(1) The Migrations of Oligocheta.

Oligocheeta may for the present purpose be divided into three

groups—limnie, littoral, and terrestrial, each with its distinctive

modes of spreading.

Limnic forms have a great diversity of means of dispersal.

They may spread directly throughout « river system, through all

the canals and into all the tanks and reservoirs supplied from it.

Their cocoons are easily transported in the mud which adheres

to the feet of wading birds; some forms are known to encyst, and

hence may be transported in this manner even in the adult state.

An Enchytreid bas been found frozen in a block of ice, and

recovered ( Beddard, 1).

As a consequence the same genera—sometimes the same species

ev y distant places. The case 1s similar to

that of the Rotifera and Protozoa, of which the same genera and

species are found in ponds and streams all over the world. ‘There

appears to be but one genus. Branchiodrilus, of the limnic

Oligocheta which is peculiar to India, while a number of species

are found both in England and India, or in Europe and India

(species of Vais, Chetogaster, Dero, Aulophorus, Pristina, etc.).

Littoral forms live on the shore, exposed at times to submersion

in salt water. Like the last group, these have a wide distri-

bution; being, unlike earthworms in general, immune to salt

water, they can be transported in masses of seaweed, or more

commonly their cocoons are so transported, entangled in masses

of weed or other detritus. Not only can they take possession

of a whole coast, and spread along the shore-line, but they may

in this way travel over sea for long distances. The most note-

worthy genus is Pontodrilus, which occurs along the coasts of

India, ane has a foininnmendeme distribution.

Terrestrial forms constitute the bulk of the Oligocheta. Here

the means of spreading are more limited; for the most part

earthworms are dependent on their own activities for reaching

new regions, and hence their wanderings must be very slow.

According to Michaelsen, worms which are found outside their

burrows apparently wandering about have for the most part been

AND ZOOGEOGRAPHY OF INDIAN OLIGOCH ETA. 125

obliged to leave their homes by illness, or by unfavourable con-

ditions such as the flooding of the burrows; many worms, if

extracted from their holes, are unable to make new ones, and

must die. Some, however, certainly possess the power of active

wandering, as is shown by the numbers sometimes found under

heaps of manure. But it is obvious that the peopling of a

territory by earthworms through their own exertions can only

be very slow. :

Not only so, but they are limited in their wanderings by

desert tracts—some degree of moisture in the soil is essential.

Snow-covered mountain ranges are another obstruction. And

especially the sea limits them, the majority of earthworms being

quite unable to pass even a narrow arm of salt water.

We have to recognise, however, that not all the terrestrial

forms are so strictly limited in their means of dispersal as the

above would imply. A tree-trunk floating down stream, or

earth between the hoofs of cattle, may transport worms or their

cocoons. More important is the part that man has played;

Lumnbricids, natives of Europe, have been introduced all over the

world along trade routes; in W. Australia they are almost the

only earthworms to be found near the towns; the indigenous

fauna is to be sought in the remoter parts of the country. One of

the commonest worms of the Punjab is Allolobophora caliginosus;

certain species of Pheretima have been carried round the globe,

far from the region where the genus is endemic. Small worms

are more likely to be carried in this way than larger ones; and

small species of Dichogaster, an African genus, are common

throughout the Malay Archipelago, and not rare in India.

Botanical Gardens are obviously likely to be centres of dispersal

for such introduced species in a new country. Records at Kew

and Hamburg leave no doubt of the reality and abundance of

these transfers through the agency of man.

There are also, of course, differences in the powers of the

worms themselves. Some species seem to be able to travel more

widely than others, and more quickly, and to adapt themselves

to new surroundings and establish themselves more easily ; and

it may thus happen that a species spreads over a large region

quite apart from human interference. It is not always possible

to distinguish between these cases and those of introduction by

man; and Michaelsen has adopted the name peregrine for the

widely wandering species, whether they owe their diffusion to

man's agency or to their own unaided powers.

For the purposes of Zoogeography, the distribution of fresh

water and littoral forms is of little or no importance; and the

same holds for the peregrine forms among the terrestrial group.

Thus, in discussions on the place of origin and past history of

the genera or larger groups, as well as in coming to conclusions

as to the past distribution of land and water, we are limited

for our facts to earthworms in the strict sense, and to those

among them that haye a definite and limited range. But,

126 DR. J. STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,

having purified our material in this way, Michaelsen holds that

we have in the Oligocheta a group which is capable of yielding

results for paleeogeography second to those of no other group in

importance and certainty; the worms cling to the soil in the

most literal way, can only make use of the earth in their wande-

ings, and by the facts of their present distribution can thus

demonstrate in the plainest manner the existence of Jand

connections where, it may be, there is now only a wide stretch of

ocean.

(2) The Facts of Distribution and their current Haplanation.

The material which is available for use in the following

discussion is derived from the following groups:—The sub-

families Megascolecine, Octochetine, and Trigastrine of the great

family Megascolecide; the family Moniligastride ; and scarcely

anything else.

(a) The Megascolecinc. For the phylogenetic relationships of

the genera of Megascolecine the previous article and its text-

figure may be referred to.

Diplotrema, from which the subfamily takes its origin, occurs

in Queensland and New Caledonia. Plutellws is found in Ceylon,

S. India, and the E. Himalayas; in Australia and Tasmania;

several species occur in the western pait of N. America. J/ega-

scolides oceurs 1n 8. India, in W. India, and in the EK. Himalayas ;

in Australia and Tasmania; and it has one species in western N.

America. Wotoscolex is found in the Indian region mainly in

Ceylon, but also in 8. India and in the EK. Himalayas; outside

India it occurs in Australia and New Zealand. Megascolew is found

especially in Ceylon, to a somewhat less extent in 8. India, and

hardly anywhere else in the Indian region ; outside India it is

found in Australia, Tasmania, the N. Island of New Zealand, and

Norfolk I. (between New Zealand and New Caledonia). Pheretima

isa genus of which many members have wandered widely ; its

proper home, however, is 8.K. Asia and the neighbouring islands—

the whole of the Malay Archipelago; from Burma on the one

side it reaches to Japan on the other; a few endemic species are

found in India proper, but they hardly entitle India to he con-

sidered as part of its proper home; one species 1s perhaps endemic

in Queensland, and perhaps one in the Comoro Is. Diporochata

is only represented by one species in India, and the record is an

old one: no locality is given, but it was probably found in §.

India; the headquarters of the genus is Victoria and Tasmania ;

species are also found in Queensland, New Zealand, and (one

species) on the Chatham Is. (east of New Zealand). Periony« occurs

as the dominant genus in the EH. Himalayas, and is also scattered

over India generally ; it occurs also in Victoria, Tasmania, and

the Auckland Is., and one species is found in Sumatra and Java

(as usual, peregrine species are omitted from this review).

AND ZOOGEOGRAPHY OF INDIAN OLIGOCHU ATA. 127

Woodwardia is found in Ceylon and 8. India, and in Burma ; also

in Australia and in Java. Comarodrilus is purely Indian—there

is only one species, found in the extreme south. Spenceriella, a

small genus, occurs in 8. India and Victoria.

A few other small genera of terrestrial Megascolecine,

characterized by an increase in the number of gizzards, do not occur

in India; two are Australian, one is common to Australia and

New Zealand, and one occurs in the Philippines and Moluccas.

It will be seen that nearly the whole of the Indian genera (all

except the small genus Comarodrilus) are represented in Australia;

a number are found also in New Zealand, a few in the islands

near New Zealand, and a few in the islands of the Malay Archi-

pelago.

The conclusion drawn from the occurrence of the parent genus

Diplotrema in Queensland is that the subfamily took its rise from

somewhere in this region, which is not very far from the centre

of the area now inhabited by the subfamily. The descendants

have travelled further afield—towards India, towards Tasmania,

towards New Zealand and the neighbouring islands, and north-

ward throughout the Malay Archipelago to Japan. And of

course the important point is that they must have travelled

by land. ‘The reason for the absence of so many of the genera

from the islands intervening between Australia and India

is that here the mighty genus Pheretima has crushed all com-

petitors; it is the youngest, most highly specialized, and most

vigorous genus of the subfamily; it is still spreading, many species

are among those most commonly introduced by man, and they

show themselves most successful colonists.

Michaelsen does not, however, assume the prolonged existence

of a broad land connection between the regions mentioned. The

relations were much more complicated, and were often changing.

Perhaps there was not a complete bridge at any time; the

normal condition of the region intervening between Australia

and New Zealand on the one hand and India on the other was that

of an archipelago, which extended to Ceylon and §S. India over

the present Bay of Bengal. The boundaries of the islands often

changed: sometimes they joined, sometimes they separated,—

and no doubt in a different place; and in this way paths

became available for the continued expansion of the various

genera.

Moreover, since certain Indian genera have such a definitely

limited area (certain of those already noticed being confined to

8. India, Periony« being chiefly an inhabitant of the Himalayan

region, and Hutypheus, to be mentioned subsequently, being

confined to the Gangetic plain), India itself was split up into

a number of large islands. Thus the Malay Archipelago is the

only remaining part of a larger archipelago which existed in

the early Tertiary, of which the middle part is submerged, and

the Western has consolidated to form the present India. ‘The

128 opr. J. STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,

occurrence of two of these genera ( Plutellus and Megascolides) in

North America is supposed to point to their having travelled

over the Angaza continent.

The other groups are less extensive; they reinforce the above

conclusions, and permit the formulation of a few more.

(b) The Octochetine. ‘This subfamily, as stated in the pre-

vious article, originated from the common Wotiodrilus ancestor

by a breaking up of the nephridial system; the lines along

which evolution has advanced have also been mentioned.

There is a form Howascolew in Madagascar in which the

breaking up of the nephridia has not proceeded far, meganephridia

coexisting with micronephridia. In Octochatus the breaking up

is complete. Dinodrilus is derived from Octochetus by a

multiplication of the sete to the number of six pairs instead of

four; the microscolecine reduction of the posterior male organs

without change in the number of sete leads to the genus

Eutypheus. A continued increase of the number of sete and

the consequent formation of complete chains was supposed to lead

to the evolution from Octochetus of Hoplochetella; and in a

previous section I have given reasons for supposing that

Eudichogaster is also derived from Octochetus by a veduplication

of the gizzard.

Octochetus is widely distributed in India, and occurs also in

New Zealand, but not elsewhere—not in Australia. Dinodrilus

occurs in New Zealand only. Hoplochetella was tirst found in

India, and species which were referred to it were subsequently

discovered in New Zealand, but these probably belong to a

different genus (20). Hutypheus and Hudichogaster are purely

Indian genera.

Here, then, we have relationships which differ from those

of the Megascolecine ; they exclude Australia, and concern only

India and New Zealand. ‘he conclusion is that at the time of

the dispersal of the Octocheetine there was a connection between

India and New Zealand which did not extend to Australia ;

perhaps it passed entirely to the north, through the great

islands of the Malay Archipelago. The Octochetine do not

occur at present in the Malay Archipelago because they have

been unable to survive in competition with the dominant

Pheretima.

as far as India is concerned. The essential character here is a

duplication or triplication of the gizzard. The parent genus

Diplocardia differs only in this respect from the Wotiodrilus

ancestor of the whole family (Diplocardia and a closely similar

genus Zapotecia have been regarded as constituting another

subfamily, the Diplocardiine). From Diplocardia is derived

Trigaster, in which the meganephridia have given place to

micronephridia ; it therefore has the original arrangement of

the male apparatus, lumbricine sete, micronephridia, and a

AND ZOOGEOGRAPHY OF INDIAN OLIGOCH ATA, 129

reduplication of the gizzard. From Zrigaster is derived Dicho-

gaster, in which caleiferous glands are developed in segments

XvV.—xvil. or thereabouts.

The geographical relations of this subfamily are quite different

from those of the preceding groups. Déiplocardia is found in

North and Central America, and its descendant Z'rigaster in

Central America and the West Indies. Dichogaster 1s endemic

in Central America and the West Indies, and also in tropical

Africa ; all the species that are found in India are introduced,

with the possible exception of one only. Hudichogaster, a purely

Indian genus, is derived from Vrigaster by Michaelsen (by the

development of calciferous glands in segments x.—x1i. or there-

abouts); but in a previous Section (No. ih of the present series)

I have given my reasons for believing that this genus belongs to

the Octochatine,

The view of Michaelsen is that Zvrigaster spread from its

original home in Central America and the West Indies by means

of a land-bridge to Africa, and thence, by a land-bridge in the

Pliocene, to India, where it gave rise to Hudichogaster, itself

disappearing in India in the transformation. Dichog gaster had

its origin from cae ati on the American side of the Atlantic,

crossed the Atlantic | by the same bridge as 7rigaster and reached

Africa; its indigenous range at present extends no further—

indeed 1t does not seem to have as yet quite reached the eastern

shores of Africa (though a large number of peregrine species

are known from farther east, including India). Vrigaster has

been exterminated in Africa by the dominant genera Hudrilus

and its own descendant Dichogaster, which between them quite

dominate this region, in the same way that so many genera of

Megascolecidee have disappeared from the Malay Archipelago in

consequence of the spread of Pheretima.

(d) Lhe Moniligastride. This family consists of only a few

genera. Without going into the relationships of these, it may

briefly be stated that Desmogaster, the supposed ancestral genus,

is found in Borneo, Sumatra, and Lower Burma, and its descen-

dant Hupolygaster has a similar distribution. Drawida, the

largest genus of the family, is predominantly §. Indian (though

its range has recently been shown to be more extensive than was

believed); dloniligaster, a small genus very close to Drawida,

belongs to the same region.

Michaelsen supposes that 8. India and Ceylon were peopled by

this family by means of a land-bridge across the Bay of Bengal,

and rejects the supposition that the forerunners of the present

S. Indian Moniligastrids could have travelled by land round the

head of the Bay; they would have left some trace of their passage

in that region (a number of endemic species of Drawida have, in

fact, been recently shown to inhabit this region). Besides, the

bridge was in existence when the Megascolecinz passed over

to S. India, and so was available for the Monili gastrida too.

Proc. Zoou, Soc.— 1921, No, IX. §

130 DR. J. STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,

(3) The Objections to the current Haplanations.

I may preface the present section by a brief statement of the

reason why I feel a difficulty in accepting the existence of land-

bridges as an explanation of the above facts of distribution.

It is this.

Terrestrial Oligocheta are, I believe, a recent group, and some

of the genera we have been considering are among the most

recent of the earthworms. They have probably arisen in the most

recent geological periods. But the general aspect of the fauna of

Australia and New Zealand shows that no land connections with

Asia have existed during these periods. It 1s necessary therefore

to find other explanations for the existence of so large a common

element in the earthworm fauna of these regions.

(a) Terrestrial Oliogocheta a recent Group.

The food of earthworms is vegetable mould; and presumably

there were no earthworms in existence until the vegetable mould

was present in sufticient quantity to nourish them. We may

thus put their rise at some time not earher than the spread of

dicotyledonous piants, which took place during the Cretaceous

period. This would limit the evolution of the first earthworms,

the differentiation of the several families, and the evolution of the

numerous genera of these along lines of descent similar to those

we have followed out in the Megascolecine and other groups, to

little more than the Tertiary and Quaternary.

The recent origin of many of the present-day genera seems also

to be indicated by the extraordinary variability of a large

number of genera and species. As examples, 1t may be mentioned

that the variability of genital papille and other markings is a

common difficulty of systematists; that the number of gizzards

in the genus Drawida varies fairly widely in many species; that

in one and the same genus of Megascolecidze we may meet with

species with testis sacs or with free testes and funnels ; in another

with the original (‘‘acanthodriline”) arrangement of the male

organs, with the microscolecine reduction, or with the “balantine”

reduction (disappearance of anterior prostates, and union of

openings of vasa deferentia with the posterior prostatic pores) ;

in another, with paired or fused genital orifices ; or with sperma-

thece varying in number from two to seven pairs or even more;

or with seminal vesicles which may vary in number or position

or both—~indeed these variations of the seminal vesicles are

sometimes found within the same species. The consequence

is that the generic and specific diagnoses are uncommonly wide

as compared with those of other groups.

Even so, the systematist often has extraordinary difficulty

in referring his specimens correctly. He seems to get so many

‘intermediate forms; in the case of single specimens it 1s some-

times impossible to say whether more ample material would

justify the erection of a new species, or would show a range of

AND ZOOGEOGRAPHY OF INDIAN OLIGOCHEDA. Wal

variability that would link it on to an existing species. The

number of described ‘‘ forms” and varieties is therefore large.

Sometimes, as in the case of Pontodrilus, cvitical examination

and the increase of knowledge results in the union of a whole

Series of species under a single name.

Not only are transitions between species common, but the

same 1s true for genera. In speaking of the Megascolecine in

the previous article it has been noted that literally all stages in

the passage from the lumbricine to the perichetine arrangement of

sete, from the meganephridial to the micronephridial condition,

and from the tubular to the racemose prostates are met with.

The same holds for other characters which have been used as

generic distinctions, e.g. the well-developed or the vestigial

gizzard, the degree of approximation of the genital apertures, ete.

The consequent ditticulty of separating genera has led Michaelsen

to fuse a number of genera, with, | think, a great sacrifice of

convenience.

In speaking of the 8. Indian earthworm fauna, I have noted

(19) that the genus Megascolex seems to have ‘recently under-

gone a notable blossoming forth, with the production of a large

number of forms and intermediate forms, and that in consequence

it is extremely difficult to separate species fom varieties, and

varieties from examples of individual variability. ‘The (Indian)

range of Megascolex is of very limited extent, yet the number of

species 1s extraordinarily large; and still every collector, wherever

he chooses to explore, brings back numerous novelties,” Of the

same kind is the discovery of what I have ealled a “nest” of

velated species of Drawida in the Chittagong district (20), and of

species of Hoplochetella in a limited region of Western India

(20). The same blossoming forth is seen in the Lumbricide,

where the distinction of species, and especially of genera, is

notoriously difficult; genus passes into genus—often into more

than one genus—and the same kinds of changes appear to be in

progress in different parts of the tree, to such an extent that the

confusion is almost inextricable. There is thus an appearance of

incomplete differentiation, and a lack of that fixity and extinc-

tion of intermediate forms which we are accustomed to associate

with old established groups.

Add to this the mere length of the line of descent from the

supposed late Secondary ancestral earthworms to such forms as

Meyascolex and Pheretima, the latter portion of which has been

traced in the previous article. It seems highly improbable that

in such a vigorous group, and one so capable of adaptation to

new environments, the differentiation of genera should have

ceased soon after its first rise—so long ago, say, as the Eocene.

(b) The Question of Land-Bridges in general.

That the outlines of land and sea have changed during geo-

logical time is of course universally admitted ; butas to how great

oy]

132 DR. J. STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,

the changes have been opinions vary; the tide sways backward

and forward, and no agreement has been reached. Zoologists.

will remember that Wallace inclined at first to the opinion of

numerous and great changes—such fundamental changes as the

bridging of the Indian Ocean by the hypothetical continent

Lemuria; but that later he came to believe in the essential

permanence of all the great ocean basins. On the whole, how-

ever, the zoologists ave to be found among the bridge-builders,

and they have the company of some distinguished geologists ;

but it is perhaps true to say that geological opinion at present

is inclining to the theory of permanence.

Needless to say, no one would deny vertical movements of the

order of 100 fathoms or so; no one could refuse to believe that

England had been united to the mainland, or that a large part of

the Malay Archipelago had been united to the continent of Asia.

And arise of 100 fathoms would unite all the large masses of

land into one, with Australia as a doubtful exception ; seen in a

N. Polar projection, we should have a mass of Jand round the

N. Pole, with three tongues, S. America, Africa, and Malaya

radiating outwards towards the 8, Pole. ‘The soundings are not

sufficient to determine whether there is a continuous bridge to

Australia above the 100 fathom line or not. New Zealand,

Madagascar, the West Indies, and numerous small oceanic islands.

would remain separate. A lowering of 100 fathoms would isolate

N. and 8, America, Asia and Africa; and EHurope would form a

complex of islands and peninsulas much like the Hast Indies.

to-day. It is changes of this order that are considered allowable

by the more conservative school, not such changes as would

bridge the N. or 8. Atlantic or Pacific Oceans.

What appears to have brought about something of a change of

Opinion in recent years is the increasing support accorded to the

theory of isostasy. _The earth’s crust is in a condition of approxi-

mate equilibrium, the crust being less dense under the mountains.

and continental masses In general, more dense under the ocean

floor—this is shown by the measurements of gravity. It may be

too venturesome to say that the mountains float like icebergs in

water; but the idea is that the land-masses project because they

are lighter, while the bed of the oceans has sunk because this.

portion of the crust is heavier ; and without the most extensive

lateral motion of the matter of the crust the general arrangement

of continental masses and ocean cannot change.

In addition to the measurements of gravity is the fact that

there are no abyssal deposits on the continental platforms

wherever these have been adequately studied; 7. e., the continents

have never been deeply submerged, though shallow seas from

time to time there may have been. And the continental shelf

is so marked, obvious, and universal a feature of the earth’s

surface that it affords the strongest kind of evidence of the

antiquity of the ocean basins and the hmits beyond which the

continents have not extended. I have mentioned the effect of

AND ZOOGEOGRAPHY OF INDIAN OLIGOCH ETA, Haye

raising the level of the land of the globe by 100 fathoms; an

elevation of five times this amount would alter the boundaries

very little more. W. D. Matthew sums up the evidence strongly

in favour of general permanency (5):—‘‘ The geologic evidence

for the general permanency of the abyssal oceans is over-

whelmingly strong. The continental and oceanic areas are now

maintained at their different levels chiefly through isostatic

balance, and it is difficult to believe that they could formerly

have been reversed to any extensive degree,”

I propose later to enumerate the several land-bridges which

have been invoked to explain the distribution of the genera of

earthworms common to India and other parts of the w orld. But

there is none of them the former existence of which seems to be

better attested than that between Australia and India; this has

almost become axiomatic in the minds of students of the Oligo-

cheta. The reason is, as has been said, the large number of

genera that are common to India and the Australian region.

There can be no reasonable doubt that the western part of the

Malay Archipelago has been joined on to the Asiatic mainland

at no distant time; according to Wallace, ‘all the wide expanse

of sea which divides the islands of Java, Sumatra, and Borneo

from each other, and from Malacca and Siam, is so shallow that

ships can anchor in any part of it, since it rarely exceeds forty

fathoms in depth”; while the eastern part of the Archipelago

has, with equal probability, formed a part of Australia. Michaelsen

assumes not only the passage of numerous genera of Megascolecidee

from the Australian side, but (or perhaps as an alternative)

suggests that some may have passed back into Australia from

outside (16).

But how does the hypothesis of land-bridges square with the

other known facts of distribution? I have given some reason

for thinking that the whole of the earthworm fauna of the world,

and in particular that pavt of it with which we are dealing at

present, is of recent origin. AJegascolex, for example, is one of

the youngest genera; its immediate ancestor MVotoscolexr is one

stage further back; both are separated by a long line of ancestors

from the earliest earthworms, which alone seems sutticient to bring

their origin down to late Tertiary times ; I/egascolea appears to be

evolving still, and has not as yet settled down to the comparative

fixity of an old-established genus.

Now it is well known that, broadly speaking, Australia has no

indigenous Hutherian population. The great groups of terrestrial

Hutherians originated in the Hecenc ome in the very early

EKocene—and spread rapidly thereafter, How, on the supposition

of a land-bridge, are we to let the Australian earthworms out to

India without letting the Asian mammals into Australia ?@ If the

door is open for the particularly slow-moving worms, it is open

134 DR. J. STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,

for the quick-moving Carnivora; in asking for land-bridges to

explain the distribution of the Oligocheta we get much more than

we want. There can never have been a land connection between

Australia and the great land mass to the north-west since

the Hocene.

Still stronger is the case of the supposed connection between

India and New Zealand. This is a necessity, according to

Michaelsen, in order to explain the occurrence of the Octo-

chetine in both lands; and since the Octochetine do not

occur in Australia, the bridge in this case avoided Australia.

New Zealand does not even contain Marsupials; yet Octochetus,

the genus common to India and New Zealand, is not a particularly

archaic genus, and its occurrence in both India and New Zealand

would, on Michaelsen’s view, have to be explained by, presumably,

late or middle Tertiary land connections. But New Zealand is

an oceanie island, and probably has never been connected at any

time* with the larger land-masses, certainly not in Tertiary

times.

It is quite possible that similar objections might be brougnt

against the other land-bridges which have been postulated to

explain the existence of velated or identical genera of earthworms

in distant lands. If have specially mentioned the above because

it is so obvious, once attention has been drawn toit. The general

principle is that, earthworms being a recent group, and requiring,

on the hypothesis of dispersal by land, connections of some

considerable permanence, other groups will have been able to

pass even more easily ; and the dispersal of earthworms by land-

bridges cannot be assumed unless there is a large degree of

similarity between other elements of the fauna also.

(4) Contributions towards a more satisfactory Solution.

I trust that, in what follows, I shall not be considered to be

treating too lightly the claims of zoogeography to a hearing in

the discussion of the problems of paleogeography. As Michaelsen

* Michaelsen’s time-scheme can be put together somewhat as follows :—The

oldest components of the Indian earthworm fauna date from the Upper Jurassic,

when India was connected broadly with both Angara and Australia; Plutellus and

Megascolides wandered off into Angara, reaching western N. America in the later

Cretaceous. The chief part of the evolution took place in the Tertiary, the period

of the changing land-bridges. In the Pliocene the now consolidated Indian

peninsula became connected on the W. or N.W. with lands which had earlier

received their earthworms from Tropical Africa (Hudichogaster).

It will be seen that he puts the evolution of the group earlier than I do; but I do

not find anything which invalidates the line of argument and general conclusions of

section 3a above, especially that of the quite recent origin of the phyletically youngest

genera such as Megascolex. The word used by Michaelsen tor the period of the

crigin of the Indian Oligochete fauna is ‘‘Malm,” which corresponds (Ziegler,

Zool. Worterbuch) to the Upper Jura. Plutellus and Megascolides ave supposed

to have then been in existence ; is there any other example of genera of a variable and

evolving group persisting since that period, especially genera, such as these, which

are connected by intermediate gradations not only with each other, but with the

genera below and above them (Diplotrema and Notoscolex), genera, ‘that i is, which

are still not sharply marked off from their ancestors and descendants ?

AND ZOOGEOGRAPHY OF INDIAN OLIGOCH ELA. 135

says, ‘‘ Since the present geographical distribution of earthworms

depends in the first place on the configuration of land and sea in

recent geological epochs, it is to be looked on as a valuable

document for the history of the earth.” We are not bound, that

to accommodate our conceptions of the wanderings of the

ancestors of the present-day fauna to the views founded on

geological evidence only ; we also are in possession of important

documents, and their evidence may perhaps be of superior cogency

to that of geology, A zoologist is not likely to underrate the

value of the evidence furnished by zoology; only we must be sure

what its value is.

And firstly, in the present case, even if there were no geological

evidence, even if we were not told that ‘the geologic evidence

for the general permanency of the abyssal oceans is over-

whelmingly strong,” it would be our duty not to introduce land

connections unnecessarily. It is an old philosophical rule that

“cause non sunt multiplicande preter necessitatem ” ; in the

present case we may substitute ‘ bridges,’ and say ‘‘pontes non

sunt multiplicandi preter necessitatem.” We have a number of

agencies which are in existence before our eyes to-day: The slow

extension of distribution by the normal wanderings of earth-

worms, the extirpation of indigenous worms by younger forms

of later introduction, the existence of natural rafts on the sea, the

known ability of certain worms and their cocoons to endure salt

water, the polyphyletic origin of certain genera, and moderate

changes of land and sea; and it may fairly be demanded that we

exhaust the possibilities of these before we have recourse to the

construction of bridges which we cannot see and which are at

any rate much more hypothetical in nature.

Again, I speak only of those bridges which have been postulated

in order to explain the distributions of Oligocheta, and especially

of those Oligochta which occur in the Indian and Australian

regions. My contention is that the greater part of these are

unnecessary in this connection; whether they are a necessary

assumption or not for other reasons, | must leave to others.

And first with regard to natural rafts. Matthew recalls the

fact that these have several times been recorded as occurring over

a hundred miles off the great tropical rivers such as the Ganges,

Congo, Amazon, and Orinoco; and for one such observed,

hundred may have drifted out unnoticed. Wallace, in his

‘Island Life,’ speaks of ‘those floating islands which are often

(italics mine) formed at the mouths of great rivers. Sir Charles

Lyell describes such floating islands ‘which were encountered

among the Moluccas” (i.e. between Celebes and New Guinea,

where there is no large river) “on which trees and shrubs were

erowing on a stratum of soil which even formed a white beach

round the margin of each raft. Among the Philippine Islands

similar rafts with trees erowing on them have been seen after

hurricanes, and it 1s easy to understand how, if the sea were

136 DR. J. STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,

tolerably calm, such a raft might be carried along by the current,

aided by the wind acting on fine trees, till after a ge of several

weeks it might arrive safely on the shores of some land hundreds

of miles away from its starting-point.”

Overseas colonization is a very remote chance, it is true, in any

given length of time; but, says Matthew, if we multiply the

almost infinitely small chance that such colonization takes place

in any given length of time, such as a year, by the almost infinite

duration of geological periods, we obtain a finite and quite

probable chance. For example, the time during which natural

rafts have been observed covers about three emnnes, while the

duration of Czenozoic time is estimated as three million years ;

if we allow that ten cases of natural rafts have been recorded

during these three centuries (the wording of the extract from

Wallace given above would, however, seem to indicate that this is

an understatement), a thousand may have actually occurred in

this time, and hence thirty million in the whole Cenozoic (this 1s

a miscaleulation—it should be ten million). He then makes

certain assumptions regarding the occurrence of living mammals

on such rafts—as to the chances of there being a couple, or a

gravid female, and as to the dangers of landing ; and his con-

clusion is that the number of cases during the Cenozoic in which

mammals will have established themselves on the larger oceanic

islands is of the order of 8300—quite enough at any ike to cover

the dozen or two known cases. With invertebrates the chances

would be much greater.

And certainly, whatever the possibilities of the transfer of

mammals by rafts, the transfer of earthworms must be far more

probable. Such rafts as have been described above may or may

not bear mammals—Matthew’s calculations ave based on the

supposition that they do so only once in a hundred times; but

every one will probably contain earthworms, in the soil, under

the bark of living trees, in the axils of their leaves, or in rotting

wood. Nor are worms restricted to the larger rafts; the sm aller

worms of euryhaline groups (those that ean withstand salt water)

and especially their cocoons, may probably be transported for

long distances in masses of tangled seaweed ; Michaelsen, himself

a br ridge- -builder, presses this point against Benham in explaining

the distribution of Microscolea in the Subantaretic regions (18).

It may be asked, too, whether edrthworms are in general so

readily killed by salt water as is assumed. Itis well known

that many Enchytreids and Tubificids are regularly found on the

shore; and among the higher groups the genera Pontodrilus,

Pontoscolen, and Microscolex have thesame habitat often, though not

always; 1 have seceived Hoplochetella from the shore of western

India, though the genus was not previously known from such

localities. It is at least possible that many worms are capable

of speedy acclimatization to salt, just as a fresh-water Amoeba

ean be acclimatized by the gradual addition of salt to its water.

And it is remarkable how difficult it is to come at any definite

AND ZOOGEOGRAPHY OF INDIAN CLIGOCH A&A. Wiz

experiment on the subject of worms and salt water; the statement

that they are destroyed by it seems to be commonly accepted,

without comment and without reference.

There are, of course, other possibilities of transport for worms,

or for their cocoons—the mud on birds’ feet for example; Benham

calculates that a strongly flying bird could pass from Australia to

New Zealand (1200 miles) in 36 hours. But the possibility that

I most wish to msist on, after that of the occurrence of rafts—

because I do not think that it has as yet received any attention

is that of the polyphyletic origin of some, at least, of the genera

common to the Indian and Australian regions. I believe that

this will go some distance towards explaining the presence of

these common elements in the two faunas; evolution has pro-

ceeded on parallel lines, and the younger genera have not wandered

from India to Australia or from Australia to Jndia—they have

been independently evolved in each region.

In my first sketch of the present argument the whole question

of polyphyly i in these genera was discussed in this place ; it was,

in fact, in reviewing Phi geographical distribution that [ was

brought up against the subject. But the space that I was obliged

to devote to it seemed too great to assign to a subordinate head-

ing, and I decided to treat it independently. The whole of the

previous article, however, may logically be placed here, as a con-

tribution towards a more satisfactory explanation of the facts of

distribution.

We may finally proceed to a separate consideration of the

several bridges which have been postulated; and here I shall

usually take the conclusions of Michaelsen (with whom Beddard

is in general agreement) as the basis of my own discussion, since

he is the author who has treated the matter most fully. Michael-

sen requires all the bridges to be mentioned, and the splitting up

of India into islands as well; however, he regards himself

conservative in this matter. In a controversy with F. Sarasin

(11) he says, “I am reproached with being too wanton in my

bridge-building, I do not think that such a charge can be sub-

stinnnenedl on the contrary, I believe that we lp not reckon

sufliciently with the mobility of the earth’s crust in this region ”

(2. e. the Indo-Austrahan region).

These bridges are as follows:—A bridge between Asia and N.

America, to explain the occurrence of Plutellus and Megascolides

in the western part of N. America ; a transatlantic bridge between

the W. Indies and Central America on the west and Africa on

the east, and a bridge between Africa and India, to explain the

occurrence in India of the Trigastrine; one between Australia

and Further India over the present Malay Archipelago, and one

across the present Bay of Bengal, said to be requisite to account

for the distribution of the Megascolecine and Micnaligastmdees are)

particular bridge, at a particular time, between India and New

138 DR. J. STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,

Zealand, to explain the distribution of the Octocheetine ; and lastly,

we may add, in the opposite sense, a number of arms of the sea,

stretching across India and dividing it into a number of islands,

which formed a western extension of what is now the Malay

Archipelago. :

The bridge to N. America may beadmitted. It would pass from

the eastern end of Siberia to Alaska, and demands no considerable

elevation of the floor of the ocean—indeed, a rise of 1000 feet

would convert the N. Pacific into dry land as far south as the

60th parallel.

The case is otherwise with the bridge between Africa and India.

What is asked for is something like the Lemuria of Wallace, or

the Gondwana continent postulated by many geologists. The

important question here concerns Hudichogaster, according to

Michaelsen a member of the Trigastrinee, and descended from

Trigaster, which is endemic in the W. Indies and Mexico.

Michaelsen supposes that either Trigaster crossed the Atlantic (by

an Americo-African bridge) and made its way across Africa, and

thence by the bridge now under discussion to India, where it

evolved into Hudichogaster (suffering extermination in the African

part of its range); or Hudichogaster originated from 7’rigaster on

the American side, passed across in the same way, and was exter-

minated in Africa but maintained itself in India. But I think

I have shown in a previous section that it is at least equally

probable that Hudichogaster originated from Octochetus (or

Ramiella); on this supposition Hudichogaster arose in India, to

whieh it has thus always been confined.

Tbave myself argued that the Indian genus Hoplocheteila

may be descended from Howascolex, found in Madagascar (20),

and Lemuria or Gondwana would form an easy path for its

transport. But Hoplochetella is—or at any rate a number of

species are—euryhaline, and are found on the shores of western

India; and we must reckon with the possibility of transport

from Madagascar i in seaweed or other tangle ; the S.W. monsoon

blows in the required direction for several’ months of the year.

Dichogqaster has reached most of the islands of the Malay

Archipelago, and some of the Polynesian islands, as well as India,

and there is no doubt that small species of this genus are

frequently transported by man in the way of trade. Itis admitted

that there is no need whatever to introduce land-bridges to

explain the wide occurrence of these species all over the Kast.

The last reason for assuming the former existence of the

Indo-African bridge would be the presence of a Moniligastrid

(though one widely different from the Oriental Moniligastride)

in tropical East Africa. This African Moniligastrid is not

descended from the ee branch of the family, nor the Oriental

from the African ; this follows from the position of the gizzards—

in front of the genital segments in the African, behind in the

Oriental worms. Thealimentary tube, without special thickening

in the common ancestor, has developed into a series of gizzards

AND ZOOGEOGRAPHY OF INDIAN OLIGOCHATA. 139

in one place in the African, in another place in the Oriental

branch. There is therefore no question of African forms having

travelled to India, or of Indian forms to Africa; the question is,

where did the common ancestor live? We can only say, we do

not know. Smith and Green, the discoverers of the African

form, do indeed suppose this ancestor to have arisen somewhere

in Gondwanaland, whence the Syngenodriline branch migrated

to Africa, the Moniligastrine to the Oriental region (17); but

so far as [ know there is no special reason for the supposition.

Wallace, as is well known, gave up Lemuria, and became a

believer in the permanence of the ocean basins. Matthew states

that there is no necessity for Gondwana, from a paleontological

point of view—not even in the Paleozoic, if the interpretation of

the facts of distribution is made along the lines he lays down

(origin of groups in the north, spread towards the south, the

more primitive groups first and furthest); the weakness of the

original evidence for the former existence of Gondwana is

forgotten, and new discoveries are interpreted in the light of it,

as if its existence were well established.

The Americo-African bridge, from Central America to tropical

Africa, does not concern us so closely, and in showing reason to

believe that Mudichogaster originated in India, we entirely do

away with the necessity for it so far as India is concerned.

Whether the large number of African Dichogasters can be

explained as easily as the large number of Indian and far Eastern

species of this genus—as having been carried to their new homes

in the way of trade or huinan intercourse—seems doubtful. At

the same time, in assuming a land-bridge we are probably getting

more than we ask for; what we want is a passage for the extremely

slow-moving earthworms, and when it is a matter of thousands

of miles this passage must be one of some permanency ; what we

actually get, therefore, is an easy and abundant passage, for a

long space of time, for all the elements of the fauna, and

mingling of the animals of the two regions to an extent which

has certainly never happened. I can only conclude that we are

probably better off, on the whole, without the Americo-African

bridge.

The objections to the Indo-Australian and Indo-New Zealand

bridges have already been sufticiently insisted on. And not only

are the objections more striking than elsewhere, ast In

the case of the Australian bridge—the difficulty in dispensing with

the connection is also smaller. The actual distance to be accounted

for, as is well known, is not great. A union of the eastern part

of the Malay Archipelago with Australia, and of the western part

with Further India, is not only a feasible but a necessary suppo-

sition on every ground; a land-bridge spanning the interval

between the eastern and western parts of the Archipelago is

objectionable except for the specific purpose of accounting for the

distribution of the Oligocheta. Wallace placed the houndar v

between the two ficonmilers faunas of the Australian and Oriental

140 DR. J. STEPHENSON ON THE MORPHOLOGY, CLASSIFICATION,

regions between the islands of Bali and Lombok; this is the

interval where the assumption of a land- bridge raises many more

difficulties than it explains. But the interval is only fifteen miles ;

and while birds’ feet and natural rafts offer a sufficient mode of

transfer for worms and their cocoons, they cannot serve to trans-

plant the mammals—not a whole mammalian fauna at any rate.

It is, too, in the genera of the Megascolecine, the group w hich is

common to India and Au stralia, that we have seen most reason

to believe in px oly phyletic origins; as bearing on the probability

of polyphyly it 1s interesting to recall w hat Michaelsen says (7).

concerning the broad differences between the Indian and

Australian groups of Megascolex-—that the Australian species are

simpler, at a lower level of evolution, and more uniform, while

the Ceylonese species are often further advanced and mm many

cases approach Pheretuma. We can thus manage quite well with

the vere cause we know, but the bridge would only embarrass us.

The distance to be overcome in the case of New Zealand is

eventer ; but the general faunistic objections to a land connection

with S. W. Asia (whieh is supposed to have avoided Australia)

are greater also. Weare compelled, therefore, to invoke the same

agencies as before.

Michaelsen’s plea for a bridge across the Bay of Bengal, by

which worms from Australia, and also from Further India, could

reach the south of the peninsula and Ceylon without going round

by the head of the Bay, depends for its force on the presence in

S. India and Ceylon of genera which are not found elsewhere in

India; the argument is that 1f these genera had passed through

the lands about the head of the Ba uy, they would have left there

some trace of their passage. Thus Drawida, a Moniligastrid,

common in 8. India, and descended from a form which was

probably not unlike Desmogaster (now found in Burma, Sumatra,

and Borneo), was, when Michaelsen wrote, unknown from the

intervening region, except for a few records of peregrine species.

But more recent discoveries have shown that, both in the EH.

Himalayas and near the coast at the head of the Bay, there are

a number of endemic species of Drawida; and it can no longer

be urged that the Moniligastridee cannot have passed round that

way because they have left no trace of their passage. Certain

genera of the Megascolecinz also were supposed to show the same

limitation of distribution. Notoscolea, was only known from

S. India and Ceylon, and the same was true of its descendant

Megascolew. Lately, however, the EK. Himalayas have been shown

to harbour three species (and a variety) of Wotoscolex (Megascolides

oneilli is a Wotoscolex); so that here again it can no longer be

claimed thata bridge across the Bay of Bengal i is necessary because

otherwise the genus would have left some trace of its passage

round the head of the Bay. And the polyphyletic origin of

Megascolex is, | think, clear enough to allow us to dispense with

the supposition that it migrated into India from outside, whether

round the head of the Bay or by a land-bridge across it.

AND ZOOGEOGRAPHY OF INDIAN OLIGOCHETA. 14}

Finally, I do not think that Michaelsen’s view that India

was, in the past, divided by stretches of sea—shallow arms of

the sea—into a number of disconnected islands, is necessary.

A number of Indian genera do show, as he remarks, a limitation

more or less definite to certain tracts of the country. These

are legascoler and Notoscolex, to the south of the peninsula

and Ceylon; Drawida, though this genus can now scarcely be

said to be even roughly limited to the south ; Hutypheus, to the

Gangetic plain; Hudichogaster, to a broad belt across the

middle; and perhaps Hoplochetella, to western India. Perion yer

has not now the strict limitation to the Himalayan region that

was previously thought; nor is Octochetus limited to any one

part of the country-—it seems to occur throughout. It is, on the

whole, the youngest genera that are limited in distribution,

and it would seem possible to explain this by supposing that they

have not as yet had time to spread very widely, rather than that

their dispersal has been hindered by arms of the sea. The con-

clusions of geolagy, moreover, seem to be against Michaelsen’s

view. “It has been conclusively proved that the peninsula of

India has never been beneath the sea since the Carboniferous

period at least.” (Encyc. Britt., xi. ed., art. Asia, section

Geology.)

heferences to Literature cited in Articles IT. and ITI.

. Bepparp, F. E.—A Textbook of Zoogeography. Cambridge, 1895.

5 Earthworms and their Allies. Cambridge, 1912.

. Bennam, W. B.—On some new Species of Aquatic Oligocheta from New

Zealand. Proc. Zool. Soc., 11. 1903.

Report on the Oligochzta, in: The Subantarctic Islands of

New Zealand, vol. i. Wellington, N.Z., 1909.

Martruew, W. D.—Climate and Evolution. Annals N.Y. Acad. Sci., vol. xxiv.

1915,

6. Micwartsen, W.—Oligocheeta, in: Das Tierreich. Berlin, 1900.

Of. Es Die geographische Verbreitung der Oligochiten. Berlin,

1903.

8. os Die Oligocheten der deutschen Tief-See Expedition, in -

Deutsche Siidpolar Expedition, Ergebnisse, vol. iii. 1905.

9. - Oligocheeta, in: Die Fauna Stid-West Australiens. vol. i.

Jena, 1907.

10. Fp The Oligocheta of India, Nepal, Ceylon, Burma, and the

Andaman Islands. Mem. Ind. Mus., vol. i. 1909.

11. 4 Die Oligochatenfauna der vorderindisch-ceylonischen Region.

Abh. Naturw. Verein Hamburg, vol. xix. 1910.

12. Me Oligochaten von verschiedenen Gebieten. Mitth. Nathist.

Mus. Hamburg, vol. xxvii. (Beihett 2), 1910.

1183, s Zur Kenntnis der Kodrilaceen und ihrer Verbreitungsver-

haltnisse. Zool. Jahrb., Syst. Veil, vol. xxx. 1911.

14. 5 Oligochaten von Travancore und Borneo. Mitth. Nathist.

Mus. Hamburg, vol. xxx. (Beiheft 2), 1913.

14a. 5 Oligochieta, in: Nova Caledonia, A. Zool. vol. i. 1913.

15. 3 Oligochaten, in: Ergebnisse der zweiten deutschen Zentral-

Afrika-Expedition 1910-1911. Leipzig, 1915.

16. 1 Oligochaten, in: Results of Dr. E. Mjéberg’s Swedish

Scientific Expeditions to Australia 1910-1913, pt. viii.

Stockholm, 1916.

142 ON INDIAN OLIGOCH ETA.

17. Swern, F., and BesstE R. Grren.—Descriptions of new African Earthworms,

including a new genus of Moniligastride. Proc. U.S.

Nat. Mus., vol. lv. 1919.

18. SrepHeEnson, J.—Oligocheta, in: Zoological Results of the Abor Expedition.

Rec. Ind. Mus., vol. viii. 1914.

19. i On some Indian Oligocheta, mainly from Southern India and

Ceylon. Mem. Ind. Mus., vol. vi. 1915.

20. be On a collection of Oligochzeta from various parts of India

and Further India. Rec. Ind. Mus., vol. xi. 1917.

ON THE STRUCTURE OF THE REPTILIAN TARSUS. 143

6. On the Stracture of the Reptilian Tarsus.

By R. Broom, D.Sc., F.R.S., C.M.Z.S8.

(Received November 9, 1920: Read February 22, 1921. }

(Text-figures 1-27.)

In 1864 Gegenbaur established the general homology of the

elements of the tarsus with those of the carpus, and gave us a

series of names for the elements which have been used by most

later writers.

The typical carpus in the higher forms was shown to be made

up of a radiale and an ulnare with an intermedium between them,

a centrale near the middle of the carpus, and five distal carpalia.

In the hind limb the tibia and fibula undoubtedly correspond

with the radius and ulna of the fore limb and the five distal tarsalia

as unquestionably agree with the five distal carpalia, but as the

proximal part of the tarsus usually has only three elements, there

has always remained some little doubt as to how to homologise

them with the four proximal carpals.

Gegenbaur considered that the two proximal elements of the

mammalian tarsus—the caleaneum and astragalus—corresponded

with the ulnare, the intermedium, and radiale of the fore limb,

and that they ought to be regarded as the fibulare and conjoined

intermedium and tibiale, while the third element, the navicular,

he looked upon as the centrale of the tarsus. As the inter-

medium and radiale are frequently united in mammals, it seems

very natural to conclude that the intermedium might be perma-

nently fused with the tibiale in the tarsus.

Gegenbaur’s view has been followed by the majority of later

comparative anatomists and paleontologists. When it was

seen, however, that there was no evidence from either paleeon-

tology or from the study of the skeletogenesis in favour of the

astragalus being a composite element, most workers came to favour

the view that the astragalus is the tibiale alone and that the

intermedium has been early lost, though some few preferred to

look on the astragalus as the intermedium and to consider that

it was the tibiale that was lost. At the present time, though

the large majority of authorities support the former view, the

question is by no means settled, and I think there are good

reasons to believe that the generally accepted view is a mistaken

one. Within recent years, paleontology has given us so much

new light that it seems necessary to reopen the question.

If we had well-preserved tarsi of all the Carboniferous and

Permian amphibians and reptiles known, there would be no

difficulty in giving the complete evolutionary history of the

tarsus in its later stages. Unfortunately, the tarsus of most of

the early Tetrapods remained largely cartilaginous, and even

where the elements are ossified it is rarely that we find them in

144 DR. R. BROOM ON THE

undisturbed positions. Still, we havea few early tarsi sufticently

well preserved to suggest to us the main lines of evolution.

The most primitive Tetrapod tarsus known is that of the

Temnospondylous amphibian, 77ematops milleri Williston, from

the Lower Permian Beds of North America (text-fig. 1). Here we

find the tarsus composed of four large proximal elements, one

of which is situated centrally, five distal tarsalia and taree centralia

lying above the first, second, and third tarsalia. The element

articulating with the end of the tibia we ought, I think, to call

the tibiale, even though it is not, as I hope to show, the same

element as articulates with the tibia in the higher forms. The

other two proximal elements must be regarded as the fibulare and

the intermedium. ‘These determinations are those of Williston,

and it is difficult to see how they can be disputed. Williston

points out that there has been a passage for vessels between the

fibulare and the intermedium,

Text-figure 1. Text-figure

Text-fig. 1.—Right tarsus and metatarsus of Trematops milleri Williston *,

slightly modified after Williston.

Text-fig. 2.—Right tarsus and metatarsus of Uranocentrodon senekalensis

vy. Hoepen*+. The tarsal elements are figured in true relative position,

as seen in two different specimens. Letters mdicate the probable

position of the elements which have remained cartilaginous.

** An American Lower Permian Temnospondylous Amphibian.

+ A South-African Upper Permian ‘Temnospondylous Amphibian.

Our South African Upper Permian Temnospondylous form,

Uranocentrodon senekalensis v. Hoepen. (text-fig. 2), gives us a

further development of the amphibian tarsus. Though the tarsus

is here imperfectly ossified, we have two specimens with the

elements in almost undisturbed relations, so that we can be fairly

sure of their determinations. As will be seen from the figure

I give, there are three large, proximal, well-ossified elements and

three imperfect ossified distal elements, If we had not the clue

afforded by Z'rematops, we might determine the proximal elements

fo)

as the fibulare, intermedium,and tibiale. We might perhaps, even

STRUCTURE OF THE REPTILIAN TARSUS. 145

more readily regard them as the fibulare, tibiale, and centrale

from their superficial resemblance to the calcaneum, astragalus,

and navicular of the mammalian tarsus. It is, however, quite

manifest that, whatever the tibia supports, it is not the element

lying on the tibial side of tle fibulare. We can readily see that

this element is the one which we have identified as the inter-

medium in Zrematops. It is similarly situated, and it has

between it and the fibulare a passage for vessels exactly as in

Trematops. The element situated distal to this intermedium is

manifestly the proximal centrale. The tibiale has remained, like

the majority of the other elements of the tarsus, cartilaginous.

The other ossified elements of the tarsus are figured as they occur.

IT identified them as the first and fourth distal tarsals and one of

the centrales. Haughton, in describing the tarsus in the Bloem-

fontein specimen, regards the distal elements as the first and

third, but the Pretoria specimen has the element in undisturbed

Text-figure 3. Text-figure 4.

Text-fig. 3.—Right tarsus and metatarsus of Scincosaurus crassus Fritsch *.

After Jaekel.

Text-fig. 4.—Right tarsus and metatarsus of Limnoscelis paludis Williston +.

The positions of the missing elements, which were probably carti-

laginous, are indicated by letters.

* A Microsaurian amphibian.

+ A primitive Cotylosaurian reptile.

articulation with the head of the fourth metatarsal. Haughton

agrees in regarding the third small ossification as the centrale.

When the cartilaginous elements are restored, it will be seen

that the tarsus is almost exactly similar to that of Trematops.

The most interesting feature of the Uranocentrodon tarsus

is the tendency that it exhibits of the elements on the tibial side

to become reduced.

The next tarsus which we are able to study is that of the Micro-

saurian Scincosaurus crassus Fritsch, (text-fig. 3). Whether the

Microsauria are to be regarded as reptile-like amphibians or as

amphibian-like reptiles need not at present concern us, as it is

agreed by most that if they are amphibians they are apparently

the nearest to the primitive reptiles. 1t is extremely fortunate

Proc. Zoou, Soc.—i921, No. X. 10

146 DR. R. BRUOM ON THE

that we have this tarsus well preserved, as it shows us how the

reptilian tarsus has been derived from the primitive amphibian

type. As figured by Jaekel, it has three proximal elements and

four distal. The elements articulating with the fibula are

manifestly those we have identified in the more primitive type

as fibulare and intermedium, and they are so identified by Jaekel.

The third proximal element articulates with the tibia and is

manifestly the tibiale. It is much smaller than the other

proximal elements. ‘The distal elements are the first four distal

tarsalia. Between the fibulare and the intermedium is the

tarsal foramen.

The great difference between the Microsaurian tarsus and that

of the Temnospondylous types is that in the former all the

central elements have disappeared or become generally reduced

and cartilaginous.

In the earliest undoubted reptiles known—the Cotylosauria—

the tarsus has only been preserved for us in a very few forms.

In Limnoscelis paludis Williston (text-fig. 4), though the tarsus

is not perfectly preserved and was probably largely cartilaginous,

we have the two principal elements preserved in position. Willis-

ton identifies them as the fibulare and the united tibiale and

intermedium, but it seems much more probable that they are

the fibulare and the intermedium, and that the tibiale is either

lost or was cartilaginous, and this latter view is admitted by

Williston as not impossible.

Another very primitive type of which we know the tarsus is

Hosauravus copet Williston (text-fig. 5), from the Middle Penn-

sylvanian of North America. Unfortunately, the head of this

animal is unknown, and weare thus in doubt whether or not it isa

Cotylosaur. The tarsus has the elements preserved in only slightly

disturbed relations. There are two large proximal elements, which

a comparison with other early types leads us to consider as the

fibulare and intermedium. ‘There are five distal tarsals, and a

small element on the tibial side of the tarsus which is

probably the reduced tibiale. The metatarsals are somewhat

displaced, and it is not improbable that the distal tarsals and

the tibiale are also a little displaced.

In Seymouria baylorensis Broili (text-fig. 6) the nearly perfect

tarsus has been discovered by Williston. It consists of two large

proximal elements and a third small one, and apparently five distal

tarsals. The two large tarsals are regarded by Williston as the

fibulare and tibiale, and the small proximal element the centrale.

T interpret them as in Scinccosawrus—the fibulare, intermedium,

and tibiale.

The only other Cotylosaur in which the tarsus is satisfactorily

known is Procolophon trigoniceps Owen (text-fig. 7), and though

Procolophon in having a roofed temporal region is usually placed

with the Cotylosaurs, yet, being a late Triassic fo:m, it has

advanced in many respects so far from the Cotylosaurs of the

Permian of North America that it ought, perhaps, really to be

STRUCTURE OF THE REPTILIAN TARSUS. 147

placed in a distinct order—the Procolophonia. The tarsus is known

with the elements in undisturbed position. There are two ossified

proximal elements and four distal tarsalia. The two proximal

elements have a passage between them, and are manifestly the

homologues of the large elements in Scincosawrus and thus the

fibulare and intermedium. There has possibly been a cartila-

ginous tibiale, which is not preserved, as it seems necessary to

have an element to the tibial side of the intermedium to support

the first tarsale. Goodrich’s figure of the Procolophon tarsus,

which is taken from Watson’s much-reduced restoration, gives

rather a misleading idea of the structure. In Procolophon the

radiale in the carpus has evidently been cartilaginous, and it is

thus not at all remarkable that the corresponding tibiale in the

tarsus should also remain cartilaginous.

When we follow the line of mammalian descent through the

American Pelycosawrs and allied forms and through the South

African Therapsids, we have as many well-preserved tarsi as we

require.

Text-figure 5. Text-figure 6. Text-figure 7.

Text-fig. 5—Right tarsus and metatarsus of Hosauwravus copet Williston*. After

Williston. The distal tarsals are evidently somewhat displaced. The large

oval element between the intermedium and the first metatarsal is probably

the displaced tibiale. The smaller element lying proximally to it is probably

the first tarsal. The largest distal tarsal is probably the fourth tarsal.

Text-fig. 6.—Right tarsus and metatarsus of Seymouria baylorensis Broilit. After

Williston.

The third and fourth tarsalia are lost.

Text-fig. 7.—Right tarsus and metatarsus of Procolophon trigoniceps Owen tf.

From a specimen in the Albany Museum. ‘The elements are preserved in

almost undisturbed relations. There was most probably a small cartilaginous

tibiale in the position indicated by the letter ‘‘t.”

** A primitive Reptile of unknown affinity.

+ A primitive Cotylosaurian reptile.

t A late Upper Triassic Cotylosaurian reptile

The most primitive type we know is Ophiacodon miraus Marsh

(text-fig.8). Here there are two large proximal elements, mani-

festly those which become the caleaneum and astragalus of the

mammal, with five distal tarsals and two small elements lying distal

to the astragalus. The caleaneum and astragalus are manifestly

again the two elements which in lower types we have identified as

fibulare and intermedium ; and the five distal tarsals present no

IO):

148 DR. R. BROOM ON THE

difficulties. The two elements in the centre of the tarsus,

however, may readily give rise to difference of opinion. By

Williston, Case, and others who have written on the type they

are regarded as first and second centralia. The inner of the two

is the element which becomes the mammalian navicular. The

outer one is an element which becomes early lost, and is only

known in a very few Permian forms.

If we identify, as I think we must do, the astragalus with

the intermedium, we must either regard the tibiale as lost, or find

it is the inner of the two supposed central elements. This inner

element supports the first tarsal, and though it appears to have

shpped away from the tibial articulation, it is still not far

removed from the tibia. If we are right in identifying the inner

proximal elements in Scincosaurus and Seymouria as the tibiale,

Text-figure 8. Text-figure 9.

Text-fig. 8—Right tarsus and metatarsus of Ophiacodon mirus Marsh *

After Williston. Slightly modified.

Text-fig. 9.—Right tarsus and metatarsus of Casea broilii Williston +.

After Williston.

* An early Theromorph.

+ An aberrant Theromorph.

then there is good reason to believe that the navicular of the

Pelycosaurs, the Therapsids, and the Mammals is also the

tibiale which by the lengthening aud narrowing of the targus

has become slightly ditered 3 mn position. In the Cotylosaurs the

distal tarsais are nearly twice as wide as the fibulare and the

intermedium. In the more active Pelycosaurs the tarsus has

become so narrowed that the distal tarsals together measure often

less and rarely much more than the width of the two large

proximal elements. If the tibiale is to be retained at all it can

only be by becoming wedged in between the intermediim and the

first and second tarsalia. This, I believe, is what has happened ;

STRUCTURE OF THE REPTILIAN TARSUS. 149

and itis remarkable, as I hope to show presently, that a similar

shifting of the tibiale is also seen in Sauropsida, and perhaps

it has arisen independently in this group.

In Casea broilit Williston (text-fig. 9) the small central element

has disappeared, and we thus have a tarsus that, except for

retaining the fifth tarsale, is essentially mammalian in structure.

In Varanops brevirostris (Williston) (text-fig. 10) the tarsus is

like that of Casea and Ophiacodon, except that not only has the

centrale disappeared, but the tibiale is evidently cartilaginous.

In the Therapsids the tarsus is almost typically mammalian

in structure. ‘lhere is never a centrale, and the tibiale is always

placed, as is the navicular in the mammal, between the inter-

medium and the first and second tarsals.

In Anomodonts the tibiale is frequently cartilaginous either

wholly or in part. Many years ago I figured the tarsus in a

small form which I referred to Udenodon gracilis Broom. Shortly

afterwards I found that the skeleton belonged to the same anima!

Text-figure 10. Text-figure 11. Text-figure 12.

Text-fig. 10—Right tarsus and metatarsus of Varanops brevirostris (Williston).

A primitive Pelycosauroid reptile. After Williston.

Text-fig. 11.—Right tarsus and metatarsus of Hmydopsis trigoniceps (Broom), a small

Anomodont reptile. The drawing is mainly from the specimen in the Albany

Museum, but partly restored from other specimens. The condition of the

tibiale varies greatly in Anomodonts. In some it is completely cartilaginous,

ae others well ossified. 1t probably affords part of the articulation for the

tibia.

Text-fig. 12.—Right tarsus and metatarsus of Galechirus scholtzi Broom, a smal]

Dromasaurian reptile. The elements are figured as found, but the inter-

medium is evidently slightly rotated.

as the skull which I had called Oudenodon trigoniceps Broom.

We now know that this small Anomodont has a few small molars,

and must be placed in a new genus, Linydopsis. I give a new figure

of the tarsus (text-fig. 11). The interesting point about it is

that the tibia probably articulates with the tibiale. In another

small Anomodont tarsus I have belonging to an undescribed

species, the tibia also appeared to articulate with the tibiale.

In the more mammal-like forms—the Gorgonopsians and the

Cynodonts—the tibia and fibula articulate only with the inter-

medium and the fibulare.

150 DR. R. BROOM ON THE

The most primitive known tarsus of the lizard-like group is

that of Broomia perpleca Watson. Though the top of the skull of

this animal is unknown, almost all the rest of the structure is

known, and fortunately the tarsus is almost perfectly preserved.

As in the large majority of early reptiles, there are two large

proximal elements. There are five distal tarsals and two other

small elements. The two proximal elements have the usual

foramen between them, and are doubtless the fibulare and inter-

medium. The two small elements are regarded by Watson as

the first and second centralia. I regard the mner one as the

imperfectly ossified tibiale. The distal end of the tibia as found

is in a position to articulate with what I regard as the cartila-

ginous position of the tibiale. Watson, in his restoration, shifts

the tibia to make it articulate with the intermedium, which he

regards as the fused intermedium and tibiale. If we articulate

the tibia as Watson has done, the three first digits seem

practically without any proximal support—a condition which

Text-figure 13. Text-figure 14. Text-figure 15.

yw Gr)

rae

Text-fig. 13.—Right tarsus and metatarsus of Paleohatteria longicaudata Credner.

A primitive Permian reptile of doubtful affinity. Regarded by Williston,

Watson, and others as a Theropsidan: by the writer and others as an early

Sauropsidan. After Jaekel.

A cartilaginous tibiale was probably present.

Text-fig. 14.—Right tarsus and metatarsus of Mixosaurus nordenskjoldi Hulke.

A Triassic Ichthyosaur. After Wiman.

Text-fig. 15—Right tarsus and metatarsus of Stereosternum tumidum Cope.

A Lower Permian Mesosaurian. After M‘Gregor.

seems very improbable. Thesecond small element is doubtless, as

Watson holds, a centrale. I give a figure of the tarsus as found,

as restored by Watson, and as I am inclined to restore it (text-

figs. 16-18).

There is another primitive reptile which one wishes one knew

more about. I refer to Paleohatteria longicaudata Credner.

Though first described over thirty years ago, and apparently

known by very satisfactory and nearly complete skeletons, we are

still in much doubt about the animal and its affinities. For many

years it was believed by every one to be a primitive two-arched

STRUCTURE OF THE REPTILIAN TARSUS. 151

reptile allied to Sphenodon. Buta few years ago Williston came

to the conclusion that it was a‘Pelycosaur or a near ally, and

Watson holdsa similar opinion, stating that “ it now seems almost

certain that Paleohatteria is really a Therapsid.” It is difficult

for one who has no chance of seeing the actual types to know

which authorities he ought to follow, and though the tendency is

always to follow the latest, I personally do not feel at all satisfied

that Williston and Watson are right in this matter. The skull

as restored by Jaekel—and his restoration seems to me the best

we have yet had—is very unlike that of any Pelycosaur or

Therapsid: the shoulder-girdle is entirely unlike and the pelvis

is only a little like. The humerus also has only a very shght

resemblance to that of a Pelycosaur or Therapsid. Fortunately

the tarsus is fairly well preserved, and, as figured by Jaekel, has

two large proximal elements, which he considers, as also I do, to

be the fibulare and intermedium (text-fig. 13). The only other

ossified elements are the five distal tarsals. Possibly there has

been a cartilaginious tibiale. If so, the only difference between

Text-figure 16. Text-figure 17. Text-figure 18.

Text-fiz. 16.—The right tarsus and metatarsus of Broomia perplewa Watson.

A primitive Permian Sauropsidan. The remains as found. Left reversed.

After Watson.

Text-fig. 17.—The tarsus and metatarsus of Broomia as restored by Watson.

Text-fig. 18.—The tarsus and metatarsus of Broomia as restored by the writer.

the tarsus of Palechatteria and that of Broomia would be that

the former had lost the small centrale.

On the evidence of the published figures of Paleohatteria 1 am

inclined to regard it is a primitive Diapsid reptile, a little more

advanced than Broomia and a little more primitive than Youngina.

Youngina is the only known Permian Diapsid in which the

tarsus is fully ossified and almost perfectly preserved (text-

figs. 19 & 20). In this tarsus there are two large proximal

elements—the fibulare and intermedium with the tarsal foramen

between them, five distal tarsalia, and a large element situated

between the intermedium and the first, second, and third tarsalia,

which I believe to be the tibiale.

The fibulare is a flat bone which has a very distinct heel

process. ‘The intermedium is very large and with a large

152 DR. R. BROOM ON THE

articular surface for the tibia. It articulates with the fibulare,

the tibiale, and the fourth tarsal. The tibiale is relatively small,

and is wedged in between the intermedium and the first, second,

and third tarsalia, Though the tibia does not articulate with

it in most positions of the foot, it seems probable that it does

when the front is turned inward, as is manifestly possible. The

three first distal tarsals are all small but well ossified. The

fourth is very large; its relations will be seen in the figure

given. The fifth tarsal is moderately large and flat; it

articulates with the fourth tarsal by a suture which allows very

little movement between the two.

Text-figure 19. Text-figure 20.

(

Text-fig. 19.—Left tarsus and metatarsus of Youngina capensis Broom *, as pre-

served. The foot has the plantar surface displayed, and the metatarsals

are somewhat distorted. Twice nat. size.

Text-fig, 20.—Right tarsus and metatarsus of Youngina capensis Broom, viewed

from the dorsal side, with the metatarsus restored in position. The

tibiale, first, second, third, and fourth tarsals of the right side are pre-

served in position, and have their dorsal aspects displayed. The fibulare

aud part of the intermedium of the left side have had their dorsal surfaces

exposed and are added to the drawing reversed.

* A Permian two-arched reptile of the order Eosuchia.

The metatarsals are long, slender, and moderately straight

bones. The fifth metatarsal, which we should have expected to

be of the Sphenodon hooked type in this undoubted two-arched

reptile, is a long slender bone, nearly as long as the fourth meta-

tarsal, and it shows no trace of the peculiar hooking. The upper

end is expanded, and the outer process probably was attached to

the fibulare by a ligament.

The remarkable points in the foot of Youngina are this

primitive generalized fifth metatarsal, and the fact that the tarsus

STRUCTURE OP THE REPTILIAN TARSUS. 153

is almost typically Therapsid or even mammal-like. A few years

ago I figured a well-preserved tarsus under the name Galesphyrus

capensis, believing it to be a Dromasaurian. The few points in

which it differs from the Dromasaurians are points in which it

agrees with Youngina, so that it is much more likely that it

is an Kosuchian.

Another very interesting tarsus is that of the South African

Upper Triassic Rhynchosaurian, Howesia browni (text-fig. 21).

The tarsus is almost perfectly preserved though doubled over, and

the restoration I give is probably nearly correct. There is a

large fibulare with a heel process, a large intermedium, and a

smaller tibiale. There are four distals, and the fifth metatarsal

has the Sphenodon-like specialization.

The tarsus of the adult Sphenodon is well known, and Howes

and Swinnerton have given us something of the embryonic

Text-figure 21. Text-figure 22. Text-figure 23.

Text-fig. 21—Right tarsus and metatarsus of Howesia browni Broom; a Triassic

Rhynchosaurian.

Text-fig. 22.— Hight tarsus and metatarsus of an embryo of Sphenodon punctatus

Gray. ‘The embryo is of Dendy’s stage R. Ossification has commenced in

the metatarsals. Though the proximal tarsal cartilaginous mass shows no

clear evidence of its nature at this stage, earlier embryos show that it is

composed of three elements, and I think only three.

Text-fig. 23.—Right tarsus and metatarsus of an embryo of Testudo sp. The

embryo is of the stage where chondrification is well advanced, but where

ossification has scarcely begun. The very marked difference between this

tarsus and that of the Sphenodon embryo is of interest.

condition. Only those who have studied developing cartilage

and procartilage know the difficulties of this mode of research.

At times we get remarkable results, but too often they are

inconclusive. I give a figure (text-fig. 22) of the tarsus in a

young embryo of Sphenodon, which I was able to examine through

the kindness of Prof. Dendy. Though ossification of the meta-

tarsals is just commencing at this stage, there is no clear evidence

of the composite nature of the large proximal element. Howes

and Swinnerton showed that there was evidence at an earlier stage

154 DR. R. BROOM ON THE

of its being made up of a fibulare, an intermedium, a tibiale, and

a centrale. I think it more probable that thereare only in it the

fibulare, intermedium, and tibiale. The evidence for a centrale

is not at all satisfactory. If, however, it is really the conjoined

fibulare, intermedium, and tibiale, then the whole tarsus becomes

strikingly similar to that of Howesia.

I give a figure of the tarsus of a very young embryo of

Testudo sp. (text-fig. 23). Here there are only seen two

proximal elements and four distal tarsals. There is no evidence of

the larger proximal elements being more than a single element,

and I am inclined to regard it as intermedium alone. Other

Chelonians are known to have an additional element between the

larger proximal element and the first and second tarsals.

Text-figure 24. Text-figure 25,

Text-fig. 24.—Right tarsus and metatarsus of Plesiosawrus rugosus. After Owen.

Text-fig. 25.—Right tarsus and metatarsus of Peloneustes philarchus Seeley.

After Andrews.

Goodrich considers this to be the centrale. JI regard it as the

much reduced and displaced tibiale. The fifth metatarsal is

shortened up.

In most of the later Diapsidans we find a tarsus which is

either of the Sphenodon type or a modification of it.

In the aquatic reptiles we find many interesting types.

Ichthyosaurus is too specialized to be of much morphological

interest, but the Triassic form, JMxosaurus, shows us how the

Ichthyosaurian paddle has originated. I believe the tarsus to

be made up of fibulare and intermedium with the five distal

tarsals, and to be thus almost identical with the tarsus of

Mesosaurus or Stereosternum, which there is some reason to

consider as perhaps its nearest allies.

STRUCTURE ON THE REPTILIAN TARSUS. 155

The Plesiosaurian tarsus (text-figs. 24, 25) presents little

difficulty. The fibulare and inter medium are well ossified, but

in the earlier forms the tibiale is often partly or wholly cartila-

ginous. In later types the tibiale is well developed. ‘here are

only three distals, which I believe to be first, third, and fourth.

The fifth metatarsal 1s shortened up as in Chelonians.

The Pythonomorphs show a most interesting type of tarsus.

In Mosasaurus (text-fig. 26) there are only three tarsal elements,

which are manifestly the fibulare, the intermedium, and be

fourth distal tarsal. In Platecarpus (text-fig. 27) there are four

tarsal elements— the fibulare, iatonmcdiurns and the third and

fourth distal tarsals. In both types there is a specialized fifth

metatarsal.

Text-figure 26. Text-figure 27.

Text fig. 26.—Right tarsus and metatarsus of Mosasaurus lemonnieri Dollo.

A European Pythonomorph. After Dollo.

Text-fig. 27.—Right tarsus and metatarsus of Platecarpus abruptus Marsh.

An American Pythonomorph. After Williston.

In the reptilian and amphibian tarsus the most remarkable

features are the almost constant presence of the fibulare and the

intermedium, and the great variability of the tibiale. Rarely is

the tibiale a large element: very frequently it remains entirely

or partly cartilaginous. In many types it is completely absent.

In the mammal-like reptiles and in the primitive Diapsidans it is

wedged in between the intermedium and the first and second

tarsalia.

The central elements, of which there are four in some

amphibians, are early greatly reduced and lest. The only one

which for a time remains in early reptiles is apparently the

homologue of the proximal centrale in Trematops. In only a few

Permian forms is it still present, and in no Triassic or later reptile

is there any trace of it.

Io Zoo WI; SONINIWNG, IE I.

ANATOMY OF THREE-TOED SLOTH (Bradypus tridactylus).

Pi Zao. | 1921) SON NDAG. = bileiis

ANATOMY OF THREE-TOED SLOTH (Bradypus tridactylus).

PAZ So 92s ON INGA Grae ealeile

ANATOMY OF THREE-TOED SLOTH (Bradypus tridactylus).

Pe Za Se 1921 SOININTA GS selves

“ANATOMY OF THREE-TOED SLOTH (Bradypus tridactylus).

ON THE ANATOMY OF THE THREE-TOED SLOTH. 157

7. A Contribution to the Anatomy of the Three-toed Sloth

(Bradypus tridactylus). By Cnas. FE. SonntaG, M.D.,

i'.Z.8., Anatomist to the Society.

[Received November 16, 1920: Read February 8, 1921. |

(Plates L.-IV. & Text-figures 10-15.)

The observations recorded here were made on the body of a

female Bradypus tridactylus which died a few hours alter admis-

sion to the Society’s Gardens. It was deposited, so I was unable

to make a complete dissection, and my investigations were, in

consequence, limited to the mouth, tongue, and organs of the

thorax and abdomen. Some of the conditions obser ved have not

been recorded before, and others difter from those which have

already been described. The organs were compared with

those of B. gularis or cuculliger, which are preserved in the

Museum of the Royal College of Surgeons,

THe Moors.

The Vestibule.

The form and communications of the vestibule depend on the

shape of the maxille and the size and distribution of the teeth.

When the skull is examined it is seen that the premaxille are

edentulous, so there is a gap in front between the first pair of

teeth in the upper jaw (Plate I. C), and there is a similar gap in

the lower jaw. Consequently, when the lips are everted, one can

see the tip of the tongue. In Cholepus, on the other hand, the

prominent rostrum, or beak, of the lower jaw diminishes the gap

in the centre.

The mucous membrane does not form a vestibular pocket in

front, as in most mammals, for it passes direct from the margins

of the jaws to the free edge of the lips. Consequently, “the

vestibule in front has no wider area than the orifice of the lips

(Plate I. A).

The first pair of upper teeth are small, and the second pair are

large and project laterally (Plate I. C). Between the mid-line

in front and the second pair of teeth the upper jaw expands, but

it narrows behind them, and, as the cheeks are not closely applied

to the sides of the jaw, the vestibule is patulous and increases in

width from before backwards (Plate I. B, v). In the ease of the

lower jaw, however, the cheeks are closely apposed against the

gum and the vestibule is reduced to a mere slit.

Posteriorly the communications between the vestibule and

mouth-cavity behind the last pairs of teeth in both jaws are

exceedingly small.

In Cholepus the vestibule and mouth-cavity communicate

through the gaps between the first and second pairs of teeth in

both jaws (Plate I. D),

158 DR. C. FE. SONNTAG ON THE

The Hard Palate (Plate I. B).

The hard palate is 2°7 em. long, ‘8 cm, wide between the first

pair of teeth, 1 cm. wide between the second pair of teeth, and

‘7 cm. wide between the last pair, so it first expands and then

contracts. It is encircled just internal to the teeth by a series of

palatal tubercles which, however, have no counterparts on the

dorsum of the tongue. On each half of the palate there are

seven of these—one opposite each tooth, one anterior to the first

tooth, and one at the side of the mid-line: the first and second

are borne by the premaxille. The tubercles increase in size from

before backwards, and the fifth pair is the largest.

The palate has the same chocolate colowr as the labial margins,

Running along the mid-line is a white streak which sends out

branches to end on the mesial sides of the tubercles. In the

posterior part this streak overlies the crest shown in Plate I. C.

Vhen the mucosa is removed from the subjacent bone, it is

seen that the palatal tubercles do not cover bony eminences, and

the bone differs greatly from that of Cholepus (Plate I. D).

In B. tridactylus there is a strong median crest on the posterior

half, and there is a gutter on each side of that ridge. Anteriorly

each gutter is converted into a tunnel by a fenestrated bridge of

bone; the tunnel on the right side reopens on to the free

surface, but the left one does not. Both gutters, however, com-

municate posteriorly with the nasal fosse.

In the mid-line in front there is a groove which widens from

behind forwards, but, as the premaxille were lost in the pre-

paration of the skull, lam unable to give a complete description

of it.

The bone is much perforated by small foramina. Some of

these open into the afore-mentioned tunnels and the others into

the nasal cavity.

No sutures are present outlining the various elements com-

prising the bony palate as in Cholepus.

The bony palate of Cholepus differs in many ways from that

of Bradypus. It is much larger and its constituent bones are

clearly outlined by sutures. There is no gutter nor ridge on the

posterior part, but many foramina of different sizes are present.

Of these, there is a large pair on the palate bones lying posterior

to the fifth pair of teeth, and a large pair on thie maxillee

in front. The former, probably, correspond to the gutters of

Bradypus.

The Soft Palate.

The soft palate, which measures 1:2 cm. from before back-

wards. is pink in colour. It is attached to the posterior extre-

mity of the hard palate in front, and to the prominent pterygoid

plates laterally. No hamular pterygoid processes are palpable

through it, but the convex free borders of the plates can easily be

felt. “Tts posterior edge is concave backwards, and its oral sur face

ANATOMY OF THE THREE-TOED SLOTH. 159

bears a small tubercle in the mid-line at a distance of 1 mm.

from the free edge (Plate I. B). There is, however, no uvula, as

was pointed out by Rapp (10).

The Teeth.

The teeth have been fully deseribed in their structural and

developmental aspects by Rapp (10), Semon (11), and others, and

T have nothing to add to their accounts.

The Tongue (text-fig. 10).

Rapp (10) has given some scanty details of the tongue, so a

detailed description i is required.

My fresh specimen has the following measurements :—Length

from the apex to the epiglottis, 3°85 cm.; length of the oral part,

2°89 em.; length of the pharyngeal part, 96 cm.; greatest width

(7. e. between the lingual attachments of the anterior faucial

pillars), 1-7 em.; width of the apex, 7 cm.; thickness in the

vallate papillary region, 1-25 em.; thickness at a point 1°4 cm.

posterior to the apex, 1-1 em.; thickness of the apex, ‘4 cm.

The tongue is, therefore, short, narrow, and thick. Its width

first decreases rapidly, but later maintains a constant dimension

to the apex. ‘The thickness remains almost uniform, on the

other hand, in the posterior two-thirds and then rapidly decreases

towards the apex.

The apex has a fine mesial notch, and is covered with small

conical and large flat fungiform papille. There is a faint mesial

sulcus which, however, 1s irregular, and the lens shows how it is

replaced in parts by fungiform papille ; the latter are, however,

invisible to the naked eye. The lateral borders ave massive, and

have a single row of large fungiform papille; they also possess

vertical rows of conical papillae with backwardly-directed points.

No lateral organs ave present, as shown by Gmelin (13).

The two circwmvallaie papille lie side by side, and both are

circular and polished. When the tongue is fresh theiv free sur-

faces are flush with the surface of the tongue. When, however,

it is placed in preserving fluid the papille are retracted within

their fosse, thus demonstrating an action similar to what occurs

in Monotremes and some Marsupials. The vallums are promi-

nent, granular, and surrounded by conical papillae. Mayer (14)

saw three papillee.

The fungiform papille ave large, flat, and not numerous. They

have the usual arrangement in clusters and rows of varying

degrees of obliquity which characterises the Mammalian tongue.

They extend from the apex back beyond the vallate papille, but

they stop short of the zone of lymphoid tissue on the base. They

form a single row on the lateral borders of the tongue, but the

papilla of the row are discrete.

The conical papille are all filiform and possess one or more

points; those in the mid-line have their points directed back-

wards, but those at the sides look backwards and inwards, They

160 DR. C. F. SONNTAG ON THE

increase in size from before backwards and from without inwards,

and they are discrete, so there is no overlapping. ‘They extend

beyond the vallate papille, and there is a sharp line of demar-

cation between them and the lymphoid tissue at the base.

On the base of the tongue there are many lymphoid nodules,

some of which have central apertures, and there is a line, convex

backwards, separating the papillary and lymphoid areas.

Text-figure 10.

tin

ity

Wa tits

The tongue. A: inferior surface. B: dorsum. C: lateral view.

No lyttéa is present as Rapp showed (10).

The inferior surface has a narrow papillary border. There is a

fine mesial notch, but no mesial ridge, suleus, nor frenum. The

surface is marked by fine horizontal ridges, but there are no

plice fimbriate. The papillary border forms thick masses at

either side of the apex.

The tongue, according to Rapp, is mechanical rather than

gustatory in function.

ANATOMY OF THE THREE-TOED SLOTH. 161

THE STOMACH.

The external appearances have been described by Cuvier (8),

Meckel (7), Carus (2), Otto, Boulart, and Pilliet (9), and the

internal appearances of the stomach of B. cweulliger. have been

described by Rapp (10) and Klinckowstrém (6). Many details

can be added, however, to their accounts. Moreover, the internal

appearances and the relative positions of the various compart-

ments are different in B. tridactylus and B. cuculliger or gularis.

The stomach bed in my specimen is unusual, for the position of

the pancreas is peculiar; that organ is contained within the

duodenal loop and none of it lies dorsal to the stomach. Again,

the stomach bed does not contain the spleen, for the latter, in

Sloths, lies on the right side of the pylorus. The kidneys lie far

back in the abdomen, and the suprarenal capsules are separated

from them, but lie in the normal position. Consequently, the

adrenal bodies are dorsal relations of the stomach, but the

kidneys are posterior. In my specimen the stomach has the

following relations :—

Anterior—liver and diaphragm.

Posterior —intestines and kidneys.

Ventral—ventral abdominal wall.

Dorsal-—vertebral column, main blood-vessels, suprarenal cap-

sules, root of the mesentery.

To the left—abdominal parietes.

To the right—spleen, pancreas, duodenum.

These peculiar arrangements make the disposition of the

peritoneum, which is described below, noteworthy.

In my specimen the gravid uterus touched the greater

curvature.

Divisions of the Stomach.—I agree with Klinckowstrém’s

division of the stomach into three groups of compartments :—

1. The Paunch, or Fundus Stomach, with its cecal appendage

(Plate WAc cland| Byimrr):

2. The Cardiac Stomach with three divisions (Plate IT. A, p

end IEE aeiced. 2)

3. The Pyloric Region composed of two parts—glandular and

muscular (Plate IT. B 3).

In B. tridactylus the main divisions, and most of the sub-

divisions, can be distinguished on the surface. The paunch is

marked off from the cardiac stomach by a notch on the left part

of the greater curvature (Plate II. A, a) and a faint ridge running

from the notch to the base of the cxcal appendage. This ridge

marks the anterior limit of a number of fissures running forward

from the greater curvature, but these are not so deep as in

B. cuculliger. The cardiac stomach is easily marked out from the

pylorus; it is capacious and has thin walls, but the pylorus is a

thick, muscular, U-shaped tube.

Proc. Zoon, Soc.—-1921, No. XI. 11

162 DR. CG. F. SONNTAG ON THE

The relative positions of these divisions are different in

B. tridactylus and B. cuculliger. The reader is referred to the

works of Rapp (10), Klinckowstrém (6), and Oppel for descrip-

tions of the latter. Klinckowstrém’s paper is the best, for it is

profusely illustrated. Moreover, there is an excellent specimen

of the stomach of B. gularis in the Museum of the Royal College

of Surgeons. Klinckowstiém described the histology.

When the stomach of 2B. tridactylus is examined from the

ventral aspect (Plate IT. A), one sees the paunch (Plate IT. A, c)

posterior and to the left; the cardiac stomach (Plate IT. A, p)

anterior and to the right; and the ventral part of the pylorus

(Plate il. A, H) lying most to the right. The entire pylorus is

not visible on the ventral surface as in B. cuculliger. When the

stomach is viewed from the right, one sees the U-shaped pylorus

consisting of dorsal and ventral limbs, and a posteriorly-placed

bend (text-fig. 14 A, py). The dorsal limb emerges from the

cardiae stomach, and the ventral limb communicates with the

duodenum. In B. cuculliger the limbs are anterior and posterior

with the bend to the left. When the stomach is viewed from

the dorsal aspect, one sees the paunch posterior, the cardiac

stomach anterior, and the dorsal limb of the pylorus to the right.

The U-shaped pylorus is seen in Plate IT. B 3.

The simple, conical, ceecal appendage of b. tridactylus is longer

and more slender than in B. cuculliger. Its position may vary,

but in my specimen it first passed from the posterior and right

part of the paunch to the right. It was then recurved on itself,

and its apex rested on the cardiac stomach. It is also more

slender than in Cholepus.

No author has described the peritoneal sheet connecting the

cecal appendage to the paunch (Plate II. A, 8). Thisis triangular,

and fills up the space between them. Its apex lies along the

anterior border of the appendage, and there is a sharp free

border looking forwards and to the right. The right gastric

vessels run between its layers as they separate to surround the

appendage (Plate IT. A, 5).

The right gastric artery reaches the ventral surface of the

stomach to the right of the cesophagus and passes posteriorly and

to the right, It ends about the middle of the cecal appendage.

It is accompanied by the right gastric vein, and all the gastric

lymphatic glands lie alongside it.

Rapp (10) and Klinckowstrém (6) have described the internal

structure of the stomach of B. cuculliger, and that of B. tridac-

tylus agrees with it in most points. Their accounts, however,

can be amplified in several ways. Speaking generally, the cardiac

stomach has a mechanical function, the paunch and appendage

are secretory, and the pylorus has both properties.

The division between the paunch and cardiac stomach is

marked by a strong, thick ridge (Plate II. B, @) and a second

ridge into which the point of the pin is inserted, corresponding to

the white line seen externally on the ventral surface. It marks

ANATOMY OF THE THREE-TOED SLOTH. 163

the transition between the rugose smooth mucosa of the paunch

anil the hard mucosa of the cardiac stomach.

The three divisions of the cardiac stomach ave separated by

partitions, and the septum between the two ventral compart-

ments runs for some distance along the inner surface of the

paunch (Plate IT. B, pin). Microscopic sections of this septum

(Plate IV. B) show the transition between the stratified epithelium

of the cardiac stomach and the glandular lining of the paunch.,

The septum between the dorsal and left ventral compartments

(Plate IT. B, c) is small and strong. The former septum is absent

in Cholepus. The epithelial transition is similar to the junction ,

of the cesophagus and a simple stomach.

The ener surface of the paunch is beset with rugs whose

characters differ on the two sides of the septum running back

from the cardiac stomach. On the left side they run in all

directions and are mostly small; on the right side they are

mostly large, and pockets are enclosed between them and the

gastric wall. One very large septum bisects the cecal appendage;

in Plate II. B one of the halves is laid open, and a glass rod

passes down through the other one. Between the ruge, which

are soft and flabby, the mucosa has innumerable small glandular

pits.

On the right side there are several small, firm ridges passing

to the large ridge between the paunch and cardiac stomach, but

they are absent on the left side, for the soft mucosa extends right

forward to that ridge (Plate II. B, a); and these small ridges are

seen above d in Plate II. B.

The interior of the cecal appendix has been incompletely

described by Rapp (10) and Klinckowstrém (6), and no illus-

trations are provided by them. Its inner structure is shown in

Plate Il. B, and a cross section is diagrammatically represented

in text-fig. 12, B. It is ridged by many longitudinal folds, but

these are reduced to six in number in the terminal third. The

upper two-thirds of the cavity is bisected by a large septum, and

small partitions attached to it enclose two pockets on either side.

These all open towards the paunch, and the arrows in text-

fig. 12 B pass through the two passages on either side of the

main septum.

A prominent muscular ridge (Plate II. B,b) runs from the

ereater curvature into the left ventral compartment of the

cardiac stomach, and ends on the septum between the latter and

the dorsal compartment. A second ridge runs from the septum

between the two ventral compartments to meet the former ridge,

and between them is the opening between the dorsal and left

ventral compartments of the cardiac stomach.

The cesophagus is continued right into the pylorus by a groove

running through the left ventral ‘and dorsal compartments ‘of the

atomach, It is shown in Plate ITI. A, a, and its characters are

the same as those already described by Rapp and Klinekowstrém.

The pyloric region, which is washadedd has dorsal glandular

164 DR. GC. F. SONNTAG ON THE

and ventral muscular divisions. Its histological characters have

been thoroughly described (6), and I have nothing to add to that

account. Running along the floor is a ridged groove continuous

with the groove from the paunch into the cardiac stomach, and

Rapp (10) thinks it has a ruminating function. The communi-

cation between pylorus and duodenum is guarded by a strong

valve (Plate IIT. C, a), but Rapp states that there is no pyloric

sphincter.

Text-figure 11.

A: Interior of the duodenum. 4.p., bile papilla; p.p., pancreatic papilla; sp.p.,

papilla of accessory pancreas.

B: The spleen with accessory spleen (a) in. the pancreas (p).

( & D: The pancreas. ¢.b.d., common bile-duct; p.d., pancreatic duet ; m., mesen-

tery ; py., pylorus. Description of other figures in text.

E: Section across the suprarenal capsules. c.m., medulla within cortex. The upper

is the left, and the lower is the right capsule.

The stomach is, therefore, very complex and contains divisions

corresponding to most of those of the Ruminantia, but the

reticulum of these animals is absent,

ANATOMY OF THE THREE-TOED SLOTH. 165

Tue Duopenum (text-fig. 11, A).

I have nothing to add to those descriptions of the macroscopic

appearances which have already been published. JI must, how-

ever, record the conditions of the bile and pancreatic ducts

which differ from those mentioned by Francaviglia (4). The

pancreatic duct opens into the duodenum 2°9 em. distal to the

pylorus ; it is situated on a papilla. Ata point 1-1 cm. posterior

to the pancreatic papilla there is a large bile papilla, and 2-7 cm.

posterior to the latter there is a small papilla for the duct of an

isolated mass of pancreatic tissue. All the papille he on trans-

verse folds of mucosa. |

The entire duodenal mucosa is thrown into circular folds

which are large at the pyloric end, and small and numerous at

the jejunal end.

THe JEJUNUM AND ILEUM.

Dr. Chalmers Mitchell (8) has described the macroscopic

appearances in B, infuscatus, and mentions the presence of a

large cecal pouch and a small rudimentary cecum opposite to it.

The latter is not present in my specimen, but the remainder of

the intestine is the same as that described in Dr. Mitchell's

paper. There is no ileo-cxcal valve.

The mucosa exhibits alternating smooth and rugose areas

(Plate IV. B, a & 8), and there is a strong development of ruge

at the posterior end of the ileum.

Tue LARGE INTESTINE.

In Plate IV. A, c & p, the entire large intestine is exhibited,

but the lower end of the ileum is also shown in the former. The

canal is, therefore, short, and maintains an almost uniform calibre

till the rectum is reached, but the latter gradually expands and

its walls become progressively thicker and more muscular. At

the anus the walls are exceedingly thick and strong.

The whole of the colon and rectum are bile-stained, especially

in the posterior part of the latter, and these parts offer a marked

contrast to the pink ileum and anus.

The interior of the large intestine presents several interesting

appearances. The colon has many circular folds of varying sizes,

but the two most posterior ones form complete diaphragms. In

the centre of each of these there is an opening surrounded by

a sphincteric valve, the anterior one being the larger. In

Plate IV. B, p, a spicule of wood is passed through the two

openings. John Hunter (12) described three valves in the Two-

toed Sloth.

The posterior septum separates colon and rectum, and the

appearances of the mucosa differ in these two divisions. In the

latter it is excavated into a number of pockets which contain

pellets of faecal matter. ‘These are deepest in the posterior part of

166 ‘DR. ©. F. SONNTAG ON THE

the rectum where the bile-staining is deepest. It also appears as

if there had been a strong septum across the rectum, for there is

a ridge round the wall.

The anus is surrounded by a tough, strong sphincter, and the

mucosa is thrown into many longitudinal folds.

The structure of the whole alimentary tube must be taken into

account, in order to understand the physiological significance of

the conditions in the rectum. ‘The stomach is complex, as im

Ruminants, but the intestinal tube is short, so it is necessary

that there be some arrangement to obtain the maximum

absorption area in the latter. This is attained by the excavation

and folding of the mucous membrane of the latter, and the

sphincteric valvular openings in the septa across the colon only

allow a small amount of material to pass at a time into the rectum.

The powerful anal sphincter muscle is also required to ensure

a suificiently prolonged stay of the intestinal contents, for absorp-

tion of water (?) may take place slowly in the rectum.

The reticulum of the Ruminants is absent in the stomach of

the Sloth, and the hollows in the rectum resemble the pockets

of a reticulum.

THe Pancreas (text-fig. 11, c & D).

Rapp (10) and others state that the pancreas extends across the

abdomen dorsal to the stomach, but in my specimen it is almost

entirely contained within the duodenal loop; none of it lies

dorsal to the stomach. It presents features, therefore, which

resemble those of the Reptilia.

It is V-shaped and has splenic and duodenal limbs (sl and dl).

The former lies between the right border of the ventral limb of

the pylorus on the left and the left border of the spleen on the

vight. The latter lies along the left border of the duodenum, and

the angle of the V runs for a short distance anterior and dorsal to

the duodenal loop. <A short distance distal to the posterior end

of the duodenal limb there is a small isolated piece of pancreas (sp).

The entire organ is surrounded by peritoneum, the splenic limb

being enveloped by the layers of the dorsal sheet of the great

omentum, and the duodenal limb and isolated mass being included

between the layers of the common mesentery.

The splenic limb consists of a cylindrical massive part and

a lateral lamina, but the latter is wrapped round the former

(text-fig. 11, D). Within the massive portion there lies a small

spherical accessory spleen. The small isolated piece of pancreas

is oval in shape.

The pancreatic duct, contained within the gland, runs along

close to the anterior border, receiving tributary ducts as it goes.

When it emerges from the end of the duodenal limb it is crossed

at right angles by the common bile-duct. It opens into the

duodenum at the summit of a prominent papilla. Francaviglia

(4), however, saw the duct open along with the common bile- duct.

ANATOMY OF THE THREE-TOED SLOTH. 167

The small isolated piece of pancreas has its separate duct and

papilla.

Histology.

When the pancreas is examined under the low power ( x 180)

it is seen to possess a loose, open texture. The glandular alveoli

are either isolated or aggregated loosely into groups, and all are

supported by delicate connective tissue. The ducts may be in-

visible when the cells are loaded with granules, but they appear

as clear circular central areas when the cells are not so loaded.

Multinucleated areas of different kinds are seen.

Under the high power (480) the gland-cells ave seen to

contain granules of different sizes and different degrees of conrse-

ness, and the bases of the cells are the most granular parts. ‘The

nuclei are circular, large, and present well-marked granules and

networks; in some the granules predominate, but in others the

reticulum is the chief feature.

The multinucleated areas differ greatly in appearance, but can

be arranged in three groups :—

1. Many nuclei and little surrounding protoplasm.

2. Many nuclei with much protoplasm which stains faintly.

3. Many nuclei with much protoplasm which stains deeply.

In all, the nuclei are of varying shapes and sizes.

The connective tissue may appear as delicate strands round the

alveoli, or it may form a strong network within the meshes of

which the alveoli are contained. The meshwork is also cellular,

and the nuclei are circular or long and narrow.

Tur Liver.

1 have nothing to add to existing accounts of the macroscopic

appearances.

Histology (Plate IV. C).

Under the low power it is seen how the cells have the usual

arrangement in columns, but each cell has golden-yellow pigment

granules in the centre. There appears, therefore, to be a yellow

axis running along the centre of the column, for all the cells are

filled from end to end with the pigment.

The high power reveals how a few cells are destitute of pig-

ment, so pigmented and non-pigmented cells are to be described.

The weprigmented cells, which are in the minority, are cubical,

hexagonal, or pentagonal in shape. Their protoplasm is faintly

erantar, and their nuclei are large, round, and granular. The

pigmented cells cannot have their limits so easily defined, for the

pigment granules are tightly packed and obscure their contiguous

walls. The granules are coarse or fine and form a central axis,

but they never obscure the large, spherical, granular nucleus of

the cell. The axis is bordered on either side by the eytoplasm of

the cell which is finely granular, but the granules do not consist

168 DR. GC. F. SONNTAG ON THE

of pigment. These granules may, however, have a pathological

significance.

THE SPLEEN.

The spleen, as Rapp pointed out (10), lies on the right side of

the pylorus. It is, in my specimen, separated from the latter by

the splenic limb of the pancreas, with which it is surrounded by

the layers of the dorsal sheet of the great omentum (text-

fig. 11, B). Its total length is 7-7 em., width of the oval part

1°3 em., and thickness :-75 cm.

Jt consists of an oval posterior part, and a long narrow anterior

part which is twice bent on itself, but the latter has a small thick

concealed process. On the inner side there is a small circular

accessory spleen lying within the splenic limb of the pancreas.

The right-sided position of both pancreas and spleen introduce

modifications in the peritoneum. In my specimen the conditions

induced by the former make the peritoneum different even to

that of forms of B. tridactylus in which the spleen and pancreas

lie as described by other authors.

The position of the spleen offers a problem for the embryologist.

THe PERITONEUM.

The Great Omentum (Plate II. A, F).

The great omentum is attached to the right side of the ventral

limb of the pylorus, but nowhere is it fixed to the greater cur-

vature of the stomach. It has dorsal and ventral sheets, each

composed of two fused layers.

The ventral sheet (text-fig. 12, A, vs) is attached to the right

border of the whole of the ventral limb of the pylorus, and the

right border of the first part of the duodenum. It separates

into two layers which surround these structures. The ventral

layer is continued from the pylorus on to the ventral surface of

the cardiac stomach, and the dorsal layer passes from the ventral

to the dorsal part of the pylorus, thereby forming the left wall of

the lesser sac. On the dorsal limb of the pylorus it meets with

the dorsal sheet of the great omentum which has returned from

the pancreas and spleen.

At the convexity of the pylorus the ventral and dorsal sheets

meet, thereby closing the lesser sac posteriorly, and the ventral

sheet fuses anteriorly with the mesoduodenum (dotted line in

text-fig. 14, A).

The dorsal sheet (text-fig. 12, A, ds) passes dorsally and to the

right from the free edge of the omentum and, at the ventral

border of the pancreas, it splits into right and left layers. The

former passes round on to the right surface of the splenic limb of

the pancreas, and is carried off round the spleen which it com-

pletely encloses; returning to the pancreas again it covers the

right surface. The left layer covers the left surface of the

pancreas and, at the dorsal border, the two layers come together

ANATOMY OF THE THREE-TOED SLOTH. 169

again and run to the dorsal limb of the pylorus. One layer

passes thence on to the dorsal surface of the cardiac stomach,

and the other unites with the dorsal layer of the ventral sheet.

Between the serous covering of the cardiac stomach and the

layers passing between the two limbs of the pylorus there is a

peritoneal pocket (text-fig. 12, A, P).

When the dorsal sheet is traced to the pyloric curvature it

separates into anterior and posterior layers. he former becomes

continuous with the part of the ventral sheet running between.

the two pyloric limbs (text-fig. 12, C, p), and the latter passes

Text-figure 12.

LESSER

SAC

A: Section across the great “omentum. Dorsal (ds) and ventral (vs) sheets

embracing spleen (Sp.), pancreas (Pa.), and ventral and dorsal parts of pylorus

(Pv. & Pa.).

B: Section across the cecal appendage of the stomach.

curvature. The layers pass to the peritoneal pocket (p) and the cardiac

stomach (cs).

round the bend and is reflected on to the right surface of the

cardiac stomach (text-fig. 12, C, cs).

The posterior pole of the spleen is connected by a fold

of peritoneum to the ventral surface of the cardiac stomach,

encircling the ventral pyloric limb and curvature. It shuts ina

peritoneal pocket containing the ventral limb.

170 Dk. ©. F. SONNTAG ON THE

The meso-appendix, which binds the cecal appendage to the

posterior surface of the cardiac stomach, is formed by peritoneum

derived from the coverings of both ventral and dorsal surfaces of

the stomach. Its attachment to the cardiac stomach is much

shorter than to the anterior border of the appendage.

The Gastro-Hepatic Omentum.

The gastro-hepatic omentum runs from the cesophagus, convex

lesser curvature of the stomach, and duodenum as far distally as

the entrance of the common bile-duct (Plate II. A) to the

diaphragm at the left side of the liver, and along the left half of

the posterior surface of the liver. It is wide and fan-shaped

with the base lying posteriorly and the narrower end fixed to the

diaphragm and liver. It fuses with the common mesentery, and

the structures forming the ventral boundary of the wide Foramen

of Winslow are common to both of them (Plate II. A, 1).

When the serous coats of the stomach are traced to the right

they are seen to separate and surround the first part of the

duedenum and become continuous with the common mesentery

(Plate II. A, a). This connection, therefore, forms an additional

means whereby the gut is fixed to liver and diaphragm.

The Lesser Sac of the Peritonewm (Plate Il. A; text-

fig. 12, A).

The long axis of the lesser sac describes an S-shaped course

from the Foramen of Winslow to the free edge of the great

omentum, and its generai direction is posterior and to the right.

It is also tapering, for the Foramen of Winslow and part abutting

against the liver are much wider than the omental end. It also

hes mostly to the right of the stomach, and part of it is anterior.

Consequently, the disposition is different from that of most

Mammals, where it lies dorsal to the stomach. ‘This curious

arrangement is due chiefly to the position of the pancreas, spleen,

and great omentum, and the relations of the layers of the omen-

tum to the limbs of the pylorus.

The boundaries of the lesser sac from behind forwards are :—

To the right—pancreas and spleen enveloped by the layers of

the dorsal sheet of the great omentum.

To the left—the two limbs of the pylorus and the beginning

of the duodenum.

Porsally—the so-called ligamentum hepato-cave-duodenale of

Klaatsch, which is a continuation of the right half of the

suspensory sheet of the liver and covers the vena cava

inferior.

Ventrally—the great omentum, pylorus, first part of the

duodenum, and the fused sheet produced by the lesser

omentum and common mesentery.

The above conditions resemble in many respects those of some

ANATOMY OF THE 'THREE-TOED SLOTH. 171

of the Reptilia, and Klaatsch (6) has deseribed the peritoneum

exhaustively in them.

The Foramen of Winslow is also very wide as in the case of

Reptiles. It has the usual boundaries.

In Plate [1. A the ventral wall of the lesser sac has been

removed round the letters MGIJK L and one sees the pancreas (P)

in the dorsal wall of the sac.

Text-figure 13.

LIVER.

a! Br

(Cr

7, ‘

aN Gem

(ANTERIOR SURFACE) aly wae

dy WN ZS

: “> wiih a= Bl"

8 ANS ,

“SKE

SED. Cc

ERI VE:

(POSTERIOR SURFACE)

The suspensory ligaments of the liver and the manner in which their continuations

enter into the formation of the anterior part of the common mesentery. Two

halves, B/ and B” of A, pass round the edges of the liver at the points +

and X. They eventually become the mesentery D of figs. Cand D. A and a,

inferior vena cava. ec: lesser omentum. E: cesophagus.

Ligaments of the Liver.

When the liver is depressed so as to show the anterior surface

(text-fig. 13, A), one can see how it is connected to the diaphragm

by a V-shaped sheet of peritoneum whose apex is dorsal. The

1} DR. C. F. SONNTAG ON THE

limbs of the sheet play important parts in the formation of

the mesentery and the ligamentum hepato-cavo-duodenale of

Klaatsch.

The right limb is inserted into the floor of a groove. It passes

round the right border of the liver and separates into two layers

which pass along the sides of the vena cava inferior. One-half

covers the right abdominal wall, and the other covers the dorsal

abdominal wall and can be traced into the common mesentery

(text-fig. 13, Cd and Dd).

In the angle between the two limbs the vena cava inferior,

arched over by peritoneum, can be seen (text-fig. 13, A, @) running

to the diaphragm.

Text-figure 14.

DUODENUM, - -

YEJUNUM

&/LEUM,

CAECUM,

The common mesentery viewed from the right (A) and left (B). Lig. Sus., suspen-

sory ligament; C.B.p., common bile-duct ; Pa., pancreas : Sp., spleen; Py.,

the dorsal and ventral limbs ind flexure of the pylorus; the unmarked dotted

line is the line of fusion of the great omentum and mesentery.

The left limb of the sheet is the true suspensory ligament. It

passes into the fissure between the left and right lobes of the

liver. It turns round the left hepatic margin, and is attached to

the posterior surface from the left border to the origin of the

common bile-duct. The lesser omentum fuses with it. Itis con-

tinued as the common mesentery, and is combined therein with

part of the ligamentum hepato-cavo-duodenale. Between the

ANATOMY OF THE THREE-TOED SLOTH. 173

two halves is the anterior part of the lesser sac (text-fig. 13, B);

which opens into the Spigelian Recess of the liver.

The Mesentery.

The mesentery has an attachment from liver to pelvic floor

which begins anteriorly to the right of the mid-line, but passes

posteriorly and to the left to reach it. It is composed at first of

the suspensory ligament, and its free edge is attached first to the

diaphragm, then along the posterior surface of the liver

(Plate IT. A,n). Its free edge then, containing the common bile-

duct, runs from the portal fissure to the duodenum. Finally, it

follows the intestine to the pelvic floor. It has attached to it

the layers of the great omentum. It sends a tubular sheath

round the oesophagus (text-fig. 13, B, #).

The stomach is attached to it anteriorly by the lesser omentum,

and to the right by the connection round the duodenum (text-

fig. 14, B).

No peritoneal bands connect the duodenum to the colon.

Dr. Chalmers Mitchell (8) and others have described the peri-

toneum from duodenum to anus, and the only fact which I have

to add to their accounts is the prominent ridge produced in the

mesentery by the posterior mesenteric vessels.

The Suprarenal Capsules.

The left capsule is flat and ovoid and measures :—

Hkeiiothy, cweerncuar canned: 1°8 cm.

Witla oi aelaraeh tae eas me a5) ke

BM CMSs Welsh ce nace ee ga O85)

The right capsule, as is shown by the following figures, is

longer, narrower, and thicker :—

Reni othis yak Saeed gee vee 2 cm.

Vall law Mi ene hi i Ech aud 56) °f

ADC OSS nee A Mts a) D

Both lie a considerable distance anterior to their corresponding

kidneys. On section the right capsule shows a round cortex and

medulla, but these are long in the case of the right one (text-

fig. 11, E).

The Thoracic Organs.

Burne (1) and Rapp (10) have described the heart and large

vessels, but the branches of the aortie arch in my specimen are

different to their accounts. It gives off from right to left the

innominate, left subclavian, and three intercostal arteries (text-

fig. 15, B).

The posterior border of the arch is connected to the left

pulmonary artery by a very prominent ligamentum arteriosum ;

this measures 1 em. long and 2 mm, wide, but is quite

impervious,

174 DR. C. F. SONNTAG ON THE

The left vagus and phrenic nerves are fused in the anterior

part of the thorax, there being only lee furrows to mark

the distinction between them (text- -fig. 15, A).

‘The lungs have large alveoli, as navn | oy John Hunter (12).

Text-figure 15.

PA.

A: The adhesion of the left vagus (v) and phrenic (P) nerves.

B: Theaortic arch (A.4.) giving off innominate (rA.), left subclavian (1.8.4.) and three

intercostal (1 4, 1, 2, 3) arteries; LA., heamentum arteriosum; PA. and LPA.,,

pulmonary and left pulmonary arteries.

Tuer GENERATIVE ORGANS.

The internal generative organs of the female have been

described by John Hunter (12), Klinckowstrom (6), and

Rapp (10). Those of my specimen were not examined, for

the gravid uterus was at once removed for embryological research.

The external genitalia and anus are all enclosed in a species of

cloaca, and the Clitori is is a small semicircular flap. ‘The male

organs also exhibit a primitive condition, and John Hunter (12)

ae described them as follows :—‘ The penis is a short flat body

enclosed in a prepuce which is within the verge of the anus. 1

is not above two tenths of an inch in length, and terminates in

an obtuse point. It has a groove which runs along the under

surface, and which makes the point somewhat for ed.”

No os penis is present.

Ballowitz (15) has deseribed the spermatozoa.

Tur Petyic Bones.

The skeleton of the specimen, described above was recently

obtained by me from Mv. Gerard, and the pelvis was compared

with that of a male animal ine was preserved in the Society’s

Prosectorium. There are numerous and striking differences be-

tween them; these are seen in the measurements of the different

ANATOMY OF THE THREE-TOED SLOTII. 11 7/5)

diameters, the shape of cross-sections of some of the bones, the

sizes of openings, and the development of crests and eminences.

Speaking generally the female pelvis is rounder and more capacious,

for it has to accommodate the large trumpet-shaped rectum, and

leave a passage for the feetus during parturition.

For purposes of measurement I have chosen the following

diameters which are made use of in human obstetric anatomy,

but have modified them slightly :—

1. Conjugate—irom the centre of the inner border of the pubis

to the front of the centrum of the first sacral vertebra.

2. T'ransverse—between the widest points of the lateral pelvic

walls: that is between the inner surfaces of the acetabula.

3. Oblique-—from the sacro-iliac joint to the mid point of the

bone between the mid line in front and the anterior border of

the obturater foramen.

4, Interspinous—between the iliac spines.

5. The measurements of the outlet are taken between the

widest points dorso-ventrally and transversely.

The proportions present in male and female pelves are as

follows :—

Diameters. Male pelvis. Female pelvis.

Interspinous diameter ............ 9-8 em. Oa emer

Conyugatevonmletiy ey neta... Clee: oe

MRAM SVIETSO a0 4 sort sesilek wersise ots Sel SD Ne Oo

Oblique Me Rate el eeve tien cane abana 7 bi (Bas

Conpugaberol Outlet) moeane es. oss DRO Sp Dilan

Dye sar 5 5-

PATS VELSO rs | Gh oMeteoehisn ules seine i) 5 Drea

\iVaWaliglau Ost FOWUOHIS jeeeesoecedonas soar 2 Me Deore

Tschiadic Poramina.

Antero-posterior diameter ......... AP) let 1ESue.

Transverse SERIO 2. S98 12Gt eS.

Obturator Foramina.

Antero-posterior diameter ......... Gee ey

‘Transverse A a Be ae aS EL: 2, 2-5

The following differences are also present between a male and

female pelvis :—

Male pelvis. Female pelvis.

Isehial tuberosities well marked. Ischial tuberosities very small.

Pubis circular on section. | Pubis flat on section. ~

Jliac spines incurved., Iliac spines not incurved,

Tnlet less defined. Inlet well defined by a sharp

| -erest.

Ischia more sloping from last:

sacral vertebra. |

Transverse processes of caudal Transverse process of caudal

vertebrie sloping. vertebrae horizontal,

176 DR. C. F. SONNTAG ON THE

THE SACRAL AND CAUDAL VERTEBRA.

Owen mentions in his ‘ Comparative Anatomy of the Verte-

brata,’ that there are six sacral and eleven caudal vertebrae. In

both my specimens there are seven sacral and nine caudal

vertebre, but there is a greater degree of fusion, and less

distinctness, between the sacral centra in the female pelvis.

Flower gives the number of caudal vertebra as 6-10.

THE Broop.

Gulliver (16) has pointed out that the red blood corpuscles are

very large, and gives their measurement as 1/2778th inch in

diameter in a young 2B. didactylus. He also states that the

Elephant is the only mammal with larger corpuscles.

SUMMARY.

The new facts recorded in this communication are >—

a. The full deseription of the mouth and tongue.

6b. The comparison between the palates of Bradypus and

Cholepus.

ce. The external relations of the compartments of the stomach,

and the distribution of lymphatic glands in the stomach-wall.

d. Details of the internal structure of the stomach, and a

complete account of the interior of the cecal appendage.

e. The unusual and reptilian-like situation of the pancreas,

which results in complications in the peritoneum. The latter is

fully deseribed for the first time in bradypus tridactylus.

f. The vessels arising from the aortic arch are different to

previous accounts.

g. The ligamentum arteriosum of Bradypus is described for

the first time.

h. The duct papille im the duodenum are different from the

form which has already been described.

i. The comparison between the male and female pelves.

BIBLIOGRAPHY.

1. Burne, R. H.—Proe. Zool. Soc. 1901, pp. 118-120.

2. Carus.—“ Erliiuterungstafeln.” Viertes Heft, p.19. T. vii.

3. Cuvipr.—‘‘ Legons d’anatomie comparée,” Paris 1805, v. 3,

p-. 388.

4, Francavietia, M.—Boll. Soc. Roman. per gli studi zool., 1894.

5. Kuaarscu, H.—Morphol. Jahrb. 1892, pp. 885-450 & 609-

716.

6. KurcKkowstrom, A.—Zool. Jahrb. Jena, 1895, p. 481.

7. MecKEeL.—Syst. der verg. Anatomie, Halle 1829, v. 4,

pp. 605-817.

8. Mircueny, P. Coaumers.—Trans. Zool. Soe. IID, p. 437.

9. PILLIET. 2G R. Soe. Biol. Paris, (9), v. 3, p. 315; Journ. de

Anat. et de la Phys., vol. xxii. pp. 402-423.

ANATOMY OF TITE THREE-TOED SLOTH. Via

10. Rarr, v.—“‘ Hdentaten,” 2. Aufl., Tiibingen, 1852.

11. Semon.—Arch. Naturg. 1902, p. 259.

12. Hunrer, Joun.— Essays and Observations,’ edited by

Richard Owen, London, 1861, vol. 11. pp. 176-179.

13. Gmevin.—Arch. f. mikr. Anat. 1892, pp. 1-28, Taf. 1.

14. Mayer, F. J. C.—Froriep’s Notizen, 1842, pp. 289-290.

15. Bautowrrz, K.—Anat. Anz. Jena, 1906, pp. 321-324.

16. Guiiver, G.—Proe. Zool. Soc. 1848, p. 37.

EXPLANATION OF THE PLATES.

Prare I.

A. The anterior end of a section through the head. 77, lips.

B. The roof of the mouth: c, cheeks; v, vestibule; sp, soft palate.

C. The bony palate of Bradypus tridactylus.

D. The bony palate of Cholvepus.

Puate II.

The external (A) and internal (B) structure of the stomach. In B the ventral

wall has been thrown upwards and to the right, and the dorsal cardiac compartment

and pylorus separated and thrown to the left.

A,c and B, 1.1.1. : paunch.

A,p and B.d, c, d, 2: cardiac stomach.

A, eu and B 3: pylorus.

Description of other letters in text.

PLATE IIT.

The interior of the dorsal compartment of the cardiac stomach (A), dorsal part of

pylorus (B), and ventral part of pylorus (C).

Aa: rod running through the ruminating gutter to the pylorus. 6, rod

communicating with the paunch (c). d, pleated mucosa.

Ba: epithelial ridges bounding glandular areas (6). ¢, ruminating gutter.

d, hard pleated mucosa continued from the cardiac stomach.

C a: pyloric sphincter with beginning of duodenum (6). ¢, cardiac stomach.

Prats LV.

A: The iuterior of the intestinal tract. A and B, pieces of the small intestines.

cand p: the lower end of the ileum, colon, and rectum.

B: Section through the septum between cardiac stomach and paunch.

©: Section of the liver.

Proc. Zoor. Soc,—1921, No. XII,

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ON DEATHS IN THE GARDENS IN 1920. 179

8. Report on Deaths which occurred in the Zoological

Gardens during 1920. By N.S. Lucas, M.B., F.Z.S.,

Pathologist to the Society.

[Received January 31, 1921: Read February 22, 1921.]

On Jan. 1st, 1920, there were living in the Gardens :—

586 mammals, 1333 birds, 411 reptiles.

(826) (2162) (486)

To these were added during the year :—

450 mammals, 1138 birds, 693 reptiles.

(446) (1356) (683)

Giving a total of :—

1036 mammals, 2471 birds, 1104 reptiles.

(1272) (3518) (1169)

The total of deaths which took place during the year 1920

was 1101. This was made up as follows :—

333 mammals, 490 birds, 262 reptiles.

(356) (857) (467)

This gives a percentage of deaths to animals living in the

Gardens during the year of :—

32°1 19-8 20-1.

(27-9) (24:3) (39:4)

Tf only those animals which had lived in the Gardens for six

months or more are considered (it being assumed that these are

acclimatized), the number of deaths is as follows :—

167 mammals, 230. birds, 50 reptiles.

Giving a percentage of deaths of :—

16:1 9°3 4-5.

(16:9) (16-4) (13:8)

As this is the first year in which the numbers have come up to

pre-war standards, or nearly so, the figures for 1913 are given in

brackets corresponding to each of the above.

Comparing these figures, those for mammals remain very

constant. The increase in 4:2 per cent. of the death-rate’ may

be considered as due to the higher ratio of unacelimatized

mammals. If only the acclimatized are taken, there is a drop of

0-8 per cent.

The total number of birds in the Gardens shows a drop of

approximately 1100. The bird population in 1913 was nearly

50 per cent. greater than now. The total and acclimatized death-

rate has diminished. In the latter case this is very marked. A

reference to the diminished figures for avian tuberculosis and

180 : DR.IN. S. TACAS ON TAFE

mycosis will shed some light on this fall. How much of this

diminution in the incidence of these liseases, 154 dug. tq the, dirds

being less crowded it is difficult to Say, ir ‘the numbers of, birds

rise to pre-war height, it will Ve interésting of fo Watch? the™ Gorre-

sponding effect on these diseases?12!008 ote od dargolod

The decrease in the death-rate among the reptiles i is still more

marked, being nearly half, even when: hewlysnnpotted: ‘individuals

are considered, This is due to the absence of an epidemic of

pneumonia, which in 1913 claimed 138 victims, while’ the total

deaths from lung diseases for 1920 is 61.

If only the more acclimatized specimens are considered, the

death-rate is remarkably low. This is probably due to the fact

that those which had been in the Gardens for more than six

months were mostly reptiles which had survived the difficult

period of the war and were rather specially well acclimatized.

Notes on the Analysis of the Causes of Death.

A. Pleurisy.—It.is interestmg tomote the varity of: this disease

among animals in comparison with its common.incidence among

humans. san BBR ; ue

One case occurred. dn a ereen ‘eahbppitheyue, the other in a

macaque.

9. This vecurtéd in a erey Widn sods MGI A

3. This was found in an Indian ce dow and WAS due ton’

poe Spiroptert' Sanguinolentat wre] [es

This occurred ina Burnett's! cereopithegue. (Bye)

: It_is difficn lt to account for the, diminution 1 in the, deaths

from nephritis. : “The figuy es this year. ave 8, 31 sin 0 for the

three classes, while’ in. al GTS. “they wee | 00, Too, and’6 TAL. fist

sight it might seem that this wits “due to a differ ence in opinion

as to the cause of death, but a8 tWere’has beén ns" ‘ediresponding

vise in any other disenee or diseases,-this, cannot, be; the,cause.

So little is known as to the cause of nephritis, that it is impossible

to assign a reason.

6. In a budgerigar.

7. Enteritis still remains the chief cause of death among the

birds... One organism, that of B. aertryck, has been isolated: from

a newly-imported grey touracou and a newly-imported parrakeet.

8. These occurred in a ring- -tailed lemur, a bad ger, and

bonnet macaque.

9. in a black-backed jackal.

10. Ina bonnet macaque. This was an old individual.

11. In a fairy blue-bird.

12. In a violet tanager.

13. The high death-vate from tuberculosis among the mammals

as compared with that of birds is of interest.

Among the mammals 69 deaths occurred, of which 43 were

those of macaques in the canal-bank aviary. Of the remaining

26, five occurred in the Lion House, leaving 21 for the other

OV ii) 2

DEATHS IN THE GARDENS IN 1920.

Analysis of Causes of Death.

181

| Diseases

Diseases of Respi-

ratory Organs. :

Diseases of

Cardiovascular

System.

| Diseases of

Genito-urinary

System.

of ali-

mentary tract.

| Various

Diseases of Liver...

Diseases of

Panereas.

Diseases of

Nervous System. §

Diseases due to

Infection.

(2)

(3)

(4)

(5)

(6)

(7)

Disease.

Lobar pneumonia .........

Lroncho-pueumonia, ¢,,...)

~ Congestion, of lungs

Bronchitis J aeeepnatte

(fidema of lungs 5..0.......

Pleurisy cat

Pleurisy with effusion.

Abscess of lung ............

Paragonomiasis of lune: i

Collapse of lung (xickets)

Granuloma of lung........-

Pleurisy and pericarditis

JPESPIGHNCUINS cas gesuacbes doo oor

Hndocarditis sue

Ruptured aneurism ......

Ruptured heart’ .2.2...22).

Rup uved artery: J... <4.

Anemia :

Anemia of spleno-n me- se)

dulliry type..

INJ@OVAKUIS: ~ Garcoososccq. 50008

@ystre kidney <..c.. 200...

Necrosis of kidney .........

Kee-binding........

Inflammation of oviduct

Carcinoma of testicle ......

Stomatitis Ravacwaceeen

(Msophageal abscess ......

Gastiitisnay eterna

Kuteritis

(8) Intussusception ............

(9)

Intestinal obstruction

? old intussusception

(10) Volvulus .........

Granuloma of intestines

(11) Ulceration of cloaca ......

(12) Fibroma of cloaca

Wane

(18)

JEPAPOUDUUIS. | Gon gdoeadcasnonces

Granuloma of mesentery

Fatty degeneration of liver

Cirvhosislof liver. .05....

Portal pyeemia Gareunsss

Kuptured liver...............

JEEWOOREEN HRHUS oro sen eah seu seeane

Cdema of brain ...........

(13.a) ; (Mubencwlosisieeesss sae

IMINO sedach ce aatsuctesencoes

Anthrax RO hee:

INE GENES, <5 con ecoccoson econ

el minnie liasiss see eee eee

Sarcoma SALARY

Abscess of ribs..............,

Ascites ? cause...............

Debility

Tnanition ..

Senility.. Buanuceens

Under anesthetic

Exposure ......

Killed by accident. or by

order.. SEPARA DOGHER

Eaten by atic cee ee

Decomposed .. Se asc:

N ot diagnosed...............

Not exannineds secre sscess

Mammals.

ee wee: e Co eS ee ee Re Re ee:

Reo:

bo Or bo

is) o

Or:

co -T TWO DO

Birds.

bo:

mee:

me poh:

| Reptiles, |

etc.

fet

182 ON DEATHS IN THE GARDENS IN 1920.

houses and enclosures. In 1913 there were 31 deaths of this

disease, of which 15 came from an epidemic in what is now

called the Rodent House, so that, deducting the epidemics, there

is a slight rise in the death-rate from tubercle this year as com- -

pared with 1913.

Of the 26 cases occurring this year, seven only had been in the

Gardens less than six months, and of two the date of arrival

was accidentally omitted. Of the remaining 17, eleven had been

in the Gardens over a year anda half. One, a blesbok, had

been there nine years.

It is therefore difficult on these figures to give any opinion as

to the method of the introduction of the disease to the Gardens.

The small death-rate among the birds may have the same

origin as that of mycosis. ;

13a. Among the cases of mammalian tuberculosis was that of

an axis deer with disease of the genital organs, which is extremely

rave in the lower mainmals. The animal had been in the Gardens

for many years.

SECRETARY ON ADDITIONS TO THE MENAGERIE. 183

EXHIBITIONS AND NOTICES.

February 8th, 1921.

Prof. KE. W. MacBrips, D.Sc., F.R.S., F.Z.S., Vice-President,

in the Chair.

The Secrerary read the following: Report on the Additions

to the Society’s Menagerie during the months of November and

December 1920.

NOVEMBER.

The registered additions to the Society’s Menagerie during the

month of November were 102 in number. Of these 48 were

acquired by presentation, 25 were deposited, 21 were purchased,

6 were received in exchange, and 2 were born in the Menagerie.

The following may be specially mentioned :—

1 Common Otter (Lutra lutra), from Lincolnshire, purchased

on November 16th.

1 Japanese Deer (Sika sika), bred in England, presented by

Ernest J. Gripper on November 4th.

3 Alpine Marmots (Marmota marmotia), from St. Moritz,

presented by the Curverein, St. Moritz, on November 29th.

DECEMBER.

The registered additions to the Society’s Menagerie-during the

month of December were 66 in number. Of these 33 were

acquired by presentation, 25 were deposited, and 8 were

purchased.

The following may be specially mentioned :—

2 Beech-Martens (Marites foina), from Denmark, purchased on

December 7th.

1 Timber-Wolf (Canzs occidentalis), from N. America, presented

by Capt. R. A. Angie on December 3rd.

Mr. E.G. Boutencer, F.Z.S., exhibited, and made remarks

upon, a Black Salamander (Salamandra atra), which, in the course

of its captivity in the Gardens, had developed yellow spots on

the body.

Prof. J. P. Htut, F.R.S., exhibited, and made remarks upon,

a series of lantern-slides of the Foetus of a Three-toed Sloth

(Bradypus tridactylus).

184 PROF. G. BLLIOLT SMITH ON EXAMPLES

. Pebruary 22nd, 1921...»

Sir S. F. Harmer; K.B,E.,.F.R.S,,:Vice President,

in the Chair.

The Seoneiane oo abe Hila Report on Atte ‘Additiol’ to

the Society’s Menagerie during the month of January 1921:

2 Vhe régistered additions to. the: Society's. Menagerie: during

the month of January were 122,in number. Of these 49 were

acquired by presentation, 64 were deposited, and 9 were

purchased.

The following may be specially mentioned :—

1 American Bison (Bison americanus), 3, bred at Woburn,

deposited by The Duke of Bedford, K.G., on January 28th.

1 Dromedary (Camelus iPaper OL d, from Mogador,

Morocco, purchased January 31st.

1 Verreaux’s Amethyst- Starling (Philodauges leucoyaster ver-

reauxr), from Durban, South Africa, new to the Collection,

presented by Harold Millar on January 15th.

1 Sundevall’s Seed-eater (Serinus scotops), from South Aifriba,

new to the Collection, purchased on January 21st.

Prof. G. Exxior Suiru, F.R:S., F.Z.S., exhibited a series of

photographs of a hving example of Deer sins, and. said :—

The accompanying photographs of a living Zarsius, taken by a

Chinese boy at Kuching, Sarawak, have inst been received from

Mr. W. E. Le Gros Gis who went out to Borneo last autumn

to take up the position of jantical officer at Kuching. Dr. Charles

Hose tells me that the usual and characteristic attitude of Tarsus

is that shown in these pictures, but during his long period of

service in Sarawak he was unable to get any satisfactor y photo-

graphs of this singularly elusive creature. Hence we owe

Mr. Le Gros Clark our deepest gratitude for making us familiar

with the somewhat unexpected appearance of this surviving

member of the Eocene family from which our own Simian an-

cestors were derived.

Before he left for Borneo last autumn, I asked Mr. Le Gros

Clark to collect all the information he could get of the hahits of

Tarsius and Tupaia, to secure photographs of the living animals,

and obtain the material for anatomical and embryological research

on these important genera. Moreover, I specially impressed

upon him the impor Ties of studying the retina of living or

freshly-killed examples of Varsius for the purpose of determining

whether or not a macula lutea was present.

Immediately upon his arrival at Kuching, Mr. Le Gros Clark

was able to secure the two photographs reproduced here, and also

the fresh corpse of an adult female (? pregnant), which he has

IP Zo So IOZil, ISLEIOW SMI Nal, Ile Me

A LIVING EXAMPLE OF TARSIUS.

OF TARSIUS FROM SARAWAK. 185

dissected. The uterus has been preserved for Professor J. P.

Hill to examine. Mr. Le Gros Clark gives me the important

information that neither with the ophthalmoscope nor by direct

examination of the retina after opening the eye was he able to

detect any sign of a yellow spot (macula lutea). At the same

time he cautiously refrained from denying its presence. Dr. H.

Woollard has been investigating the specimens of Varsius given

to me many years ago by Dr. Charles Hose for the purpose of

preparing a monograph on the anatomy of Jarsiws. He was

unable to find the macula lutea in his preparations of the retina,

nor was he able to discover any trace of it in a series of excellent

histological preparations lent me by Dr. W. L. H. Duckworth of

Cambridge.

The macula lutea is a very obtrusive feature of the retina in

monkeys, and can easily be seen even in badly preserved material.

Hence it could hardly have been overlooked in the fresh retina

of Tarsius, if it had been present. Hence Mr. Le Gros Clark’s

observations, taken in conjunction with Dr. Woollard’s work at

University College, make it practically certain that in Tarsius

this distinctive feature of the retina of the Anthropoidea (among

mammals) is lacking. In this respect Tarsiws agrees with the

Lemurs, and differs from all monkeys.

In the Anthropoidea the development of the macula lutea is

associated with a distinctive arrangement of the fibres of the

optic tract. Instead of a complete decussation of the optic

nerves there is only a semi-decussation, the fibres coming from

the lateral half of each retina remaining uncrossed. Dr. Woollard

tells me that he is unable to find any evidence of such an arrange-

ment in Varsiws, which, like the Lemurs, seems to have either a

complete or almost complete decussation.

These observations are of far reaching significance from the

light they shed on the cardinal factors that brought the

Anthropoidea into existence.

I have repeatedly called attention to the influence of the

adoption of an arboreal life upon the cerebral cortex and the

behaviour of mammals, which is so clearly displayed in the

Menotyphlous Insectivora by comparing Macroscelides with

Tupaia. Inthe Tree-Shrew the importance of the sense of smell

is diminished, and those of vision, touch, and hearing greatly

enhanced. The further emphasizing of these adaptations brought

the Primates into existence. Among the Prosimiz the reduction

of the size of the nose in one group, the Tarsioidea, allows the

eyes to come to the front of the face so that their fields of vision

overlap. The enormous enhancement of the importance of vision

which is thus effected leads to the sudden expansion of the

cortical area for vision and its further specialisation in structure.

The sense of sight now completely supplants that of smell as the

dominant guide to the animal, and the alteration of the position

of the eyes enables the animal to look forward, both in the litera]

and the metaphorical senses of the expression.

Proc. Zoou. Sec.—1921, No. XIII. 13

186 EXAMPLES OF TARSIUS FROM SARAWAK.

But though Varsius has binocular vision, 1t has no macula lutea

to enable it to appreciate the details of the objects seen, nor has

it any automatic mechanism for producing the conjugate move-

ments of the eyes necessary for bringing the two images of objects

seen on to the corresponding areas of the two retin. In other

words, Varsius has not yet acquired true stereoscopic vision.

But the wide range of movement of the head on the neck shows

that the co-ordination of the two eyes is becoming biologically

useful to the animal; but as it has no mechanism for automatically

regulating the positions of the eyes the one to the other, it moves

its head as the cat does.

These enhanced powers of observation opened the possibility

for one branch of the Eocene Tarsioidea to guide its hands with

greater precision for the performance of skilled movements, and

so incidentally enhanced the sense of touch. Hence both the

tactile and the motor areas in the cortex underwent a great

expansion and elaboration; and at the same time the prefrontal

cortex began to grow rapidly as a mechanism was built up for

co-ordinating the movements of the eyes. When this happened

specially sensitive areas (macule lute) were differentiated in

the two, retinz, and the optic tracts became rearranged for the

purpose ci stereoscopic vision. These far-reaching changes led

to the transformation of the brain and converted a Tarsioid into

a monkey.

PROCEEDINGS

ee THE

| =% Nek

GENERAL MEETINGS FOR SCIENTIFIC Vefenamiua’> ~)

OF THE

| ZOOLOGICAL SOCIETY

OF LONDON.

1921.

PART i,

CONTAINING Pages 187 ro 446, with 10 PLaTEs AND

83 TEXxtT-FIGURES. :

JUNE 1921.

PRINTED FOR THE SOCIETY,

SOLD AT ITS HOUSH IN REGENT’S PARK,

LONDON:

l MESSRS. LONGMANS, GREEN, AND CO,

PATERNOSTER ROW.

Ore [Price Twelve Shillings.] SE)

a Ee ETT STR a On a ee 4 .

LIST QF CONTENTS.

1921, Parr II. (pp. 187-446).

EXHIBITIONS AND NOTICES. =

W. Suvurevpr, C.M.Z.8. Photographs of the “last ae the Passenger Pigeons

“pistes migratorius). (Text-figure 1.) 2.1... cece cree cece cece teense ees oe

». BR. I. Pococr, F.R.S., F.Z.S. Exhibition of, and remarks upon, the skull of a —

Shi okae IE orG comudeaGunoes To OnaG oo do anon KgosOabOUmbadoEe ac coce soc’

Messrs. Gerrarp & Sons. Exhibition of a Cheetah skin from Tanganyika Territory ..

The Sucretary. Report on Additions to the Society’s Menagerie during ume Bat of

Hebruary sl OO eects ssi eid a welaiats nin Sots wi te 1c ee eee ola sue ope aces Ae rs en pa See ce

Dr. P. Cuatmurs Mircnext, 0.B.E., F.R.S. Exhibition of, and remarks upon, lantern-

slides of a baby Chimpanzee born in the New York Zoological Park ..... Boucbotcs

Sir 8. F. Harmer, K.B.E., F.R.S. Exhibition of, and remarks upon, a photograph of

Elephant Twins ...... Sin ois \ciataisl Mejetele isis: peverave okescssiatsr isbeyeetelntevnr a citerens saan ecto meeenee

Mr. E. G. Bovtenerr, F.Z.S. Exhibition of, and remarks upon, some Reptiles and

BaveACMiAnen ac tc eictishettee sie Wigtel Mist Aee Gee ae ee Be EOS COS ROOTS seta aunieoenees

Miss L. E. Cunusman, F.1.S, Exhibition of, and remarks upon, a nest of Anapha venata

(epic op teres) eiscen cco c-e arcrcrnacte ole alee seaTauel steele) cetera tele or eeape ete efer et aRen sce eater rete oe

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

Marla VO 2M oie oie cecsctisl el sper les stab epeheve) fe chcrhens oitecGis ar shasehen casi ier tice hontai ae rae fod

Mr. R. H. Burne, M.A., F.Z.S. Exhibition of, and remarks upon, specimens of NS

JQEVBal Mies Come da Am orinOS docu MEA nC ER Hohe soOSeUO COCK be Saou Haag oO oc

Dr. P. Cuba Mircuett, C.B.H., F.R.S. Letter from Mr. E. H. Bean, Director of

the Washington Park, Milwaukee, Wis., U.S.A., describing the successful rearing of

Paw OlaIsD CAI OUD Gace acarare ere ieles inlets, Nato eke ciciayers oe Are ererese ee ayer eee cee

Mr. ©. Davizs Suerzorn, ¥.Z.S. Exhibition of, and remarks upon, a coin of the Saka

Dynasty (Punjab Region), showing the so-called ‘‘ maneless lion” of Asia ....se+s

Mr. F. Martin Duncan, F.Z.S.. Exhibition of, and remarks upon, photographs of a nest

of the Wiasp (Vespa: geringmtai ts) rercicles says ieee eae ana ieiaiaiets wislaraileisTiavefaoie

442

443

443,

443

444

445

Mr. D. Setn-Smirn, F.Z.S. Exhibition of, and remarks upon, a series of skins of the.

ZT Nae Budgerigar (Melopsittacus undulatus) 0+....eeeseeee Nate seitelere lars eacens

Major HE. E. Austmy, D.S8.0., F.Z.8.. Remarks on an app change of habitat of the

Common Cricket (Gryllus AOMESLIGUS). raiaicie wale vispesarsnetayticieie RIE

445

Contents continued on page 3.0f Wrapper.

THE REPRODUCTIVE ORGANS OF KUKENTHALIA BOREALIS. 187

9. On the Reproductive Organs of the Ascidian Atikenthalia

borealis (Gottschaldt). By Dr. Avausra ARNBACK-

CHRISTIE-LINDE *.

(Text-figures 1-8.)

"Received October 7, 1920: Read March 8, 1921.]

The subfamily of the Polyzoine, to which Kikenthalia borealis

(Gottschaldt) has been referred, is regarded as an intermediate

group between the Styelide and the Botryllide. ~ According to

Michaelsen 7, who has given descriptions of this group of Ascidians

in several papers, it comprises about ten genera, distributed in

almost every region all over the world.

In the Arctic region it is represented by a single genus with

one species, Kiikenthalia borealis.

The species was first described by Gottschaldt + under the name

of Goodsiria borealis. It was re-described by Hartmeyer§ in

‘Fauna Arctica’ under the new generie name Kiikenthalia.

As it thus appears, the species has been the object of thorough

examination by several eminent zoologists. However, it has not

hitherto been possible, so far as | know, to make out its internal

structure, owing to the unsatisfactory knowledge of the repro-

ductive organs.

Gottschaldt (7. c.) merely writes as follows: ‘‘ Die Geschlechts-

organe liegen in der Tunica (es wurde ein einziges ziemlich grosses

Ei gefunden).”

Two years later Bonnevie || described a few colonies of the

species which had been dredged by the Norwegian Atlantic

Expedition, 1876-1878. They were described under the name of

Goodsiria coccinea (Cunningham). With regard to the repro-

ductive organs, the author states that they ‘are found in small

polycarps in the muscular stratum,” a statement which proves

to be a mistake.

Hartmeyer writes (/. c. p. 260) :—‘“ Weiblicher Geschlechts-

apparat diffus (grdssere und kleinere Hizellen tiberall in den

Innenkorper eingestreut sowie in den Blutbahnen der Darm-

wandung und der jungen Knospen) ; minnlicher Geschlechts-

apparat “pisher nicht nachgewiesen.” ‘* Kin besonderes Ovarium

ist nach unseren Untersuchungen nicht vorhanden” (J. c. p. 258).

In the paper of Michaelsen we find the same statement.

* Communicated by Dr. F. A. Barner, F.R.S., F.Z.S.

+ Michaelsen, W., “Revision der compositen Styeliden oder Polyzoinen.”’

Mitteilungen aus d. Naturhist. Museum in Hamburg, Jahrgang 21. Hamburg,

1904.

t Gottschaldt, R., “ Die Synascidien der Bremer Expedition nach Spitzbergen im

Jahre 1889.” Jen. 7%. vol. xxviii. Jena, 1894.

§ Hartmeyer, R., ‘Die Ascidien der Arctis.” Fauna Arctica, Bd. iii. Jena,

1903-1904.

|| Bonnevie, K., “ Ascidie simplices and Ascidie composite. From the North

Atlantic Expedition.” The Norwegian North Atlantic Expedition, 1876-1878.

Christiania, 1896.

Proc. Zoou, Soc,— 1921, No. XLV. 14

188 DR. A. ARNBACK-CHRISTIE-LINDE ON THE

According to this author, male gonads, as well as female ones, are

unknown. And he is of opinion that, with regard to the repro-

ductive organs, the genus in question differs from all other

Polyzoine, and even from all other Styelids.

Nor did Bjerkan * succeed in enlightening us on the matter,

though, as he says, he had a great mass of colonies for examination.

Being occupied with studies on the Northern and Arctic

Ascidians belonging to the collections of the Swedish State

Museum, I have had opportunities of examining Avikenthalia

borealis.

My investigations of the species are not yet completed. How-

ever, as important facts concerning the reproductive organs have

been established, I publish the results already arrived at, hoping

that they will augment our knowledge of an organic system of

which one has hitherto tried in vain to get a clear conception.

The material examined was collected off Spitzbergen, Waygat

Tslands, 60 fathoms, August 1861, and in Ice Fjord, Safe Harbour,

30 fathoms, June 1864. I have also had at my disposal a colony

from Greenland, 65° 15’ N. lat., 53° 30’ W. long., collected in

June 1883; depth 75 fathoms.

The specimens are preserved in alcohol.

SPICULES.

Introductorily I will here mention that small spicules occur in

the test of Kikenthalia borealis.

As is known, the species forms colonies which are massive,

usually rounded, and not very large; they are attached by a

Text-figure 1.

ji ff

LOD ge

=) EZ NS Den

SQ Sy oh,

Oy SSeS,

FoI SShse

Lf ope

Kiikenthalia borealis (Gottsch.).

Spicules. X 110.

narrow base. The zooids are not arranged in systems, and the

atrial as well as the branchial aperture of each zooid opens

independently on the surface. The test of the colony is well

developed. Internally and at the base it is soft and gelatinous,

externally it is leathery. To this might be added also that the

capsules which surround the individuals of the colony are firm

and tough.

The test owes its consistency, partly at least, to the presence of

the above-mentioned spicules which are scattered in it. In

larger colonies the spicules are abundant, in smaller ones they

seem to be less numerous.

% Bierkan, P., “Die Ascidien des nérdlichen Norwegens.” Tromsé Museums

Aarshefter 24-25. 1908.

REPRODUCTIVE ORGANS OF KUKENTHALIA BOREALIS, 189

They show some variability with regard to size and shape.

The general plan of the structure is, however, the same. Text-

figure 1 shows two varieties of spicules which occur in different

colonies. From a comparison it appears that the crystals are

arranged about a common point forming like a sphere, studded

“with irregular, short, sharp points, or they radiate forming a

stellate group with rays of irregular length. Sometimes a ray is

elongated like a stalk.

According to the literature, spicules often occur in different

groups of Ascidians, but such structures do not seem to have been

observed before in any genus of the Polyzoine. The possibility

is, however, not excluded that they have been overlooked, since

their presence has been until now unobserved in Kwkenthalia.

But if this should prove not to be the case, the spicules, having

been found only in Aiikenthalia, are to be regarded as a charac-

teristic of the Arctic form.

As spicules. might: be of some value for comparative and

systematic studies, an investigation of their occurrence in the

different genera of Polyzoinw would be of interest.

THE REPRODUCTIVE ORGANS.

A more important result of this investigation is, as will be

shown, that the reproductive organs, testes as well as ovaries, have

been found in Aukenthalia borealis.

Text-figure 2,

Iiikenthalia borealis (Gottsch.).

Zooid seen from the left side, test partly removed. X 8,

at. Atrial aperture. ~67. Branchial aperture. 6. Bud. 6p. Brood-pouch. gp. Genital

pouch with testis and ovary. a. (Esophagus. r. Rectum. s. Stomach.

The organs are in the form of a hermaphrodite structure, the

male glands as well as the ovary and the ducts being enclosed in

-a long, sac-like outgrowth of the mantle which extends into the

common test, This outgrowth is here named the genital pouch.

14*

190 DR, A. ARNBACK-CHRISTIE-LINDE ON THE

Tn some individuals another sac-like structure, generally of

greater size, has been observed. It is situated at the upper side

of the above-mentioned one, and projects like the latter into the

common test. When an individual contained developing

embryos, they were enclosed in this sac. No doubt it develops to

receive the embryos, and it is consequently to be regarded as a

brood-pouch,

In the zooids most advanced in development there are thus

two pouches which are to be referred to the reproductive organs

(cf. text-fig. 2).

Tn the individuals examined, reproductive organs have been

observed only on the left side of the body; they are placed ventrally,

on the outer side of the mantle, and are embedded in the test.

Only one genital pouch—consequently only one brood-pouch—

has been observed in each zooid. If the individuals are seen from

the inside of the colony, the position as well as the number of the

organs in question is easily determined. The members of the

Text-figure 3.

or at

Kiikenthalia borealis (Gottsch.).

Zooid seen from the left side, test partly removed. X 7.

at. Atrial aperture. br. Branchial aperture. 6. Bud. gp. Genital pouch with

testis and ovary. ce. (sophagus. 7. Rectum. s. Stomach.

colony are arranged in one layer and are placed close to each

other side by side; only the ventral part is free. From here the

reproductive organs extend between the individuals, and only one

genital pouch is visible between them, projecting from the left

side of each zooid.

As in the genus Gynandrocarpa, the reproductive organs of

RKiikenthalia seem thus to consist of a single hermaphrodite

structure. In the former it is placed on the right side, in the

latter on the left.

In the different zooids of the same colony the reproductive

organs show various degrees of development. In some zooids the

genital pouch with the gonads was very large and of considerable

length, in others it was hardly distinguishable (cf. text-figs. 2 & 3).

REPRODUCTIVE ORGANS OF KUKENTHALIA BOREALIS, 191

A brood-pouch was developed only in few zooids. It could not

be decided whether all the zooids of the colony were mature, the

material being little fit for dissection. From the above-mentioned

facts one might, however, conclude that the members of the

colony do not attain sexual maturity at the same time, probably

owing to different age.

As appears from text-figures 2 and 3 the part of the genital

pouch which contains the genital glands is of a bulb-shaped

form. On one side a large swelling, enclosing the distal part of

the oviduct, is visible, and its form varies, owing to the degree

of development of the brood-pouch, as will be described in the

following.

Mate.—The testis is composed of numerous spermatic vesicles

and a long vas deferens.

As has been mentioned above, the testis is placed on the out-

side of the body-wall, invested by an outgrowth of the mantle,

projecting into the common test. The vas deferens, which is a

duct of considerable length, extends into the peribranchial

cavity and opens into it.

Text-figure 4.

[

Kiikenthalia borealis (Gottsch.).

Vas deferens. X 33.

A. Vas deferens, usual form. | B. Vas deferens, coiled up.

vd. Vas deferens.

The spermatic vesicles are pyriform glands which are arranged

in two bundles, on account of which the testis appears to be

bipartite. The male glands open into two sperm-ducts, each

bundle having its duct, and these unite as a rule into a main

sperm-duct, the vas deferens (cf. text-figs. 4 4 & 5).

Tn one individual examined another arrangement was observed.

Here the two ducts did not join, but entered separately into the

peribranchial cavity, thus forming two vasa deferentia.

After its entrance into the peribranchial cavity, the sperm duct

192 DR. Av ARNBACK-CHRISTIE-LINDE ON THE

generally bends somewhat to the side, extending along the inner

wall of the cavity. This arrangement was observed in all

specimens but one. In the latter, a rather large individual with

a strongly developed testis, the distal part of the vas deferens was

coiled up (¢f. text-fig. 4B), and its opening lies close to that of

the brood-pouch.

As this arrangement has been found in a single individual only,

I do not venture to draw any conclusions from it. The possibility

seems, however, not excluded that it might be of some impor-

tance for the fertilization, in this case probably the self-fertiliza-

tion of the individual. The fact that ova and spermatozoa are not

seldom found mature at the same time in the same individual

appears also to support the view that self-fertilization can take

place; whether it is usual is another question.

Though most Ascidians are hermaphrodite, it is, however, not

probable that self-fertilization generally takes place. On the

contrary, in many groups arrangements have been observed which

seem to prevent it. As an instance of such an arrangement, J

will mention that ova and spermatozoa are often produced at

different times, and are consequently not found mature together.

From sections it appears that, in zooids belonging to colonies

caught in June, masses of spermatozoa fill up the sperm-duets.

Other cells are macerated, but the spermatozoa are in good con-

dition—a fact which ought to be noticed, the material being

preserved in alcohol and dredged so many years ago. It confirms

the observation made many times before that the spermatozoa

are more resistant than other cells, and that they are highly

resistant, even to maceration.

FemaLe.—In zooids most advanced in development the female

organ consists of a small ovary with a wide oviduct and a very

large brood-pouch.

Asis shown by text-figure 5, a small rounded vesicle with a

wide duct is situated at the side of the male glands in the genital

pouch. Sections through it did not show much of its structure,

the tissues being rather macerated. ‘The presence of eggs makes

it, however, evident that the vesicle is to be regarded as an ovary.

In coveral individuals one very large egg, covered with follicular

epithelium, was observed here, Apparently ready to pass through

the oviduct (cf. text-fig. 7).

As mentioned above, the ovary is placed at the side of the

male glands and is quite separated from them. Seen from the

left side of the zooid, the one bundle of male glands is partly

covered by the other, and the ovary has its position opposite their

middle line. If one imagines the gonads spread out, the ovary

would thus have its position between the two bundles of male

glands.

The oviduct extends from the above-described ovary between

the two sperm-ducts towards the brood-pouch, and opens into the

distal part of the latter with a very wide aperture (c/. text-fig. 5).

The lumina of the brood-pouch and the oviduct become thus

REPRODUCTIVE ORGANS OF KUKENLHALIA BOREALIS. 193

distally confluent, and they comm-nicate with the peribranchial

cavity by a short, narrow aperture which is well observable near

the vas deferens (cf. text-fig. 8).

These facts are of a special interest; then, owing to the above-

mentioned arrangement, the passage of the egg is secured: the

oviduct opening widely into the brood-pouch, t the egg, having left

the oviduct, must be laid into the brood-pouch, where it pr obably

remains until it is fully developed.

The oviduct is of considerable width, especially the distal

part, which is rather sac-like, and the wall is deeply folded on

one side.

Text-figure 6 represents a transverse section of the genital

pouch on a level with the bifurcation of the vas deferens. The

latter is on the point of forking, on account of which it appears

somewhat broad. The figure shows how the oviduct is somewhat

compressed from side to side: it is lined with a low epithelium.

Text-figure 5.

Kiikenthalia borealis (Gottsch.).

The reproductive organs from the outside. X 23.

bp. Brood-pouch. o. Ovarium, od. Oviduct. ¢. Testis. vd. Vas deferens.

More distally, on certain parts of the wall, a strongly ciliated

epithelium was observed. Asappears from the figure, the ciliated

epithelium of the vas deferens is also well developed.

The brood-pouch has the form of a large double-walled sac

with a constricted neck. It generally contained one embryo.

In some individuals it is of considerable [one ih and width, pro-

jecting beyond the genital pouch, Text- fisure 5 5 represents ib at

a stage somewhat less developed.

In individuals containing no embryos a brood-pouch has: also

been observed, though highly reduced with regard to form and

size. Such an interesting stage is shown by text-figure 7, where

the brood-pouch is represented by a short blind sac or an appendix

in connection with the oviduct; and in zooids with rudimentary

reproductive. organs, as. for instance.in that represented by

text-figure 3, the presence of such a blind sac can also be

stated, as will be seen from text-figure 8. The last-mentioned

194 DR. A, ARNBACK-CHRISTIE-LINDE ON THE

figure shows three transverse sections of the genital pouch with

the ducts, which are cut obliquely. The brood-pouch is repre-

sented by a short blind sac ; like the oviduct, 1t is compressed from

side to side. In text-figure 8@ the oviduct and the brood-pouch

Text-figure 6. Text-figure 7.

Text-fig. 6.—Kiikenthalia borealis (Gottsch.).

Transverse section of the genital pouch. X 110.

od. Oviduct. wd. Vas deferens.

Text-fig. 7—Kikenthalia borealis (Gottsch.).

The xeproductive organs from the outside. X 23.

bp. Brood-pouch. o. Ovarium. cd. Oviduct. t. Testis.

vd. Vas deferens.

Text-figure 8.

Kiikenthalia borealis (Gottsch.).

Sections through the genital pouch, showing the distal part of the oviduct and

the rudimentary brood-pouch; cut obliquely. > 60.

bp. Brood-pouch, od, Oviduct. o. Opening into the peribranchial cavity.

vd. Vas deferens,

are still separated ; fig. 6 shows how the oviduct opens into the

brood-pouch, and in fig. c the opening into the peribranchial cavity

is visible.

Whether this blind sac is to be regarded as a vestigial trace or

possibly as a rudiment of the brood-pouch, is a question which

REPRODUCTIVES ORGANS OF KUKENTHALIA BOREALIS. 195

ought to be examined in connection with that of the origin and

formation of the whole complicated female organ. As it is

difficult to state anything for certain about the age of the

individuals hitherto examined, we cannot draw any conelusions

from the above-mentioned facts as to the presence of a rudi-

mentary brood-pouch, even before the first egg has been laid.

Another matter of interest 1s the question of the homology of

the brood-pouch in Aiikenthalia. Before deciding it, it ought to

be investigated whether the brood-pouch of the species in question

arises as a direct projection from the peribranchial cavity, or

whether it is possibly formed in connection with the oviduct with

which it, when fully formed, is so intimately connected.

In other Ascidians in which a brood-pouch develops to receive

the embryos, it is generally a diverticulum of the peribranchial

cavity.

Though the systematical position of Avikenthalia borealis will

not be discussed in this paper, I will, however, point out that,

as the structure of the reproductive organs has been made the

chief character in distinguishing the genera belonging to the

group Polyzoime, the facts which have been ascertained by this

investigation will no doubt be of decisive importance for the

systematic position of Avikenthalia. Comparing the reproduc-

tive organs of the last-mentioned genus with those of the other

genera of the group, we find that the most striking difference is

the presence of a brood-pouch in Avikenthalia. Though the

Polyzoine have been the object of thorough investigation,

especially with regard to the structure of the reproductive organs,

nothing which corresponds to a brood-pouch seems to have been

observed. One might thus conclude that it occurs only in the

Arctic form, of which it is consequently characteristic.

With respect to the genital pouch, the beginning of analogous

structures may possibly be found in some genera, though they

have not attaimed the same degree of development as in

Kikenthalia.

In certain respects, points of agreement may be found between

Kikenthalia and Gynandrocarpa, as, for instance, they agree with

regard to the reduced number of the gonads and the structure of

the testis {ef. Herdman™, pl. 44. fig. 4).

NoN-GONADIAL SEXUAL PRopDUCTS.

Though I now venture to maintain that the presence of gonads,

testis as well as ovary, in Aitikenthalia borealis is put beyond a

doubt, and that it has been proved that ova and spermatozoa are

produced in special organs, there are, however, certain facts

concerning the reproduction of the species which still seem

* Herdman, William A., ‘‘ Report on the Tunicata collected during the voyage of

H.M.S., ‘Challenger’ during the years 1873-76,” Part 2, vol. xiv. London, 1886.

196 THE REPRODUCTIVE ORGANS OF KUKENTHALIA BOREALIS.

unexplained. Such a fact is, for instance, the presence of sexual

products, ¢. e. ova, in the mesoderm.

Isolated ova have been observed before in the mesoderm of

Kiikenthatia, but it has not been made out where they originate.

As no gonads, only isolated ova were met with, Michaelsen

concluded that the female organs of Kiikenthalia consist of ova

diffusely placed in the mesoderm. Michaelsen writes as follows:

“Diese Gattung weicht durch den Bau der weiblichen Geschlechts-

organe von allen tubrigen Polyzoinen, ja, von allen Styeliden

iiberhaupt, ab. Ich bezeichnete die Gestaltung desselben als

‘diffus,’ da die sich entwickelnden EHizellen an kein bestimmt

lokalisiertes Organ gebunden sind, sondern sich weit zerstreut

im Innenkorper und in den Blutbahnen anderer Organe, so

besonders des Darmes, sowie in den jungen Knospen, vor-

finden. Der urspringliche Ort der weiblichen Gonaden. ist»

unbekannt...” (d. ¢. p. 112).

Though female organs of other structure have now been

found, the observation that isolated eggs occtir in the mesoderm

proves to be right. According to my observation, they are,

however, met with less abundantly than has been stated above.

As it hardly ¢ can be thought that they originate in the above-

described ovary, one must suppose that they are produced in

the mesoderm, where they occur, though in no special gonads,

2. ey MONE aonadially. This supposition seems to be verified by

my observation of small ovaries or, rather, groups of eggs which

are situated in the mesoderm close £0 the wall of the peribranchial

cavity and in the vicinity of bud-rudiments.

In Kiikenthalia, buds occur on both sides of the body ; on the

left side they are to be seen in the vicinity of the reproductive

organs above described. In the mesoderm of those bud-rudiments

eggs have been observed, and even in very small bud-rudiments

very large eggs are often visible. In all probability the eggs

originate in the parent animal, the generative cells being sup-

posed to wander from the latter; but the question is whether

they are differentiated in the bud or have wandered at an

advanced stage from the parent animal into the bud. But this

ig a problem which can be decided only after thorough investi-

gation of the further development of the non-gonadial ova as well

as of the formation of the reproductive organs of the bud.

VARIATION OF THE SCAPULA IN THE BATRACHIA. TSG

10. On the Variation of the Seapula in the Batrachian

Groups Aglossa and Arcifera. By Joan B. Procrmr,

F.Z.8.

[Received October 29, 1920: Read March 8, 1921. |

| (Text-figures 1-10.)

The variation of the scapula in the Batrachia Heaudata is of

considerable interest, both on aecount of its wide range and of

its comparative stability within a genus. The present study has

been confined to the Aglossa and the Arcifera as representing the

most primitive types now living. JI have examined all the

skeletons in the collection of the British Museum, but this

material, although large, is not as complete as one could wish.

Text-figure |.

(b)

Bufo hematiticus. X2.

(a) Pectoral girdle from without ; (6) scapula from within.

p.a., pars acromialis. .2., pars glenoidalis. sh., shaft. sup.art., superficial

articulation. f.a., fovea acetabulum. c.f., central foramen. p.c., paraglenoid

cartilage.

In the majority of species the scapula is about equal to the

preecoracoid in length ; it is usually rather slender in the middle

and expanded at each end. The proximal end consists of two

parts, generally separated by a cleft: the upper or pars acronualis

is either in contact with the head of the preecoracoid, or narrowly

separated from it by acromion cartilage ; the lower or pars glenoi-

dalis is much depressed and coneave, forming the bottom of the

glenoid cavity—it may be in contact with the end of the coracoid,

or separated from it by paraglenoid cartilage. ‘This cartilage also

forms a subtriangular knob projecting below the head of the

precoracoid, and attached to the head of the humerus by a liga-

ment. The portion of the scapula between the glenoid cavity

198 MISS JOAN B. PROCTER ON THE VARIATION

and the supra-seapula will be alluded to as the shaft. A circular

depression is often situated in the thickness of the shaft, where

it becomes one with the pars glenoidalis: it is called the seper-

ficial ar ueulation, and forms the distal wall of the glenoid cavity 5

opposite to it is the fovea acetabulum, a similar depression i in the

extremitas scapularis of the coracoid, which forms the proximal

wall of the glenoid cavity.

In order that the varying proportions in the different genera

may be fully Sp Pregaied: the length of the precoracoid will always

be given as 10 units*, and all bther measurements standardized to

this scale; in this way the comparative reduction and develop-

ment can be seen at a glance, as the normal length of the scapula

should be about 10 also.

In fossil frogs of the genus Paleobatrachus, the scapula was

shorter and more compact than in the majority of modern genera,

being three- to four-fifths the length of the precoracoid, and about

once and a half as long as broad at its expanded end. The pars

Text-figure 2.

Scapula of Paleobatrachus sp.? %2. (After Wolterstorff.)

acromialis was not distinct from the pars glenoidalis, but, accord-

ing to Wolterstorff, a groove can be seen on the inner surface of

the bone, in place of the cleft shown by modern forms.

The following figures are standardized from actual measure-

ments given by Wolterstortff slic

Examples.

P. fritschii. | P. grandipes. . P. wetzleri.

Length of precoracoid ... 10 10 10 10

a Seapulann eerie 6 6:4 ice 8:5

Breadth of ,, (proximal) 3 3°6 4°5 57

a oy (asta) xt A A-5 ay) 6:7

Although the size of the scapula varies in comparison with

the precoracoid, its own proportions are remarkably stable,

Wolterstorff says of it: ‘Ks finden sich nur wenige vielleicht

nicht einmal specifische Verschiedenheiten.”

* Units of length.

+ “Ueber fossile Frésche, insbesondere das Genus Pal@ohatrachus’ (Theil 1.

1886; Theil i1. 1887).

in Opucuts

OF THE SCAPULA IN THE BATRACHIA. 199

Preip#,

As in Paleobratrachus, the genera of this family have the

scapula entire, without the cleft which separates the pars

acromialis from the pars glenoidalis, at least partially, in all other

families. This, together with the marked reduction of the

scapula, constitute diagnostic characters for the Pipide,

Xenopus Wael.

X. levis Daud.—Seapula vestigial ; subtriangular. Shaft and

pars acromialis completely absent; pars glenoidalis containing

the circular depression of the glenoid cavity. Head of precora-

coid in contact with supra-scapula. Occasionally the scapula is

Text-figure 3.

Pectoral girdle of Xenopus levis. X2.

(a) scapula distinct ; from without.

(5) scapula not distinct; from within,

apparently wanting, having fused with the head of the preecora-

coid; in most specimens this fusion is more or less marked,

making the boundaries of the vestigial scapula difficult to trace.

Example.

Menethvonmorecoracoldinca tresses. see. 10 units.

t. Scajpul ay (at ase eee a ecto. oe Ase

Breadth,of 5; | (expanded:end) ..........: Pree irene

Weng thvol suprasscap ula resqee serene. cs LOrSy

X. muellera Ptrs.—Scapula as in X. levis. A figure of the

pectoral girdle is given by Peters*, in which it is absent. In

* Peters, *‘ Reise nach Mossambique’ (1882) Taf, xxvi. 12 a.

200 MISS JOAN B. PROCTER ON THE VARIATION

his specimen it is evidently incorporated in the precoracoid, as

described above.

X. calearatus Buchh. & Ptrs.—Scapula as in X. levis, but

more easily distinguished, as the glenoid cavity extends to the

upper border.

X. clivii Peracca.—Scapula with a distinct, but vestigial,

triangular shaft, the apex of which narrowly separates the head

of the preecoracoid from the supra-scapula ; pars glenoidalis as in

X. levis; pars acromialis absent.

Example.

eneth on ipreecoracovd ican ore. eet cere 10. units.

RS Scapulax(@it base), 2... a eis 352 5

“ at (upperiedge)) 7a cee enue DI) a

a 3 SHOE WSRG VaBer i ias MECeRSe ROC many lise se

Breadth of _,, (expanded@entd))ime an cccecr BS)

Ment of cuprasseapulla 220.02. ceanias oe Orns

In the tadpoles and young of X. levis, X. calcaratus, and

X. muellert the reduced condition of the scapula is somewhat

less marked than in the adults; an extremely short shaft is

present, precisely as in the adult of X. clwu. If ontogeny may

be taken as a guide to phylogeny, Y. clivii should be considered

the most primitive species in the genus.

This vestigial form of scapula, which even in YX, clivii does not

exceed two-fifths of the preecoracoid in length, and which may

be completely suppressed in other species, renders Xenopus

unique both among the Aglossa and Arcifera.

Hymenocuirus Blgr.

H. boetiegert Tornier.—Scapula much reduced, but less so than

in Xenopus. Distal end expanded, proximal end narrowed,

entire; the whole wedge-shaped. Proximal end strongly over-

lapped on outer surface by head of precoracoid. No glenoid

cavity.

Hxample.

ddenieth of preecoracoid; backnet- dere iiscs- dye: 10 units.

ae Seajouilan(@reawesty)i aterm ounce sacra Drie,

mi sg (appemiedine)) gas itiear eee sald ie

Breadth of _ ,, (expanded end). ............ Oro,

uy He (least sor. e8 de Balcola sts DHL

Length of supra-scapula ...............0seeseaee ore) 4

Pipa Laur.

P. americana Laur.—Scapula compact, very much reduced,

almost triangular when seen from within. Shaft vestigial; pars

acromialis strongly overlapped by head of precoracoid on inner

surface; a strong crest and protuberance on outer surface, bor-

dering the glenoid cavity ; pars glenovdalis much depresssed when

seen from without, not distinct from pars acromialis when seen

from within,

OF THE SCAPULA IN THE BATRACBIA. 201

Heample.

dhemaiih: of joreecome. Goudy snes ans css ee RS. 10 units.

53 Scapullace as eeenoe. Sowa) phate gor. Bye) ian

x i (tree-upper adge) .......... eg)

as Bites SUE OR A te 2 Se ee Medea

Breadth afi cp... (Gusheiitomel \isteihy te... see. DeGrres

4 SM MUPASD) pace WE ERiR tT, 1689,3

ength of Sippra seaplane iostece ce. coccsace. ons SF iiee

Text-figure 4.

Pectoral girdle of Pipa americana. X2.

(a) from within; (5) scapula from without.

ARCIFERA.

In all the Argiferous genera in which I have studied the

scapula, its three main branches are all present, even if one or

more of them is reduced. The pars acromialis is more or less

separated from the pars glenoidalis by a cleft or foramen.

DISCOGLOSSID®.

As in Pipide the scapula is greatly reduced. The pars

acromiatlis more or less overlaps the pars glenoidalis, so that the

cleft which separates them can only be seen when viewed

obliquely.

DiscoaLossus Otth.

D. pictus Otth.—Seapula much reduced, its greatest length

only half that of the precoracoid. Shaft vestigial ; pars

acromialis well developed, but shorter than pars glenoidalis ; not

overlapped by. head of przcoracoid; pars glenoidalis well de-

veloped, strongly depressed, the separating cleft pierced hori-

zontally. In eight skeletons of this species there appears to be

no individual variations,

202 MISS JOAN B. PROCTER ON THE VARIATION

Hxample.

enethiot jrecoracoid (fhe. -chenacse eee 10 units.

SS scapula (lowersborder)\.u-ceeeee: Spee

5 ». (free-upper edge) ......... alas

if Mane S| OIE oh acnapeee Ria Sealey PIE

Breadth of). )...//(distallfewd) sine oe os ag 3°D 4,

ength of supra-seapulla:ons.2 5m ee ae an epee oe

Text-figure 5.

Pectoral girdle of Discoglossus pictus, X23; from within.

BomBinator Merr.

B. igneus Laur.—Scapula much reduced, subtriangular in

shape; upper edge shorter than distal edge. Shaft vestigial ;

pars acromialis much reduced, forming an oblique suture with

head of precoracoid; pars glenowdalis much depressed, longer

than pars acromialis, from which it is separated by a notch

rather than by a cleft.

Example.

ene th¥ok preecoracordin ei esnecesn eee astele cn 10 units.

. scapula (lower border)... )\s0.0. 4... A3 ,,

Ae Bey ms) 0) OX racine Clatly)aoe aa eur aes A DAA) > 5

ie Sy MO AL Deets mice aneent neonates sion

Breadthvote sy.) (distalvend) yr ne ee eee Soo ee

Wengethiot supra-sca pula ey. cwsec. contents os Sale

Text-figure 6.

Pectoral girdle of Bombinator maximus, X2; from without.

B. pachypus Blgr. and B. orientalis Blgr—Scapula as in

B, wgneus.

B. maximus Bler.—Scapula as in B. ignews in shape and pro-

portions, but differing in that the upper border and adjoining

OF THE SCAPULA IN THE BATRACHTA. 203

head of precoracoid are thin almost to transparency, forming a

semicircular crest, the diameter of which follows the free upper

edge.

The proportions of the scapula are remarkably stable within

the genus Bombinator, and are highly characteristic.

AtytTEes Wagl.

A. obstetricans Laur.—Scapula short, but less reduced than in

Discoglossus and Bombinator. Upper border forming a semi-

circular crest thin to transparency, resembling Bombinator maxt-

mus in this respect. Shaft reduced; pars acromialis narrow, in

contact with head of precoracoid; pars glenoidalis broad,

depressed, equal in length to pars acromialis; the two over-

lapping, so that the dividing cleft is pierced horizontally and

only visible obliquely.

Example.

Wenoth: of preecoracord a. eaaeecereetat ee 10 units.

3h scapula (lower edge) ............... Ar).

‘3 yo.% \CUpperd aaah) tas: LSee se BAO)

ids * SAVIN Aun ALAS 5 Canin n is Dns ha ae 22h.

iBreadthiofy 15°, (distalkemd)) Wee ases.cr ss Aoi

luength of supra-scapula .............-.-. dean aD es

PELOBATID®.

In this family there is a wider generic diversity in the propor-

tions of the scapula, which is not, however, greatly developed in

any genus. In all, the glenoid cavity is excessively deep, some-

what under mining the pars acromialis, the lower edge of which

circumscribes it above.

Scarutorus Holbr.

S. solitarius Holby.—Scapula well developed, longer than

precoracoid ; as long as the supra-scapula. Proximal end well

developed, subcircular. Shaft long, wedge-shaped ; pars acromialis

large, with crest-like upper border, lower border projecting over

and round glenoid cavity in a strong ridge; pars glenoidalis

small, strongly depressed, the dividing cleft short and broad,

forming a small oval foramen in the glenoid cavity, pierced

vertically.

Hxamople.

Ientethyok. preecoracoid Meurer nsjacuiseca- sce 10 units.

re scapula (upper or lower border)... 13 _ ,,

= soja MAS GUL eee reecatiee crate ahd 9-4 ,,

Breadth of ,, (distal end)............ Breer Heo) Be

Pomme (0TUKCCGIKS)) orn eno eteee seem are ES)

Length of supra-scapula ......... Seer carat aia.

Proc. Zoou. Soc.—1921, No. XV. 15

204 MISS JOAN B. PROCTER ON THE VARIATION

PELOBATES Wagl.

P. fuscus Laur.—Scapula well developed, thick, once and a

half as long as precoracoid. Shaft oblong, twice as long as

broad; pars acromialis broad, the lower border of its outer

surface with a ridge which runs round the glenoid cavity ; pars

glenoidalis much depressed ; dividing cleft short, forming an

oval foramen pierced horizontally.

Example.

Length of preecoracoid ............... Leis iS dee 10 units.

s scapula (lower edge) ............... iD. hres

uk Wigs NeW i We ganeee AE me ec NOTA: ;

Breadthroh am = (distalend)) ii ee-castes pees. Oe. eae

Length of supra-scapula (lower border) ...... HS 0,

P. cultripes Cuv.—Scapula precisely as in P. fuscus.

Pretopytes Fitz.

P. punctatus Daud.—Scapula much reduced, squarish in shape.

Shaft shorter than broad at distal end; pars acromialis as broad

as long, lower border of its outer surface forming a prominent

ridge running round the glenoid cavity ; pars glenoidalis much

Text-figure 7.

Pectoral girdle of Pelodytes punctatus. 2.

(a) from without; (6) scapula from within.

depressed, subcireular ; dividing cleft short, pierced horizontally,

only visible when viewed obliquely.

Vacample.

eno thton preeconacoids cp -cnun ce sseers ee eee 10 units.

is scapula (upper border) ...... sate DEO) es

M SRT S C1 Clie, aay unary ot 4-4 ,,

Breadth ole teen a (Gistealiend)) nena saseemetner 4-4 ,,

Length of supra-scapula (upper edge)......... 9-4 ,,

In shape the scapula of Pelodytes is very similar to that of

Paleobatrachus, which it resembles in many ways. Paleeba-

trachus, however, lacks the proximal dividing cleft, which,

though outwardly invisible, is present in Pelodytes.

Barracuopsis Bler.

B. melanopyga Dovia.—Scapula well developed, about onee and

a quarter as long as precoracoid ; slender in the middle,

OF THE SCAPULA IN THE BATRACHIA. 205

distal end broader than proximal. Shaft long, wedge-shaped ;

pars acromialis with straight oblique upper border, upper half of

proximal end free; pars glenoidalis much depressed, slightly

undermining pars acromialis; dividing cleft short, pierced hori-

zontally, only visible when viewed obliquely.

Heample. +

eng th of preeconacord inc nc ses cteece 10 units.

3 scapula (upper or lower edge) ... 12:2 ,,

7 See GEST ENCE" Gato eae ae JA.

Breadthvote ¢.g. (Gistaljend). s..n2 4. . seceas. Onl.

i Mie (att CICS)" Gener, Senn enrane 2 OL ae

ue St a((D ROD STI GrOVal)) Ae Dies

ene tia of suppral-scaouilapeeereeereer er cen laces: WleGiae

Mecatorurys Kuhl.

M. pelodytoides Blgr.—Scapula well developed in all its parts,

as long as the precoracoid. Shaft wedge-shaped, as long as broad

at distal end; pars acromialis with an oblique straight crest

along its upper border, rising to a sharp point proximally; lower

ridge of outer surface with a prominent ridge running round the

glenoid cavity ; upper half of proximal end free; pars glenoidalis

much depressed; dividing cleft short and broad, pierced

obliquely, forming a foramen, which is longer when viewed from

within than when viewed from without.

Text-figure 8.

Pectoral girdle of Megalophrys parva. X2.

(a) from within; (6) from without.

Example.

Wenethiotpreconacordansney. cee eens 10 units.

A S@a ula 655 Pee ee had tabs LORS ce

% BAe SUN DY Ga coceace : SneaRee ee GoD

Breadth ofs;. 5,9) s(distalwemel)rentaetss 0s. c00 Os25e

Weng throb supra-scapulamy eae eee eh «ccc... Plein.

M. major Blgr.—Scapula similar in form to that of JL. pelody-

toides with the exception of the pars acromialis, the crest of

which has a paracentric upper edge; upper half of proximal end

free.

M. parca Blgr. (= Xenophrys menticola Gthr.).—Scapula as

15*

206 MISS JOAN B. PROCTER ON THE VARIATION

in M. major. In this species the preecoracoids are reduced, and

do not meet each other in the middle; the scapula is, therefore,

longer than the precoracoid, but not longer than its distance

from the edge of the epicoracoidal cartilage.

M. montana Kuhl.—Form of scapula as in MW. major, but

slightly larger than the precoracoid, which is normal in length.

M. longipes Blgr. —Scapula as in M. major.

M. fee Blgr.—Form of scapula similar to that of M7. major,

with the exception of the distal end of the shaft, the lower corner

of which is somewhat produced, making the lower border of the

scapula once and a quarter the length of the upper border ;

lower border of scapula describing an inverted semicircular

curve, ~>. Pars acromialis with a semicircular upper edge.

Although there is specific differentiation within this complex

genus, the form of the scapula is highly characteristic. The

equal development of the shaft, the pars acronualis, and pars

glenoidalis, together with the oval central foramen, pierced

horizontally, and the highly developed crest of the pars acromialis,

are characters common to all the species.

Hynip&.

In this family the scapula is usually normal in its proportions,

and about equal in length to the preecoracoid. In some instances,

however, it is greatly reduced in length, especially the shaft.

The cleft between the pars acronialis and pars glenoidalis is

always broad and open, pierced vertically, and visible from with-

out and within as a large oval foramen in the middle of the glenoid

cavity. The pars ylenoidalis is reduced in size, and always

shorter than the pars acromialis, part of it presumably giving

place to the central foramen.

Hyta Laur.

H. maxima Laur.—Scapula normal in its proportions, equal in

length to the precoracoid, and having all the family charac-

teristics already described. Shaft once and a quarter as

long as broad at distal end; pars glenoidalis separated from

coracoid by cartilage; superficial articulation and fovea aceta-

bulem very distinct.

EKaample.

ene thiol qorcecoracoid jene..-ceccescecnee teres 10 units.

. scapula (upper border)............... MO 5

s pa SOT G 6 iionse ee area Ole

iBReadtheot } apne (distal em) eaeeeersass.chy 6-4 ,,

. am. (Tard dle), 5. <5 eteel eerie ren Zane

zs OOHRS OMIA WBSendanes sabe assess ses 56 WSS) a

ee DOE ONACOCITUOTD- Sa 9G aoncoeasouore st eS) ay

Length of pars acronvialis............)..0.2-20 +2. Melee.

mi TOCUPS GUA OOUKHNS) “scposcecacosacsasnee: 18h,

ie SIU OT N-SGAY OWE, Sowkonbocbaec soo oe oaeKos hose

OF THE SCAPULA IN THE BATRACHIA. 207

In the following species the scapula is as in /H. maxima :—

H. fauber Wied, H. albomarginata Spix, H. boans Daud., 1. taurina

Fitz., H. venulosa Laur., H. nigromaculata Tschudi, H. domini-

censis Bibron, H. versicolor Lec., H. pulchella D. & B., H. arborea

L., H. macrops Blgr., H. cwrulea White, H. dolichopsis Cope,

HI. aurea Less., H. lesweurit D. & B., H. nasuta Gray, H. gratiosa

Lec.

In H. lichenata Gosse and H. baudini D. & B., the scapula is

a little longer than the precoracoid.

Although Hyla is a very large genus, it will be seen that the

scapula is remarkably stable in its proportions. lt is not of

much use, however, as a diagnostic character, as it is of the form

most commonly met with in this family.

Nororrema Gthr.

NV. marsupiatum D. & B.—Scapula elongated, once and a

quarter as long as precoracoid, very slender in the middle. Shaft

as long as precoracoid; pars acromialis and pars glenoidalis as in

Hyla maxima.

PrernouyLa Bler.

P. fodiens Blgy.—Scapula as in H. maxima,

Nycrimantis Blgr.

NV. papua Blgr.—Scapula as in H. maxima.

AGALYCHNIS Cope.

A. moreletii A. Dum,—Seapula as in H. maxima.

PuyLLomEDusA Wazgl.

P. dacnicolor Cope.—Scapula abnormally developed, about twice

as long as precoracoid, and about twice and a half as long as

broad at distal end. Shaft about once and a quarter as long

as precoracoid, very slender, expanding at distal end; pars

acromialis not in contact with head of precoracoid; pars

glenoidalis very much reduced, branching widely from pars

acromialis ; central foramen very large, oval.

P. bicolor Bodd.—Scapula similar in form to that of P. dacni-

color, but even more developed in length, being twice and a

quarter the length of the precoracoid. Shaft once and three-

quarters the length of precoracoid; pars acronvialis separated

from head of precoracoid on outer surface by acromion cartilage,

in contact on inner surface.

Eeample.

Wengthyek precoracovdiy ceeds. cree. deel. . 10 units.

fy scapula (lower border)............ we Loe fs

# hue | N(USPe Te WEAN) ta. oo Seis i DA nos

SInaititny) Hees aoe ei te ae Lb ihe 33

”? ”)

208 MISS JOAN B. PROCTER ON THE VARIATION

Breadth of scapula (distal end)................. ~ 10:2 units.

i spbvenn(gantclclile))b es atsts feces see ey B2kOF a.

Ms DONS \ACRONMULLIS He tncat. . So. seeie ts os aH oe

ws ars Glemotdaliswar. .hcrtya-aasceneee Me Rens

i central foramen get enh) .caeeeee DON

Length of — ,, Fs ee ha Ae aa se loca ea 3 4:4 ,,

(QUES CEROMLENS osebacssecnoncneosn>: By) 5,

iy WATS GlLEROUAALIS \...n.cen eee Reee ny: Lahishvee

A SUpra scapula pa sttensesnere AO Re i are a Sie

Text-figure 9.

Pectoral girdle of Phyllomedusa bicolor, X2; from without.

P. burmeistert Blgr.—Scapula as in P. daenicolor.

This form of scapula is highly characteristic of the genus

Phyllomedusa.

TRIPRION Cope.

T. petasatus Cope.—Scapula as in Hyla maxima.

BUFONIDA.

In this family the scapula is usually normal in its proportions ;

the pars glenoidalis is equally developed to the pars acromialis.

No important variations occur.

EuPEMPHIX Steind.

HL. natterer’ Steind.—Scapula normal in its proportions ;

slightly longer than precoracoid ; slender in the middle; similar

in shape to that of Batrachopsis melanopyga. Dividing cleft

between the two pars short.

Buro Laur.

B. vulgaris Laur.—Scapula normal, upper border about as long

as precoracoid, Jower border a little longer. Shaft stout, sub-

triangular ; pars acromialis somewhat diamond-shaped, with a

straight, oblique upper edge; pars glenoidalis much depressed,

narrower, but equal in length to the pars acromialis, from which

it branches, forming a Y of which the shafts represent the tail;

pars glenoidalis widely separated from coracoid by paraglenoid

OF THE SCAPULA IN THE BATRACHIA. 209

eartilage ; superficial articulation and fovea acetabulum strongly

marked.

Hxample.

Length of precoracoid (greatest)......... yas 10 units.

Ki scapulai(upper border)’ ].2200. ... VOSA Oe

a Kane (LOW EMRE Ot carse'srdoe Ne TO 2s

“i Sa ugar LAER Gs cot aun Sone a ees ene (ES Nes

Breadt hyo. 951 (distalneid) meeseeesseeeese ee. (Onn

by Apia {ONG CUO) soda aes MERoar Eman 3°60)

Ry FOES CXAROIMOUTUOS » niciotsdnbadouocbunnunds Zo ace

ue DANS, GUCNOULCIISN Meee anehe ees taces Os INI ays

ene thin sujpra=scayoullar teases ssc tase a: 253)

In the following species the scapula is like that of B. vul-

garis :—B. hematiticus Cope, B. calamita Laur., B. viridis Laur.,

B. intermedius Gthr., B. lentiginosus Shaw, B. marinus L., B.

crucifer Wied.

In Bb. latifrons Blgr., B. tuberosus Gthr., BD. melanostictus

Schn., 4. claviger Ptrs., B. quadriporcatus Blger., B. typhonius L.,

and BL. valliceps Wiegm., the seapula is a little longer than the

precoracoid, but of the same form as that of B. vulgaris.

In B. granulosus Spix the scapula is a little shorter than the

precoracoid.

There is remarkably little variation in this genus, in some

species the pars glenoidalis is somewhat shorter than in others,

and there is slight variation in the length of the shaft.

EHaample to show Range of Variation in Bufo.

B. granulosus. B. melanostictus.

Length of preecoracoid............ .. 10 units. 10 units.

rr scapula (lower border). 7:7 ,, 14°20 55

a Seorshabkh: yewiAvnes Ghee RO, 3%

Breadth of ,, (distal end) ... 6:2 ,, Glas:

2 itn (muatdl clike)) eet. A BB) ep =) ae

He pars acromialis ......... A’ Gone ey caer

“ pars glenoidalis ......... Si hn 5) ee

Length of supra-scapula ............ 10-4 ,, 14°3 .,

B. melanostictus has been chosen as an example of the more

developed type of scapula, as, being a common Malayan form, it

forms an interesting comparison with 3. granulosus from

E. South America.

The equal development of the pars acromialis and pars

glenoidalis in length, together with the fact that the latter is

always widely separated from the coracoid by cartilage, dis-

tinguishes the scapula of Bufo from nearly all other genera of the

Arcifera (see text-fig. 1).

RurnopuryNus D. & B.

R. dorsalis D, & B.—Scapula as in Bufo.

210 MISS JOAN B. PROCTER ON THE VARIATION

CYsTIGNATHID#.

In this family the scapula is extremely variable. It may be

slender or stoutly formed, shorter than the precoracoid or

abnormallly elongated. The range of variation is large as in

Hylidae, and very much greater than in any other families of the

Arcifera.

Pseupis Laur.

P. paradoxa* L.—Scapula equal in length to the precoracoid,

rather stout, parallelogrammic in shape; superficially similar in

shape to that of Pelodytes punctatus. Upper border of shaft

bearing a thin straight-edged crest; pars acromialis broader

than long ; pars glenoidalis narrow, obtusely pointed, partially

separated from pars acromialis by a notch forming part of a

small oval foramen pierced obliquely.

Exanvple.

ene thvotepneconacond ses: e ence eee: 10 units.

a SICH) DIDI E ats Manet terreus aehers ae Oe

es si ST TY GR baa UR Soe Online

Breaditheo tae. gees (GuSbal eml ieee eee eer ereee Ons

pea aCIMT CLG LG) 5 itis Maes seus otc Ondine

Length Of Supra-scapulay ccec-cecee pee ceeeces 13-4 ,,

CatyprocepHALus D. & B.

C. gayi D. & B.—Scapula normal in its proportions, but large

in comparison with the other bones of the pectoral eirdle.

Longer than precoracoid.

Tretmarosius Wagl.

T. jelksti Ptrs.—In this species all the bones of the pectoral

girdle are extremely stout in comparison with their length.

Scapula equal to the precoracoid in length, stout. Shaft

squarish ; pars ucromialis and pars glenoidalis both weil de-

veloped, separated proximally by a very small cleft or central

foramen.

Hxample.

Beneth of precoracordl sees. teas eect cree 10 units.

Be scapula (upper border) ............ LO pps

a un eG MleUbbis vended cetera tas comeeee are Ocome

iBreadthyot. 4 1 (distaliemd)cp ay.) esse: Oras.

x oe me) (ae ete serene wee A has

i DONS UCTONUUGL USA e ath Pepe tenet Asti

ie FOG GHA OWIGIS. Seocdsosenngocsneb at ae.

Tene thvot supta-scapulagyey snare. pees eek LES Yo

Kxosta Tsch.

E. bufonia Gir.—Scapula equal to the precoracoid in length,

‘ather slender, but of normal proportions.

* Bones pale green, as in Rappia chlorostea K. Blgr.

OF THE SCAPULA IN THE BATRACHIA. PAM

Hytopes Fitz.

H. raniformis Blgr.—Scapula a little shorter than the pre-

coracoid, but well developed. Pars glenoidalis smaller than pars

acromialis ; dividing cleft short.

H. fleischmanni Boett.—In this species the scapula is subject

to slight individual variation. It may be equal to or a little

longer than the precoracoid.

H., lineatus Schn. and H. martinicensis Tsch.—The scapula is as

in 7, raniformis.

The scapula in Hylodes is somewhat like that in Hyla as

regards the relative proportions of the shaft, pars acromialis, ancl

pars glenoidalis. ‘They differ, however, in that the dividing cleft

is short in Hylodes, forming a very small central foramen instead

of a very large one, as in Hyla.

CrRATOPHRYS Boie.

C. boie Wied.—Scapula greatly developed, once and a half

as long as precoracoid. Distal end of shaft about twice

width of proximal end; pars acromialis and pars glenoidalis

normal; dividing cleft short, forming a very small central

foramen; a small portion of acromion cartilage exposed between

head of precoracoid and end of pars acromialis.

In C. ornata Bell and C. americana D. & B. the scapula is as

in OC. bore.

Ewample.

Qength of precoracoid ..........--....--.-.----- 10 units.

3 OD OU EN Ganangoacougsous seoaoebe Joudodbe 15225

an BWR S| ce) AME obo ce coonep ooo eac Ia eae ae

Breadth Of i (distal ex) ites sie-adascccce yaar

= 55° AGEINCICIIS) See ose conc coksednones 2Roes

Length of supra-scapula .........-......:-2se0ee WUBPM op

LepropactyLus Fitz.

L. pentadactylus L.—Scapula equal to or slightly longer than

precoracoid ; stoutly formed. Shaft broad, half length of pre-

coracoid; pars acromialis well developed, partially separated

from head of precoracoid by acromion cartilage ; pars glenoidalis

well developed, strongly depressed; dividing cleft short and

broad, forming an oval foramen, pierced horizontally, hardly

visible from outer view.

L. ocellatus L.—Scapula as in L. pentadactylus.

Hytoruina Bell.

H. silvatica Bell.—Scapula as in Hyla.

LIMNODYNASTES Fitz.

L. peronii D. & B.—Scapula well developed, longer than prz-

coracoid ; distal end twice the width at the middle. Shaft as

long as precoracoid ; pars acromialis somewhat diamond-shaped,

212 MISS JOAN B. PROCTER ON THE VARIATION

having a /\-shaped upper edge; pars glenoidalis also well

developed ; dividing cleft short, forming a small central foramen.

L. dorsalis Gray.—Scapula as in L. peronia.

L. ornatus Gray.—Scapula more elongated than in L. peronit.

Example.

enteral Ot pprseCONA COIN esr .n eee ear 10 units.

a SCapulay gereegi te \s-ec sare sere sertyales se Las

a % FoI Rae eters Meas PES MEO

Breadth of _ ,, (distaljend)) py. saycreatecnss Daan

Me a (omic) WR GRES ee eengoonab yee EPS

Length of supra-seapula ............ Seb aise ae iteret LD h ec

Cryproris Gthr.

C. brevis Gthr.—Scapula equal in length to the preecoracoid ;

normal in its proportions. Dividing cleft pierced horizontally as

in Megalophrys.

Curroueptis Gthr.

C. australis Gray.—Scapula greatly developed, about once and

three-quarters length of precoracoid; slender in the middle.

Shaft twice as long as broad at distal end, about once and a

quarter length of precoracoid ; pars acromialis well developed,

subtriangular, similar to that of Bactrachopsis melanopyge ;

pars glenoidalis smaller than pars acromialis; dividing eleft

pierced obliquely.

Example.

engthvor preecoracoid eee ge sec cctaees sees 10 units.

x Scaypouiley a ee seeteectas ectite setae steer Lis

ye NARS) cen alee ata ta A a 2 Nar tess

iBreadthiol, ™,, )7M(distalvend) 253 s5. 2 see (hsiinces

a 5) aunt dle). 228s. Seek ee cee Ay et

Mengthvof supra-scapulla 2n...20 0-0. cee secre 12(%) ,,

C.. platycephalus Gthr.—Scapula similar in form to that of

©. australis, but much shorter in proportion.

Example.

Length of [SHERACOUANGONG! yas Lan adnabsoce At ales AU e a LOmumibss

a SCH OUIRY Gas didoonocodacsodananoobdecodce ce alee aes

HELErororus Gray.

H. albopunctatus Gray.—Scapula abnormally developed ; twice

and a half as long as precoracoid; slender in the middle,

greatly expanded at distal end, where it is almost the length of

the precoracoid in width. Shaft once and three-quarters length

of precoracoid; pars acromialis well developed, completely in

contact with crescentic head of precoracoid; pars glenoidalis

also in contact with head of precoracoid, partially separated

OF THE SCAPULA IN THE BATRACHTIA,. Dilley

from end of coracoid by cartilage, longer than pars acronualis ;

dividing cleft pierced horizontally, only visible when viewed

obliquely.

H, pictus Ptrs.—Seapula as in H. albopunctatus.

Example.

Length of precoracoid .............. naeccoruane ete: 10 units.

- Scapular c s.merere Sic NY Hee DO as

a TUL ISH EME GENRE cise sit taoces Weoaee

Breadithy ofa yj (alstalmenc))\nan serene: 8°8 ,,

as say. t (TAGS) Bienes ei sarc IPS i 8

a pars acromialis + pars glenoidalis 88 ,,

ibenethiot suprarscapulaye seen en ere. eis:

Text-figure 10.

Pectoral girdle of Heleioporus pictus, X2; from within.

The abnormal development of the scapula in this genus is

similar to that in Phyllomedusa as regards length. In Phyllo-

medusa, however, the pars glenoidalis is much smaller than the

pars acronialis, which is separated from the head of the przecora-

coid by acromion cartilage; also the dividing cleft is long and

broad, forming a very large central foramen, pierced vertically.

Summary and Conclusion.

These data are sufficient to show that the form of the scapula

is remarkably stable within a genus, and that in many instances

it is so highly characteristic that a glance at this bone alone

suffices to recognize the genus to which the skeleton belongs.

This is the case throughout the Pipide and Discoglosside, and, to

a great extent, in other families. In the most recent families,

however, there are many genera conforming to one pattern of

scapula, which makes the character, in these cases, worthless for

diagnostic purposes.

Of the genera which [ have studied, the greatest specific varia-

tion occurs in Bufo; in Hyla, which is a very large genus, there

is practically none.

Although in many ways the Pelobatid genera resemble Palco-

batratchus, the Aglossa must be regarded as having the most

primitive type of scapula, as in Pipa, Hymenochirus, and Xenopus

the proximal end of the bone is entire. In all other genera*

* Genera of the Arcifera.

214 THE VARIATION OF THE SCAPULA IN THE BATRACHIA.

the pars acromialis and pars glenoidalis are separated from each

other by a cleft, or at least a notch (Bombinator and Pseudis).

This character seems to be of much greater importance than

mere size, and corroborates the conclusion of Wolsterstorff, who

says*: “ Paleobatrachus verbindet im Systeme die AGLossa mit

den ARCIFIRA.”

In spite of generic stability the variation is enormous, par-

ticularly in the length of the shaft. Xenopus, on the one hand,

in which the scapula is absent or represented by the pars

glenoidalis only, ranging to Phyllomedusa, Ceratophrys, and

Heleioporus, in which it is enormously developed, in Heleioporus

being twice and a half the length of the precoracoid.

It is astonishing that such a useful bone as the scapula should

have been overlooked for so long. The following key is based on

this character alone :—

Acabroximaliendjormscapulaxenticempeiec: soneceacnce maceereeee eee LE ED

B. Proximal end of scapula cleft.

I. Length of scapula $ or less than } length of precoracoid. D1scoGLossipZ.

II. Length of scapula more than } length of praecoracoid.

a. A large oval foramen pierced vertically in glenoid

cavity ; pars glenoidalis greatly reduced ......... Hy Lip.

&. Foramen, if pierced vertically in glenoid cavity,

moderate or small; often represented by a cleft

pierced obliquely or horizontally ............... 0c 00 PELOBATIDZ.

BuFONIDS.

CYSTIGNATHID&.

{ am unable to find reliable family characteristics which

differentiate between the Pelobatidw, the Bufonide, and the

Cystignathide.

* Op. cit.

ON MARINE WCOD-BORING ANIMALS. 215

11. Notes on Marine Wood-boring Animals.—II. Crustacea.

By W. T. Cautmayn, D.Sc.

(Submitted for Publication by permission of the Trustees of the British Museum.)

[Received November 17, 1920: Read March 8, 1921. ]

The Crustacea collected for the Committee of the Institution

of Civil Engineers include, besides species already known to bore

into wood, a number of others, the presence of which in the

damaged timber is probably accidental. The possibility, however,

that some of these also may prove to be destructive makes it

desirable to record their names. The actual wood-boring species

in the collection are all well-known, and little of importance is

added to our knowledge of their distribution, but the opportunity

has been taken to confirm, by direct comparison, the suggested

identity of the Indo-Pacific Spheroma terebrans with the Atlantic

S. destructor.

As in the case of the Teredinide, the occurrence of Kuropean

species of wood-boring Crustacea (Limnoria and Chelura) in

Australia and New Zealand has been attributed to introduction

by wooden ships. Itis true that neither of them, so far as I know,

has been recorded as living in ships’ timbers, but their appearance

in widely-separated localities, while distinct species of the same

genera occur at intermediate points (e.g., at Christmas Island,

J‘alman, Ann. Mag. Nat. Hist. (8) v. 1910, p. 181), is suggestive

of some such means of transport.

Order ISOPODA.

Sub-order FLABELLIFERA.

SPH#ROMA TEREBRANS Spence Bate.

S. terebrans Spence Bate, Ann. Mag. Nat. Hist. (3) xvii. 1866,

p- 28, pl. ii. fig. 5; Stebbing, Spolia Zeylanica, ii. 1904, p. 16,

pl. iv.; Hansen, Q. J. Micr. Sci. xlix. 1905, p. 116; Stebbing,

Ann. 8. Afric. Mus. vi. 1908, p. 49; Chilton, N.Z. Journ. Sci.

Technol. ii. 1919, p. 12; Calman, Marine Boring Animals, Brit.

Mus. (Nat. Hist.) Economic Ser. No. 10, 1919, p. 21, fig. 113 id.,

Committee on Structures in Sea-water, Inst. Civ. Engineers, 1st

Rep. 1920, p. 70, pl. i. fig. 5; Barnard, Ann. 8. Afric. Mus. xvii.

1920, p. 358.

S. vastator Spence Bate, Ann. Mag. Nat. Hist. (3) xvii. 1866,

p- 28, pl. i. fig. 4.

S. destructor Richardson, Proc. Biol Soc. Washington, xi. 1897,

p- 105, text-figs. ; 7d., Amer. Nat. xxxiv. 1900, p. 223 ; ¢d., Proce.

U.S. Nat. Mus. xxiii. 1901, p. 534; id., Bull. U.S. Nat. Mus,

liv. 1905, p. 282, text-figs.

Locality— Brisbane, Queensland. Specimens forwarded by

Mr. E. A. Cullen, Engineer for Harbours and Rivers. From

216 DR. W. 'T. CALMAN ON

Jetties, 5 miles from river entrance in Moreton Bay, in Swamp

Mahogany (Tristania suaveolens) and Ironbark (Hucalyptus

paniculata). Many specimens.

Specimens are in the Museum collection from Wyong River,

New South Wales, and Brisbane (Prof. Chilton), Travancore

(Crivandrum Mus.), Isipingo River, Natal (Durban Mus.), and

St. John’s River, Palatka, Florida (U.S. Nat. Mus., syntypes of

S. destructor Richardson).

Remarks.—This species has already been recorded from Bris-

bane by Prof. Chilton.

Barnard has recently adduced further evidence in favour of

Stebbing’s view, disputed by Miss Richardson, that the Indo-

Pacific species is identical with S. destructor Richardson, from

Florida. No one, however, has hitherto been able to base this

identification on a direct comparison of specimens. By the

courtesy of the authorities of the U.S. National Museum, who

have presented syntypes of S. destructor to the British Museum

(Nat. Hist.), I have been able to compare these with specimens

from all the other localities mentioned above. I have failed to

find any difference that can be regarded as specific. Both Stebbing

and Barnard have indicated a certain amount of variation in the

tubercles and ridges of the dorsal surface, and this is shown more

conspicuously by the specimens now examined. The specimens

from Brisbane grow to a greater size (11°5 mm. total length)

than the syntypes, and have the tubercles on the sixth and

seventh pereon-segments very low ; the submedian tubercles on

the telsonic segment are distinctly elongated in an antero-

posterior direction, and the granulation of the telsonic segment

is less close than in the syntypes. The Travancore specimens

agree better with the syntypes in their smaller size, generally

more prominent tubercles, and closer granulation of the telson,

but they differ in having the submedian tubercles of the telson

distinctly smaller than those of the lateral pair. The Natal

specimen bears a close resemblance to those from Brisbane. I

have only been able to observe one character in which the Indo-

Pacific specimens agree with one another, while differing from

the syntypes. Seven specimens from Wyong River, Brisbane,

Travancore, and Natal were found, on dissection, to agree with

Stebbing’s account in having three large sete and a small one on

the inner plate of the maxillula. Miss Richardson found in her

specimens “‘ five strong plumose sete and occasionally a sixth one

that is feeble.” The single syntype that I have dissected has five

large setee and a small one on the maxillula of one side, but on

its fellow there are six sete of equal size. In none of the other

characters mentioned by Miss Richardson is there any constant

difference to be detected *.

* §. peruvianwm Richardson (Proc. U.S. Nat. Mus. xxxvini. 1910, p. 81, text-figs.

3 and 4), found boring in the roots and stems of mangroves on the coast of Peru, is

very similar to the species here discussed, but the broadly-rounded terminal segment,

the longer antennules and antennie, and other characters indicated by Miss Richard-

sen may justify its separation. I have seen no specimens.

MARINE WOOD-BORING ANIMALS. 217

Hansen has already stated that this species is to be included in

the genus Spheeroma as restricted by him. Ovigerous females

in the Brisbane collection agree with his definition of the genus

in having large overlapping oostegites, while the eggs (and

embryos) are contained in internal pouches opening by four

pairs of large slits on the sternal surface, as in S. rugicauda and

S. serratune.

A large number (well over a score) of males have been ex-

amined, which, from their size (up to 9°5 mm. in length), from

the fact that they were found in the same gathering with

ovigerous females and from the presence on the last thoracic

sternite of a pair of penes, would ordinarily be regarded as

adults. Two individuals were dissected and found to have the

vasa deferentia distended with bundles of filiform spermatozoa.

Nevertheless, in no case was there a trace of an appendix

masculinag or even of a marginal thickening on the second

pleopods *. Hansen states (op. cit. p. 88) that an appendix

masculine is present in adult males of all the genera of Sphero-

mide except Dynamene and Ancinella, but he notes that in the

subfamily Spheromine it “does not appear before the animals are

nearly full-grown,” while the penes are present at a much earlier

stage. His account of Cymodoce pilosa (op. cit. p. 89) suggests

caution in deciding as to the sexual maturity even of large

individuals. Nevertheless, the conclusion seems to be justified

that the appendia masculina is not developed in Spheroma tere-

brans, which in this respect forms an exception among the

species of the genus.

Crustacea associated with 8. terebrans.—Stebbing has recorded

the occurrence of the little Asellotan ats pubescens in association

with S. éerebrans at Ceylon. He had previously given an extended

description, with figures, from specimens taken on Eaospheroma

gigas at the Falkland Islands (Proc. Zool. Soc. 1900, p. 549,

pl. xxxviii.). The collection of S. terebrans from Brisbane includes

numerous specimens,and that from Natala solitary young specimen

of what is probably the same species. Some of the Brisbane speci-

mens carrying eggs do not exceed 2:24 mm. in length (Stebbing’s

Falkland specimens reached 2°5 mm.). They have not more

than 13-14 segments in the flagellum of the antenna, the uropods

are nearly half as long as the dbdomen, and the exopod of the

uropods is much longer than the peduncle and definitely longer

than the endopod.

Specimens found on Lwospheroma gigas from the Auckland

Islands differ from these and agree with Stebbing’s Falkland

Island specimens in having more numerous segments in the

flagellum of the antenna (22, while Stebbing records “ attaining

to 25”), the uropods not more than one-third as long as the

telsonic segment, and the exopod of the uropods equal to the

* Barnard notes the absence of the appendix masculina in the single male (9 mm,

in length) examined by him.

218 DR. W. T. CALMAN ON

peduncle and shorter than the endopod. The distal segments of

the antennules are a good deal less slender than in the Brisbane

specimens.

Tam unable to perceive any other differences of importance,

and, while it might be considered that those enumerated would

justify giving a new varietal or even specific name to the Brisbane

specimens, | prefer to await further evidence from other localities

regarding the range of variation in this widely-distributed species.

Among the Brisbane specimens of Spheroma terebrans were

numbers of another Spheromid, apparently belonging to the

genus Hxospheroma, but differing from all the described species

of that genus. There seems to be no reason for suspecting this

species of complicity in the destruction of the timber. The

body is more depressed than one would expect to find it in a

burrowing animal, and the mouth-parts are much less prominent

than they are in Spheroma terebrans.

Somewhat more suspicious is the case of an Isopod of which

a collection was sent from H.M. Dockyard, Simon’s own, by

Lieut. L. H. A. Shadwell, R.N.V.R. The specimens were from

‘‘Greenheart camber piles, W. yard,” and were labelled as

Spheroma. They proved to belong to the species Parisocladus

stimpsoni (Heller), agreeing closely with Barnard’s description

and figures (Ann. 8. Afric. Mus. x. 1914, p. 399, pl. xxxil. G),

and being of the smaller size (male about 7°5 mm. long) which he

mentions as characteristic of specimens from the east side of the

Cape Peninsula. Here, again, the general form of the body and

the disposition of the mouth-parts are not such as to suggest a

boring habit. On the other hand, the timber in question must

have been attacked by some boring animal, and if Spheroma

terebrans had been present in numbers sufficient to cause visible

damage it could hardly have been entirely overlooked by the

collector.

The only other animals in this gathering were two specimens

of Cirolana sulcata. Like its congeners, this species is no doubt

predatory and not at all likely to attack wood.

LIMNORIA LIGNORUM (Rathke).

Cymothoa lignorum Rathke, Skrivt. Naturh, Selsk. Kjpbenhavn,

ve(L) 1799 ps LOI, jolyauetie es asd:

Limnoria terebrans Leach, Edinburgh Encye. vil. 1814, p. 433 ;

Coldstream, Edinburgh New Philos. Journ. xvi. 1834, p. 316,

all, Wale

Limnoria lignorum White, Pop. Hist. Brit. Crust. 1857, p. 227,

pl. xii. fig. 5; Spence Bate and Westwood, Brit. Sessile-eyed

Crust. ii. 1868, p. 351, figs.; Harger, Rep. U.S. Comm. Fish. vi.

1880, p. 373, pl. ix. figs. 55-57; Hoek, Verh. K. Akad. Wet.

Amsterdam (Tweede Sectie), i. No. 6, 1893, 103 pp., 7 pls.; Sars,

Crust. Norway, ii. 1897, p. 76, pl. xxxi.; Chilton, Ann. Mag. Nat.

Hist. (8) xiii. 1914, pp. 380 & 448; id., N.Z. Journ. Sci. Technol.

ii. 1919, p. 3, figs.; Calman, Marine Boring Animals, Brit. Mus.

MARINE WOOD-BORING ANIMALS. 219

(Nat. Hist.) Economic Ser. No. 10, 1919, p. 17; id., Committee

on Structures in Sea-water, Inst. Civ. Engineers, lst Rep. 1920,

p- 68, pl.i. fig. 4.

Localities. — Leith. Specimens forwarded by J. Dalgleish

Easton, Deputy Superintendent, Port of Leith. From Jetties,

in White Pine and Pitch Pine. Many specimens.

Southampton. Specimens forwarded by F. E. Wentworth

Shields, Docks Engineer, L.& S8.W. Railway. From Outer Dock,

in American Elm. Few specimens.

Simon’s Town, Cape of Good Hope. Specimens forwarded by

Lieut. L. H. A. Shadwell, R.N.V.R., Officer in charge of works,

H.M. Dockyard. From creosoted Danzig timber, No. 3 slip.

Few specimens.

Auckland, New Zealand. Specimens forwarded by Mr. W. H.

Hamer, Engineer to the Auckland Harbour Board. From

Totara timber. Many specimens.

Remarks.—As the European LZ. lignorwm has already been

recorded from 8S. Africa (Port Hlizabeth) and Auckland, the

records given above do not extend its known range.

In stating that “in Kurope... Zimnoria lignorwm seems to

be constantly associated with Chelura terebrans” Chilton (I. ce.

1919, p. 6) is repeating a statement frequently made, but cer-

tainly erroneous. At Leith, and elsewhere on the east coast of

Scotland and England, while Limnoria is abundant, there seems

to be no record of Chelura.

Order AMPHIPODA.

Suborder GAMMARIDEA.

CHELURA TEREBRANS Philippi.

C. terebrans Philippi, Arch. Naturgesch. li. 1839, p. 120, pl. iu.

fic. 5; Stebbing, Das Tierreich, xxi., Gammar idea, 1906, p. 693

(with references) ; Chilton, Ann, Mag. ee Hast. (8) xili. 1914,

p. 380; 7., N.Z. Journ. Sel Mechnol! LORD. Osis. 5 Cal-

man, Marine Boring Animals, Brit. Woe ‘ON at. Hist.) Economie

Ser. No. 10, 1919, p. 20; zd., Committee on Structures in Sea-

water, Inst. Civ. Engineers, Ist Rep. 1920, p. 71, pl. 1. fig. 6.

Localities—Southampton, Specimens forwarded by F. EH.

Wentworth Shields, Docks Engineer, L. & S.W. Railway. From

Outer Dock, in American Elm. Few specimens.

Simon’s Town, Cape of Good Hope. Specimens forwarded by

Lieut. L. H. A. Shadwell, R.N.V.R., Officer in charge of works,

H.M. Dockyard. From creosoted Danzig timber, No. 3 slip.

Many specimens.

Remarks.—This species has been recorded by Chilton (0. ¢.) as

destructive in Auckland Harbour, but specimens forwarded from

Auckland as Chelura proved to be Corophiwm contractum G. M

Thomson, a species, no doubt harmless, also recorded by Chilton.

Proc. Zoot, Soc,— 1921, No, XVI, 16

220 ON MARINE WOOD-BORING ANIMALS.

T learn from Mr. Hamer, however, that this is merely due to

a transposition of labels, the Chelwra, whose identity with the

European species is vouched for by Prof. Chilton, being abundant

and well known at Auckland.

It appears doubtful whether there is any trustworthy record of

Ohelura occurring apart from Limnoria. The gathering from

Simon’s Town seemed at first sight to consist exclusively of

Chelura, but on closer examination 13 specimens of Limnoria

were discovered among more than 300 of the other genus, If

these numbers are at all representative of the proportion in

which the two species were living in the wood, the great pre-

ponderance of Chelura is very remarkable. The only previous

record of Chelura from South Africa appears to be that of

Hammersley-Heenan, who found it at Port Hlizabeth in 1893

(Trans. 8. African Phil. Soe. v. p. 316), and it is noteworthy that

no mention is made of its being accompanied there by Limmnoria.,

COLOUR-PRODUCTION IN THE FEATHERS OF BIRDS. 22:1

12. Colour-production in relation to the Coloured Feathers

of Birds. By A. Matuocg, F.R.S., F.Z.8.

[Received February 7, 1921: Read February 22, 1921.]

(Text-figures 1-4.)

The brilliant colouring of many birds and the forms of the

feathers on which this colouring appears, present many points

of interest both in regard to the means by which the eolour-effect

is produced, and to the processes of selection which have led to

their development.

In the present note I touch only on the physical side of the

problems, and hope to give some idea of the many ways in which

colour may be produced by the action of white or composite light

on matter, and of the effect of the forms of various feathers in

modifying the appearances which the more intimate structure of

the material produces on the incident light.

If any object appears coloured when viewed in white light, it

shows that the matter of which it is composed exercises some

selective action on the composite light falling on it, absorbing or

transmitting certain colours and reflecting or scattering the

remainder. This selection may be of two kinds: namely, a

relation between the periods of light of various wave-lengths and

the molecular periods of the matter on which it falls, or on a

relation between some distance or spacing in the structure of the

substance and the wave-lengths themselves.

The first of these relations includes all pigment colours, and

the second those which are known as interference effects.

In the following table I have given a list of all the types of

colour-production with which I am acquainted, and I believe

that these will cover every known case, although the actual

dynamics of'a large proportion are very doubtful, involving as

they do the dynamics and constitution of the molecule.

Colour may be produced by :—

Te:

A relation between the periods of waves of light and the

molecular periods of the colour-producing matter :

Haeamples.

Dispersion, f | Prismatic Colours, Rainbows, etc.

ee He Ch mare elegy Powdered glass or other colourless trans-

wave-length. | parent material immersed in a fluid of

| the same mean refraction index but

| . . .

| different dispersive power.

All dyes and pigments, which may be} The greater number of coloured solids

transparent and scatter or transmit} and fluids.

waves of certain periods and absorb

the remainder. Or:

Reflect certain periods and transmit the | Aniline colours in crystal or dry films,

remainder, and many other crystals.

| 16*

Die, MR. A. MALLOCK ON COLOUR-PRODUCTION

Are opaque and reflect certain periods

Opaque pigments such as lead chromate,

and absorb the rest.

ete.

Complete opacity may arise either from

the absorption of all wave periods or

by complete reflection.

Lamp-black.

Silver and most white metals.

Fluorescence and Phosphorescence.

A relation between the wave-lengths of light and the

structural dimensions of the matter which appears coloured :

Examples.

Reflection or transmission from or Colours of thin plates.

through a striated or laminated | Mother-of-pearl.

structure. Lipmann films.

| Diffraction gratings.

Pitted surfaces with pits of uniform

| depth.

Scattering or transmission of light by

particles of sizes comparable with the

Red of sunset.

Light, seen through vapours or emul-

_wave-length, but irregularly distri-| sions.

buted. Blue sky and the colours of the sea and

rivers.

Glass coloured with gold.

Supernumerary rainbows.

The colours of polarization in most cases depend both on

molecular structure and on the linear dimensions (measured

along the paths of the rays) of the bodies which exhibit them.

With regard to the first class, little is known concerning the

intimate structure of matter. It is a fact, however, that lght-

waves travel more slowly in solids and liquids than i vacuo, and

that the velocity is in some unknown way dependent on the wave-

length.

In most cases this is best represented by assuming that the

ether is, as it were, loaded by matter, while in some others

it would appear that the elasticity of the ether is affected. The

difference may be exemplified by a stretched string which has a

definite period settled by its length, tension, and mass. If the

string is loaded, the period is increased by an amount depending

on the added load, but an equal increase in period may be pro-

duced by relaxing the tension. If the shape of a transparent

body is such that the direction of light after passing through it

depends on the velocity of the waves in the interior, as for

instance in a prism or sphere, the emergent light will appear

differently coloured in different directions.

In the case of pigments, it is most probable that the individual

molecules have a natural period identical with that of some of the

IN THE FEATHERS OF BIRDS. 223

periods of the incident light, and that the colours which they

reflect or transmit are due to resonance.

The actual dimensions, forms, and rigidities of molecules are

unknown, but their diameters are apparently of the order of a

ten-millionth of a centimetre; and one may inquire what the

longest natural period of a sphere 1/10,000,000 cm. diameter

would be if it were as rigid as steel, or, which comes to the same

thing, what is its least natural frequency. Without going into

the details of the calculation, it may be stated that this least

natural frequency is somewhere about 5x10” vibrations per

second. All the other natural modes of vibration would have

higher frequencies, and there is good reason to suppose that the

rigidity of molecules far exceeds the rigidity of the matter formed

by their aggregation. ‘The frequency of yellow light is 5x10"

vibrations per second. Thus itseems that the natural frequencies

of molecules and of visible light-waves are at any rate of the

same order.

All these pigmentary and dispersion colours depend on the

constitution of the molecule itself. In the second class it is the

relation of the size of particles, or on their disposition in space

as compared with the wave-length which determines the selective

influence of the matter on white or composite light.

The origin of the colouring presented to view as the result of

selective action of the structure on wave-length can be fairly well

determined by the following tests :—

(a) Mechanical compression or extension.

(6) Immersion in various fluids.

Of these, the compression test is the most decisive; for, if the

mechanical distortion of the structure changes or obliterates the

colour, it may be assumed that the colour itself depends on some

special arrangement of the parts, and not on the molecular pro-

perties of the material of which it is built up.

There may be some apparent exceptions, as for instance when

a material transmits one colour with less loss than another, so

that the predominating colour is dependent on the thickness of

the layer through which the light travels (e.g. manganese glass

or a solution of chlorophyll).

In the circumstances, however, in which this test is applied

to organic structures, such as feathers, these exceptions will

hardly operate.

The greater part of the colours of feathers have their origin in

pigments of the nature of which little is known. Except in

one instance, no solvent has been found for them, and the

pigments themselves vary much in physical properties. Some

are nearly opaque, while others are transparent and transmit the

complementary colour. Many, again, polarize the incident light,

and this is especially noticeable with transparent yellows. ‘These

224 MR. A. MALLOCK ON COLOUR-PRODUCTION

form rather brilliant objects when viewed between crossed .

nicols.

But by far the most brilliant colouring of birds has its origin

in interference, that is to some periodic structure in the sub-

stance of the feather, where the spacing of the parts is a multiple

of the half wave-lengths of the light they reflect. Such is the

case among humming-birds, sun- birds, peacocks, birds of paradise,

and ducks, to mention only a few instances of what are spoken

of as “‘ metallic” colouring.

All these colours disappear when subjected to pressure, and in

all cases the colour-producing substance is confined to a very thin

layer overlying an iatensely opaque black or brown substratum.

The general effect when viewed from a distance depends to a

great extent on the form of the surfaces on which the colouring

layer is disposed. If these surfaces are planes, the relative

positions of the eye and source of light with regard to the feather

has to be rather carefully adjusted, in order that any colour may

be visible.

When, however, the surfaces are rounded, the range of incidence

is much extended, and from almost any point of view some

colour appears, although the intensity is lessened, just as a tray

of small glass beads will scatter sunlight in all directions

although the intensity in any one direction is much less than

what would be produced by a plane mirror adjusted to. reflect

lights in that direction only.

The examples chosen all show distinctive structure peculiar to

the orders to which the examples belong.

A feather may be described as consisting of a stem, branches,

and leaves (text-fig. 1) (named by zoologists respectively vachis,

rami or barbs, and barbules, the latter. sometimes as carrying

barbicels). It is in the modifications of the leaves that the

distinctive features are found.

I will notice these in order.

In all the ornamental feathers of Humming-Birds the branches,

but especially those of the gorget, are so bent that their ends

are parallel and the colour-bearing surface is on the leaves. A

cross-section of the branches and leaves forms a succession of

hollows in text-fig. 2 e.

The section of the leaves themselves is something like the

numeral 7 (text-fig. 4c), and the colour-producing material les

in a thin layer on the upper surface of the leaves, and is nearly a

plane, so inclined that the normal makes an angle of about 60°

with the branch, but lying in a plane parallel to the latter. (This

angle varies in different species.)

To keep the leaves in this position, each barb terminates in a

curved plate (text-fig. 4d), which interlocks with the similar plates

of the two or three leaves in advance.

Each branch ends in a long bristle (fig. 2 6), which, when the

plumage is in good order, lies in the trough ef the valley formed

IN THE FEATHERS OF BIRDS. . 225

by the leaves and branches of the succeeding feather. This

bristle is only found in the head- and gorget-feathers. The body-

feathers, where coloured, end as shown in text-fig. 2c.

From these arrangements it happens that the best «display of

colour occurs when the body of the bird is in the nearly vertical

position it assumes when hovering.

In the Sun-Bird each leaf has a curious rasp-like shape, and

the upper colour-producing surface is in the form of 3 or 4

convex plates, in each of which the virtual image of the sun or

other source of light appears as a coloured spot.

The chief peculiarities of the metallic feathers of Birds of

Paradise are that the leaves are only developed on one side of the

branch, and are so disposed with reference to it as to be parallel to

the stem (text-fig. 36). The upper surface consists of more or less

rounded lobes, an nd the cross-section is shown in text-fig.3.¢. This,

so far as [ have observed, is the rule in all the metallic feathers,

whether on the head, wings, or tail.

In the Rifle-Bird, which is a near connection of the Birds of

Paradise, the leaves are only developed very slightly on that side

of the branch on which, in the true Birds of Paradise, they

are absent.

In the Peacock and all other pheasant-like birds, the leaves

ave of the form shown in text-fig. 4a, 6. The cross-section of the

leaves is comma-shaped and the whole structure is transversely

corrugated. The colour layer lies on the upper surface of the

“dot” of the comma, which thus presents a series of rounded

knobs to the light, each giving rise to a spot of colour.

In the head-feathers of Ducks the colour is developed on the

leaves on both sides of the branch, but in the speculum, on one

side only, the uncoloured leaves lying below the coloured part of

the adjacent branch and serving to lock the two in position.

As before stated, all these feather colours disappear when the

structure is compressed. For this test I place the feather

between a quartz plate and a plano-econvex lens of the same

material, of a foot radius, these being mounted to fit on the stage

of a microscope. The only difficulty in applying this test is

to separate a suitable part of the coloured material on which to

operate. The result is a conclusive proof that the colours are

not due to any form of ene and strong evidence that they are

due to interference, ost feathers are extremely impermeable

to fluids, but in certain cases (the Peacock for instance), when

immersion takes place, the colour changes at once to one of a

longer wave-length: blue becomes green, green yellow, and so

on. Where this happens it is evidence either that the feather is

to a certain extent permeable by the fluid used, or, more probably,

that the colour-production depends on some quality or grain of

the outside surface.

The greater number of the metallic feathers which I have

experimented with show no change on immersion in any ordinary

MR. A. MALLOCK ON COLOUR-PRODUCTION

Text-figure 1.*

* For description of the figures see next page.

Fig. 1.

Rig. 2a.

2b.

2c.

Fig. 4a.

IN THE FEATHERS OF BIRDS. OAT

Text-figure 4,

(0 Fe one.

4c.

Diagrammatic drawing of a Feather, showing Stem, Branches, and Leaves.

Feather from the gorget of Humming-Bird (X20).

Termination of single branch of same feather (X30).

a 6 of coloured body-feathers from the same

bird (x30).

. Termination of gorget-feather (130), showing the leaves and their prolon-

gations ( p-p), which serve to keep the former regularly spaced, and the

planes of their upper surface inclined at a constant angle to the axis of

the branch.

. Cross-section of gorget-reather, parallel to the line X X of 2a (X20).

. Cross-section, parallel to X X of 2d, of single branch of the same feather

(X130). Note the sections of the prolongations of the leaves at p p.

Longitudinal section of the same, parallel to Y Y of 2d.

. Feather from the wing of King Bird-of-Paradise by reflected light (X40).

. Part of a single branch of same feather (X130), showing leaves developed

on one side of the branch only.

. Cross-section of leaves, parallel to X X in 36 (220).

. Longitudinal section of one of the leaves of same feather.

Branches of blue feather from Peacock’s neck by transmitted light (30).

. The same by reflected licht (X40).

. Cross-section of single branch of same (X 1380).

. Part of a single leaf of same, seen in perspective.

228 COLOUR-PRODUCTION IN THE FEATHERS OF BIRDS.

fluid (alcohol, xylol, chloroform, oil, etc.), but strong acids in

time cause the colours to change towards the red end of the

spectrum.

The most penetrative fluid which I have tried is the solution

of iodide of mercury in iodide of potassium. ‘This, when concen-

trated, rapidly destroys the feather substance, but in dilute

solution merely penetrates into the interior. The gorget-feathers

of the Humming-Bird (from Costa Rica) reflect a brilliant lilac,

i.e. a mixture of red and blue, but after a few hours’ immersion

in the iodide solution, the red disappears and the blue changes to

a very bright green.

All these metallic colours shift towards the blue as the angle

of incidence of the light increases, as do the ordinary colours of

thin plates ; but this is not a proof that both have the same origin,

for many of the aniline colours when in thin dry films show

somewhat similar changes depending on the angle of incidence.

Michelson in America has compared the metallic colour of

some beetles with those of the anilines, and has given reasons

(connected with similarity of the polarization of light reflected by

both) for believing that the origin of the colours in the two cases

is of the same kind.

The pressure test, however, seems to make this conclusion

invalid.

The colours seem to me to be more allied to those of Lipmann

films, in which layers of reduced silver are spaced at half wave

intervals, and in the case of metallic feathers I believe that one

or two layers of optically dense material are the sources of

interference.

Although half wave-lengths can be readily resolved by high-

power microscopic objectives, it is almost impossible to cut sections

thin enough (viz. less than *00002 in.) to use with such powers.

At least I have cut many hundred sections, but although in some

cases a laminated structure seemed to be present, this was due to

a diffraction effect. as was evident from the changes in the dimen-

sions of position of the apparent lamina which occurred with the

change of focal adjustment.

In the case of the Lipmann films, the layers of reduced silver

are readily seen if the sections are expanded by wetting, though

I have not been able to resolve them satisfactorily when dry.

ON FOSSIL BIRD-REMAINS. 229

13. Fossil Bird-remains collected by Dr. Forsyth Major in

Sardinia, Corsica, and Greece. By E. T. Newton,

BUR S2UE. GIS EY ZeS:

[Received February 8, 1921: Read March 22, 1921. |

In the year 1891 Mr. R. Lydekker * gave an account of some

Pleistocene Bird-remains which had been collected by Dr. Forsyth

Major in Sardinia and Corsica. These remains were from three

localities—(1) Tavolara, an island on the N.E. coast of Sardinia ;

(2) Monte San Giovani, in the 8.W. of the same island; and

from (3) Toga, near Bastia, Corsica. From these localities some

score of species were recognized more or less definitely as follows:—

(1) Tavolara.

Coracias cf. abyssinicus.

Corvus corone.

Coccothraustes vulgaris.

Fringilla ccelebs.

Serinus hortulanus ¢

Columba cf. livia.

Coturnix communis.

Puftinus ef. fuliginosus, | Most

> . ef. anglorum. common.

» alliedto selbrauitneRTe|

(2) Monte San Giovani.

-Bubo ef. cinerascens.

Milvus ef. ictinus.

Aquila sp.

Vultur ef. monachus.

Pyrrhula europea.

Alauda arborea @

Turdus musicus.

Hirundo rustica.

Putiinus cf. fuliginosus.

(3) Toga, near Bastia.

Turdus merula.

Columba cf. livia.

Aquila sp.

Some years later Dr. Forsyth Major made further explorations

of the Pleistocene deposits in the Mediterranean islands, and

obtained a large number of Mammalian and other vertebrate re-

mains, not only from the above-named places, but also from several

new localities. Among these were many bones of birds, which

* Proc. Zool. Soc. 1891, p. 467.

230 MR. E. T. NEWTON ON

Dr. Forsyth Major in the year 1908 (just before he left England)

submitted to the present writer with a view to their identification.

This agreeable task was accomplished some years ago, and the

results have been awaiting Dr. Forsyth Major's description of

the Mammalia; but, as he is still absent from England, it seems

desirable that a_ brief account of the Avian remains should be

published.

Some of these are from the same localities as those described

by Mr. R. Lydekker, and, as they include some additional

interesting species, these will be first considered.

(1) From Tavolara, N.E. Sardinia, there are in the present

series several vertebrae belonging to Puffinus, a genus which

-Mr. Lydekker found so abundant; but besides these I have

recognized bones of the Common Swift (Cypselus apus Linn.)

and Alpine Swift (C. melba Linn.), also a small Crake (Porzana

parva? Scop.) and the Storm-Petrel (Procellaria pelagica Linn.).

(2) From the second locality, Monte San Giovani, there is only

one Passerine humerus.

(3) From Toga, near Bastia, [ have a nearly perfect humerus

of a Chough (Pyrrhocorax graculus Linn.), which is distinctly

larger than the humerus of the Alpine Chough.

(4) Grotta di Funtanedu.

Specimens from this and the following localities were not

included in the series sent to Mr. R. Lydekker and alluded to

above.

The Grotta di Funtanedu is near Bastia, N.E. Corsica, and is

situated 400 metres above sea-level. It has yielded a good

number of birds’ bones, and some 34 species have been identified ;

but each of these is represented by only a few bones.

As no account of Avian fossil remains from this cave has

hitherto been published, it will be well to give a detailed list of

the species recognized :—

PASSERES.

Missel-Thrush. Zurdus viscivorus Linn.

Song-Thrush (or Redwing). 7. musicus? Linn.

Fieldfare. 7’. pilaris Linn.

Blackbird. 7’. merula Linn.

*Large Thrush. Turdus sp. (? 7. varius Pallas).

Wheatear. Saaicola ananthe Linn.

Whinchat. Pratincola rubetra Linn.

Redstart. Ruticilla phenicurus Linn.

Robin. Lrithacus rubecula Linn.

Willow-Wren. Phylloscopus trochilus Linn.

*Dipper? Cinclus aquaticus? Bechst.

Wagtail. MWotacilla lugubris ? Temm.

Titlark. Anthus pratensis Linn.

Rock-Pipit 2? A. obscurus ? Lath.

FOSSIL BIRD-REMAINS. Dips

Linnet. Acanthis cannabina Linn.

Chaftinch. ringilla calebs Linn.

Starling? Sturnus ?

Chough ? Pyrrhocorax graculus ? Linn.

Jay. Garrulus glandarius Linn.

Jackdaw. Corvus monedula Linn.

Skylark? <Alauda arvensis ? Linn.

*Shore-Lark? Otocorys alpestris ? Linn.

PICARIA.

Swift. Cypselus apus Linn.

STRIGES.

Short-eared Owl? Asio accipitrinus ? Pall.

ACCIPITRES,

Kagle. Aquila sp.

Sparrow-Hawk. <Accipiter nisus Linn.

Kestrel. Falco tinnunculus Linn.

ANSERES.

Wild Duck? Anas boscas? Linn.

CoLUuMB&.

Wood-Pigeon. Columba palumbus Linn.

GALLINA.

Red-legged Partridge. Caccabis rufa Linn.

Quail. Coturnix communis Bonnat.

FULICARIA.

Crake (small form). Porzana parva ? Scop.

LIMIcoLa.

Redshanks? Yotanus calidris? Linn.

GAVIA.

Tern. Sterna fluviatilis? Naum.

(5) Grotta di Brietta.

This cave is situated in the same locality of N.E. Corsica as

Funtanedu, but about 500 metres lower down—that is, about

100 metres above sea-level. From this I have identified bones of

Thrush, Blackbird, Wheatear, and *Magpie (Pica rustica Scop.),

the last-named species being an addition, to the fauna of these

islands.

(6) Gradicchia Cave.

This is another cave near Bastia, which may possibly be pre-

historic ; but probably even more modern. From it are

recognized bones of Blackbird, Crow (or Rook), Fowl, and (it may

be) Pheasant,

(7) Margine Cave.

The Margine Cave is situated near Nouga, Corsica, and from

it remains of only two birds have been identified—the Chough

(Pyrrhocorax graculus L.) and a Pigeon,

232 ON FOSSIL BIRD-REMAINS.

(8) Grotta del Capo.

This cave is at Sagro, near Sisio, Cape Corso, N. Corsica, and

among the fossil remains Thrush and Blackbird are the only

Avian species recognized.

(9) Monte Santa Cave.

Monte Santa is near Guisba in N.W. Sardinia, and from this

cave we have representatives of two birds—the Red-legged

Partridge (Caccabis rufa Linn.) and the Woodcock (Scolopax

rusticula Linn.).

10. Nesakia, Isle of Cerigo, Greece.

The remains of four species of birds are recognized among

these Grecian island fossils—namely, Blackbird, Rook (?), Pigeon,

and Barn-Owl (Strix flammea Linn.).

The species mentioned in these notes are without exception

living forms; but those marked with a star (*) have not hitherto

been “recorded from these Mediterranean islands tf. Dr. Forsyth

Major appears to have no doubt as to the deposits at all the

above localities, excepting No. 6, being of Pleistocene age, and

doubtless the Mammalian remains which he has collected will

justify this opinion; but the Birds, being only referable to

modern species, offer no sure grounds for a judgment in this

respect. As remarked by Mr. R. Lydekker =, many modern

species of birds, especially among Passerine forms, are distinguished

by their plumage rather than by any osteological differences, and

possibly these fossil bones, although lke fiose of modern species,

may have been differently clothed in Pleistocene times, and in

this way have differed from their descendants of the pre esent day.

Since these notes were written, I have had the pleasure of

seeing the valuable and detailed work of M. M. Boule on the

Vertebrate fossils from the Grimaldi Caves §, in which many

Birds’ bones are beautifully figured and carefully described. As

might have been anticipated, ‘these caves of the mainland have

yielded a very similar Avian fauna to those of the Mediterranean

Islands.

Dr. Forsyth Major’s specimens will be deposited. at the

Natural History Museum, South Kemsington.

‘+ See John Whitehead, “Ornithological Notes from Corsica,’ Ibis, 1885,

pp. 24-48 and plate.

t Proc. Zool. Soc. 1891, p. 468.

§ ‘Les Grottes de Grimaldi’ (Monaco, 1919), Tome i. Fasciculus iv. p. 299.

P.Z.S. 1921. Andrewes. PI. I.

3 * Fa Ss VITTY & SEABORNE LTD.

JER. C. Knight, del. LONDON,

ORIENTAL SPECIES OF CALLISTOMIMUS.

THE ORIENTAL SPECIES OF CALLISTOMIMUS. 233

14. The Oriental Species of the Genus Callistomimus

(Coleoptera, Carabide). By H. EH. ANpREwsEs*.

[Received December 13, 1920: Read March 22, 1921.]

(Plate I.)

Having some new species to describe in this genus, I thought

it would be desirable, if it were alsc possible, to deal in one

paper with all the species at present known. I have been

fortunate in securing the loan of a number of types, and in

recelving permission to see others; in this connexion [ have to

offer my thanks to Dr. Gahan of the British Museum, Dr. Gestro

of Genoa, Dr. W. Lundbeck of Copenhagen, Mr. René Oberthiir,

Mr. EK. Fleutiaux, and Mr. Guy Babault. To Mr. Severin of

Brussels my thanks are not less due, for at my request he made

an exhaustive search in the Brussels Museum—unfortunately with-

out result—for the types of the two species described by Putzeys.

In 1872 Chaudoir described the genus Callistomimus (Bull.

Mose. i. p. 382), which he differentiated from Callistus by the

edentate mentum and some other characters. In the following

year Bates described a new genus from China under the name of

Pristomacherus (Trans. Ent. Soc. Lond. p. 323) for an insect

(P. messit), which was widely different, especially in the form of

the prothorax, from any species of the group then known. In

describing a number of new species in 1892 (Ann. Mus. Civ.

Gen. pp. 303-7}, Bates discusses the relationship of the two genera,

remarking in conclusion ‘‘so that there remains only the

prolonged hind angles of the thorax, separated from the base by

a deep sinuation, to distinguish Pristomacherus.”

In examining various species, including some hitherto

undescribed, I find a number of links connecting the extreme

forms of the prothorax; the difference is one of degree not of

kind, and in such species as Callistomimas coarctatus Laf. and

C. littoralis Motch. the hind angles of the prothorax are sharp

and the sides of the base distinctly, though only slightly,

emarginate. | have endeavoured for some little time past to

keep the genera distinct, but I have now seen intermediate forms

which render this no longer possible, and I therefore treat

Pristomacherus as a synonym.

As in Callistus the outer maxillary lobe is in one piece. Bates

pointed out, when describing Pristomacherus, that the maxille

projected far beyond the mandibles, and this is sometimes the

case. In connexion with this, however, there is frequently an

optical illusion, which struck me when examining a long series of

a new species. Normaliy the mandibles are tightly closed, and the

maxille project as long slender shafts in front of them; when

the mandibles are opened, it is seen that the relative difference

in length of the two organs is considerably less than at first

* Communicated by S. A, Neave, B.A., F.Z,S,

934 MR. H. E. ANDREWES ON THE

sight appeared to be the case. There is also a point to note in

connexion with the ligula, which, as in Callistus, is said to be

bisetose. Actually there are two long sete placed close together

on the middle of the apical margin and directed a little down-

wards, but there are also some very fine sete on each side,

These latter seem very liable to abrasion, but apparently there

are about three such minute hairs on each side of the apical

margin,

Chaudoir thought that Callistus and its allies were related

to Anchonoderus, and in this he was followed by Lacordaire.

Schaum and G. H. Horn put them near Chlenius, and both

Bates and Ganglbauer have followed this example. IJ agree with

the latter, but think that the unjointed outer lobe of the

maxilie and the elongate inner one, together with the

pubescent palpi necessitate a separate group, in which I follow

so good an authority as Bates.

I propose in the following pages to give first of all a brief

account of the generic characters, then a catalogue of the species,

followed by a dichotomic table, and finally descriptions of the

new species, together with additional notes on some of the old

ones, where the information afforded by their authors appeared

inadequate. In differentiating closely allied species I am aware

that it is generally undesirable to rely too much upon colour, but

in the insects which I am here considering, especially the smaller

ones, the structural characters vary little, and I have been com-

pelled to make use of the colour scheme in my table of species.

GENERIC CHARACTERS.

LTigula dilated and truncate at apex, bisetose but with some

additional minute sete at sides of apex; paraglosse membranous,

adnate to near apex, extremities rounded, extending rather beyond

ligula. Mentwm edentate, lobes shar ply contracted in fr ont, not

much rounded, pointed at apex; epilobes narrow, extending

slightly in advance of lobes. Mawille exceptionally long and

narrow, with relatively few bristles on inner margin, especially i in:

upper part, sharply hooked close to apex, outer lobe in one piece.

Palpi \ong, slender, setose, pointed at apex, penultimate of labials

plurisetose on inner margin. Mandibles long, almost edentate,

wide at base, slender and very sharp, but hardly hooked, at apex.

Labrum transverse, generally emarginate, sexsetose at apex,

minutely setose at sides. Clypeus truncate, a seta not far from

front angle on each side. yes prominent, one supraorbital seta.

Antenne thick, setose, densely so from joint 4.

Upper surface densely punctate and setose; hind angles of pro-

thorax usually acute, with a more or less deep emargination

between them and middle of base; hind body subpedunculate ;

elytra 9-striate, with a fairly long seutellary str iole between

suture and stria 1, base unborder ed) over first three intervals on

each side, sides secant sinuation or visible internal fold near

ORIENTAL SPECIES OF CALLISYOMIMUS. 235

apex; under surface shiny, coarsely but not densely punctate,

ventral surface more finely punctate, metepisterna not much

longer than wide; legs slender, tarsi setose on upper surface,

joint 1 longer than 2+3, ¢ front tarsi with three dilated joints,

more ov less quadrate with rounded angles (as in Chlentus),

joint 1 narrowed towards base, densely pilose beneath, claws

sunple.

CATALOGUE.

acuticollis (Callistus), Fairm. Ann. Soc. Ent. Fr. 1889, p. 6=yunnanus

(Pristomacherus) Maindr. Bull. Soc. Ent. Fr. 1909, p. 275. Yunnan.

belli, sp. n. S. India.

cauhops (Pristomacherus) Bates, Ann. Mus. Civ. Gen. 1892, p. 306.

Burma.

ecylonicus Dupuis, Ann. Soc. Ent. Belg. 1913, p. 6. Var. of littoralis,

Motch. Ceylon.

chalcocephalus (Panagseus) Wied. Zool. Mag. 11.1. 1823, p. 57; Chand.

Bull. Mose. 1861, iv. p. 351; id. Ann. Soc. Ent. Belg. 1878, p.84;

Bates, Scientific Results Sec. Yark. Miss. 1891, p. 4 = messii

(Pristomacherus) Bates, Trans. Ent. Soc. Lond. 1873, p. 324. S. China,

Indo-China, N. India.

‘ ; P Java.

chlorocephalus (Panageus) Kollar, Ann. Wien. Mus. 3. 1835, p. 335,

t. 31. f. 4; Chaud. Bull. Mose. 1861, iv. p. 351; id. Ann. Soe.

Ent. Belg. 1878, p. 84. P India.

coarctatus (Callistus) af. Ann. Soc. Ent. Fr. 1851, p. 280. N. India.

@abreui, sp. n. N. and Cent. India.

dicksoni Wat. Ann. Mag. Nat. Hist. (5) xiv. 1884, p. 429; Andr.

Trans. Ent. Soc. Lond. 1919, p. 197. Formosa.

dux, sp. n. Burma.

eucharis (Pristomacherus) Bates, Ann. Mus. Civ. Gen. 1892, p. 305. _ Burma.

humeralis, var. nov. Var. of modestus Schau. Indo-China, Burma, N. India.

insularis, var. nov. Var. of modestus Schaum. Java.

jucundus, sp. n. S. India.

lebioides (Pristomacherus) Bates, Ann. Mus. Civ. Gen. 1892, p. 306. Burma.

littoralis (Callistus) Motch. Et. Ent. 1859, p. 33; Bates, Comp. Rend.

Soc. Ent. Belg. 1891, p. 827 = westwoodi (Callistus) Schaum,

Berl. Ent. Zeit. 1863, p. 85. Tndia.

modestus (Callistus) Schaum, Berl. Ent. Zeit. 1863, p. 85; Bates,

Trans. Ent. Soc. Lond. 1878, p. 246; id. Ann. Mus. Civ. Gen.

1892, p. 303; Fairm. Ann. Soc. Ent. Fr. 1888, p. 336. Japan, 8. China,

Indo-China, Burma, N. India.

nair (Pristomacherus) Maindr. Bull. Soc. Ent. Fr. 1909, p. 276. S. India.

nilgirinus, var. nov. Var. of nair Maindr. S. India.

quadricolor (Pristomachwrus) Putz. Stett. Ent. Zeit. 1877, p. 101.

? =eucharis Bates.

quadriguttatus (Pristomacherus) Putz. Stett. Ent. Zeit. 1877, p. 101.

? =chalcucephalus Wied.

quadristigma (Pristomacherus) Bates, Ann. Mus. Civ. Gen. 1892,

p. 303. Var. of chalcocephalus Wied. Indo-China, Burma.

rubellus (Pristomachwrus) Bates, Ann. Mus. Civ. Gen. 1892, p. 305.

N. India.

Burma.

sikkimensis, sp. n. N. India.

subnotatus, sp. n. Cambodia,

suturalis Fleut. Ann. Soc. Ent. Fr. 1887, p. 61, t. 4. f. 2; Bates, Ann.

Soc. Ent. Fr. 1889, p. 265. Annam.

venustus, sp. n. Laos.

virescens, sp. n. Burma.

vitalisi, sp. n. Tonkin,

Proc. Zoou, Soc.—1921, No. XVII, ia

[SU we}

—

es)

—

15 (22).

16 (17).

17 (16).

18 (19).

19 (18).

20 (21).

21 (20).

22 (15).

23 (28).

MR. H. E. ANDREWES ON THE

Key to the Species.

.). Upper surface black to brassy green (prothorax

sometimes reddish), elytra with red or yellow

spots (sometimes fasciew), prothorax closely

and coarsely punctate.

. Prothorax green, elytra with red or yellow spots.

- Prothorax half as wide again as long, sides hardly

sinuate before hind angles. Length 7:v-

775 mm.

- Yellow spots on elytra smaller, front one from

stria 5 to margin

. Yellow spots on elytra larger, ‘rout one from

stria 4 to margin

- Prothorax not more than a . third a as s wide a again

as long, sides strongly sinuate before hind

angles. Length 5°0-6°0 mm.

. Elytral spots red, front one covering shoulder and

extending inwards to stria 5, two minute ones

behind on intervals 4 and 6 respectively ......

. Elytral spots yellow, front one small, rather

behind shoulder, extending inwards to stria 7,

hind one small, quadrate, on intervals 6-8,

another minute spot on interval 4 .

. Prothorax reddish, elytra with yellow fascie.

. Prothorax half as wide again as long, elytra

coarsely and confluently ‘punctate.

. Prothorax dark red, elytra without sutural red

patch; eyes small, very prominent, pedunculate

. Prothorax light red, elytra with sutural red patch

reaching to midway between fascie ; eyes

moderately prominent, not pedunculate ......

. Prothorax a third as wide again as long, dark

red, elytra moderately and not contluently

punctate, without sutural red patch ............

. Upper surface variegated: head metallic green

or blue, prothorax red (at least at base), elytra

dark (at least on apical half) with two inter-

rupted whitish fascia (exceptionally with

white spots); prothorax closely but not

coarsely punctate.

Elytra with a red sutural stripe from base to

apex, merging in white spot (when present)

at apex.

Prothorax blue with red base, elytra violet- blue,

with wide dirty-white fascixe from sutural

stripe.to side margins ; Li OL ale Ra

Prothorax red, elytral fascie not reaching

margin.

Shoulders and epipleuree of elytra coarsely

punctate; front fascia hardly Rava extern-

ally beyond stria 6

Shoulders and epipleure of. elytra. not coar: arsely

punctate; front fascia reaching at least to

stria 8.

Hlytra maoderatels convex, front fascia reaching

stria 8, hind fascia as wide as front one,

oblique... :

Elytra flat on “disk, “front fascia reaching stria

9, hind fascia very narrow, bent sharply back-

wards soon after leaving sutural SOuIDC Re peer:

Elytra with or without “red sutural ‘stripe or

scutellary patch ; when present, not reaching

beyond hind fascia.

Elytra without red sutural stripe or scutellary

path (though the scutellum itself is red).

chalcocephalus Wied.

v. quadristigma Bates.

subnotatus, sp. n.

Jucundus, sp. n.

cauliops Gates.

rubellus Bates.

sikkimensis, sp. 0.

venustus, sp. N.

virescens, Sp. N.

dicksoni Wat.

suturalis Fleut.

28 (23).

29 (30).

30 (29).

31 (36).

32 (38).

33: (32).

34 (35).

35 (34)

38 (31).

37 (42).

38 (39).

9 (38).

40 (41).

41 (40).

42 (37).

43 (44).

44, (43).

45 (46).

46 (45).

47 (48).

‘48 (47).

49 (50).

50 (49).

ORIENTAL SPECIES OF CALLISTOMIMUS.

. Elytva green, with fairly wide yeliow-white

fascie ; prothorax red, with two dark spots on

disk, one on each side of median line.........

. Elytra black with narrow fasciw, prothorax

*. concolorous.

. Prothorax considerably wider than head, elytra

witi a small light spot at apex, sides of front

fascia tapering inwards..

. Prothorax very “little wider than | head, elytra

without light spot at apex, front fascia of

nearly equal width throughout

Elytra with a red sutural “stripe or scutellary

patch.

Ground colour of elytra green, each elytron with

4 white spots. IL ngth 40 mm.

Ground colour of elytra black or very dark blue.

Length at least 4°5 mm.

Elytra with the whole of the basal area red.

Elytra with red basal area reaching hind fascia,

without dark line at margin between shoulders

and front fascia, ee often blue with

base only red .

Elytra with red ‘basal area not ‘yeaching hind

fascia, a dark line at margin between shoulders

and front fascia.

Prothorax red, ground colour of elytra dull black.

Length 7-0 mm..

Prothorax with base red and apex ‘blue (Gn v vary-

ing proportions), ground colour of elytra dark

blue, faintly shining. Length 4 mm. ......

Elytra with sutural area only red (to a greater

or less width), shoulders always dark.

Red sutural area on elytra joining hind fascia.

Elytra half as long again as wide, shoulders not

prominent; hind angles of prothorax acute, a

distinct emargination between them and

middle of base :

Elytra about a fourth as s long : again as ‘wide,

shoulders prominent ; hind angles of pro-

thorax right, with only a slight emargination

between them and middle of base.

Elytra with red sutural stripe covering intervals

1-3 at base, 1-2 between fascie . F

Elytra with red sutural stripe covering intervals

‘1-5 at base, tapering to 1-3 at hind fascia

Red sutural area on elytra not joining hind fascia.

Red sutural area on elytra reaching level of hind

fascia, but separated from it Be

Red sutural area on elytra not or hardly extending

behind the front fascia.

Elytral fascize reaching margin, neck red .........

Elytral fascia not reaching margin, neck dark.

Hind angles of prothorax right, quite incon-

spicuous, a minute indentation between them

and middle of base

Hind angles of prothorax acute, a a deep emargin-

ation between them and middle of base.

Front elytral fascia covering intervals 5-8, hind

one evidently oblique, moderately wide, cover-

ing intervals 3-8 55600

Front elytral fascia covering intervals 4-8, ‘hind

one only slightly, blige narrow, covering

intervals 2-8 . vasivddecenates sauors

belli, sp. n,

acuticollis Fairm.

lebioides Bates.

@abreni, sp. n.

modestus Schaum.

dus, sp. n,

v. insularis, nov.

naiy Maindr,

littoralis Motch.

v. humeralis, nov.

v. ceylonicus Dup.

vitalisi, sp. n,

coarctatus Laf.

v. nilgirinus, nov.

eucharis Bates.

237

238 MR. H. E. ANDREWES ON THE

CALLISTOMIMUS CHALCOCEPHALUS Wied. (Plate I. fig. 1.)

Callistomimus messic Bates.

do. do. var. quadristigma Bates...

Wiedemann’s type, which I have recently seen at Copenhagen,

was alleged to come from Java and may have done so, though I

have seen no other example from the Malay region, He was in

the same paper also describing species from Bengal, and I think

it possible that his insect came from that locality.

Bates described his C. messti from Hong Kong, and C. quadri-

stigma from Burma. The former is identical with Wiedemann’s

species: of the latter, which has rather larger yellow spots on

the elytra, Bates says ‘“‘ Probably not more than a local race of ”

C. messit, with which I quite agree.

T have before me specimens of the type form from Hong Kong,

and from Tonkin—Than Moi (2. Vitalis de Salvaza), of the variety

(in addition to cotypes from the Fea collection) specimens from

Laos— Vientiane (2. Vitalis de Salvaza), Cochin-China— Bien-hoa

(Coll. Fleutiaux), and Burma—Rangoon (H. L. Andrewes); also

intermediate forms, as regards the size of the yellow spots, from

Burma—Maymyo (i. L. Andrewes), Sikkim—Gopaldhara (#.

Stevens), and Dehra Dun (/ndian Musewm and Forest Res. [nst.).

An example from Kumaon—W. Almora (Hf. G. Champion) has

very small elytral spots, and the upper surface, as in all Indian

specimens I have seen, is more strongly punctate than in the

China and Burma forms. The example taken by Dr. Stoliczka in

the Jhelam Valley (Second Yarkand Mission), and determined

by Bates as C. chalcocephalus, is now fragmentary, the head and

prothorax having disappeared, but the elytra seem to be those of

the species under consideration, with the yellow spots well

developed. Bates indicates in his paper no feeling of doubt

regarding this determination, but he put two marks of interro-

gation on his label. With such slender Indian material available

it seems useless to put names to any of the slightly varying forms.

C. chlorocephalus Kollar has not to my knowledge been identi-

fied as yet, and, like Wiedemann’s species, it was evidently

unknown to Chaudoir. ‘The locality is uncertain, but Kollar

thought the specimen probably came from India. The figure is

a poor one, but, judging by the hind angles of the prothorax, I

think the species will prove to be identical with chalcocephalus*.

C. quadriguttatus Putz. is probably also the same thing, but I

have unfortunately not yet ascertained the whereabouts of

Putzeys’ types.

CALLISTOMIMUS SIKKIMENSIS, sp. n. (Plate I. fig. 5.)

Length 5°5 mm.

Black. Head metallic green ; prothorax red, with a vague dav

faintly eneous patch on disk'on ech side of median line; elytra

* Dr. Holdhaus has kindly compared a specimen of C. chalcocephalus, which I

sent to him, with Kollar’s ty pe in the Vienna Museum, and finds the species to he

different. The type is unfortunately a “ruin,” so that he cannot send it to me tor

examiiation.

ORIENTAL SPECIES OF CALLISTOMIMUS. 239°

with a very faint brassy tinge, each with two interrupted yellow

fascie; joints 1-3 of antenne, palpi, and legs flavous ; buccal

organs, underside of head, epipleuree of prothorax, and a faint

common spot at apex of elytra more or less testaceous.

Head (1:0 mm. wide) convex, moderately and closely punctate,

hardly less closely on middle of front than elsewhere, labrum

truncate, maxilla not longer than mandibles; antenne nearly

half the length of body, the 8 apical joints wide, flattened, and

quite black. Prothoraw (1:2 mm. wide) not much wider than

long, convex, sides moderately rounded in front, sinuate before

hind angles, which are acute, reflexed, and point both outwards

and backwards ; median line and basal fovex deep, surface closely

and coarsely punctate. Hlytra (2:10 x 3:0 mm.) ovate, convex,

with well marked shoulders, striz fairly deep and finely punctate,

intervals moderately but closely punctate, the punctures hardly

confluent; front fascia from stria 3 to margin, transverse,

widening a little at margin towards shoulder and tingeing the

epipleura with yellow; hind fascia from stria 3 to 9, a little

oblique outwards and backwards. Underside shiny, moderately

punctate, rather more closely and finely on ventral surface.

Practically the whole body is shortly pubescent.

Much smaller than C. chalcocephalus; head more finely and

more uniformly punctate, eyes less prominent, prothorax red (not

eeneous), much narrower and more strongly sinuate before hind

angles, elytra relatively longer and narrower, more finely punctate,

the yellow markings differently shaped.

Sikkim, Gopaldhara (H. Stevens), 1 ex. ¢. Mr. Stevens has

kindly allowed me to retain the type.

CALLISTOMIMUS SUBNOTATUS, Sp. N.

Length 5°75 mm. Width 2°50 mm.

This species bears so strong a likeness to C. yucundus m., next

described, that I need do no more than point out the characters

in which it differs.

Colour (except for elytral markings) identical. Head and

prothoraw more coarsely and less closely punctate. #lytra more

coarsely and confluently punctate, a little more shiny: front spot

reddish, larger, elongate, covering the whole shoulder from

margin to stria 5 (and including the epipleura), produced back-

wards along the two outer intervals to very nearly half the length

of the elytra; two minute reddish ‘hind spots, one placed on

interval 4, as in some examples of C. jucundus, the other a little

further back on interval 6.

Cambodia, Kompong Kedey, April 1914, lex. 2 (2. Vitalis de

Salvaza). The type is in the British Museum.

CALLISTOMIMUS JUCUNDUS, sp.n. (Plate I. fig. 2.)

Length 6°0 mm. Width 2°50 mm.

Black. Head neous, prothorax above and beneath with

bluish reflection, elytra with a faint green tinge; mouth-parts

240 MR. H. E, ANDREWES ON THE

joints 1-3 of antenne, two spots on each elytron, epipleure. of

elytra adjacent to front spot, and legs testaceous yellow.

Head (1:20 mm. wide) convex, shiny, pubescent, very coarsely

punctate ; clypeus, neck, and a small foveiform area on middle of

front and vertex smooth and polished, eyes prominent. Prothorax

convex, rather narrow, not much wider than head (with eyes), a

third wider than long, sides rounded, sinuate before hind angles,

which are acute, reflexed, and directed backwards; median line

rather deep; surface closely, coarsely,and confluently punctate, with

yellowish pubescence. Hlytra convex, rather square at shoulder,

widest behind middle, nearly twice as wide and two and a.

half times as long as prothorax, punctate-striate, intervals 2. and

4 a little narrower than the adjacent ones; surface densely

punctate (but much less coarsely than prothorax), punctures more

or less laterally confluent; front spot small, triangular, behind |

shoulder, widening from stria 7 to margin, hind spot small,

square, at a fourth from apex, covering intervals 6-8. . In some.

examples there is a small additional spot on interval 4, a little in

advance of the hind spot. All sterna coarsely, ventral surface

more finely punctate, pubescent.

A good deal smaller than C. quadristigma Bates, head and

prothorax much narrower, the latter darker in colour and less

contracted at extremities, elytral puncturation similar, but

yellow spots much smaller.

Nilgiri Hills, Ouchterlony Valley, 3500 ft., July, “running

among beams on the ground” (//. L. Andrewes), 3-ex. 3 Q.

Bombay: N. Kanara (7. R. D. Bell), 2 ex. The type (¢) is in

my collection.

CALLISTOMIMUS VENUSTUS, sp. n. (Plate I. fig. 6.)

Length 6°0 mm. Width 2°25 mm.

~ Black. Mouth-parts (exe. last joint of palpi fuscous), antennal

joints 1, 2, and underside of 3, and epipleure of elytra (exe. at

shoulder) testaceous; head and prothorax blue or blue-green,

with red base and side-margins; elytra violet-blue, suture red

over intervals 1 and 2, and for basal third over interval 3; two

interrupted fascie on elytra, apical margin, and legs yellow-white

(apex of femora, tibie, and tarsal joints fuscous); metasternum

and ventral surface (last segment darker) brown.

Head (about 1:0 mm, wide) shiny, a little less convex but more

pubescent and punctate than in allied species, the usually

smooth space on middle of front with one or two coarse punctures,

neck smooth, eyes prominent. Prothorax relatively flat, rather

small, hardly wider than head with eyes, nearly half as wide

again as long, sides rounded in front, contracted behind, and

-sinuate before hind angles, which are acute and project both

laterally and a little backwards; transverse impressions, median

line, and basal fovex all well marked; surface closely and con-

fluently punctate, and pubescent. lytra square at shoulders,

nearly parallel, rather flat, twice as wide and three times as long

ORIENTAL SPECIES OF CALLISTOMIMUS. 24)

as prothorax, weakly striate, the whole surface rather dull and

finely aciculate-punctate; front fascia at a third from base, not

very wide, but widening outwards from stria 3 to margin; hind

fascia at a third from apex, of about same width as front one,

widening from stria 2 to margin. Underside moderately

pubescent, last ventral segment more strongly so and trans-

versely subrugose.

Distinguished from its congeners by its small prothorax, square

shoulders, and especially by the fact that the colour of the red

scutellary patch extends along the suture to apex.

Laos: Pak Tha, Paklung and Ban Saleun, 16 ex. ¢ 2

(R. Vitalis de Salvaza). The type is in the British Museum.

CALLISTOMIMUS VIRESCENS, sp. n. (Plate I. fig. 7.)

Length 5mm. Width 2 mm. ;

Black. Head blue, prothorax red with purplish reflections on

upper surface, a sutural stripe on the elytra yellow-red, elytra

velvety blue-black with metallic blue-green lustre on shoulders

and along sides; two fascie on elytra (interrupted by the sutural

stripe) and apical margin, joints 1—3 of antenne, palpi (exc. apex),

front margin of labrum, and legs (exc. apex of femora) yellow-

white.

Head (about 1-0 mm. wide) convex, shiny, coarsely punctate

at sides, smooth and highly polished along median line and on

neck; maxille as long as mandibles. Prothorax very couvex,

a little transverse, just wider than head, sides bordered and

strongly rounded, more contracted behind than in front, hind

angles in form of a minute right-angled tooth, with sides of base

advancing towards them, median line short but deep ; surface

coarsely but not very closely punctate, faintly pubescent. Llytra

punctate-striate, intervals flat, punctate at sides but not evidently

so on disk, pubescent, base witha strongly marked border, region

of the shoulder irregularly punctate and rugose, epipleure

coarsely punctate; sutural stripe covering intervals 1 and 2 (more

or less), front fascia at a fourth from base, narrow, reaching

stria 6, hind fascia at a third from apex, wider, reaching stria 9.

Allied to C. sutwralis Flt., but a little smaller, hind angles

of prothorax less evident, elytra more convex, sutural stripe

narrow at apex and lighter in colour, hind fascia much broader,

shoulders and sides much more rugosely sculptured. The very

coarse puncturation of the elytral epipleure at once distinguishes

the species from its congeners.

Burma: Tharrawaddy, 2 ex. ¢ 2 (G. Q. Corbett). The type

is in my collection.

CALLISTOMIMUS BELLI, sp.n. (Plate I. fig. 8.)

Length 4:25 mm. Width 1:75 mm.

Black. Head dark metallic green with purplish reflections;

prothorax brick-red with a small rather faint purplish spot on

each side of disk; elytra dark green; mouth-parts, joints 1—3 of

242 MR. H. FE. ANDREWES ON THE

antenne, underside of head and prothorax (exe. prosternal

process and region round front cox), epipleure of elytra; two

interrupted fascize on elytra (with a Be plish border), a common

apical spot and margin, testaceous; legs (exe. apex of femora,

which is fuscous) yellow- white.

Head (barely 1:0 mm. wide) shiny, convex, pubescent, coarsely

punctate over whole surface (exe. clypeus and labrum), though a

little less closely on middle of front, neck smooth, eyes prominent.

Prothorax moderately convex, salir: wider than head, sides

rounded and a little sinuate before hind angles, which are acute,

slightly retlexed, and directed outwards and backwards; ; surface

densely but not coarsely punctate and covered with a close, dark

pubescence. LHlytra rather ovate, about half as wide again as

prothorax, punctate-striate, surface aciculate-punctate and

pubescent ; interval 2 narrowing towards base and 6 towards

apex; front fascia at a third from base, a little wider than in the

allied species, extending from the middle of interval 3 to margin,

where it widens out both in front and behind, an indentation of

the ground colour in front on interval 6; hind fascia oblique

(outwards and backwards), extending from stria 2 to margin ;

apical spot a little larger than in the allied species. Underside

moderately punctate and pubescent, prosternal process bordered,

last ventral segment transversely substrigose.

Not unlike (. lebioides Bates in form, but quite different in

coloration. Prothorax less transverse, less coarsely punctate,

hind angles much more evident; fascise on elytra much wider and

extending to margin, an apical white spot (absent in lehzoides).

The species are alike in the absence of a red scutellary patch.

Bombay: North Kanara, Kadra, 11. vii.1907, 2 ex. ¢ 2

(T. k. D. Bell), The type is in my collection.

CALLISTOMIMUS AGUTICOLLIS Fairm.

CALLISTOMIMUS YUNNANUS Maindr.

Both these species were described from Yunnan, and I have at

different times seen the types of both, though I have not been

able to compare them. I have before me the type of yunnanus,

which agrees with Fairmaire’s déscription and with some notes I

made on the type of acuticollis. I think the species are identical.

T have seen further examples from Djoukoula, Yunnan, in the

collection of Mr. H. de Touzalin.

CALLISTOMIMUS D’ABREUI, sp. n, (Plate I. fig. 9.)

Length 4:0 mm. Width 1-5 mm.

Black. Head dark neous with purplish reflections ; prothorax

and a scutellary patch red, former with an ill-defined purplish spot

on each side of disk; elvtra deep green, each with four white

spots; mouth-parts, underside of joint | and base of joints 1 and

2 of antenne, epipleure, and margin of elytra testaceous, sterna

and cox reddish, trochanters aad femora yellow-white. In the 3

ORIENTAL SPECIES OF CALLISTOMIMUS. 243

specimen (type) the tibie and tarsi are fuscous, in the two Q 2

specimens they are yellow- white.

Head (0:90 mm. wide) shiny, convex, slightly pubescent,

coarsely punctate at sides and back, vertex and middle of front

smooth and polished, eyes moderately prominent, maxille a little

longer than mandibles. Prothorawv narrow, just. wider than head,

not much wider than long, sides rounded in front, strongly

contracted behind, sinuate just before hind angles, which are

right; surface closely and coarsely punctate, pubescent. Hlytra

about half as wide again as prothorax, square at shoulder, widest

at about middle, punctate-striate, intervals rather convex, the

whole surface finely aciculate-punctate, dull viewed from above,

but a little shiny when viewed sideways; the red scutellary patch

extending half-way to apex, bounded on each side at base by

stria 2, but tapering behind; the front white spot, at a fonrth

from base, extends from interval 4 to margin (it is rounded on

intervals 5-7, contracted on 8, and expanded again on 9); the

hind spot, also rounded but a little larger, at a fourth from apex,

covers intervals 5-7: the two remaining spots are minute, one

on interval 3 just behind the end of the scutellary patch, the other

at extreme apex. Underside sparsely, ventral surface (exc. at

sides) more strongly punctate and pubescent.

India: Central Provinces, Mandla district, Chiriadangra,

Nerbudda bank, 7.xii.1916 (#. A. D’Abrew), 1 ex. fg (type).

U.P.: Sitapur, July 1917 (#1. G. Champion), 1 ex. 2 (defective)

Bihar: Pusa, 1 ex. 9 — Agric. Res. Inst. Pusa. Mr. D’Abreu

has kindly allowed me to keep the type in my collection.

C/ALLISTOMIMUS MODESTUS Schaum.

Callistomimus amabilis Redt. (in litt.) Reis. Novar. ii. 1867,

Col. 20.

The species was described from Hongkong,and I have not seen

the type; there seems to be no doubt regarding the identification,

and, as it is probably the best known species in the genus, I think

it unnecessary to prepare a fresh description.

The name amabilis was introduced by Redtenbacher, who

attributed it to Chaudoir; the latter mentions the name when

describing the genus, but attributes it to Redtenbacher. I

cannot find that either author published a description, and it

seems a pity that it should have got into circulation.

Var. humeralis, nov.

Schaum gives no hint of a dark spot on the shoulder, but Bates

refers toa small black shoulder-spot on some of the examples

taken by Mr. Fea in Burma, one of which is in my collection ;

the specimen described by Fairmaire from Tonkin is similarly

marked. A specimen in the Paris Museum, also from ‘Tonkin,

another from Bengal—Sarda (/. W. Champion), and other Indian

examples in the British Museum, all have this spot more fully

developed, and J think the form is worthy of a name.

244 MR. H. E. ANDREWES ON THE

Var. insularis, nov.

Mr. Guy Babault has kindly sent me an example (there being

others I understand in his collection) which comes from Java,

which has a narrow black longitudinal line behind the shoulder.

and in which the two parts of the hind fascia are quite detached

from the reddish area in front, so that they appear as transverse

whitish spots on. a blue-black ground, which covers the apical

half of the elytra. In the Brussels Museum there are also three

examples of this form labelled “Semarang ( Drescher)” * ; in these.

the hind fascia is less widely separated from the red area in.

front.

CALLISTOMIMUS DUX, sp. n. (Plate I. fig. 10.)

Length 7:0 mm. Width 2°60 mm.

Black. Head dark blue, prothorax (above and beneath) dull

red with faint purplish reflections, basal two-fifths of elytra brick-

red, the coloured area bulging in the middle and reaching to half-

way between base and apex, two transverse fascie (front one very

short, submarginal) and extreme apex yellow-white, joint 1 of

antenne (rest wanting) and legs (exe. apex of femora) testaceous,

margin of elytra (exc. just below shoulders) infuscate.

Head (2°40 mm. wide) convex, shiny, very coarsely punctate ;

vertex, middle of front (exe. for one or two punctures), and

clypeus smooth, eyes prominent. Prothorax (2°60 mm. wide)

convex, very transverse, sides strongly rounded, much contracted

at extremities, but more so behind than in front, hind angles in

the form of a small right-angled tooth, a little in front. of

apparent base; median line faint, basal fover round and deep,

surface coarsely and confluently punctate. lytra oval, flat,

punctate-striate, intervals nearly flat, finely punctate, (presumably)

pubescent ; front fascia from stria 5 to 8, merging in colour

of basal area, hind one moderately broad, oblique, from stria 1]

to 9. Underside moderately and rather coarsely punctate, more

finely on ventral surface.

Burma: Prome (G@. Q. Corbett), 1 ex. g. Type in ss Lee ious

The species is so widely different from all others known to me

that I have described it with a view to making my note on the

Oriental species of the genus as complete as possible. ‘The unique

exam ple, however, 1s defective, especially in regard to the PUUISS DILEE.,

tarsi, and buccal organs.

CALLISTOMIMUS NAIR, Maindr., var. nilgirinws, nov.

IT have in my collection an example of Maindron’s species,

which I have had the opportunity of comparing with a typical

specimen kindly sent to me for examination by Mr. Guy Babault.

My specimen agrees almost exactly with Maindron’s, which came

from Mahé on the Malabar Coast: the only difference I detect is

in the elytral fasciz, which are narrower, the hind one being also

* Tn reply to an enquiry of mine, Commandant Dupuis writes: “ Etique te d’un

enyoi recu du Musée de Sarawak (Bornéo), sans autre précision, mais les. insectes

provenaient tous, je crois, de récoltes faites dans la région.”

ORIENTAL SPECIES OF CALLISTOMIMUS. 245

less regular in outline and wider near the suture than near the

margin, ‘This specimen was taken in the Nilgiri Hills by Mr...

H. L. Andrewes, and his note reads ‘ Ouchterlony Valley,

December, 3500 ft. Flooded out during irrigation of tea.”

A second specimen ( ¢ ) was also ellen by Mr. Andrewes in the

Nilgiri Hills, but I have no note of the exact locality. It is of

the same size as C. nair and does not seem to differ structurally,

but there are distinct differences in the coloration. Joints 5-11 of

the antenne and the front tarsi are brown instead of black; the

margin of the elytra is only vaguely reddish, there is only

suggestion of a scutellary red patch, and this does not extend

perons the frout fascia; this latter is very small and its sides

taper inwards from stria 8 to 4, the hind fascia is rather wider

than the front one and extends from stria 8 to 2, neither of the

fascive reaching the margin. Whether this form will prove to be

a local race, or a mere aberration, remains to be seen, and for the

present I call it var. nilgirinus.

CALLISTOMIMUS LIrToRALIS Motch. (Plate I. fig. 11.)

Callistomimus westwoodi Schaum.

Length 4°55 mm. Width 2:0 mm.

Black. Head and front margin of prothorax greenish-blue ;

vest of prothorax (upper sidg) and a sutural stripe, bounded at

sides by stria 2 and extending backwards to the hind fascia, dull

red (the sutural stripe widens to stria 3 at the front fascia);

elytra dark blue; two interrupted fasciz on elytra (exe. on

intervals 3,4 and 9 of hind one, where reddish), apical margin,

a common apical spot, and legs (exc. apex of femora and tibie

fuscous) yellow- white 5 underside of prothorax, meso- and meta-

sterna, side margins and epipleure of elytra, and buccal organs

testaceous.

Head (0°38 mm. wide) convex, closely and coarsely punctate,

except on neck, clypeus, and middle of front, surface finely rugose,

eyes prominent, maxille elongate but not really reaching beyond

mandibles, labrum emarginate. Prothorax (1:10 mm, wide)

convex, a little transverse, sides strongly rounded and contracted

behind, hind angles small and sharply rectanguiar, surface

moderately but closely punctateand pubescent. Hlytra (2°75 mm.

long) shortly ovate, punctate-striate, pubescent ; front fascia ata

fourth from base, widening out from sutural line to margin;

hind fascia extending obliquely backwards to margin, eouctticked

on stria 4. Underside finely punctate and pubescent.

For a comparison, with C. coarctatus Laf., see under that

species.

I have seen examples from various localities in North, Central

and Southern India, but not from Sind. The var. ceylonicus

Dupuis. from Ceylon is apparently a variety of this species

and not of C. coarctatus*.

* T have recently seen a cotype of the variety, which Dr. W. Horn kindly sent to

me for examination. It proves as [ anticipated to be a var, of Jittoralis.

246 MR. H. E. ANDREWES ON THE

CALLISTOMIMUS CoARCTATUS Laferté. (Plate I. figs. 12, 124,

and 12 d.)

Length 5°50-5:75 mm. Width 2°0-2°25 mm.

Black. Head metallic blue; prothorax above (with purplish

reflections), and beneath (exc. middle of sternum), a scutellary

patch on elytra (tapering rather abruptly at extremity, reaching

nearly half-way to apex, and bounded at sides by stria 4), and

mouth-parts red; elytra blue-black with faint metallic lustre ;

two interrupted fasciz on elytra, apical margin and a common

apical spot, palpi (exc. apical joint), underside of joints 1-3 of

antenne, trochanters, femora (exc. apex), upper side of tibiz

(exc. apex), and tarsal joints (exc. apex) yellow-white.

Head (about 1:0 mm. wide) convex, shiny, more or less

pubescent, coarsely but not closely punctate, smooth and highly

polished on middle of front, vertex, and neck; eyes prominent,

maxille of approximately same length as mandibles. Prothorax

(about 1:30 mm. wide) convex, transverse, strongly rounded at

sides and contracted behind, sinuate before the sharp

acute—hind angles; surface densely and coarsely punctured,

pubescent. lytra punctate-striate, intervals moderately punc-

tate, pubescent ; front fascia, at a third from base, extending on

each side from the sutural red patch at stria 4 to 8; hind fascia,

slanting a little backwards from suture towards margin, extending

from stria 2 to 9, but not reaching margin. Underside shiny,

lightly punctate and pubescent.

Closely allied to @. littoralis Motch., but larger. Head less

closely punctate, prothorax more ‘transverse and more coarsely

punctate, hind angles a little more evident. In littoralis the

front elytral fascia extends to the margin, and the red sutural

patch (which in coarctatus stops half-way between the two

fascie) extends backwards and joins the hind fascia.

The species seems to be confined to the Himalayas. It has

been taken in Kumaon by Dr. 8. W. Kemp and (in considerable

numbers) by Mr. H. G. Champion, in Spiti by Mr. Guy Babault,

and in Sikkim by Mr. H. Stevens and Mr. F. H. Gravely. An

example from Kurseong is in the collection of Mr. H. de Touzalin,

CALLISTOMIMUS VITALISI, Sp. 0.

Length 65 mm. Width 2°5 mm.

Black. Head (exc. neck) eneous green; upper surface of neck

and prothorax (latter with some ill-defined small purplish spots),

and a scutellary patch on elytra red; underside of neck and

prothorax, joints 1-3 of antenne, pron coxe and mouth-parts

(exc. apical joint of palpi, which is fuscous) testaceous ; legs (exe.

middle and hind coxe, and apex of all femora, which are black),

two interrupted fascize and a common apical spot on elytra,

together with margin, yellow-white.

“Head (about 1:0 mm. wide), shiny, convex, slightly pubescent,

moderately punctate, but smooth on neck, middle of front, and

elypeus; eyes prominent, maxilla hardly longer than mandibles.

Prothorax (about 1°50 mm. wide), a third as wide again as long,

moderately convex, sides gently rounded in front, more strongly

ORIENTAL SPECIES OF CALLISLOMIMUS. 247

behind, hind angles acute, reflexed, and projecting a little back-

wards ; surface closely and coarsely punctate and pubescent.

Hlytra striate, the striz faintly crenulate, intervals flat, finely

punctate and pubescent; red scutellary patch ill-defined, extend-

ing about one-third towards apex and to stria 4 at sides; front

white fascia at a third from base, narrow, extending on each side

from stria 4 to margin, where it widens out towards apex; hind

fascia at a third from apex, a little wider than front one, extend-

ing from stria 1 to margin, curving alittle backwards near margin,

but hardly increasing in width. Underside shiny, pubescence

faint, a little more evident on ventral surface, sterna coarsely but

not closely punctate.

In appearance strongly resembling C. coarctatus Laf.; a little

larger and less brightly coloured ; prothorax wider, less convex,

much less contracted at extremities and with more conspicuous

hind angles; the hind elytral fascia less oblique, and extending

inwards to stria 1 (instead of 2). For Bates this species would

have been a Pristomacherus, on account of the acute and reflexed

hind angles of the prothorax.

Tonkin: Hoabinh, 6 ex. d Q (&. Vitalis de Salvaza). The

type is in the British Museum.

CALLISTOMIMUS EUCHARIS Bates.

Putzeys’ C. guadricolor came from Darjiling, and I do not know

where the type is to be found. Bates differentiated his Burmese

species from it by the puncturation of the prothorax, apparently

the only point of difference which struck him. Mr. H. Stevens

has lately sent me from Gopoldhara in Sikkim some specimens

which agree exactly with cotypes of C. ewcharts in my collection,

and I think the two species are the same. There must, however,

remain some doubt until Putzeys’ type is available for examination.

A single defective example in the Brussels Museum, labelled

“Central Java, Mt. Oengaran (4. Aoller),” is evidently a very

close ally of this species, but the elytra are a little wider and less

deeply striate, the fascize being more oblique.

Index.

Page Page

ACULICONIS agri n-ne LOU Zeen | IMeSSIL ates) | nes eeee eee. 238

amabilis Redt. ...............00. 00 243 | modestus Schawm ............... 237, 243

Delliy Spins eerenesse acess een ZOMeele |) Malm Mandira one eee) DOTRO4A

CMMI OOS: TEVRAS Goons doo codecs eeecnc 236 | nilgirinus, var. nov. (nair) ...... 237, 245

ceylonicus Dupwis var. (litto- quadricolor Pwtz................... 247

THINS) Gooanodobeuocceronidoness boven PEG ein | CuexbieainmnnS J2iee, 238

chalcocephalus Wied. ............ 286,238 | quadristigma Bates var. (chal-

chlorocephalus Kollar ......... 238 cocephalus) Seiuecowadean Ely ABs

CoarciatusPMajam east ee eeeee eS OUe + OM Lube lluSm Bites) saan nnn nne 236

Wabreui, sp. n. ............ 4+... 287,242 | sikkimensis, sp. n. ............... 236,238

Gicksoni ates see eee eee 236 | subnotatus, sp. mn. .................. 236, 239

GWtSs Sse sodusagsocsecceonsnondcs Parcel I OEMS TOG oo 236

eucharis Bates ..................... 237,247 | venustus, sp. n...................... 236, 240

humeralis, var. nov (modestus) 237,243 | virescens, sp. 0. oo. .cccce cee cee eee 237, 246

insularis, var. nov. (modestus)... 237,244 | vitalisi, sp. m....... 00. cece eee eee 237, 246

jucundus, sp. n..... Pepe seen eens SLO ONABO! HN WEStIVOOdM SCatiD Wess... 245

lehbioideswBGitesisss.eesseeeeeesess 237 | yunnanus Maindr. ©.............. 242

Iittoraliswotciy wes. cs een LOMO

248 THE ORIENTAL SPECIES OF CALLISTOMIMUS.,

EXPLANATION OF PLATE I.

Fig. 1. Callistomimus chalcocephalus Wied.

2. oi jucuudus Andy.

3 os cauliops Bates.

4, rubellus Bates.

5. 3 sikkimensis Andr.

6 9 . venustus Andy.

7 Ss virescens Andy.

8, xa belli Andry.

oY: i @abreut Andy.

10. due Andy.

11. 3 littoralis Motch.

12. Aa coarctatus lat.

12 a. vf aA Upper side of head.

12 6. 3s aS Lower side ot head.

ON THE MOLLUSCAN GENUS COCHLITOMA. 249

15. On the Molluscan genus Cochlitoma and its Anatomy,

with Remarks upon the Variation of two closely-allied

Forms. By G. CU. Rosson, B.A.*

[Received December 10, 1920: Read March 22, 1921.]

(Ovtered for Publication by permission of the Trustees of the British Museum.)

(Text-figures 1-9.)’

CONTENTS.

Page

I LVS Ware LS (et Ta) tame Ppt aes opceab oc dash cud god cascbeeon cece ato PRE a

II. Structural: Anatomy of Cochlitoma zebra var. obesa and fulgurata. 250

ILI. Genetic Affinities and Variation of var. obesa and fulgurata ......... 263

LV. On the Classification of the Achatinin€ .....cccecccsecceececeeceeseasesesne 264

I. Iytropucrion.

In 1914 Mis, G, B. Longstaff brought from South Africa to

England some living examples of the big land-snails usually

referred to the genus Achatina. A series of observations was

planned by herself and the author upon the habits and later

development of these forms. ‘These observations were started

early in 1915, and, thanks to the satisfactory way in which the

animals acclimatized themselves, have been successfully continued

and some interesting results obtained which are given elsewhere

(infra, p. 379), As certain phases of the behaviour of the two

forms represented differed rather sharply and as their specific and

generic position was uncertain, an examination of their anatomy

was undertaken by the author. The principal object of this

paper is to provide additional knowledge of the anatomy of the

Achatinine ; but it also affords an opportunity for discussing how

far the individuals of two closely-allied forms agree or differ in

all the details of their structure.

The material for this study was presented by Mrs. Longstaff

to the Zoological Department of the British Museum (Natural

History); and the author is indebted to Mrs. Longstaft, and to

Major M. Connolly for information accessory to the study.

The genus Cochlitoma was established by Férussac (1821) for

_a heterogeneous body of forms mostly referable to Achatina (s.s.).

‘The name was restricted by Pilsbry (1904) to a small group of

the Achatinine. Some thirty-five species were enumerated by

Pilsbry, and no substantial modification of the genus has been

made since. The group is restricted to 8.E. and 8. Africa though a

few forms straggle up the West Coast, and one species is apparently

erroneously recorded from Brazil. Our knowledge of the anatomy

consists of a very brief description of the late embryo and embry-

onic shell, shell-lobes, kidney, jaw, and genitalia of O. zebra, due

* Communicated by Dr. W. T. Cauman, F.Z.S

250 MR. G. C. ROBSON ON THE

to Semper (1870), a brief description by the same author of

C. granulata (Krauss), and a short account of some of the internal

parts of C. erawfordi (Morelet) due to Pilsbry (1904).

II. Srrucrura..

1. ANATOMY OF COCHLITOMA ZEBRA (Brug.) var. OBESA.

Two specimens were available for dissection, both almost fully

grown and referable to Pfeiffer’s variety abesa. ‘They measure

83 x 47 mm. and 82x 47 mm. respectively, and are thus a little

shorter than Pfeiffer’s type (1854). In the following description

the two examples are referred to as z, and z, respectively.

Keaternal Features.

(1) Shell.—The conchological status and synonymy has been

discussed by Pilsbry (1904). It is necessary, however, to refer

in some detail to these features in order to show what relation

there is between the variation of external and internal characters.

In Pilsbry’s description the columella is “arcuate.” In z, it is

perfectly straight and contrasts strikingly with the ar cuate form

seen in %,. In z, there is a well-marked callus which 1s absent

SUD FA The coloration, invariably regarded as an inconstant

feature, is markedly different in the two forms. Z, agrees with

Pilsbry’s description, but in z, the dark stripes are often twice

and thrice the size of the white ones. Both examples have 73

whorls. The aperture varies. In z, it is wider and shorter

than that of z,, measuring 48 x27°5 mm. as compared with

50°5 x 26 mm.

The sculpture is, as Pilsbry says, granoso-decussate. But in

both forms the spiral striz are weak, so that, as a result, the

transverse ribbing is more marked than in the variety fulgurata.

In z, the spiral striation is stronger than in z,. This sculpture

starts at about half-way round the second whorl in z, and three

quarters of the way round the second whorl in z,. It is con-

tinued in both cases to the end of the sixth whorl. <A peculiar

character seen in the var. fulgurata as well, viz. the enlargement

of some of the longitudinal bands of granules, is seen in a weak

condition in z,, but is absent in z,. There is, however, in the

same position an abrupt enlar cement of all the granules,

(2) Coloration.—This is more fully dealt with in Mrs. Long-

staffs paper.

(3) Mantle-folds, ete—The cervical lobe was thick, narrow, and

pointed in z,. That of z, differed only in being much longer.

The posterior lobe is irregular in shape and has two main sur-

faces, one vertical (between which and the cervical lobe the anal

canal runs) and one horizontal. The latter is continuous with

the shell-lobe.

(4) Anws, etc.—The anus and pneumostome lie more or less

closely together. The latter is interior to and rather below the

MOLLUSCAN GENUS COCHLITOMA. 251

former. The wreter opens by a wide and non-plicate aperture

anteriorly to the anus. In front of the ureter lies a small lobe.

A certain amount of distortion made it impossible to compare Zz

and z, in this respect. i

Internal Anatomy.

(1) Alimentary Canal.

The jaw was examined only in the case of z,, that of z, being

previously damaged.

Viewed anteriorly the jaw is markedly lunate and its antero-

dorsal surface is very wide. The interior surface is covered by

a fairly dense cuticle which is continued backwards and round

the sides and bottom of the mouth. It is more dense dorsally

and is continued posteriorly for about 5-6 mm., but is not con-

tinued on to the antero-dorsal plate. There is on this cuticle

a faint but definite striation. The weak ribbing seen in var.

JSulgurata is represented only by a few irregular faint dark lines

which can be seen under a high power.

Text-figure 1.

Cochlitoma zebra var. obesa.

Radula; median and admedian teeth. Tosition of degenerate entocone shown

by dotted line.

The radula. (Text-fig. 1.)

Hocmnl a0 2h ee aee eee 80. Ts 80. SGlisiae

COM ie 69.) Sel33.

Another example from the

Gwatkin collection ...... Lo. ool MEA INI os

These figures represent maxima.

For the purpose of this study only the median, the first two

admedian and the adlateral teeth were compared.

Proc. Zoon. Soc.—1921, No. X VILI. 18

DARD MR. G. GC. ROBSON ON THE

All the specimens examined showed a slender median tooth

with simple, rather piriform cusp. In z, this is abnormal and

irregularly bicuspid. The admedian teeth have large heavily

conical mesocones, rather weak ectocones, and degenerate ento-

cones. In z, the latter were very meagrely developed. The base

of the mesocones is very heavy, broad, and deep. Its inferior

edge is usually, but not always, turned upwards and outwards.

The adlaterals (e. g., about the fortieth) are bicuspid with low

stout mesocones and a very deep narrow bagal plate.

The phary yne.—The attachment of the pharyngeal muscles is

semilunar in shape. It will be more fully discussed under var.

Sulgurata. There is an exceptionally short @sophagus passing

Text-figure 2.

Cochlitoma zebra vay. obesa.

Salivary glands.

into a crop measuring approximately 20 mm. +3. Interiorly

the latter exhibits a number of longitudinal folds with transverse

secondary connecting ridges, which form a rough trabecular

system. In both examples the crop 1s covered by the salivary

glands. The ducts of the latter originate in the usual position.

The glands themselves are in both | cases asymmetrical, the left

being larger than the right. They are joined together in the

median line by two organic connections, one anterior, the other

posterior. Pilsbry (1919) mentions that the glands of Achatina

schweinfurthii von Marts. are similarly joined, but only an-

teriorly. Between z, and z, are several considerable differences

in detail.

MOLLUSCAN GENUS COCHLITOMA. 253

The crop is succeeded by a narrow continuation of the cso-

phagus which fairly rapidly widens into the anterior part of the

stomach, which is thin-walled and crescentic. At the base of

the third whorl this gives off a large reniform, muscular sac

which may provisionally be regarded as the representative of

the cecum found in many Prosobranchs, Opisthobranchs, and

certain Basommatophora. This structure has been noticed by

Wiegmann (1898) in Achatina panthera. It is not present in all

Stylommatophora, but occurs in Bulimus (Borus) ovatus and

proximus according to Plate (1896). Owing to the fact that

the anterior part of the stomach opens close to the pylorus of the

intestine, we may fairly assume that the large muscular part

does represent the originally smaller cecum. The muscular

portion receives the posterior hepatic duct and is very thick-

walled, its inner surface being furnished with characteristic

ridges. A more detailed description of these organs is given

under var. fulgurata. ;

(2) Circulatory and Respiratory System.

As both animals had been dead for some time previous to

dissection, it was impossible to make an injection of the circu-

latory system. ‘This is all the more to be regretted as some

Text-figure 3.

Cochlitoma zebra var. obesa,

Anterior end of pulmonary vein.

interesting results might be expected from the study of the

elaborate cerebral vascular system noticed when dissecting the

circumeesophageal nerve complex.. The pericardiwm is under

one-third the size of the kidney, a smaller proportion than usual.

The auricle, as usual, is thin-walled and is attached to the adrenal

wall of the pericardium for about one-third of its length. It is

irregular in shape, but presents a broad end to the ventricle.

The latter is of a narrow, ovoid shape and is slightly shorter

18*

254 MR. G. C. ROBSON ON THE

than the auricle. Paired auriculo-ventricular valves of a simple

nature were found; but apparently there is no aortic valve.

The course of the aorta was not followed, but its anterior branch

was found passing into the pharynx in the usual position;

before doing which, it gives off several branches which con-

tribute to the cerebral vascular system referred to above. The

pulmonary vein (text-fig. 3) is very large. On its left-hand

side and particularly towards the anterior end of the pulmonary

cavity, it sends out several large branches which become very

much concentrated anteriorly. On the left-hand side there were

no particularly large vessels. In z, the pulmonary vein lies close

to the ureter—on an average )mm.from it. In the last respect

z, differed from z,, being much further away from the ureter.

The latter condition is seen in Archachatina bicarinata (Deshayes

(1821)) and also apparently in Limicolaria. The anterior con-

centration of large secondary vessels is seen in Achatina chryso-

lewca (Pilsbry (1904)) and in Arch. bicarinata.

(3) Renal System.

The kidney measures 58 x 11-12 mm. im z, and 42 x 9-10 mm.

in z, Anteriorly it narrows off slightly to an obtuse point.

Posteriorly it exhibits a curious diagonal groove on its under

surface which separates off a sort of posterior lobe. This occurs

in both z, and z,; but it 1s doubtful whether it is of systematic

value, though it was absent from both specimens of var. fulgurata.

Another interesting feature is the presence of a sort of ‘ heel”

projecting from the posterior end of the kidney. In both forms

this was disposed in prolongation of the main axis of the organ.

(4) Nervous System. (Text-fig. 4.)

General.—In the adult the central nervous system is covered

over by a dense layer of connective tissue which involves the

main ganglia and roots. This layer, which has been commented

upon by previous authors (Nabias (1894)), is exceedingly difficult

to dissect away even in young forms, while in most adults it

appears to have complete histological union with the surface

of the ganglia. In young forms it is less extensively developed,

less closely applied to the ganglia and more hyaline. The growth

of this sheath involves the anterior aorta and its cerebral

branches. It is hoped that a future work may be devoted to the

study of this sheath. This much may, however, be stated that,

whatever functions it may or may not have acquired in the

course of its intimate association with the surface layer of the

ganglia, it has certainly developed a great importance as pro-

viding additional holdfasts for the muscles of the head and

anterior part of the body. For example, the rhinophorial

retractors appear to be partly dependent on the postero-lateral

extension of this sheath. Interesting analogies are suggested

by this structure, particularly on the subject of its function as a

MOLLUSCAN GENUS COCHLITOMA. 255

protection for the brain and support for muscles, but they must

be deferred to a later date.

The cerebral ganglia are joined by a stout commissure. The

several elements of the visceral commissure are very closely

united, and on the right side no distinction between abdominal

and visceral ganglia could be made out. On the left, the line of

demarcation between the visceral and pallial ganglia was fairly

plain, that between the visceral and abdominal less plain. ‘There

are no pallio-pedal connectives, the centres being closely approxi-

mated. The pedal centres are likewise intimately fused to form

Text-figure 4.

Cochlitoma zebra vay. obesa.

Central neryous system (left moiety).

cpde. = cerebro-pedal connective. eple. = cerebro-pleural connective. eg. = cerebral

ganglion. pg- = pedal ganglion. plg. = pleural ganglion. vag. = viscero-

abdominal ganglion. wn.=visceral nerves. plz.=pallial nerves. pdn.=pedal

nerves.

a single mass, all that indicates an originally separate condition

being a faint median ventral furrow. The right side of this

complex is larger than the left.

Pedal Nerves.—In counting the number of nerve-roots arising

from all ganglia, allowance has to be made for a certain amount

of error due to the density of the sheath and the consequent

impossibility of being certain that adjacent roots are separate.

In z, there are eighteen pedal roots on the left and sixteen (three

256 MR. G. C. ROBSON ON THE

fused) on the right. In both specimens a grouping of the pedal

nerves into two tiers, one above the other, could be made out.

The upper tier consists of five roots, one anterior three median

(usually fused) and one posterior. Certain of the lower tier

roots are separate in origin but enclosed together for the first

2-3 mm, of their length in common sheaths of connective tissue.

The arrangement of these differs from side to side and in

individuals. In z, there were seventeen roots on each side.

Pallial and Visceral Nerves.—In neither example can it be

asserted that any nerve arises from the pallial centres. The

latter (as defined by the position of the cerebral-pallial connective)

is clearly marked off posteriorly from its neighbour and no

pallial root arises from it on either side. Pallial nerves arise,

however, from the posterior ganglion mass (viscero-abdominal).

We can only conclude that fusion of ganglia has gone so far that

the slight external divisions are not reflected internally. On the

right there is only one pallial nerve, on the left three. There

are three main abdominal and visceral nerves arising from the

posterior end of the complex. Z, and z, agree in these particulars.

Cerebral Nerves—Fyvom the cerebral ganglia there arise on

each side nine roots in z,, eight in z,. ines consist of three

large labial and pharyngeal nerves, two Bea laes optic,

buccal connectives, and two (one in z,) of uncertain distribution.

There is a certain amount of unimportant difference in the

position and distribution of the nerves between z, and z,. The

buecal commissure is symmetrical and the ganglia closely fused.

On each side six pharyngeal nerves originate from these ganglia,

the origin of which is asymmetrical.

The optic nerve is exceedingly slender and arises a little

posterior to the rhinophorial nerve. It enters the sheath of the

rhinophore above the latter nerve. In z, the base of the rhino-

phore is innervated by a separate nerve arising just anteriorly

to the first mentioned rhinophorial nerve. Amaudrut (1886)

refers the innervation of the base of the rhinophores in Achatina

panthera to a nerve arising from the anterior lobe. There is in

C. zebra no distinction ce three distinct lobes as Amaudrut

described for his species, though traces of a fairly distinct lobe

corresponding to Amaudrut’s “lobe de sensibilité” could be seen

in Z,. Again, there is no fine branch of the rhinophorial nerve as

there is in A. panthera.

(5) Reproductive System.

Neither specimen dissected was apparently in a state of

reproductive activity. The gonad which is situated in the 2-3

whorl, is succeeded by a slender duct which is straight for

about 6 mm. and then becomes stouter and more convoluted.

In the third whorl this passes into the spermoviduct. The male

portion of this, as in the Helicide, forms a sort of gutter incom-

pletely covered by two more or ‘less overlapping ridges. The

duets of the numerous prostatic follicles open into the bottom of

MOLLUSCAN GENUS COCHLITOMA. 257

this. The prostate invests one side of the spermoviduct over

practically its whole length and projects into the lumen of the

uterus. The upper end of the latter is much folded, the lower

end smooth.

The vagina is short (22 mm. in both cases) as compared with the ,

Jong spermoviduct. The spermatheca has a characteristic duct

with a broad and stout basal portion narrowing to a short thin

Text-figure 5.

pr.

en/s in

sheath

Cochlitoma zebra var. obesa.

Genitalia. For lettering see text-fig. 9, p. 262.

section which expands to the long, truncheon-shaped sperma

theca. The duct measures 17 mm. in z, and is longer in z,. The

latter appears not to have attained its full development, so that

the spermatheca may grow at the expense of the duct. The

common duct of vagina and spermatheca measures 15°D mm. in

z, and only 8 mm. in z,.

The vas deferens measures 52 mm. in z, and 37 mm. in z.

It passes to the base of the penis and 1s inserted into the latter

258 MR, G. GC. ROBSON ON THE

at a point about two-thirds of the way from its apex downwards.

In z, the point of insertion is lower down. The penis sheath is

complete, and the retractor is a branch of the right ocular band.

In the internal structure of the penis there is a marked

difference between the two forms. In z, the penis fits closely

into its sheath. In z, tne neck of the sheath through which the

retractor passes is very much thickened and the retractor passed

down it some way before it meets the penis, which is thus shorter

than in z..

2. COCHLITOMA ZEBRA var. FULGURATA (Pfeiffer).

Two specimens were dissected, measuring 115 mm. and 94 mm.

These are designated f, and f, respectively.

Keternal Features.

(1) Shell.—The aperture is in both cases more effuse ventrally

than in the typical form, and the columella is sinuous in both

cases. The sculpture starts in f, on whorl 13-13, in f, on whorl

13-2. The character of the sculpture was cy fale y based upon f,

as f, was very much worn. It consists of flattish blister-like

decussations of a more delicate texture and quality than in zebra.

As in var. obesa they are crowded at the upper sutures. There

is a band formed of enlarged granules running spirally round the

upper whorls. In f, this is lower down on each whorl than in ine

There are eight whorls in each case, and the granules extend

as 3-5 sutural rows on to the last but one whorl in ead a) 715

these are only very slightly represented, though the shell is less

worn. The character of the colour banding varies a good deal

between the two specimens, but it is substantially different from

var. obesa.

(2) Coloration.—A detailed account of this feature will be

found in Mrs. Longstaff’s paper.

(3) Mantle-folds, etc—The cervical lobe is broad and flat in f,,

narrow and pointed in f,. The posterior lobes are not different

from those of var. obesa.

The anus and pneumostome lie very much as they do in var.

obesa and their general character is the same. <A. special lobe is

found parallel to and below the cervical lobe which more or less

screens the urinary aperture as in var. obesa.

Internal Anatomy.

(1) Alimentary Canal.

The jaws agree in both cases. They are flatly lunate when

viewed anteriorly. The upper surface is of the same average

width all the way round and is rather narrow in proportion to its

width. There are faint and irregular signs of broad ribbing and

traces of faint striation.

The jaw is at once distinguished from that of var. obesa by its

narrowness and the parallel edges of its upper plate.

MOLLUSCAN GENUS COCHLITOMA. 259

The radula (text-fig. 6) has the formula :—

he ee Hels ile. 143:

f (Santee 78s. 122"

The median tooth is short-cusped and much wider than that

of var. obesa. The first admedian has a stout but rather low

mesocone, a fairly well-developed ectocone, and only the faintest

trace of an entocone. It is probable, though not quite certain,

that it is less well-developed than in var. obesa. The basal plate

is uniformly less deep than in the latter, broader and more

rectangular. Very little difference can be noticed in the ad-

laterals. A curious abnormality was to be observed in f..

Commencing at the sixteenth transverse row, a short series of

about three rows of teeth is dwarfed and crowded closely together.

Text-figure 6.

——_s

Cochlitoma zebra vay. fulgurata.

Radula; median and admedian teeth.

The pharynx.—Four main pharyngeal retractors pass through

the subcerebral orifice and spread out in a broad fan-shaped

fascia which, as in var. obesa, has a semi-lunar insertion on the

postero-ventral surface of the pharynx. There is a short ceso-

phagus which passes insensibly into the crop. Internally a

greater complication of the internal longitudinal folds, a greater

frequency of transverse ridges, and a greater general thickness

enables us to assume that the crop starts about 10 mm. from the

pharynx. Measured from this point the crops of f, and f, are

39+3 and 26-3 mm. in length respectively.

The salivary glands (text-figs. 7 & 8) differ extensively in

f,andf,. In general they correspond to those described above.

The crop passes into the anterior part of the stomach, in f,

abruptly, in f, gradually. The stomach exhibits considerable

differences in f, and f,. Thus in f, the shape is piriform and

in f, irregular. But there is no character which differentiates

these forms from var. obesa.

The relation between the anterior and posterior parts of the

stomach (or stomach and cecum) has not been commented upon

at any length save in a general way by Wiegmann (1898). The

stomach as a whole is bent on itself, and the apex of the bend is

260 MR. G. C. ROBSON ON THE

principally occupied by the muscular portion. The floor of the

latter shows a distinct ridge which separates the entrance of the

fore gut from the intestinal pylorus. The longitudinal muscular

folds of the stomach cross this ridge at right angles. In both

forms (and in obesa as well) the folds begin abruptly on the

anterior side of the ridge, cross it and pass down the intestine

for a short way. Very characteristic is a strongly developed

Text-figure 7. Text-figure 8.

—->.

Text-fig. 7.—Cochlitoma zebra var. fulgurata.

Salivary glands (fz) (semi-diagrammatic).

Text-fig. 8.—Cochlitoma zebra var. fulgurata.

Salivary glands (f,) (semi-diagrammatic).

ridge commencing close to the posterior hepatic orifice and

running straight down the proximal part of the intestine.

Possibly an incomplete or degenerate typhlosole, it begins as a

thick pillar and gradually diminishes in size. On one side it is

bent on itself and accompanied by a smaller narrow ridge, which

is separated from it by a narrow space. This space is continued

up to the posterior hepatic orifice and round the well-marked

angle between the ridge on the floor of the muscular portion and

its anterior wall. It passes over on to the anterior side of the

ridge and communicates with the anterior hepatic orifice.

MOLLUSCAN GENUS COCHLITOMA, 261

In these general features there was no difference between the

two varieties or the examples of each.

Neither ¢ntestine nor anus differs from those of var. obesa.

Circulatory and Respiratory System.

The pericardium is slightly less than half the size of the

kidney. In both cases the ventricle is broader and shorter than

in zebra. The pulmonary vein of f, was affected by disease.

That of f, had its secondary branches more evenly distributed

than z, and z,. As in z, the pulmonary vein was a considerable

distance from the ureter.

(3) Renal System.

The kidney measured :—

3 X 13 mm.

Ste

+2 x 14 mm.

Ts Capeuteee: 36

Even with this variation the kidney is in general appear-

ance broader than that of var. obesa. There is no posterior

groove and the posterior appendage tends to lie at right angles

to the main axis of the kidney.

(4) Nervous Systen.

The remarks made above (p. 254) about the connective tissue

sheath and its general significance are relevant in the case of

this variety.

Cerebral Ganglia.—The general shape resembles those of var.

obesa. ‘The position and origin of the main roots are more or

less identical though there are differences of detail. The buccal

ganglia are fairly closely approximated in f,, but widely separated

by a distinctly differentiated commissure in f,. Each buccal

ganglion gives off four antero-lateral and one posterior nerve.

As in zebra, there are three large labial and pharyngeal nerves.

The innermost supplies the lower lip, the next innervates the

sides of the mouth and sends a branch to the anterior tentacle

(a branch found in obesa), while the third gives off lateral and

superior labial and a mandibular branch. Behind and above

there arises a very slender auditory nerve and behind and above

this the rhinophorial nerve.

Between this variety and obesa the most important differences

appear to be the greater apparent differentiation of the “lobule

de sensibilité” and the fact that there are only seven nerves aside

in this variety.

Pedal Ganglia, etc.—The cerebro-pedal connectives are long

and slender. As in var. obesa they enter the pedal ganglia in an

antero-dorsal position. There is the same tiered arrangement

of the pedal nerves, and in f, there are seventeen roots aside and

probably the same number in f,.

There was not nearly so much enclosure of adjacent pedal roots

in common sheaths as in var. obesa.

262 MR. G. C. ROBSON ON THE

Pallial Ganglia, etc.—The cerebro-pallial connectives join the

pallial gangha in exactly the same position in the two varieties, and

in this area the arrangement is very similar to that of var. obesa.

There appears to be no visible distinction between the pallial,

visceral, and abdominal ganglia. There is the same asymmetry

with regard to the pallial roots, there being three on the left

and only one on the right. This asymmetry was observed by

Amaudrut in A. panthera (1886). The three main abdominal

nerves came off a common root in f,.

Text-figure 9,

’ b)

(A OU

NEA

Cochlitoma zebra var. fulgurata.

Genitalia.

hd. = hermaphrodite duct. spv. = spermoviduct. spth. = spermatheca.

vd. = vas deferens. pr. = penis retractor. p. = penis.

(5) Reproductive System.

The gonad les between the second and third whorls. The

hermaphrodite duct, spermoviduct, and albumen gland show no

difference from those of obesa. The vagina is long, measuring

27 inf, and 34 mm.inf,. The spermatheca is spatulate in both

examples and the duct is long (68-2 in f, and 35+.2 mm. in f,).

The vas deferens measures 56-+-2 mm. in f, and 41+2 mm. in f,,

and therefore may be regarded as long, though the anomalous

MOLLUSCAN GENUS COCHLITOMA. 263

condition seen in var. obesa shows that this is not a character to

be relied upon. It enters the penis sheath in f, half-way down

its length. In f, its point of insertion 1s doubtfult With regard

to the actual junction of the vas deferens and penis a general

approximation is made in f, to the condition seen in z,. The

length of the vagina and the form and proportions of the sperma-

theca differentiate both examples dissected from var. obesa.

III. Genetic AFFINITIES AND VARIATION OF COCHLITOMA

ZEBRA VAR. OBESA AND FULGURATA,

The author does not feel justified in altering the position

assigned by Connolly (1912) and others to C. fulgurata Pfr. as a

variety of C. zebra. Although it differs from var. obesa in certain

conchological and anatomical features, conchological intermediates

(British Museum, Zool. Dept.) have been found which cannot be

disregarded.

It is a commonplace among systematists that between two

varieties of a “species’’ or two “closely allied species” there may

be complete agreement in certain respects and difference in

others. It is plain, however, that in the case of the two varieties

of C. zebra, the individuals of a variety do not differ from those

of a closely allied variety in respect of the same characters.

Though they may all differ to the same extent from the typical

form in respect of, for example, two characters, they do not all

show the same degree of difference in their other characters. In

the present instance var. obesa differs from var. fulgurata in

having a truncheon-shaped spermatheca with a short duct

as opposed to a spatulate spermatheca with a long duct, and a

shell with narrow brown and white stripes as opposed to broad

and jagged stripes of brown and yellow. But one individual of

fulgurata agreed with obesa in having a narrow and _ pointed

cervical lobe, while the other obesa had a broad flat one. Of the

three forms that agreed in this respect one (z,) differed from the

two fulgurata and z, in having a straight columella as opposed to

a sinuous one. Again, one example of fulgurata and one of obesa

agreed approximately in the length of the vas deferens, while one

fulgurata agreed with the two obesa in the relation of kidney to

pericardium and differed from its fellow. Again, the same obesa

agreed with fulgurata (cf. above) in respect of the columella, and

yet the relationships of the two obesa are reversed in respect of

the form of the rhachidian tooth of the radula. Two conclusions

may be drawn from the above data :—

(1) That the amount of correlated variation in this case is

very slight.

(2) That the degree of difference or agreement is independent

of the type of organ or structure involved; though gland-

ular tissues appear to be more unstable than the others.

The size and possibly the shape of glands, however, are

264 MR. G. C. ROBSON ON THE

probably conditioned by the amount of secretion which is usually

subject to periodic variation.

The late B. F. Cummings and the present author put forward

a suggestion (‘¢ X” 1914) that internal structures would probably

turn out to be better guides for ordinal and generic characters,

while the external structures register specific and varietal

differences more readily. ‘There would now be grounds for

considering that the internal anatomy is neither more nor less

variable than the external structures.

The small amount of correlated variation in these forms, and

the irregular fashion in which individuals of a variety agree or

differ from those of a closely allied variety, make it very difficult

to distinguish the “species” and “ varieties” of current taxonomy

in this case. The author has come to the same conclusion as

Pilsbry (1912), that ‘a philosophic method of dealing with intra-

specific differentiation is one of the greatest needs of systematic

zoology.” ‘‘Species” and ‘‘variety” appear to the author, and

also apparently to Dr. Pilsbry, to be an association of forms

which in their mutual relationship may be singularly unstable

and loosely associated. It is not merely a question of the

extreme linear or graduated variability with which all naturalists

are familiar. In that case species and varieties are equivalent

for example to the divisions of time into hours and minutes as

convenient, if arbitrary, units. In the present case, however,

the large amount of uncorrelated variation between imdividuals

of a “variety” leaves a very strong suspicion in the mind that

along with linear (or graduated) variability there may exist a

radial * tendency of association which cannot be expressed in

arbitrary subdivisions. If this impression is substantiated by

other evidence it will remain to discover which of the two ten-

dencies corresponds more closely with the results of cytology and

genetics. The author is indebted to Professor Duerden for per-

mission to state that certain conclusions arrived at by him in

the course of work upon the Ostrich are in line with the facts

recorded above.

TV. On THE CLASSIFICATION OF THE ACH AT/NINA AND

THE POSITION OF THE GENUS COCHLITOMA.

Pilsbry (1904) has taken the shell as his basis for arranging the

twelve genera of Achatinine, and his first subdivision into two

main groups depends upon the presence or absence of sculpture

on the embryonic whorls. We cannot, however, disregard the

fact that examination of a number of examples in the British

Museum collection has made it clear that the sculpture of the

embryonic whorls of these forms differs in individuals of the same

* The antithesis between linear and radial variation used in this connection may

be criticized as vague. It is difficnlt, however, to find two terms which express

adequately the fact that an individual of a “‘species” placed in a series presumed

to express descent may, however, exhibit characters which quality it for a place in

another series not necessarily expressing the same line of descent,

MOLLUSCAN GENUS COCHLITOMA, 265

species. Thus, in two examples of Burtoa nilotica the commence-

ment of the sculpture varied from whorl 2; to whorl 33. The

result of this is that it is possible to find examples of species

placed by Pilsbry in his first group (“embryonic whorls seulp-

tured”) with sculpture starting in exactly the same spot as it

does in species placed in the second group. For example, in the

two examples of (. zebra var. fulgurata in the British Museum

the sculpture starts on whorl 23, the same position as in an other-

wise typical Burtoa nilotica. A more serious consequence is that

Pilsbry’s arrangement cuts right across several other potential

arrangements of equal and possibly greater importance. The

character of the penis retractor which is either inserted on the

diaphragm or on the right ocular bend, that of the penis ring-

muscle, whether a complete sheath or annular, and the character

of the admedian teeth, whether having three, two, or one cusps,

are three out of many characters which might be used as a basis

and, if used, would give a different arrangement of the genera

and species.

Tf a classification based upon the shell gives (as it certainly

does) a different result from that based upon the radular

characters, and if the latter yields a different scheme from that

based upon the insertion of the penis retractor, it is plain that

any classification based upon a single character must be arbitrary *.

What is required is a complete knowledge of the variation and

mode of inheritance of all the important structures in each genus

and a classification based upon the greatest measure of agreement

among the various forms.

The anatomy of all the genera and species of this subfamily is

not known. It may be, therefore, argued that a provisional

arrangement based upon the only organ known in the species,

viz. the shell, is justified.

If, however, the object of classification is a correct arrangement

expressed in terms of relationship, it would seem best, in cases

such as the present, to select from all the available data a com-

bination of the more important characters as a basis and to

arrange the species upon it as far as possible, even if some are

left vaguely designated as ‘“‘Achatina.” The selection of a

character (in this case the shell) that will enable every species,

whether known completely or not, to be given a definite place,

involves the production of a scheme that sooner or later has to

be revised, if not abandoned altogether.

With regard to the conchological genus Cochlitoma itself,

only three species are known anatomically, and of these only the

two varieties of zebra are known in any detail. But it is

very doubtful whether these four forms are referable to the

same genus. ©. crawford: differs from the others in the

remarkable insertion of its penis retractor. C. granulata

* WNote—lIt is conceivable that intensive study may show that, within a given

genus, one organ or structure retlects genetic modifications better than others. But

this has yet to be proved,

266 ON THE MOLLUSCAN GENUS COCHLITOMA.

does not fall into line in the character of its penis sheath. These

two forms also appear to be separate from the other forms in

respect of the jaw (v. supra). Conchologically some of the forms

included are indistinguishable from Achatina (s.s.). We there-

fore conclude that the present conception of the genus Cochlitoma

is unsound. Fresh data are, however, necessary before a final

arrangement of the forms involved can be made.

LInterature cited.

Amauprut. 1886.—‘‘Systéme nerveux des quelques Mollusques

pulmonés.” Bull. Soc. Philomath. Paris, (7) x,

in MON

Connouiy. 1912.—“ Reference list of South African non-marine

Mollusea.” Ann. South African Mus. xi. pt. 3,

p. 09.

DesHAves. 1821.—Histoire nat. Moll. Atlas (2) pl. 127.

Firussac. 1821.—Prodromus (Tabl. Syst. Moll.) p. 52.

Nastas. 1894.—‘‘ Recherches sur les centres nerveux des Gasté-

ropodes.” Actes Soc. Linn. Bordeaux, (5) xlvii.

Evade joer NEUE

Premrrer. 1854.—‘‘ Neue Heliceen.” Malak. Blatt., Cassel, 1,

p. 224.

Pirspry. 1904.—Tryon’s Manual of Conchology, Philadelphia,

(2) xvil. passim.

1912.—The same, (2) xxii. p. xxix.

1919.—‘“ Land Mollusks of the Belgian Congo.” Bull.

Am. Mus. N. Hist. New York, xl. art. 1, p. 1.

Prare. 1896.—‘‘ Anatomie des Bulimas ovatus, etc.” Sitz. Ges.

Naturf. Freunde, Berlin, p. 232.

Semper. 1870.—Reisen... Archipel d. Philippinen, Wiesbaden,

Th. ui. Bd. 3. p. 144.

WircMANN. 1898.—(v. Martens & Wiegmann), ‘‘ Mollusken der

Seychellen.” Mitth. Zool. Mus. Berlin, 1. p. 38.

“X ” (Cummings & Robson). 1914.—“'Taxonomy and Evolution.”

American Naturalist, New York, xlviu. p. 369.

PZS2 92122 Arrows allel

VITTY & SEABORNE LTD

LONDON.

BEEREES OF me GENUSsECRINGEOREIA

ON MELOLONTHINE BEEYLES OF THM GENUS ECTINOHOPLIA. 267

16. A Revision of the Melolonthine Beetles of the Genus

Kietinohoplia. By G. J. Arrow, F.Z:8., FHS.

(Plate I.)

[Received January 6, 1921: Read April 5, 1921.]

The genus Hoplia contains an enormous number of species,

often of great beauty but extremely variable and difficult to

discriminate. Attempts to divide it into smaller genera have

not been very satisfactory, but Hetinohoplia, containing a number

of Oriental species of mederately large size, distinguished by the

propygidium not being covered by the elytra, in addition to the

existence (generally) of a tuft of bristles at the extremity of the

elytral suture, 1s fairly well defined. .Some of its species,

however, have not been referred to it, and others have been

described under more than one name. I have therefore

attempted a revision of the genus, and the accompanying key to

the species includes all those at present known to me as members

of it. The striking difference in the coloration of the scales

which distinguishes the two sexes of some of the species has

hitherto not been noticed. Contrary to the general tendency in

insects, the females of some of the species are remarkable for the

occurrence of splendid golden, silvery, or iridescent scales,

replaced by dull scales in the other sex.

Although differences of this kind are not very unusual, it is

almost invariably the male which shows the brighter coloration ;

and it is therefore interesting to observe in the exceptional

instances here brought to notice that the brilliant clothing of

the females is almost confined to parts of the body where it is

concealed in the ordinary position.

In the male of Z. sutwralis (Plate I. fig. 1) the whole of the

scales, of the upper and lower surface alike, are of a uniform pale

non-brilliant blue, in striking contrast to the yellow and golden

scales of the female (Plate I. fig. 3), in which, however, those of

the upper surface are not brilliant, while upon the abdomen they

are of the richest gold and pale metallic blue. Under a high

power of the microscope the difference is seen to be due to a

fundamental difference in the individual scales. The golden scales

of the female ave deeply pigmented and completely smooth, but

the dull blue scales.of the male are devoid of pigment and the

surface is covered with a dense clothing of fine hairs or hairlike

outgrowths, to which the opacity and blue colour are evidently

due. A remarkable change is effected by moisture upon

the opaque scales, which immediately become metallic and

glistening.

E. auriventris Moser has a similar sexual difference, the

scales of the lower surface, pygidium, and propygidium being

golden in the female and opaque orange-coloured in the male.

Proc, Zoou. Soc,—1921, No. XIX. 19

268 MR. G, J. ARROW ON THE MELOLONTHINE

In several of the new species here described the females have the

median part of the body beneath decorated with beautiful silvery

or iridescent scales.

The first to deal with any species of Hetinohoplia was Wollaston,

who bestowed the name Hoplia paive on specimens taken by

Fortune in “N. China.” These specimens actually consisted of

two species, typical specimens of both of which were bought by

the British Museum in 1855 and 1858, labelled ‘‘N. China

(Shanghai).” Wollaston regarded them as belonging to a single

species, but the form which he described as typical, and to which

T restrict his name, is that subsequently called Kctinohoplia

variegata by De Borre, and again described by Reitter as

E. variabilis, by Fair maire as nigrotincta and guttaticollis, and

lastly by Moser as E. tonkinensis. Wollaston’s second form is

K. hispidula Reitter.

E. suturalis De Borre was re-named chrysura by Reitter and,

again, luteostriata by Brenske, while H. diabolica, of Reitter, is a

dark variety of it. All these names were applied to the female

form alone, this being apparently the more numerous sex, for I

have seen 46 specimens as compared with 14 males.

I do not know FZ. davidis and pictipes, of Fairmaire, EL. hutten-

bacheri and mus, of Nonfried, or H. trichota, of Jakowleff, some of

which may not belong to the genus, and these are therefore

omitted from the following Table of Species. H. latesuturata

Fairm., also omitted, belongs to the genus Hoplia.

Synoptical Table of the Species.

(32). Hind claw entire.

(29). Mesonotum not covered with glistening blue scales.

(22). Not very elongate; hind tibia not much dilated.

. Pygidium not brilliant.

. Pronotum without tubercles.

. Pronotum without close erect sete.

. Pronotum without black patches.

(9). Clypeus straight in front

rufipes Mots.

(8). Clypeus evenly rounded.

OMNIA nTkWNe

10 (11). Hind femora densely scaly formosana, sp. n.

11 (10). Hind femora not densely scaly.. gracilipes Lewis.

12 (7). Pronotum decorated with black ‘patches... paive Woll.

13 (6). Pronotum clothed with close erect sete.

14 (15). Median elytral bar angular ........................... hispidula Reitt.

15 (14). Median elytral bar straight . SG reteeceeeeseen) | Oneplagnaca Martin,

16 (5). Pronotum bearing tubercles above.

17 (18). Pronotum with two tubercles ........................ sinuaticollis Moser.

18 (17). Pronotum with four tubercles.

19 (20). Surface pale... 4-tuberculata De Borve.

20 (19). Surface dark inscripta, sp. 0).

21 (4). Pygidium brilliant... dodda aoscocsocne. CUES NOS,

22 (3). i ‘longate, with dilated hind tibize.

23 (28). Py oidium dull-coloured.

24 (27). Hind femora dull.

25 (26). Elytra with three transverse bars vee. SOrOr, Sp. N.

26 (25). Elytra with two transverse bars..................... latipes, sp. n.

27 (24). Hind femora brilliant) y)022..0... 2 ain. te eee. COCA, Sp. me

28 (23). Pygidium bright-coloured : auriventris Moser.

29 (2). Mesonotum clothed with glistening blue scales.

30 (31). Pygidium bearing two black spots .. oculicauda, sp. n,

BEETLES OF THE GENUS ECTINOHOPLIA. 269

31 (80). Pygidium not spotted ...........csessecerseerrseeeeee Susturalis De Borre.

32 (1). Hind claw cleft.

33 (44). Scutellum small.

34 (37). Lower surface entirely dull.

351(86)a eleadidenselyscallysuensneee teeta nee eae eelen tem OU ducta Mots:

36 (35). Head scarcely scaly ...... vistssssssseseee Sulphureiventris Redt.

37 (34). Lower surface more or less brilliant.

38 (43). Pygidium not shining.

39 (40). Pronotum deeply trisuleate ..... 00.0... flavicauda, sp. n.

40 (39). Pronotum feebly trisulcate.

41 (42). Sutural line green, complete ........................ a@ffinis, sp. n.

42 (41). Sutural line blue, incomplete ........................ nitidiventris, sp. n.

413} (BS). ey entalnbuan) JommMIbE NGC. sooo ccobscors coospnoocsaded-oounbeer nitidicauda, sp. nN.

ANAL (BS). SOUTH bonds MEHHERE) ooo boocegacesobods doo canddeoossuy oopocncem | CROCIAAMMGAHely Soo ae

KicrinoHOPLiIA FORMOSANA, Sp. Nn.

Picea, supra et subtus squamis fulvo-cinereis, aliis pallidioribus

interspersis, densissime vestita, capite pedibusque, femoribus

posticis exceptis, squamis opalescentibus minus dense tectis; parum

elongata, depressa, clypeo nudo, nitido, margine antico semicircu-

lari; pronoto convexo, quam longiori paulo latiori, lateribus

medio-leviter angulatis, antice et postice contractis, angulis

anticis acutis, posticis obsoletis, scutello minuto; elytris depla-

natis, postice separatim rotundatis, ad suture finem setis munitis ;

pedibus gracilibus,, tibia antica dentibus tribus approximatis

armata, pedum posticorum ungue integro

3, fronte squamis rotundatis parum nitidis sat crebre vestito ;

Q,fronte setis angustis sat parce vestito, metasterni et

abdominis subtus medio squamis argenteis ornato,

Long. 8°5 mm.; lat. max. 4 mm.

Hab, Formosa.

This is intermediate between H. rufipes Mots. and E. gracilipes

Lewis, and is closely similar to each. It is rather smaller than

Lf. rufipes, and the narrower, more rounded, clypeus will readily

distinguish 1t. From 4. gracilipes it differs by its more dull-

coloured scales, the rather less slender and more densely scaly

legs of the iets. and the glistening silvery-blue and pink scales

at the middle of the sivetierstneranetre and abdomen of the female. In

the latter sex the forehead is clothed with rather seattered narrow

opalescent sete, replaced in the male by more closely-set large

round scales.

EcrINnoHoPLiA INSCRIPTA, sp. n. (Plate I. fig. 12.)

Nigra, vel fusco-brunnea, antennis pedibusque rufis, femoribus

pallidioribus, corpore subtus, propygidio, pygidio, femoribusque

posticis dense squamis oleae: et flavis inter aR vestitis (sed

foeminze corporis medio et femorum posticorum squamis aureo-

viridibus), femoribus anticis et intermediis haud dense aureo-

squamosis, fronte parce aureo-squamoso, pronoti lateribus et

angulis anticis dense, disco laxe, squamis olivaceis et flavidis

ornato, elytrorum marginibus anguste fasciisque tribus undulatis

squamis similibus orn natis ; parum elongata, capite setoso, clypeo

semicirculari, pronoto quam longiori paulo latiori, dorso transverse

19*

270 MR. G. J. ARROW ON THE MELOLONTHINE

4-tuberculato, lateribus arcuatis, medio vix angulatis, angulis

anticis acutis, posticis obsoletis; scutello sat minuto, acuminato ;

elytris quam prothorace multo latioribus, deplanatis; pedibus

gracilibus, posticorum ungue integro :

3, tibia antica elongata, minute tridentata ;

Q , tibia lata, fortiter tridentata.

Long. 8°5-9°5 mm.; lat. max. 4°5 mm.

Hab. 8, Cusna: Fo-kien.

This belongs to the group in which the claw of the hind tarsus

is not cleft at the tip. Itis a little more sienderly built than

most of that group, darker in the colour of the upper surface,

without definite pattern upon the thorax and with the elytral

pattern reduced. Sometimes, as in 1. obducia and suturalis, the

paler scales are entirely wanting above and the surface a uniform

blackish brown. As in #. 4-tuberculata, the pronotum bears

four tubercular elevations in a transverse straight line across the

middle, but that species is quite different in its surface, being

almost covered by pale scales. In “. inscripta these pale scales

are of two similar shades intermixed, but they are scattered upon

the pronotum and consist upon the elytra of a fine marginal line

and three narrow incomplete transverse bands.

EctInoHOPLIA soroR, sp. n. (Plate I. fig. 2.)

Nigra, corpore subtus, propygidio, pygidio, femorisque postici

parte majore dense sulphureo-squamosis, pronoti margine (postice

medio interrupto), linea angusta mediana maculaque parva

utrinque, scutello, elytrorum. margine angusto (basi excepto),

fasciis tribus transversis, prima posthumerali recta, secunda

mediana, tertia anteapicali, biarcuatis, lineisque tribus exiguis

longitudinalibus, femoribus anticis et intermediis sat dense,

capitis fronte, tibiis, tarsis, femorumque posticorum extremita-

tibus disperse argenteo-ceeruleo-squamosis; sat elongata, capite

rugoso, clypeo semicirculari ; pronoto quam longiori paulo

latiori, lateribus medio angulatis, antice et postice valde con-

tractis, angulis anticis acutis, posticis obtusis; scutello minuto ;

elytris quam prothorace latioribus, dorso deplanato, ad suture

finem longe setosis; pygidio parce setoso; pedibus longissimis,

posticorum ungue integro :

¢, tibia antica dentibus tribus minutis, approximatis armatis,

posticis compressis, basi excepto valde dilatis, abdomine subtus

fortiter curvato.

Long. 11 mm.; lat, max. 5 mm.

Hab. ©. Cutna (Fo-kien): Kuatun.

Very closely related to H. latipes and with the legs exactly as

in that species. It is a little less elongated, and the sides of the

prothorax are a little more sharply angulated in the middle. The

sealy clothing of the propygidium, pygidium, lower surface, and

legs is identical, as well as the yellow pattern of the pronotum.

The elytral pattern is almost the same as in &. variegata De

Borre, There are three well-marked transverse bars, the first

BEETLES OF THE GENUS ECTINOHOPLIA. Dil

nearly straight, placed just behind the shoulders, and the other

two curved and angulated at the suture. The inner, outer, and

hind edges are narrowly margined, and there are traces of three

slender lines crossing the transverse bars longitudinally.

I know only a single male.

EcTiNnoHOPLIA LATIPES, sp.n. (Plate I. fig. 5.)

Nigra, corpore subtus, propygidio, pygidio femorisque postici

parte. majori dense sulphureo- squamosis, pronoti margine (hoe

antice et postice medio interrupto), linea mediana, angusta

maculaque utrinqgue parva, scutello, elytrorum margine angusto

(basi interrupto), lineis longitudinalibus discoidalibus tribus, fasciis

duabus transversis, nonnunquam interruptis, internexis, harum

antica posthumerali, recta, postica antemediana, ad suturam

angulata, leviter arcuata; femoribus anticis et intermediis

posticorumque extremitatibus sat dense, capitis fronte, tibiis

atque tarsis disperse argenteo-cceruleo-squamosis; elongata,

angusta, capite rugoso, subnitido, clypeo semicirculari; pronoto

quam longiori parum latiori, lateribus medio angulatis, antice

et postice valde contractis, angulis anticis acutis, posticis

obsoletis ; scutello minuto; elytris quam prothorace latioribus,

elongatis, dorso deplanatis, ad suturee finem setis longis instructis;

pygidio erecte setoso, pedibus longissimis, posticis crassis, horum

ungue integro:

3, abdomine subtus fortiter curvato, pygidio elongato, tibiis

anticis dentibus tribus minutis parum remotis armatis, posticis

elongatis, compressis, basi excepto valde dilatatis.

Long. 11-12°5 mm.; lat. max. 5 mm.

Hab. Tonwxin: Chapa (Rk. Vitalis de Salvaza—May, June,

July), Paklay (Laos) (August).

This is a very slender-bodied species with a pattern similar

to that of ZH. nitidicauda and paive. In its undivided hind

claw and the metallic scales confined to the head and legs 16

shows a closer relationship to the latter. The thoracic pattern

is almost the same, but the yellow bars upon the elytra,

both longitudinal] and transverse, are very slender, and only two

of the latter, both situated before the middle, are recognizable.

The most distinctive feature of the species is found in the very

broad hind tibie, especially in the male. The abdomen in that

sex is very strongly arched and the pygidium longer than it is

broad.

EcriNoHOPLIA TIBIALIS, sp.n. (Plate I. fig. 8.)

Nigra, corpore subtus, propygidio pygidioque dense sulphureo-

squamosis, pronoti linea. integra mediana, margine toto (antice et

postice excepto), puncto utringue, scutello elytr orumque margine

angusto integro(basinonnunquam inter rupto), lineistribustenuibus

longitudinalibus fasciaque transversa interrupta paulo post basin

pallide viridi-squamosis, femoribus sat dense, tibiis capitisque

G2, MR. G. J. ARROW ON THE MELOLONTHINE

fronte disperse aureo-viridi-squamosis; valde elongata, angusta,

capite rugoso, setoso, clypeo semicirculari, pronoto quam longiori

paulo latiori, lateribus medio angulatis, antice et postice valde

contractis, angulis anticis acute productis, posticis obtusis ;

scutello minuto; elytris prothorace vix latioribus, dorso depla-

natis; pedibus longissimis, posticorum ungue integro :

3g, abdomine subtus fortiter arcuato, tibiis anticis dentibus

duabus parum remotis armatis, tertio obsoleto, tibiis posticis

elongatis, compressis, basi excepto valde dilatato.

Long. 9°5 mm.; lat. max. 4 mm.

Hab, ASSAM.

Like #. latipes, soror, and auriventris, this has the hind tibia

dilated beyond its base, especially in the male. It is smaller than

any of those species, of the narrowly elongate form of the first

and almost identical in colouring and pattern, the scales being

quite dull, except upon the head and legs, where they are silvery,

those of the hind tibia included. The longitudinal lines of pale

scales upon the elytra are almost entire, and there is an incomplete

transverse band just behind the base. The scales of the tibiz

and tarsi are much narrower than those of H. latipes.

A second specimen, a female, apparently belonging to the

species, is from the Khasi Hills, but the single male bears no

precise locality. In the female the longitudinal lines of the elytra

are interrupted in the middle.

EcTINOHOPLIA OCULICAUDA, sp. n. (Plate I. fig. 4.)

Nigra, pronoti angulis anticis extremis lineaque angusta,

mediana, scutello toto, elytrorum margine suturali angusto

maculisque minutis externis posthumeralibus, corpore subtus

pedibusque (his parce) squamis metallico-ceeruleis vestitis, ab-

dominis lateribus, propygidio et pygidio lete aureo-squamosis,

hoe basi maculis duabus nigris; elongata, capite rugoso, subnitido,

nudo, clypeo parum transverso, margine arcuato, proncto erecte

sat longe setoso, quam longiori tertia parte Jatiorl, antice et

postice fortiter angustato, angulis anticis acutis, posticis

obtusissimis; scutello minuto; elytris dorso deplanatis, postice

separatim vrotundatis, ad suture finem setis munitis; pedibus

gracilibus, tibiis anticis dentibus tribus distantibus armatis,

tarsorum posticorum ungue integro.

Long. 10 mm.; lat. max. 4-5 mm.

Hab. Tonxin: Chapa (2. Vitalis de Salvaza—May, June). |

T have seen females only. The male will no doubt be found to

be strikingly different, as ig the case with the closely-related

EL. suturalis De Borre (Plate I. figs. 1 & 3), of which also the

female only has hitherto been described. H. oculicauda (female)

is similar in shape and pattern to the latter, but the median

stripe, instead of being opaque yellow, is brilliant pale blue, like

the lower surface, and the beautiful golden or silvery-blue scales

of the pygidium are contrasted with a black eye-like spot on each

side at the base.

BEETLES OF THE GENUS ECTINOHOPLIA. 273

KCTINOHOPLIA FLAVICAUDA, sp. n. (Plate I. fig. 7.)

Nigra, squamis ceeruleis et flavidis ornata, capite nigro, pronoti

linea meiliana duabusque lateralibus obliquis, elytrorum sutura

late, lateribus angustius, squamisque sat numerosis interspersis

ceruleis, opacis, pedibus, metasterni eb abdominis medio squamis

ceruleis nitentibus, pronoto angulis anticis, propygidio, pygidio

corporisque subtus lateribus squamis sulphureis vestitis ; parum

elongata, capite rugoso, setoso, vix squamoso, clypeo semicireulari,

pronoto parum lation! quam. longiori, 4-carinato, carinis externis

obliquis, abbreviatis, lateribus medio angulatis, antice et postice

valde contractis; elytris pronoto multo latioribus, deplanatis,

ad suturve finem setis longis instructis; pedibus modice gracilibus,

posticorum ungue fisso.

Long. 9-10 mm.; lat. max. 5 mm.

Hab. ToNK1: Chapa (LR. Vitalis de Salvaza—May, June, July).

This resembles #. sulphureiventris rather closely, but is smaller,

rather shorter, with the pronotum much more deeply channelled

along the middle and on each side, so that four distinct carine

are formed, the two median ones nearly straight and the two outer

strongly oblique. ‘These are always black, whilst the depressed

parts of the surface contain blue scales. The scutellum is small

and may be with or without blue scales. The elytra are shorter

and broader than those of #. sulphureiventris, with a similar and

equally undefined and variable pattern formed by scales of

rather brighter blue colour, which are densest around the scutellum

and along the inner and outer margins. ‘The exposed part of the

propygidium, the pygidium, and the lower surface of the body are

densely covered, as in the Chinese species, with dull sulphur-

yellow scales, but the entire median part of the metasternum and

abdomen bear instead scales of a beautiful glistening blue, with

which the legs are also adorned, but less densely. There is a

denuded patch on each side of the metasternum. In the male the

front tibize are more elongate and the teeth shorter and much

closer together than in the female.

ECTINOHOPLIA AFFINIS, sp.n. (Plate I. fig. 9.)

Nigra, squamis pallide viridibus supra ornata, pronoti linea

integra mediana margineque laterali, ab angulo postico intus

retrorsum producto, scutello, elytrorum margine integro suturali,

sepe cum margine tenul extenno connexo, viridibus, corporis

subtus lateribus pygidioque dense viridi-squamosis, illius parte

mediana argenteo-cceruleo-squamosa; modice elongata, capite

rugoso, setoso, vix squamoso, clypeo semicirculari, pronoto longi-

tudine sua parum latiori, trisuleato, lateribus antice et postice

contractis, angulis anticis acutis; elytris postice separatim

rotundatis, ad suture finem setis longis instructis; pedibus modice

gracilibus, posticorum ungue fisso.

Long. 7-7°5 mm.; lat. max. 4 mm.

Hab. Assam: Khasi Hills; 8.W. Curya: Yunnan.

274 MR. G. J. ARROW ON THE MELOLONTHINE

Of the same size, shape, and general appearance as #7, nitidiven-

tris, but with the pattern of the upper surface composed of uniform

pale green scales. These form a continuous median line from the

front margin of the pronotum to the hind margin of the elytra,

round which they are continued, and generally unite with a lateral

margin extending from the shoulder. There is also a lateral loop

of similar scales on each side of the pronotum formed by a rather

broad border, continued round the front angle and almost reaching

the hind angle, where it remains open. The pygidium and sides

of the body beneath are densely covered with similar green scales,

and the median part is clothed with silvery-blue scales, also

sprinkled thinly over the legs.

EcTrINOHOPLIA NITIDIVENTRIS, sp. n. (Plate I. fig. 6.)

Nigra, squamis supra ceruleis, subtus viridibus ornata, capite

nigro, pronoti vittis tribus exiguis, elytrorum vitta suturali pos-

tice abbreviata squamisque nonnullis posthumeralibus cceruleis,

opacis, pedibus, metasterni et abdominis medio squamis argenteo-

viridibus, pronoti angulis anticis, propygidio, pygidio corporisque

subtus lateribus squamis pallide viridibus opacis vestitis; modice

elongata, capite rugoso, setoso, vix squamoso, clypeo semicircular,

pronoto longitudine sua parum latiori, trisulcato, lateribus

antice et postice contractis, angulis anticis acutis; elytiis postice

separatim rotundatis, ad suture finem setis longis instructis ;

pedibus modice gracilibus, posticorum ungue fisso.

Long. 7°5-8°5 mm.; lat. max. 4 mm.

Hab. Assam (Khasi Hills): Shillong, 3000-5000 ft. (A. &.

Turner—April, May), Gaubati, 1000-.3000 ft.

Closely related to #. flavicauda and sulphureiwentris. As im

the former, the seales covering the median part of the metasternum

and abdomen are extremely brilliant, but of a pale silvery green

instead of blue. Jt is rather more elongate in shape than that

species, the elytra being narrower, and the three sulci of the

pronotum are less deep. The scales forming the pattern of the

upper surface are of the same bright blue colour but are reduced

in number. Those in the thoracic grooves are very few except at

the base of the median groove, where they form a triangular

patch. The scutellum may or may not bear similar blue scales,

and the elytra have a rather broad, elongate, indefinite patch upon

the suture, beginning at the scutellum but not reaching the hind

margins. There are also scattered scales of the same colour at

the sides behind each shoulder, usually not extending beyond the

middle. The exposed part of the propygidium, the pygidium and

the sides of the body beneath, as well as the front angles of the

pronotum, are densely covered with scales of a very pale green

colour. The front tibize of the male are more slender than those

of the female. Their teeth are less close together than in the

male of #. flavicauda.

BEETLES OF THE GENUS ECTINOHOPLIA. 275

EcrInomcPLiA NITIDICAUDA, sp.n. (Plate I. fig. 10.)

Nigra, squamis flavis vel fulvis et aureis ornata, capite haud

dense aureo-squamoso, clypeo fere denudato, pronoti marginibus

anticis et lateralibus, linea angusta mediana vittaque utrinque

obliqna, plerumque abbreviata vel disrupta, elytri singult

marginibus (antico excepto) lineisque duabus angustis nonnun-

quam fasciis tribus transversis interruptis connexis, fulvisvel flavis,

pedibus, propygidio, pygidio corporisque subtus medio pallide

viridi-aureis, hujus lateribus sulphureis; parum elongata, capite

setoso, clypeo subnitido, semicireulari, pronoto sat brevi, medio

sulcato, lateribus medio obtuse angulatis, antice valde con-

tractis, angulis anticis acutis, posticis obtusis, elytris prothorace

latioribus, haud elongatis, postice separatim rotundatis, ad suture

finem setis longis instructis, pedibus gracilibus, rufis, posticorum

ungue fisso.

Long. 9-10 mm.; lat. max. 5 mm.

Hab. Tonxin: Chapa (2. Vitalis de Salvaza—June).

Similar to #. variegata De Borre and #. obducta Mots., but

easily distinguished by the brilliant silvery-green scales covering

the pygidium and part of the lower surface. The scutellum also

is larger. The scales forming the pattern upon the upper surface

are generally ochre-coloured, but sometimes pale yellow or of both

tints Intermixed. The head bears rather scattered golden scales ;

the pronotum has a narrow median line, a marginal line, which

generally includes the front margin but does not extend past the

hind angles, and an oblique mark on each side, sometimes extend-

ing from the base to the front margin, sometimes abbreviated

in front and sometimes reduced to basal and median spots. The

scutellum is larger than in the two species just mentioned, and

has usually only a few scattered yellow scales. The elytra, in

additior, to a marginal line not including the base, have each two

narrow longitudinal straight lines on the dise extending from

the base nearly to the extremity. There may be also fragments

of three straight equidistant transverse bars connecting the

longitudinal ones. The scales of the propygidium, pygidium,

and lees are entirely pale silvery green in both sexes, and those

of the whole median part of the metasternum and abdomen in

the female. Inthe male the latter region is covered with dull

yellow and brilliant scales intermixed.

EXCTINOHOPLIA SCULELLATA, sp.n. (Plate I. fig. 11.)

Nigra, pedibus antennisque rufis, corpore toto squamis ochraceo-

fulvis atque brunneis dense vestito, corpore supra setis minutis

erectis nigris sat equaliter intersperso, pedibus et capite (femi-

neque pronoto antice) aureo-squamosis ; convexa, clypeo semi-

circulari, pronoto fere duplo latiori quam longiori, supra haud

suleato, lateribus medio sat fortiter angulatis, autice et postice

valde angustato, angulis anticis acutis, posticis obtusis; scutello

276 ON MELOLONTHINE BEETLES OF THE GENUS ECTINOHOPLIA.

magno; elytris postice separatim rotundatis, ad suture finem

haud fasciculatis; pedibus giacilibus, posticorum ungue fisso.

Long. 7°5 mm.; lat. max. 3:5 mm.

Hab. Tonkin: Chapa (2. Vitalis de Salvaza—June).

I refer this species to Metinohoplia because the propygidium is

almost entirely uncovered by the elytra, but the latter are without

the usual tuft of bristles at the sutural angles. The nebulous

pattern of the elytra is unlike that of any species of Hctinohoplia

except H. mdica Moser, but is of a type common to many species

of Hoplia (e18. H. Negantila White, aurantiaca Wat., aurotincta

Fairm., etc.). The ground-colour is an ochreous yellow, some-

times with a tinge alt green, and there are vague markings of a

fawn or pale Bhocolate colour, consisting of a transverse patch

on each elytron behind the aeratlls se a paler, less-defined,

and narrower arcuate patch before the middle. The scutellum,

the outer margins, and a patch at the end of the suture are

also generally darker than the general surface. The lower

surface is densely clothed with yellow scales, and those of the legs

and the clypeus and anterior part of the head are of a pale golden

colour. The species is elongate, with the pronotum distinctly

transverse but narrower than the elytra, its sides strongly con-

tracted before and behind, the angles well-marked, the front

angles acute, the hind obtuse, and the base lobed in the middle.

The scutellum is large and the elytra are little flattened. The three

teeth of the front tibia are strong and sharp, and not placed close

together. The claw of the hind tarsus is cleft. In the female

the scales of the median anterior part of the pronotum are

metallic, as well as the whole of those upon the head.

EXPLANATION OF PLATE If.

Vig. 1. Hetinohoplia suturalis De Borre, male.

2 ss soror, sp. 0.

3 5 suturalis De Borre, female.

4, Me oculicauda, sp. i.

6. op latipes, sp. 0.

6 a nitidiventris, sp. n.

7 3 Jlavicauda, sp. n.

8 op tibialis, sp. n.

9 Ms affinis, sp. 1.

10. oe nitidicauda, sp. n.

Ant 55 scutellata, sp. n.

12. a inscripta, sp. .

THE ANATOMY OF THE 'TONGUES OF THE MAMMALIA. 277

17. The Comparative Anatomy of the Tongues of the

Mammalia.—IIJ. Family 2. Cercopithecide : . with

notes on the comparative physiology of the tongues and

stomachs of the Langurs. By CHARLES F. Sonnraa,

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

[Received January 24, 1921: Read April 5, 1921.]

(Text-figures 16-36.)

ConrTENntTs.

Page

Umbroduction = wast natu sees seca nec ney eal

Genuseresbybesinss era eae heen eee recto 278

EWS CHOHOS soos ncn coocos ceowen gooued ano. cbh denonagsancoaga.| CASE

Genusi@enconathecusien ser entee ses re ee ne ee ec cy o

Genusiacacusm ee eee ene ee eee ao

(Gress). CRADAANTS sp sohovedoaonanne cdoodevogarondgoneacusons GAUL)

Genus iver. op icliccu sme aan een ener eee ee OL

Genus Ba pion orn ee ee cece eeneace alee OS

Summary and Conclusions: Aip-s.0secee ees OAL

Bibliosrap ly, ayiveete cam eeere ee eemece ae TOou

Introduction.

The papers dealing with the tongues of the Cercopithecide

which have already appeared are divisible into two groups. In the

first group are included papers by Mayer (7), Munch (8), Chatin

(4), and Tuckerman (11) on the number and arrangement of

the vallate papille, and the taste-buds. In the second group the

papers give short accounts of the entire tongue, as part of

the description of the anatomy of the whole animal or its

alimentary canal; and the best examples of papers of this kind

are Flower’s ‘Lectures on the Organs of Digestion of the

Mammalia” (5).

Most of these papers have the defect that they state one form

of vallate papillary grouping for each tongue. I found, however,

that all the different forms of vallate papillae occurring through-

out the Cercopithecide will be found in most species if a sufficient

number of examples of each are examined. Consequently,

one should qualify each account by a statement that such and

such a type occurs in the majority of examples of, say, Macacus

rhesus, but other types may appear.

The tongues of the Cercopithecidee possess characters which

unite them to the tongues of the Cebide, and separate them from

those of the Simiide. Their vallate papille, at least in my one

hundred and forty-nine examples, are never arranged in the

Y-form, as in most of the Simiide, but adopt the triangular,

V-arrangement or double-pair type. The orifices of gland ducts

278 DR. C. F. SONNTAG ON THE ANATOMY

are more numerous than in the Simiide, and the lateral organs

possess very different characters. In the Simiide, as was shown

in my last paper (18), the lateral organs consist either of convex

bodies on the sides of the tongue, or of fissures and laminze on

the dorsum ; in the Cercopithecide they consist of rows of fissures

and lamine on the lateral borders.

In all the Cercopithecide the following structures are absent :—

1. Foramen cecum.

2. Apical gland of Nuhn or Blandin.

3. Lytta.

4. Plice fimbriate.

The plicee may, however, be present in the very young animal

(see page 291), and disappear as age advances.

The species of different genera have been arranged in groups,

according to their external characters, by Pocock and others, but

they cannot be so arranged according to the characters of their

tongues. I have arranged my species of Cercopithecus according

to Pocock’s grouping, and it will be seen how the lingual

structures vary, Sometimes considerably, in each of the groups.

As regards classification, I will not go farther than state that

the members of the Althiops-group are the only species of

Cercopithecus in which the vallate papille exhibit the double

pair type in a pure or unmodified form.

In most tongues the conical and fungiform papille exhibit the

usual arrangement in clusters and rows of varying degrees of

obliquity, and the conical papillae have the usual distribution

according to size; their points, as a rule, have the usual

direction. The fungiform papille stretch across the entire dor-

sum or are absent from the centre, thereby forming a dorsal

bounding zone.

Genus PRESBYTES (= SEMNOPITHECUS).

Tur Purpie-racep Lancur (P. cephalopterus).

Habitat : Ceylon.

Measurements.— Total length 3:7 em.; length of the oral part

3cm.; length of the pharyngeal part ‘7 cm.; width between the

lingual attachments of the palato-glossal folds 1-4 em.; width of

the anterior part 1°5 cm.; thickness in the vallate area 1-1 cm.;

thickness of the apex ‘2 cm.

The spatulate tongue has a flat apew; the latter is devoid of a

notch, and is covered by small conical and fungiform papille, but

the latter are not numerous. The lateral borders ave rounded,

and have papille disposed in the usual manner. The fungiform

papille thereon are neither numerous nor prominent. They also

lodge the lateral organs, which are well-developed, thus agreeing

with the description of Boulart and Piliet (1). The structures

mentioned above are absent.

OF THE TONGUES OF THE MAMMALIA, 279

The Circumvallate Papille (text-fig. 16 D).

Three prominent, circular, vallate papille form an isosceles

triangle whose acute vertical angle is directed backwards, All

are bluish-black in colour, granular, and have white secondary

papille at their anterior poles. All the fosse (text-fig. 16 D, e)

and vallums are well-marked, and the posterior vallum is coarsely

nodulated (text-fig. 16 D, 6 and c).

The vallate triangle contains several large conical papille, and

to)

a large fungiform papilla bisects the base.

Text-figure 16.

The tongue of Presbytes cephalopterus.

A. Dorsum; B. Inferior surface ; C. Lateral organs; D. and E. Plan and elevation

of the posterior vallate papilla (a, anterior part; d. main part). Other letters

in the text.

The Fungiform Papille.

The fungiform papille form a dorsal bounding zone on which

they have the usual arrangement in clusters and rows, but they

have not got the usual distribution according to size, .The

280 DR. C. F. SONNTAG ON THE ANATOMY

posterior rows are close together. All are hemispherical, and

their surfaces are smooth or granular.

The Conical Papille.

The arrangement, distribution according to size, and course of

their points follow the usual type, and the interpapillary dorsum

appears as narrow lines. ,

The Lateral Organs (text-fig. 16 C).

The prominent lateral organs extend over the lateral borders

from the dorsum to the inferior surface. The primary sulci are

deep, and the lamin, which possess secondary sulci, appear as

small rods. Each organ is concave towards the lateral vallate

papilla of its own side, for the lamins run in different directions.

The lateral vallate papille (x.v.p.) are level with the posterior

lamine. The anterior laminze run upwards and backwards, the

middle lamine are vertical, and the posterior lamine run upwards

and forwards. The limits of the organs are shown, in text-

figs. 16 A, fand B, f. Hach organ is 1 cm. long. The right one

has 13 lamine and 14 sulci, and the left one has 13 Jamine and

14 sulci. At the anterior end a small fold of mucosa is seen.

The Frenal Lamella (text-fig. 16 B, g).

The triangular lamella has a bifid apex. Its crenated edges

run postero-laterally as far as the middle of the lateral organs.

The median ventral sulcus is shallow and wide, the frenwm is of

moderate length, and the ventral papillary zone is narrow.

Glands and Lymphoid Tissue (text-fig. 16 A, 2).

The whole of the dorsum, behind and at the sides of the

vallate triangle, is covered with white circular or oval areas

containing round or slit-like orifices. These glands increase in

size from before backwards, and their secretions keep the tongue

viscid. ‘They are more developed than in the Cercopitheques,

Macaques, Mangabeys and Baboons. The viscidity of the tongue

remains for a long period, even in preserved specimens.

These large lingual glands are accompanied by an enormous

development of the salivary glands (text-fig. 16 B,/& 4h).

The lingual and salivary glands of the Langurs form a larger

glandular apparatus than in all other Primates. In the case of

the lingual glands ceular inspection and microscopic examination

must both be employed to estimate their area.

The Physiology of the Lingual Glands and Stomach.

To understand the significance of the great development of the

lingual and salivary glands, one must take into account the

nature of the diet, the presence or absence of cheek-pouches, the

degree of complication of the stomach, and the size of the caecum.

The Laneurs have no cheek-pouches, large lingual and salivary

glands, a complex stomach, and a cecum of moderate length,

They eat leaves almost entirely.

OF THE TONGUES OF THE MAMMALTA. 281

Owen (18) considers that the first part of the stomach replaces

the cheek-pouches. JI believe his opinion to be correct and

would add that the leaves, mixed with the copious secretions of

the lingual and salivary glands, lie in the first part of the stomach

till insalivation is completed.

The AnrHroroip Apes have no cheek-pouches, a very high

development of the basal lingual glands, and a simple stomach.

They eat fruits and shoots which are more succulent than the

food of the Langurs, and a complex digestive apparatus is not

So necessary.

Vhe CrrcoprrHeques, MAcAgurs, Basoons and MancABrys

have cheek-pouches, a moderate degree of development of lingual

and salivary glands, and a simple stomach. ‘hey live on a

succulent diet and store food in their cheek-pouches. <A little

food can be removed from time to time, insalivated thoroughly

and swallowed. I would suggest that cheek-pouches are, conse-

quently, part of the digestive apparatus, and not only store-

houses.

In the Uneutata the process of rumination obviates the

necessity for largely-developed lingual glands.

The Turer-rorp Store (Bbradypus tridactylus) resembles the

Langurs in the nature of its diet, but the physiology of

the tongue and stomach is different. The tongue is mainly

mechanical in function, for its gustatory and secretory organs are

small. Prehension is its strongest mechanical action. As the

stomach contains many hard, aliost entire leaves, mastication

and insalivation cannot be very complete. Moreover, there are

no cheek-pouches to prolong the stay of the food in the buccal

area. Consequently, the stomach must soften the leaves for it

gets little assistance from the tongue and salivary glands. It

also cannot share the process of digestion with the cecum. The

ruminating gutter running through the stomach takes on the

regurgitant function of the esophagus of the Ruminantia.

In the Koata (Phascolarctos cinereus) there are large cheek-

pouches, the lingual glands are well-marked, and the salivary

glands, especially the parotids, are large. ‘The stomach is simple

and the cecum enormous. The leaves remain long in the buccal

area, in virtue of the cheek-pouches, and can be thoroughly

moistened and softened. They then pass to the stomach. but the

effect of the peculiar gastric gland has not been worked out.

The stomach, however, does not play such an important part as

in the Langurs and Sloths, for much of its work is taken away

by the enormous caecum.

These remarks indicate that the cheek-pouches are not entirely

storehouses, and show how the functions of the different parts are

interdependent. They also demonstrate how a diet of leaves

requires a complex stomach and smal] cecum, or a simple stomach

and a large complex cecum, for its digestion,

282 DR. C. F. SONNTAG ON THE ANATOMY

Tue Enretitus Lancur (P. entellus).

Habitat: India.

The tongue differs from that of P. cephalopterus in the following

respects :—1. It is thicker. 2. The vallate papille are smaller

and the posterior vallum is more nodulated. 35. The fungiform

papille are more numerous. 4. The conical papille on the base

are larger. In other respects the tongues are similar.

In the Capped Langur (P. pileatus) the frenal lamella is a

broad bilobed plate.

in the Madras Langur (P. priamus) the frenal lamella is

triangular and bifid, and the apical vallate papilla is small.

Genus CoLopus.

Flower (5) has briefly described the tongue of the White-

thighed Guereza (C. vellerosus) as follows :—

“The tongue is long and narrow, with three large cirenm-

vallate papillee forming the corners of a triangle, with the apex

directed backwards; close behind each of the large anterior ones

is a smaller one of the same form. At the lower part of the

frenum is a short, thick, fleshy salivary papilla, constricted at the

base, then dilating, and pointed and bifid at the extremity.”

Flower does not describe the condition of the glands on the

base of the tongue, but pomts out that the salivary glands,

especially the submaxillaries, are very large, and he gives

measurements. As the diet and stomach closely resemble those

of the Langurs, and cheek-pouches are absent in tle Guerezas,

the physiology of the tongue and stomach is similar to those of

the former, which I have described on page 280.

The tongues of the members of the genera Presbytes and

Colobus, as will be shown later, differ from those of the other

Cercopithecide chiefly in the nature of the glands on the base.

Genus CERCOPITHECUS,

The species are here arranged in the groups described by

Pocock (9) :—

THE ALBIGULARIS GROUP.

Species examined: C. preussi; C. albigularis.

Preuss’s CERCOPITHEQUE (C. prewss?).

Habitat : Cameroons.

The conical tongue has the following measwrements :—Total

length 5-4 cm.; length of the oral part 4°6 em.; length of the

pharyngeal part -8 em.; width between the lingual attachments

of the palato-glossal folds 2°5 em,

OF THE TONGUES OF THE MAMMALIA, 283

The apex is rounded, and bears many closely-aggregated conical

papillee, and a few small fungiform papille. It has no notch.

The lateral borders are rounded and have the lateral organs at

their posterior extremities. The structures mentioned in the

introduction are absent.

The Circumvallate Papille (text-figs. 17 A & C).

Three papille form an equilateral triangle with the apex

behind. The two anterior papille are smaller than the posterior

one, the fossz are all well-marked, and the vallums appear as

clear zones. All are prominent, especially the posterior one, and

the surfaces are granular.

Within the vallate triangle there are many large conical

papille, and a fungiform papilla bisects the base.

Several long sulci and Jamine of the lateral organs converge

towards the lateral vallate papillae and end on the outer borders of

the vallums. In no other species of Cercopithecus is this condition

present.

The Fungiform Papille.

On the dorsal bounding fungiform zone the papille have the

usual arrangement, as also on the sides and inferior surface.

They are hemispherical or bossed, and tne surfaces are smooth or

granular, -There is no distribution pattern according to size, and

the posterior rows are close together. One of them bisects the

base of the vallate triangle,

The Conical Papille.

The papille have the usual arrangement in clusters and rows

of different degrees of obliquity.

Appearance.—VThe papille of the oral part are dark grey in

colour. All of them appear, to the naked eye, like small nodules.

Under a pocket lens they appear surrounded by zones of inter-

papillary dorsum (text-fig. 17 D), and have bodies of different

shapes, with or without processes.

Glands and Lymphoid Nodules.

Small nodules are present on the base, but no duct orifices are

visible,

The Lateral Organs (text-fig. 17 C).

The lateral organs begin anteriorly as a number of small ill-

defined fissures and lamine (text-fig.17 C,a). These are followed

by a series of long tapering laminz and sulci (text-fig. 17 C, 6)

converging towards, and ending on, the vallums of the lateral

vallate papille, and they are also convex forwards. ‘These are

succeeded by a number of lamin separated by sulci which are

all parallel to one another (text-fig. 17 C, c). Most of the

laminz are traversed by secondary sulci. The measurements, etc.,

are shown as follows:—The right organ is 1’4 em. long, and has

Proc. Zoou. Soc.—1921, No. XX. 20

284 DR. C. F. SONNTAG ON THE ANATOMY

Th laminee and 12 sulci. The left organ is 1-4 em. long, and has

9 lamine and 10 sulei.

The ventral papillary zone (text-fig. 17 B, d) is wide anteriorly

and narrow posteriorly, and its papille have the usual arrange-

ment.

Text-figure 17.

The tongue of Cercopithecus preussi.

The Frenal Lamella (text-fig. 17 B,e).

The triangular frenal lamella has a bifid apex, and a plain

upper surface. Its edges, which extend postero-laterally as far

back as the level of the middle of the lateral organs, are fused

posteriorly with the under surface of the tongue. They have

pointed processes anteriorly and tubercles posteriorly. In no other

species of Cercopithecus did I find such long, sharp processes.

The median ventral sulcus is narrow and deep anteriorly, but

widens out posteriorly. It lodges a crest derived from the

frenum (text-fig. 17 B, /).

In the fresh tongue the vessels on the base were seen, and they

OF THE TONGUES OF THE MAMMALIA. 285

resembled the condition which I have already described and

figured for MJacacus (10).

The outstanding features are, therefore, the great length of

some of the laming of the lateral organs, and the sharp processes

on the frenal lamella.

Sykes’s Cercopitheque (C. albigularis) has an entire, oval, un-

divided lamella (text-fig. 20 D). Owen (15) described a vallate

triangle.

THE MONA GROUP.

Species examined: 1. C. burnetti (two examples).

2. C. mona ( two examples).

I did not observe any pigmented forms in this group.

BurnNett’s CERCOPITHEQUE ((. burnetii).

Habitat: West Africa.

When the tongues were examined fresh they did not exhibit

the vessels running between the base and the vallatie papille.

Measurements.—Total length 3°2 em.; length of the oral part

2-5 em.; length of the pharyngeal part :7 em.; width between the

lingual attachments of the palato-glossal folds 1:5 em.

The apex and lateral borders ave the same as in C. preussi, and

the same structures are absent.

The Circumvallate Papille (text-fig. 18 D).

The vallate triangle has an acute or obtuse vertical angle.

It lodges many conical and a few fungiform papille.

The three papillz, especially the posterior one, are prominent,

and the latter appears as if it had been pushed through the

vallum, thereby stretching it. All are granular, the fossie are

well-marked, and the vallums look like clear zones.

The Fungiform Papille.

The fungiform papille are the same as in ©. preussi, but ave

more numerous at the sides. None are overlapped by conical

papille.

The Conical Papille.

The arrangement is similar to that of C. prewssi. Most of them

are cylindrical, and each one is surrounded by a zone of inter-

papillary dorsum (text-fig. 18 C). Their surfaces are granular,

and there are few points.

A few orifices of ducts ave present on the base of the tongue.

The Lateral Organs (text-fig. 18 KE, F, & G).

The sulci are narrow, and the laminzx lie on both dorsum and

lateral borders (LB.) All thesulei and lamine are convex

forwards, and secondary grooves traverse the latter. The right

organ is ‘65 cm. long, and has 6 lamine and 7 sulci. The left

20%

286 DR. C. F. SONNTAG ON THE ANATOMY

organ is ‘7 cm. long, and has 5 lamine and 6 sulci. None of

them reach near the vallums of the lateral vallate papillw. False

folds of mucosa (a) may be present. The lamine are flat on

plan (G).

Text-figure 18.

The tongue of Cercopithecus burnetti.

The Frenal Lamella (text-fig. 18 B).

The triangular lamella has an entire apex, and its edges are

devoid of tubercles and processes.

The inferior surface is otherwise similar to that of C. preusst.

When the dorsum is examined by the naked eye it has a finely

nodulated surface as in C. preussi, and the lens reveals how the

interpapillary dorsum forms zones round the conical papille

(text-fig. 18 C).

OF THE TONGUES OF THE MAMMALIA.

“I

Tue Mona CrercorirHEQue (C. mona).

Habitat: West Africa.

The fresh tongues did not reveal the vessels running between

the base and the vallate area.

The following measurements refer to an adult specimen :—

Total length 3-5 em.: length of the oral part 2°9 em.; length of

the pharyngeal part °6 em.; width between the lingual attach-

ments of the palato-glossal folds 1:9 em.

One of my specimens (adult) has an apical notch whence a

median sulcus runs back for 1:05 cm., but the young specimen

has neither. In other respects, however, the apex and lateral

borders have the same characters, and the lateral parts of the

dorsum have the same clusters of fungiform papille as in

C. burnetti.

Text-figure 19.

Cea ees Ole

a te ’

987654321

The tongue of Cercopithecus mona.

The Circumvallate Papille (text-fig. 19 C, D, & E).

The vallate triangle is isosceles. The three papille, especially

the posterior one, are prominent as in C. burnetti. One tongue

has a finely-granular posterior vallum, but the latter is coarsely

nodular in my other specimen (text-fig. 19 D). Within the

vallate area there are more fungiform papille than in C. burnetii.

The anterior papille are oval, but the posterior one is circular,

Miinch (8) described a vallate triangle,

288 DR. C. F. SONNTAG ON THE ANATOMY

The Fungiform Papille (text-fig. 19 A & F, ¥.p.).

On the middle of the oral part of the dorsum the papille are

small, but those on the lateral parts are large and prominent.

The apical cluster contains many large ones. They are hemi-

spherical or umbilicated, and nene are overlapped by conical

papillee.

Text-figure 20.

C. mona. C. mona.

CG. WN BD. (a) E. |

C. nisnes. Cl. albigularts. C. schmidti.

iF na Ge ix

C lalandiy C. patas.

The frenal lamella of the Cercopitheques.

The Conical Papille.

The conical papille have the usual arrangement, but the

posterior rows of the oral part of the dorsum are less oblique

than in most species of Cercopithecus. They are surrounded by

zones of interpapillary dorsum (text-fig. 19 B). They also have

the usual distribution according to size, and the points run in

OF THE TONGUES OF THE MAMMALIA. 289

the usual directions. ‘The papillary bodies are flat or cylindrical,

and the number of points per papilla varies.

Glands and Lymphoid Nodules (text-fig. 19 A, a).

At the sides of the epiglottis there are prominent nodular

masses with a few minute orifices of gland-ducts and pits.

The Lateral Organs (text-fig. 19 F & G).

The lateral organs lie on the lateral borders (.B.) and inferior

surface. They consist of short, broad, furrowed lamine separated

by short, wide sulci. The laminz do not project (text-fig. 19 G).

The right organ is 1:45 em. long, and has 9 lamin and 10 sulci.

The left organ is 1:3 em. long, and has 7 laminz and 8 sulci.

The Frenal Lamella (text-fig. 20 A & B).

The apex is entire and reaches a point 3 mm. from the apex of

the tongue in my adult specimen, and 2 mm. in my young one

It is pointed or rounded. The edges reach as far back as the

middle of the lateral organs.

THE PATAS GROUP.

Species examined: C’. patas; C'. pyrrhonotus.

Tue Paras CERCOPITHEQUE (C. patas).

‘Habitat: West Africa.

Three specimens were examined, and the following measurements

refer to the largest one:—Total length 4-8 cm.; length of the

oral part 4°3em.: length of the pharyngeal part °5 cm. ; width

between the lingual attachments of the palato-glossal folds 19 em.

In two examples the tongue is yellowish-brown in colour, but

in one it is unpigmented. The apex and lateral borders are

similar to those of the preceding forms, except that the apex has

4 median notch asin ©. mena. There is no median dorsal sulcus,

and the structures mentioned on page 278 are absent.

The Cireumvallate Papille (text-fig. 21 A & F to I).

Three papillary patterns are present in my three examples :—

Specimen No. 1.—The three vallate papille form an. isosceles

triangle with an acute vertical angle. All are prominent.

especially the posterior one; they are circular on plan and conical

on elevation, the broad ends of the cones projecting beyond the

vallums (text-fig. 21 A, H, & F). All have granular surfaces.

The vallate area contains several prominent fungiform papille,

and is roughened by conical papille.

Specimen No. 2.—Four vallate papille form a V. On the left

limb, including the posterior papilla, there are three, but the right

limb has only two (text-fig. 21 I). The mid left papilla is small.

Specimen No. 3.—Six papille are arranged in the form of a V,

each limb of which has three elements, The most anterior

290 DR. C. F. SONNTAG ON THE ANATOMY

papilla of each limb is large, and the middle one is small. The

left posterior papilla is large, and the right one is small, but

both stand on the summit of a common elevation (text-fig. 21 G).

Text-figure 21.

« afade¥ a ede ¥eynccy L

= J.

The tongue of Cercopithecus patas.

B. Ventral surface of an adult specimen; C. Ventral surface of

a young specimen. Other letters in text.

This specimen is an example of a double-pair type greatly modified

by the addition of small papille,

OF THE TONGUES OF THE MAMMALIA, 291

The Fungiform Papille.

The fungiform papille stretch right across the dorsum, but

have the usual arrangement in clusters and rows.

All of them have granular surfaces and are hemispherical,

bossed or collared. Those which are included within the vallate

area may be mistaken for vallate papille, but a careful examina-

tion reveals the absence of fossee and vallums.

There is a variable amount of overlapping or concealment by

long conical papille (text-fig. 21 M, 2), especially in the case of

the fungiform papille within the vallate triangle where the

conical papille form prominent processes.

The Conical Papille (text-fig. 21 L & M).

The conical papille have the usual arrangement in clusters and

rows, and their disposition according to size takes the usual form.

Their points, however, face in all directions.

Most of them are filiform and make the dorsum shaggy, but

not to such an extent as in C. ethiops. The points form rows,

and between them one can see strips of interpapillary dorsum

(text-fig.21 D&E). The number of points to each papilla

varies greatly from one to a brushwork.

The Lateral Organs (text-fig. 21 J, a, b, c, & K).

The appearance presented by the lateral organs differs greatly

from those of C. preussi and C. burnetti. The sulci are wide, so

the lamin, which are small and rounded, appear like a row of

small oval bodies. Each lamina is traversed from without

inwards by a broad, shallow, secondary suleus, and a row of

conical papilla runs backwards and inwards across the dorsum

from the inner end of each ridge. The measurements, ete., are

given as follows :—

Organ. Length. Lamine. Sulci.

Ei oiiGe = saeco i2oreme ahd) 11

Wein series: EBS 9 10

The organs may appear as straight rows or curved lines (text-

fig. 21 J,b &c).

The Frenal Lamella (text-fig. 21 B& C; 20 G).

The apex is bifid, and the halves taper from a broad base. The

plain edges run postero-laterally to the level of the middle of the

lateral organs. ‘The points are rounded or sharp, and may or may

not be equal in size.

Plice Fimbriate (text-fig. 21 B & Ca).

Plicee are absent in all my adult specimens, but are present in

the young one, so they may disappear as age advances. ‘They begin

anteriorly at the level of the apex of the free part of the frenal

lamella, and run postero-laterally for 1°8 em. The mucosa

292 DR. G. F. SONNTAG ON THE ANATOMY

between them is more smooth and polished than that of the rest

of the inferior surface of the tongue.

The median ventral sulcus contains a median crest, which,

however, is narrow.

Tuer Nisnas CercopirHEque (CO. pyrrhonotus).

Habitat: Sudan.

Mr. Pocock’s sketch shows an oval lamella with two apical points

(text-fig. 20 G). It is very similar to that of C. patas.

THE AMTHIOPS GROUP.

Species examined: C. ethiops; C. sabeus; C. tantalus ;

C. rufoviridis.

The only pigmented specimen observed is my young one of

C. tantalus.

THe ABYSSINIAN CERCOPITHEQUE (C. ethiops).

Habitat: Sudan.

Measurements.—Total length 4°3 em.; length of the oral part

3°5cm.; length of the pharyngeal part -8 cm.; greatest width

1:8 cm. These figures refer to the largest of five examples.

The apex has no mesial notch, no dorsal sulcus is present,

and the structures mentioned on page 278 are absent. The whole

dorsum is very rough.

The Circumvallate Papille.

Three small plane or umbilicated papille are arranged in the

form of an isosceles triangle with an acute vertical angle. They

are surrounded by prominent conical papillae which may conceal

the fossee and vallums, or be a little distance from them. The

two-pair type of vallate papillee is rare (1 in 5).

The Fungiform Papille.

These stretch right across the dorsum and have the usual

arrangement, but many are concealed by the long conical papille

(text-fig. 22 C). All are hemispherical, smooth and polished, and

some have a small central pit.

The Conical Papille.

The arrangement cannot be made out easily as their long and

numerous points conceal the papillary bodies and interpapillary

dorsum. Their distribution according to size is typical, however.

All are filiform and have one or more points.

Lymphoid Nodules.

There are many large nodules, and several large duct orifices

are visible in some specimens,

OF THE TONGUES OF THE MAMMALIA. 293

The Lateral Organs.

The sulci are wide, but not to such an extent as in C. patas, so

the lateral organs appear to be composed of a row of small bodies

each of which is furrowed. These are not, however, so prominent

as in O.. patas.

The measurements, etc., are given as follows :—

Organ. Length. hidges. Sulci.

Lesa a ednnss 1-5 cm, 9 10

ehts sien Qh 8 9

Text-figure 22.

The tongue of Cercopithecus ethiops.

The Frenal Lamella (text-fig. 22 B).

The triangular frenal lamella has an entire apex, and smooth

plain edges which are free throughout their entire length.

The Median Ventral Sulcus.

The sulcus is short and opens into a wide triangular space. It

lodges « small median crest.

THE GREEN CERCOPITHEQUE (C. sabeus).

Habitat: Sierra Leone.

Several authors have described the patterns exhibited by the

vallate papille. Mayer (7) described four pairs of papille, and

Miinch (8) recorded the two-pair form; he also mentions cases in

which there were three papille with an additional small papilla

on each limb of the V. Im six cases I observed the two-pair

type.

Se of my largest specimen :—-Total length 4:9 em. ;

length of the oral part 3°9 cem.; length of the pharyngeal part

294 DR. C. F. SONNTAG ON THE ANATOMY

‘lem.; width between the lingual attachments of the palato-

glossal folds 2 em.

The apex has a mesial notch, and it and the lateral borders are

covered with fungiform papille which are more numerous and

prominent than in the species already described. The structures

mentioned on page 278 are absent.

The Circumvallate Papille (text-fig. 23 HE & F).

Four vallate papille are arranged in two pairs—anterior and

posterior. The former correspond to the lateral papille of the

triangular type, and the latter occupy the position of the posterior

papilla. The posterior papilla are close together (text-fig. 23 K).

Text-figure 23.

The tongue of Cercopithecus sabeus.

The anterior papille are surrounded by clear, smooth vallums.

The posterior vallums are similar to the anterior ones, or are very

prominent and crowded with conical papille (text-fig. 23 F).

The vallate area contains many conical and a few fungiform

papille. The former vary greatly in size, and they were very

large in a young specimen which I examined.

The papille vary greatly in their degree of projection (text-

fig. 23 BH), hee

OF THE TONGUES OF THE MAMMALIA, 295

The Fungiform Papille (text-fig. 23 G).

Fungiform papille are absent from the centre of the dorsum,

and do not form a cluster in front of the vallate papille. In this

respect the tongue resembles those of ©. preuwssi, C. burnetti, and

C. mona, and differs from that of C. wthiops. They form, there-

fore, a papillary zone on which they have the usual arrangement.

Some are smooth and glistening, others are granular, and all are

hemispherical or cylindrical in shape.

The Conical Papille (text-fig. 23 H).

Conical papille make the entire dorsum rough, but not to such

a degree as in C. patas and C. ethiops. They have the usual

arrangement in clusters and rows, and the direction in which

their points run is typical. Between the rows there are strips of

interpapillary dorsum.

None of them overlap the fungiform or vallate papille.

When they are examined through a lens it is seen how the

papille vary greatly in shape and appear flattened out. The

surface may be plane or excavated (text-fig. 23 H), and one or

more processes are present.

The Lateral Organs (text-fig. 23 D).

The lateral organs consist of a series of short laminz and sulci

convex forwards. Hach ridge is bisected by a longitudinal

sulcus. The measurements, lamine, and sulci are shown as

follows :—

Organ. Length. Lamine. Sulci.

Rieibies er 1-6 cm. 8 9

lhe hipaa sse -- ions 5 6

The ridges decrease in size from before backwards, and the

lateral vallate papille are level with the posterior lamine.

Lymphoid Nodules and Glands (text-fig. 23 A, a).

Prominent lymphoid nodules are present on the base of the

tongue, and a number of orifices are visible.

The Frenal Lamella (text-figs. 23 Ba & Ca).

The frenal lamella is triangular and entire or bifid, being divided

in the latter (text-fig. 23 B,a) into large left and small right

parts. The edges are devoid of processes and tubercles and

extend almost to the posterior fissures and lamine of the lateral

organs.

The Frenwm.

The frenum occupies the whole of the median ventral sulcus

as a ridge, and the part stretching from the suleus to the upper

surface of the frenal lamella is very short, as in all Cereopithecide.

Plice fimbiiate are absent, even in the young specimen. The

animal was older, however, than my young C. patas in which

plicee are present.

296 DR. C. F. SONNTAG ON THE ANATOMY

Tue TANTALUS CERCOPITHEQUE (C. tantalus).

Habitat: Nigeria.

Measurements.—Total length 4°6 em.; length of the oral part

4 em.; length of the pharyngeal part °6 em.; width between the

lingua! attachments of the palato-glossal folds 2 cm.

The apex and lateral borders are the same as in C. sabeus, and

there is no median dorsal sulcus.

Phe Circumvallate Papille.

Four vallate papille are arranged in the two-pair type, but the

elements of the posterior pair are more widely separated than in

O. sabeus. ‘The left papille are oval and all the others are

circular on plan. In some specimens both posterior papille are

circular (text-fig. 24 KX).

The vallums of the anterior pair are clear zones, or have the

appearance of nodulated and granular bands (text-fig. 24 C), and

those of the posterior papille are raised, as in C. sabeus (text-

fig.24.D), but are not so thickly crowded with conical papille.

All the papille project beyond the vallums, and they are conical

on elevation, with the broad ends free (text-fig. 24 L).

The Fungiform Papille (text-fig. 24 K).

The fungiform papille are placed on a zone round the anterior

two-thirds of the dorsum, and they are arranged thereon in the

typical manner. On the lateral borders they form a row of

prominent bodies. ‘hey have, therefore, the same appearances

as in C. sabeus.

They are hemispherical, cylindrical or lobulated, and their

surfaces are smooth or granular.

The Conical Papille (text-fig. 24 F & G).

The conical papille have the usual arrangement, the common

distribution according to size, and the typical mode of direction

of the points. Through the lens they appear very similar to those

of O. sabeus, but the additional forms are shown in text-fig.

24F &G. Between the different rows the strips of ijnter-

papillary dorsum are visible.

Lymphoid Follicles.

Several lymphoid follicles and duct orifices can be seen en the

base of the tongue.

The Lateral Organs (text-fig. 24 H).

The lateral organs resemble those of C. preussi. They consist

of a series of lamine and sulci running from behind forwards

and inwards, and each of the former is traversed by a secondary

sulcus. The whole organ diminishes in size from before back-

wards. Behind them are folds of the mucosa. The lamin do

OF THE TONGUES OF THE MAMMALIA. 297

not touch the lateral vallate papille as in OC. preussi, but are

separated from them by a papillary zone.

The measurements, ridges and lamine are shown as follows :—

The right organ is 1 cm. long, and has 5 lainine and 9 sulci.

The left organ is 1:2 em. long, and has 8 lamine and 6 sulci.

In a second specimen there are 9 lamine and 10 sulci in each

organ.

Text-figure 24,

The tongue of Cercopithecus tantalus.

The Frenal Lamella.

The triangular freval lamella has a bifid apex and crenated

edges running backwards as far as the lateral organs.

No plice fimbriate ave present.

298 DR. C. F. SONNTAG ON THE ANATOMY

THE MozAMBIQUE CERCOPITHEQUE (C. rufoviridis).

Habitat: Mozambique.

The tongue, which is widest between the anterior limits of the

lateral organs, has the following measurements :—Total length

4-6 cm.; length of the oral part 3-8 cm.; length of the pharyngeal

part -8 cm.; greatest width 1°8cm. These figures refer to the

larger of two specimens.

The apex and lateral borders are similar to those of C. sabwus

and C. tantalus.

The Circumvallate Papille.

There are two pairs of vallate papillee—anterior and posterior —

and the posterior elements are close together. All are circular,

prominent and granular, the fosse are well-marked, but the

vallums are not easily distinguishable from the surrounding

dorsum. In my second specimen small papille lie between the

four large ones of the double pair.

The Fungiform Papille.

Fungiform papille are absent from the centre of the dorsum, so

they form a papillary bounding zone. The apical cluster is large,

and the oblique posterior rows are close together. The former

set contains large papille. On the inferior surface the apical

papillz form a prominent cluster, but there is a single row of

papille posteriorly.

All are hemispherical, smooth and polished, and none are over-

lapped by conical papille.

The Conical Papille.

The papille have the typical arrangement in rows and clusters.

Most of them have cylindrical bodies surrounded by a zone of

interpapillary dorsum, and they give the tongue a sago-grain

appearance.

The Lateral Organs.

Deep sulci separate short, flat lamine. The right organ is

1 em. long, and has 7 lamine and 8 sulci. The left organ is

1:2 em. long, and has 5 lamine and 6 sulci.

A few orifices of gland ducts and pits are present on the base of

the tongue.

The Frenal Lamella.

The triangular lamella has a bifid apex, and its smooth edges

extend back to the anterior limits of the lateral organs.

The median ventral sulcus is shallow and triangular.

The Vervet Cercopitheque (C. pygerythrus or lalandit) has a

long, narrow, bifid lamella (text-fig. 20 F).

OF THE TONGUES OF THE MAMMALIA, 299

THE DIANA GROUP.

Tuckerman (11) has described the tongue of C. diana and

pointed out that it has a vallate triangle. He states that it has

no frenal lamella, but does not explain whether the tongue was

carefully removed. The lamella is easily cut away unless great

care is taken in removing the tongue.

THH PETAURISTA GROUP.

Minch (8) has pointed out that the tongue of C, petawrista has a

triangular vallate area.

Mr. Pocock’s sketch of the tongue of Schmidt’s Cercopitheque

(C. schmidti) has a conical, or oval, undivided frenal lamella

(text-fig. 20 E).

Summary of the Tongues of the Species of Cercopithecus,

1. The tongues are conical or spatulate.

. Apical notches and median dorsal sulci are variable.

3. The foramen cecum, lytta, and Apical Gland of Blandin

are absent. .

4, Plicee fimbriate are absent in the adult, but may be present

in the young animal, as Deniker described in the Gorilla

and Gibbons.

5, The lateral organs appear as flat lamine and sulci, or as

rows of oval bodies.

6. The vallate papille form a V, T, triangle or double-pair

arrangement, and it must be stated that the type described

in each species 1s the commonest in a large series of

examples which were examined by me or recorded by

others.

7. The frenal lamella has an entire or bifid apex.

8. Fungiform papille form a dorsal bounding zone or stretch

across the whole dorsum.

9. It is not possible to group the Cercopitheques by the nature

of the tongues as one can by their external characters,

and I have arranged them in their zoological series to

show this. Most of the species in the Athiops Group

have a double-pair of vallate papille, but others have the

triangular type, and that is as far as one can go.

10. Some examples of C. patas and C. tantalus are pigmented.

Genus Macacus.

Tue Ruesus Macaque (I, rhesus). —

Habitat: India.

Tuckerman (11) has briefly described the tongue, and most of

his account deals with the gustatory papille. He has described

Proc, Zoou, Soc,—1921, No. X XJ, 21

300 DR. C. F. SONNTAG ON THE ANATOMY

a vallate triangle, but I only observed that form once in fifty-eight

cases. I give the following measurements as those of my largest

specimen :—Total length 5-4 em.; length of the oral part 4-7 cm. 5

length of the pharyngeal part ‘7 cm.; width between the anterior

limits of the lateral organs 2°3 cm.

The rounded apex and the lateral borders are the same as in

C. preussi, and the structures mentioned on page 278 are also

absent. ;

The Circumvallate Papille.

In fifty-eight. examples | observed three papillary patterns

which are described as follows :—

Specimen No. 1 (type present in fifty-six examples) :—There

are two pairs of vallate papille, and the members of the posterior

pair lie close together, but their relative positions differ (text-fig.

95 0, D, G). The fosse are well-marked, and those of the

anterior pair have recesses at their anterior and posterior poles

(text-fig. 25 J). The fosse of the posterior pair have no such

recesses. All the vallums are prominent and granular.

Von Ebner has published an illustration of the histology of

the posterior papille (14).

Within the vallate area there are both conical and fungiform *

papillee.

Specimen No. 2 (type observed only in one case) :—Three

vallate papilla form an isosceles triangle, the posterior papilla of

which is oval. The sides are filled in by ridges of the mucosa.

Within the vallate triangle there is a triangle of fungiform

papille (text-fig. 25 E).

Specimen No. 3 (occurring once) :—Five papilla form a V with

the apex behind. The three terminating papille are large and

prominent, and the middle papille of the limbs are small, but

prominent. All the fossee and vallums are well-marked, and the

vallate area contains V-shaped rows of fungiform papille

(text-fig. 25 F).

The Fungiform Papille (text-fig. 31 A).

The entire oral part of the dorsum is covered by fungiform

papille, but these have the usual arrangement in clusters and

rows. Occasionally, however, they may be absent from the centre

of the dorsum. They are small, but prominent, on the lateral

borders, and form a well-marked cluster behind the apex on the

inferior surface.

Tuckerman (11) has shown that the apical papillae have well-

developed taste-buds.

The Conical Papille (text-fig. 31 B).

The conical papillze have the usual arrangement, but the close

aggregation and mutual compression of the elements somewhat

obscure the pattern. They have one or more points which are

OF THE TONGUES OF THE MAMMALIA. 301

directed in the usual manner. The inter-papillary dorsum forms

strips (text-fig. 25 M & N).

On the base of the tongue the glands and lymphoid nodules

form elevations of different sizes, and a few minute orifices of

ducts and pits are present.

The Lateral Organs (text-fig. 25 K & L).

The laminz and sulci are short and either straight or convex

forwards. The lamin are mostly traversed by secondary sulci,

and the main sulci vary in width. The left organ is 1-1 em. long,

Text-figure 25.

The tongue of Macacus rhesus.

and has 8 lamine and 9 sulci. The right organ is 1:2 em. long,

and has 11 lamine and 12 sulci. At each end of each organ

there are small folds of mucosa.

The Frenal Lamella.

The triangular lamella has a bifid apex, and the upper surface

of the free part is devoid of sulci, The edges, which extend

postero-laterally as far as the anterior limits of the lateral organs,

are plain anteriorly and tuberculated posteriorly.

913

“

302 DR. C. F. SONNTAG ON THE ANATOMY

Toe Common Macaque (IZ. fascicularis).

Habitat: Malay States.

Tuckerman (11) has briefly described the tongue, and the

following details supplement his account. Measurements :—

Total length 4°6 em.; length of the oral part 3-9 cm.; length of

the pharyngeal part ‘7 cm.; width between the anterior

extremities of the lateral organs 1:9 cm. These measurements

refer to the largest of twelve examples.

The apex and lateral borders ave the same as in JZ. rhesus, and

the usual structures are absent.

The variations in the vallate papille are not so numerous as in

M. rhesus, but the fungiform and conical papillae are very similar.

The former are absent from the centre of the oral part of the

dorsum in most cases, whereas that is the occasional form in

M., rhesus.

Text-figure 26.

a mr

The circumvallate papille of Macacus fascicularis.

The Circumvailate Papille (text-fig. 26).

Two types can appear:—l. There are two pairs of papille

whose appearances, and their variations, resemble those of

M. rhesus. 2. There is an anterior pair of papilla, and a

posterior cluster of three which stand either on a plane surface

or on the summit of an elevation. The vallums consist of one or

more rows of nodules, and may contain fungiform papille.

A few minute orifices of ducts and pits are present on the base

of the tongue.

The Lateral Organs.

The short laminz and sulci are arranged in a straight line or

in a curve convex downwards. Many of the lamine have small

secondary sulci. The right organ is *9 em. long, and has

6 lamine and 7 sulci. The left organ is :9 cm. long, and has

5 lamine and 6 sulci.

The frenal lamella is triangular and has a bifid apex.

Tue Bonner Macague (M. sinicus).,

Habitat: Continental India.

Tongues of Bonnet Macaques differ in size and shape. They

may taper from the palato-glossal folds towards both apex and

base, or their widest point may be between the anterior

OF THE TONGUES OF THE MAMMALIA. 303

extremities of the lateral organs. My largest specimen belongs

to the latter cles and has the following measurements :—Total

length 6°6 cm.; length of the oral part 5:4 em.; length of the

pharyngeal part 1:2 em.; greatest width 2°4 em.

The apex is rounded, has. no mesial notch, and bears conical

and fungiform papille, but the latter are neither numerous nor

prominent, The fungiform papillae on the lateral borders and

inferior surface are also insignificant and few in number.

The following Senuchures™ are absent :—mesial dorsal sulcus,

dorsal ridges, foramen cecum, lytta, and Apical Gland of Blandin

or Nuhn.

The Circumvallate Papille.

The vallate papille are arranged in the form of the letter Y or

V, and I subjoin detailed descriptions of the vallate areas of

three specimens.

Text-figure 27.

cont ae S230 © @e,o ®

A ® C.

— ®

wil

li) AWS

)

amu te

The vallate papille and lateral organs of Macacus sinicus.

Specimen No. 1 :—The vallate papille form a V, with the angle

posterior. Hach limb has two papille, so, including the posterior

‘papilla, there are five altogether. Hach papilla is circular and

prominent, its fossa is well-marked, and its vallum appears as a

clear zone. On examination through a lens the papille and

vallums appear granular. The middle papilla of the right limb

is displaced mesially, and there is a fungiform papilla between the

two mesial papille, so a false appearance of a Y is produced

(text-fig. 27 A).

Specimen No. 2:—The papille also form a V, but the limbs

differ greatly. Those of the left limb are close together, but the

mesial papilla of the right limb lies close to the ‘posterior

304 Dk. C. F, SONNTAG ON THE ANATOMY

papilla (text-fig. 27B). All but the posterior papilla are

circular; the fosse are plain and the vallums appear as clear

zones. ‘The posterior papilla is compound, and consists of two

elements included within the same fossa. The right element is

reniform, and the left one is oval. These two elements may

represent a process of fusion of two elements of a posterior pair.

If that be the case we may consider a single vallate papilla as the

ultimate stage.

Specimen No. 3:—Six papille form the letter T. The hori-

zontal limb consists of five elements, and the vertical limb

consists of the middle horizontal papilla and one posterior papilla

(text-fig.27C). All are circular, prominent, and granular.

The Fungiform Papille (text-fig. 31 A).

Although fungiform papille are scanty in the centre of the

dorsum, they cover the entire oral part. They do not form,

therefore, a wide dorsal papillary zone, and they have the usual

arrangement in clusters and rows.

They are hemispherical or globular, and their edges may be

overlapped by conical papille.

On the inferior surface they are clustered round the apex, but

farther back they form a single chain.

The Conical Papille (text-fig. 31 B).

'Vhe conical papille have the usual arrangement in rows and

clusters, and the usual distribution according to size. ‘Those

lying between the posterior vallate papille are large and

well-marked.

They belong to the cylindrical and filiform series and have one

or more points.

Lymphoid Tissue and Glands.

The glands and nodules on the base are well-marked, and

several small orifices are present in front of the epiglottis.

The Lateral Organs.

The descriptions given below of the lateral organs of three

examples correspond respectively to the three specimens whose

vallate papille have already been described.

Specomen No. 1 (text-fig. 27 D):—A series of short laminze

commence on the sides of the tongue, taper on the dorsum and run

towards the lateral vallate papille, and are separated in the latter

situation by wide shallow sulci. Many of the lamine, which are

convex forwards, are traversed by secondary sulci. Some of the

outer borders are rounded, but others merge into the under

surface of the tongue. The right organ is°9 cm. long, and has

13 lamine and 14 sulci. The left organ is 1 cm. long, and has

12 lamine and 13 sulci. At either end there are small folds of

mucosa.

Specimen Vo. 2 (text-fig. 27 K):—The short, rounded, furrowed

lamin are separated by wide primary sulci. The right organ is

OF THE TONGUES OF THE MAMMALIA. 305

1:4 em. long, and has 10 lamine and |i sulci. The left organ is

1:3 em. long, and has 9 lamine and 10 sulci.

Specimen No. 3 (text-fig. 27 F) :—The lamine and sulci are

short and convex forwards, and the latter look like series of

short incisions into the lateral borders of the tongue. The right

organ is *8 em. long, and has 12 lamine and 13 sulci. The left

organ is *7 cm. long, and has 11 lamine and 12 sulci.

The Frenal Lamella.

The frenal lamella is triangular. The apex expands and is

either rounded and entire, or pointed and bifid. The edges

extend postero-laterally almost to the posterior ends of the lateral

organs.

Tue Toque Macagus (I, pileatus).

Habitat. Ceylon.

The spatulate tongue has the following measurements :—Total

length 4:7 cm.; length of the oral part 3°9 cm.; length of the

pharyngeal part *8 em.; width between the lingual attachments

of the palato-glossal folds 1°7 cm.

The apex and lateral borders are the same as in MW. rhesus, and

the same structures are absent.

Text-figure 28.

cb a.

c. 1 t

@ @ \ >) ~ 7) 209 Tr

eo B. C.

The vallate papilla and lateral organs of Macacus pileatus.

The Circumvallate Papille (text-fig. 28 A).

The isosceles vallate triangle has an obtuse vertical angle; the

two anterior papille are small, but the posterior one is large and

oval. The fosse are well-marked, and the vallums are clear,

prominent zones. Within the vallate triangle there are both

conical and fungiform papille.

The conical and fungiform papille have the usual arrangement,

but the latter are large anteriorly and small posteriorly. Large

clusters of fungiform papille are present behind the apex and in

front of the lateral organs on the inferior surface. .

Few orifices of gland ducts ave present on the base of the

tongue.

The frenal lamella has a bifid apex and plain sides.

The Lateral Organs (text-fig. 28 B & C).

The lamin and sulci are all short, and are either straight or

306 DR. GC. F. SONNTAG ON THE ANATOMY

bent with the angles directed forwards. The right organ is

lem. long, and has 8 lamine and 9 sulci. The left organ is

1-1 cm. long, and has 9 lamine and 10 sulci. At the anterior

extremities are fungiform papille (@) and false folds (0), the true

lamine and sulci being shown at c. The organs are flat on

elevation (C).

THe Pic-rAILED Macaque (M/. nemestrinus).

Habitat: East Indies and Malay States.

Measurements.—Total length 8:'4cem.; length of the oral part

7-3 cm.; length of the pharyngeal part 1:1 cm.; width between

the lingual attachments of the palato-glossal folds 2°7 cm.

The apex and lateral borders are the same as in M. rhesus, and

the same structures are absent.

Text-figure 29.

@ x)

ee LOpoLo Lose

The vallate papillee (A) and lateral organs (B) of Macacus nemestrinus.

The Circumvallate Papille (text-fig. 29 A).

Seven papille are arranged in the V-type. All are very pro-

minent, the fosse are clearly defined, and the vallums form well-

marked zones. In this formation my specimen resembles that of

Miinch (8), but differs from Mayer’s example (7) in which there

were four vallate papille. The vallate area contains both conical

and fungiform papillae. In my second specimen the apical papilla

is replaced by three papille deeply sunk within the common

fossa. It is unusual to have the three papille of a cluster deeply

recessed ; they are usually prominent.

The Fungiform Papille (text-fig. 31 A).

The distribution is the same asin M. rhesus, but they are not

so numerous in the centre of the oral part of the dorsum. They

are very numerous behind the apex and in front of the vallate

area. ‘The ventral apical cluster is also large, but the posterior

part of the ventral papillary zone has few papille.

The Conical Papille (text-fig. 31 B).

The arrangement of the papille and inclination of their points

take the usual form.

The papillary bodies are flat, conical or cylindrical, and the

number of points is variable.

A few minute orifices of gland ducis and pits are present:

OF THE TONGUES OF THE MAMMALIA. 307

The Lateral Organs (text-fig. 29 B).

The short,. wide sulci, which run forwards and upwards,

separate flat, furrowed lamine. The right organ is 14 em. long,

and has 11 lamine and 12 sulci. The left organ is 1:2 cm. long,

and has 10 lamine and 11 sulci.

The apex of the triangular /renal lamella is slightly cleft and

rounded.

I have not observed the row of glandular bodies under the

tongue described by John Hunter (16).

THE STUMP-TAILED MAcAQuE (WV. speciosus).

Habitat: Burmah.

Measurements.—Total length 5:8 em.; length of the oral part

4-7 cm.; length of the pharyngeal part 1-1 cm.; greatest width

2°6 cm.

The apex has a mesial notch. The lateral borders have the

usual characters, and the structures mentioned on page 278 are

absent.

Text-figure 30.

@. @

° P wow

The vallate papillze and lateral organs of Macacus speciosus.

The Circumvallate Papille (text-fig. 30 A & B).

Four vallate papille are arranged in the V-type. Including

the posterior papilla, the right limb has three elements, but the

left one has only two. The left anterior papilla is round and

umbilicated, the fossa is clearly marked, and the vallum is flat

and granular. The right anterior papilla is oval, granular, and

recessed, the fossa is clearly marked, and the vallum appears as

a prominent circular band. ‘The posterior papilla is large and

prominent ; it has a central club-shaped papilla, and is surrounded

by a lobulated vallum.

The Fungiform Papille (text-fig. 31 A).

The fungiform papille are neither numerous nor prominent,

but they stretch right across the dorsum. They have the usual

arrangement in clusters and rows. On the lateral borders and

inferior surface they are discrete. They are hemispherical or

pedunculated, and none are overlapped by conical papille.

The Conical Papille (text-fig. 31 B). ;

Their arrangement and disposition according to size and

direction of their points follow the usual plan, and the inter-

papillary dorsum appears in the form of strips. There is no

great variation in their types.

308 DR. C. F. SONNTAG ON THE ANATOMY

The Lateral Organs (text-fig. 30 C).

The lateral organs appear as rows of short, furrowed, flat

lamine separated by wide sulci. The rows may form a straight

line, or be convex downwards. The right organ is 17 cm. long,

and has 13 lamine and 14 sulci. The ieft organ is 1°5 cm. long,

and has 6 lamine and 7 sulci.

Lymphoid Tissue and Glands.

On the sides of the base there are many small round eminences

with central openings, and a few are situated in front of the

epiglottis. They are more numerous than in any other Macacus

tongue which I examined, but they do not occupy such a large

proportion of the tongue as in the Langurs.

The Frenal Lamella.

The triangular lamella has a bifid apex, the halves of which

taper from a wide base. ‘The edges extend postero-laterally as

far as the middle of the lateral organs. The upper surface of the

free part is smooth.

The ventral papillary border is narrow and has few papille, but

these have tne usual disposition.

The median ventral sulcus lodges a triangular crest which is

larger than in all other Primates.

Toe Barpary Are (J. inwus).

Chatin (4) and Miinch (8) have described the tongue of M.

inuus or ecaudatus, and recorded that it has a vallate triangle.

The latter examined thirteen tongues of Macaque monkeys and

only found the vallate triangle in this species.

Text-figure 31.

The fungiform (A) and conical (B) papille of the Macaques.

Tue Parippmxs Macaque (M. philippinensis).

Mr. R. I. Pocock has lent me the sketch of the frenal lamella

which is simply bifid.

The fungiform papille (text-fig. 31 A) and conical papille

OF THE TONGUES OF THE MAMMALIA. 309

(text-fig. 31 B) present a uniformity of types, as shown in these

figures, in all species.

Summary of the Genus Macacus.

1, The vallate papillae are arranged in the triangular, V-type,

or double-pair formation, and all forms may appear in any

species if sufficient examples of each are studied, as shown

in the introduction to this paper, and in the description of

Macacus rhesus.

2. The pharyngeal part of the tongue is relatively larger in

the genus J/acacus than in the genus Cercopithecus.

3. The glandular orifices are, in most cases, like pin-holes.

They are largest in MJacacus speciosus.

4, The vallate papille may have recesses at their anterior and

posterior poles, and in no other genus of the Cercopithecidze

did I see these.

5, The frenal lamella is more frequently bifid than entire.

Genus CEROCEBUS.

WHIvi-COLLARED MancaaBeEy (C. wthiopicus).

Habitat: West Africa.

Measurements.—Total length 4:8 em.; length of the oral part

3°6em.; length of the pharyngeal part 1-2 cm.; width between the

lingual attachments of the palato-glossal folds 2 em.; thickness

in the vallate area -95 cm.

The apex is rounded, has a delicate median notch, and is

roughened by closely-set conical and fungiform papille, and the

fungiform papille on the lateral borders are prominent and close

together. These lateral papillae are followed by a row of larger

oval bodies which compose the lateral organs.

A fine mesial sulcus runs along the dorsum.

The Circumvallate Papille.

Three large white circular vallate papille are arranged in the

form of an acute-angled isosceles triangle, with the apex posterior.

They offer a marked contrast to the yellow dorsum.

All the papille are smooth, polished and glistening, the fossie

are well marked, and the vallums appear as clear flat zones.

The Fungiform Papille.

The fungiform papille are absent from the centre of the oral

part of the dorsum; on the sides they are arranged in clusters

and rows in the usual manner.

All are hemispherical, smooth and polished, and there is no

overlapping by conical papille.

The fungiform area is bounded posteriorly by a V-shaped row

of papille running backwards and inwards from the anterior

extremities of the lateral organs to the middle of the vallate area.

Ble DR. C. F. SONNTAG ON THE ANATOMY

The Conical Papille.

The conical papille are arranged in the typical form, and the

direction of the points takes the usual course. Those on the base

are large.

The papille haye cylindrical, conical or compressed bodies, and

have one or more points.

At the insertions of the palato-glossal folds there are clusters of

pedunculated papille, but histological examination alone reveals

whether these are conical or fungiform in type.

Text-figure 32.

ITY v.

The tongue of Cercocebus ethiopicus.

Lymphoid Follictes and Ducts.

On the base of the tongue there is on each side a large cluster

of lymphoid nodules, and several large ones have orifices of ducts

or pits. The two masses are separated by the median glosso-

epiglottic fold.

The Lateral Organs (text-fig. 32 C & D).

On each side of the tongue there is a row of lamine and sulci.

The latter are short and wide, and the former are small and

rounded, so the organs look like rows of beads. -The lamine are

smooth, furrowed, or crossed by a ridge, and are single or double

(text-fig. 32 C, Nn).

The measurements, lamin, and sulei are shown as follows :—

Organ. Length. Lamine. Sulci.

Baty ea ult cieidossy 1 Senn: 9 10

2 Dea yaaa elie ae aes 8 9

OF THE TONGUES OF THE MAMMALIA. dll

The Frenal Lamella.

The apex is bifid, and the edges, which are tuberculated

posteriorly, extend as far as the middle of the lateral organs.

The ventral papillary zone is widest in front. It has a single

‘row of fungiform papille.

The structures mentioned on page 278 are absent.

Minch (8) has shown that the tongue of the Sooty Mangabey

(C. fuliginosus) has a vallate triangle.

THe WHITE-CROWNED Mancasey (C’. lunulatus).

Habitat: West Africa.

Measurements.— Total length 5 cm.; length of the oral part

4-4 cm.; length of the pharyngeal part *6 em.; width between

the lingual attachments of the palato-glossal folds 2°3 em.

The apex is flat and devoid of a notch. It is very closely

studded with small fungiform papillae. The lateral borders are

more rounded than in C. ethiopicus, but their fungiform papille

are more numerous, smaller, and arranged in more than one row.

The usual structures are absent (see page 278).

Text-figure 33.

e® @

deleted Ww)

The vallate papille (A) and lateral organs (B) of Cercocebus lunulatus.

The Circumvallate Papille (text-fig. 33 A).

Three papille form an isosceles triangle with an acute apex,

but they are not so large nor prominent as in C. ethiopicus.

Moreover, they are not thrown, into prominence by the contrast

between their white colour and pigment of the dorsum, for

the tongue is devoid of colour. All are circular and vary in

_ protrusion or retraction, the fosse are clearly-cut, but the vallums

are not prominent.

The Fungiform Papille.

The character and distribution are the same as in C. ethiopicus,

but the papille are smaller and more numerous.

All the papillae are hemispherical, and their surfaces are

smooth or granular.

The Conical Papille.

Their arrangement and distribution according to size follow the

usual plan.

The papillary bodies are cylindrical, irregular or tapering, and

312 DR. C. F. SONNTAG ON THE ANATOMY

they have one or more points. The interpapillary dorsum appears

as streaks.

Glands and Lymphoid Tissue.

The base of the tongue is nodulated, and a few orifices of gland-

ducts and pits are present, as in C. ethiopicus.

The Lateral Organs (text-fig. 33 B).

The lateral organs present a very different appearance to those

of OC. ethiopicus, for they do not look like rows of small oval

bodies. They consist of a series of flat, furrowed, lamine

separated by wide primary sulci, and those in the centre of the

organs are larger than those at the sides. The. right organ is

1-1 em, long, and has 10 lamine and 11 sulci. The left organ is

1-1 em. long, and has 8 lamine and 9 sulci. One of the right

laminz has three furrows (text-fig. 33 B, a).

The Frenal Lamella.

The triangular frenal lamella has a bifid apex, and the plain

edges extend back as far as the posterior ends of the lateral

organs.

The ventral papillary zone maintains an even width, and the

median ventral sulcus is narrow and deep.

Genus THEROPITHECUS.

Garrod (6) has pointed out that the Gelada Baboon (7. gelada)

has a triangular vallate area.

Genus Papto (==,CYNOCEPHALUS).

Tue GuiInEA BABoon (P. sphinv).

Habitat: “Africa.

The conical tongue has the following measwrements :-—Total

length 6 cm.; length of the oral part 46 cm.; length of

the pharyngeal part 1-4 em.; width between the lingual attach-

ments of the palato-glossal folds 2°6 em.

The rounded ape has no notch, and is very thickly clustered

with very small fungiform papille. The lateral borders are

rounded and possess many fungiform papille arranged in the

usual manner. At their posterior ends the lateral organs are

situated.

The Circumvallate Papille.

Type—double pair.

The two anterior papille are large, circular, smooth and

prominent, the fosse are clearly defined, and the vallums are

nodulated, Both posterior papille, of which the left one is the

OF THE TONGUES OF THE MAMMALIA. al}

larger, are smooth, oval, and contained within well-marked fosse ;

both stand on a nodulated elevation (text-fig. 34).

Briicher (3) has figured a tongue with six papillw arranged

in the form of a V.

The Fungiform Papille.

The fungiform papilla form a dorsal bounding zone which is

broad, and they have the usual arrangement thereon. They are

very numerous and prominent, and, with the exception of a

cluster of small ones in the mid-line behind, they increase in size

in the usual manner. Those on the lateral borders and inferior

surface are also numerous, prominent, and close together.

They are hemispherical or bossed, their surfaces are mostly

smooth, and none are overlapped by conical papilli (text-fig. 30).

The Conical Papille (text-fig. 35).

The conical papille have the usual distribution and the usual

arrangement according to their size and the direction of their

points. They have flat or tapering bodies and one or more

points. Between them the interpapillary dorsum appears as

strips.

At the sides of the base of the tongue there are a few small

orifices of glandular ducts and pits.

The Lateral Organs (text-fig. 34).

The lateral organs are confined almost entirely to the lateral

borders. They consist of a series of wide sulci separating

lamine which are short, traversed by secondary sulci, and more

or less rounded. The organs, therefore, look like rows of small

oval bodies. The left organ is 1-2 em. long, and has 9 lamine

and 10 sulci. The right organ is 1-3 em. long, and has 8 lamin

and 9 sulci.

The Frenal Lamella (text-fig. 36).

The triangular lamella has a deeply-cleft apex, and the upper

surface of the free anterior part is smooth. The edges, which

extend postero-laterally as far as the levels of the anterior limits

of the lateral organs, bear tubercles and pointed processes

anteriorly, and are undulating posteriorly.

The ventral papillary zone narrows from before backwards, and

is crowded with conical and fungiform papill arranged in the

usual manner.

The median ventral suleus is narrow and deep throughout its

entire length, and a ridge passes into it from the upper surface

of the frenal lamella. No median dorsal sulcus is present,

however, in the fresh tongue.

Mr. R. I. Pocock has lent me a sketch of the frenal lamella in

which the apex is divided into two large diverging processes

(text-fig. 36),

314 DR. C. F. SONNTAG ON THE ANATOMY

THe ARABIAN Bapoon (P. hamadryas).

Habitat; Arabia and Abyssinia.

The spatulate tongue has the following measurements :—Total

length 7:4 cm.; length of the oral part 5-7 cm.; length of the

pharyngeal part 1:7 em.; width between the lingual attachments

of the palato-glossal folds 2°6 cm.; width of the anterior third

2°9 em.

The flat apex has no mesial notch, and its papille are insigni-

ficant. The lateral borders are rounded, and their papille are also

small; their fungiform papille are discrete (separate). Running

back from the apex for 2-4 cm. there is a deep mesial dorsal

sulcus.

Text-figure 34.

e €

& @ @ @

6° @ @ «6

P. sphinx. P sphinx P mormon.

CANS CLORT, Gp

P hamadryeas. P anubis.

The tongues of the Baboons.

The upper figures are vallate papillary patterns, and the lower figures of P. sphina,

P. hamadryas, and P. anubis represent lateral organs.

The Circumvallate Papille (text-fig. 34),

Pour papille form a V with a backwardly-directed apex. The

left limb consists of three papille, including the posterior

one, whereas the right limb has two. ‘The posterior papilla is

large, oval, smooth and glistening, the fossa is clearly cut and has

OF THE TONGUES OF THE MAMMALIA. 315

a posterior straight prolongation; its vallum is prominent,

granular and coarsely nodulated. All the other papille are small

and circular; their bodies are smooth and polished, their fosss

are sharply cut, and their vallums appear as clear zones. The

two small papille on the left limb appear to stand on a common

vallum.

Within the vallate area there are several large fungiform

papilla which can easily be mistaken for the small anterior vallate

papille, but the poeket lens reveals how they have neither fosse

nor vallums. The area is also considerably roughened by conical

papillee.

The Fungiform Papille (text-fig. 35).

The fungiform papille are absent from a small area on the

middle of the dorsum, but they have the usual arrangement

elsewhere. They are all hemispherical or bossed, and are

surrounded, but never concealed, by the conical papille.

The ventral apical cluster contains small elements.

The Conical Papille (text-fig. 35).

The conical papille have the usual arrangement, and the usual

direction for their points. They have flat, cylindrical or tapering

bodies, and the number of points which each possesses varies.

Between the rows the interpapillary dorsum appears as strips.

The papille on the base are prominent.

Glands and Lymphoid Nodules.

Several prominent elevations are formed on the base by

lymphoid nodules, and several orifices are present. These are

minute in front of the epiglottis, but there is, on each side of the

base of the tongue, a row of prominent round or slit-like orifices

stretching along the whole length of the palato-glossal folds.

These are more pronounced than in P. sphinw,

The Lateral Organs (text-fig. 34).

On each lateral border there is a series of short furrowed

lamin separated by wide sulci. The laminz do not project,

however. Some of the more posterior laminze appear as oval

bodies incised by deep secondary sulci, but these must not be

mistaken for glandular pits, from which they are separated by

a short interval. The right organ is 1:9 em. long, and has

11 lamine and 12 sulci. The left organ is 1:8 em. long, and has

12 lamine and 13 sulci.

The Frenal Lamella (text-fig. 36).

The lamella has a rounded, entire, finely crenated apex, and

the edges, which are not prominent, extend almost as far back as

the middle of the lateral organs.

Proc. Zoou, Soc.—1921, No, XXII. 22

316 DR. C. F. SONNTAG ON THE ANATOMY

THe Cuacma Baxpoon (P. porcarius).

Habitat: South Africa.

The tongue is spatulate and has the following measurements: —

Total length 10°3 em.; length of the oral part 8°6 cm.; length of

the pharyngeal part 1:7 em.; width between the lingual attach-

ments of the palato-glossal folds 3:2 em.; width of the anterior

third 4 em.; thickness in the vallate area 2:4 em.

The apex is flat and has a mesial notch whence a mesial dorsal

suleus runs back for 1 em. It is covered by small conical and

fungiform papille which are all visible to the naked eye; they

are not, however, so prominent as in some species of Cercopithecus.

The lateral borders are similar to those of the Cercopitheques

in general both as regards papille and lateral organs. The fungi-

form papiilee are discrete.

The Circumvallate Papille.

Three large prominent vallate papille form an equilateral

triangle (each side=1°5 cm.) with the apex behind. Within

that there is a small vallate V consisting of three very small

papille on the left limb and two on the right. Minch (8) has

described small papille as being remnants of an anterior V-row,

and Briicher (3) has figured them as connecting the large papille

in P. sphinw, but my specimen is more in support of Miinch’s

views than is Briicher’s illustration.

All the large papille are circular and have granular surfaces ;

their fosse are well-marked, and the vallums appear as white

eranular rings raised above the level of the rest of the dorsum.

The small papille are circular and prominent, but their vallums

are not raised,

Within the vallate area there are many conical and a few

fungiform papille which can be mistaken for the small vallate V.

To distinguish the latter I have coloured them black in

text-fig. 35. In my second specimen five large papille form a V.

The Fungiforn Papille.

Fungiform papille cover the whole of the oral part of the

dorsum, but are few in number in the centre. They have the

usual arrangement in clusters and rows, but the anterior cluster

covers a larger area than in any species of Cercopithecus; it

covers the whole of the anterior third of the tongue. On the

lateral borders and inferior surface they form a single row of

discrete elements.

All the papille are hemispherical and a few have central bosses.

Their surfaces are smooth or granular.

The Conical Papille.

The conical papillae extend back on to the epiglottis. They

have the usual arrangement in clusters and rows, but the

OF THE TONGUES OF THE MAMMALIA. BY ¢

directions of the latter are not so clearly marked as in other

Cercopithecide. In their type and arrangement, and their

enormous development on the base of the tongue, they resemble the

conical papille of the Gorilla. They differ in the latter respect

from the conical papille of P. anubis. The points of the papille

are directed backwards, or backwards and inwards in the usual

manner,

Text-figure 35.

Jit ATO

The dorsum, inferior surface, and lateral organs of Papio porcarius, and the

fungiform and conical papillz of the Baboons.

When they are examined through the lens it is seen how they

present a large variety of forms which are shown in text-fig. 35.

Behind the apex, on the dorsum, there are many cylindrical

and a few filiform papille. In the middle third the papillary

bodies are flat and seale-like, and their margins are plain or

prolonged into a variable number of processes. On a zone lateral

and posterior to the large vallate papillae the conical papille are

. 22*

318 DR, C. F. SONNTAG ON THE ANATOMY

of medium size; and their bodies are cylindrical, conical or

fusiform, and are plain or have processes. Behind the zone and

extending on to the epiglottis are large papille which are

spherical, shield-like or tuberose; they may or may not have

processes. On the sides and inferior surface the papille are

small and filiform.

Lymphoid Nodules and Glands.

The base of the tongue has glands and lymphoid nodules of

variable size, and large duct orifices are present in rows on the

elongated lateral masses. It is difficult, without preparing

sections, to determine whether a small nodule is a large papilla

or a lymphoid nodule.

The Lateral Organs.

The lateral organs consist of small fissures and lamine on the

lateral borders and dorsum. All the lamine have small secondary

fissures, and the primary or interlaminar sulci are wide. All the

fissures of the left organ run from below forwards and upwards,

but those of the right lateral organ run forwards, upwards, or

backwards. .The measurements, lamine, and sulci are shown

as follows :—

Organ. Length. Lamine. Sulei.

ARielait pease P:9hcemns i 12

ebb Veoeeeeces: Shes 10 11

The Inferior Papillary Zone.

The papillary bounding zone of the inferior surface is wide

round the apex, but narrows as it is traced posteriorly. It is, in

the former situation, closely dotted by small conical and fungiform

papillee.

Plicee fimbriate are absent in my specimen.

The mesial sulcus is shallow and wide.

The frenwm is long and lax.

The Frenal Lamella.

The lamella is triangular in shape, but the edges sweep round

to the bifid apex, and the dividing fissure is deep. The edges

algo run back to below the middle of the lateral organs and are

crenated or bear processes. The upper surface has a mesial

longitudinal sulcus from which short, horizontal fissures pass out

across the lamella.

Tue AnuBis Bapoon (P. anubis).

Habitat: West Africa.

Measurenents.—Total length 9:2 cm.; length of the oral part

6-6 em.; length of the pharyngeal part 2°6 cm. ; width between

the lingual attachments of the palato-glossal folds 3 em.; width

of the anterior third 3°2 cm.

OF THE TONGUES OF THE MAMMALIA. 319

The apex has a mesial notch whence a mesial dorsal sulcus runs

backwards for 2°3 em. It has many fungiform papille.

The structures mentioned on page 278 are absent.

The Circumvallate Papille (text-fig. 34).

The yallate area consists of two anterior papille, and a mesial

cluster of three small papille standing on a plane surface. The

Text-figure 36.

fj \

Phamadryas. PR. sphinx P sphinx.

P babouin.

Banudis. PR mormon.

The frenal lamellee of the Baboons *.

anterior papille are oval and retracted, the fossee are clearly

defined, and the vallums appear as prominent zones. All papille

are smooth, but the vallums are granular.

* Several of the figures are reproduced from sketches lent to me by, Mr. R. I,

Pocock.

320 DR. C. F. SONNTAG ON THE ANATOMY

The Fungiform Papille (text-fig. 35).

The fungiform papille have the usual arrangement, but the

rows are maintained far forwards. They form a dorsal bounding

zone on which they are numerous, but they are scanty on the

ventral zone.

All are hemispherical, and smooth or granular. None are

overlapped by conical papille.

The Conical Papille (text-fig. 35).

Arrangement.—Behind the apex there is a dense cluster, but

behind this the papille are all in oblique chains. No transverse

rows are present. Between them the interpapillary dorsum

appears in the form of strips.

The papillary bodies are flat, conical, cylindrical or tuberose,

and they have one or more processes which are directed in the

usual manner. They are not, however, arranged in zones as

in P, porcarius.

The Lateral Organs (text-fig. 34).

Small rounded lamelle are separated by wide primary sulci, so

the organs look like rows of small oval bodies. ‘They lie entirely

on the lateral borders. The right organ is 1°3 cm. long, and has

: aes See

8 lamine and 9 sulci. The left organ is 1:2 cm. long, and has

10 lamine and 11 sulci.

The Frenal Lamella (text-fig. 36).

The triangular lamella has a rounded apex with a small fissure,

but no transverse sulci are present on the upper surface. The

edges have small tubercles posteriorly.

A few orifices of ducts and pits are present on the sides of the

base.

The mesial ventral sulcus has a mesial crest, and the frenwm is

long and lax.

Flower (5) states that there is one posterior vallate papilla, but

there may be small ones merging into fungiform papille, and the

latter are large.

Toe MAnpRILL (P. mormon).

Miinch (8) has pointed out how the tongue of P. mormon has

a well-developed lateral organ.

In the specimen which I examined there are five vallate

papillz in the V-formation. The mesial papilla of each limb is

small, the two anterior papille are large and round, and the

posterior papilla, which is the largest of all, is oval (text-fig. 54).

The left lateral organ has seven lamine and eight sulci, and the

‘right organ, whose ridges are all furrowed, has five lamine and

six sulci.

OF THE TONGUES OF THE MAMMALIA. all

The frenal lamella has a bifid apex, and the edges have long

processes (text-fig. 36).

No plice fimbriate are present.

From the descriptions of different species of Papio described

above, one can see that plice fimbriate are absent. Meckel, as

reported by Oppel, found traces, however (17). As the mucosa

of the under surface of the tongue is sometimes very lax in the

Baboons, and thrown into small irregular folds, a false impression

of the plice may be obtained.

SUMMARY.

1. The tongues of the Cercopithecide do not contain many

pigmented forms, and those which are cojoured vary in the dis-

tribution of the pigment. The tongue of Cercopithecus patas may

be yellow or colourless.

9. The conical and fungiform papille, with few exceptions,

exhibit the usual type of arrangement. Only in Cercopithecus

aethiops was it different.

3. The vallate papille form a triangle, V, or double pair, but

the pattern is not characteristic in any genus.

4. The lateral organs appear as rows of sulci and flat lamine, or

as rows of oval or rod-like bodies.

5. The frenal lamella is entire or bifid, even in different

examples of the same species, so it is not of value for purposes of

classitication.

6. The glands on the base of the tongue are largest in the

Langurs, in which they occupy a large area. They are not very

prominent in the Cercopitheques nor in the Macaques ; in the

former the duct orifices are larger than in the latter. In the

Mangabeys and Baboons they form prominent masses with large

orifices on the sides of the base of the tongue. These structures

are, therefore, of the greatest value in classifying the tongues of

the Cercopithecide.

7. Plice fimbriate, lytta, foramen cecum, and Apical Gland

of Nuhn or Blandin are absent in the adult tongue. Plice

fimbriatz may, however, be present in very young tongues.

BIBLIOGRAPHY.

1. Boutarr & Pruurr. “ Note sur Vorgane folie de la langue

des Mammiferes.” Journ. de l’anat.

et de la physiol., T. 21. pp. 387—- .

345. 1885.

Pe eee 3 Ke CR. de Ja Soc. de biol., T. 36 (ser. 8,

T. 1), pp. 626-627. 1884.

3. Bricurr, C. Deut. Zeit. f. Tiermed. u. vergl. Pathol., Bd.

10. pp. 938-111. 1884.

322

THE ANATOMY OF THE TONGUES OF THE MAMMALIA.

Cuatin, J. ‘“‘ Les organes des sens dans la série animale.”

Lecons d’anat. et de la physiol. compareées,

faites 4 la Sorbonne, pp. 726. Paris, 1880.

. Frower, W.H. ‘Lectures on the Comparative Anatomy of

the Organs of Digestion of the Mammalia.”

Medical Times ead Gazette, 1872, vol. i.

p. 451-553.

. Garrop, A. H. P. Z.8. 1879, pp. 451-457.

7. Mayer, F.J.C. Nov. Act. Acad. Leop.-Carol. Nat. Cur.,

T. 20. P. 2. pp. 721-748. 1844.

8. Mtneu, F. ‘Die Topographie der Papillen der Zunge

des Menschen und der Siaugethiere.”

Morphol. Arb. 1896, pp. 605-690.

9. Pocock, R. I. P. Z.8. London, 1907, pp. 677-746.

10. Sonn'rac, C.F. 1 1920, p. 121.

11. TUCKERMAN, In Journal of Morphology, 1890, pp. 152-193.

12. 1891, pp. 188-189.

13. Owen, R. Tr. Z.8 ¥. London, 1833-35, p. 65.

14. von EBNER. Kolliker’s Handbuel der Gewebelehre des

; Menschen, 6. Aufl. 3. Bd. J]. Halfte.

Leipzig, 1899.

15. Owen, R. ‘P.Z.8. London, 1822, p. 20.

16. Hunvrer, Joun. ‘“ Essays and Observations,” edited - by

Richard Owen. London, 1861, vol. 1

page 14,

1%. Opprrn, A. Lehrbuch der verg. mikr. Anat., vol. ii

p- 256.

18. Sonnrac, C. F. P.Z.8. London, 1921, pp. 1-29.

PiZS! #19212 Womness iets

+ at a ° VITTY & SEABORNE LTD,

B.C. knight, det. onset

NEW MOTHS FROM SOUTH EAST BRAZIL:

P:ZSs 2192) Jomescuntginale

Ee Knight dell VITTY & SEABORNE LTD,

ete : Y ce LONDON.

NEW MOTHS FROM SOUTH EAST BRAZIL.

PZS: 1921) Jones api Il.

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NEW MOTHS FROM SOUTH EAST BRAZIL.

ON NEW MOTHS FROM SOUTH-EAST BRAZIL. 32

18. Descriptions of New Moths from South-East Brazil.

By H. Dugtyrietp Jonzs, F.Z.8., FHS.

[Received October 15, 1920: Read February 22, 1921. |

(Plates I.-III.*)

Fam. NocTutID &.

MicrocHRoA PAULATA, sp. n. (PI. I. fig. 17.)

Female.—Palpi rufous brown; legs brown, the fore and mid

tarsi fuscous ringed with ochreous; head and body light brown ;

anal tuft rufous. Fore wings brownish ochreous, suffused with

rufous at base and on medial and terminal areas; dark basal

spots on costa, subcostal nervure, and inner margin ; a wavy dark

antemedial line, dilated on costa and followed by broad whitish

band, which includes the orbicular; orbicular whitish, distally

defined by black; reniform whitish, incurved and defined proxi-

mally and distally by black lunules confluent with a black diffused

fascia in cell and beyond it; a fine wavy postmedial line excurved

beyond cell, incurved from vein 4 to inner margin, followed by

rufous shade; a large fuscous triangular subapical spot on costa

containing three white points; a sinuous whitish subterminal

line incurved from apex to discal fold, angled inwards on the

fold, excurved from discal to submedian fold and bent outwards

to tornus; termina! area rufous: a terminal line of. fuscous

lunules on the interspaces; cilia rufous and fuscous brown. Hind

wings brown ; cilia with ochreous spots opposite the veins.

Expanse 27 mm.

Hab. Sao Paulo, 8.E. Brazil.

MICTOCHROA PALLIDULA, sp.n. (PI. I. fig. 18.)

Male.—Palpi and antenne light brown ; body and wings creamy

white; abdomen slightly irrorated with light red-brown. Fore

wings suffused and lightly irrorated with pale reddish brown ;

the lines very obscure; antemedial line excurved from costa to

submedian fold, then incurved to inner margin; orbicular and

reniform creamy white defined by pale reddish brown ; indica-

tions of medial dark shade; postmedial line sinuous, excurved

from costa to vein 2, incurved from 2 to inner margin; termen

and cilia coneolorous. Hind wings slightly irrorated with pale

reddish brown.

female similar to male.

Expanse: male 21 mm., female 27 mm.

Hab. Castro, Parana, Brazil.

* For explanation of the Plates see p. 356.

324 MR. E. DUKINFIELD JONES ON

BRYOCODIA PAULINA, sp. n. (Pl. I. fig. 3.)

Female.—Palpi reddish brown; pectus white; legs ochreous

brown ; frons, head, and antenne reddish brown ; tegule ochreous

with dark brown bar; thorax ochreous and brown mixed ; abdo-

men ochreous irrorated with fuscous, some fuscous seales on dorsal

tufts. Fore wings reddish brown; double dark subbasal lines

from costa to median nervure, angled outwards on costa; dark

subbasal patches below cell and on inner margin; a dark ante-

medial line, distinctly marked from vein 1 to median nervure,

obscure thence to costa, some rufous before it from 1 to median

nervure ; a median dark shade on costa, outlined on proximal

side by orbicular, a whitish disc below median vein and a prolonged

second dise reaching nearly to postmedial line on submedian

fold; orbicular large, grey, ringed with white and outlined with

fuscous on proximal and distal sides; reniform centre white sur-

rounded with grey, and an outer ring of white outlined with

rufous, some dark scales below it; some rufous in cell and on

discal fold; postmedial line oblique from costa, excurved beyond

cell, sharply angled outwards on vein 6, then incurved to inner

margin, followed by a broad white curved fascia with distal pro-

jections on yeins 3 and 4, a narrow diffused dark line on the

white close to the postmedial ; a very dark fascia beyond the white

from tornus to vein 3; a large brown apical spot with white at

the lower end; termen dark brown with rufous line before it; a

broad white fascia on inner margin from base to postmedial.

Hind wings ochreous suffused with brown.

Expanse 23 mm.

Hab. Sao Paulo, 8.K. Brazil.

TARACHE PARANA, sp. n. (PI. I. figs. 4, 5.)

Male-—Palpi white, third joint brown; pectus white; frons

brown ; head, cheeks, tegule, patagia, and thorax white ; antenne

brown; abdomen dorsally brown, ventrally white, anal tuft

luteous. Fore wings white; a bluish-grey basal spot followed by

two larger ones on costa and subcostal nervure; two still larger

confluent spots on costal and submedian areas and a smaller one

below vein 1; a sinuous bluish-grey antemedial band, broad from

costa to submedian nervure and narrow thence to inner margin,

separated from the above by a narrow white line ; a small bluish-

grey spot on middle of costa; a minute black spot in cell; a

broad dark medial band in and below cell to inner margin,

incurved from cell to vein 1 and followed by similarly shaped and

broader steel-blue space; an irregular dark. fuscous-brown post-

medial band followed by steel-blue to near apex; an irregular,

broken, rufous-brown subterminal band ending in large fuscous

subapical spot on costa; cilia rufous brown and white, a large

purple-brown spot above the tornus and at the end of vein 5.

Hind wings white; a terminal fuscous-brown suffusion, broad at

apex ; cilia white with brown band.

NEW MOTHS FROM SOUTH-EAST BRAZIL. 325

Female.—The basal spots as in male; the antemedial dark

band confluent with the medial steel-blue space above inner

margin; the dark shades on medial area suffused with olivaceous

brown. Hind wings fuscous brown, lighter at the base.

Expanse: male 24 mm., female 22 mm.

Hab. Castro, Parana, Brazil.

ACANTHODICA FRIGIDA, sp. n. (PI. I. fig. 14.)

Female.—Palpi red-brown, with a fine dark longitudinal streak

on outer side; legs red-brown; frons anteriorly red-brown,

frontal tufts brown mixed with greenish grey; back of head

dark red-brown ; antenne brown; tegule light and dark brown

mixed, a few scattered white scales; thorax and abdomen red-

brown; shoulders and patagia white, a few greenish-grey scales

on thorax and inner fringe of patagia. Fore wings light red-

brown ; base white; a small annulate black basal spot; indica-

tions on costa of double subbasal, antemedial, and medial lines ;

a diffused postmedial line, oblique on costa, bent outwards along

vein 11, then strongly excurved and marked by double row of

minute dark points on the veins from 7 to 2; a diffused dark

subterminal shade; a dark brown fascia on discal fold beyond

postmedial ; a large irregular white space extends from the base

to the postmedial line, extending above subcostal nervure from

subbasal to antemedial line, basal half of cell, submedian area to

below vein 1, interrupted by brown triangle at one-third from

base, continued to postmedial as far as vein 7; distal half of cell

brown, containing # small white spot ; a large fuscous spot below

cell at origin of vein 2; a large semicircular ochreous apical spot

defined by fuscous on proximal side ; inner margin brown strigu-

lated with dark fuscous; twe minute white spots above tornus

followed by black points. Hind wings suffused with reddish

fuscous; an elongated white spot with black bar across it at

tornus.

Expanse 40 mm.

Hab. Sao Paulo, 8.E. Brazil.

PHURYS FASCIATA, sp.n. (PI. I. fig. 16.)

Male.—Palpi, legs, and antenne light brown ; head and thorax

brown mixed with lilacine grey ; abdomen ochreous. Fore wings

lilacine grey irrorated with brown ; costa brown; a broad, dark

fuscous streak on submedian interspace from near base to near

subterminal dark band; a subterminal dark band from before

tornus to apex, evenly incurved on proximal and wavy on distal

edge, followed by light brown; a lance-shaped dark streak, with

base at origin of vein 3 and point near the dark band at vein 6;

a terminal brown shade. Hind wings ochreous white with

diffused dark subterminal band.

Expanse 28 mm.

Hab. Castro, Parana, Brazil.

326 MR. E. DUKINFIELD JONES ON

Fam. NOTODONTIDE.

LoBEZA IRRORATA, sp. n. (Pl. I. fig. 13.)

Female.—Palpi fuscous and white; pectus tawny ; femora and

tibiz fuscous, tawny, and white ; tarsi dark fuscous and white ;

head and thorax fuscous, tawny, and white; abdomen black,

some tawny and white on anal segment, ventral surface fuscous,

some tawny hairs on distal ends of segments, anal segment

bright tawny. Fore wings white, heavily irrorated with fuscous

and tawny; an obscure geminate basal line, distinct on costa ; a

wavy geminate antemedial line angled outwards on subcostal and

median nervures; the basal area dark and heavily suffused with

tawny ; a slight tawny medial shade; a white discocellular spot ;

postmedial line well defined, very wavy, dark, diffused, angled

outwards on veins 8 and 7, incurved from 6 to 4, angled outwards

on 3, thence incurved to inner margin, followed by narrow light

shade and a broad fuscous and tawny suffusion to subterminal

line; subterminal line very wavy, with double distal projections

between veins 2 and 4 and 6 and 8; a dark lunular terminal

line; cilia a crenulate line of fuscous and white. Hind wings

fuscous ; cilia white at ends of veins.

Expanse 72 mm.

Hab. Alto da Serra, Santos.

Fam. MELALOPHIDE,

RosEMA PALLIDA, sp.n. (PI. I. fig. 10.)

Female.—Palpi, legs, and antenne ochreous; head green;

tegule reddish ochreous; thorax green; abdomen ochreous.

Fore wings pale green; costa ochreous ; a minute white point on

discocellulars. Hind wings white. Underside white.

Expanse 38 mm.

fab. Castro, Parana, Brazil.

MoresA mona, sp.n. (PI. I: fig. 15.)

Female.—Palpi and legs light brown sprinkled with fuscous ;

frons green; vertex of head and antenne light brown; tegule,

patagia, and thorax green; abdomen light buff. Fore wings

green ; costa ochreous irrorated with fuscous; a large white spot

irrorated with fuscous on discocellulars; a smaller similar spot

above vein 2, and a still smaller one above vein 3; a minute

white point on vein 1 at one-third from base. Hind wings

uniform light buff.

Eixpanse 44 mm.

Hab. Alto da Serra, Santos.

NEW MOTHS FROM SOUTH-EAST BRAZIL,

Fam, GEOMETRIDA.,

Subfam. BoaARMIIN#.

Merrocausta FELINARTA, sp. n. (PI. I. tig. 20.)

Female.—Palpi: 1st joint ochreous buff, 2nd and 3rd purple-

brown; legs ochreous buff speckled with purple-brown; frons

anteriorly ochreous buff, posteriorly purple-brown; vertex of

head purple-brown; patagia and thorax ochreous; a minute

dorsal and two larger subdorsal spots on metathorax ; abdomen

ochreous speckled with purple-brown, Fore wings ochreous

sparsely irrorated with purple-brown, the medial area suffused

with light brown extending to termen between veins Gand 7; a

wavy diffused antemedial line angled outwards on median ner-

yure; an obscure postmedial line, wavy from inner margin to

vein 6, interrupted between 5 and 7, bent inwards from 7 to

costa; a subterminal series of minute dark points; an obscure

brown discocellular bar; termen and cilia reddish brown. Hind

wings ochreous irrorated with purple-brown ; a diffused medial

line in continuation of postmedial of fore wings; terminal area

slightly suffused with reddish brown.

Expanse 27 mm.

Hab. Castro, Parana, Brazil.

APLOGOMPHA CASTRARTA, sp.n. (PI. I. fig. 21.)

Male.—Palpi and legs ochreous mixed with brown ; head,

thorax, and abdomen purplish brown; antenne purplish brown,

the shaft ringed with ochreous. Fore wings purplish brown suf-

fused with fuscous, the medial area lighter than the rest; dark

antemedial, medial, and postmedial bands, the two former nearly

straight, the latter evenly excurved; terminal area darkest,

especially towards apex ; apex lighter; costa crossed by minute

ochreous-orange strigule; a ferruginous discocellular spot sur-

rounded by dark purple-brown confluent with medial band.

Hind wings the same shade as the medial area of fore wings; a

rather broad dark antemedial band; a narrow wavy postmedial

line strongly excurved between the discal and submedian folds ;

an obscure subterminal line more clearly indicated on inner

margin; apical area dark; termen dark; cilia light purple-

brown. Underside bright ochreous orange closely covered with

bright purple-brown strigule ; the bands and lines as on upper

side, but much brighter, terminal area with orange strigulated

spots at costa, Inner margin, and between veins 2 and 4.

Female.—The markings as in the male, but the whole surface

of the wings is heavily suffused with fuscous; cilia somewhat

reddish. Underside suffused with reddish purple-brown.

Expanse: male 16 mm., female 17 mm.

Hab. Castro, Parana; Sao Paulo, §,E. Brazil,

328 : MR. BE. DUKINFIELD JONES ON

APLOGOMPHA FUMARIA, sp.n. (PI. I. fig. 8.)

Male.—Palpi ochreous and fuscous; legs ochreous speckled

with fuscous; antenne shaft fuscous ringed with ochreous,

pectinations fuscous; thorax and abdomen purplish fuscous.

Wings purplish fuscous: fore wings with antemedial, medial,

postmedial, and subterminal dark bands; a yellow bar across

costa beyond antemedial and four beyond medial (in some

specimens there is another bar beyond the postmedial) ; a ferru-

ginous spot on discocellulars; termen dark fuscous; cilia dark

fuscous, tipped with whitish below apex: hind wings, the bars

very obscure; a ferruginous discocellular spot surrounded by

dark fuscous. Underside: fore wings purple fuscous, suffused

with reddish at apex; three bands of ochreous-white strigule,

the medial one broad and the postmedial narrow; the disco-

cellular spot brighter than on upper side: hind wings similar, but

the strigule more evenly scattered over the lower half of the

wing ; discocellular spot as on fore wings.

Expanse 15 mm.

Hab. Castro, Parana; Aragatuba, Sao Paulo, Brazil.

APLOGOMPHA SETINARIA, sp. n. (PI. I. fig. 9.)

Male.—Palpi and legs rufous brown; frons and antenne dark

rufous brown; tegule, patagia, thorax, and abdomen purplish

grey speckled with darker shade. Fore wings glossy purplish grey;

three rather broad sinuous ferruginous lines excurved on upper

half and slightly incurved on lower half of wing; a whitish

streak before the antemedial on costa; the postmedial with

outward points on veins 3, 4, and 6; costa yellow barred with

purplish grey; costal area thickly and rest of wing sparsely

striated and irrorated with whitish. Hind wings the same shade

as the fore wings, a medial and a postmedial line broader and

more diffused than on fore wing; the irrorations very much

scattered.

Expanse 18 mm.

Hab. Castro, Parana, Brazil.

APLOGOMPHA YAPONARIA, sp.n. (PI. I. fig. 7.)

Male.—Head, palpi, pectus, and legs ochreous orange, tarsi

suffused with purple-brown ; antenne, shaft light brown ringed

with purple, the pectinations purple; thorax and abdomen

brownish orange; fore wings bright brownish orange, heavily

suffused with fuscous brown on terminal area; fine dark brown

strigule on costal area; a dark antemedial line broad at costa,

narrow on inner margin, bent outwards on median nervure and

vein 1; postmedial oblique from costa to vein 7, thence wavy

and obscure; an obscure series of subterminal spots; a dark

discocellular streak; cilia dark fuscous brown. Hind wings

brighter than the fore wings; an obscure antemedial line; a

well-defined medial line from inner margin to vein 7 close to

NEW MOLHS FROM SOUTH-EAST BRAZIL. 329

origin; a subterminal line of spots; cilia concolorous excepting

at apex where they are fuscous brown. Underside brighter

than upper, sparsely irrorated with dark brown; a heavy suff usion

of ferruginous brown at apex and tornus ; a large triangular ochre-

ous spot on apical suffusion from below vein 5 to above 6, suffused

with ferruginous on the veins; a well-defined antemedial line on

fore wings, angled outwards on median nervure; a diffused wavy

postmedial line on fore wings continued medially across hind

wings.

Expanse 18 mm.

Hab. Castro, Parana, Brazil.

OPHTHALMOPHORA COLUMBARIA, sp. n. (PI. I. fig. 23.)

Female—Palpi: 1st joint white, 2nd and 3rd buff; pectus

and femora white; tibize and tarsi ochreous; frons bright buff;

vertex of head and tegule white; antenne ochreous; patagia

pale ochreous brown, white on the shoulders ; thorax pale ochreous

brown; abdomen: first segment ochreous brown at base, then

white and posteriorly ochreous; the rest of abdomen ochreous.

Fore wings white, heavily suffused with reddish brown; costal

area, pale ochreous yellow with white below ; outer half of inner

margin white; cilia pale primrose-yellow. Hind wings white; a

very broad band at base the same colour as the fore wings ; a white

band separating this from the rest of the wing, which is suffused

with yellow and thickly irrorated with red-brown ; a postmedial

line of iridescent gold scales from beyond the cell to tornus,

nearly meeting on costa a subterminal line of similar scales ;

terminal area orange; some scattered metallic scales on inner

margin near base; a black spot ringed with white and centred

with iridescent metallic scales between veins 6 and 7; cilia pale

primrose-yellow.

Expanse 24 mm.

Hab. Castro, Parana, Brazil.

CALLURAPTERYX PAULARIA, sp.n. (PI. IT. fig. 15.)

Male.—Palpi fawn-colour with white at the ends of the joints ;

pectus white ; legs ochreous ; frons fawn ; vertex of head white ;

antenne light brown; tegule, patagia, and thorax white ; abdo-

men fawn. Fore wings white ; costa light brown; the lines ight

brown, straight, expanding on costa, basal narrow, ante- and post-

medial geminate, enclosing lighter shade, subterminal narrow,

terminal broad; a brown streak on discocellulars ; cilia a lighter

shade. Hind wings white; a broad postmedial line, geminate,

enclosing lighter shade, angled outwards on vein 6 and above

vein 7; a narrow subterminal line from before tornus to discal

fold; a terminal line from tornus dilating to vein 4, where it

suddenly becomes narrow and is bent upwards and changes to pale

tawny ; apex pale tawny, an irregular metallic silvery-white ring

on the tawny from discal fold to near costa, within this ring there

is a black spot surrounded by white and with a few metallic scales:

330 MR. E. DUKINFIELD JONES ON

in the centre; a large black terminal spot above vein 6 and a

lunular one above 7; cilia pale brown from tornus to vein 4,

white from there to apex.

Expanse 38 mm.

Hab. Sao Paulo, S.E. Brazil.

EARIODES FLAVICILIA, sp. n. (PI. II. fig. 26.)

Male.—Palpi light brown; legs ochreous, the fore- and mid-

tibie shaded with fuscous; antenne brown; frons and vertex of

head yellow; back of head, tegulz, and shoulders yellowish green ;

patagia green; abdomen pale ochreous, some green at base of

dorsum. Fore wings bright yellowish green with a few scattered

black scales ; costa orange at base, then white with narrow yellow

shade below to near apex, where it changes to light brown with

bright rufous in place of the yellow; a series of postmedial bright

rufous spots on veins 2, 3, and 4; cilia yellow. Hind wings

creamy white.

Expanse 23 mm.

Hab. Alto da Serra, Santos

HARIODES BIMACULATA, Sp. N.

Female.—Palpi, frons, and antenne brown; vertex of head and

thorax green; tegule light brown; abdomen ochreous, dorsally

green at base. Fore wings yellowish green; costa white except

at base which is yellow, at the apex there is a reddish streak

below the white; a large pale purplish-brown lenticular spot

irrorated with whitish and red-brown from below vein 2 to above

vein 4; a bar of the same colour from inner margin to submedian

fold at one-third from base; cilia yellowish white. Hind wings

ochreous white. Underside: the lenticular spot on the fore wings

is fuscous.

Expanse 28 mm.

Hab. Castro, Parana, Brazil.

Very close to #. variomaculata Warren.

APICIA STRIGULARIA, sp. n. (PI. IT. fig. 24.)

Male.—Palpi ochreous with some brown scales, third joint

brown ; pectus and legs ochreous; hind femora and tibize with

fuscous spots, spurs ringed with fuscous; frons light ochreous

brown speckled with darker scales ; thorax, patagia, and abdomen

ochreous irrorated with black. Fore wings pale buff irrorated

with black, the basal area suffused with tawny and the terminal

area heavily striated with fuscous ; sub-basal line represented by

a black spot in the ceil; an obscure brown antemedial line suf-

fused with whitish, angled outwards in cell and marked by black

spots on subcostal and median nervures, on vein I, and inner

margin; a minute black discocellular spot; a postmedial band

from middle of inner margin to apex, consisting of a dark line,

light shade, dark line, whitish line, and dark line, the second

NEW MOTHS FROM SOUTH-EAST BRAZIL. 331

dark line punctuated on the veins; a dark suffusion at the apex ;

some strigule on inner half of wing from vein | to inner margin ;

termen dark; cilia light at base, dark at tips. Hind wings

striated over the whole; a broad medial band similar to post-

medial of fore wings; a minute discocellular spot.

Expanse 30 mm,

Hab. Alto da Serra, Santos.

APICIA GEMINIMACULA, sp.n. (PI. IT. fig. 17.)

Male.—Palpi and frons brown; pectus and legs light brown ;

head, antennze, and thorax light brown, two black spots on meta--

thorax ; abdomen light brown, two black distal subdorsal spots

on basal segment, the spots confluent on the following segments.

Wings light brown irrorated with darker brown and _ black.

Fore wings: antemedial line tawny brown from inner margin to

median nervure with dark spot on vein 1 and median, above

median invisible except black dot on subcostal nervure ; a minute

black discocellular spot ; postmedial line tawny brown, slightly

wavy, dark points on the veins; two large subterminal spots on

veins 2 and 3: termen and cilia concolorous, a dark spot at ends

of the veins. Hind wings: a broad diffused tawny medial shade ;

a black discocellular spot; a fine dark postmedial line, diffused

on veins, preceded by diffused tawny shade, followed by light

shade; a subterminal row of small spots on veins 1-3; termen

angled on vein 4; cilia with black spots at ends of veins.

Expanse 25 mm.

Hab. Castro, Parana, Brazil.

CRATOPTERA FENESTRARIA, sp.n. (PI. II. fig. 16.)

Male.—Palpi rufous brown; pectus light brown; legs ochreous

speckled with black, tarsi ringed with black; frons and head

light brown, white between the antenne ; antenne light brown;

tegule, patagia, thorax, and abdomen light brown, the latter

with a few scattered black scales. Fore wings light ochreous

brown, suffused with purplish brown and striated with purplish

brown and black strigule; antemedial line strongly angled out-

wards in cell, preceded by white points on costa, on median

nervure, and vein 1; a medial dark shade strongly angled

outwards below costa and merging into a straight dark band

from middle of inner margin to apex followed by narrow tawny

and broader whitish shade; a black spot followed by white on

costa near apex; a large ochreous semihyaline spot beyond the

band from below vein 2 to above 3 with some strigule and

the veins dark; beyond the spot is a dark diffusion, leaving a

light space below apex; a black discocellular spot with a short

black fascia above it: cilia dark brown. Hind wings ochreous

brown with brown and black strigule and suffused with tawny

brown on lower part of outer half of wing: an antemedial band

of brown, tawny and whitish in continuation of band on fore

Proc, Zoon, Soc.—1921, No, XXIII, 23

332 MR, E. DUKINFIELD JONES ON

wings; a dark medial shade; three black points at tornus on

veins 1 and 2 and submedian fold; cilia dark brown.

Expanse 42 mm.

Hab. Alto da Serra, Santos; Castro, Parana.

TETRAGONODES GEMINARIA, sp.n. (PI. II. fig. 9.)

Female.—Palpi ochreous, outwardly brown; pectus ochreous ;

legs ochreous speckled with brown ; frons, head, antenna, thorax,

and abdomen light yellowish brown, obscurely strigulated with a

darker shade and irrorated with black; a minute black disco-

cellular spot; a very obscure dark medial shade; a geminate

postmedial line from costa just before apex, incurved to vein 7,

where it is acutely angled outwards, slightly incurved to 6 and

thence straight to inner margin one-third from tornus, the inner

member rufous brown and the outer dark brown, with whitish

between; termen concolorous; cilia dark brown with whitish base.

Hind wings ochreous suffused with light brown and irrorated

with black, the terminal area with a darker suffusion; a medial

geminate line similar to postmedial on fore wings, slightly and

evenly excurved ; cilia as on fore wings.

Expanse 29 mm.

Hab. Castro, Parana, Brazil.

BAGODARES CASTRA, sp. n. (PI. I. fig. 22.)

Female.—Palpi, head, and antenne reddish brown, a white

bar between the antenne; legs ochreous suffused with reddish

brown; tegule reddish brown mixed with white; patagia, thorax,

and abdomen light brown. Fore wings white ; costa light brown

irrorated with darker shade; two narrow brown bands from inner

margin near the base converging and meeting in the cell, thence

a single line to below costa just before apex, where it bends

violently inwards to costa; a brown fascia from one-third on

costa to join the other band in the cell; two brown bands from

middle of inner margin converging and meeting at apex ; a sub-

terminal single band meeting the preceding below apex; termen

dark brown; cilia light brown. Hind wings: a straight ante-

medial brown band; two parallel postmedial bands; a subterminal

band meeting terminal line at apex; terminal line dark brown ;

cilia light brown.

Expanse 22 mm. e

Hab. Castro, Parana, Brazil.

Potts AcUTARIA, sp.n. (PI. II. fig. 10.)

Female.—Palpi brown, outwardly fuscous ; pectus white; legs

ochreous speckled with fuscous; frons rufous-brown ; head,

antennie, thorax, and patagia light brown mixed with white ;

abdomen darker brown. Fore wings light brown with fuscous

strigule ; a well-defined dark brown antemedial band from inner

margin to cell at origin of vein 2, followed by tawny suffusion ;

NEW MOTHS FROM SOUTH-EAST BRAZIL. 333

a diffused tawny postmedial line, fuscous on costa, bent outwards

below costa, followed by series of indistinct dark spots diminish-

ing in size from inner margin’ to vein 4; a pale submarginal

band, preceded by indistinct dark spots at tornus and on vein 2 ;

a minute dark discoeellular spet; cilia dark brown. Hind wings

light brown, heavily strigulated ; a subbasal dark brown band

forming continuation of antemedial of fore wings; an obscure

medial line of spots, indistinct except on inner margin, where

there is a large dark spot surrounded by white; a wavy dark

subterminal line on lower half of wing, expanding into large

patch at tornus; a very minute discocellular spot.

Expanse 39 mm.

Hab. Alto da Serra, Santos.

TROTOGONIA CASTRARIA, sp. n. (PI. IT. fig. 18.)

Male.—Palpi, pectus, and fore- and mid-femora dull orange ;

legs ochreous ; frons rufous; antenne ochreous orange speckled

with purple-brown ; tegule whitish; patagia bright ochreous,

the scales tipped with white; abdomen ochreous, dorsally rufous

with some white scales. Fore wings bright ochreous; costa

olive-brown; an olive-brown antemedial band between fine

whitish lines from subcostal to inner margin ; a narrower rufous-

brown band, evenly incurved from vein 6 through end of cell to

middle of inner margin; a dark brown subterminal band irrorated

with a few whitish scales ; termen dark brown shading to rufous

at apex; the medial area suffused with rufous; subapical area

rufous, containing two dark subcostal spots surrounded with

whitish ; terminal area from vein 5 to tornus heavily suffused

with Biive: brown, containing ochreous spot below vein 2; cilia

dark olive-brown shading to rufous at apex, a white spot above

submedian fold. Hind wings bright ochreous slightly irrorated

with rufous; a dark antemedial band with some whitish scales

slightly incurved from middle of inner margin, obscure on costa ;

a minute spot at apex below vein 7; cilia bright ochreous except

at apex, which is dark olive-brown with a few whitish scales.

Underside brighter than upper; fore wings: costal area rufous,

terminal area broadly rufous brown with yellow spot below

vein 2; hind wings bright ochreous yellow, a rufous spot at

apex below vein 7.

Expanse 29 mm.

Hab. Castro, Parana, Brazil.

NUMIA STRIGULARIA, sp.n. (PI. II. fig. 11.)

Male.—Palpi dull ochreous yellow, the third joint rufous-

brown; legs ochreous yellow speckled with rufous; frons and

antenne rufous brown; thorax and abdomen dull ochreous

yellow. Fore wings dull ochreous yellow striated with minute

rufous strigule which become black on costal edge; a rufous-

brown antemedial band, broad at costa and narrowing to- inner

Q3*

334 MR, E. DUKINFIELD JONES ON

margin, angled outwards below costa and preceded by a whitish

line; a narrow postmedial band, indistinct except from inner

margin to vein 2, followed by some whitish scales; a subapical

brown spot on costa containing a few white scales; a subterminal -

spot on vein 3 and below vein 2; termen andl cilia rufous brown.

Hind wings ochreous yellow striated with rufous strigule; a

postmedial brown band, broad at costa and narrowing to inner

margin, followed by whitish scales; apex rufous brown; cilia

ochreous yellow except at apex. Underside: the colours are

brighter and the markings more distinct.

Expanse 27 mm.

Hab. Castro, Parana, Brazil.

CAMPATONEMA, gen, nov.

Palpi upturned, roughly scaled, third joint minute ; frons with

conical prominence ; antenne of male fasciculate ; thorax clothed

with scales and hairs; patagia fringed with long hairs; abdomen

smooth. Fore wings: costa evenly arched; apex rectangular ;

termen evenly curved; cell more than half the length of the wing ;

vein 2 from middle of cell, 3 from well before angle, 4 from

angle, 5 from middle of discocellulars, 6 from upper angle,

7 from cell close to angle, 8 and 9 stalked from 7, 10 from 7,

11 from cell. Hind wings: cell more than half the length of the

wing; termen curved, slightly angled on vein 4; vein 2 from

middle of cell, 3 close to angle, 4 from angle, 5 absent, 6 from upper

angle, 7 from close to angle.

Type, C. marginata.

CAMPATONEMA MARGINATA, sp.n. (PI. II. fig. 21.)

Male.—Palpi brownish ochreous, 2nd joint with some fuscous

scales on outer side ; legs, head and antenne brownish ochreous ;

thorax and abdomen ochreous. Fore wings ochreous sparsely

irrorated with ferruginous brown; costa ferruginous brown ;

termen broadly ferruginous brown at tornus, diminishing to apex ;

traces of postmedial and subterminal lines, conspicuous on inner

margin ; inner margin ferruginous brown at tornus; a minute

dark discocellular point. Hind wings ochreous, the outer half

irvorated with ferruginous brown; a medial band wide at costa

and angled outwards above vein 7, narrower on inner margin ;

termen broadly ferruginous brown.

Expanse 24 mm.

Hab. Alto da Serra, Santos.

OXYDIA PALLIDARIA, sp.n. (PI. IT. fig. 12.)

Male.—Palpi dark brown, the tip of third joint ochreous white;

pectus ochreous white; legs light reddish brown spotted and

ringed with dark brown ; frons ochreous white with two brown

vertical streaks; vertex of head light brown with white border ;

NEW MOTHS FROM SOUTH-EAST BRAZIL. 335

antennx dark brown, the basal joint. white; tegule brown mixed

with white; patagia tawny in front, ochreous white behind ;

thorax ochreous white, two small dark subdorsal spots ; abdomen

ochreous white irrorated with black. Fore wings ochreous white

sparsely irrorated with black; costa tawny ; medial shade indi-

cated by tawny mark below costa; a minute black discocellular

spot; postmedial line black, evenly incurved from beyond middle

of inner margin to near termen above vein 6, where it is bent

violently inwards and incurved to costa, the portion above vein 4

is tawny; a large cluster of black irroration beyond the post-

medial on vein 3; terminal area darker than the rest of the wing

and somewhat suffused with fawn-colour. Hind wings ochreous

white sparsely irrorated with black; a minute black discocellular

spot; postmedial similar to that of fore wing and continuous with

it, vanishing at vein 7; outer half of wing more heavily suffused

with fawn-colour ; indications of a wavy tawny subterminal line.

Underside more heavily irrorated than upper; the postmedial

lines diffused black and tawny ; minute discocellular spots.

Expanse 58 mm.

Heb. Castro, Parana, Brazil.

In a second specimen the whole of the outer area of the fore

wings beyond a line drawn from tornus to costa a little beyond

the cell is heavily irrorated.

ISOCHROMODES ELEGANTARIA, sp.n (PI. IT. fig. 19.)

Female.—Frons and palpi rufous brown, 3rd joint darker ; legs

brown; vertex of head and tegule light rufous brown ; patagia,

thorax, and abdomen light ochreous brown. Fore wings lght

brown slightly irrorated with black ; antemedial line light rufous

brown, excurved from costa to just before origin of vein 2,

slightly incurved to vein 1, then bent inwards to inner margin ;

a minute black discocellular spot; postmedial line dark rufous

brown outwardly shaded by light brown, sinuous, excurved on

veins 7 and 4, incurved from 4 to inner margin, dark points

on the veins, preceded by rufous-brown shade broad at costa and

narrow on inner margin; a wavy ochreous-white subterminal

line only visible above vein 5, below 5 the whole area from post-

medial to termen is ochreous white on which are three small

rufous-brown spots on veins 1 and 2 and the submedian fold, a

larger and more distal spot above vein 2; termen dark brown ;

cilia ochreous white with dark points at the veins. Hind wings

light brown, irrorations more numerous than on the fore wings ;

antemedial line obscure; postmedial rufous brown followed by

whitish, slightly wavy, evenly excurved and preceded by a slight

rufous-brown suffusion reaching to cell; a subterminal series of

obscure diffused rufous-brown spots; termen and cilia as in

fore wings.

Expanse 25 mm.

Hab. Alto da Serra, Santos.

336 MR. E. DUKINFIELD JONES ON

AZBLINA HANEBARIA, sp. n. (PI. II. fig. 13.)

Male.—Palpi dark rufous brown, the third joint tipped with

ochreous ; legs bright creamy ochreous suffused and spotted with

purple-brown; head and antenne rufous brown; patagia and

thorax ‘purplish brown; abdomen rufous brown. Fore wings:

base lilacine with brown striations; a fine black antemedial line

outwardly shaded with dark brown, obliquely excurved from

costa to vein 1, where it bends outwards and then makes a

minute and violent curve to inner margin ; the black line follows

the inner margin and connects with the postmedial line, which is

slightly excurved from inner margin to submedian fold, then

incurved obliquely outwards to vein 4 well beyond the cell,

thence almost straight to well before apex; the medial area

above the median vein creamy buff slightly irrorated with brown,

some reddish-brown suffusion bevond the cell; below the median

vein dark purplish brown striated with lilacine ; a diffused lilacine

subterminal line joining postmedial on costa, the space between

postmedial and subterminal creamy buff striated with brown ;

terminal area suffused with red above vein 2 to apex; termen

lilacine; a series of dark subterminal spots between the veins,

those above veins 3 and 6 containing ochreous points; cilia dark

brown. Hund wings lilacine grey; an obscure pale wavy post-

medial line; inner margin golden buff to postmedial, beyond this

bright creamy ochreous to tornus and extending to vein 3;

black subterminal spots between the veins, the one above vein 6

with ochreous point; cilia brown tipped with white.

Expanse 33 mm.

Hab. Alto da Serra, Santos.

Nearly allied to A. habenaria Gueneée.

AZELINA CETANA, sp. n. (PI. II. fig. 14.)

Male.—Palpi light brown, outwardly fuscous, third joint

fuscous tipped with light brown; legs light brown heavily irro-

rated and suffused with fuscous; frons and antenne reddish

brown; vertex of head and tegule light brown mixed with

ochreous white; patagia reddish brown suffused with fuscous, a

few ochreous-white scales; thorax and abdomen red-brown.

Fore wings light brown ; an indistinct double inner line; a black

sinuous antemedial line outwardly shading to red-brown, strongly

angled inwards and preceded by ochreous-white spot on vein 1

and median vein; a wavy black postmedial line angled inwards

on the veins and strongly excurved on the interspaces, shaded

inwardly with brown and followed by ochreous white at costa,

preceded by a nearly straight dark shade confluent at costa and

at veins 3 and 4; a wavy subterminal line from before apex, to

vein 4, where it joims the terminal line, strongly angled inwards

on vein 6; a terminal line of spots between the veins; the inner

area suffused with violaceous and irrorated and striated with

brown; the medial area above the median vein yellowish brown,

below the median violaceous ; outer area pale olive-green suffused

NEW MOTHS FROM SOUTH-EAST BRAZIL. 337

with brown except the part beyond the subterminal line at apex,

where it is violaceous brown; cilia reddish brown. Hind wings

suffused with golden brown, lighter on inner margin; a pale

postmedial line, obscure on costa and clearly defined on inner

margin; a series of subterminal spots between the veins, the

three near tornus much larger than the rest. Underside reddish

brown slightly irrorated with darker shade and suffused with

fuscous on central area; costa crossed with brown strigule ;

ochreous white at apex, above tornus, and on discocellulars; a

slightly sinuous postmedial line; on the hind wings the post-

medial line is wavy, dark, followed by ochreous white.

Expanse 34mm.

Hab. Alto da Serra, Santos.

NepHopIA BoNTTARIA, sp.n. (PI. LI. fig. 22.)

Female—Palpi light brown, third joint dark ; legs, head,

antenne, thorax, and abdomen brownish grey; patagia dark

brown on shoulders. Wings brownish grey. Fore wings: costa

dark brown at base, becoming paler towards apex ; a dark diffused

antemedial shade, clearly defined on costa ; a dark diffused disco-

cellular bar ; a narrow postmedial line parallel with termen ; cilia

dark with darker spots at ends of veins, Hind wings: an indis-

tinct discocellular spot ; a diffused postmedial line excurved from

costa to vein 3, incurved from 3 to inner margin ; cilia with dark

spots at ends of veins.

Underside :—Fore wings suffused*on costal and terminal areas

with broad brown strigule; postmedial line and discocellular bar

as on upper side. Hind wings more heavily suffused with brown

strigule ; postmedial as on upper side.

Expanse 34 mm.

Hab. Castro, Parana, Brazil. é

NEPHODIA PAULARIA, sp.n. (PI. II. fig. 23.)

Male.—Palpi light brown; pectus grey; legs grey suffused

with brown ; frons, head, and antenne grey ; tegule and patagia

suffused with brown; thorax and abdomen grey. Wings grey.

Fore wings: costa and outer half of wing suffused with brown ;

indistinct diffused dark medial shade beyond cell, confluent with

postmedial on vein 2; an indistinct diffused dark postmedial

shade; a well-defined dark discocellular bar; termen and cilia

concolorous. Hind wings: a minute dark discocellular spot; a

very faint medial dark shade; terminal area darker than the

rest of the wing.

Underside :—Fore wings: costa strigulated with darker brown ;

a well-defined dark discocellular bar ; a well-defined double fuscous

postmedial shade. Hind wings evenly strigulated with broad

diffused strigule; a well-defined discocellular spot ; dark diffused

medial and subterminal shades.

Expanse: male 47 mm., female 48 mm

Hab. Siio Paulo, 8.E. Brazil.

338 MR. E. DUKINFIELD JONES ON

BoaRMIA NIGRARIA, sp. n. (PI. II. fig. 1.)

Female.—Palpi white, some fuscous scales at end of 2nd and

3rd joints; legs ochreous speckled with fuscous, fore tibize with

large fuscous patches on upper side; fore tarsi ringed with fuscous,

mid- and hind-tarsi fuscous; pectus ochreous white; frons

fuscous in front and ochreous above; back of head ochreous ;

antenne ringed with fuscous and white; tegule and patagia

fuscous, light brown, and ochreous; thorax fuscous brown and

ochreous, a large ochreous-white spot on metathorax with

fuscous band across it; abdomen ochreous white irrorated with

fuscous, dark subdorsal spots, a wide black dorsal band at base.

Fore wings ochreous white heavily suffused with fuscous and

black; an obscure dark subbasal line; antemedial line narrow,

black, angled outwards on vein 1; an obscure double dark

medial shade, distinct on inner margin, where there are a few

reddish-brown scales; a discocellular bar of raised bluish-grey

scales; postmedial only visible from costa to below vein 5, black,

ineurved, followed by whitish and then reddish brown; a very

wavy whitish subterminal line; terminal area ochreous white

with a dark fuscous space irrorated with bluish grey from 4

to 7; veins 2, 3, and 4 black from medial shade to subterminal

line; a white space on inner margin from antemedial line to

medial shade; termen black; cilia ochreous white with dark

central band. Hind wings ochreous white irrorated with

fuscous ; an antemedial dark shade, double from inner margin

to cell, single from cell to costa; a black postmedial line,

punctuated on the veins, slightly incurved from costa to

discal fold, then more strongly incurved to inner margin; an

elliptical bluish-grey discocellular bar surrounded with black ;

a very wavy white subterminal line; the space between post-

medial and subterminal from inner margin to discal fold heavily

suffused with fuscous. Underside ochreous white; fore wings

slightly strigulated with fuscous brown; a dark diffused disco-

cellular bar; a large fuscous subapical spot; apex ochreous

white with two minute terminal spots above 7 and 8; hind

wings: diffused fuscous discocellular spot, some dark suffusion on

terminal area.

Expanse 44 mm.

Hab. Castro, Parana, Brazil.

H{YMENOMIMA SINUOSARIA, Sp.n. (PI. LI. figs. 2, 3.)

Male.—Palpi brown; legs ochreous speckled with brown ;

frons pale reddish brown, a dark bar in front of antenne ; vertex

of head brownish ochreous; antenne brown; tegule ochreous

mixed with reddish brown ; thorax and abdomen ochreous white

sparsely irrorated with black; a pair of subdorsal tawny spots

irrorated with black on second segment of abdomen. Fore wings

ochreous white irrorated with black; costa narrowly tawny; a

narrow, tawny, sinuous antemedial line, excurved from costa to

NEW MOTHS FROM SOUTH-EAST BRAZIL. 339

median vein, slightly incurved thence to inner margin ; a tawny

wavy medial shade; postmedial line narrow, sinuous, excurved

from costa to vein 3, incurved thence to inner margin, black on

the veins and tawny on the interspaces, followed by a narrow

pale line, a broad tawny band and a rather obscure zigzag pale

line angled inwards on the veins and outwards on the inter-

spaces; terminal area suffused with pale ferruginous brown; a

terminal row of minute black points on the interspaces; cilia

brown. Hind wings similar to the fore wings, with the exception

that there is no antemedial line. Underside ochreous white

more heavily irrorated than upper, outwardly suffused with pale

ferruginous brown; a black discocellular spot ; a postmedial line

of black spots on the veins.

_Female-—The wings purplish grey irrorated with black; the

lines and suffusions as in male, but darker and duller.

Expanse: male 29 mm., female 31 mm.

Hab. Castro, Parana, Brazil.

HYMENOMIMA NIVACARIA, sp.n. (Pl. II. fig. 4.)

Male.—Palpi rufous brown; pectus white; legs brown; frons

rufous brown, white in front; vertex of head rufous brown

mixed with white; antenne light brown, white at base; tegule

rufous brown: thorax, patagia, and abdomen ochreous with

scattered rufous-brown scales. Fore wings ochreous irrorated

with rufous brown; edge of costa dark brown: antemedial line

light rufous brown, strongly bent inwards in cell; medial line

slightly excurved; postmedial line dark rufous brown, excurved,

strongly dentate, the points outwards on the veins; an obscure

dentate subterminal line; terminal line dark rufous brown ;

cilia ochreous. Hind wings similar to fore wings, but the lines

not so well defined ; postmedial angled outwards on vein 4.

Expanse 26 mm.

Hab. Nivae, Matto Grosso, Brazil.

LARENTIOPSIS, gen. nov.

Palpi porrect, short, clothed with scales, third joint minute ;

frons with a sharp-pointed prominence ; antenne of male bipec-

tinate, the pectinations short and dilated at the ends; tibial

spurs short ; thorax and patagia clothed with hairs and hair-like

scales; a metathoracic dorsal tuft; abdomen smooth, a small

dorsal tuft on second segment. Fore wings: costa straight ; apex

rounded; termen evenly curved, slightly crenulate; vein 2 at

two-thirds from base, 3 and 4 from angle, 5 from middle of

discocellulars, 6 from upper angle, 7, 8, and 9 stalked from close

to angle, 10 and 11 from cell. Hind wings: vein 2 at three-

quarters from base, 3 from just before angle, 4 from angle,

5 absent, 6 from upper angle, 7 from close to angle, 8 approxi-

mated to cell near base only.

Type, LZ. costiplaga.

340 MR. E. DUKINFIELD JONES ON

LARENTIOPSIS COSTIPLAGA, sp. n. (PI. II. fig. 5.)

Maie.—Palpi and head fuscous grey ; pectus white; legs brown

suffused with fuscous grey; tegule grey; patagia white mixed

with olivaceous; thorax dark fuscous grey, the metathoracic tuft

olivaceous ; abdomen dark fuscous grey. Fore wings fuscous grey

heavily suffased and irrorated with black and olivaceous-green

scales with iridescent reflection ; basal area fuscous grey; ante-

medial line wavy, black, nearly straight from costa to subcostal

nervure, where it is angled inwards, excurved below cell, angled

inwards on vein 1, exeurved to inner margin, followed by

some olivaceous scales above and below vein 1; postmedial wavy,

black, angled outwards on the veins; a dark, obscure subterminal

line; terminal line black; medial area darker than the rest of

the wing; a large white spot on costa beyond cell reaching to

vein 4, outwardly striated with black on the costa; an obscure

white apical spot heavily suffused with black ; a similar spot at

termen between veins 1 and 2. Hind wings uniform dull smoky

black.

Expanse 39 mm.

Hab. Castro, Parana, Brazil.

Subfam. LARENTIINA.

RHODOMENA SANTARIA, sp. n. (PI. IT. fig. 6.)

Female—Palpi fuscous brown; pectus luteous; legs brown ;

head and tegule luteous brown; antenne dark brown ; patagia

and thorax olive-green and fuscous brown, a metathoracic tuft of

black curved scales with metallic lustre; abdomen light brown

suffused with greenish. Fore wings luteous olive-green; base

fuscous; antemedial line geminate, wavy, slightly excurved

through base of cell, very dark on costa, followed by olive-green

on which there is a ferruginous-brown band from inner margin

to subcostal vein; a geminate dark wavy medial line enclosing

darker shade of green, very dark on costa; medial area pale green

suffused with luteous except on costa; a black discocellular spot ;

postmedial wavy, geminate, the outer member denticulate, the

points inwards on the veins, enclosing darker shade of green,

very dark on costa, followed by a narrow pale line, a diftused

dark line and a broad ferruginous-brown band which is confluent

with a large black subapical spot and has distal projections above

tornus and veins 2 and 3; terminal area suffused with dark

olive-green at apex and tornus, leaving a luteous spot below

vein 2 and a luteous space from vein 3 to the subapical spot; a

terminal row of black lunular spots between the veins; cilia

luteous olive-green with dark centre line. Hind wings pale

luteous slightly suffused with fuscous on inner margin; an

obscure geminate postmedial line, excurved beyond the cell ; ter-

men crenulate, black, interrupted at the veins; cilia luteous

suffused with fuscous. Underside luteous: fore wings suffused

NEW MOTHS FROM SOUTH-EAST BRAZIL. 341

with pale olive-green at base; a dark olive-green discocellular

spot; apex dark olive-green, cut by a fine luteous line from

above vein 5 to before apex; a crenulate dark green postmedial

line, broad at costa; hind wings: a dark gveen discocellular

spot; a crenulate diffused dark green postmedial line and similar

subterminal ; a brown terminal suffusion.

Eixpanse 32 mm.

Hab. Alto da Serra, Santos.

RHODOMENA PAULARIA, sp. n. (PI. II. fig. 7.)

Female.—Palpi ochreous white, the first joint outwardly black ;

legs buff heavily suffused with black, the tarsi black ringed with

buff; frons lustrous whitish fawn, a large central purple-brown

spot; vertex of head pale buff-white with brown central spot ;

antenne slightly metallic purple-brown, the basal joint ochreous ;

tegule pale buff-white with a dark brown line; patagia ochreous

white, fuscous brown on the shoulder; a large black and brown

metathoracic tuft with metallic reflection; abdomen dark ochreous,

ringed with tawny and dorsally suffused with black. Fore wings

ochreous white heavily suffused with tawny and fuscous brown ;

a black basal line from costa to vein 1; a dark diffused ante-

medial line, narrow on inner margin and broad at costa; a dark

diffused medial shade, narrow below cell, broad in and above cell,

enclosing black discocellular spot and preceded by a narrow

whitish shade; a second dark shade before postmedial ; post-

medial line narrow, denticulate, excurved from vein 2 to 6,

thence straight to costa, slightly bent outwards to vein 2, followed

by rather broad ochreous white ; terminal area heavily suffused

with fuscous and brown, the apex very dark; an obscure wavy

subterminal line indicated by lilacine scales. Hind wings

ochreous suffused with luteous brown; obscure diffused post-

medial and subterminal lines; termen brown.

Expanse 27 mm.

Hab. Alto da Serra, Santos.

HYPoOLEPIS BELLA, sp.n. (PI. II. fig. 25.)

Male.—-Palpi, pectus, and legs tawny, the latter heavily streaked

and suffused with dark reddish brown; frons brown tawny ;

vertex of head golden tawny ; antenne tawny, the shaft streaked

and suffused with brown; tegule tawny mixed with golden

yellow ; patagia outwardly golden brown, inwardly golden yellow;

abdomen golden yellow heavily suffused with purplish brown

except at base, a dorsal and two sublateral dark spots on first

segment, dorsal only on second and third. Fore wings rich

golden brown; costa and subcostal vein barred with golden-

yellow strigule, costa almost black on basal half; a silvery-white

wavy antemedial line angled outwards on subcostal vein and

inwards on vein 1; a silvery-white medial band passing through

cell, angled outwards on vein 2, the middle of the band occupied

by a narrow band of yellow irorated with brown, the white

342 MR. E. DUKINFIELD JONES ON

band preceded by an oblique golden-yellow spot in the cell and a

dark suffusion in and below cell ; a wavy postmedial line straight

from costa to vein 7, exeurved to 5, excurved to 3, diagonally

inwards to 2, below which it makes an § and is angled inwardly

on vein 1; a broad silvery-white band follows, much dilated at

costa, and includes denticulate subterminal line, the points

outwards on the veins, the middle of the band is suffused with

yellow irrorated with brown; the space between the white bands

heavily with dark brown, the suffusion extending to termen

between veins 2 and 4 and nearly obliterating the outer white

band ; terminal line fuscous brown, crenulate ; cilia golden yellow

interrupted at the veins with fuscous brown. Hind wings

golden brown; an obscure diffused dark sinuous postmedial line

and discocellular spot ; terminal line and cilia as on fore wing.

Expanse 25 mm.

Hab. Alto da Serra, Santos.

PSALIODES AURANTARIA, sp.n. (PI. III. fig. 18.)

Female.—Palpi white beneath, ochreous above; legs ochreous ;

antenne brown; frons dark brown; vertex of head and tegule

ochreous; patagia and abdomen ochreous irrorated with brown.

Fore wings ochreous white; basal area suffused with fuscous; a

wavy dark inner line followed by white; central area orange;

a broad suffused fuscous medial band preceded and followed by

white; a black discocellular spot; an obscure postmedial line

followed by broad dark band, fuscous at costa and brown above

tornus ; terminal line fuscous ; terminal area near apex suffused

with orange. Hind wings ochreous slightly suffused with orange ;

a dark discocellular spot.

Expanse 21 mm.

Hab. Alto da Serra, Santos.

KUPITHECIA CUPREARIA, sp. n. (PI. III. fig. 9.)

Male.—Palpi pale reddish brown ; pectus white ; legs ochreous

brown; head reddish brown ; antenne brown, the shaft ringed

with light and dark; tegule light brown; thorax and abdomen

ochreous white. Fore wings ochreous white, suffused with

fuscous on costa and medial area and with coppery brown on

outer area; base ochreous white; a dark diffused wavy basal line

angled inwards on median vein and outwards on submedian fold ;

a dark diffused wavy antemedial line angled inwards below costa,

outwards on subcostal vein, inwards in cell and outwards on

median vein and vein 1; indications of a dark medial shade; a

diffused black lunular discocellular spot ; postmedial very obscure,

preceded by light space on costa; a subterminal line of obscure

spots, followed by white spot above vein 3 and a few scattered

white scales on the other interspaces; broad dark bars on costa 5

a fuscous space at tornus. Hind wings ochreous white, the

medial area below cell suffused with fuscous, other areas suffused

with coppery brown and fuscous; base ochreous white; a round

NEW MOTHS FROM SOUTH-EAST BRAZIL. 343

dark discocellular spot ; postmedial line broad, diffused, excurved

beyond cell, straight to inner margin, followed by a light band

containing a line of diffused coppery spots; a subterminal row of

obscure dark spots followed by white spot above vein 3; termen

dark ; cilia ight brown.

Expanse 23 mm.

Hab. Alto da Serra, Santos.

KUPITHECIA MAUVABIA, sp. n. (PI. IIT. fig. 1.)

Female.—Palpi, legs, and head grey tinged with light brown ;

antenne brown; tegule, patagia, and thorax lilacine grey;

abdomen greenish grey. Wings lilacine grey. Fore wings:

costa dark fuscous at base; antemedial line straight from costa

to cell, where it forms a right angle, thence straight to inner

margin; medial area, a large dark fuscous patch on costa, out-

lined with whitish and including a discocellular bar of raised,

iridescent scales, a whitish patch on inner margin suffused with

fuscous; an obscure pale subterminal line preceded by some

greenish-grey suffusions; terminal area suffused with greenish

grey. Hind wings: medial area whitish with irregular fuscous

diffused lines, including discocellular spot; some greenish-grey

suffusions on outer half of wing.

Expanse 20 mm.

Hab. Castro, Parana, Brazil.

HUPITHECIA AQUANIVARIA, Sp.n. (PI. IIT. fig. 2.)

Male.—Palpi, frons, head, tegule, and patagia white with

sight rufous tinge, the latter with anterior fuscous spot ; fore

legs brown, tarsi ringed with brown; abdomen ochreous white.

Wings lustrous greenish grey. Fore wings: base of costa fuscous

black ; a large niedial patch of fuscous black on costa, surrounded

by whitish and containing discocellular bar of marae iridescent

seales ; a whitish dentate "subterminal line preceded by minute

fuscous spots on the inters paces, those at tornus and costa larger ;

cilia very long, lustrous grey with diffused greenish fuscous spots

on base at the ends of the veins. Hind wings: postmedial and

subterminal dentate whitish lines preceded by dark shade, angled

outwards on vein 4, thence incurved to inner margin ; oe) minute

fuscous discocellular spot.

Female similar.

Expanse: male 16 mm., female 2] mm.

Hab. Castro, Parana, and Alto da Serra, Santos. -

Subfam. STERRHINA.

ANISODES BIZARTA, sp.n. (PI. ITI. fig. 3.)

Male.—Palpi light rufous brown, whitish beneath ; legs, head,

antenne, thorax, and abdomen pale rufous brown. aire wings

pale rufous boa finely striated with darker purplish brown ;

344 MR. E. DUKINFIELD JONES ON

a narrow purplish-brown antemedial line angled outwards on

subcostal nervure, thence straight to inner margin; a similar

postmedial line, slightly sinuous, incurved below costa to vein 6,

excurved to below discal fold and straight to inner margin ;

costa, heavily striated with purplish brown on medial and terminal

portions; a terminal series of minute lunules between the veins ;

cilia light reddish brown, some purple at the tornus. Hind

wings similar to the fore wings, the lines straight in continuance

of those of the fore wings; cilia of inner margin purple.

Hxpanse 19 mm.

Hab. Alto da Serra, Santos,

ANISODES PARANARIA, sp. n. (PI. III. fig. 4.)

Female.—Palpi reddish brown above, whitish beneath; legs

and head pale ochreous brown; vertex white; antenne ochreous ;

thorax, abdomen, and wings ochreous brown. Fore wings

irrorated with rufous brown ; well-defined ante- and postmredial

narrow dark rufous-brown bands; termen and discocellular bar

dark rufous brown ; cilia ochreous brown. Hind wings similar;

discocellular spot elongated, whitish, surrounded by rufous

brown.

Expanse 23 mm.

Hab. Castro, Parana, Brazil.

ANISODES CAROLINA, sp. n. (PI. III. fig. 5.)

Female.—Palpi pale ochreous brown; legs ochreous brown,

the trochanters and femora glistening whitish; head roseate ;

vertex ochreous white with roseate bar; antenne ochreous

white; tegule light rufous brown ; patagia with roseate anterior

spot; metathorax with some roseate scales ; abdomen ochreous

brown. Wings ochreous brown, striated with roseate brown,

both wings similar ; antemedial line represented by fuscous spots

on vein 1, median neryure, and cellular fold; postmedial line of

diffused fuscous spots on the veins from inner margin to vein 7,

preceded by broad fuscous suffusion from inner margin to vein 4 ;

discocellular spot whitish surrounded by fuscous; termen con-

colorous, with fuscous spots at ends of veins.

Expanse 22 mm.

Hab. Castro, Parana, Brazil.

ANISODES JAPARIA, sp. n. (PI. ITI. fig. 6.)

Female.—Palpi purple-brown above, white beneath, pectus

white; legs light ochreous brown; frons white; vertex light

brown with purple-brown bar; antenne light brown; tegule

purple-brown; patagia, thorax, and abdomen light ochreous

brown. Fore wings light ochreous brown irrorated with dark

purple-brown ; antemedial line represented by fuscous spot on

yein 1 and on median and subcostal nervures; a wavy dark

NEW MOTHS FROM SOUTH-EAST BRAZIL. 345

medial shade; a postmedial line of fuscous spots on the veins; a

subterminal line of diffused brown spots on the interspaces; a

terminal row of fuscous spots on the interspaces; cilia ochreous.

Hind wings light ochreous brown ; antemedial line represented

by spots on vein 1, median nervure, and in cell; a wavy medial

dark shade; postmedial, subterminal, and terminal lines as on

fore wings; discocellular spot silvery white surrounded by brown.

Expanse 28 mm.

Hab. Castro, Parana, Brazil.

ANISODES ANTENNARIA, sp.n. (PI. III. fig. 7.)

Male—Palpi and legs light ochreous brown, fore femora and

tibize heavily suffused with purple-brown ; frons purple-brown ;

head, antenne, tegule, and patagia light ochreous brown ; thorax

spotted with roseate brown ; abdomen light ochreous brown with

row of roseate dorsal spots. Wings light ochreous brown

irrorated with roseate brown. Fore wings: antemedial line

wavy, strongly angled outwards in cell, defined by dark spots on

median nervure, in cell, on subcostal nervure and above it; a

diffused narrow medial shade ; a postmedial line of spots on the

veins, followed by narrow ochreous brown shade; terminal area

suffused with ochreous brown; a terminal row of roseate fuscous

spots on the interspaces ; discoceliular spot surrounded by roseate

spots at ends of veins. Hind wings similar; discocellular spots

larger.

Expanse 35 mm.

Hab. Castro, Parana, Brazil.

ANISODES VIGORARIA, sp. n. (Pl. III. fig. 8.)

Female.—Palpi outwardly dark brown, inwardly ochreous

white ; legs ochreous brown ; frons dark brown; head, antenne,

and body ochreous. Fore wings ochreous, irrorated with fuscous ;

antemedial line wavy, angled outwards in cell, inwards on median

nervure, outwards on submedian fold, thence oblique to inner

margin; a medial line angled inwards on subcostal, outwards on

vein 6, thence straight to lower angle of cell and slightly in-

curved to before middle of inner margin, followed by narrow

dark shade; a diffused postmedial line with dark spots above

veins 4-7; an obscure subterminal shade; a terminal line of

lunular spots; discocellular spot ochreous, surrounded by orange

and fuscous; cilia ochreous. Hind wings similar, the medial

shade well defined on inner margin, not reaching costa.

Expanse 36 mm.

Hab. Castro, Parana, Brazil.

HL@MALIA MANTARIA, sp. n. (PI. IIT. fig. 10.)

Female.—Palpi brown; pectus white; legs ochreous brown ;

frons, vertex, and antenne dark brown; tegulae, patagia, and

thorax light ochreous brown; abdomen dark brown with large

346 MR. E. DUKINFIELD JONES ON

metallic dorsal spots on first segment. Fore wings ochreous,

heavily suffused with ferruginous and irrorated with dark purple

metallic scales; costa purple-brown; antemedial, medial, and

postmedial lines narrow, wavy, purple-brown ; a wavy subterminal

line broadly diffused at tornus, on and above vein 4 and at costa ;

termen dark purple-brown; cilia brown. Hind wings heavily

suffused with ferruginous and fuscous and irrorated with dark

metallic scales, the lines very obscure.

Expanse 26 mm.

Hab. Castro, Parana, Brazil.

Near H. micaceata W1k.

H@MALIA MAGITARIA, sp.n. (PI. III. fig. 11.)

Female.—Palpi and frons rufous brown ; pectus silvery white,

iridescent ; legs ochreous, iridescent ; head and antenne rufous

brown; vertex whitish; tegule brown with some iridescent

scales; thorax, abdomen, and wings ochreous brown. Fore

wings irrorated with darker brown, costa brown, dark at base

and lighter at apex; antemedial line very obscure; postmedial

slightly excurved beyond cell; subterminal wavy, followed by

light shade : terminal area slightly suffused with brown; a

fuscous discocellular spot ; termen brown ; cilia ochreous. Hind

wings more heavily irrorated; a medial shade in continuation of

postmedial of fore wings, passing on proximal side of disco-

cellular ; subterminal line excurved on veins 6-7.

Expanse 20 mm,

Hab. Castro, Parana, Brazil.

Eors LUTEARIA, sp. n. (PI. IIT. fig. 12.)

Male—Bright buff with rufous irroration; frons rufous;

antenne brown. Fore wings: antemedial line rufous, excurved

across middle of cell; medial line evenly excurved ; postmedial

excurved, approximating to medial on inner margin, all the lines

somewhat diffused, narrow on inner margin and broader at

costa ; subterminal line much diffused ; termen and cilia suffused

rufous. Hind wings similar, more heavily suffused; only two

conspicuous lines, antemedial and medial.

Expanse 17 mm.

Hab. Siio Paulo, 8.E, Brazil.

PryCHOPODA TERMINARIA, sp. n. (PI. III. fig. 13.)

Female.—Palpi and legs ochreous; frons reddish brown;

vertex of head white; antenne ochreous; tegule, patagia, and

thorax pale reddish brown; abdomen ochreous brown. Fore

wings pale ochreous, almost semihyaline; traces of brown ante-

medial and postmedial lines; a dark discocellular spot ; subterminal

line wavy diffused dark brown, incurved on submedian fold,

nearly straight from vein 2 to costa; terminal area a lighter

NEW MOTHS FROM SOUTH-EAST BRAZIL. 347

shade of brown except at apex which is ochreous; termen and

cilia brown. Hind wings pale ochreous; traces of postmedial

line ; subterminal line fer brown and terminal area a rather

lighter shade; cilia brown.

Expanse 15 mm.

Hab. Sao Paulo, S.E. Brazil.

PrYCHOPODA DELICATARIA sp.n. (PI. III. fig. 14.)

Male.-—Palpi, legs, head, and body light rufous brown ; vertex

of head and antenne ochreous white. Wings ochreous slightly

irrorated with light brown; the lines light brown, diffused ;

antemedial excurved in cell; an almost straight medial shade ;

wavy, slightly dentate postmedial and subterminal lines, the

former marked on costa by large fuscous spot; a fuscous disco-

cellular spot; terminal line fuscous, broken at the veins; a

lightly defined suffusion just before termen. Hind wings similar,

but the inner lines very indistinct.

Expanse 15 mm.

Hab. Castro, Parana, Brazil.

PrYCHOPODA CAMPINARIA, sp.n. (PI. III. fig. 15.)

Female.— Palpi, legs, head, andantenne ochreous brown; vertex

of head whitish ; tegule and patagia ochreous brown with lilacine

iridescence ; abdomen ochreous brown. Wings pale ochreous

brown irrorated with darker brown and irridescent scales; the

lines dark brown. Fore wings: antemedial line wavy, strongly

excurved in cell; a medial line passing well within the cell;

postmedial line well defined, straight from costa to vein 2,

incurved on submedian fold; costal and terminal areas lightly

suffused with brown; terminal line brown, broken at the veins;

a fuscous discocellular spot; cilia long, qlee Hind wings:

the postmedial line well defined, continuous with that of fore

wings. Underside : the postmedial lines well defined.

Expanse 18 mm.

Hab. Sao Paulo, 8.E. Brazil.

PryCHOPODA LILACARIA, sp. n. (PI. III. fig. 16.)

Male.—Palpi, legs, and head rufous brown; vertex and

antenn ochreous brown ; tegule and patagia rufous brown with

lilacine iridescence ; abdomen dorsally purple-brown, ventrally

rufous. Wings ochreous brown with lilacine iridescence. Fore

wings: an obscure purple-brown antemedial line, excurved from

costa to submedian fold; a well-defined diffused dark medial

band; postmedial line wavy, narrow and obscure at inner

margin, broader at costa, followed by light shade; terminal

area heavily suffused with purple-brown, with light space divided

by a fine brown line from vein 3 to apex; a minute fuscous

discocellular spot; termen purple-brown with conspicuous spots

Proc, Zoou, Soc.—1921, No, XXIV. 24

348 MR. E. DUKINFIELD JONES ON

below apex; cilia- purple-brown. Hind wings: three diffused

bands; terminal line of elongated spots; cilia ochreous.

Expanse 20 mm.

Hab. Alto da Serra, Santos.

HiYRIA GEMMARIA, sp.n. (PI. III, fig. 17.)

Male.—Palpi ochreous and purple-brown; legs ochreous, the

fore pair suffused with purple-brown on upper side; frons and

vertex of head dark purple-brown ; antenne light. brown suffused

with purple near base; tegule white; patagia, thorax, and

abdomen light brown with lilacine reflection ; a large black spot

on first segment of abdomen. Fore wings lilacine grey sparsely

irrorated with purplish brown, a strong lilacine reflection ; costa

pure white from base to postmedial line; indications on costa of

a dark medial shade and postmedial line; a luteous white disco-

cellular spot shaded inwardly and outwardly with a few dark

seales; a dark purple-brown subterminal line from tornus to

two-thirds on costa, slightly and evenly excurved beyond the cell

and dilating on costa, followed by lilacine.and reddish purple to

near termen; termen and cilia bright luteous. Hind wings

lilacine grey irrorated with purple-brown; medial dark shade

well defined, angled inwards on submedian fold, outwards on

median vein and incurved in the cell; postmedial line less dis-

tinct; discocellular spot luteous white surrounded with purple ;

subterminal line parallel with termen and followed by reddish

purple; termen and cilia luteous.

Female.—Similar to male, but all the colours more vivid; the

inedial shade and postmedial line clearly defined on the fore- as

well as on the hind wings.

Expanse: male 13 mm., female 15 mm.

Hab, Castro, Parana, Brazil.

HiyrioGONA SANTARIA, sp.n. (PI. IIT. fig. 19.)

Male.—Palpi purple-brown; legs light brown, the fore pair

suffused with purple-brown ; head purple-brown; antenne light

brown ringed with darker shade; tegule and thorax lght

brown; abdomen light brown heavily suffused with purple.

Fore wings ochreous suffused with light brown with lilacine

reflection and very sparsely irrorated with purple-brown; ante-

and postmedial lines indicated by spots on costa; a large purple-

brown space with lilacine reflection from just before middle of

inner margin to tornus, the inner edge incurved to origin of

vein 2, an abrupt outward projection between 3 and 4; this

space is bordered by darker diffused lines and crossed vertically

by a medial line; a triangular ochreous space with base on vein 3

and apex at tornus; traces of diffused subterminal bands near

apex; a black discocellular spot; cilia pale ochreous, a rather

large black point opposite end of vein 3; outer half of inner

margin strongly lilacine. Hind wings ochreous heavily suffused

with purple-brown ; a large ochreous space on inner margin just

before tornus with tawny shade on inner- and upper sides

NEW MOTHS FROM SOUTH-EAST BRAZIL. 349

extending to discocellular spot; a dark purple-brown subterminal

spot above vein 3 preceded by ochreous; terminal area from

tornus to vein 6 dark lilacine grey ; termen from vein 6 to apex

ochreous; an ochreous line divides the grey from the purple

from apex to subterminal spot above vein 3; cilia ochreous,

black points opposite ends of veins.

Expanse 17 mm.

Hab. Alto da Serra, Santos.

HYRI0GoNA MONTARIA, sp.n. (PI. IIT. fig. 20.)

Male.—Palpi and head purple-brown; legs light brown, the

fore pair suffused with purple; antenne, tegule, and patagia

light reddish brown; thorax posteriorly purple; abdomen light

reddish brown, a few purple scales. Fore wings light reddish

brown with lilacine reflection irrorated and suffused with purple-

red; costa slightly suffused with dark brown; narrow diffused

dark medial and postmedial lines; a dark discocellular spot; a

diffused dark purple line from tornus to two-thirds on costa

shaded outwardly with purple-red, which expands violently

above vein 5 to close to apex; terminal area luteous, angled

inwards on vein 5; a terminal row of minute purple points on

the ends of the veins; cilia luteous. Hind wings similar to the

fore wings, but the subterminal area is much more broadly

suffused with purple and the termen narrowly luteous.

Expanse 17 mm,

Hab. Alto da Serra, Santos.

METPASIOPSIS PROUTARIA, sp.n. (PI. II. fig. 8.)

Male.—Palpi brownish ochreous ; legs ochreous irrorated with

light brown; frons and head rufous brown; vertex ochreous

white ; antenne white; tegule pale rufous brown; thorax and

abdomen ochreous irrorated with brown. Wings ochreous white.

Fore-wings: indistinct subbasal, antemedial, and medial rufous-

brown bands, the latter enclosing fuscous discocellular spot;

postmedial line pale rufous brown, wavy, marked by fuscous spot

on costa; terminal area fuscous from tornus to vein 7, the inner

edge excavated from 2 to 4, the dark space traversed by wavy

pale subterminal line: a terminal line of fuscous lunules on the

interspaces, a fine wavy whitish line dividing them from the dark

area; cilia ochreous white. Hind wings: medial, postmedial,

subterminal, and terminal wavy diffused fuscous brown bands,

all of them darker on inner margin,

Expanse 22 mm.

Hab. Castro, Parana, Brazil.

Subfam. GEOMETRIN&.

OosPILA ALTONARIA, sp.n. (Pl. IIT. fig. 21.)

Male.—Palpi inwardly white, outwardly tawny; pectus white;

legs ochreous, fore legs suffused with tawny ; frons purple-brown,

24%

350 MR. E. DUKINFIELD JONES ON

a whitish anterior bar; vertex and antenne shaft white, pectina-

tions ochreous; thorax green; abdomen white; dorsum rosy

brown. Wings emerald-green irrorated with darker green scales.

Fore wings: costa tawny yellow; a large black discocellular spot

with white centre and surrounded with white, the white scales

having rosy reflection ; termen crenulate, dark purple-brown ;

cilia rosy brown, white on the interspaces. Hind wings: a

smaller discocellular spot. Underside white.

Expanse 32 mm.

Hab. Alto da Serra, Santos.

TACHYCHLORA FLORA, sp.n. (PI. III. fig. 22.)

Male.—Palpi white; pectus and fore trochanters green; legs

ochreous white; frons green, anteriorly white; vertex white;

back of head green ; antenne shaft white, pectinations ochreous ;

abdomen green at base, segments 4 to 6 grey, anal segments

white. Fore wings emerald-green; costa white; a minute

black discocellular spot; a subterminal line of minute brown

spots on the interspaces, the one above the discal fold larger than

the rest; cilia white. Hind wings emerald-green ; a large ante-

medial yellow space from vein | to subcostal nervure, outwardly

nearly covered by a rosy-brown patch inwardly excavated in cell

and containing dark discocellular bar; a postmedial series of

minute brown spots on the interspaces, the one above discal fold

the largest. Underside white, a very conspicuous discocellular

bar on hind wings.

Expanse 25 mm.

Hab. Castro, Parana, Brazil.

Fam. URANIIDA.,

EPIPLEMA ORNATA, sp.n. (PI. III. fig. 23.)

Female.—Palpi and frons fuscous brown; legs brown, the fore

pair suffused with fuscous, mid- and hind-tarsi ferruginous

brown ; vertex of head and body greyish brown ; antenne shaft

ringed with purplish. Fore wings ochreous grey, the costal area

suffused with greyish brown to postmedial line, the suffusion

expanding downwards on medial area, forming a triangle with

apex at vein 3 defined inwardly by a dark line from cell to post-

medial line on vein 2; a fine dark wavy indistinct antemedial

line excurved through cell; postmedial line wavy, inwardly

ferruginous, outwardly dark, followed by whitish shade; outer

area grey with light strigulate suffusion of greyish brown; a fine

dark subterminal line and ferruginous suffusion; cilia light

brown with dark centre line. Hind wings ochreous grey ; ante-

medial line very dark, slightly wavy, forming a right angle on

fold in cell; a medial line parallel with antemedial from inner

margin to vein 4, thence slightly diverging to excavation on

NEW MOTHS FROM SOUTH-EAST BRAZIL. 351

costa; the area enclosed between the two lines very dark grey-

brown and ferruginous, extending in a rounded projection beyond

medial at costa; a wavy dark postmedial line on dark suffusion

with some ferruginous brown; cilia ferruginous with dark

scattered scales.

Expanse 18 mm.

Hab. Guaruji, Santos.

EPIpLEMA RECTANGULARIA, sp.n. (Pl. III. fig. 24.)

Female.— Palpi, legs, head, antennee, and tegule dark brown ;

base of antenne ochreous white in front; thorax and abdomen

greyish brown suffused with rufous, base of abdomen dorsally

purple-brown. Fore wings ochreous brown suffused with fuscous

on upper half and rufous on lower half; obscure traces of an

antemedial line angled outwards on subcostal nervure and vein 1 ;

a wavy dark diffused postmedial line bent inwards on discal fold,

followed by ochreous streak from inner margin to vein 2; apex

creamy white; a dark purplish-brown suffusion at tornus; a sub-

terminal series of black points on the interspaces ; cilia ochreous

and brown, black points at the veins. Hind wings ochreous

heavily suffused with rufous brown on upper half of basal and

medial areas ; a dark basal line; a dark antemedial line forming

a right angle in the cell; postmediai line dark purple-brown

from excavation on costa to vein 4 where it forms a right angle

straight to inner margin, followed by rufous shade containing a

fine strigulate steel-grey line; a dark suffusion on outer area

between veins 3 and 6; a dark wavy subterminal line preceded

by greyish-brown strigule and followed by rufous; a whitish

terminal line; apex creamy white with a few dark irrorations ;

cilia ochreous brown and fuscous.

Expanse 24 mm.

Hab. Castro, Parana, Brazil.

SACCOPLOCA STRIGARIA, sp. n, (PI. ITT. fig. 25.)

Male.—Palpi, legs, head, and antenne rufous brown; body

light brown. Fore wings light brown striated with purplish

brown; some white strigule on basal area; inner half of wing

darker brown ; a ferruginous-brown antemedial line oblique from

costa to cell, where it makes a rounded right angle and then runs

straight to one-third on inner margin ; postmedial line oblique

from costa to vein 7, thence straight to middle of inner margin ;

cilia concolorous. Hind wings light brown, less heavily striated ;

the basal area darker than the rest of the wing; costalarea white

excepting at apex; antemedial and medial lines from inner

margin to subcostal nervure, the medial being in continuation of

postmedial of fore wing.

Expanse 27 mm.

Hab. Castro, Parana, Brazil.

Bea - MR. E, DUKINFIELD JONES ON

NEDUSIA CASTRA, sp.n. (PI. III. fig. 26.)

Male.—Palpi and frons purplish brown; legs light fuscous

brown; vertex of head and antenne brown; patagia light brown ;

thorax and abdomen purplish brown, the anal tuft golden buff.

Fore wings light brown suffused, striated, and irrorated with

fuscous brown; a dark wavy medial line strongly excurved

through the end of the cell, slightly bent inwards from cell to

submedian fold, angled outwards on submedian fold and inwards

on vein 1, some white before it-in the cell; postmedial line wavy,

dark ferruginous brown, slightly incurved from costa to vein 8,

excurved from 8 to 2,then bent outwards and downwards to

inner margin; terminal area lighter than rest of wing, suffused

with darker from inner. margin to vein 4; a curved subterminal

dark streak between 4 and 7 followed by ferruginous; a dark

point between 7 and 8; cilia dark. Hind wings light brown

suffused with fuscous brown; the lines dark, parallel, angled out-

wards above median nervure, the inner preceded and the outer

followed by white; medial area dark fuscous brown; cilia dark.

Expanse 28 mm.

Hab. Castro, Parana, Brazil.

ALAPLENA, gen, nov.

Proboscis short; palpi slightly upturned, evenly scaled, third

joint moderate; frons smooth; eyes large, round; antenne of

male simple; thorax clothed with scales; patagia terminating in

hairs and long spatulate scales; legs smoothly clothed with

scales; spurs moderate, the outer three-fourths of inner;

abdomen smooth. Fore wings triangular ; costa slightly arched

at base and hollowed at middle; apex rounded; termen straight ;

inner margin slightly excavated before tornus; a small fovea

below vein le, distorting it and the median nervure, vein 1¢

becoming obsolete before origin of vein 2; vein 2 from near end

of cell; veins 3 and 4 separate from lower angle of cell; vein 5

from just above middle of discocellulars; veins 6 and 7 from

upper angle; veins 8-9 and 10-11 stalked from cell. Hind

wings: costa highly arched; apex square; termen nearly

straight; inner margin folded over on the underside, the fold

fringed with long spatulate hair-like scales; on the terminal area

of the upper side the scales are raised and minute; the median

nervure, submedian veins, and discocellulars aborted, the only

veins remaining being 6, 7, and 8; 6 and 7 from subcostal ner-

vure at middle of wing; vein 8 approaches subcostal only at base,

where it bends violently towards costa and arched to costa.

Type, A. casiraria.

ALAPLENA CASTRARIA, Sp.n. (PI. III. fig. 27.)

Male.—Palpi and frons dark rufous brown ; legs light brown ;

vertex of head white; antenne shaft white ; tegule, patagia, and

NEW MCTHS FROM SOUTH-FAST BRAZIL. 353

thorax dark rufous brown; abdomen lighter brown. Fore wings

light brown suffused and striated with fuscous; a medial fer-

ruginous brown line outwardly shaded with fuscous, straight

from inner margin to lower angle of cell, then bent to disco-

cellulars; a whitish spot in the end of the cell; postmedial line

sinuous, ferruginous brown inwardly shaded with fuscous, ex-

curved from costa to vein 3, incurved from 3 to 1, followed by

broad diffused whitish band; a fuscous-brown subterminal line

from apex to vein 3, broad at apex, narrowing to vein 3 and

broken at the veins, a narrow whitish shade. on inner side;

terminal area broadly grey; termen dark; cilia ferruginous.

Hind wings light brown suffused with dark ferruginous on basal

half above inner margin; terminal area brownish grey; a fer-

ruginous brown antemedial line from costa to subcostal nervure ;

a medial line straight from before middle of costa to just beyond

origin of vein 7, incurved below 7; obscure traces of a fine post-

medial line.

Expanse 24 mm.

Hab. Castro, Parana, Brazil.

Fam. LASIOCAMPID4.

ARTACE REGALIS, sp.n. (PI. I. fig. 12.)

Male.—-Pure white. Palpi outwardly black, inwardly white; fore

tibie and tarsi black, ringed with white ; antennz rufous more.

Fore wings: subbasal black spots at origin of vein 2 and below

costa; antemedial line of black spots on veins 1-2, on median

nervure at origin of vein 3, subcostal nervure, vein 11, and costa ;

a black point on the angles of the cell; a postmedial line of spots

from middle of inner margin nearly straight to vein 8, where it

bends inwards to costa; a subterminal line of ue elongated

spots slightly incurved from inner margin to vein 3, thence ex-

curved to vein 10, the spot on costa distally displaced ; a sub-

apical row of rather obscure spots above veins 5-8; a terminal

row of black spots on the interspaces; cilia white. Hind wings

pure white.

Female similar.

Expanse: male 43 mm., female 70 mm.

Hab, Castro, Parana; Alto da Serra, Santos.

Fam. PEROPHORID.,

PEROPHORA THERMESIA, sp.n. (PI. I. fig. 11.)

Male.—Palpi dark brown, rufous beneath ; legs brown suffused

with rufous and irrorated with lilacine white scales tipped with

black; frons anteriorly brown, posteriorly rufous; vertex and

antenne reddish brown ; tegule pale*reddish brown with a few

black irrorations; thorax light brown suffused with Jilacine ;

abdomen light yellowish brown, Fore wings lilacine irrorated

with white scales tipped with black ; costa light brown, becoming

354 MR. E. DUKINFIELD JONES ON

darker before apex; a very wavy antemedial line followed by

light brown suffusion on medial area; a dark brown postmedial

line excurved between costa and vein 8, and abruptly angled out-

wards below 8, thence straight to inner margin at two-thirds

from base, followed by narrow lilacine and a brown suffusion,

which is confluent with a dark subterminal shade from discal

fold to tornus, a red suffusion beyond the line below apex ; two

square brown spots on costa at one-third and two-thirds; termen

dark brown ; cilia tipped with lilacine; a minute hyaline spot at

end of cell. Hind wings ochreous liJacine, deeply suffused with

reddish brown, the veins on outer half suffused with red; a dark

medial line excurved from costa to vein 2, thence straight to

inner margin, followed by light shade; a diffused dark sub-

terminal shade; a minute hyaline spot at end of cell; termen

brown; cilia lilacine. Underside: fore wings heavily suffused

with reddish brown; a dark somewhat diffused postmedial line,

oblique from costa to below vein 8, where it is angled outwards,

straight to 5, then wavy to inner margin; a red subterminal

space from 4 to 8; the veins beyond the cell slightly suffused

with red. Hind wings less heavily suffused; a slight red suf-

fusion beyond upper angle of cell; a dark diffused wavy post-

medial line.

Expanse 41 mm.

Hab, Alto da Serra, Santos.

Fam. MEGALOPYGIDS&.

CARAMA INCOLORATA, sp.n. (PI. I. fig. 2.)

Male.—Grey ; palpi, legs, head, and thorax suffused with light

brown ; antennz lght yellowish brown; abdomen grey ringed

with yellowish. Wings satiny grey, rather darker at costa.

Expanse 31 mm.

Hab. Castro, Parana, Brazil.

NaRope CINGULATA, sp.n. (PI. I. fig. 1.)

Male.—Palpi and pectus black; fore legs black with white

hairs, mid- and hind-legs white, the tarsi fuseous with white

scales on upper side; head very pale primrose yellow; antenne

tawny, the shaft white; thorax pure white; abdomen white

with a broad band of orange on each segment. Fore wings pure

glossy white; costa narrowly ochreous; antemedial, medial, and

postmedial depressions and ridges. Hind wings pure white.

Expanse 30 mm.

Hab. Alto da Serra, Santos.

Fam. THYRIDID4&.

RHODONEURA OXYDATA, sp.n. (PI. I. fig. 19.)

Male.—Palpi light reddish brown, third joint dark; legs

ochreous, the fore pair suffused with fuscous; head and antenne

NEW MOTHS FROM SOUTH=HAST BRAZIL. 575)

light reddish brown ; tegule fuscous brown; patagia anteriorly

dark, posteriorly light brown; thorax dark brown; abdomen

dark brown, the first segment and a dorsal line on segments 3-5

ochreous ; anal tuft ochreous ; ventral surface ochreous with dark

central suffusion. Fore wings light brown striated with silvery-

blue strigule; a series of inverted V-shaped marks on costa; a

broad reddish medial shade from near tornus to upper angle of

cell, with dark suffusion at end of cell ; discocellulars circled with

silvery blue; a blue circle between veins 2 and 3 at lower angle

of cell; two small silvery-buff spots beyond the medial shade

between veins 5 and 6; a large spot of the same colour and

crossed by brown bar and striations from 5-7 and _ two

triangular spots below 5; a large silvery-buff spot above tornus

and some small ones on termen and inner margin ; a subterminal

series of silvery-blue spots below veins 4, 6,7, and 8; cilia brown.

Hind wings silvery buff heavily suffused with brown and striated

with silvery blue; a reddish medial shade most conspicuous

above vein 1; a golden suffusion on the submedian fold contain-

ing a black spot and some black points beyond ; some subterminal

black points below the fold ; apical area dark brown.

Expanse 25 mm.

Hab. Alto da Serra, Santos.

Fam. PyRALID&.

PoLYGRAMMODES EXrMIA, sp. un. (PI. I. fig. 6.)

Female.—Palpi: Ist joint white, 2nd and 3rd black; legs

white, the fore tibiae tinged with purple and ending in black ;

tarsi ringed with black; frons white with black at sides; head

and antenne white; tegule white with large chocolate central

spot; thorax white with large posterior dorsal and two subdorsal

chocolate spots; patagia white with central chocolate spot ;

abdomen white with transverse chocolate bands and short lateral

transverse streaks on segments 2, 4, 5, and 6, segment 7 orange,

anal tuft white. Wings white. Fore wings: a large basal

chocolate spot; three subbasal spots on costa, in cell, and on

inner margin; an antemedial chocolate band, broader on costa

and angled outwards on median nervure, followed by spot in cell ;

a medial series of three bars, one on discocellulars, one below

vein 2 to submedian fold and one from vein ! to inner margin ;

a postmedial bar from costa to vein 5 and series of elongated

spots between the veins ; a subterminal series of elongated spots

from below vein 8 to inner margin, confluent with postmedial

series between 1 and 2 and approximate between 3 and 5; a

terminal orange band expanding below vein 2 and broad at apex ;

an orange shade beyond lower angle of cell; cilia white, black at

the veins. Hind wings: a medial series of four chocolate spots ;

a postmedial series of spots above veins 2-5; a subterminal

series of elongated spots; a terminal orange band, expanding

356 ON NEW MOTHS FROM SOUTH-EAST BRAZIL,

below vein 2 and at apex; a slight orange shade beyond lower

angle of cell.

Expanse 37 mm.

Hab. Alto da Serra, Santos.

Fam. ZyG @NID &.

HARRISINA MEPHISTO, sp. n.

Male.—Black with purple, blue, and green reflection ; tegule

red; red lateral patches at base of aionen almost meeting on

orcuae

Expanse 28 mm,

Hab. Alto da Serra, Santos.

EXPLANATION OF THE PLATES.

Pruate I.

Fig. ) Big.

1. Narope cingulata. ¢. 13. Lobeza irrorata. 9.

2. Carama incolorata. 6. 14. Acanthodica frigida. 2.

3. Bryocodia paulina. &. 15. Moresa mona. :

4. Tarache parana. 6. | 16. Phurys fasciata. 6.

5. ‘i 4h : | 17. Mictochroa paulata. @

6. Polygrammodes eximia. &. | 18: a pallidula. 9.

7. Aplogompha yaponaria. 6. | 19. Rhodoneura oxydata. 6.

8. 5 fumaria. 6. | 20, Merocausta felinaria.

9. setinaria. 6. | 21. Aplogompha castraria. 6.

10. Rosema pauladaa wD) as 22. Bagodares castra. 9.

11. Perophora thermesia. @. 23. ee columbaria. &.

12. Artace regalis. 2. , . }

Puate II:

1. Boarmia nigraria. &. 14. Azelina cetana. 6.

2. Hymenomima sinuosariaz. 15. Callurapteryx paularia. 6.

3. >» > 2, 16. Cratoptera fenestraria. 6.

4. ee nivacaria. . 17. Apicia geminimacula. g.

5. Larentiopsis costiplaga. 6. 18. Trotogonia castraria. 6.

6. Rhodomena santaria. &. 19. Isochromodes elegantaria. @.

ihe i paularia. 2. 20. Paracomistis tristaria.

8. Metasiopsis proutaria. ¢. 21. Campatonema marginata. 6.

9. Tetragonodes geminaria. &. 22. Nephodia bonitaria. §.

10. Polla acutaria. &. 23. » paularia. 6.

ll. Numia strigularia. 6. 24, Apicia strigularia. 6.

12. Oxydia pallidaria. @. 25. Hypolepis bella. 8. °

13, Azelina hanebaria. 6. 26. Hariodes flavicilia. 6.

Prate IIT.

1. Hupithecia mauvaria. &. 15. Ptychopoda campinaria. &.

2. 5 aquanivaria. &. 16. % lilacaria. 6.

3. Anisodes bizaria. 3. | 17. Hyria gemmaria. &.

A. ss paranaria. &. 18. Psaliodes awrantaria. §.

5. Ay carolina. ®. 19. Hyriogona santaria. a

6. ty Jjaparia. @. 20. Ss montaria. 6.

fe uf antennaria. 6. 21. Oospila altonaria. 6.

8. . vigoraria. &. | 22. Tachychlora flora. &

9. Hupithecia cuprearia. ¢. | 23. Epiplema ornata. §&.

10. Hemalia mantaria. 9. 24. fe, rectangularia. 2.

1 i magitaria. &. 25. Saccoploca strigaria. 6.

12. Eoislutearia. 6. 26. Nedusia castra. 6.

13. Ptychopoda terminaria. %. 27, Alaplena castraria. 6.

14. i delicataria. §. | :

ON NEW OR LUPTLE-KNOWN ACARI. 357

19. On some new or little-known Acari, mostly Parasitic

in Habit. By Sranuey Hirsr, F.Z.S.

(Submitted for Publication by permission of the Trustees of the British Museum.)

[Received January 18, 1921: Read April 5, 1921. |

(Text-figures 1-15.)

The present note deals with a miscellaneous series of Mites

from various localities, including a number of new Gamasid

mites found by Mr. James Waterston in the nasal cavities of

Scottish birds, also several new ectoparasites from various sources.

Demodex is recorded from several new hosts, viz., deer, long-eared

bat, mole, and water-rat, three of the species being regarded as

new. An interesting pseudoparasite of flies is also described,

and a new English jumping-mite (Speleorchestes),

Text-figures 1-7 were drawn by Mr. E. J. Engel Terzi, nos. 8-15

by Mr. Percy Highley.

Family GAMASLID&.

RHINONYSSUS CALEDONICUS, sp. n. “(Text-fig. 1.)

2. Abdomen not elongated. Hairs on venter mostly only

slightly thickened at the base, the posterior ones situated just in

front of the anus are not dentiform, and there is only one pair

in this position. Genito-ventral plate shorter and wider than in

R. neglectus and R. waterstoni. Legs of moderate length, those of

the first pair apparently sometimes longer than the others, Legs

furnished with spinules, but they are minute. Claws of first leg

with a distinct short process or denticle dorsally.

Length *92-1:01 mm.

Habitat. Nasal cavities of Uria grylle (Black Guillemot), North

Mavine, Mainland, Shetland Islands: Gluss Voe (25. 11. 1912),

and Gluss Point (17. x. 1912), and Voe, Mainland, Shetland

(2. 111. 1912). Specimens collected by James Waterston.

RHINONYSSUS LEVINSENI Tragardh.

Sommatericola levinsent Triigirdh, Monograph arktisch. Acar.

in Rémer and Schaudinn’s Fauna Arctica, 1905, iv. pp. 28-30,

text-figs. 42-47, also pl. 1. figs. 1, 3, and 8.

Trigirdh’s genus Sommatericola must, I think, be regarded as

a synonym of hinonyssus Trouessart. His species Sommaterr-

cola = Rhinonyssus levinseni (from nasal cavities of Sommateria

mollissima) seems from his description to be closely allied to 2.

scoticus, but he depicts more spinules round the anus than are

yresent in that species, and also shows the lateral spines on the

palp as much stronger than in 2. waterstont.

Text-figure 1.

pavalita

. fle

Gy SS

Rhinonyssus waterstoni, sp.u., 2. Ventral view.

ON NEW OR LITTLE-KNOWN ACARI. 359

RHINONYSSUS WATERSTONI, sp. n. (Text-fig. 2.)

2. Abdomen not elongated. Very minute spinules are present

on the venter in this species, instead of the hairs that are present

in R. caledonicus. Capitulum short; segments of palp very

short, being very much wider than long; tarsal segment very

small, the conical tubercle on it well developed. Legs not very

long, the first pair apparently slightly shorter than the fourth.

Coxe with very short spinules or hairlets instead of the fairly

long hairs present in &. caledonicus. Spines on legs much weaker

than in R. neglectus. Claw of first leg apparently without any

dorsal process.

Length °96 mm.

Habitat. Nasal cavities of the Razorbill (Alcea torda), Ollaberry,

North Mavine, Shetland Islands. Specimens collected by James

Waterston (15. xii. 1913).

RHINONYSSUS ECHINIPES, sp. n. (var. of A. neglectus?). (Text-

figs. 3 & 4.)

2. Abdomen not very elongated in the female sex. There are

three pairs of minute denticles or tubercles anteriorly on the

venter. The three pairs of denticles in front of the anus are

much smaller and weaker than in &. neglectus. Capitulum

apparently shorter than in 2. neglectus. Segments of the palp

also shorter ; the minute conical tuberele on the tarsus is un-

usually large in this species. Legs. Denticles on posterior cox

weaker than in R. neglectus or replaced by hairs with a somewhat

thickened base; the other segments of the legs furnished with

numerous strong denticles as in R. neglectus.

$. Abdomen apparently more elongated (conical) than in the

female.

Length, § 1°36 mm., 2 1°36 mm.

Habitat. Nasal cavities of Ringed Plover (Wgialitis hiaticola),

Queyfirth, North Mavine, Shetland Islands (5. i. 1914). Speci-

mens collected by James Waterston.

RHINONYSSUS NEGLECTUS, Sp. n. (Text-fig. 5.)

2. Abdomen not elongated. Three pairs of strong but short

spinules are present anteriorly on the venter, and three pairs of

very strong spinules (grouped closely together) are placed imme-

diately in front of the anal aperture. Capitulwm unusually

elongated. Palpi with the segments more slender than in &,

echinipes. Legs. First and fourth pairs of legs rather long, being

considerably longer than the second and third pairs. Denticles

on legs much stronger than in #. caledonicus and Rh. waterstoii.

Length 1°36 mm.

Habitat. Nasal cavities of Tringa striata (Purple Sandpiper),

myer

u ‘ds ‘sadenyaa snsshuo

“MOTA [BIQUAA 5 &

maya

MOTA [BIJUIA "PMT

( Ly

"€ OINSY-4X9q,

'p OINSH-4Xxoq,

ON NEW OR LITTLE-KNOWN ACARI. 361

Yell Sound, Ollaberry, North Mavine, Shetland Islands: Little

Roe Island (13. ii. 1913). Specimens collected by James

Waterston.

Text-figure 5.

Rhinonyssus neglectus, sp.n., 2. Ventral view.

RHINONYSSUS CONIVENTRIS Trouessart. (‘Text-figs. 6 & 7.)

@. Abdomen elongated (conical), Anterior spinules on venter

similar to those present in 2. neglectus, etc. ‘There is also a pair

of spinules situated not far behind the genito-ventral plate and

two pairs of minute spinules or tubercles near the anal orifice.

Legs. Last pair of legs greatly enlarged and furnished with very

strong spinules.

3. Fourth leg about as long as the first and not much stouter.

Length, 2,1:45 mm.

Habitat: Nasal cavities of the Turnstone (Strepsilas interpres) ;

North Roe, Mainland, Shetland. Specimens collected by James

Waterston (27. 11. 1911).

362 MR. STANLEY HIRST ON

Text-figure 6.

one: A ae

Rhinonyssus coniventris, Y. Ventral view.

LIPoNYSsUS BERLESEI, sp.n. (Text-figs. 8 & 9.)

2. Body fairly long oval, and sometimes slightly constricted

near the middle (behind the last pair of legs); the abdomen is

sometimes rather swollen, and may exceed the width of the

anterior part of the body; posterior end with a slight notch.

Dorsal shield wide anteriorly, but narrowed and coming to a

point posteriorly. There are some quite short hairs on the

scutum, chiefly placed near the margin, and also a few very

minute and inconspicuous hairs in the middle of its surface,

NEW OR LITLLE-KNOWN ACARI. 363

Text-figure 7.

Rhinonyssus coniventris, ¢. Ventral view.

Hairs on uncovered part of dorsum not very numerous, and

mostly shorter than those on the venter. Sternal plate wider

than long and with three pairs of long hairs, its anterior margin

is not well defined, being continued forwards by transverse linear

markings that reach the front end of the body. Genito-ventral

plate short, of moderate width, and the posterior end is blunt

(not pointed). Anat plate of the usual pyriform shape, but more

elongated than usual. Perttreme reaching forwards almost to the

level of the middle of the coxa of the first leg. Zegs normal in

appearance, being of moderate length. Second coxa with a long,

sharp anterior spine and a very slight rounded spur posteriorly,

Third coxa posteriorly with a sharp, well-developed inner spur,

and an inconspicuous outer denticle. Tarsus of second leg with a

pair of short pointed denticles at the end. The hairs on the legs

are shown too strong (spiniform) in the figures of this species.

They are more hair-like.

Length (including capitulum) «87-90, width -36—44 mm.

Habitat. Parasitic on Myospalax scansus, Shensi, North China ;

several specimens collected by Capt. H. EK, M. Douglas (24, x1.

1908).

Proc. Zoou, Soc.—1921, No. XXV, 25

~—

‘mata Testo, * 3 “u ‘ds ‘¢saz.taq susshuodyyT

+S “u'ds ‘psaquaq snsshuodvy

“MOIA [RI}TO A

1p fayySiyy

cS)

9 dINSY-4XOT,

6 9INST-}Xoq,

ON NEW OR LITTLE-KNOWN ACARI. 365

LIPONYSSUS SERPENTIUM Hirst.

This species was described from examples found on a Couper’s

Snake at the Society’s Gardens. I have examined some additional

specimens from the Horned Viper (Vipera cerastes), also from

‘our Gardens. The first-named host is a Mexican species, whilst

the latter is a North African form. This mite has also been

found on snakes that have died at the Bronx Zoological Park,

N.Y. It is believed that this parasite was introduced with the

Malayan Python (Python reticulatus), and, whilst it does not

harm that host, may leave it and attack American snakes of an

kind, giving rise to a kind of blister or eruption under the edge

of the scale, causing death. According to Mr. Ditmars, several

snakes have been lost from this cause. The above details, relating

to the American case, are taken from Mr, P.S. Falshaw’s letter

accompanying the specimens.

Liponyssus serpentium is easily distinguished from the allied

L. natricis by the presence of two dorsal shields instead of only

one. The genito-ventral plate is ornamented with longitudinal

linear markings, as in LZ. natricis.

LIPONYSSUS ARABICUS, sp. n.

2. There are two plates on the dorswm in this species, the

posterior being very minute and oval as in ZL. serpentiwm mihi,

but the anterior one is long and wedged-shaped, reaching far

backwards, and only separated from the minute posterior plate

by a comparatively short space (whereas in L. serpentium the two

plates are widely separated from one another). Anterior plate

furnished with a number of hairs. The minute posterior plate

does not bear any hairs. Hairs of body lacking minute off-shoot

present in Z. bacote mihi. The minute platelets present between

the main shields in L. serpentiwm appear to be absent in the

new species. Besides the usual fine striations of the integument,

there are a number of fairly long, but very fine chitinous (2)

linear markings, especially numerous at the sides of the body.

Venter. Sternal shield trapezoidal in shape, and with two pairs

of hairs on its surface. Genito-ventral shield long and narrow.

Anal plate pear-shaped ; the paired hairs on it shorter than the

unpaired one. Legs. Coxe without spurs; the other segments

with the usual fairly stiff hairs, but without any strong spines.

Length *73 mm.

Habitat. A single specimen from a lizard (Agama adramitana) ;

Ad Dthala, Upper Hushabi, South Arabia.

Lironyssus BURSA Berlese.

During a recent visit to the Natural History Museum, Mr. M.

Koidzumi brought me two specimens of a mite found on a

Chinese patient at Hoko T6 Island, Formosa. They are referable

to the tropical fowl-mite (Liponyssus bursa Berlese).

25%

366 MR. STANLEY HIRST ON

Family ListTROPHORIDS.

LISTROPHORUS FRONTALIS, sp.n. (Text-fig. 10 A & B.)

3g. Dorsal (frontal) process of capitulum much more strongly

salient than in ZL. argentinus, being fairly long and conical (text-

fig. 10 A). Abdominal lobes rather shorter than in LZ. argentinus,

and somewhat differently shaped (text-fig. 10 B).

Q. Process of capitulum similar to that in the male.

Length, 3 330-345 pw, 9 365 p.

Habitat. Parasitic on the rodent Orizomys delticola, from Isla

Ella del Delta, Parana.

Text-figure 10.

A, Anterior end of Listrophorus frontalis. B. Posterior end of abdomen of male of

the same species. C. Anterior end of Listrophorus argentinus. D. Posterior

end of abdomen of male of this mite.

LISTROPHORUS ARGENTINUS, sp.n. (Text-fig. 10C & D.)

&. Dorsal (frontal) process wider than long and rounded in

outline (text-fig. 10 C). Abdominal lobes ending in a delicate,

almost truncate process, the posterior edge being, however,

slightly oblique; the gap between the two lobes long and

narrow (text-fig. 10 D).

Q. Capitulum of female very like that of the male.

Length, 3 345-360 pb, 9 415-450 p.

Habitat. Argentina; on a rodent (Scapteromys tomentosus).

TrrcHacrus BREVIPES Can. & 'T'rt.

According to Canestrini and Kramer in ‘ Das Tierreich’ (Sar-

coptide, ete.), this interesting little Listrophorid mite has only

been found in Thessaly, and I cannot find a later rererence to the

species. As a matter of fact, it is not uncommon on English

rodents. IL have collected numerous specimens off Evotomys

glareolus britannicus ab Exeter, and off Microtus agrestis at

Barnes, Surrey.

A. Demodex chiropteralis. Ventral view

capitulum of same

view of capitulum of same species

very strongly chitinised, and furnished with unusually large

NEW OR LITTLE-KNOWN ACARI 367

Family DeEmMoDICID”&

DEMODEX CHIROPTERALIS, sp. n. (Text-fig. 11 A, B & C.)

2. This species differs from the small form of Demodea:

(D. soricinus) sometimes present in the skin of the long-eared

bat in its much larger size and stronger legs, the latter being

Text-figure 11.

1B,

5 |

Highley del

B &C, Enlarged view of foot and of

D. Demodex melesinus, 2 ;

ventral view. E. Dorsal

claws, the genual joint, or articulation, is very pronounced, and

the distal half of the limb readily ele

Vulvar opening

368 MR. STANLEY HIRST ON

elongated. Body wide, the abdomen pointed posteriorly. (Both

my specimens of this species are slightly damaged, and it is ditti-

cult to give the exact proportions.) Capitulum apparently

without any dorsal tubercle or spinule; it is rather like that of a

nymph in some respects.

Measurements. Votal length 229 pu, length of cephalothorax+

eapitulum 114-115 pw, of abdomen 115 »?; greatest width of

cephalothorax 59-61 jw, of abdomen ?.

Host. Long-eared bat (Plecotus auritus); two specimens col-

lected by the author (October 21st, 1919).

DEMODEX SORICINUS Hirst.

®. I have found several specimens of a small form of Demodea

in the skin of long-eared bats, which is very similar to that

found in rats and shrew-mice described under the name soricinus.

The capitulum in these specimens from bats is apparently without

any dorsal tubercle or spinule. The body is somewhat differently

shaped, the posterior end of the abdomen being blunt (almost

rounded), instead of pointed as in typical specimens of the

species. Capitulum considerably wider than long. Abdomen a

little shorter than cephalothorax+capitulum ; bodya little more

than three times as long as its width.

Measurements. Total length 111-128 1; length of cephalo-

thorax +capitulum 63-75 p, length of abdomen 52-53 pw, width

of cephalothorax 33-38 p, width of abdomen 33-36 p, of ceapitulum

18-23 w, length of capitulum 13-19 p.

Material. Several specimens from live long-eared bats (Plecotus

auritus), collected by the author, October 1919.

DEMODEX GLIRICOLENS, sp. n. (var. of D. arvicole?). (Text-

fig, 12.)

3. Body varying rather considerably in shape. One specimen

is rather narrow and elongated, and about five times as long as

wide. Abdomen of this example distinctly longer than the

cephalothorax+capitulum. Capitulum rather narrow and fairly

elongated (about as long as wide).

Another specimen is much shorter and comparatively wide,

being about three times the width of the cephalothorax. The

abdomen is slightly shorter than the united length of cephalo-

thorax and capitulum. Capitulum of this specimen rather wide,

being wider than long.

Posterior end of abdomen rounded off in this species. Spinule

on dorsal surface of capitulum smaller and more siender than is

usually the case in D. arvicole.

Genital opening of male situated on the dorsal surface above

the interval between the first and second pairs of legs, in this

respect resembling D. erminew; the opening is, however, very

simple, consisting merely of a curved line.

Measurements. Slender example. Total length 144 p, length

of cephalothorax+capitulum 70 p, of abdomen 74 pw, greatest

NEW OR LITTLE-KNOWN ACARI.

_ Text-figure 12.

Demodex gliricolens, 3.

A & B. Dorsal views, showing variation in shape of body

369

370 > MR. STANLEY HIRST ON

width of cephalothorax 29 p, of abdomen 30 p, length of

capitulum 18 yu, its width 174. The short wide form measures

as follows :—Total length 103 jy, length of cephalothorax+

capitulum 56 yc,-0f abdomen 47 pv, greatest width of cephalothorax

35 , of abdomen 36 p, length of capitulum 15 p, its width 15 p.

Host. Water-rat (Arvicola amphibius); two male specimens

collected by the author from freshly-killed examples of this

mammal (July 1919). E

Text-figure 13.

Highley del

Demodex talpe.

A. Dorsal view of female. B. Ventral view of male. C. Penis greatly enlarged.

Note.—The sexual orifice of the male is really dorsal in position and

the penis situated in the median line.

DeMoDEX TALP&, sp.n. (‘Text-fig. 13.)

2. Body about four times as long as the width of the cephalo-

thorax. Cephalothorax usually rather narrow in front, becoming

a little wider posteriorly. There is a well-defined longitudinal

NEW OR LICTLE-KNOWN ACARI. 371

groove running down the middle of the cephalothorax, which

always seems to be present, and is the chief distinguishing

feature of the species. This groove is limited laterally by a

rather sharp line, which sometimes has the appearance of being

double (owing to the depth of the groove). Abdomen sometimes

with the sides slightly convex, and it is then distinctly wider

than the cephalothorax, in other specimens the abdomen is of

fairly uniform width throughout its length; it is a little longer

than the cephalothorax. Capitulum fairly elongated, but still

it is slightly wider than long. Spinule on its dorsal surface

flattened in much the same way as in D. arvicole, and pointed

distally (internally), the outer angle distinct. The spinule is

fairly thick, for under very high magnification a distinct posterior

edge can be seen; it is fairly large and wide as compared with

the size of the capitulum. One of the little claws or denticles on

the ventral surface of the terminal segment of the palp is much

longer than the others.

3. Male very like the female in general appearance and with

the dorsal groove well-developed. Body about four times as long

as width of cephalothorax, the abdomen being longer than the

cephalothorax+capitulum. Male sexual orifice situated on the

dorsal surface above the interval between the second and third

pairs of legs. Penis quite long and slender.

Measurements. 9. Total length 128-130; length of cephalo-

thorax+eapitulum 59-62 mw, of abdomen 68-74 4; greatest

width of cephalothorax 31-34 », of abdomen 34-41 p; length of

capitulum 15-19 w, its width 19-20. o¢. Total length 126 yp;

length of cephalothorax+capitulum 54, of abdomen 72 p, of

penis 224; greatest width of cephalothorax 30-33 yw, of abdomen

37“; greatest width of capitulum 21 p, its length 16 p.

Material. Several specimens obtained by the author from

freshly-killed moles (Z'alpa europea); May 1919.

DEMODEX BOVIS Stiles.

I have examined specimens apparently referable to this species

collected by Dr. E. Brumpt from deer. The material consists of

a small piece of skin, preserved in spirit, containing a number

of pustules filled with whitish matter and numerous parasites in

all stages of development. ‘The material is not very well pre-

served. The following is a description of the adult specimens :—

a little more than three times to about three and a half times as

long as the cephalothoracie width. Abdomen not quite so sharply

pointed as in typical specimens of D. bovis; it is nearly always

distinctly longer (sometimes considerably longer) than the

cephalothorax+capitulum, but occasionally only about the same

length. ‘Capitulum wider than long. Spinule on its dorsal sur-

face moderately long and distinctly bent or angular near the

distal end.

Si MR. STANLEY HIRST ON

3. Male sexual orifice situated well forward on the cephalo-

thorax above the first pair of legs; the minute lobes, etc., round

it are exactly as in specimens from cattle. Males are very rare

in the slides mounted from the material, and the measurements of

the male sex given below are based on very few examples.

Measurements. 9. Total length 180-238 yp; length of cephalo-

thorax+capitulum 90-101 pu, of abdomen 90-143 ww; greatest

width of cephalothorax 58-65 yw. (rarely 70), cf abdomen 53-60 pm.

3. Total length 199; length of cephalothorax + capitulum

79 wu, of abdomen 1204; greatest width of cephalothorax 61 jp,

of abdomen 64 p.

_ Host. Material from a deer (species?) at Chantilly, France

(Dr. LE. Brumpt).

DEMODEX MELEsINUS Hirst. (Text-fig. 11 D & E.)

The Demodex of the badger is of unusual interest, for of all the

known forms it most closely resembles that present in man (D.

folliculorum), but is smaller in size and has the capitulum more

elongated.

DEMODEX CANIS var. ERINACEI Hirst.

Since writing my paper on Demodex, | have obtained a number

of specimens of this variety from the head of a hedgehog caught

by the Museum taxidermist (Mr. A. H. Bishop) at Dymchurch,

Kent (8. vii. 1919). They are elongated specimens, agreeing

fairly well with those on which my description was based. These

additional examples measure as follows :— 9. 312-370 p (a large

bent example of this sex probably measures about 400).

3. 232-292 u.

Family TROMBIDIID&.

TETRANYCOPSIS HORRIDA OC. & F.

This ‘* Red Spider” has not been recorded from England. I

have seen an example collected by Mr. K. G. Blair, on hazels at

Shoreham, Kent, and during an excursion to that locality I

obtained another specimen myself. The species, apparently, is

not very abundant, even where it occurs. Numerous specimens

of the lime-tree mite (Zetranychus tiliariwm) occurred on the

hazels in this locality.

Genus PsorERGATES.

In some details of structure, the immature stages of the mites

of the genus Psorergates resemble those of the genus Demodeu.

For instance, the feet of the larval and nym/phal stages are in-

complete, consisting of an epimeron, shaped almost exactly as in

Demodex, both its ends being angular and supporting a roughly

circular disc-shaped segment, bearing processes or claws fur-

nished with denticles of a similar type to those present in the

immature stages of Demodex. In spite of the difterence in shape

NEW OR LITTLE-KNOWN ACARI. 373

and general appearance, it seems probable that the genera

Psorergates and Demodex are related to one another.

SPELEORCHESTES VENTRIOSUS, Sp. n.

@. This new saltatorial mite differs from S. podwroides, mihi

in being much stouter in appearance; the abdomen is especially

wide (hence the specific name ventriosws), being much stouter

than in S. poduroides. Ovipositor when protruded provided with

a basal collar-like structure or sheath as in S. poduroides, but it

is longer than in that species, its length exceeding the distal part

of the ovipositor that projects beyond it (whereas in S. poduroides

the collar is much shorter than the part of the ovipositor that

projects beyond it). Shape of hairs and their distribution much

as in S. podwroides.

Owing to its rather squat appearance, this new species of

Speleorchestes somewhat resembles Vanorchestes, but the anterior

part of the cephalothorax is quite separate and distinct from the

posterior portion, not being enclosed by it laterally ; posterior

part of cephalothorax much narrower than the abdomen and

divided from it by a distinct constriction (whilst in Manorchestes

the posterior part of the cephalothorax is almost as wide as the

abdomen and only separated from it by a slight groove).

Length (including chelicer) 317 » ; width of anterior (cephalic)

part of cephalothorax 62-67 ; width of posterior (thoracic) part

of cephalothorax 107-110 » ; width of abdomen 155-160 p.

Habitat. Hindhead, Surrey; a number of specimens collected

by the author under stones near the path running above the

natural hollow known as “the Devil’s Punch Bowl” (just

below Hindhead Beacon), May 1918.

SPELEORCHESTES PODUROIDES Hirst.

I have added a few more measurements for comparison with

those of S. ventriosus, sp. n.

Length (including chelicere) 278-290; width of anterior

(cephalic) part of cephalothorax 58-60; width of posterior

(thoracic) part of cephalothorax 82-85 x; width of abdomen 106—

1154.

Key to British Species of Saltatorial Mites (Genera Vanorchestes

and Speleorchestes).

Body short and wide; anterior part of cephalothorax en-

closed laterally by the posterior part........ ............. Nanorchestes Tops.

& Trouess.

Dorsal hair on chelicera slender and divided into two

PluMTOSe | PLAT CMeS css seeeee cee seeeeeceeeeeeree ea eceee reese ONCOL US PEUUUS

(Mendip Hills).

Dorsal! hair on chelicera rather short, stiff, rod-like, and

TAOVW CONIC! jn ch pga abc suede beoncodac use ses novde Jypesonbanescns | ENG GAP AGORS IOS:

& Trouess.

(Littoral species.)

Body more elongated; anterior part of cephalothorax

free (not enclosed laterally by posterior part) ......... Speleorchestes Trgdh.

374 MR. STANLEY HIRST ON

Body fairly narrow; the part of the ovipositor projecting

beyond the collar-like sheath longer than the sheath

IbSEL TS) jase SEEMS Mae Ry on MEA a AMEE NL Seren Ne SM MOU UROL EORELINS b

(Malvern Hills.)

Body wider ; sheath of ovipositor longer, the part pro-

jecting beyond the sheath shorter than the sheath

TUUSeLE SAMAR he RRR ROOT ern. Oe Forti eeneaah ad S. ventriosus, sp. n.

(Hindhead, Surrey.)

Family TARSONEMID &.

PYGMEPHORUS TARSALIS, sp. n. (Text-fig. 14.)

% Pigmephorus stercoricola Berlese, Redia, 1911, vii. p. 184.

Q. Tarsus of first leg ending in a large conical protuberance

situated above the claw, which is moderately developed. Striated

Text-figure 14.

Highley del.

Pygmephorus tarsalis, sp. n., §.

A. Dorsal view. 3B. Tarsus of first lee much enlarged,

sensory (so-called olfactory) hairs on first tarsus all slender, and

none of them are especially large; the distal one is not very

long, being very slender ; the two proximal ones are placed close

together, they are club-shaped, and one of them is longer than

€

NEW OR LITTLE-KNOWN ACARI. 375

the other (also slightly longer than the distal one), The remain-

ing (second) sensory hair is prone and closely applied to the

segment, and therefore is inconspicuous. Second tarsus with a

single striated sensory hair dorsally near the proximal end, and

it is comparatively stout. The pair of tiny oval structures

situated in front of the pseudostigmata are smaller and further

part than in P. americanus. Pseudostigmata not rounded off

distally (as is the case in P. spinosus and P. pilosus), but ending

in a point. There is only one pair of long hairs on the cephalo-

thorax (instead of three pairs as in P. americanus); each hair

being situated just in front of, and slightly to the side of, the

pseudostigmata. An exceedingly short and inconspicuous hair is

also present close to each of these two long hairs. The posterior

hair present on the cephalothorax in P. americanus is missing in

this species from mushrooms. ‘Tarsus of fourth leg with quite

long hairs. Hairs on body long and very slightly feathered, the

feathering being scarcely visible even under very high magnifica-

tion (instead of quite distinct as in P. spinosus and P. pilosus).

Measurements. Total length 217-297 4; width 100-132.

Habitat. Numerous specimens from off the top-surface of culti-

vated mushrooms; received through the kindness of Professor H.

Maxwell Lefroy.

PyGMEPHORUS AMERICANUS Banks. (Text-fig. 15.)

Pigmeophorus americanus Banks, Treatise on Acarina, P.U.N,

Mus. 1904, xxviii. p. 77, fig. 151 (figured without description).

Pigmeophorus americanus Banks, Proc. Ent. Soc. Washington,

vii. p. 139 (1905).

@. This species is chiefly distinguished from the others of the

genus by the position of the little oval structures or accessory

stigmata *? (situated on the cephalic segment in front of the

pseudostigmata), which are larger than usual and placed quite

close together (instead of widely apart as in P. tarsalis). Another

character is the shape of the pseudostigmata, which are not

elongated but short and globular, lacking the slight terminal

(apical) point present in P. tarsalis.

Two rather long hairs are present on each side of the cephalo-

thorax (instead of one long hair and a very short one as in

P. tarsalis), There is a third pair of very long hairs on the

cephalothorax situated just in front of the posterior margin.

The following details of structure may also be useful :—

Shape of body similar to that of P. tarsalis ; some specimens

appear to be much wider than others (as compared with their

length), but this is perhaps merely due to the pressure of the

cover-slip. Hairs on body distinctly plumose. First leg stouter

* Oudemans apparently considers these minute paired oval structures to be organs

of sense (see Arch. Naturg. 79, Abt. A, Heft 9, p. 113, 1913), but it seems to me to

be more probable that they are stigmata, for they appear to be connected with the

respiratory trachez,

376 MR. STANLEY HIRSY ON

than in P. tarsalis, resembling that of P. estivus in this respect.

First tarsus produced beyond and above the claw, but apparently

not so strongly as in P. tarsalis.

Striated sensory (‘‘ olfactory”) hairs on first tarsus rather

similar to those of P. tarsalis; they are four in number, all of

them being more or less club-shaped; the one situated near the

distal end is very short and slender, the next is the largest, being

rather stout, close beside it there is another slender but not very

short sensory hair, the sensory hair nearest the proximal end of

the tarsus is very short and slender. Second tarsus with a short

but fairly stout sensory hair near the proximal end ; the claws of

this leg are distinctly bifid.

There is an unusually stiff bristle on the dorsal surface of the

first leg (especially well developed in examples from Lyperosia

irritans (= Hematobia serrata)).

Length of body 205-225 w; width 110-140 pm.

Habitat. Pseudoparasitic on J/usca domestica (slides in collec-

tion of Laboratoire de Parasitologie, Paris) and Stomoxys calcitrans

(the specimen from the latter was collected by Dr. J. Burton

Cleland, and therefore is presumably Australian in origin). Also

numerous specimens (reddish in colour) found on an English

specimen of Lyperosia irritans.

Prof. T, Harvey Johnston and M. J. Bancroft mention in their

paper on the life-histories of Musca australis (MU. fergusoni) and

M. vetustissima (Proc. Roy. Soc. Queensland, 1920, xxxi. footnote

on p. 183) that ‘‘ These flies may be parasitised by larval mites,

a red one probably Acarus muscarum Linn., and also a minute

whitish species.” The latter is probably the species identified as

P. americanus in the present note.

PYGMEPHORUS AMERICANUS var. SOCOTRENSIS, var. noy.

@. Very like the typical P. americanus in most respects, but

with the sensory (striated) sete on the tarsi of the first and second

legs different in shape, the largest one on the first tarsus being

considerably elongated and cylindrical, whereas in 7. americanus

(typical form) it is club-shaped. The sensory seta of the penul-

timate segment of the second leg differs in a similar manner.

The paired oval structures situated near the front of the cephalo-

thorax are placed close together as in P. americanus, but are

constricted in the middle (perhaps differences like this are merely

due to the presence of air or gas in the preparation). Pseudo-

stigmata globular as in P. americanus. There is a stiff dorsal

bristle on the second free segment of the first leg, much as in

P. americanus.

Measurements. Total length 210 « ; width 125 wp.

Hab. Musca domestica: Socotra. A single specimen found on

a dry example of a fly from that locality.

NEW OR LITLLE-KNOWN ACARI. 377

Text-figure 15,

Highley det

Pygmephorus americanus, § .

A. Dorsal view. . B. Tarsus of first leg much enlarged.

PYGMEPHORUS SPINOSUS Kramer.

2. The first tarsus is very characteristic in this species, being

much enlarged, the claw also very large and moving against a

strong process or spine. There are four striated sensory hairs

(the so-called olfactory hairs), a distal pair of fairly long cylin-

drical ones being placed close together (one of them is slightly

stouter than the other), and a proximal pair of very short rather

club-shaped ones. Second tarsus with a single sensory hair

dorsally, which is not very long but fairly stout, situated near

the proximal end of thesegment. Pseudostigmata rounded at the

distal end. Hairs on body long and more distinctly feathered

than in P. tarsalis.

Habitat, Talpa europea; England and the Continent,

378 ON NEW OR LITTLE-KNOWN ACARI,

PyYGMEPHORUS PILOSUS Oudmns.

@. First tarsus not produced nor enlarged. The second

striated sensory hair (from the distal end) is much larger and

stouter than the others, the first being of moderate length and

slender; the two proximal sensory hairs are quite short, one of

them is slender but club-shaped, the other very slender, straight,

and cylindrical.

Second tarsus with a rather long spine in the middle of the

dorsal surface, and with a short spine and also a short club-

shaped sensory hair near the proximal end. Claws of this leg

slender and not bifurcated. Pseudostigmata rounded distally

(not pointed). Hairs of body and limbs more distinctly feathered

than in P. tarsalis.

Habitat. Arvicola agrestis and Talpa europea, England ; several

specimens off these hosts.

ACARAPIS, gen. nov.

I propose the new generic name Acarapis for Tarsonemus woodi

Rennie, 1921, the principal structural differences between this

new genus and Zarsonemus being as follows:—Anterior leg of

larva of Acarapis well-developed and furnished with a pair of

claws and a pulvillus, but the second and third legs are very

short (almost rudimentary) and without either claws or pulvillus.

(In Tarsonemus the larval stage has all three pairs of legs well-

developed and all end in claws and pulvillus.) The globular

pseudostigma always present between the first and second legs in

the females of Zarsonemus is absent in Acarapis. Fourth leg of

female shorter and stouter than in Zarsonemus, and furnished

with a larger number of hairs, resembling Scwtacarus in this

respect.

J. Green, del.

COCHLITOMA

PAYS. 19er Monistarty wolge

ors cnc

Bemrose, Derby.

ZEBRA var OBESA:and van FULGURATA.

PZ.S. 1921. Longstaff PLT

J. Green, del.

COCHLITOMA ZEBRA var OBESA.

Longstaff. PI. Ill.

PZ-S:

Green, del

COCHLITOMA.

COCHUTOMA.

ZEBRA, VAR. OBESA.

ZEBRA, VAR. FULGURATA.

VITTY

& SEABORNE

LONDON

LTO.

THE HABITS OF COCHLITOMA ZEBRA IN CONFINEMENT. 379

20. Observations on the Habits of Cochlitoma zebra, var.

fulgurata. (Pfeiffer), and Cochlitoma zebra, var. obesa

(Pfeiffer), in Confinement *. By Jane Lonasrarr,

FT. S., EGS.

(Communicated by Lieut.-Colonel RatpH Lonestar, D.S.O., F.Z.S.)

| Received January 24, 1921: Read April 19, 1921.)

(Plates I-III. +)

When visiting Cape Colony, during 1914, I obtained six

specimens of Cochlitoma zebra, var. obesa, at Humewood, Port

Klizabeth, on March 31st. From these I selected two examples,

which I brought home alive in company with a fine specimen

of the var. fulgurata (the latter was presented to me by

Mr. O’Connor, of the “ Woods and Forests,” who found it at

Keiskamma Hoek, about nineteen miles north-west of King

William’s Town).

On leaving Cape Town, on April 11th, I put all three together

in a tin box, which I opened frequently ‘during the voyage, so as

to feed them and allow them to crawl about. Upon arrival, on

April 28th, I placed them in the conservatory.

In bringing these molluscs to England I had no intention of

breeding them, but only surmised they might be of interest for

anatomical purposes—therefore I did not keep the two varieties

separate ; and subsequently there was great mortality among the

young through lack of preparation and inexperience.

Broods of original Specimens.

April 29th, 1914.—-The smaller example of var. obesa had depo-

sited during the night sixteen young molluscs and about thirty

CLES ; seventeen of the latter were either broken or punctured.

The two varieties were now placed in different pots, with sandy

soil, tufts of grass, and varied food-leaves ; also the parents were

separated from the eges and young in every case.

May 4th.—The larger specimen of the same variety deposited

a dozen or more living young ones and numerous eggs. No

fragments of egg-shells were then observed, but the next day I

noticed that two of the young were much smaller than the

others, and that one of them had a small bit of ege-shell

adhering to it—also that there were some pieces of egg -shell lying

near them. None of these fragments looked large enough to |

have contained the biggest of the young, but they might have

* T have followed Major Connolly (‘Annals of the South African Museum, vol. xi.

pt. iii. 1912, pp. 204, 205) in regarding these forms as varieties of Cochlitoma zebra

(Chemnitz).

+ For explanation of the Plates, see p. 387.

Proc. Zoou. Soc,— 1921, No. XX VI, 26

380 MRS. J. LONGSTAFF ON THE HABITS

held the smaller ones. Thus the two latter had evidently

hatched from the eggs after they were laid. I did not ascertain

the exact numbers at first, as I was afraid of disturbing them,

but I now counted seventeen young molluses and about fifty eggs,

many of which were cracked.

May 4th.—I discovered that var. fulgurata had deposited two

living young ones, which were buried in the soil, without any

eggs or fragments of egg-shell associated with them. These died

that day when on a visit to the British Museum (Nat. Hist.),

where a drawing of one was made.

May 7th.—Var. fulgurata laid about seventy eggs near the

surface, without any living young molluscs.

May 15th, 16th, 17th, 21st, 22nd, and 25th.—-On each of these

days an egg hatched, thus there were six young var. fulgurata

from the eggs laid on the 7th. Some of the other eggs cracked,

but with one exception no more inolluses came out.

June Ist.—This one hatched, but the animal died immediately

and the shell was cleared out, either by Agriolimax agrestis Linn.

(which had been introduced, probably on lettuce) or else by its

brethren.

September 9th, 11th, and 12th, 1915.—The larger specimen of

var. obesa was observed to have deposited another brood. On

each of these days a single young mollusc appeared on the sur-

face. I then scraped up the soil gently, to the depth of one or

one and a half inches, and discovered nine additional young ones.

Ten were about the same size, while two were smaller—there

were no eggs or fragments of egg-shell.

October 8th.—Two additional young ones were observed with

the parent. It is not evident whether these were overlooked

before (for young shells, when covered with soil, greatly resemble

pebbles), or whether they had been deposited more recently. One

was of about the same dimensions as that of the medium size, and

the other was about equal to, or rather less than, the smallest of

the previous ones.

October 31st, 1916.—This brood was entirely killed by a single

specimen of Arion hortensis, which I found beside them. Thus

they existed little more than a year.

Hibernation and Mortality.

May 7th—July 13th, 1914.—The var. fulgurata, after laying

egos, formed an epiphragm and between these dates buried itself.

Then it came up at night and ate lettuce, but went below during

the day, and continued doing this for about a fortnight, when it

remained completely buried till the beginning of October. At

this time it rose, and died on November 26th, after having been

in confinement eight months.

Winter, 1914-15.—The four young var. fulgurata, which alone

were living, commenced to hibernate in October and remained

below till March 4th, One, however, had protruded its head

OF COCHLINOMA ZEBRA IN CONFINEMENT. 381

and eaten lettuce on February 8th, and then retired again.

Only two survived the winter.

November 25th.—The two adult var. obesa, as well as the

young ones, commenced to hibernate, and they remained more or

less, but not continuously, inactive till February 8th, 1915.

May 24th, 1915.—The smaller adult var. obesa buried itself,

and as it did not rise again I dug it up on July 12th, when I

found it was dead, after being from fourteen to sixteen months

in confinement.

Winter, 1915-16: October 9th.—The two young var. fulgurata

hibernated. One came up January 6th, the other on the 29th.

November.—During this month the single adult and the young

var. obesa buried themselves till January. The former rose on

the 20th and remained on the surface till March 2nd, when it

died, after about two years in confinement.

Winter, 1916—-17.—Two young var. fulgurata began hibernat-

ing in September, but roused up and ate at the end of February;

the weather, however, becoming very cold caused them to go

below again till late in March. One died in June, when about

three years old.

November.—There were now only six young var. obesa living,

these began to hibernate during this month. They did not,

however, remain continuously below, but came up at different

times, though they did not eat. Three died in February and

two in March, thus only one specimen remained which belonged

to the first brood.

Winter, 1917—18.-—-The solitary specimen of var. fulgurata

began hibernating at the end of September and roused up at the

end of January. The single example of var. obese was only

dormant from December 15th to January 24th.

Winter, 1918-19.—Var. fulgurata hibernated from the first

week in September till February 22nd. Var. obesa from the

first week in October till February 21st.

Winter, 1919-20.—Var. fulgurata hibernated from September

29th till February, when it came up, but retired again below till

the middle of March. Var. obesa was buried from October 6th

till the middle of March.

The hibernation of the original var. fulgurata during our

summer may be explained by the fact of that season being dry

and cooler in the district whence it came, when it would naturally

rest. ‘There being no marked difference between the dry and

wet seasons at Port Elizabeth would cause var. obesa to be more

adaptable. It must be remarked, however, that the young var.

Sulgurata did not follow the example of their parent, but that of

var. obesa.

The conservatory in which.the molluses lived was heated, and

did not fall below 45° F. at night in winter. Nevertheless, they

seemed to feel variations in temperature, for there was greater

mortality in cold weather, and if they roused up earlier from

hibernation during a inild spell they generally went down again

26%

382 MRS. J. LONGSTAFF ON THE HABITS

when there was a return of cold. During hibernation they

invariably closed the aperture with an epiphragm, which had a

slit at the posterior end (PI. I. fig. 13).

Broods of Progeny.

June 21st, 1918.—As there was only one specimen of each

variety living I placed them both together, but they did not

appear to like one another at first, for they went as far apart as

possible. Later on, however, they became more friendly.

September 16th, 1919.—Several young ones and also some eggs

were-discovered below the surface, and the adults were separated

from them. On the 19th I counted eleven young molluses, twelve

eggs, and three broken bits of egg-shell. On the 26th I found

another and still smaller young one, with two bits of egg-shell.

It had, therefore, hatched from an egg, but there is no evidence

to show whether the whole of those first found had done so.

Var. fulgurata appeared to have deposited this brood.

July 23rd, 1920.—I scraped up the soil to a depth of about

two inches, and discovered fifteen living young molluscs and two

dead ones, accompanied by some fragments of egg-shell. ‘The

two dead shells were very small, had no coloured markings, and

were without any trace of the animals.

August 4th, 6th, and 12th.—Other living ones were found,

making nineteen in all, and the eggs numbered between fifty and

sixty. It is not clear which variety deposited the young ones.

After finding these I actually saw var. obesa lay some of the

smaller eggs; it is, however, quite possible that var. fulgurata

may have laid some of the larger ones.

Hibernation.

Winter, 1919-20.—October 6th to the middle of March.

Winter, 1920-21.—Both broods commenced to hibernate the

middle of October. During mild spells in January they came up

and ate. There were nine living of the first brood and seventeen

of the second, all grandchildren of the original specimens.

Summer, 1920.—-Var. fulgurata was not so active as usual,

appeared sickly in August, and finally died on September 22nd

at the age of six years and four months.

October 24th.—Var. obesa died after being inert several days,

when six and a half years old.

Reproduction.

Unfortunately, the cases here described do not afford clear

evidence of the length of time elapsing between fertilization and

birth. The two original specimens of var. obesa travelled together

with one of var. fulgurata for seventeen days. The former may

have fertilized each other, or they may have been fertilized pre-

viously by their fellows, or, on the other hand, they may have

been impregnated by var. Julgurata on the journey. With regard

OF COCHLITOMA ZEBRA IN CONFINEMENT. 383

to the var. fulgurata, five or six weeks had elapsed since it had

the opportunity of copulation with its own kind, but there is the

possibility of its having been crossed with var. obesa; fertilization

in transit was unlikely as the specimens were tightly packed.

' The second brood (September 1915) of the larger specimen of

var. obesa must have been produced either from the prolonged

effect of the previous copulation, or else the molluse must have

been fertilized afresh at least four months before, as that length

of time had elapsed since separation from its only fellow. As far

as can be judged from the appearance of the animals and shells,

cross-breeding does not seem to have taken place. It is, however,

a question whether the two broods of the progeny of the original

specimens may not be the result of cross-fertilization. The brood

of var. fulgurata (September 1919) was deposited about two years

and three months after the death of the last individual of its

own kind, aged three years, and after it had been living with

var. obesa about fifteen months. The actual parent of the young

ones deposited July 1920 is doubtful. . If it was var. obesa, and

fertilized by its own form, three years and four months had

elapsed since the death of its last companion, which was then

barely three years old. If fertilization was by var. fulgurata it

may have taken place any time after June 1918. If var. fulgurata

was the parent, either the effect of the first copulation must have

continued or else there must have been crossing with var. obesa.

The latter variety certainly laid some of the eggs, but it is not

evident whether any hatched.

These last two broods are as yet too young to ascertain whether

there are signs of cross-breeding. The older one, however, bears

most resemblance externally to var. julgurata. In connection

with this I would point out that Mv. Darbishire*, in an abstract

of Prof. Lang’s experiments with Helix hortensis and Helix

nemoralis, quotes an example of one of the snails being fertilized

in 1900, which, without further copulation, produced four families

in the years 1900, 1901, 1902, and 1903. Dr. Cooke t gives

instances of the prolonged effect of a single copulation in Helix

aspersa and Arion ater. In the case of the former the first batch

of eggs was laid after the lapse of only four days, and the second

the following summer. In the latter a month elapsed, and then

this pair of Arion ater both laid batches of eggs at intervals

during a period of seven weeks. He also quotes from a paper

communicated by Mr. J. 8. Gaskoin te the Zoological Society

in November 1850, to the effect that Helix dactea produced young

after being isolated four and a half years.

Eggs and Young.

It is of interest to note that these two varieties of Cochlitoma

have been observed to be at the same time both ovoviviparous

* Jour. Conch. vol. xi. no. 7, 1905, p. 194.

+ “ Molluses,” Cambridge Nat. Hist, 1895, p. 42.

384 MRS. J. LONGSTAFF ON THE HABITS

and oviparous, and also that var. obesa had one brood entirely

ovoviviparous. A dead specimen of the latter variety, found

associated with the living ones, contained a number of dead

young shells, with which no eggs were noticed.

Dr. Pilsbry * states that Cochlitome i is Ovoviviparous, and gives

an instance of Semper having found in the uterus of a specimen

of C. zebra sixty undeveloped eggs with calcareous shells, the

largest measuring 6 mm. in length and the smallest 3 mm.,

together with twenty-five embryos whieh had crawled out of the

egg-shell. Further, he remarks that Mr. Clapp found young

shells of about 8 mm. in diameter inside a specimen of C. craw-

fordi (Morelet), with which there were no traces of calcareous

ege-shell.

The dimensions of the eges of the different broods vary:

var. fulgurata has the largest eggs—these are preserved in such a

manner as to show the structure best. The eggs are calcareous,

yellowish in colour, and short-oval or rounded in form. The

biggest measure 9 mm. in length by 7-5 mm. in width; the

smallest 3°75 mm. in length by 3°25 mm. in width. Inside the

outer calcareous shell there is an amber-coloured layer which

does not cover the whole of the interior, and which appears to

give rise to the embryonic mollusean test. Fully a dozen of the

broken eggs of var. fulgurata contain the embryo and exhibit a

very thin grey layer, more or less calcareous, covering the aper-

ture and the greater part, if not the whole, of the shell. The

young shell consists of little more than three whorls, of which

the earliest is smooth, pale horn-colour, with the apex slightly

sunk; the succeeding whorls have spiral lines crossing the lines

of growth and rendering the surface granulated; the brown-

coloured markings begin on the last third of the body-whorl,

before it leaves the egg, and they form an interrupted spiral

band with a few longitudinal brown lines near the aperture. The

soft parts of the animal have entirely disappeared from all the

eggs. A young shell, released from a broken egg, had a length of

7-5 mm. and width of 8 mm.

The later brood of var. fulgurata (offspring of the original

specimen) containea fewer eggs but more living young molluscs.

I broke one of the twelve eggs, and found it was not so far

advariced as some of those of the first brood, as there was only

the amber-like layer inside the calcareous shell. The largest

measures 8°25 mm. in length by 6°5 mm. in width ; the smallest

6-5 mm. in length by 5°5 mm. in width.

One of the biggest eggs of the larger example of var. obesa

measured 8 mm. in length and 6 mm. in width; one of the

smallest, 3 mm. in length and 2:5 mm. in width. The biggest

egg of the smaller parent is less, measuring 5°5 mm. in length by

4 mm. in width.

The latest brood, which is probably mixed, contains eggs

* Man. Conch. vol. xvii. ser. L195 Ue

+ Ibid. p. 93.

OF COCHLITOMA ZEBRA IN CONFINEMENT. 385

varying from 7 mm. to 3°75 mm. in length, and from 5° mm.

to 2°75 mm. in width.

Growth.

The young molluscs of the different broods varied in size and

rate of growth; specimens of each of the earlier broods have

been figured at intervals to show the latter (Pls. 1. & II.). As far

as possible the same individual was represented, but when one

died another of the same brood was substituted. The rate of

growth was not always uniform, and sometimes a smaller specimen

surpassed a larger companion. The difference in size was shown

in a remarkable manner in the case of the latest brood of var. /ul-

gurata (September 1919), which consists of the grandchildren of

the specimen originally brought home. Three days after discovery

the largest measured 20 mm. in length and 15°5 mm. in width,

while a week later the smallest, which was probably hatched from

an egg, had a length of 10mm. and width of 9mm. On May 4th,

1920, about six weeks after they ceased hibernating the smallest

was the same size as before, while the largest had a length of

22 mm. and width of 16°5 mm. There was also another remark-

ably small one. These two small examples were not long-lived,

for the first-mentioned died on May 27th, when it measured

10 mm. in length and 9°5 mm. in width. The second died in

October, when it was 13 mm. in length by 10°5 mm. in width.

A living companion of average size at this time measured 42 mm.

in length and 29 mm. in width. This last, and the rest of the

brood, were then commencing to hibernate, so I did not wish to

disturb them to make further measurements.

The drawings represent the first broods up to the age of three

years and four months, when (September 1917) only one specimen

of each form was living. Neither of these had been figured

before, and they were then figured with the animals (Pl. IIT.

figs. 1 & 2). At that date this specimen of var. fulgurata

measured 71 mm. in length and 41 mm. in width; var. obesa

57 mm. in length and 39 mm. in width.

In September 1918 var. fulguratw measured 81 mm. in length

and 46 mm. in width; var. obesa 77 mm. in length and 49 mm.

in width.

In September 1919 var. fulgurata measured 88 mm. in length

and 49 mm. in width; var. obese 80 mm. in length and 52 mm.

in width. At death var. fulgwrata had attained a length of

89 mm. and width of 50 mm.; var. obesaa length of 80 mm. and

width of 52mm. It must be noted that both grew most between

September 1917 and September 1918, and least during the last

year of their lives. The increase in all cases was actually made

after hibernation, between the months of January and September

of each year. Var. fulgurata did not attain the length of its

parent, which was 107 mm. ; but var. obesa exceeded the size of

its parent (smaller specimen), which only measured 68 mm. in

length and 47 mm. in width.

386 MRS. J. LONGSTAFF ON THE HABITS

Resemblances and Differences.

In extreme youth the shells of the two varieties greatly

resemble one another, but the brown markings, so far as I have

observed, appear to commence earlier in var. fwlgwrate than in

var. obesa. ‘The embryos of the latter form, found inside a dead

specimen, do not show them at all, und I have only noticed the

faintest trace of them in one embryo. As the molluscs grow

older, however, they differ in character and colour of the shell,

as well as in the colour of the animal. The specimens of var.

obesw examined vary somewhat in the spiral angle of the shell,

but on the whole it is wider than that of var. fulgurata. The

shell of var. obesa is lighter in colour, being white or yellowish

white, while that of var. fulgurata is deep hore colour ; both are

ornamented by brown, longitudinal, more or less zigzag streaks,

which are less closely packed on var. fulgurata, and are also

broader on the body-whorl. The appearance of the latter is like

horn or tortoise-shell, while var. obesa is more porcellanous, and

is thus similar to the typical C. zebra.

The colouring of the animal of var. fulgurata is pale greenish

or yellowish grey, reticulated with a darker shade, and th ere are

three almost black stripes running down the head, with a banana-

yellow stripe on each side of the central one. In var. obesa the

animal is lead-grey reticulated with a darker shade; it also has

three dark stripes, which are not so nearly black as those of var.

Sulgurata, the central stripe is broader, and those on each side

are light grey. The fringe and sole are darkest in var. obesa.

The eggs of var. obesa are proportionately longer than those of

var. fulgurata, but on the whole they are smaller.

The habits of the two varieties also differ somewhat. Var.

fulgurate is more nocturnal, moving about and eating most at

night, while var. obesa moves about during the day as muchas at

night, and lives more continuously near the surface. The former

is more vigorous in its movements, making rapid and deep

excavations. It has also. been noted that the progeny of the

original var. fulgurata generally hibernated earlier, seeming to

require a longer period of rest. Contrary to its usual custom,

the solitary example of this variety lived on the surface during

the last summer of its existence, and did not appear to have the

energy to move much or to burrow.

Food.

The favourite food of both forms is lettuce, and next to that

dandelion and the leaves of the Life Plant (Bryophyllum calycinum

Bee They ate the skins of grapes that had been sucked, but

ould not manage the entire fruit; also banana and the outer

pee of cauliflower, when there was nothing else. They posi-

tively disliked cabbage and sprouts, either: fresh or cooked.

All had a habit of dragging leaves into their burrows, and some-

OF COCHLITOMA ZEBRA IN CONFINEMENT. 387

times the molluse would remain buried while merely the head was

protruded for feeding.

I am greatly indebted to Mr. G. C. Robson for several valuable

suggestions, also for affording me every facility when studying

specimens under his care.

EXPLANATION OF THE PLATES.

Prats I.

Cochlitoma zebra, var. obesa (Pfeiffer).

Fie. 1. Brood I. Drawn June 17th, 1914. 2. Born April 29th, 1914.

2. Brood II. Dvawn shortly after birth, May 4th, 1914. > 2. (Shell lost.)

2a. Brood I. Sculpture, greatly enlarged.

3. Brood II. Different specimen, drawn June 17th, 1914. x 2.

4,4a. Brood II. Eggs of medium and small size. X 2. Larger one showing

inner amber-like layer.

Cochlitoma zebra, var. fulgurata (Pfeitter).

5. Drawn May 4th, 1914, soon after birth. > 2. (Died after visit to the

Museum.)

5a. Sculpture greatly enlarged.

6. Drawn June 17th, 1914. X 2. Hatched from an egg, May 15th or 17th, 1914.

7. Kee, laid May 7th, 1914, showing inner amber-like layer. X 2.

8. Embryo, with egg-shell partly broken away, and showing inner thin cal-

careous layer, also the brown markings on the last whorl. X 2.

9. Specimen previously figured June 17th (tig. 6), drawn again July 28th,

1914. X 2.

10. The came drawn September 15th, 1914. X 2. (Died immediately after-

wards.)

11. Another shell, drawn April 9th, 1915. X 2.

12. Thesame, drawn June 30th, 1916. Nat. size. (Died June 1917.)

13. Portion of epiphragm, showing shit. X 2.

Prate II.

Cochlitoma zebra, var. obesa (Pfeifter).

Fig. 1. Brood I. Same as Pl. I. fig. 1, drawn July 28th, 1914. x 2.

2. 3 3 5 » september 15th, 1914. X 2.

3. ” “5 - » April 9th, 1915. x 2.

A, a a is » June 30th, 1916. Nat. size. (Died

September 1916.)

5. BroodII. Sameas Pl. I. fig. 3, drawn July 28th, 1914. x 2.

6. 55 ap . » september 15th, 1914. X 2.

Ue Ge 3 3 , April 9th, 1915. x 2.

8. 5 5 3 » dune 80th, 1916. Nat. size. (Died

February 1917.)

PratE III.

Cochlitoma zebra, vay. obesa (Pfeiffer).

Fig. 1. Brood I. Drawn September 5th, 1917. Nat. size. (Not figured previously.)

Born April 29th, 1914. Died September 1920. Dissected by Mr. G. C.

Robson.

Cochlitoma zebra, var. fulgurata (Pfeiffer).

2. Drawn September 5th, 1917. Nat. size. (Not figured previously.) Hatched

from an egg, between May 15th and 25th, 1914. Died September 1920.

Dissected by Mr. G. C. Robson.

All the specimens figured have been given to the British Museum (Nat. Hist.).

THE EXTERNAL CHARACTERS OF THE PROCYONIDA. 389

91. The External Characters and Classification of the

Procyonide. By R. I. Pocock, F.R.S.

[Received February 5, 1921: Read April 19, 1921.]

(Text-figures 1-13.)

ConTENTS.

Page

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The External Characters as a Guide to Classification ... 416

The Position of Ailwropoda .........c..ccccceee ce tenees 4LT

Classification of the Urside, Ailwropoda, Ailwrus, and

Eheabrocyonida varies eerie ries ees aeae nace Eh rr scrap oO

Introduction.

To show the wide divergence of opinion on the classification of

the genera assigned in the current text-books to the Procyonide,

it is needless to go to an earlier date than 1869, when Gray pub-

lished his ‘ Catalogue of Carnivorous ... Animals.’ Using in the

main the carnassial teeth, this author divided the Fere into two

suborders, Carnivora and Omnivora, the latter, with the teeth in

question non-sectorial, comprising the forms now relegated as a

rule to the families Urside and Procyonide. Gray, however,

admitted six families of Omnivora:—(1) Urside with several

genera; (2) Nasuide for Vasua; (3) Procyonide for Procyon

with two subgenera Procyon and Huprocyon; (4) Cercoleptide

for Cercoleptes (Potos); (5) Bassaride for Bassaris; (6) Ailuridee

for Ailurus.

In the same year, however, Flower (P. Z. 8. 1869, pp. 4-37)

classified the genera in question into the three families Procyo-

nide (Procyon, Bassaris, Naswa, Cercoleptes); Ailuride (Arlurus) ;

and Urside (Ursus) *.

In 1872, Gill, whose estimate of the value of characters was far

in advance of his time, made a classification which may be

described as a combination of Gray’s and Flower’s (Smiths. Mise.

Coll. xi. pp. 58-59, 66-67, 1872). It was as follows :—Fam.

Urside (with the genera admitted by Gray); Fam. Hluridee

(Hlurus); Fam. Cercoleptidee (Cercoleptes = Potos) ; Fam. Pro-

eyonide with the subfamilies Nasuine (Vaswa), Procyonine

(Procyon) ; Fam. Bassaridide (Bassaris).

* This grouping, however, was the same in substance as that proposed by Turner

about twenty years earlier (P. Z. S. 1848, p. 86) with the conversion of Turner's

subfamilies Ursina, Ailurina, and Procyonina of the family Urside into families of

the Arctoidea.

390 MR. R. I. POCOCK ON THE EXTERNAL

In 1876 (Proc. Acad. Nat. Sci. Philad. pp. 20-23), J. A. Allen

described the genus Bassaricyon and remarked: ‘“ As the species

[B. gabbr] differs more from either Vaswa or Procyon than the

latter do from each other, it seems to form a type quite as well

entitled to rank as a subfamily of the Procyonide as do either of

the others, and may hence be called Bassaricyonine.”

In 1883 (Hneycl. Brit. (9) xv. p. 441), Flower repeated his

classification of 1869 with the addition of Bassarieyon to the

Procyonidx and of Ailuwropus to the Urside.

In 1885, Mivart (P. Z.8. 1885, pp. 392-394) adopted Flower’s

views, with one or two important exceptions. He admitted only

two families; the Urside and Procyonide, fusing the Ailuridee

with the latter and relegating Ailuropus to a place alongside

Ailurus. His Procyonide, therefore, were grouped as follows :—

(1) Procyonine (Procyon, Nasua, Bassaris, Bassarieyon, Cerco-

leptes); (2) Ailurine (Ailurus, Ailuropus). It may be noted

that after comparing the dental and cranial characters of Ailurus

and Ailuropus, he summed up by saying “....so that on the

whole it appears to me that there is more decided natural affinity

between Arluropus and Ailurws than between Ailuropus and

Ursus.” .

Mivart was, I believe, the first author definitely to assign

Ailurus and Ailuropus to the Procyonide. Nevertheless, when

Flower and Lydekker published their volume on the Mammalia

in 1891, they quoted Blanford as the authority for the opinion

they adopted that Adlwrws belongs to that family. <Ailwropus

they retained in the Urside.

Similarly, in their paper upon “luropus, published in 1901

(Tr. Linn. Soe., Zool. viii. pp. 161-173), Lankester and Lydekker

do not appear to have consulted Mivart’s paper or to have been

acquainted with his views; and the result was that Lydekker

put forward a classification of the Procyonide identical in every

particular with that of Mivart.

Trouessart (Cat. Mamm. Suppl. pp. 183-184, 1904) referred

Aduropus and Ailwrus to the Urside, grouping them in the

subfamily Ailurine as opposed to the Ursin containing the

genera of true bears. The Procyonide he divided into two

subfamilies, the Potosine for Potos (olim Cercoleptes) and the

Procyonine for Bassaricyon, Bassariscus, Nasua, and Procyon.

With the substitution of Potosinz for Cercoleptine, this classi-

fication is the same as that published by Trouessart in 1898

(Cat. Mamm. i. p. 248). -

In 1914, Bardenfleth (Mindeskrift, ete., for J. Steenstrup’s

Wodsel, Copenhagen, no. xvii. pp. 1-15) reconsidered the question

of the affinities of Ailwrus and Ailuropoda, and, deciding that the

resemblances between them are purely adaptive, left the former

in the Procyonide and adopted Flower’s view that Ailuropoda is

‘an aberrant member of the Urside. The author tabulates in

three columns, devoted respectively to Ailuwrus, Ailuropoda, and

Ursus, no fewer than 58 characters by which these genera may

CHARACTERS OF THE PROCYONIDA. 391

be compared and contrasted, the only character quite in-

adequately dealt with being the feet, wherein he would have

found considerable corroborative evidence supporting his opinion

of the Ursine aftinities of Ailuropoda.

The latest classification was published in 1916 by Hollister

(Proce. U.S. Nat. Mus. 49, pp. 141-150), who, following Gray and

Gill, separated Bassariscus from the rest of the genera, his

grouping being as follows:—Fam. 1. Bassariscide (Bassariscus) ;

Fam. 2. Procyonide (Ailurus, Procyon, Nasua, Nasuella * Bas-

saricyon, Potos). This grouping involves the conclusion that

the affinity between Ailuwrus and Procyon or Potos is greater

than the affinity between Procyon or Potos and Bassariscus, a

conclusion which, in my opinion, is quite indefensible. With

regard also to the affinities of Potos and Bassariscus with the

other genera, I cannot agree with Hollister, whose opinion with

regard to the dentition of Bassariscus may be contrasted with

that of Flower when he wrote in 1869 “‘ Cercoleptes [Potos|

deviates in its dentition from the more typical members of the

group far more than Bassaris [Bassariscus], though in a precisely

opposite direction.”

From the above-given review it will be evident that there is

no sort of unanimity on the three following points:—(1) The

position of Aiduropodat. Some authors claim that the genus

belongs to the Urside; others place it in the Procyonide.

(2) The position of Adlurust, some authors regarding it as the

type of a special family, others, in a majority, placing it in the

Procyonide. (3) The constitution of the so-called Procyonide

of America. Hyery genus has at various times been made the

representative of a special family or subfamily. Justification for

this course is to be found in the much greater differences that

exist between them than between the genera of Felide, Canide,

and other compact families of Carnivora.

This variety of opinions calls for a revision of the questions at

issue in the light of other characters than those supplied by the

teeth, skull, and skeleton, which have mainly been used. Gray,

it is true, employed the feet to a certain extent, but he was

compelled to depend very largely upon dried skins, and most

authors attach comparatively slight importance to external

characters.

During the past few years I have been making sketches of the

feet, ears, rhinaria, and other external organs of the Carnivora

that have died:in the Gardens of the Zoological Society : and the

* To illustrate the differences between Nasua and Nasuella and justify the

creation of the latter, Hollister seems to have selected skulls exhibiting extremes or

variation. I have skulls of Naswa almost intermediate between the two figures.

+ This was the name originally given to the genus by Milne Edwards; but he

subsequently changed it to Ailwropus because Gray had previously employed the

name Aluropoda for a section of the Carnivora comprising the Felidae, Mustelide,

and other families. Gray’s action, however, did not invalidate the use of Ailwropoda

in a generic sense. I have, therefore, retained it.

+ The original spelling of this name is also adhered to in this paper,

392 MR. R. I. POCOCK ON THE EXTERNAL

present paper is based mainly upon the Procyonide examined in

the Prosectorium,

The following material has passed through my hands :—

Ailurus fulgens.—Several specimens of both sexes.

Procyon lotor or closely-allied forms.—Several specimens of

both sexes.

Nasua nasua (ring-tailed Coati Mondi, with a dark snout and

distinctly ringed tail).—Several specimens.

Nasua narica (the white-nosed Coati Mondi, with a white

snout and indistinctly ringed tail).—Several specimens.

Bassaricyon sp.—A. single specimen, sent on approval by

Mr. Cross, the animal dealer, which died on the day of its

arrival and had to be returned to the vendor. Of this

only the feet and rhinarium were sketched. My observa-

tions on this genus have been supplemented by an

examination of the dried skins in the British Museum.

The example of this genus dissected by Mr. Beddard

(Proc. Zool. Soc. 1900, pp. 661-675) was converted into a

skeleton.

Potos caudivolvulus or closely-allied forms.—Several specimens

of both sexes.

Bassariscus astutus.—A single immature male preserved in

aleohol — probably the specimen whose anatomy was

described by Mr. Beddard (Proc. Zool. Soc. 1898,

pp. 129-131).

Jentinkia * sumichrasti.—Dried skins in the Natural History

Museum. This Cacomistle is sometimes admitted as a

subgenus of Bassariscus. I have provisionally quoted it

asa distinct genus, the material available being insufficient

to establish the absolute constancy of the differences in

the feet observable between astutws and swnichrasti.

Ailuropoda melanoleuca.—Mounted specimen and skull in the

Natural History Museum.

The skulls examined are in the collection of the Zoological

Society and in the Natural History Museum.

The External Har.

As recorded by previous authors, the pinna of the ear varies

in shape and size, being short and rounded in Nasua, Potos,

and Bassaricyon, longer, more pointed, and more expanded in

Procyon, Bassariscus, Jentinkia, and Ailurus. The cartilages

strengthening the base of the ear are simple and normally

carnivorous in their arrangement, requiring no special deserip-

tion, especially as they vary in development to a certain extent

* Proposed by Trouessart (Cat. Mamm. Suppl. p. 184, 1904) to replace the pre-

occupied name Wagneria Jentink (Notes Leyd. Mus. vii. p. 129, 1886) based on the

differences hetween B. variabilis [sumichrasti] and astutus pointed out by Peters

(Mon. Ak. Wiss. Berlin, 1874, pp. 704-705).

CHARACTERS OF THE PROCYONID. 393

within generic limits. Much greater interest attaches to the bursa.

In Bassariscus and Jentinkia this is developed as in the Canide,

Felide, and some Viverride, its posterior half being formed by

Text-figure 1.

A. Head of Potos caudivolvulus (vel sp. aff.). Sad. Xd.

B. 4, Bassariscus astutus. g immat. X34. (From specimen

in spirit with ear half closed and hair wet.)

C. ,, Procyon lotor, gad. XH.

D. Lateral view of rhinarium of Bassaricyon sp. ?

E. Ear of Bassaricyon alleni. (Sketched without measurement from

dried skin.)

394 MR. R. I, POCOCK ON THE EXTERNAL

a semicireular flap of integument rising behind the adjacent free

edge of the pinna, its anterior rim being deeply emarginate.

Judging from dried skins, the bursa of Jentinkia 1s similar to

that of Bassariscus. J also find it present in dried skins of

Bassaricyon, but it appears to be less well developed than in

Bassariscus. The upper portion of the posterior flap arises

behind the margin of the pinna, but its lower portion is confluent

with it, and its anterior rim is lower and apparently not emar-

ginate. The only other genus in which it is present is Vasua,

where it has the form of a shallow pouch near the edge of the

pinna which bulges slightly at this point. A similar bulge on

the ear may be seen in Procyon, but in no example examined

was there a trace of the pouch, and there is no trace of it in

Potos or Ailurus. There is also no trace of it visible in the

mounted example of Ailuropoda in the Natural History Museum,

and it is absent in all the genera of Urside,

Tf, as I believe, a well-developed bursa, such as is seen in the

Felide and Canide, is a primitive feature in the Carnivora—and

its presence in some of the genera of all the recognised families,

except the Urside, suggests that conclusion,—it follows that the

ear of Bassariscus is the most primitive type in the Procyonide.

Next comes that of Bassaricyon, then of Vasua showing stages

in its suppression which has been achieved in Procyon, Potos,

and Adurus.

The following table summarises the characters of the ears in

the genera examined :—

a. Bursa retained.

b. Bursa marginal, at least the upper end of the posterior flap

attached behind the adjoining edge of the pinna.

c. Lower end of posterior flap behind the edge of the pinna;

anterior flap emarginate; ears large ................ Bassariscus, Jentinkia.

c’. Lower end.of posterior flap confluent with edge of pinna;

anterior flap with low straight edge; ears small ......... Bassaricyon.

b’. Bursa a shallow pouch in front of posterior edge of pinna.. Nasua.

a’, Bursa suppressed.

d. Kars large, expanded, and more or less pointed ............... Procyon, Ailurus,

diseWarsismalidandereunde ditra-eeceene teste teeerere eeeeeee me AOLOSS

By this table Ailuropoda would fall alongside Pofos under the

heading d’. In the Urside also no trace of the bursa remains

and the ears are always rounded. They may, however, be com-

paratively large and rounded as in. Selenarctos * tibetanus, or

small as in Helarctos malayanus.

* Mr, A. de Carl Sowerby has pointed out to me that Heude applied this generic

name to this Oriental bear in 1901 (Mém. Hist. Nat. Chinois, v. pt. 1, p. 2). The

later name Arcticonus, which I gave to it in 1917 (Ann. Mag. Nat. Hist. (8) xx.

p. 129), lapses, therefore, as.a. synonym of Selenarctos. My excuse for missing this

point is that the then recorder of the Mammalia in the ‘Zoological Record ’ pur-

posely refrained trom quoting Heude’s work on account of its general Wonnalcs=ness

in his opinion, :

CHARACTERS OF THE PROCYONIDA. 395

Facial Vibrisse.

Mivart described the facial vibrisse of Procyon as follows :—

“The whiskers on each side have 5 or 6 bristles grouped together,

and there are four sets of such groups. There is also a tuft over

the eye, one behind the angle of the jaw, and one under the

Text-figure 2.

ayo? ))

Sie

A. Head of Nasua nasua. gad. Xi.

BB. ,, Ailurus fulgens. Gad. Xi.

middle of the chin.” That is quite true; but, in addition, the

superior genal tuft is also present, although sometimes it is hard

to detect when the normal tuft is reduced to a single vibrissa on

Proc, Zoot, Soc.—1921, No. XX VII, 27

396 MR. BR. I. POCOCK ON THE EXTERNAL

each side. Submental vibrisse are also present. In this genus,

therefore, the tufts of facial vibrissee are normal for the Carni-

vora, consisting of mystacials, superciliaries, two tufts of genals,

the lower behind the angle of the mouth, the upper below and

behind the angle of the eye, submentals, and an unpaired

interramal tuft. ;

I have no notes or sketches of the facial vibrissee of Bassart-

cyon, but from Beddard’s. remark that there are five or six long

vibrissee forming the whiskers, three or four upon each cheek and

two or three under the chin, I think it may he inferred that this

genus falls into line with Procyon and Potos in having these

tactile hairs normally developed.

In Ailurus the tufts of vibrisse are also normal in number and

distribution, and the upper genal tuft, represented by one or two

vibrisse, is situated low down on the cheek, only a little above

the lower, and is often difficult to detect amongst the normal

hairs which thickly clothe this portion of the head.

Similar tufts are present in the genera NVasua, Bassariscus,

and Potos, being better developed and consisting of stouter

vibrisse in Vasua than in the others. In two examples of Potos

the upper and lower genals are represented by a single vibrissa

each. In Bassariscus the upper genal tuft is set high up rather

close to the angle of the eye, as is usual in predatory Mustelide.

In Potos and Nasua it is lower down the cheek.

In Ailuropoda the mystacial vibrisse are all short and ob-

viously of little import as in the Urside. On the single stuffed

specimen examined I was unable to satisfy myself as to the

presence of the other tufts.

The Muzzle and Rhinarium.

In Procyon the snout is moderately lengthened, and projects

considerably beyond the lower jaw. The rhinariwm is large and

naked above. In front it has no central groove or, at most, a

short indistinct one below. The upper margin is flat or evenly

convex, not biconvex, and the width is much greater than the

depth, but less than the combined depth of the rhinarium and of

the upper lip. There is no trace of a philtrum, the upper lip

being continuously, although comparatively scantily, hairy across

the middle line, and the inferior edge of the rhinarium transverse

in the middle and upeurled laterally. The internarial septum is

broad and the infranarial portions moderately deep.

In Bassariscus astutus the muzzle is almost as elongated as in

Procyon, the width of the rhinarium being much less than the

combined heights of the rhinarium and of the upper lip; but

the lip is cleft by a long narrow philtrum. The rhinarium itself

is very like that of some Felide and Viverride, its upper edge

being horizontal, with rounded angles from the anterior aspect

and its inferior edge strongly angled mesially, where it passes into

the philtrum, ‘The internarial septum is moderately wide and

CHARACTERS OF THE PROCYONID®. 397

the infranarial portion comparatively shallow. There is a faint

median groove marking the rhinarium from its summit to the

bottom of the philtrum.

In Vasua the snout is much more elongated and mobile than in

Procyon, and, as in that genus, the upper lip is entire, the rhina-

rium showing no trace of philtrum. The rhinarium differs from

that of the other genera in certain particulars. Its upper anterior

Text-figure 3.

A. Rhinarium of Ailurus fulgens ; front view.

B. 5 Potos caudivolvulus; front view.

Cc: Py Bassaricyon sp.?; front view.

DE s Bassariscus astutus ; tront view.

EK. . Nasua nasua; front view.

19% of Procyon lotor; front view.

All figures X 4 approx.

(D, E, F. Drawn with upper lip in same plane as anterior surface of rhinarium.)

margin is strongly produced. Its greatest width in front across

the infranarial portions is less than the combined heights of the

rhinarium and of the upper lip. The inferior edge is commonly

strongly sinuous, owing to the median emargination. The nostrils

27%

398 MR. R. I. POCOCK ON THE EXTERNAL

instead of being subcircular are markedly elongated from above

downwards, and the internarial septum is long and narrow. The

infranarial portions are deep, and are defined by a groove from

the adjoining central portion, which has no trace of median

groove.

To Beddard’s description of the rhinarium of Bassaricyon as

naked and marked with a median groove which also cleaves the

upper lip and is visible on the dorsal side, | may add that in

the specimen examined by me, the upper edge, when seen from

the front, was lightly biconvex, the median groove was well

marked and complete, the philtrum was short but barely as wide

as the internarial septum, and that the infranarial portion on

each side was exceedingly deep and marked with a faint trans-

verse groove, meeting its fellow of the opposite side in the middle

line; and that the total width of the rhinarium exceeded the

height from the edge of the upper lip to the summit, the snout

being short as in Potos, and not produced as in Bassariscus and

Procyon.

In Potos the snout is not produced or slender, and the rhina-

rium is remarkable for the great width of the philtrum, which is

as wide as or wider than the internarial septum, its sides often

diverging below, and the greatest width of the rhinarium exceeds

its height from the summit to the edge of the upper lip. The

upper edge, seen from the front, is convex or lightly biconvex,

the internarial septum is wide, the infranarial portions are very

deep, gradually narrowing beneath the slit-like portions of the

nostrils laterally. There isa well-marked median groove running

from the edge of the lip and sometimes extending on to the

upper surface in front, and a faint subhorizontal groove runs

inwards beneath the nostril to meet its fellow of the opposite

side in the middle line.

In Ailurus the snout is not produced, and the upper lip is

divided by a short narrow philtrum, much narrower than the

internarial septum, and the greatest width of the rhinarium

considerably exceeds its height from the summit to the edge of

the upper lip. The upper edge is tolerably evenly convex or

markedly biconvex, the internarial septum is wide, and the

infranarial portions are comparatively shallow, each being marked

by a faint groove which extends obliquely downwards and in-

wards to meet its fellow of the opposite side in the middle line.

The median groove is well marked, and extends from the edge of

the upper lip often on to the summit of the rhinarium.

So far as it is possible to judge from the mounted specimen of

Ailuropoda, the rhinarium resembles that of <Adlwrus in all

respects, especially in the shortness and narrowness of the

philtrum, except that its upper surface is entirely covered with

hair. In this respect it differs from that of all the genera of

so-called Procyonide and Urside.

I was unable to satisfy myself as to the protrusibility of the

lips in Ailuropoda, although the lower lip, as mounted, is slightly

CHARACTERS OF THE PROCYONIDA. 399

compressed and pouted in the middle line. The point is not

without importance, because all the Urside differ from all the

Procyonide and from Ailwus in having the lips protrusible or

capable of being pouted. In the Procyonide and Ailwrus the

hips above and below the incisor teeth are comparatively closely

adherent to the gums as in normal Carnivora.

By the characters supplied by the snout and rhinarium the

genera may be classified as follows :—

a. Snout greatly elongated; upper edge of rhinarium produced be-

yond the nostrils so that its anterior surface is nearly flat and

slopes obliquely downwards and backwards ; anterior portion of

nostrils vertically elongated, separated by a high, narrow, un-

grooved septum, and almost concealed in profile view. Upper

lip longandsumdivided by, plultmumi ssc ee ee cee eee.

a’, Snout not, or only moderately elongated; upper edge of rhi-

narium not produced, its anterior surface lightly convex, not

noticeably receding ; anterior portion of nostrils subcircular,

separated by a low, broad, generally grooved septum, and visible

in profile view.

6. Snout moderately produced; upper lip long so that the width

of the rhinarium is less than the combined heights of the

rhinarium and upper lip; median groove on rhinarium weak,

or indistinct.

e. Upper lip undivided by philtrum ; infranarial portion of rhi-

narium deep and wide, median groove abbreviated or

Indisbincbisgeepee e eerece Nos) oso ee. csc Procyon.

cl. Upper lip divided by long, narrow, groove-like philtrum;

infranarial portion of rhinarium shallow and narrow, median

eroovelcompleten Seneca eeeer eee eee ess) Bassariscuss

b!, Snout not produced; upper lip short, width of rhinarium ex-

ceeding its height from the summit in front to edge of upper

lip; median groove strong.

d, Infranarial portion of rhinarium very deep.

Nasua.

é> Ehiltmumimioderately broad |.1c-c. veces eeee eee ee cate ee: Bassaricyon.

é:. Philtraty yerysbrordieeany cee «noch. cika cee lt eae sac Potos:

@’. Infranarial portion of rhinarium comparatively shallow and

philteem Marto ws eked eee cok ew eee Atrus.

By this key Azluropoda, so far as it is possible to judge, would

fall under heading d' alongside Ailurus, from which it differs in

having the upper surface of the rhinarium covered with hair.

The rhinarium is variable in the Urside, being comparatively

highly specialised in Melwrsuws (Ann. Mag. Nat. Hist. (9) i,

pp. 378-379, 1918),

The Feet.

In describing the feet of the American genera, those of Vasua

may be taken as a standard for comparison.

The claws of the fore foot are long, powerful, blunt and not

greatly curved, and the digits are united by web, as described by

Mivart, up to the proximal end of the digital pads. The under-

side of the digits and of the webs is entirely naked. The plantar

pad is broad, moderately well defined and four-lobed. The two

conjoined carpal pads, of which the outer is about twice as

large as the inner, are together as wide as the plantar pad and

400 MR. R. I. POCOCK ON THE EXTERNAL

approximately equal to it in area. Th ey are respectively in contact

with the inner and outer lobes of the plantar pad; but in the

middle line they are separated from it by a depressed area of

smooth wrinkled integument. Above the carpal pad on the

Text-figure 4.

. Right fore foot of Nasua nasua.

» hind foot of same.

. Claw of fore foot.

nh hind foot.

Right fore foot of Procyon lotor.

» hind foot of same.

X 2;

HOO p

CHARACTERS OF THE PROCYONID#, 401

ulnar side there is a tuft of carpal vibrisse. The claws and

digits of the hind foot resemble those of the fore foot, except

that the claws are somewhat shorter. The plantar pad is four-

lobed ; and above it the whole of the tarso-metatarsal area is

Text-figure 5.

A. Right fore foot of Potos.

B. ,, hind foot of same.

C. Claws and toes of fore foot of same,

D. Right fore foot of Bassaricyon sp.; digits not spread,

E. ,, hind foot of same. ,

F. Claw of fore foot of same.

wie

naked as far back as the tip of the caleaneum, although the

proximal third of the naked area is considerably narrower than

the distal two-thirds. There are no definite metatarsal pads.

The feet of Procyon are a little less robust than those of

402 MR. R, I. POCOCK ON THE EXTERNAL

Nasua, the pads are less well defined and the claws rather smaller;

but the great and essential difference lies in the complete

absence of interdigital webbing, the digits being entirely free as

far back as the plantar pads. This very rare phenomenon in the

Carnivora is paralleled only, so far as I know, in the Crab-eating

Mongoose Atilax, and in both cases it is probably subservient to

delicacy of touch in finding and handling food.

Text-figure 6.

A. Right fore foot of Jentinkia sumichrasti.

B. ,, hind foot of same.

C. 4, fore foot of Bassariscus astutus ; immat.

D. ,, hind foot of same.

(A, B. Sketched from dried skin.)

role

The feet of Potos* differ from those of Vasua in that the

claws are shorter, sharper, and more curved; approximately the

distal two-thirds of the digits are free from webbing, and, in

the hind foot, the lower side of the caleaneum is covered with

hair, the hair-covered area corresponding to the narrow naked

area in Vasua.

* The feet of Potos weve figured by Kidd to illustrate the development of the

sensory ridges on the pads. The feet from which these figures were taken closely

resemble those examined by me, but the metatarsal pads seem to have been better

eee (‘The Sense of Touch in Mammals and Birds,’ pp. 24-25, figs. 9 & 10,

1907).

CHARACTERS OF THE PROCYONIDA. 403

In Bassaricyon * the feet closely resemble those of Potos, the

ealeaneum being hairy beneath; but the interdigital webs are

nearly as well developed as in Vasua. The claws are compara-

tively short and strongly curved and unlike those of Vasua.

In Jentinkia sumichrastit the feet are seemingly very like

those of Bassaricyon in the extent of the interdigital webbing

Text-figure 7.

A. Right fore foot of Ailurus fulgens.

B. The same with hair cut short and digits spread.

C. Right hind foot of same.

D. The same with hair cut short and digits spread.

5 Bio

and the length and curvature of the claws, but possibly they are

somewhat narrower. The great difference lies in the lower

* Beddard’s figure of the fore foot of this genus showing the presence of the

carpal vibrissze agrees closely with my figure of this foot in the example I examined

and sketched (see Proc. Zool. Soc. 1900, p. 663).

I have unfortunately oniy seen of this species dried skins with the feet

shrivelled or stretched. ‘he main features of the pads were discernible, but I could

not assure myself of the constancy in shape and size of the naked metatarsal area.

404 MR. R. I. POCOCK ON THE EXTERNAL

surface of the metatarsus, which is for the most part covered with

hair, except for a relatively short, naked, proximally narrowed

area just above the plantar pad. In this respect the feet of

Jentinkia differ from those of the above-mentioned genera.

In Bassariscus astutus the feet are short, compact, and some-

what like those of Genetia, with short, curved, sharp claws and

interdigital webs extending nearly up to the digital pads. They

differ from those of all the genera previously described in having

the lower side of the digits and of the webs covered with short

hair almost as far back as the plantar pads, which are thick and

strongly lobed. Also in the fore foot there is a single submedian

carpal pad, which is elongated and narrowed proximally and much

narrower at its distal end than the plantar pad, with which it is

almost in contact. Furthermore, in the hind foot, the meta-

tarsal area is covered with hair right down to the plantar pad.

It is mainly on the strength of the differences observed between

the feet of Bassariscus astutus and of Jentinkia sumichrasti, which

by most authors is referred to the genus Bassariscus, that I have

preserved the genus Jentinkia. It is necessary to repeat, how-

ever, that I have only seen dried skins of J. swmichrasti and only

one spirit-preserved example of B. astutus. Feet of dried skins

of the latter seem, however, to agree with those of that example ;

but admittedly more material is required.

The feet of Adlurus differ from those of the American genera

in the complete concealment of all the pads beneath the clothing

ef long and thick hair which everywhere covers the soles and in

the consequent reduction of the pads to apparently functionless

vestiges. Separation of the hair reveals vestiges of the pads as

areas of naked, somewhat thickened skin, those on the digits

being small subtriangular patches behind the base of the claws.

The carpal pad is represented by a small subcircular patch remote

from the plantar pad; but there is no trace of metatarsal pads.

The plantar pads are reduced to transverse recurved areas,

expanded at each end towards the base of the first and fifth

digits and sending forwards two angular processes, longer on the

hind than on the fore foot, approximately in line with the third

and fourth digits. A further difference between these pads is

that the inner (preaxial) arm is longer than the outer (postaxial)

on the fore foot, the converse obtaining on the hind foot. The

digits are united by webbing to approximately the same extent as

in Bassariscws, and the claws are short, curved, and very sharp *.

* This description of the feet agrees in a general way with that published by

Flower (P. Z. S. 1869, p. 754). He does not, however, mention the carpal pad, and

describes the plantar pad as a ‘larger, transversely oval, bare space 4” [less than

half an inch] across covered by pink soft skin.” I assume that he did not trace the

details of these pads to their full extent. Mivart would never have described the

claws of the Panda as “blunt” if he had ever been scratched by them. The hairs

on the soles of the foot of the Panda are a great disadvantage in climbing smooth

branches. It is only with considerable difficulty that the animal can slowly ascend a

stout branch, stripped of its bark, even when inclined at an angle of 45 degrees. It does

so by means of the penetrating power of the sharp claws. It is not easy to suggest

a meaning for the hairiness of the feet, since the animal does not live in a region of

perennial snow. Possibly, however, it is to obviate slipping on wet branches.

CHARACTERS OF THE PROCYONID&. 405

-Text-figure 8,

Way tf

Wid

HEV uh

1 iyi

SSS

SSsss

. Right fore foot of Ailuropoda melanoleuca.

- 5, hind foot of same. (Qopied from Milne Edwards’s figure.)

» fore foot of Euarctos americanus.

+ 5, hind foot of same,

406 MR. R. I. POCOCK ON THE EXTERNAL

From the above-given description it is clear that the feet of

Ailurus differ very matevially from those of all the American

genera, to which it is supposed by some authors to be tolerably

closely allied.

The feet of Ailuropoda differ from those of Atlurus and of the

American genera in being essentially Ursine in three par-

ticulars :—(1) The fore and hind feet are approximately equal in

length, owing to the shortening of the hind; (2) the five naked

digital pads form a slightly curved transverse line, the second

and fifth lying respectively alongside the third and fourth, and

the first (pollical and hallucal) touching the second when the

digits are in contact; (3) the plantar pad is a wide naked trans-

verse cushion, at least twice as wide as long and separated from

the digital pads by a short area overgrown with hairs.

The fore foot further resembles that of Ursus and Huarctos in

possessing a large carpal pad separated from the plantar pad by a

long area overgrown with hair; but it differs from the fore foot

of all the Wisides in the mariod preaxial projection of the

plantar pad beyond the line of the pollex. This gives additional

width to the foot, and is doubtless correlated with the great

development of the radio-carpal bone, described by Lankester,

which simulates an additional metacarpal. The hind foot differs

from that of the typical Urside in that the entire sole is covered

thickly with hair from the caleaneum to the plantar pad. It

must be remembered, however, in this connection that in

Thalarcios the corresponding portion of the foot is hairy, the

metatarsal pad being reduced to a comparatively small lozenge-

shaped area. The skeleton of the digits differs from that of. the

Ursidx, as Lydekker pointed out, in the presence of a bony hood

at the base of the terminal phalange. In the latter particular,

as in the remoteness of the carpal pad from the plantar pad, the

feet resemble those of Azlwrus, but they differ therefrom in the

large size and exposure of the pads and in the alignment of the

extremities of the digits.

Excluding those of Ailuropoda, the characters of the feet may

be summarized as follows :—

a. Soles of feet entirely and thickly covered with woolly hair con-

cealing the pads, which are reduced in extent and thickness to

apparently functionless areas of naked skin; carpal ped a minute

naked area remote from the plantar pad ............... . Ailurus.

a’. At least the digital, plantar, and carpal pads need ands Ww Fall dogs.

loped ; carpal “pad, or pads, comparatively large and in contact

with the plantar pad, or nearly so.

6. Area between digital and plantar pads hairy ; carpal pad single,

much narrower than plantar pad; metatarsus covered with

hair down to plantar pad.......... . Bassariscus.

b’. Area between digital and cone ails melede car omar waadls oil,

conjointly as w ride as plantar pad ; at least a small naked area

on metatarsus above plantar pad.

c. Naked area of metatarsus reduced to a Oe el small

patch, narrowing above .......... . dentinkia.

ce’. Naked area of metatarsus sxisnaling ¢ over ine renter “a oF

that area,

CHARACTERS OF THE PROCYONIDA. 407

d. Digits entirely free from webbing down to plantar pad... Procyon.

d’, Digits united more or Jess by webbing beyond plantar

pad.

e. Claws fossorial, long, strong, and blunt; digits fully

webbed up to digital pads; caleaneum naked mesially.. Naswa.

e’. Claws shorter, curved, and sharp; digits less fully webbed ;

- calcaneum coyered with hair.

f. Digits united by webbing for about two-thirds of their

Teng thine wi wamcaues em ean bese io pteali ke Bassaricyon,

f’. Digits united by webbing for about one-third of their

lengthy beyondtplantar padi ie see ets. oes) Rotos:

Text-figure 9.

Se eee Se ak

Ventral view of Potos, showing the median cutaneous glands, the position

of the prepuce, etc.

Cutaneous and Anal Glands.

The only genus, known to me, which has specialized cutaneous

glands, apart from those connected with the anus, is Potos, In

408 MR. R. I. POCOCK ON THE EXTERNAL

this genus there are two in the middle ventral line, one anterior

and the other posterior. The former consists of a comparatively

small patch of skin, scantily covered with short hairs and

situated just in front of the anterior end of the sternum; and

the latter, which is structurally similar to it, of an elongated

tract on the abdomen, extending in the male-from a point just in

Text-figure 10.

ANIA i if) Wi big

A “oy

aN ah q\)

TAY

wes

A. Anus and vulva of Ailurus fulgens.

B. = scrotum of same.

C. os vulva of Procyon lotor.

D. mi scrotum of same.

(In A and D the orifices of the anal glands are shown; in D the size and

position of the gland of the left side is dotted in.)

front of the prepuce, where it is narrowed, over the umbilicus to

the posterior end of the sternum. They are similarly placed and

equally well developed in the female. These areas are not

CHARACTERS OF THE PROCYONIDA. 409

sharply circumscribed at the margins, and the glandular portion

is composed of well-developed normal skin-glands *,

Potos also has a considerable amount of nearly naked skin on

the lower jaw and round the mouth. This, however, does not

appear to be especially glandular. Probably the absence of the

hair serves to keep the lips and chin clean from such sticky

substances as honey, on which the Kinkajou feeds.

In Procyon, Nasua, and Potos the anus, like that of the

Urside, Canide, and others, opens in the centre of a circular,

slightly protuberant area of naked skin, separated by hair from

the scrotum or vulva. As Mivart stated, there is a pair of normal

anal glands in Procyon like those of the Canide, but relatively

a little smaller. Mivart also states that these glands are present

in Potos; but [ entirely failed to find even a trace of them in

two examples, a male and female, the result of my examination

confirming Owen's statement that they are absent (Proc. Zool. Soe.

1835, p. 122). Beddard does not appear to have looked for them

in Bassaricyon. Their presence in Bassariscus was recorded by

Gervais in the following passage (Voy. de la ‘ Bonite,’ i. p. 19) :—

“Une petite plaque crypteuse circumanale existe 4 la terminaison

de Vintestin, comme dans les Mangoustes, mais elle est moins

grande que dans ces animaux et asa surface débouchent de méme

les deux conduits des glandes anales.” Although this passage

suggests the presence in Lassariscus of a circumanal pouch like

that of the Mongooses, I am disposed to think Gervais was merely

describing the button-like area of naked skin immediately sur-

rounding the anal orifice, such as is seen in Procyon or Nasua.

This supposition is borne out by what he says as to the position

of the orifices of the anal glands; and is further supported by the

absence of a circumanal pouch in the male example of Bassariscus

LT examined.

In Naswa anal glands are present, but in a much modified

form, as Mivart pointed out. When the anus is opened, they

appear as a series of four or five parallel slits, each series being

set just within the orifice. The slits are the apertures of ag

many narrow sacs formed by the folding of the anal integument.

These glands are quite different from those of any Carnivore

known to me, and serve to differentiate Vasuwa tolerably sharply

from its allies.

Flower described the anus of the male of Ailurus, pointing out

the presence of a pair of normal anal glands and of a glandular

area round the anal orifice. In the male I examined, the

integument round the orifice corresponding to that seen in

Procyon, for example, is highly glandular, and partially insunk so

as to suggest the anal pouch of the Mungotide. Round it there

is a considerable area of naked skin extending inferiorly to the

* These glands are conveniently placed for rubbing the secretion along the

branches of trees to enable Kinkajous to track each other by scent. Although I

have never noticed these animals behaving in a way to suggest that that is the

function of the glands, I do not doubt that it is so, ;

410 MR. R. I. POCOCK ON THE EXTERNAL

scrotum, and from it the glandular central area is somewhat

sharply marked off.

Text-figure 11.

A. Anus and scrotum of Nasua nasua.

B. Anus of Naswa opened to show the glands; the upper margin of the

anus below.

C. One of the anal glands of Nasua in vertical section, showing the

series of pouches opening upwards and the underlying muscle.

Dp. Anus and vulva of Potos.

The adult female possesses a similar glandular depression round

the anus, but the surrounding area of naked skin is not so wide

above and laterally, although extending inferiorly to the vulva *.

Tn an immature female, six months old, the anus is like that of

Procyon, showing no special development of glands.

The four genera I have examined may be tabulated as follows

by their specialized scent-glands :—

a. Anus opening in the centre of a shallow glandular pouch like

that of the Mungotide ; normal anal glands retained ......... Ailurus.

a’. Anus as in ordinary Carnivora .

6. Anal glands suppressed; two ventral cutaneous glandular

ATOAS Aye sa ON a UPR Re EERO OLN Ee

b’. Anal glands retained; no ventral cutaneous glands.

c. A pair of anal glands of the type normal in the Carnivora.. Procyon and

Bassariscus.

Potos.

c', Anal glands peculiar, consisting of a series on each side of

five shallows litle jp OWeHeS| ieeer cee eaceitce eae nayh i receeayer Nasua.

* Adults of both sexes of the Panda have the habit, so well known in the

Mongooses, of rubbing the secretion of the anal region on the branches or other

parts of their cage.

+ On the assumption that my interpretation of Gervais’ description of the anus of

Bassariscus is correct,

CHARACTERS OF THE PROCYONIDA. 411

Nothing is known of Ailuropoda with respect to specialized

cutaneous or anal glands. The latter are probably present and

the former absent. No specialized cutaneous glands have been

discovered in the Urside; but the anal glands are present or

absent. Owen declared them to be absent, and I failed to find a

trace of them in Ursus arctos, the type he probably examined.

In an example of Huarctos americanus, however, I detected the

normal pair, though relatively much reduced in size. The anus

in this family resembles that of the typical Procyonide.

Heternal Genitalia,

In Procyon, Nasua, Potos, and Bassariscus, and presumably in

Bassaricyon, the prepuce is abdominal and far in front of the

scrotum, as in the Canide, Urside, and Mustelidee, and the penis,

which is supported by a ‘long bone (baculum), is susceptible of

protrusion from the preputial “orifice for the entire length of the

bone. The aperture of the urethra is on the ventral side of the

bone in some vascular tissue which can be stretched to a certain

extent beyond the apex of the bone.

In Ailurus, however, as recorded by Flower, the penis is com-

paratively quite short, the prepuce being close to the scrotum as

in the Felidze and Mungotidee, although the penis is longer than

in those families and structurally resembles that of the genera

referred to the Procyonide.

Hodgson declared that Ailurus has no scrotum; and in the

male examined by Flower that sac was represented by a pair of

swellings between the anus and prepuce, the testes being internal.

But in the male seen by me the testes had descended into the

scrotum, which formed a quite distinct swelling below the anus,

though not so constricted at the neck as in the examples of

Procyon, Nasua, and Potos examined by me*.

The baculum of Procyon lotor has long been known. It was

figured by Blainville (Ost. des Mamm. neenee i., Subursus, pl. viii. )s

and refigured and described by Gilbert (Morph. Jahrb. xviii.

p. 818, vl. xxvil. fig. 8, 1892). It is relatively longer than in

any other species referred to the Procyonide, surpassing 100 mm.

along the upper curvature, the distal third of the bone being

bent downwards. ‘The apex is expanded both transversely and

vertically, and forms a pair of condyle-like lobes separated by a

deep notch.

According to Hollister, the baculum of Procyon cancrivorus

(referred to the subgenus Huprocyon) is less curved than in the

typical form, P. lotor ; but the curvature is subject to a great

deal of variation in the latter, 7. e. from an angle of 135° to 90°.

In Nasua the baculum, as shown in Blainville’s figure (Joc. cit.),

* Tt may be added that Flower appeared to be quite unaware of the interest of

the fact when he published the discovery of the proximity of the prepuce to the

scrotum—a peculiarity in which Aizwrus differs from all other Arctoid Carnivora.

Proc. Zoou. Soc.—1921, No. XXVIII. — 28

412

a OmQ

MR. R. I. POCOCK ON THE EXTERNAL

Text-figuie 12.

——

—=

SSS

—=

—

———

Uy)

) Zi,

A vy iJ,

: Mi

Nae

N : i /

. Ventral view of scrotum and prepuce of Procyon lotor, showing

the abdominal position of the prepuce.

. Lateral view of penis of same partially protruded from prepuce, with

bristle thrust into urethral aperture; baculum dotted in.

. Anterior view of extremity of penis of same, with urethral orifice

between condyles of baculum.

. Inferior view of distal portion of penis of same, showing course of

urethral canal.

. Lateral view of penis of Maswa nasua partially protruded; the

slack ventral tissue pulled forwards and bristle thrust into

urethral aperture.

. Upper view of same.

. Lower view of same.

penis of Potos partially protruded.

9 99

. Ventral view of scrotum and prepuce of Ailurus fulgens, showing

scrotal position of prepuce.

. Lateral view of penis of same protruded almost to full extent, with

bristle passed into urethral orifice with loose ventral tissue.

All figures except H_X 3.

CHARACTERS OF THE PROCYONID. A138

is straight or slightly curved and normally * subcylindrical and

attenuated, with the apex expanded transversely and somewhat

abruptly to a considerable extent. The sides of the expanded

portion are rounded, and the distal margin is tolerably evenly

truncated or mesially concave, so that the tip is slightly bilobate.

The lower surface of the expanded termination is lightly convex ;

the upper to a corresponding degree concave.

In an adult male of each of the two species examined, namely

Nasua nasua and N. narica, there is a difference in the length

of the baculum. In the former species it measures 63 mm., in

the latter 77 mm. Possibly this is a specific character 7.

The baculum of Vasuella is, I believe, unknown.

The baculum of Bassariscus was figured by Blainville (Ost. des

Mamm., Atlas 1., Justela, pl. x.), and described and figured by

Gervais (Voy. de la ‘ Bonite,’ i. p. 20, Atlas, pl. iv.). Disregard-

ing the club-shaped base, it is tolerably evenly attenuated up

to the apex, which has a simple undivided slightly depressed

thickening, like the button on a foil. In the specimen figured by

Blainville it was rather strongly upcurled in its basal third, with

a slight downward distal curve, and measured about 50 mm. In

Gervais’ specimen it was straighter and measured the same,

which, judging from the figure of the skeleton, was about two-

thirds the basal length of the skull. Im the young example of

Bassariscus examined by me—an example in the stage of the tooth

change, the milk premolars being still in place—the baculum,

resembling in all essential particulars that figured by Gervais, is

less than half the basal length of the skull, the latter beng

65 mm. and the baculum 30 mm.

The baculum of Jentinkia sunuchrasti as figured and described

by Lonnberg (Anat. Anz. xxxvili. p. 232, fig. 2, 1911) measures

43 mm., is straight and subcylindrical, 2. e. only gradually nar-

rowed from the distal to the proximal end. ‘The tip is a simple

rounded condyle-like thickening ; but close behind it, rather on

the underside of the baculum, there is a pair of smaller rounded

condyle-like tubercles, recalling the similar accessory processes in

the baculum of Potos, which, however, project upwards and out-

wards.

As I have elsewhere remarked, the baculum of Jentinkia is

very like that of the Musteline Girison, except that the post-

apical processes in the latter are dorsal in direction. The pre-

sence of the two tubercles in question distinguishes the baculum

of Jentinkia from that of Bassariscus.

The only account of the baculum of Bassaricyon known to me

is that of Hollister, who described it as ‘32 mm. in length,

* Tn one example of Naswa nasua the bone shows an abnormal flattening on ths

right side.

+ In an adult but castrated male of Naswa nasua the baculum measures only

52 mm., and is exceedingly slender without any thickening at the proximal end.

The penis was correspondingly short as compared with that organ in the entire male.

One would perhaps expect the operation in question, if performed early in life, to

affect the development of the baculum.

9R*

“

A414 MR. R. I. POCOCK ON THE EXTERNAL

slightly bowed, and much less distinctly bilobed anteriorly than

in Procyon or Huprocyon.’ IL infer from this description that the

baculum is of a simple type, resembling somewhat closely that of

Nasua. I presume the bone was taken from an adult animal,

Tf so, its small size as compared with that of Potos and the

difference in the formation of the tip are interesting, although

there is, of course, no reason to suppose that BDassaricyon is in

any way nearly related to that genus despite their superficial

resemblance in some particulars. In connection with the com-

parative shortness of the bone, it must be remembered that

Bassaricyon is the smallest member of the so-called Procyonide.

Text-figure 13.

= ee:

L K G

A. Upper side of baculum of Potos.

B. The same of Bassariscus astutus (immat.).

OB chia che Ailurus fulgens.

TD) Oe hae ats Nasua nasua.

E. Proximal end of baculum of Az/urus from below.

F. Lateral view of baculum of the same.

G 5 ty bs Bassariscus astutus (after Blainville).

Jal. x, 55 apex of baculum of Nasua.

I. The same of Jentinkia (after Lénnberg).

Ree yess ay LEOUOSS

L. Upper side of tip of baculum of Jentinkia (after Lénnberg).

Figs. A, D, H, K X 4; the rest natural size approximately.

f=) 5) 3

In Potos the baculum is somewhat shorter than in Vaswa, but, as

in that genus, it is straight or slightly curved, subeylindrical, and

attenuated. The apex differs from that of all the genera referred

to the Procyonide. It ends in four condyle-like processes, one

smaller, directed upwards and outwards on each side, and two

at the end separated by a wide and deep notch. The two

CHARACTERS OF THE PROCYONIDAI, A415

bacula of this genus which I possess differ a little in length and

in the development of the terminal processes. One measures

65 mm., the other 60 mm. The former is narrower at the base,

but slightly thicker at the neck behind the two proximal pro-

cesses, and these are smaller, a little less salient, and directed

slightly less upwards. The baculum is wider across them than

across the terminal processes, which project straight forwards and

converge slightly at their inner angles, so that the notch between

them is a nearly complete oval. In the shorter of the two the

proximal processes are larger and the terminal processes diverge

slightly from the middle line, so that the width across them is

equal to the width across the proximal processes, and the notch

between them is wider, and not narrower, at its mouth.

Flower figured and deseribed the baculum of Adlurus. Con-

forming to the size of the penis, it is relatively very short, being

only about an inch in length*. Its upper side is markedly con-

cave, owing to the slightly upturned apex and the rather strongly

elevated base. Its proximal half is flat below but compressed

and carinate above. The tip is a little expanded and truncated,

with rounded angles and only very obscurely bilobed, and there is

a short median groove on the upper side just behind the tip.

The epithet spatulate” applied by Flower to the tip of the

specimen he examined, does not express the shape of the termi-

nation of the bone in my example. This bone measures about

23 mm.

By the penis and baculum the genera may be distinguished as

follows :—

a. Penis short, prepuce close to scrotum; baculum short, its apex

simple and upturned, its base with a high keel ..................... Ailurus.

a’. Penis long, prepuce abdominal; baculum long, not upcurled

apically, its base clavate, not str ongly keeled.

6. Baculum ending in four short, radiating condyle-like branches. Potvs.

6’. Bacnlum ending otherwise.

c. Baculum curved downwards distally, its apex fonene two

rather deeply cleft condyle-like processes .......... eee ROCU OI.

. Baculum straight, or nearly so, distally, its apex oe anost

indistinetly bifid.

d. Baculum with sur flattened, ae indis-

tinctly bifid extremity .. ogee ae GI Wee ey ... Nasua

“(and ? Bassaricyon).

d’, Baculum with rounded apex.

e. ‘Two small condyle-like processes just behind the apex ... Jentinkia.

e’. No accessory processes behind apex .................000+... Bassariscus.

The length of the penis and the position of the prepuce, two

very important points in the classification of the Carnivora,

are unknown in Ailwropoda. The Urside resemble the typical

Procyonide in those characters; and in all the species I have

examined the baculum is a long, stout, distally tapering bone,

with a simple termination.

* That is to say about one-third of the length of the baculum in Pogos, a smaller

animal.

416 MR. R. I. POCOCK ON THE EXTERNAL

There is very little to say about the external genitalia of the

female. In Procyon and Ailurus the vulva occupies the same

relative position as in Canide and Urside. In both genera it is

an oval or egg-shaped exerescence, with hairy labia surrounding a

central pit, the clitoris being a small exerescence near the lower

edge of the pit, and above the clitoris is the orifice of the vagina.

In Ailurus the clitoris contains a small bone; but I have no note

of this in Procyon. In Potos the vulva is relatively a somewhat

shorter angular prominence, with the orifice forming a transverse,

not a vertical, rima—an adaptation possibly to the width of the

tip of the baculum of the male. I found no clitoris within the

orifice. Beddard did not describe the vulva in the female example

of Bassaricyon he examined.

The Haternal Characters as « Guide to Classification.

A review of the above-recorded characters brings to light

some interesting facts bearing on the classification of the genera

examined,

There is nothing special to say about the vibrisse except their

high development in Vusua.

The external ear shows in its variation generic features, but

practically no evidence of close intergeneric kinship.

The muzzle and rhinarium also exhibit good generic characters ;

while the rhinarium of Vaswa differs markedly in its specialization

from that of the rest.

In the case of the feet, however, a marked difference may be

noticed between Ailurus and the rest. In the latter a gradation

may be traced from Bassariscus through Jentinkia to Bassaricyon,

from the latter to Vaswa on the one side and to Potos on the

other, and from Potos to Procyon.

Ailurus also stands alone in possessing a specialized glandular

area round the anus. ‘lhe presence of normal anal glands both

in Adlwrus and Procyon merely indicates the mutual inheritance

of a primitive feature; but the modification of these glands met

with in Vaswa is an acquired differentiating peculiarity, as also

is their loss by Potos. The latter is also peculiar in the possession

of the ventral glands.

Finally, Adluwrus differs markedly from the rest of the genera

in the scrotal position of the prepuce, the shortness of the penis

and baculum, arid the strueture of the base of the baculum itself.

The others retain the primitive abdominal position of the prepuce

and the large baculum seen in typical Arctoids, all Cynoids, some

AAluroids (e. g. Cryptoprocta), and the Pinnipedes.

The above-mentioned peculiarities of Ailurus, coupled with the—

better-known peculiarities of the skull and teeth, justify the

severance of that genus from the American Procyonide as. pro-

posed by Gray and upheld by Gill, Turner, and Flower, though

CHARACTERS OF THE PROCYONIDA. 417

subsequently abandoned by the latter and by most recent

authors.

Potos, too, has several peculiarities in its external characters as

well as in its skull and teeth; and probably no one will cavil at

the adoption of Trouessart’s view that it should stand as the

representative of a special subfamily of Procyonide, namely

Potosine.

The status of the remaining genera is not so easy to settle.

Taking first the older known forms, Procyon, Nasua, and Bassa-

riscus, there does not seem to be evidence of any close affinity

between them, a fact clearly perceived by Gray and Gill, although

by making Vaswa and Procyon the types of special subfamilies of

Procyonid, Gill expressed his idea of closer kinship between

them than between either of them and Bassariscus, which he kept

in a family apart. Even quite recently Hollister has adopted the

view that Bassarisews should rank as a separate family.

The discovery of Bassaricyon, since the time of Gray and Gill,

does not help matters, since the genus is equally isolated and

serves in no respect to affiliate any two of the other three. The

four genera, in fact, differ in a large number of characters, to any

one of which full generic value would be granted nowadays; and

the logical inference to be drawn from this argument is that the

sum of the characters demands supergeneric recognition, which

should be expressed systematically by elevating the genera to the

rank of subfamilies. Further justification for this course may be

found in following the present day tendency to grant full generic

value to the characters upon which such forms as Luprocyon,

Nasuella, and Jentinkia were founded. We shail then have the

Procyonine, the Nasuine, and the Bassariscinze with two genera

each and the Bassaricyoninz with one.

The Position of Ailuropoda.

The question of the systematic position of Aduropoda cannot

be passed by in a paper dealing with the Procyonide, since the

genus has been referred to that family. Milne Edwards con-

tented himself with pointing out the resemblances between

Ailuropoda and Ailurus on the one hand, and Aiwropoda and

the Urside on the other. And, so far as I am aware, Mivart was

the first author definitely to state the opinion that Ailuropoda is

more nearly akin to the Procyonide, with which it is affiliated

through Ailwrus, than to the Urside; and this opinion found

practical expression in the ascription of Ailuropoda to the

Procyonid, under a special subfamily also including Ailurus.

Mivart’s view was adopted, with the support of much additional

evidence, by Lankester and Lydekker, and Mivart’s classification

was independently reached. Finally, Bardenfleth attempted to

show that Flower was right in classifying Ailwropoda in the

Urside.

418 MR. R. I. POCOCK ON THE EXTERNAL

Bardenfleth appears to have been much influenced by Winge’s

opinion on the homologies of the cusps in pm. 4 of the upper

jaw, which, admitting its correctness, shows that the structure of

the tooth in question lessens the evidence for the affinity between

Ailuropoda and Ailurus and strengthens the evidence for aftinity

between the former and the Urside. The points are:—(1) The

antero-internal cusp of Azlurws corresponds to the postero-

internal cusp of Arluropoda; (2) the postero-internal cusp cf

Ailurus is unrepresented in Ailuwropoda and the antero-internal

cusp of Ailuropoda is unrepresented in Ailurus; (3) the postero-

internal cusp of Ailwropoda corresponds to the postero-internal

cusp of the Urside; (4) the antero-external and antero-internal

cusps of Ailuropoda are unrepresented in the Ursidee—in other

words, pm. 4 of the maxilla of the Urside corresponds to the

posterior half of the same tooth in Aiduropoda, with the further

difference that the inner and posterior roots of this tooth in the

Urside are, except in abnormal cases, fused. Nevertheless, the

resemblances between this tooth in Adluropoda and Urside do

not, in my opinion, justify the conclusion that Ailuropoda is a

member of that family. For, if the tooth in Ailwropoda is not

Ailurine or Procyonine, it is certainly not Ursine. It is peculiar

and stands by itself, so far as living Carnivora are concerned.

Some of the characters cited as of systematic value by Lydekker

and Bardenfleth are too inconstant to be of use. Steno’s fissure,

the cleft between the anterior palatine (incisive) foramina, is

cited by Lydekker as diagnostic of the Procyonine, restricted to

the American genera, which have it, and the Ailurine, including

Ailurus and Ailuropoda, which ave without it; and Bardenfleth

states that it is present in Ursus and absent in Ailurws and

Ailuropoda, The actual facts are as follows :—Even in the com-

paratively small series of skulls belonging to the Zoological

Society, I find this fissure present in an example of dAdlwrus,

where it is represented by a groove terminating in a foramen

which passes up into the nasal passage—a condition which it

commonly presents, even within the limits ef the genus Procyon.

As for the Urside, it varies in size from a long cleft to a

minute orifice, through which a needle-point can only just be

passed.

Lydekker, endorsing Lankester’s statement, also draws attention

to the “important fact that Aiurus and Ailuropoda resemble

the American Procyonide, and thereby differ from all other Car-

nivora, in the presence of two lobes or cusps on the inner or

tubercular portion of the upper carnassial,” Even if he had said

“some of the American Procyonide,” the statement would not

have been true, because both Helictis and Taaidea, two of the

Mustelide, have two cusps on the inner portion of the tooth.

Again, ‘“&luropus approximates to the Racoon in the absence of

postorbital processes of the frontals.” This is not true, the frontal

postorbital processes being well developed in all the skulls of

CHARACTERS OF THE PROCYONIDA. 419

Procyon I possess*. Farther on we read: “A point of resemblance

between Ursus ft and dluropus is to be found in the circumstance

that the maxillary [preorbital] foramen opens on the side of the

skull well in advance of the zygomatic root, whereas in Zlurus

and Procyon it perforates the zygoma itself.” So far as this

statement is applied to the Urside, it is contradicted by the

Malayan Bear, in which the foramen perforates the zygoma exactly

as in Adlwrus and all the American Procyonide. I may also add,

in this short-headed species of Ursidee the zygomatic width of the

skull bears to the basal length about the same proportion as in

Ailuropoda, thus contradicting Lankester’s statement that ‘in all

Bears the skull is much longer and narrower both in its facial

and cranial regions than in the skull of Zluropus.” Bardenfleth

similarly wrongly contrasts Zluropus with Ursus when he says

of the former “zygomatic arches exceedingly wide” and of the

latter “zygomatic arches moderately wide”; and his statements

that the basioccipital is broad and the bulla not inflated in Ursus

are not always true of the species in question. These cor-

rections, however, are of no great moment in settling the degree

of kinship between the Urside, Ailwropoda, and Ailurus—the

point at issue between the authors quoted. What they prove is

that the resemblances between the genera concerned are closer

than either author claimed.

My own opinion about the matter is that Lankester and

Liydekker, as Bardenfleth held, overrated the resemblances be-

tween Atluropoda and Ailurus and underrated the differences ;

and that Bardenfleth underrated the peculiarities of Ailwropoda

which distinguish it from the Urside. The attempt to place

Ailuropoda in the Procyonide makes the definition of that family

an impossibility. For example, every character used by Lydekker

for defining that family has its exception. The same criticism

cannot be advanced against its inclusion in the Ursidew, because in

one or two well-marked characters, like the structure of the feet,

the length of the tail, and the presence of m. 3 in the mandible,

Ailuropoda and the genera of Ursidz resemble each other, and

differ from <Ailurus and the American genera assigned to the

Procyonide. But since the assignment of Ailuropoda to the

Urside disturbs the homogeneity of that family, which already

has some half-dozen well-defined genera, I prefer to regard

Ailuropoda as the representative of a distinct family. The genus

is neither Ursid nor Procyonid, but something distinct from

both.

* Bardenfleth also is wrong in stating that Melursus is without frontal post-

orbital processes.

+ Fig. 3, Pl. xix. of Lydekker’s paper (Tr. Linn. Soc., Zool. vol. viii.) is part of the

skull of an Ursus, not of a Procyon as labelled.

A420 MR. BR. 1. POCOCK ON THE EXTERNAL

Dichotomous Classification of the Urside, Ailuropoda, Ailurus,

and the Procyonide.

Although this method of classifying animals has its disadvan-

tages on account of its deceptive simplicity and its concealment

of cross resemblances, it has the convenience, if not admittedly

artificial, of presenting clearly the characters relied upon by the

author and of expressing his views as to the kinship of the

groups concerned.

a. Feet short and broad, digits subequal, their pads forming a lightly

curved line in front of the broad plantar pad. Tail reduced to an

anal operculum. J. 3 of mandible retained. Carotid foramen

situated at posterior end of bulla, close to foramen lacerum

posticum.

a’. Cheek teeth comparatively small, especially pm. 1—pm. 38, which

are mostly unicuspid and one-rooted, but pm. 1 larger and moie

persistent than pm. 2; upper pm. 4 at most tricuspid, its inner

root normally fused with the posterior root. Lower pm. 4 short,

unicuspid. Zygoma arising approximately above the middle of

m. 2, which is tar in advance of the mesopterygoid fossa, the

latter preceded by a long edentulous posterior palate. Alisphe-

noid canal present. External auditory meatus with its floor

produced to reach approximately to end of thick mastoid process.

Mandible not thickened on inner side of coronoid, which does

not conceal m. 3 and is comparatively low and but little hooked ;

angular well developed, condyle and glenoid surface not abuorm-

ally wide. Fore foot without long radio-carpal sesamoid etc.*... URsipa.

a>, Cheek teeth excessively developed, except pm. 1 of upper jaw,

which is minute in maxilla, and absent in mandible; pm. 2 of

maxilla and pm. 2 and pm. 3 of mandible tricuspid; pm. 3 of

maxilla sexcuspid ; pm. 4 of same, with three large outer and two

large inner cusps, its inner root not fused with posterior root ;

lower pin. 4 long, tricuspid. Zygoma arising approximately above

middle of m.1; posterior end of m.2 reached or overlapped by

anterior end of mesopterygoid fossa; no long edentulous pos-

terior palate. Alisphenoid canal absent. External auditory

meatus with its floor abbreviated and falling far short of

long compressed mastoid. Mandible thickened on inner side

of coronoid, which conceals m.3 and is high and hooked ; angular

much reduced ; condyle and glenoid surface abnormally wide.

Fore foot with long radio-carpal sesamoid ete.* ......... AILUROPODIDS.

6. Feet comparatively long and slender }, digits unequal, with their pads

arranged in a strongly curved line round the comparatively narrow

plantar pad. Tailat least long enough to reach the ground. W.3

of mandible suppressed. Carotid foramen on inner side of bulla

well in advance of foramen lacerum pesticum.

6’. Penis short, prepuce close to scrotum. Pads of feet reduced and

functionless, completely concealed by woolly hair; carpal pad

remote from plantar pad. Anus in centre of glandular depressed

area. Pm. 2 and pm.3 of maxilla large and three-rooted; pm. 3

quinquecuspid and closely resembling pm. 4. Alisphenoid canal

present. Foramen rotundum minute, lying beneath for. lac.

ant., the two separated by a very thin plate of bone and sunk

in a common pit; foramen ovale elongate. Anterior edge of

coronoid imelinedstouwands) J. 22 .o sehen | ste eeeetae acecyote- cists) AML URAL MeAaye

* For other skeletal characters distinguishing the Urside and Ailuropodide, see

the papers by Lankester, Lydekker, and Bardenfleth.

+ It is hardly an exaggeration to say that there is no such thing as a plantigrade

carnivore. They all run and walk on the digital and plantar pads, whether they are

cats, dogs, bears, or badgers. Bears, when standing on their hind legs, and some-

times when walking, place the metatarsal pads on the ground; but generally these

pads, like the carpal pads, are raised from the ground in ordinary progression. A

naked metatarsal area does not indicate platigradism, as has been supposed.

a CHARACTERS OF THE PROCYONID. 49]

62. Penis long, prepuce abdominal, remote from scrotum. Feet with

digital and plantar pads normally developed and quite naked;

carpal pad or pads also well developed and close to the plantar

pad. No Bingen area round anus. Pm. 2'and pin. 3 of

reese comparatively small and one- or two-rooted*; pm. 3

unlike pm. 4, at most with one main cusp and three ‘minute

cusps. Alisphenoid canal absent. Foramen rotundum of

normal size, well separated from the for. lac. ant. and not sunk

in a common pit with it; foramen ovale rounded. Anterior

edge of coronoid inclined backwards .......................... PROCYONIDA.

c. Mandible very massive, the rami early fused by a long sym-

physis, the lower edge straight by the growth posteriorly of a

large lamina, sometimes rounded, sometimes angled, beneath

the angular process, which is reduced to a small excrescence

beneath the condyle+; inner dentary foramen beneath base of

anterior edge of coronoid. Palate parallel-sided, as wide in

front as. behind, depressed behind molars; paroccipitals and

mastoids standing away from bulla, which has carotid fora-

men set forwards much nearer for. lac. med. than for. lac.

post.; molars even when newly cut fiat-crowned ; pm. 1 absent

above and below. ‘Tail prehensile. ‘Iwo ventral cutaneous

glands Anal glands aborted .. Mel: . Potosine.

. Mandible slender, the rami weparable arial: Wied, itemnall & sym-

physis, lower edce arched and inclining upwards without

expansion towards angular process, which is well developed

and close beneath the condyle; inner dentary foramen about

midway between the condyle and the anterior margin of the

coronoid. Palate with arcuate sides, wider behind than in

front, not depressed behind molars; paroccipitals and mastoids

arising close to the bulla, which have the carotid foramen

about equally distant from for. lac. med. and for. lac. post.,

or nearer the latter. Molars cuspidate when newly cut; pm. 1

present. Tail not prehensile. No ventral glands. (Anal

glands retained in Bassariscus, Procyon and Nasua, unre-

corded in Bassaricyon.)

d. Mesopterygoid fossa long, extending to posterior molars ;

molars and premolars with poimted or blade-like cusps ;

canines rounded in section, not grooved and not mutually

sharpened. Bursa of ear with anterior flap emarginate,

posterior flap attached behind edge of pinna. Metatarsus

with at most a small naked areaabove plantar pad; claws

short, sharp, curved; digits fully webbed or nearly so.

Vacial portion of skull and zygoma as under e; hamular

in front of foramen ovale; no supplementary foramen

behind carotid ............... .... Bassariscine §.

d’. Mesopterygoid fossa short, Ronee ae olan by a long

posterior palate. Molars with subequal conical compara-

tively blunt cusps ; canines grooved, mutnally sharpened.

Bursa absent or reduced, its anterior flap at most very low

and not excised. Metatarsus naked beneath.

* Hxceptionally in Procyon, pm. 3 is three-rooted.

+ This at all events is, 1 believe, the correct interpretation of this region of the

mandible.

t Huet figures a mandible with it present on the right side.

§ Hollister g gave full family rank to this group, because “ The Cacomistles (Bassa-

riscus), while exhibiting many of the characters of the Procyonide, differs so

ereatly in the nature of all the teeth that it seems inpossible to retain them in the

family. ‘The dog-like premolars and molars, the rounded canines, and the evident

though small secondary lobes on the incisors ..... all show unmistakable characters

of ‘the teeth of the Canidse.” Nevertheless, by a series of comparatively simple

changes, the teeth of Bassariscus can be derived from those of such a Procyonid as

Nasua, for example, or vice versa, and the secondary grooves and lobes on the

incisors are not always more marked in Bassarviscus than they are in Potos, and the

grooves are often traceable in other genera. ‘The teeth of all the genera attest the

extreme plasticity of those organs. (See above, p. 418.)

22 THE EXTERNAL CHARACTERS OF THE PROCYONIDA.

e. Skuil with muzzle short, not compressed above, preorbital

foramen close to anterior rim of orbit, which is above

pm.3 or pm.4; inferior edge of anterior nares normally

formed ; canines only moderately sharp im front; pos-

terior root of zygoma not abnormally expanded;

typically no supplementary foramen on inner side of

bulla between carotid and foramen lacerum posticum.

Upper edge of rhinarium not produced, its anterior

surface convex, with normal nostrils and wide septum.

Ff. Mastoid small, not bigger than paroccipital and

hardly surpassing auditory orifice; carotid foramen

not behind middle of bulla; foramen ovale well

behind tip of bamular; ridges of posterior palate

marginal. Claws short, sharp, much curved; digits

webbed for two-thirds of their length. Snout not

produced; upper lip divided by wide philtrum.

Bursa retained and marginal ................... Bassaricyonine.

Ff. Mastoid large, larger than paroccipital and projecting

far beyond auditory orifice ; carotid foramen behind

middle of bulla; foramen ovale not behind tip of

hamular; ridges invading lower surface of posterior

palate. Claws longer, blunter, and less curved;

digits entirely free from webbing. Snout produced,

upper lip undivided, no philtrum. Two normal anal

glands. Bursa absent ().-..0.02.. 22-2. c2-c0--tees--ee LProcyonine:

e’. Skull with muzzle elongate and compressed above; pre-

orbital foramen remote from orbit, the anterior rim of

which is above m’; interior edge of anterior nares

thickened and produced; glenoid portion of zygoma

much extended antero-posteriorly; a very distinct sup-

plementary foramen between carotid and foramen

lacerum posticum. Upper edge of rhinarium produced,

its anterior surface nearly flat, with vertically elongated

nostrils and narrow septum. Digits with powerful

claws, fully webbed. Anal glands a series of small

pockets on each side. Mastoid as under f: palate as

under f’; position of carotid foramen and f- ovale

intermediate. Bursa retained but not marginal ...,.. Naswine.

IQA SIMOOU SI, IP,

Poa Zas:

oa!

REPTILES AND BATRACHIANS FROM SOUTHERN ANNAM.

my ZS. 1921, SIMONA, JP Wk.

BATRACHIANS FROM SOUTHERN ANNAM.

ON REPTILES AND BATRACHIANS FROM SOUTHERN ANNAM. 423

22. New or Little-known Reptiles and Batrachians from

Southern Annam (Indo-China). By Matcorm A.

Simms MeReCise lak: Ciba ZnS.

(Plates I.-IT. and Text-figures 1-2.)

[Received February 22, 1921: Read April 19, 1921.]

INDEX.

Page Page

Fimbrios klossi, gen. & sp. n....... 425 | Lygosoma stellatum ........ 1... 431

ZAMenisS MOt, SP. Ne ...vesesveereeeeee AZ5B Fé corpulentum, sp.n. ... 431

Tropidonotus johannis............... 426 | Dibamus montanus, sp.n. ......... 431

Coluber oxycephalus .............5. 426 hana millvetis Sp. settee AO

Dendrelaphis subocularis ......... 426 | 5 COLGTROOTATED,” 5otnn oaoasoces tes 433

Calamaria pavimentata » sautert

Var. wniformis, NOV............. 426 Wiareohns7,.ROVA \.netpeees hs Ok

Trimeresurus monticola ...........- 427 BS) LOIMULUAO ASPs Ms) ees riec state 436

Gymnodactylus pegquensis ......... 427 6) GIPEMUIDED scosoRecsece ogasoeccune AST!

Warsianiguloniss Nove tse ooo Mncrony lo pictay wake .e soe .eeseeen es AOU

Var. irregularis, nov. ......... 428 | Bufo galeatus ....... . 438

Calotes microlepis .................. 428 | Megalophrys inter medius, sp. n. . 439

Bue RINYSCACEUS eracve nsec: 4.29 a hasseltit

Liolepis belliana Var. pullus, NOV. ..ccceeeeeese 440

Var. annamensis, nov. ......... 429

The collection of Reptiles and Batrachians of which this paper

is the subject was made in Southern Annam, chiefly on the

Langbian Plateau, in March, April, and May, 1917. I was

fortunate in having as my companion on the trip Mr. C. Boden

Kloss, of the Selangor Museum, Federated Malay States, and as

he was able to remain on in the country for several weeks after

T had left, was the means of adding many valuable specimens to

my collection.

The famous Plateau had long attracted us, as zoologically it was

almost unknown; and as, owing to the war, we were unable to

obtain home leave, of which we were badly in need, we decided

to take advantage of a short holiday and visit this region. The

enterprise of the French Government, too, in building a sana-

torium at a high elevation, and in constructing fine roads up to it,

rendered the “plateau easily accessible, so that intitle time was

wasted in travelling.

Our expectations of rich material were fully justified. The

report on the birds by Messrs. Robinson and Kloss has already

appeared in ‘ The Ibis’ (July 1919), and a good account of the

general conditions on the plateau has been given there by

Mr. Kloss. It is unnecessary for me to repeat his remarks here,

but for convenience I have recapitulated the camps at which

collections were made. Starting from the sea-coast at Tour Cham,

we gradually made our way up into the hills, our final camp

being underneath the Langbian peaks at 2000 metres elevation.

T was accompanied by two trained native assistants. Altogether

about 700 specimens were obtained, Many more of the common

424 DR. M. A. SMITH ON REPTILES AND

forms could have been taken, but with a good series in hand, and

the difficulties in transporting heavy collecting-tanks about the

country, it was necessary to place some limit upon what was

caught.

The number of snakes obtained was not great, and most of

these were caught after the rains had commenced in the middle

of April. Mountain-streams abounded everywhere, and frogs,

chiefly of the genus Rana, were plentiful. A fine series of

Rhacophorus and Ixalus were also taken, During the day these

tree-frogs were seldom seen. their small size and fine protective

colouration rendering them almost invisible among the bushes in

which they lived. Their shrill cries by night, however, guided one

quickly to them, and by means of a lantern they were easily found

and taken. — ae

Perhaps the two most interesting discoveries of the trip were

a new genus of snake, allied to Yenodermus, and a new species of

the degraded skink, Dibamus. A new form of Gymnodactylus

peguensis was. obtained on the plateau, but, with the exception of

the common house-geckoes and the equally common Phyllo-

dactylus siamensis, the family Geckonid# was remarkable for

its absence.... The same can be said of Tropidophorus, a single

specimen only being obtained, although in search of frogs the

collectors were daily working along streams. Three new forms

of Rana and two of Megalophrys are also described in this paper.

Want of time has prevented me, for the present, from completing

my examination of the Rhacophorus, the Jxalus, and the smaller

species of Lygosoma.

On the whole, the Reptilian and Batrachian fauna of the

plateau, so far as my examination extends, approximates most

nearly to that of the hills of Siam and Southern Burma and the

higher hills of the Malay Peninsula. A few species only are

related to those of more northern origin.

Types of all the species here described have been presented to

the British Museum of Natural History.

Finally, I wish to express my thanks to Mr. G. A. Boulenger,

E.R.S., for his valuable help in several difficult determinations.

The following localities were collected in :—

Tour Cham, on the sea-coast (lat. 12° N.).

Daban, in the foot-hills at 200 metres altitude. Dry, deciduous

jungle, but fairly dense.

Sui Kat and Dran, localities about 6 kilometres apart in the

hills, at 1000 metres elevation. Chiefly evergreen jungle, with

some useful small swamps.

Dalat, Camly, Le Bosquet, Arbre Broyé, localities on the

plateau at 1200 to 1800 metres. The country at the two first-

named camps was chiefly open pine-forests, but at the two last

dense evergreen jungle was met with.

Langbian peaks, 2000 metres. Mixed forest, some pine, more

oak,

BATRACHIANS FROM SOUTHERN ANNAM. 425

Fimprios, gen. nov. (PI. I. fig. 1.)

Teeth subequal, 30-32 in each maxillary; head not very

distinct from neck, covered with large shields; eye small, with

round pupil; loreal very large, touching the eye, nostril in the

anterior part of a large nasal; body slender, scales elliptical,

keeled, juxtaposed anteriorly, feebly imbricate posteriorly, those

of the outer row larger than the others ; ventrals large, rounded ;

tail rather short, subeaudals single.

Allied to Xenodermus Reinhardt.

FIMBRIOS KLOSSI, sp, n.*

Nostril in the anterior part of a large, concave nasal; rostral

triangular, concave, not visible above ; internasals much smaller

than the prefrontals and separated from the rostral by a

horizontal ridge of the skin; frontal as broad as long, longer

than its distance to the end of the snout, much shorter than

the parietals; supraocular very small and narrow; preocular

small, just touching the frontal; a large square loreal in contact

with the eye; two postoculars and a subocular; temporals

small, 3+4; 9 or 10 supralabials, the first 5 very small, with

strongly raised edges, the last one much elongated; no mental ;

12 infralabials, the first 7 very small and with their edges raised

like the supralabials, Ist and 2nd pairs im contact with each

other ; a pair of very large chin-shields.

28 to 30 scales round the anterior part of the body, 30 to 32

round the middle; ventrals 162 to 167; anal 1; subcaudals

43 to 58.

Dark grey above, yellowish (in life white) below, the edges of

the posterior ventrals and subcaudals tinged with grey.

Three specimens obtained at Dalat and Camly at 1500 metres.

Measurements of the type series in mm. :—

Author’s No. Total length. Tail. Ventrals. Subcaudals..

DAA OO vaemae 395 50 166 43

DANAUS a) nee 345 68 162 57

yA 3 a 310 60 167 58

These remarkable snakes were caught beneath fallen timber.

They were quiet and gentle in their movements, and made no

attempt to bite when handled. I kept one alive for a few days

in the hopes of learning something of its habits, but difficulties

of transport prevented my doing this as long as I should haye

wished.

ZAMENIS MOTI, sp. 0.7

Maxillary teeth 18; eye moderately large; rostral consider-

ably broader than deep; internasals shorter than the prefrontals ;

* Named after Mr. C. Boden Kloss, to whom I am indebted for two of the

three specimens.

+ Named after the Moi people, the aboriginal inhabitants of the country in

which it was found,

426 DR. M. A. SMITH ON REPTILES AND

frontal 11 times as long as broad, longer than its distance to the

end of the snout, shorter than the parietals; loreal twice as long

as high ; one pre-and two post-oculars ; no subocular ; temporals

1+2; 8 supralabials, 4th and 5th touching the eye; 6 infra-

labials, 4th very large; 4 infralabials in contact with the anterior

chin-shields, which are as long as the posterior; posterior chin-

shields in contact anteriorly. Scales in 15 rows throughout,

entirely smooth; ventrals, rounded, 168; anals 2; subcaudals

103 pairs.

Olive-greenish above, with indistinct, narrow, pale, dark-edged

cross-bars on the posterior part of the body and tail. Below

yellowish, speckled with grey on the posterior two-thirds; a dark

median streak between the subcaudals.

Total length 1000 mm., tail 290.

Allied to Z. korros Schleg.

A single male specimen collected at Dran (1000 metres) by

Mr. Boden Kloss in May 1917. Author’s number, 2153.

TROPIDONOTUS JOHANNIS Blgr.

Ann. & Mag. Nat. Hist. (8) ii. 244, Sept. 1908.

Ten examples from the Plateau differ from typical yohanmis

only in the supralabial shields. Two of them have 8 on one side,

9 on the other; all the rest have 9. From 7. modestus Ginther,

which it resembles very closely, it differs in the fewer caudal

shields and in the colouration of the belly.

Variation in my series:—Scales 19-17, ventrals 149-159,

caudals 83-98. Five out ef the ten specimens have the tail more

or less docked. Largest: total length 640 mm., tail 185(¢ ).

Colour. Brown above, with small black spots, and a series of

small yellow ones also present in most. Labials with black

sutures, and a yellow streak from the last labial to meet its fellow

on the nape. Belly yellowish white, with a black spot at the

outer side of each ventral.

CoLUBER OXYCEPHALUS Boie.

Bouleng., Rept. Malay Pen. p. 143 (1912).

1 ex. from. Daban. Scales 23, 23, 15. V. 245. C. 130.

Total length 1880mm., tail 480. Green above, yellowish below,

tail pale reddish-buff (in spirits).

DENDRELAPHIS SUBOCULARIS Bler.

Cat. Sn. B. M. ii. p. 89 (1894).

1 ex., Dran. Scales 15, 15,11. V. 165. C. 98. 8 supra-

labials.

CALAMARIA PAVIMENTATA D. & B.

Cat. Sn. B. M. ii. p. 348 (1894). ‘

Var. UNIFORMIS, nov.

Differs from the present known forms in its distinctive

colouration.

BATRACHIANS FROM SOUTHERN ANNAM, 497

Olive-brown above, uniform (no longitudinal lines or collar),

below yellowish-white, with a median line along the tail and

usually another down the belly. Labials yellow.

10 examples examined. Variation: 4. V. 143-149; C. 30-34.

O77 MeetGG) Voss. 8. 19%

Type locality, Langbian peaks at 2000 metres.

Type series, Author's Nos. 2135, 2136, 2137, and 2139.

'TRIMERESURUS MONTICOLA Giinther.

Lachesis monticola, Cat. Sn. B. M. iti. p. 548 (1896).

lex. The specimen differs from the recognized description in

having all the subcaudal shields single, and in having only two

rows of scales between the eye and the labials. Wali, however,

records one from the Chin Hills (Journ. Bombay N. H. 5S. xx.

p- 775), in which the subeaudals are irregularly single and paired,

and J find in the British Museum an example in which there are

only two rows of suboculars. For the present, therefore, I regard

my specimen as 7’. monticola. Scales 21, 15. Anall. V. 132.

C. 38. Six scales between the supraoculars.

GYMNODACTYLUS PEGUENSIS and subsp.

G. peguensis, Bouleng. Rept. Malay Pen. p. 36 (1912).

Two specimens of a Gymnodactylus obtained at Camly agree

well in characters with the typical form of G’. peguensis, but

differ distinctly in colouration. Both are males, and in both the

tails are missing, and it is possible, with more complete material

for examination, that the present diagnosis will be found in-

correct. I refer them here to G. pegwensis, and at the same

time take the opportunity to describe another colour race which

T have obtained in Eastern Siam.

The three forms may be described as follows :—

GYMNODACTYLUS PEGUENSIS, forma typica. (Text-fig. 1, A.)

7-8 preanal pores;. 9-11 upper and 7-9 lower labials; two

series of (6—8) large round spots on the back, or with the spots

confluent transversely. Head in the adult with large rounded

spots.

Type locality, Palon, Pegu. Has been found also in other

parts of Pegu and in Peninsular Siam as far north as Nakon

Sri Tamarat.

Var. ANGULARIS, nov. (Text-fig. 1, C.)

10-11 upper and 9-10 lower labials; two series of (4) large

angular spots connected mesially, Head in the adult with

indistinct angular spots. .

Habitat. Dong Rek Mts., Eastern Siam.

Proc. Zoou. Soc,—1921, No, XXIX. 29

428 DR. M. A, SMITH ON REPTILES AND

Text-figure 1.

Gymnodactylus peguensis.

A. Forma typica. B. irregularis. C. angularis.

Measurements of specimens known, in mm. :—

Head &

body. ‘Tail. Arm. Leg.

5364. Lat Bua Kao, Korat (Type). 92 68 80 27 32

D028. Pal’ Jone Ae ene hee tec eekee Q 65 70 25 32

PAUP Hi le bay Bay oj. ane se dads SRE nes juv. 34 35 13 eA

Var. IRREGULARIS, nov. (Text-fig. 1, B.)

5-7 prenal pores; 8-9 upper and 8-9 lower labials; small

angular spots not arranged in any very marked pattern; head

with well-marked angular spots.

Habitat. Langbian Plateau.

Head &

body. Arm. Leg.

A875. Camily (Type).-.......... 6 09 28 36

AST6. “LE CWB 5a ayer 3 80°. 29 37

CALOTES MICROLEPIS Bler.

Fauna Brit. India, p. 134 (1890).

1 ex., Camly. The specimen, a 2, agrees well with the type

in the British Museum. It has 70 scales round the middle of

BATRACHIANS FROM SOUTHERN ANNAM. 499

the body. Head and body, 80mm.; tail, 180. Brown above,

finely speckled with black and yellow. Below whitish, speckled.

Black lines radiating from the eyes.

The characters given by Boulenger (Rept. Malay Pen. p. 70)

to separate this form from floweri, namely narrower head and

more compressed tail, will not stand the test of my specimens.

I have examined two examples of flowert from Chantabun, in

addition to the two in the Museum. They have from 50-55

scales round the middle of the body,

I separate microlepis as having more scales, 65-70, round the

body, smaller ventrals and smaller tympanum ; it is possible that

flowert is only a Southern form of this species.

I have examined pregnant females of both forms. The eggs

are oval,

CALotES mystaceus D. & B.

KY, BEL. p. 138i (L890),

Specimens obtained at Saigon and on the Langbian Plateau

lack the three chocolate spots on the back which appear to

characterize the form found west of the Mekong River. Enough,

however, is not yet known of this handsome lizard in Burma to

name races definitely. The plateau is a considerable extension

eastwards of its known habitat.

LIOLEPIS BELLIANA Gray.

Bouleng., Rept. Malay Pen. p. 73 (1912).

Var. ANNAMENSIS, Novy.

When collecting upon the sea-coast at Tour Cham, before

ascending the Plateau, our attention was attracted by the marked

difference in colour between the form of Z. belliana which we

observed there, and the one which we knew so well. from Siam

and the Malay Peninsula. The vivid orange bars upon the flanks

of the typical form were replaced by bands of pure white.

A detailed examination of the specimens obtained showed

further that, in the number of femoral pores, and in the size of

the seales behind the tibia, they differed from the typical form.

I distinguish the two as follows :—

Forma typica.

13-20, usually 15-18, femoral pores.

(ealbwadenr 8-10, scales across the back of the middle of

the tibia.

Flank with orange and black bars alternating.

Hab. Burma, Siam, Malay Peninsula and Archipelago, and

S. China*.

* Annandale has shown (Ree. Ind. Mus. vii. p. 90, Feb. 1912) that the occurrence

of this lizard in S, India is incorrect,

29%

430 DR. M. A. SMITH ON REPTILES AND

Var. ANNAMENSIS, nov,

19-26 femoral pores.

14-22, usually 17-20, post-tibial scales.

Flanks with black and white bars alternating.

Hab. Coast of S. Anna.

Specimens examined.

Forma typica :-—

Femoral pores. Tibial scales.

SEIN ORNL ne URIS SEES RIT AY ce 15—15 4)

SR Me aie con ORR: SK LASS a 16—18 10

soca Eee ae NUE EN) Ree ES ae i -—alel 9

Remamee raite iyi ie ovo 34a seat aa RN e 15— 16 9

erally ies eos. nak eae eee 20—20 9

Salou wBacania-ci.). -ee eee es 17—17 9—10

- MMP READE ie. 3) ai ah 16==h8 10

Momascerimy «2! ...c.eem ane: 16 12

MMC UE ao ea oc iets « perme locus 15 9

TBAUTES AUER, Seeeees MMMM SSIES AS 20—20 13

ROHS OOY OR RRMA RERIAL LSA) Voy cl 16—16 9

Sie: (ECO) MISEDTOUT) ab eneransnsouae 16—17 8

ar udsol lak): 0h... .eeeikornsecee 16—17 9

3 iad) Us ea nr Pare 16—16 10

Ie Bayo ake cee eRe Weaeee Be Ny tle 16—16 9

Chiara sac un aoe een ewaeid ck ROE 9

PPS Reenter esl) SALA LEG sn Ale eG al

Var. annamensis :—

Femoral pores. Tibial scales.

ZAG San LouiakChamngserecn see SB HO—AS 16

2469. ,, pias actin bees 3. 24-24 16

2470. ,, mis HOltcRO ASAI of Q. 24—25 18

DA Mls o> ig, oi pl rina eae Q. 21—22 17

2472. ,, Sli Rae ase A 22—21 19

2473. ,, ast THR TSAR 3. 23—24 20

2474. ,, poly asta RES 21—22 22

DAT Oe 55 sis Set cee ela 3. 24--24 16

DAE i siege Meta NA ra 20—20 18

DAES ae ie. Ea) Qa niN 3. 24—24 I)

DAO te. Pei aa 3. 23—24 17,

2482. ,, He UA a eee 20-—21 14

CCC Maman k leaiiat na ml 4 Vila) 17

2481. Cap St. Jacques ...... 19—20 12

ZAS0 0 0 3s SPORE Ya Bate 22-—29 18

Types from Tour Cham. Type series, Author’s Nos. 2473,

2475, 2480, 2481, 2482, and 2483,

BATRACHIANS FROM SOUTHERN ANNAM. 431

LyGOsoOMA STELLATUM Bler.

Bouleng., Rept. Malay Pen. p. 87 (1912).

1 ex., Dalat. Before known only from two specimens, the

types in the British Museum, from the Larut Hills, Perak.

My example differs in that the prefrontals just miss contact, and

the black spots on the neck and shoulders are arranged to form a

broad vertebral band.

LyGosoMA CORPULENTUM, sp. 0.

Section Riopa. Distance between end of snout and arm twice

in distance between axil and groin. Limbs well developed,

short, pentadactyl, widely separated when adpressed. Snout

obtuse, eyelids scaly, supranasals in contact behind rostral ;

frontonasals forming a good suture with frontal; prefrontals

small; frontal broader than the supraocular, longer than the

frontoparietal and interparietal together; parietals in suture

behind the interparietal; 4 supraoculars; 2 loreals, posterior

longest; 7 supralabials, 6th subocular; temporals small and

scale-like ; ear-opening small, subcireular, about half the size of

the eye-opening, without projecting lobules ; a large azygos post-

mental; 36 smooth scales round the middle of the body;

preanals slightly enlarged; digits short, compressed, 4th toe a

little longer than the 3rd, 12-13 keeled lamella inferiorly.

Head and body, 170; tail, 150 mm.

Colour in life. Light chocolate-brown above, mingled with

yellowish on the flanks; lips, sides of neck, and throat yellow.

Belly brownish-white. Labial shields edged black.

A single specimen obtained at Dalat. Author’s No. 2128.

Closely allied to LZ. bamfyidii Bartlett, from Borneo, Sumatra,

and the Malay Peninsula, from which it differs in the absence of

lobules to the ear-opening, number of scales round the body,

colour of the head, and size.

DIBAMUS MONTANUS, sp. n. (Text-fig. 2.)

Snout covered with 7 more or less complete shields; a high,

narrow rostral, well visible above ; a pair of prefrontals, separated

Text-figure 2.

Dibamus montanis. Side and upper view of head.

from the lst labial by a suture running backwards from the

nostril, but fused with the labial anteriorly, the nostril thus

432 DR. M, A. SMITH ON REPTILES AND

lying between the Ist labial and prefrontal; a long 2nd labial*.

Other head-shields as in D. argenteus Taylor and D. novee-guinece

D. & B., viz. an enlarged frontal and a larger interparietal, an

ocular and an enlarged scale behind the 2nd “biel on either side.

Mental narrow, trapezoid, with a pair of long infralabials.

24—26 scales round the middle of the body, scales imbricate

and subequal. Preanals enlarged. Light chocolate-brown above,

paler below.

Types, ¢& 2. Author’s Nos. 4864, 4865. From Le Bosquet.

Measurements of specimens in mm.

Le Bosquet.

Total Diameter,

length. Tail. mid-body.

CO ABCD MUAMEEL, TIN? Orie JOBLAD 23 5

OIPAS CAIN JOLT OURO Od OM, 145 21 5

Daban

Die ZO ODM cir iced situ lout com te 112 2 3

@ MUZOOS taste. alas eel! 115 20. 3

OnyOsOowel + .2 2ahiwlaledtinant i) 11 2°5

Op ROOOO. BL tet. clei tee 86 16 2°D

Osho SOmaer canons ahinse. hls. 65 12 2

In the length of the tail (which from this table appears to be

compar: atively longer-in the young than in the adult) D. montanus

resembles D. a7¢ genteus from the Philippines. From 1 also, as well

as from D. nove-guinee, it differs in the greater differentiation

of the shields of the snout.

RANA MILLETI, sp. n. (Pl. II. fig. 2.)

Vomerine teeth in oblique series, commencing from the

anterior borders of the choane and extending beyond their

posterior borders, the distance between them equal to their

distance from the choane. Head a little longer than broad,

snout obtusely pointed, projecting beyond the mouth, longer than

the eye; canthus rostralis distinct; loreal region slightly oblique,

strongly concave; nostril distinctly nearer the tip of the snout

than the eye; distance between the nostrils greater than the

interorbital width, which is equal to or a pactle greater than

the upper eyelid; tympanum very distinct, $—-? 3 the diameter of

the eye, and 91-3 times its distance from the. ane.

Fingers rather long, Ist longer than 2nd; tips with small but

distinct dises, which may bear a feeble groove separating the

upper from the lower surfaces; subarticular tubercles large and

prominent; discs of the toes larger than those of the pasos and

with a distinct groove separating the surfaces; toes 4+ webbed ;

outer metatarsals separated nearly to the base; subarticular

* In the two adult examples from Le Bosquet (1200 metres) the shields are es

described, but in five juveniles from Daban (200 metres) the sutures are feebly

evident or entirely absent.

BATRACHIANS FROM SOUTHERN ANNAM. 432

tubercles prominent; no tarsal fold; inner metatarsal tubercle

23-3 times in length of inner toe; a small, prominent rounded

outer tubercle; tibio-tarsal articulation reaching to the tip of

the snout or not quite so far; tibia 17-123 in distance from snout

to vent; heels overlap when the limbs are folded at right angles

to the body.

Skin of the back finely granular; a prominent and fairly broad

dorso-lateral fold from the eye to the hip.

Yellowish-brown or greyish-brown above, sometimes with

indistinct darker markings; sides of the head dark brown ; limbs

with indistinct dark bars; below yellowish.

Males without vocal sacs, with a large, flat, humeral gland, and

a small pad on the first finger.

Allied to R. adenopleura Bier., from Formosa, from which it

differs in the more slender habit, less extensive webbing to the

feet, the web not reaching the discs of any .of the toes, and in

colouration.

Eggs pigmented, the vitelline sphere measuring 2mm. in

diameter.

Type locality, Dalat. Numerous specimens were obtained in

the type locality, from the Langbian peaks, and from Dran.

T have named this frog after Monsieur Millet, Conservator of

Forests to the French Government, who gaye us every help

possible while travelling upon the Plateau.

Measurements of type series in mm.

Author’s Nos. 5119 5128 5129 2571 2609 2602 4818 5103 5107 5176

45 46 37 36 36 37 39

Snout tovent.. 48 47 47

Heade-es ieee ako i7/ 16 16 17 145 14 14. 14 14,

Width of head 15 16 15 15 16 13 ors IBA BMS) alt

SHOU sscoseco 7 7 7 7 8 6 6 6 6 6

1 DWI tanettecee eee a) 5 5 5 5 45 45 A A 4

Interorbital ... 45 4& 4 5 5 om sa 98 4, 4,

Tympanum... 4 4. 4, 4 4 5 so era) 4 3°5

Armies. F289 27 28 27 27 22 21 23 22 24,

Mies HARI 185 79 79 81 81 64 64, 62 63 67

A MISVEW Se beard be al an 20, 25 27 27 26 20 20 20 20 21

ootwsseueee ss ae 25 25 26 26 20 20 20 20 21

2 9 2 2 a 3 3 3 3 3

Dalat. Dalat. Dalat. Dran. Dalat. Dalat. Dran. Dran. Dran. Lang

Bian.

Rana NigRoyirrara Blyth.

Bouleng., Rec. Ind. Mus. xx. p. 144, June 1920.

Boulenger’s Monograph on the Asiatic Rane omits any state-

ment of localities, or measurements of specimens of this species.

It is known from Southern Burma to the Man Son Mts.,

Tonkin. On the bills in Northern Siam it is very common, and

T obtained specimens on the plateau at Sui Kat, Dran, and Dalat.

Boulenger further describes the male as haying internal vocal

vesicles, but I have many specimens in which the skin of the

434 _ DR. M. A. SMITH ON REPTILES AND

throat is pigmented, and sufticiently differentiated, to entitle

the sacs to be called external. ;

In general appearance it. nigrovittata very closely resembles

R. mortensent from §.K. Siam. A. mortensent, however, has no

groove at all to the finger discs, grows to a larger size, and the

male has internal voeal sacs. Adult males cannot be confused,

but immature specimens or females could be mistaken. &. mor-

terseni appears to be confined to the Island of Chang. ‘The

specimens identified by Boulenger as having been obtained in the

Karin Hills (N. Sian), Monograph, No.5 & 6, p. 136, should, I feel

sure, be referred to negrovittata. he tadpole also, as originally

described by me under nigrovittata, should stand as correct.

I give measurements of some specimens in the British Museum

which I have examined.

a ep On a a

Be oer ueeaues a2 48 (alae

rc - A——_ cC_—™ Cc

Snout to vent... 49 50 49 52 42 44 54 46 56 49 42

TER OY a souddaceycadaen LO Wy Mee UB yeh AWG 0 TUS UG) TS} ly AG

Width of head...... WS USP VAG RS cep WR TB) URS a US} yp AWG)

SMOUi Meee en eins 8 7 8 6 RS) 2 Gis) Zi 6

JOKE) | Gas asdoneansbacadl Oe. Gray Gey. G ANDI | 655 6 5 6 5 45

Tympanum ......... 4 4 4 AS 25 A 35 4 35 3'5

PNRM, anGay bccn encod! PAS eA OAS) WOR OIA AN oO) UBS 8 Bi) 30 3=—- 28

Weer tia hee een 2a ICA lea SON as oli Onn Oamae Clima id annoo 84 75

Tabi aes Seas eee OX reba, PUP OS Oa) Day Oe. bai 25 = 24

Hootees eres pee PAS Oey Oy OA ON ORT ar Oe 19) 25 24

Rana SAvuTERI Boulenger.

Rec. Ind. Mus. xx. p. 143, June 1920.

Var. JouNsI, nov.* (PI. II. fig. 1.)

Eleven specimens of a frog from the Plateau differ sufficiently

from R. sauteri Boulenger, from Formosa, to be entitled to

vacial distinction. I have compared them with types in the

British Museum, and separate them on the following grounds :—

More pointed snout, longer leg, tibio-tarsal articulation to well

beyond the snout, very prominent glandular dorso-lateral fold,

and smaller size.

In other points also, Boulenger’s description, drawn up from

four females, shows small variations, and I therefore describe my

specimens in detail. ‘

Vomerine teeth in oblique groups between the choanz and

extending beyond their posterior borders, as far from the choane

as from each other, or a little farther.

Head as long as, or a little longer than broad, depressed ;

snout obtusely pointed, prejecting beyond the mouth, longer than

* Named after Mr. Johns, British Consul in Saigon at the time of our visit, and

who was cf the greatest assistance to us in arranging many details of our expedition.

BATRACHIANS FROM SOUTHERN ANNAM. 435

the eye; distance between the nostrils considerably greater than

the interorbital width, ue is equal to the upper eyelid ;

tympanum very distinct, 2-2 the diameter of the eye, which is

3-4 times as long as its distance from the latter.

Fingers moder: ate, the tips swollen into very small dises which

do not bear a groove; lst longer than 2nd; subarticular tubercles

large and prominent.

Hind limb long and slender, the tibio-tarsal articulation

reaching far beyond the snout; heels strongly Ore lang when

the limbs are folded at right angles to the body; tibia 5-6 times

as long as broad, 14-14 times in length from snout to vent, longer

than the foot. ‘Toes with small dises, more developed than those

of the fingers, and bearing a groove separating the upper from

the lower surfaces; web reaching to the dise of the 5th toe and

to the 5rd on its outer side, last two phalanges of 4th free.

Outer metatarsals separated nearly to their base; no tarsal

fold; subarticular tubercles aenely developed ; inner metatarsal

pence oval, prominent, 3-4 length of inner toe; a very small

round, distinet tubercle at the base of the 4th toe.

Skin smooth or very finely granulate, a few enlarged scattered

tubercles on the back, a /A-shaped glandular fold between the

shoulders, and short oblique folds crossing the thighs and tibie ;

a narrow, prominent, dorso-lateral fold from the eye to the hip,

and two short folds from behind the tympanum enclesing a

triangular black patch.

Greyish-brown above, uniform or faintly mottled with darker;

a well-defined, rich, dark brown patch from the eye to the

shoulder, enclosing the tympanum; snout below the canthus

rostralis darkish ; limbs with narrow dark cross-bars s; a brown

streak along the back of the arm and another along the front of

the tibia. Below whitish, the chest and throat speckled with

ervey.

Males with internal vocal sacs, and a brown nuptial prominence

on the Ist finger.

Types from Sui Kat. Type series, Author’s Nos. 2638, 2639,

2640, 2641, 2644, 2657.

Measurements in min.

Author’s Nos. 5031 2657 2642 2639 2640 2641 5087 2638 2644 2645 2457

Snont to vent. 45 43 42 43 40 50 43 42 42 41 40

Head) ..eee 15 14 14 14 13 16 15 15 14, 13 13

Width of head 14 145 145 14 12 15 14 14 14 13 12°

NSHAVOUNUE bocuooaes 6 65 6 6 a5 a 65 65 65 6 6

Bivelee trees 4°5 45 45 45 4, 55 45 45 4°5 4

Interorbital.. 3:5 3 3 3 3 4 35 3 Simo 3

Tympanum.. 35 35 3 3 NO oon) TOONS 30) 3 3

AMIE eee, WO 25 25 25 25 29 26 24 25 23 22

Leg 87 81 84 82 Mie LOO 81 78 81 77 75

Otloy aenanatce 28 27 27 27 26 34 28 2 27 26 25

Foot 26 25 25 25 23 30 25 2 25 23 22

EW oe gordo Sodas noma Wenn

436 DR. M, Ae SMITH ON REPTILES AND

RANA MONTIVAGA, sp. uu. (PI. I. fig. 2.)

Vomerine teeth in short oblique groups between the choane,

and extending beyond their posterior margins. Head broader

than long; snout rounded, scarcely projecting beyond the mouth,

longer than the eye; canthus rostralis strong; loreal region

oblique, deeply concave; nostril a little nearer the tip of the

snout than the eye; distance between the nostrils greater than

the interorbital width, which is equal to the width of the upper

eyelid; tympanum very distinct, about half the diameter of the

eye, 14-23 times its distance from the latter.

Fingers moderate, terminating in small dises which bear a

groove separating the upper from the lower surface; Ist a little

longer than the 2nd; subarticular tubercles large and prominent.

Hind limb long and shapely; tibio-tarsal articulation reaching to

far beyond the snout; heels strongly overlapping when the limbs

ave folded at right angles to the body; tibia 14-14% times in

length of head and body, much longer than the foot. Toes with

well-developed dises, which are larger than those of the fingers and

bear a strongly-marked groove; web reaching to, and including

a portion of the dises of all the toes, but in the ease of the 4th it

is continued on as a narrow fringe from the 2nd phalanx; outer

metatarsals separated nearly to the base, subarticular tubercles

large and prominent; a feeble tarsal fold; inner metatarsal

tubercle oval, 24 times in length of inner toe; no outer tubercle.

Skin above granular, with larger tubercles; a narrow, promi-

nent, dorso-lateral fold from the upper eyelid to the hip, strongest

in front, sometimes broken up. Lower parts smooth.

Yellowish-brown or greyish-brown above, usually with indis-

tinct blackish markings; sides of the head darker ; limbs with

dark cross-bands. Below yellowish-white, usually powdered with

brown on the throat and chest.

Males smaller than females, with internal vocal sacs.

Types from Dalat, Langbian Plateau, at 1500 metres.

Allied to &. varians Blgr., from Celebes and the Philippine

Islands; from which it differs in the stouter habit, broader

Measurements of type series in mm.

Author’s Nos. ... 5124 5123 5126 2572 5013 4826 2529 25380 4829 65022

Snout to vent... 48 48 48 46 48 75 72 71 66 58

fend) aioe gece lh 16 18 16 17 25 23 QA 22 21

Width of head... 18 18 75) NG 17 26 25 26 24 20°5

Suout ESE GXO, is, 7 7 Uf 10 11 ih, 10

Bye Boa aus Gio Gio, 46 6 6 8 8 9 8 75

Interorbital ...... 5 5: 5 4, 4 6 6 7 6

Tympanum....... A A 4 3 3 Aco) Loon sO 5 45

ATO te 3 see O28 30 30 28 3L 43 44 47 40 34

Deon eae) 298 94, 95 93 96) 140 Sos IS Bails Simmel

Uiaibiawse fe oe 32 32 30 31 44, 45 45 42 39

Woot pate eee OO) 2 29 27 Qi. 37 AL 37 34

BATRACHIANS FROM SOUTHERN ANNAM. 437

head, shorter*and more rounded snout, and absence of external

mnetatarsal tubercles.

‘This frog was common on the Plateau, and numerous specimens

were obtained at between 1500 and 2000 metres elevation.

RANA GRAMINEA Boulenger.

P. Z.S. 1899, p. 958, pl. Ixvii. fig. 1; id., Rec. Ind. Mus. xx.

p. 204, June 1920.

Boulengew’s description was drawn up from male specimens

only. A fine series from the Plateau shows that the females

ave much larger than the males, some of them being twice as

large. The specimens differ from the types in the snout being

longer than the eye, and in the nostrils being distinctly nearer

the tip of the snout than the eye. Females have a propor-

tionately smaller tympanum than males.

Colour. Above bright green to dark clive, or greyish-brown,

uniform or with large darkish spots. Below white, uniform or

powdered with grey. Females usually light greyish-brown above,

seldom bright green, and conspicuously spotted on the back and

limbs.

This frog was common on the Plateau at all elevations above

1500 metres.

Measurements of &. graninea in mm.

Author’s Nos.... 2515 2513 2518 2526 2647 2506 2509 2510 2648 2649

Snout to vent... 58 53 56 45 40 105 101 89 79 72

Fleadiioye. .ueik e220 20 20 17 15 35 34 29 27 25

Widthof head... 20 18 18 16 145 35 35 32 27 25

SuOwiy sees 10 9 8) a 65 17 16 14: 13 11

Biyey ssdissez et2nt 9 o 7 51) oxen al 11 105 «8 8

Intevorbital ...... 45 45 45 ore) 9 8 8 6

Tympanum.. ... 45 A 45 Sha de 6 5 45 4,

ALIS essa atch OO) 34 Br 3l 30 66 66 60 52 49

Ab Css Pee ayeacoetasode oe 94 94 85 76 190 190 168 150 150

AMMO Goaaooetcoroaas 224 32 @) 29 65 65 59 52 53

3 25

TOC loon secasseeones el 2 27 24, 22 54 54 46 46 deh

MicrouHyLaA pPicra Schlegel.

Verh. Nat. Ges. Basel, xiii, 1901, p. 151, fig.

Known from a single specimen in the Basel Museum.

Dr. Roux has kindly compared one of my specimens with it,

and considers them to be identical. In general characters

M. picta agrees so closely with I. rubra from India that, were it

not for the distinctive colouration of the two forms and their

seographical distribution, it would be difficult to separate them.

Comparing my specimens with examples of J/. rubra in the

British Museum, I find the following differences :—Snout (of

pict) a little more obtuse, tibio-tarsal articulation reaching to

posterior border of eye, web of toes slightly fuller.

438 DR. M. A. SMITH ON REPTILES AND

Colour. Greyish or yellowish above, with a large dark white-

edged mark on the back, commencing between the eyes and

diverging about the middle into two arms which 1un to the

groin. Dark markings along the sides parallel to it, and often a

second A over the sacral region, Limbs with dark bars. Below

yellowish, the males with blackened throats.

M. picta is found in Cochin China (type locality unknown),

M. rubra in India and Assam.

I found this little frog one night in Aprii at Cap St. Jacques,

where if was breeding in the ditches beside the road, not far from

the sea. We were attracted by the strident voices of the males,

and without difficulty captured large numbers.

Buro GALeatus Giinther.

Rept. Brit. India, p. 421 (1864).

B. galeatus was described by Gtinther from a single specimen

obtained by Mouhot in Cambodia. A series of eight specimens

obtained at Dran in April enables me to enlarge his original

diagnosis, and I take this opportunity to re-describe the species.

Canthus rostralis with strong bony ridges, which are continued

backwards more or less distinctly over the supraorbital and

parietal regions. A thick, arched, elevated orbito-tympanic

ridge, separated by a slight depression from the parotid gland,

which is about as long as the bony ridge; snout short, blunt ;

tympanum very distinct, two-thirds diameter of eye. Ist finger

distinctly longer than 2nd; toes one-third to nearly half webbed ;

two well-marked metatarsal tubercles, the inner nearly twice as

large as the outer; no tarsal fold; tibio-tarsal articulation reach-

ing to tympanum in male, not so far in female. Upper parts with

prominent warts, which become spiny on the flanks; lower parts

coarsely granular; parotids prominent, elongate, about as long

as the orbito-tympanic ridge.

Reddish or greyish-brown above, with dark marblings on the

back, the limbs with dark bars; bars upon the lips. Dirty

yellow or whitish below, more or less distinctly spotted with

black.

Males considerably smaller than females and with black nuptial

asperities on the inner two fingers.

Measurements in min.

AmbhouisyNiosy esate e426 2497 2428 2492 2132 2592 Type

Snowbetonventiencascece | Meco 70 61 62 50 43 63

Snout to ant. border 2?

oftympanum § 23 WW WW 17 is a te

Width of head ......... 35 27 25 24 18 15 27

INOW INGA. Bousceckonedeon oO 4A 40 45 36 33 42,

Jalnnol Wino ccotcobcelsondes 99 81 72 83 67 59 82

NPE oe one aa nae 3o2 26 24 Q7 22 19 25

AH YON GARE NAMteIA ears 32 25 23 25 20 18 24,

BATRACHIANS FROM SOUTHERN ANNAM, 439

MEGALOPHRYS INTERMEDIUS, sp. n.

A form intermediate between Jf. carinense and M. fee,

differing from the latter in the web between the toes, and from

the former in the more posterior position of the vomerine teeth,

and from both in several other small points.

Tongue feebly nicked behind. Vomerine teeth present in two

widely-separated groups J behind the level of the choane.

Head large and depressed, 1{—2 times as broad as long; snout

rounded, hardly as long as the eye, not projecting beyond the

lower jaw; canthus rostralis very distinct; loreal region slightly

oblique, feebly concave; nostril equidistant from the eye and the

snout; interorbital space 13 times to nearly twice as broad as

the upper eyelid; tympanum hidden. Fingers short, with feebly

swollen tips, Ist and 2nd about two-thirds length of rd; no sub-

articular tubercles; metacarpal tubercles very indistinct ; toes

with feebly swollen tips one-third to one-half webbed, the web

extending as a fringe along either side ; no subarticular fiber cles ;

a large oval, flat, inner metatarsal tubercle; tibio-tarsal articu-

Jation reaching to nearly, or quite, the commissure of the jaw ;

tibia one-third to two-fifths in length of head and body; foot

longer than the head. Upper eyelid with conical tubercles, one

of which is enlarged to form a short horn; an oblique glandular

fold on the back parallel with the supratemporal fold usually

present ; more or less distinct oblique folds across the limbs,

throat finely granulate, belly nearly smooth.

Colour in life. Above bronze or dark coppery, the head and

fore part of the body, outside the dorsal glandular fold, usually

lighter. Back usually with indistinct light and dark markings ;

arms and legs with dark cross-bands; a dark patch over the

region of the tympanum and dark bars below the eye. Below

dink brown to bronze, paler on the belly. Dorsal fold and

tubercles on the flanks black on their inferior aspect, light

Measurements of type series in mm.

Author’s Nos....... 2070 2073 2075 2078 2067 2086 AN 2085 2076 2084 2083

Snout to vent ...... Oy Oe NOs) OE SHE By 8B) TOS IB GRA 283

Head (tolocciput) En 2ip eco) One o ome Zonm2o. 28h Ze 18s 4:

Width of head...... ByO) aS) aS) lyf NS ANS Als) fal, a Yo)

encth of Snouti sea) selZe oe Sela SiS Zs Re ele oS IT 9 65

Diam. of eye......... 9 9 9 9 8 9 SK) fel 5 5

IbMRAAOS GyhNA s., die} 8} 1B} TI, NB} aI ley ale aI 9 a

Hore lim Devereeeeeee: 52P ble 60) Ashe 54m boi bah eb8 9 50) 32) 223

la hos eeerneeseeaupacns Disy AON ML MN OI Bl OY ire GIT alte} 5 bee

Hind limb } we M4 WG. HO Tides ies Iie) We 116) Loo) 69) 60

(to artic.)

MT abtes ha teeteanes SG 9 SOM oO" HtoOMmTSONNNOUL move oz 0 wen lb

HOOtS antatitenens B/S) SiS SI. piel BE Sis I SPY BS alley

440 ON REPTILES AND BATRACHIANS FROM SOUTHERN ANNAM.

superiorly, Young with paler (yeliowish) head and shoulders,

and with the markings more clearly defined.

21 specimens examined.

This fine Megelophrys was common on the Plateau above

1500 metres, and the loud, harsh croakings of the males could be

heard at all times of the day and night. It was by means of

their call that most of them were discovered and finally tracked

down to their hiding-place in some deep crevice between the

rocks or boulders of the streams in which they lived.

MEGALOPHRYS HASSELTI 'schudi.

Bouleng., Fauna Malay Pen. p. 282 (1912).

Var. PULLUS, nov.

Differs from the typical form in the longer and more pvro-

minent metatarsal tubercle (at least twice as long as broad),

longer leg (to tympanum), smaller size, and in colouration.

Toes 3 webbed.

Dark grey above, sometimes with indistinct darker markings.

Limbs with alternate dark and light ecross-bars. Below whitish

or brownish. Upper half of iris (in life) scarlet.

Type locality, Arbre Broyé. Also found at Camly.

Altogether 20 specimens were obtained, which I have been

able to compare with some 20 examples of the typical form from

Siam and the Malay Peninsula.

Measurements of type series in mim.

Author’s Nios:......... 2093 2101 2108 2091 2103 2090 2105 5192

Snout to vent ......... 49 4A 52 45 4A, 45 4A. 49

dB oer0 LON a Sey eae ie il7/ 15 18 15 15 16 15 17

Width of head ...... 21 19 22 20 19 19 18 21

Intterorbitaleecces ee O 5 7 55 5 oy G5) bid

PAIN) he ee eee eee ea 30 27 38 27 29 28 29 34

Hand 12 10 13 10 10 10 10 12

Leg SUR aeRO O 50 62 51 52 51 50 56

MDa Ta) seas iepaaeee sen co epi Lts 15 18 15 15 15 15 17

TOOL e Areeeene aden ete, oli) 16 18 15 15 15) AUS If

EXPLANATION OF THE PLATES.

Puate I.

Fig.1. Fimbrios klossi.

2. Rana montivaga. (Nat. size.)

Prate II.

Fig. 1. Rana sauteri var. johnsi. (Nat. size.)

2, Rana milleti. (Nat. size.)

ON THE PASSENGER-PIGEON OF THE UNITED STATES. 44]

EXHIBITIONS AND NOTICES.

March 8th, 1921.

Sir 8S. F. Harmer, K.B.E., F.R.S., Vice-President,

in the Chair.

In the absence of the donor, Dr. R. W. Suurrtpt, C.M.Z.S.,

three photographs of the last of the Passenger- Pigeons (/ctopistes-

nugratorius) presented by him to the Society, were exhibited b

Dr. P. Coatmers Mircuetn, C.B.E., F.R.S., who read the follow-

ing notes from Dr. Shufeldt :—‘‘ The specimen was a female which

Text-figure 1.

Passenger-Pigeon (Hetopistes migratorius).

died in the Zoological Gardens of Cincinnati in September 1913,

where she had lived in captivity for a number of years with

442 ‘THE SECRETARY ON ADDITIONS TO THE MENAGERIE.

others of the same species. From time to time these others died,

until only the female remained—the sole living representative of

the endless millions that originally were to be found in the

United States.

““ When the dead specimen came to the United States National

Museum, I posed it in three different positions, that anterior,

posterior, and lateral views of the body might be photographed.

New York State, when I was a boy, had annually in its avifauna

its millions of these pigeons (Hctopistes migratorius), and they.

were slaughtered there every season, as they were all over the

country and in all the States.”

Mr. R. I. Pocock, F.R.S., F.Z.S., exhibited, and made remarks

upon, the skull of a Sumatran Tiger.

On behalf of Messrs. Gerrard & Sons, Mr. R. IL. Pocock

exhibited, and made remarks upon, a Cheetah skin from Tan-

ganyika Territory, showing an interesting modification in the

arrangement of the markings on various parts of the coat.

March 22nd, 1921.

Sir S. F. Harmer, K.B.E., F.R.S., Vice-President,

in the Chair.

The Secrerary read the following Report on the Additions to

the Society's Menagerie during the month of February 1921 :—

The registered additions to the Society's Menagerie during

the month of February were 143 in number. Of these 50 were

acquired by presentation, 53 were deposited, 36 were purchased,

and 4 were born in the Menagerie.

The following may be specially mentioned :—

1 Nylghau (Boselaphus tragocamelus), 3 , from India, deposited

by H.G. The Duke of Bedford, K.G., F.R.S., on February 24th.

2 Capybaras (Hydrocherus hydrocherus), from 8. America,

deposited on February 5th.

1 Collared Peceary (Vayassu tajacu), from 8S. America, pur-

chased on February 10th.

2 Pied Starlings (Spreo bicolor), from South Africa, new to the

Collection, purchased on February 15th.

A collection of 20 Pheasants and 17 Peafowl, presented on

February 21st by the Hon. Nathaniel Charles Rothschild, F.Z.S.

THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 443

April 5th, 1921.

Prof. E. W. MacBrips, D.Se., LL.D., F.R.S., Vice-President,

in the Chair.

Dr. P. Coaumers Mircneyt, C.B.H., F.R.S., Secretary of the

Society, exhibited, and made remarks upon, a series of lantern

shdes of a Baby Chimpanzee born in the New York Zoological

Park.

Sir S. F. Harmer, K.B.E., F.R.S., exhibited, and made remarks

upon, a photograph of Elephant Twins.

Mr. E.G. Boutencsr, F.Z.S., exhibited, and made remarks

upon, the following Reptiles and Batrachians:—a young speci-

men of Testudo nigra, Monopeltis capensis, Siphonops annulatus,

and Phryniscus nigricans.

Miss L. E, Curesman, F.E.S., exhibited, and made remarks

upon, a nest of Anapha venata (Lepidoptera).

April 19th, 1921.

Sir 8S. F. Harmer, K.B.E., F.BS., Vice-President,

in the Chair.

' The Secretary read the following Report on the Additions to

the Society’s Menagerie during the month of March 1921 :—

The registered additions to the Society’s Menaterie during

fhe month of March were 276 in number. Of these 73 were

acquired by presentation, 34 were deposited, 153 were pur-

chased, 7 were received in exchange, and 9 were born in the

Menagerie.

» The following may be specially mentioned :—

3 Wapiti (Cervus canadensis occidentalis), 3¢ 2 2, from Van-

couver, presented by the Govt. of British Columbia through the

Game Preservation Board on March 30th.

1 Persian Gazelle (Gazella subgutturosa), 3, from Mesopo-

tamia, presented by Maj.-Gen. Sir Percy Cox, K.C.M.G. on

March 29th.

' 1 Bennett’s Gazelle (Gazella bennetti), 2 , from Feluja, Mesopo-

tamia, presented by Major Chadwick on March 29tb.

Proc. Zoou. Soc.—1921, No, XXX, 30

444 : ON THE BIRTH OF A POLAR BEAR CUB.

1 Black Wallaby (A/acropus walabatus), from New South Wales,

presented by H.R.H. The Prince of Wales, K.G.,on March 4th.

1 Searlet-breasted Sunbird (Cinnyris guituralis), from Durban,

South Africa, new to the Collection, presented by Harold Millar,

C.M.Z.S., on March 18th.

1 Southern Tree-Snake (Dryophis dispar), from India, new to

the Collection, presented by A. P. Kinloch, on March 26th.

Mr. R. H. Burne, M.A., F.Z.8., exhibited, and made remarks

upon, a series of mounted specimens of young Flat Fish, demon-

strating various stages in the transference of the left eye to the

right side of the head.

Dr. P. Cuatmers Mrircuext, C.B.E., F.R.S., read the following

extract from a letter he had received from Mr. E. H. Bean,

Director of the Washington Park, Milwaukee, Wis., U.S8.A.,

describing the successful rearing of a Polar Bear Cub :—‘t Our

cub was born December the 2nd, 1919. I am quite sure the

period of gestation was eight months. I removed the female

bear in September. She was placed in a smaller compartment of

our bear den, alone. The sleeping den is small, about 6x8 ft.

The outside enclosure was also small, with a bathing tank in

same. The mother bear bathed regularly up to about three

weeks before the cub was delivered. We attempted to give her

bedding prior to the birth of the cub, but she always put it out-

side and preferred the bare boards to sleep upon. She stopped

eating about three days before the cub came. About nine o’clock

in the morning, December the 2nd, I passed her cage, detected a

strange sound, which at first [ thought to be a stray cat, but

upon investigation, found to be coming from the polar bear cage.

It happened to be twelve below zero that morning, and the den

door faced the north, from which direction a very stiff wind was

blowing. The mother bear had her back to the opening, and it

seemed as though she had the baby cuddled up against her

abdomen, and covered it as best she could with her great paws,

I called the keeper, and he took a bale of straw on the rocks

above the den door and gradually dropped a few handfuls down.

I watched to see the effect it would have upon the mother bear.

She did not seem to be disturbed at all, just raised her head up,

looked around, and then settled back. We continued until we

had shaken up two bales of straw, pushing same well up into the

mouth of the den. During the night she took all of this straw

inside, and at intervals of a few days we continued until she had

seven bales of straw. It was very interesting to note her ability

to determine approaching storms. When a storm was on the

way, she would pack the opening from the inside, just leaving a

. ON SKINS OF THE AUSTRALIAN BUDGERIGAR. 445

small vent at the top. When mild weather was on, she would

take her paws and pat the straw down until the opening was

practically full size. She left the nest at the end of seven days

for food and drink, and continued to do so until along in April,

when she permitted the baby to come out. The nest was so con-

structed that it resembled a large wicker basket. For a few

weeks the baby would crawl out and look over the side, but

mother was always there to discourage any attempt on the baby’s

part to leave the nest.

“We have four adult polar bears that came to us direct from

Bergen, Norway, in 1912. I have accustomed them to what

might be considered a peculiar diet for this class of animals,

We feed them on bread, apples, carrots, boiled rice, dried stock-

fish, a mess of fresh fish seldom, and no meat whatever. They

get a pan of cod-liver oil now and then, grass of all kinds during

the summer months, also dog biscuits. The adult bears have

a sleeping den on the hillside 8 x 47 ft. deep and 7 high, and an

outside enclosure 50 x 60 ft., containing ledges, grottos, ete., also

a bathing tank 15 ft. wide, 30 ft. long, and 8 ft. 5 inches deep.

The bears bathe continuously, and are one of our main attractions

in the collection.

* We used every precaution not to disturb our mother bear.

The day that she was out for food I peeped into her nest, and

this baby was about as large as a full-grown guinea-pig, fully

furred, and with two little black eyes like beads. I cculd not

determine when it had opened its eyes. It is a wonderful cub,

only had one sick spell, and that lasted only for a few hours.”

Mr. C, Davies SHErpory, F.Z.S8., exhibited a coin of the Saka

Dynasty (Punjab Region) of King Arzes (Ayasa), showing the

so-called ‘“‘maneless lion” of Asia, about B.c. 30. This coin

belongs to the Bactrian Series, and is inscribed in Greek on one

side and in Pali on the other.

Mr. F. Martin Duncan, F.Z.S., exhibited, and made remarks

upon, a series of lantern-slides of a remarkable nest of the Wasp

( Vespa germanica),.

Mr. D. Seru-Smire exhibited a series of skins of the Australian

Budgerigar (Melopsittacus undulatus), showing the various colour-

varieties that have been produced in captivity. He remarked

that the species was apparently first imported alive in Europe by

John Gould in 1840, since when not only had it been freely

brought home, but it had bred in captivity so readily that now it

might be said to be a domesticated species, and, like other domes-

ticated species, certain colour-varieties had been produced. There

446 ON CHANGE OF HABITAT IN THE COMMON CRICKET.

was the well-established yellow variety, the rarer blue variety,

and a so-called olive variety, and recently a nearly pure white

bird (which was shown) had been bred by Mr. H. D. Astley.

These varieties were produced by the elimination or partial

elimination of the two primary pigments, yellow and black, which

were present in the normally-coloured green bird. The yellow

variety showed entire lack of black pigment, the blue variety was

entirely lacking in yellow pigment, the blue colour being pro-

duced by a combination of black pigment and prismatic feather

structure.

The olive variety Mr. Seth-Smith was unable to show, but it

appeared to be a case in which both yellow and black pigments

were weak, while the unique white specimen was an instance of

the elimination of both the black and yellow pigments.

Major KE. HE. Ausrey, D.S.0O., F.Z.8., drew attention to an

apparent change of habitat on the part of the Common Cricket

(Gryllus domezsticus), and described a recent visit he had paid to

a large refuse dump where he had observed the Crickets in con-

siderable numbers,

ZOOLOGICAL SOCIETY OF LONDON.

Tas Society was founded in 1826 by Sir Sramrorp Rarriss,

Mr, J. Sasine, Mr. N. A. Vieaors, 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.

WPatrow.

HIS MAJESTY THE KING.

COUNCIL. eco

HIS GRACK THE DUKE OF BEDFORD, K.G., E.R.S., President.

AxrreD H. Cocks, Ese., M.A. || Cor. Sim Henry McManon,

G.C.M.G.,, K:CaE.

KE. G. B. Meaps-Watpo, Esa.,

Vice-President.

Caarces Drummonp, Ese,

Treasurer.

ATED Lips Be. Oo 8 i. P. Cuanmers Mrrcnett, Esa.,

Tue Rieat Hon. rae Viscount CBE MA. DSc, iD.

Greve KeG. RC; E.RS., Secretary.

Srr Srpwey F, Harwer, K.B.E., || fae Eant or Onstow, O.B.E.

M.A., D.Sc. F.R.S., Vice- || Mason Atserr Pam, O.B.E.

President. Tae Lorp QurenyBoroucs.

Pror. James: P. Hirt, D.Sc., || 4 G Tue Duxsor Rurtanp, K.G.

F.R.S., Vice-President.

Masor Ricuarp 8S. Tayror.

A. Trevor-Bartysr, Ese., M.A.

Mayor tHE Lorp Atasrair || ayrpony H. Winerrexp, Esa.,

Rosurr Iynes-Krzr. Vice-President.

Witiram Huntsman, Esa.

Pror.ERnestW. MacBripe,D.Sc., || A. Siri Woopwarp, Esa.,

LL.D., F.B.S., Vice President. LL.D., F.R.S., Vice-President.

The Society consists of Fellows, and Honorary, Foreign, and

Corresponding Members, elected according to the By-Laws. It

carries out the objects of its foundation by means of its collection

of living animals, by its Library, and by its Scientific Publications.

The Office of the Society, Regent’s Park, N.W.8, where all com-

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 half-past

Five o’clock p.m.

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.LS., is the resident Super-

intendent and Curator of Mammals, Mr. D. Seth-Smith is Curator

ef 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-

caticns for anatomical material or facilities for work in the

Prosectorium should be addressed to Dr. R. T. Leiper, Director of

the Society’s Prosectorium.

TERMS FOR THE ADMISSION OF FELLOWS.

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

bution of £3, due on the Ist of January, and payable in advance,

or a Composition of £45 in lieu thereof; the whole payment,

including the Admission Fee, being £50.

No person can become a Frrtow until the Admission Fee and

first Annual Subscription have been paid, or the annual payments

have been compounded for.

Fertiows elected in November and December are not liable for

the Subscription for the vear in which they are elected.

PRIVILEGES OF FELLOWS.

Fettows haye Personal Admission to the Gardens upon signing

their names in the book at the entrance gate, and may introduce

Two Companions daily.

The Wire or Hussanp of a Futtow can exercise these privileges

in the absence of the Fellow.

Until further notice, Frttows will receive 40 undated Green

Cards, available on any Sunday or week-day up to the end of

February of the year following the year of issue, and 20 White

Cards available on any week-day up to the same date. Twenty

of the Green Cards may be exchanged for a book containing two

Orders for each Sunday in the year. Twenty White Cards may

be exchanged for a book of dated Week-day Orders, each Order

available for any day during the week except Sunday. Special

children’s tickets are no longer issued, but the Green and White

Cards are perforated, and each half is valid for a Child under twelve

years of age. It is particularly requested that Fellows will sign

every ticket before it goes out of their possession. Unsigned tickets

are not valid.

Frttows are not allowed to pass in friends on their written

order or on presentation of their visiting cards.

Frttows have the privilege of receiving the Society’s ordinary

Pubhecations issued during the year upon payment of the additional

Subscription of One Guinea. This Subscription is due upon the

Ist of January, and must be paid before the day of the Anniversary

Meeting, after which the privilege lapses. Fertiows are likewise

entitled to purchase these Publications at 25 per cent. less than

the price charged to the public. A further reduction of 25 per

cent. is also made upon all purchases of Publications issued prior

to 1881, if above the value of Five Pounds.

Frtiows also have the privilege of subscribing to the Annual

Volume of ‘The Zoological Record,’ which gives a list of the Works

and Publications relating to Zoology in each year, for the sum of

One Pound Ten Shillings. Separate divisions of volumes 39 onwards

can also be supphed. Full particulars of these publications can

be had on application to the Secretary.

FrLtows may obtain a TRransreraBLe Ivory Ticker admitting

two persons, available throughout the whole period of Fellowship,

on payment of Ten Pounds in one sum. A second similar ticket

may be obtained on payment of a further sum of Twenty Pounds.

Any Fstiow who intends to be absent from the United Kingdom

during the space of at least one year, may, upon giving to the

Secretary notice in writing, have his or her name placed upon the

“ dormant list,” and will then be called upon to pay an annual

subscription of £1 only during such absence, but after three years

must make a further application to be retained on that list.

Any Frtiow, having paid all fees due to the Society, is at liberty

to withdraw his or her name upon giving notice in writing to the

Secretary.

Ladies or Gentlemen wishing to become Fellows of the Society

are requested to communicate with ‘“‘ The Secretary.”

P. CHALMERS MITCHELL,

Secretary

Regent’s Park, London, N.W. 8.

June, 1921.

MEETINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON

FOR

SCIENTIFIC BUSINESS.

1921

Atioaasinysn’, DUNN saschecnvose he

‘5 OCTOBER ......... 18.

ih NOVEMBER ...... 8 and 22.

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

ZVOROGICAES SOCIETY OFF KON DOIN:

LIST OF PUBLICATIONS.

Tue scientific publications of the Zoological Society of London

are of two kinds—“ Proceedings,” published in an octavo

form, and “ Transactions,” in quarto.

According to the present arrangements, the “‘ Proceedings”

contain not only notices of all business transacted at the scien-

tific meetings, but also all the papers read at such meetings

and recommended to be published in the “Proceedings ” by

the Committee of Publication. A large number of coloured

plates and engravings are issued in the volumes of the

“ Proceedings,” to illustrate the new or otherwise remark-

able species of animals described therein. Amongst such

illustrations, figures of the new or rare species acquired in a

living state for the Society’s Gardens are often given.

The “ Proceedings”’ for each year are issued in four parts,

paged consecutively, during the months of March, June,

September, and December. From January 1901 they have

been issued as two half-yearly volumes, indexed separately.

An “ Abstract of the Proceedings’ is published by the

Society on the Tuesday following the date of the Scieutific

Meeting to whichit refers. It is issued along with the “ Pro-

ceedings,” free of extra charge, to all Fellows who subscribe to

the Publications, but it may be obtained on the day of publi-

cation at the price of Sixpence, or, if desired, sent post free

for the sum of Six Shillings per annum, payable in advance.

The ‘‘ Transactions” contain such of the communications

made to the Scientific Meetings of the Society as,on account of

the nature of the plates required to illustrate them, are better

adapted for publication in the quarto form. They are issued

at irregular intervals.

Fellows and Corresponding Members, upon payment of

a Subscription of One Guinea before the day of the Anni-

versary Meeting, are entitled to receive the Society’s

Publications for the year. They are likewise entitled to

purchase the Publications of the Society at 25 per cent. less

than the price charged to the Public. A further reduction

of 25 per cent. is made upon purchases of Publications

issued prior to 1881, if they exceed the value of Five

Pounds.

Fellows also have the privilege of subscribing to the

Zoological Record for a sum of One Pound Ten Shillings

(which includes cost of delivery), payable on the Ist of July

in each year; but this privilege is forfeited unless the

subscription be paid defore the 1st of December following.

The following is a complete list of the publications of the

Society already issued.

TRANSACTIONS OF THE ZOOLOGICAL SOCIETY OF LONDON.

4to. 20 vols. and Index. Price to Price to the

Fellows. Public.

Vols. I.-IV. (out of print).

Vol.) WV. containing, 67, /Plates)) @iS62=66) i bas ee Ono mao

vile! ea Gye eye | 51) TO

Ope U9. HEN 6 (1875277) 0.) oT eM eee mo nmn

Index. Vols. Xie seiearerirety es (1833=79) ey NO)! G7 Gye hn Ons OmanO

Vol. XT, containing, 97 Plates’ .. (880-85) 2.) 9/12) 0). 52 allio

eerie WeXeleLe. 65 pil (eke CL SSE=90)\ ie ete o ne Ome (ocak @)

son OU 5 62 yi oo (WEI) 5255 °O GhiG Sie

eM EXGNY a ns 52 my oo. USPS) 4 BIG 8. 714 0

OVE Se. FST 1908): 5 08 0 Oem ame

Vag OO Wes oe 11908 51906)0. 518.6 eneecalemmel

i Ow NAIDU, at yA oe (CBO e Iie a! Ole oS) a0

RK | sea s S09-1910)i94 10954) Om eal mloMaRT

POC cn me ORION a 1s 7 O

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

(une ONC) Meer rec 2.5 VON yee oun Onn,

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

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

separate papers, of which a list can be supplied, at about one-fourth their published price.

PROCEEDINGS OF THE COMMITTEE OF SCIENCE AND

CORRESPONDENCE OF THE ZOOLOGICAL SOCIETY OF

LONDON. 8vo. 2 vols. (Letterpress only).

Price to Price to the

Fellows. Public.

Part I. 1830-31. 1 vol. 8vo., out of print.

ali 832, Less oral Nae Oe SANs Ala 4s, 6d.) 2, Mest

PROCEEDINGS OF THE ZOOLOGICAL SOCIETY OF LONDON.

First SERIgEs.

Parts I—XV. (1833-1847). 8vo. 15 vols. (Letterpress only.) Price to

Fellows : 4s. 6d. each part ; to the Public, 6s.

Index 1830-1847. Price to Fellows : 4s. 6d.: to the Public, 6s.

Parts I., VII—-IX., XI., XIV., XV., out of print.

SECOND SERIES.

Parts XVI.-XXVIII. (1848-1860). 8vo. 13 vols. (Letterpress only.)

Price to Fellows: 4s. 6d. each part ; to the Public, 6s.

Index 1848-1860. Price to Feliows: 4s. 6d.; to the Public, 6s.

The Parts of this series containing Coloured Plates are out of print.

PROCEEDINGS OF THE SCIENTIFIC MEETINGS OF THE

ZOOLOGICAL SOCIETY OF LONDON. 8vo. 40 vols. and 4 Indices.

Letterpress only. With Plates uncoloured. With Plates coloured.

Price to Price to the Price to Price to the Price to Price to the

Fellows. Public. Fellows. Public. Fellows. Public.

USGI AS Oda: sian) GSS. since sce 9s. Boe mt She, aetna SB Bb ooog CRAP

MSGQM AMA SH Game) (BSs. arate ots 9s! BOON opeon a See thlsios 6 4ie

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SOL=1900 se Renkvolumesieny je achimeerneiscieietnten 36s. Geno oaks:

Imdexs SOT 1900) even ac ee Ast Oda ee cy wos:

* No perfect copies in stock. t Out of print.

PROCEEDINGS or tar GENERAL MEETINGS ror SCIENTIFIC

BUSINESS or tut ZOOLOGICAL SOCIETY OF LONDON.

8vo. 32 vols. and Index.

Price to Price to the

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GOIN Twenty volumesie seer aera Hachillesie seers 24s,

iheKtlehe) RON ee see boo condor oad date tale dbaac oY On ad 0 As 60. ome MOS:

TASH oie nannies Simian. i AMR Eero ot oid cl ain ad Sis een ee DAs.

ne Pte Caer ae CMC be Wel 1 a tine cANCK at Gin aid 18s. ee 24s.

HOM OP Mate tats echt gee Ue CaM coat eie Oee 186). 24s,

dient Pee | aren iar Wines eerie Cc c SORD poe re o ieReL aS ay A 4 LSS. teen 24s

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TO al et Ot sl eae er RRA EO re 168.5. nee eee 20s.

wy Biba py ud UR 2 en el are vane Eemech ec eiriear sa eaaties, ta 18sec eee 24s,

LISTS OF THE ANIMALS IN THE SOCIETY’S GARDENS.

List of the Vertebrated Animals now or lately Living in the Gardens

of the Zoological Society of London. (Kighth Edition.) 8vo.

1883. Cloth, 4s. 60.

List of the Vertebrated Animals now or lately Living in the Gardens

of the Zoological Society of London. (Ninth Edition.) 8vo.

1896. Cloth, 6s.

CATALOGUE OF THE LIBRARY of the Zoological Society of

London (Fifth Edition.) 8vo. 1902. Cloth, 6s.

THE OFFICIAL ILLUSTRATED GARDEN GUIDE— 18th Edition

(Revised)—with (1) a Railway and Street Map; (2) a Plan of

the Grounds; (3) a short description of some of the principal

animals in the Collection, with 52 Photographic Illustrations

and Index. Price 1s. in Stiff Paper Cover, postage 2d.

THE HOUSE-FLY CAMPAIGN. 3rd Edn. Illustrated. vo.

1916. 3d.

PRACTICAL ADVICE ON THE FLY QUESTION. 8yo. 1915.

2d.

ZOOLOGICAL RECORD.—Vol. 55, containing literature relating

chiefly to the year 1918, was published in November 1920.

Vol. 56, for the year 1919, is being prepared as usual.

P. CHALMERS MITCHELL,

Secretary.

Regent's Park, London, N.W. 8.

June, 1921.

These publications may be obtained at the Socrery’s OFrricz

or through any bookseller.

PAPERS.

1. The Comparative Anatomy of the Tongues of the Mammalia.—II. Family 1. Simiidz.

By Cnaruzus F. Sonnrac, M.D., Oh.B., F.Z.S., Anatomist to the Society. (Text-

HGR IED). “do ine e pno Bigg GO DUO OO Ub OG Bape Gd ooe aa aeide ec Pe ee pet a ois Sb!

2. Note on the Capture (in London) of a rare Parasitie Fly, Hammomyia (Hylephila)

unilineata Lett. By Lt.-Col. 8S. Monckton Copeman, F.R.S., M.D., F.R.O.P., F.Z.S.

(GRextaficurres1s) ei. weiss 3 6 SUE ERISA a NESE SA DI Co RIA enone Clio NC Fa Se ees

3. The Bases of Classification of the Theriodontia. By D. M. 8. Warson, F.ZS.,

University College, London. (Text-figures 1-29.) .... 00... eee ee ce ee eee ee ee

4. Experiments on Colour-changes of the Spotted Salamander (Salamandra maculosa),

condueted in the Society's Gardens. By E. G. Bounencnr, F.Z.S. (Curator of

Reptiles). (Plates I. & II., and Text-figure 1.).... 2.0... .... ce sees ce ee teens

5, Contributions to the Morphology, Classification, and Zoogeography of Indian Oligo-

cheta. By J. Srmpuenson, D.Sc., F.Z.S., Lecturer in Zoology in the University of

IBehhal onan, “(Aiea tfai an iy ibis cig pho obind oo nomd Joncon de cbne aoe

6. On the Structure of the Reptilian Tarsus. By R. Broom, D.Sc., F.R.S., C.M.Z.8.

(Text-figures 1-27.) ........00+. Be eseoraiepoict cua foleteie nine eVereliel suave denote Sistah cnet es oer ae

7. A Contribution to the Anatomy of the Three-toed Sloth (Bradypus tridactylus). By

Cuarves F. Sonnraa, M.D., Oh.B., F.Z.S., Anatoinist to the Society. (Plates I-IV.,

and Text-figures 10-15.) .......... DE NPS SOCAL ISD LUG SOLETIC OLD UO OSCR SOIE EO

8. Report on Deaths which occurred in the Zoological Gardens during 1920. By N. 8.

Lucas, M.B., F.Z.8., Pathologist to the Society.......-.-+eee++ «s- Siete Sisiatel stones

Page

103

143

179

List OF PLATES.

1921, Parr I. (pp. 1-186).

; Page

E. G. Bounencer: Pls. I. & II. Colour-changes of the Spotted Sala-

mander (Salamandra maculosa) .. 99

C. F. Sonntag: Pls. I-IV. Anatomy of the Three-toed Sloth

(Bradypus tridactylus) ......:.+. 157

G. Exuiot Sarrn: = PIT. Photographs of a living example of

LOPSUUS sos hn Aa e crore Eee eee 184

NOTICE,

The ‘ Proceedings’ for the year are issued in four parts, paged consecutively,

so that the complete reference is now P. Z. §. 1921, p.... The Distribution

is usually as follows :—

Part I. issued in March,

wlacfiate i) ena eR June.

sever GE Eanes September.

wernt) RG Tees December,

‘ Proceedings,’ 1920, Parts I. & II. (pp. 1-194), were published together on

July 21st, 1920. Part III. on September 15th, 1920. Part IV. on January

15th, 1921.

The Abstracts of the ‘ Proceedings,’ Nos. 211-212, are

contained in this Part.

13.

id,

16.

27.

18.

19.

20.

21,

22.

PAPERS.

. On the Reproductive Organs of the Ascidian Kiikenthalia borealis (Gotischaldt). By

Dr. Aveusta ArnpAck Onristiz-Linpr. (Text-figures 1-8.) .......... mVblehaaesieree

. On the Variation of the Scapula in the Batrachian Groups Aglossa and Arcifer. By

Joan B. Procrnr, F.Z.S. (Text-figures 1-10.) ............ efoieieitecne's SAGAR OCOD

. Notes on Marine Wood-boring Animals.—II. Crustacea. By W. T. Catan, D.Se.,

IAS Sr Reames BaP GHC ADO HOR OCH RGHE Odd SAR OUMOAIRA Gan Sa OC b Aaa oes

. Colour-production in relation to the Coloured Feathers of Birds. By A. Matiocg,

E.R.S., F.Z.S. (Text-figures 1-4.) ........-.... eo anche eee t tenn wes ececreenrs

Fossil Bird-remains collected by Dr. Forsyth Major in Sardinia, Corsica, and Greece.

By E. T. Newroy, F.R.S., F.G.S., F.Z.S.

sete were et ee ee ee sees seer ee eese Sess oe

The Oriental Species of the Genus Callistomimus (Coleoptera, Carabide). By H. E.

ANnpREWES. (Plate I.)

Ce CC i i me a ir a |

. On the Molluscan genus Cochlitoma, and its Anatomy, with remarks upon the Variation

of two closely-allied Forms. By G. C. Rozsson, B.A. (Text-figures 1-9.)

A Revision of the Melolonthine Beetles of the Genus Hetinohoplia. By G. J. Arrow,

Sipe mOn Hes eB iaenle uch eee ge eas pee nov eye eae

The Comparative Anatomy of the Tongues of the Mammalia.—III. Family 2.

Cercopithecids : with notes:on the comparative physiology of the tongues and

stomachs of the Langurs. By Cuartzes F. Sonnraa, M.D., F.Z.S., Anatoiist to the

Society. (Text-figures 14-36.) ..... Gino Oeste COREG SEO OER a bh OOe

Descriptions of New Moths from South-East Brazil. By E. Duxiyriexp Jones, F.Z.S.,

HIP So ss Ca lavcersllle abit les) etapeevarevaiever rier sros elesel vuci-esik es wolamricwerave oats sincere ete RAAT on Ne

On some new or little-known Acari, mostly Parasitic in Habit. By Sranuey Hirst,

B.Z.S. (Text-figures 1-15.) ................

Observations on the Habits of Cochlitoma zebra, var. fulgurata (Pfeiffer), and C. zebra,

var, obesa (Pfeiffer), in Confinement. By Janu Lonesrarr, F.L.S., F.G.S8. (Plates

AM = PESTS Pr aNenebeees eocraite ele veinier teins SHIPS scSpalersbaxclle dnote ce albeit, auticrm em ae lacah eh vo Me aime iaecbiac ars

The External Characters and Classification of the Procyonide. By R. I. Pocock,

B.R.S., F.Z.S. (Lext-figures 1-13.)

New or Little-known Reptiles and Batrachians from Southern Annam (Indo-China).

By Matcorm A. Switn, M.R.C.S., L.R.C.P., F.ZS. (Plates I-II. and Text-

figures 1-2.) .... Peete eee ee etre eee ee cece ete ee teen teen as OOS Sc ickartar

Mitlepage ..e6.. +s MENG She ia ee ae Seana Vaairke eesti Beare Sater ct. Palate igiars canta

Maso @ourciland Officers,” si. iverslsiccre « o:cpisigle cia a velev satis afer aie Rife rare) els) si cterstalatalcuers

Alphabetical Aint of Contributors gee bre pes: Reeth oe ig abe Gees aes ae

Index of Illustrations ........ aSincoe oon aFattes Whaleleinceniel aexciererene eke aee Salona c Yoel cia’ y nett ate

Index ..... Ri (eeeTeicie ial s Basal abalcnstelaiaes Srojstatelbrelaie ale atsreleiste 7 ie afoeiaere

187

197

215

221

229

233

249

267

277

323

357

379

389

423

LIST OF PLATES.

1921, Parr IL. (pp. 187-446).

Page

H. E. ANDREWES: Pl. I. Oriental Species of the Genus

Callistomimus .......s+0 eeoo

G. J. Arrow: Pia Beetles of the Genus Eetinc-

Hop iaen So okie Geto eee te 267

BK. Durinrietp Jones: LPls.J.-II1I. New Moths from South-East

Brazil) oes he es ee

JANE LONGSTAFF: Pls. 1.-I1I. Cochlitoma zebra, var. obesa and

Cochlitoma zebra, var. ful- _

GUT OLL Nate scien otal caine eee 379

Maucoum A, Satu: Pls. 1-II. New or little-known Reptiles

and Batrachians from

Southern Annam.......... 423

NOTICE.

The ‘ Proceedings’ for the year are issued in four 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.

Apia 6d (eases June,

Pye Obi meee September.

Pema! bi eeu December,

‘Proceedings, 1921, Part I. (pp. 1-186), was published on April 2nd,

1921.

The Abstracts of the ‘Proceedings, Nos. 213-216, 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 ‘ Transactions,’ 1833-1869, will be found in the

‘ Proceedings’ for 1913, page 814. ag

&1i-.